The Barbary Macaque A Case Study in Conservation

Barbary macaque mother and infant (E.M.)

The Barbary Macaque A Case Study in Conservation

Edited by

John E. Fa Animal Ecology Research Group

University of Oxford Oxford, United Kingdom

Plenum Press • New York and London

Library of Congress Cataloging in Publication Data

International Conference for the Conservation of the Barbary Macaque (1982: Gibraltar) The barbary macaque.

"Based on papers presented at the International Conference for the Conservation of the Barbary Macaque, held June 16-20, 1982, in Gibraltar"-Verso of t.p.

Includes bibliographical references and indexes. 1. Barbary ape-Congresses. 2. Wildlife conservation-Congresses. 1. Fa, John E. II. Title.

QL737.P96157 1982 639.9'7982 84-11618

ISBN-13: 978-1-4612-9718-5 DOl: 10.1007/978-1-4613-2785-1

e-ISBN-13: 978-1-4613-2785-1

Based on papers presented at the International Conference for the Conservation of the Barbary Macaque, held June 16-20, 1982, in Gibraltar

© 1984 Plenum Publishing Corporation

Softcover reprint of the hardcover I st edition 1984

233 Spring Street, New York, N.Y. 10013

All rights reserved

No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical,

photocopying, microfilming, recording, or otherwise, without written permission from the Publisher

Aux peuples nord~africains

FOREWORD

The Barbary macaque (all too often mistakenly called an ape) was first brought to the attention of the Conservation Working Party of the Primate Society of Great Britain late 1979 when John Fa reported that 'surplus' animals were being sent from Gibraltar to dubious locations, such as an Italian safari park. Since there had been no scientific input into the Army's management of the monkey colony on Gibraltar, and there was concern about inbreeding, nutrition and health - about the long-term viability of the colony, it was felt that the Society could help. The Gibraltar Scientific Authority and the Army were very receptive to our offer and ideas, and this topic occupied successive chairmen over the last few years - Robin Dunbar and Richard Wrangham, myself and now Miranda Stevenson - with constant prompting and help from John Fa.

Considerations soon extended to the status of the species as a whole, so that there have been three main aspects:- (1) the improved health of a larger self-sustaining population on Gibraltar, (2) the status and behavioural biology of natural populations in North Africa (Morocco and Algeria), and (3) the breeding achievements in European parks and zoos, and their potential for reintroduction to suitable areas in North Africa, along with other possibilities.

Robin Dunbar organized the compilation of recommendations for managing the Gibraltar colony with regard to numbers, age-sex struc­ture and behavioural relationships, with some observations on diet to avoid obesity and infertility. Following concern over the pro­cedures adopted in reintroducing 200 Barbary macaques from a French park to Morocco, I encouraged Julian Caldecott and Michael Kavanagh to produce guidelines for the reintroduction of ~rimates to the wild, which played an important part in the Gibralter conference in June 1982.

This conference represented the peak of our efforts, especially John Fa's. With help from the Gibraltar authorities, World Wildlife Fund and the Fauna and Flora Preservation Society, John Fa organized an excellent meeting, where all three topics were thoroughly consid­ered and integrated. It was especially stimulating for the successful

vii

viii FOREWORD

blending of the expertise of field and laboratory workers, zoo people and conservationists. At the time its success was reflected by the production of eight hard-hitting Resolutions that effectively covered the field and which have been distributed widely, along with the reintroduction guidelines.

The key issues that emerged during the conference were the needs to make further progress in (1) evaluating distribution and habitat preferences, (2) assessing population densities and carrying capaci­ties in different areas, (3) studying population dynamics and repro­ductive biology and (4) relocating surplus animals in the natural habitat, parks or zoos. These required (1) the identification of conservation areas and the creation of national parks, (2) long-term socio-ecological studies in different habitats, (3) changes in national and international law and (4) increased coordination of activity and funding by national and international agencies. John Fa's exciting proposals for a Monkey Park on Gibraltar, which would provide a sanctuary for all native plants and animals as well as an educational and research centre, also merit acceptance and implementation.

Now we have this substantial volume that presents our current knowledge of this monkey and publicizes its plight even more widely. It represents an integrated approach to the problems of a primate and its biology, its habitat and the needs of the local people, and its captive maintenance. It should serve as a model for comparable problems that are all too common nowadays throughout the tropical world. I have been pleased to be associated with such developments, and congratulate John Fa and his colleagues on their achievements, hoping that they will go on to greater successes for the benefit of the forests and wildlife of North Africa, and, hence, for the people.

David J. Chivers

Cambridge October, 1983

PREFACE

It is ironical that a species which has figured so largely in the history of Mediterranean civilizations as the Barbary macaque should still be in need of appraisal. It is unfortunate that this appraisal has become an urgent matter only because the very survival of the species is now in jeopardy. To develop steps to maintain or improve the status of a threatened species requires thorough research into the species' requirements as well as into the pressures acting upon it. This book presents all known. data on the Barbary macaque in the hope that by promoting a deeper understanding of its precar­ious status conservation work on the species will be facilitated. It is a compilation of papers on the position of the Barbary macaque in the wild and in captive and semi-natural environments, most of which were presented at a conference on the conservation of the species held in Gibraltar from 16-20 June, 1982. The meeting was sponsored by WWF/IUCN, ffPS, New York Zoological Society and the Gibraltar Government.

The data presented here are not only relevant to the Barbary macaque, they exemplify most importantly the situation of most species in the interface between the vulnerable and endangered category and, in fact, relates to all forest species in North Africa. The book is a first in that both sides of the conservation effort, captive propagation and protection in the wild, are discussed as complementary issues.

Chapter 1 is a general introduction to the Barbary macaque. It provides a review of the taxonomy, behaviour and general ecology of the species. Part I looks at the situation of the monkey in the wild. In Chapter 2, Thirgood lays the groundwork for the section by describing the status and history of the North African forest, the Barbary macaque's main habitat. He emphasizes the role that man has played in determining the present day distribution of species. Taub (Chapter 3) outlines the changes that have occurred in the dis­tribution of the macaque throughout its range from the historical literature, to explain the present day extent of the monkey. Fa, Taub. Menard and Stewart (Chapter 4) present the most recent survey figures on monkey numbers and habitat condition for the Barbary

ix

x PREFACE

macaque throughout North Africa. These figures forecast a bleak future for most monkey populations except those in the Moyen Atlas and in the Algerian Kabylies.

The demography of the more viable monkey populations in North Africa, those in the Moyen Atlas cedar forests, is discussed by Deag in Chapter 5. Drucker (Chapter 6) exposes the conflict that exists in conserving the macaque in timber forests, where the monkey is con­sidered a forestry pest. He rejects this claim on the basis of quantitative data on the feeding ecology of the monkeys in a cedar forest in the Moyen Atlas. Mehlman (Chapter 7) takes up the theme of the effect of man on the habitat and monkey population. His study site is also a conifer forest, in the Moroccan Rif mountains, a prime example of human disturbance and of overt competition between humans and the macaque. This habitat is of special significance in that it is unique to Morocco.

Part II is a forum for discussion and description of the Barbary macaque in captive and semi-natural environments. Stevenson (Chapter 8) starts by examining the roles of zoos and wildlife parks in pro­moting conservation of primates and comes to the unfortunate con­clusion that, in general, captive propagation is not successful as yet. As regards the Barbary macaque the species is numerically safe but there are concomitant problems which Stevenson discusses. Hornshaw (Chapter 9) considers the keeping of monkeys in captivity and how this influences their behaviour and their social structure. Turckheim and Merz (Chapter 10) follow by describing the unprecedented success of three large enclosures in Europe in breeding Barbary macaques. They argue that these enclosures could be of significant value not only as a reserve stock for reintroduction into the wild but also as centres for conservation education and scientific study, which are economically independent.

In Chapter 11, Fa discusses the status of the Barbary macaque population in Gibraltar and presents data on the demography of the two component troops since 1936. The structure and dynamics of the two separate monkey troops that compose the population are seen in relation to the amount of provisioning and human disturbance each have received since their start. Burton and Sawchuk (Chapter 12) give the genetic condition of the Gibraltar population of Barbary macaques through estimates of the effective population sizes and loss of heterozygosity during a period of more than 20 years.

The final part of the book, conclusions and recommendations, summarizes the foregoing information and formulates objectives and regions where future action could be taken. These recommendations are drawn largely from those which were outlined at the conference in Gibraltar in 1982.

PREFACE xi

Although this book does not set out to provide answers to specific practical management problems it is hoped that in examining the progress and possible lines of action for the conservation of the Barbary macaque and its habitat it will go some way towards its aim of bridging the gap between conservation biology as a scientific discipline and practical social and ecological solutions.

John E. Fa

ACKNOWLEDGEMENTS

Acknowledgements and thanks are due to a large number of persons and organizations for collaboration and financial support in the pro­duction of this book. The following persons are thanked for their encouragement and advice in making this publication possible: Dr. David J. Chivers (University of Cambridge and Chairman of Primate Society of Great Britain Working Party for Conservation), Dr. Russell Mittermeier (WWF-US), Mr. Bill Konstant (WWF-US), Mr. Pierre Portas (WWF-Switzerland), Dr. Harmut Jungius (WWF-Switzerland), Dr. Michael Kavanagh (IUCN Monitoring Centre, Cambridge), Dr. Walter Angst (Affen­berg Salem), members of the PSGB Working Party for Conservation and all delegates that attended the Gibraltar Barbary Macaque Conference.

Thanks are due to the World Wildlife Fund, the International Union for the Conservation of Nature and Natural Resources, the Flora and Fauna Preservation Society of Great Britain, the Gibraltar Govern­ment and the New York Zoological Society for financial help in the convening of the Barbary Macaque Conference at Gibraltar. This has been the main venue from which most papers included in this book are taken.

The chapters were revised by Dr. F. Alvarez, Dr. M.S. Boyce, Dr. F. Braza, Dr. M.J. Coe, Dr. J.M. Deag, Dr. B.A.C. Don, Dr. P.G.H. Evans, Dr. A. Gautier, Dr. C.W.D. Gibson and Dr. W.C. McGrew. All are gratefully thanked for their helpful discussion. To Mr. Michael Amphlett lowe my greatest appreciation for his excellent draftsman­ship and Mrs. Mary Aldworth and Mrs. Joy Johnson have more than earned the highest esteem for their careful typing and well-placed creativity in producing the book.

My very special thanks go to my wife, Ms. M. Williamson, for neverending encouragement and assistance in correcting and editing this publication. To her I am most deeply indebted.

CONTENTS

List of Plates

GENERAL INTRODUCTION

Chapter 1. The Barbary Macaque J.E. Fa

PART I: THE BARBARY MACAQUE IN THE WILD

Chapter 2. The Demise of Barbary Macaque Habitat -Past and Present Forest Cover of the Maghreb •

J.V. Thirgood

Chapter 3. A Brief Historical Account of the Recent Decline in Geographic Distribution of the Barbary Macaque in North Africa • • • • • • •

D.M. Taub

Chapter 4. The Distribution and Current Status of the Barbary Macaque in North Africa • • • •

J.E. Fa, D.M. Taub, N. Menard, and P.J. Stewart

Chapter 5. Demography of the Barbary Macaque at Ain Kahla in the Moroccan Moyen Atlas ••• •

J.M. Deag

Chapter 6. The Feeding Ecology of the Barbary Macaque and Cedar Forest Conservation in the Moroccan Moyen Atlas .. . . . . . . • . . . . . . . • .

G.R. Drucker

Chapter 7. Aspects of the Ecology and Conservation of the Barbary Macaque in the Fir Forest Habitat of the Moroccan Rif Hountains ••••

P.T. Mehlman

xiii

xv

3

19

71

79

113

135

165

xiv

P ART II: THE BARBARY MACAQUE IN CAPTIVE AND SEMI-NATURAL ENVIRONMENTS

Chapter 8. The Sense and Direction of Captive Breeding Programmes - The Position of the Barbary Macaque • • • • • • • • • • • • •

M.F. Stevenson

Chapter 9. A Comparison of Proximity Behavior in Two Groups of Barbary Macaques - Implications for the Management of the Species in Captivity

S.G. Hornshaw

Chapter 10. Breeding Barbary Macaques in Outdoor Open Enclosures . ... . . . . . . . . . • . . .

G. de Turckheim and E. Merz

Chapter 11. Structure and Dynamics of the Barbary Macaque Population in Gibraltar • • • •

J.E. Fa

Chapter 12. The Genetic Implications of Effective Population Size for the Barbary Macaque in Gibraltar • • • • • • • • • • • • • • • • •

F.D. Burton and L.A. Sawchuk

CONCLUSIONS AND RECOMMENDATIONS

Chapter 13. Conclusions and Recommendations • • J.E. Fa

Appendix I: Definition of Age-Sex Classes for the Barbary Macaque • • • •

Appendix II: Diet of the Barbary Macaque in the Wild •

Appendix III: Variant Spelling of Place Names Mentioned in the Text ••••

Contributors • •

Index . . . . . . . . . . . . . .

CONTENTS

203

221

241

263

307

319

335

347

357

359

361

LIST OF PLATES

Plate I. Early 19th century plate of a Barbary macaque.

Plate II. Diagrams from an Islamic treatise on human anatomy depict­ing dissections based on Galen's anatomical drawings of the Barbary macaque (from the Tashri-i-mansuri by Mansur ibn Muhammed Ahmad, 11th/17th century Persian}ffi No 5266, in the Majles Library, Tehran).

Plate III. Male care of Barbary macaque infants, a) adult male sitting with infant; b) teeth-chattering interaction between adult male and infant (EM).

Plate IV. Triadic interactions between Barbary macaque males and infants (GdeT).

Plate V. Barbary macaque habitats, a) scrub, Rif, Morocco; b) cliffs, Rif, Morocco (JEF); c) scrub, Haut Atlas, Morocco (JMD); d) rocky slopes, Haut Atlas, Morocco (JMD); e) mixed oak forest, Rif, Morocco (JEF); f) fir forest, Rif, Morocco (JEF); g) fir forest and high mountain rocky slopes, Rif, Morocco (JEF); h) cedar forest, Moyen Atlas, Morocco (JMD).

Plate VI. Ain Kahla, a) aerial view of the Ain Khala study site showing the distribution of the cedar forest and grassland areas. Forestry post is encircled. b) and c) two views from the Ain Kahla forest looking towards the forestry post in the centre of the study site. The foreground shows a typical spur running down from the forest (JMD).

Plate VII. Barbary macaques in the Ain Kahla cedar forest, a) and b) Barbary macaque troop travel feeding below the forest canopy (JMD); c) use of tall cedar trees for social interaction (JMD).

xv

xvi LIST OF PLATES

Plate VIII. Cedar forest habitat at Bou Jirrir, a) semi open cedar forest (GRD); b) relict cedar and holm oak forest (GRD); c) use of forest area by sheep (GRD).

Plate IX. Barbary macaques feeding in cedar forest at Bou Jirrir, a) adult male feeding on Portuguese oak acorns (GRD); b) adult female feeding on underground food items (GRD). c) adult male debarking conifer (JH).

Plate X. Rhomaran fir forest, a) view of the Talassemtane fir forest, north side (PTM); b) south-facing slopes of the Talassemtane fir forest with holm oak scrub (JEF); c) typical forest structure of the Talassemtane fir forest (JEF); d) grazing area around stream in the interface between north and south facing vegetation at Talassemtane forest (JEF).

Plate XI. Two views of the cliff habitat at Talassemtane forest (PTM) •

PlateXII. Human activities in the Rhomaran fir forest, a) pole logging (PTM); b) beam logging (PTM); c) goat herding (PTM) .

Plate XIII. Damage caused to the Rhomaran fir forest by a) timbering and pruning of trees (PTM); b) winter snows and low temperatures (PTM); and c) erosion and overgrazing (PTM).

Plate XIV. Spacing behaviour of Barbary macaques in a) Gibraltar, adult female Dorothy with her infant I and infant 2 -animals part of Matriline A (SGH); b) Toronto Zoo, adult female and infant 2 threatening infant 2 of another group (SGH).

Plate XV. Enclosure habitats, a) La Montagne des Singes, Kintzheim (GdeT); La Foret des Singes, Rocamadour (GdeT); c) Affenberg, Salem (GdeT).

Plate XVI. Interaction between Vlsltors and monkeys in the French and German Barbary macaque enclosures (GdeT).

Plate XVII.Gibraltar, a) eastern scarp slope and b) western dip slope overlooking the town area (JEF).

Plate XVIII. Use of Barbary macaques in street shows and as pets in North Africa (JEF) and (GRD).

Plate XIX. Birth to two years of age, a) newborn baby (AP); b) 6th week (AP); c) 11th week (AP); d) 14th week (AP); e) 18th week (EM); f) 2-3 yrs. (EM).

LIST OF PLATES

Plate xx. Males, a) 5-6 yrs. (EM); b) 7-8 yrs. (EM); c) and d) 10-11 yrs. (EM); e) 18+ yrs. (EM).

xvii

Plate XXI. Development stages of dentition in Barbary macaque males, a) 4-5 yrs. (EM); b) 5-6 yrs. (EM); c) 6-7 yrs. (EM); d) 18+ yrs. (EM); e) example of fully developed male dentition (EM) •

Plate XXII.Females, a) 4-5 yrs., primiparous (EM); b) 4-5 yrs., nulli­parous (EM); c) 6-7 yrs., mUltiparous (EM); d) 18+ yrs. (EM); e) 20+ yrs. (EM).

Plate XXIII.Female sexual swellings, a) 3-4 yrs. (EM); b) 4-5 yrs. (EM); c) 5-6 yrs., maximal swellings, during mating season (AP); d) 8-9 yrs. (EM); e) 11-12 yrs., charac­teristic presenting pose of swollen female (EM); f) 15-20 yrs. (AP); g) very old female (20+ yrs.) with only slight swelling (AP).

Picture credits: AP = Andreas Paul; EM = Ellen Merz; GdeT = Gilbert de Turckheim; GPn = Graham R. Drucker; JEF = John E. Fa; JH = Jurgens Hesse; JMD = John M. Deag; PTM = Patrick T. Mehlman; SGH = Susan G. Hornshaw.