Gorilla Pathology and Health: With a Catalogue of Preserved Materials

Gorilla Pathology and Health

John E. Cooper, DTVM, FRCPath, FRSB, CBiol, FRCVS

RCVS Specialist in Veterinary Pathology Diplomate, European College of Veterinary Pathologists Diplomate, European College of Zoological Medicine, Honorary Research Fellow, Durrell Institute of Conservation and Ecology, The University of Kent, Canterbury, United Kingdom

Wildlife Health, Forensic and Comparative Pathology Services, United Kingdom

With a Catalogue of Preserved Materials

Gordon Hull

Gorilla Pathology Study Group, London, United Kingdom

Table of Contents

Cover image

Title page

Copyright

Dedication

Quotations

Foreword

Preface

Acknowledgements

About the Authors

About the Contributors

List of Abbreviations

Part I: Gorilla Pathology and Health

Chapter 1. The Genus Gorilla – Morphology, Anatomy and the Path to Pathology

Abstract

Introduction

Primate and Comparative Anatomy

The Current Status of Anatomy

Terminology

Conclusions

Chapter 2. The Growth of Studies on Primate Pathology

Abstract

Introduction – The Growth of Pathology

Studies on the Pathology of Primates

African Studies

The Importance of Comparative Pathology

Primates in Biomedical Research

Studies on Microscopic Anatomy and Cellular Pathology

The Continuing Relevance of Studies on Pathology

The Advantages and Dangers of Extrapolation

Access to Data

The Relevance of Pathology to Health

Chapter 3. Infectious Disease and Host Responses

Abstract

Introduction

Disease and Death in Gorillas

The Aetiology of Disease

What Is ‘Pathology’?

Understanding Host Responses – A Comparative Approach

Endogenous Factors Influencing Host Responses

Discussion of Infectious and Noninfectious Diseases

Bacteria

Viruses

Parasites (By Ian Redmond)

Microbiota, Microbiome and Normal Flora

Emerging Diseases

Chapter 4. Noninfectious Disease and Host Responses

Abstract

Introduction – Noninfectious Conditions

Trauma – Causes and Pathogenesis

Specific Responses – Trauma

Fighting

Infected Wounds

Attacks on Humans

Wounds – General

Ageing of Wounds

Healing and Regeneration

Sepsis

Snares

Firearms, Exposure to Blasts, Explosions and Unexploded Ordnance

Predation

Exposure to Temperature Changes

Stress and Stressors

Chapter 5. Methods of Investigation – Observation, Clinical Examination and Health Monitoring

Abstract

Introduction

Surveillance of Syndromes

Clinical Work With Gorillas

The Importance of Observation

Indicators of Ill Health

Restraint and Handling

Clinical Examination

Anaesthesia

Medical and Surgical Treatment

The Importance of Multidisciplinary Collaboration

Health Monitoring of Gorillas

Why Is Health Monitoring Important?

Health Monitoring Techniques

Some Conclusions

Chapter 6. Methods of Investigation – Postmortem Examination

Abstract

Introduction

The Nature and Role of the Postmortem Examination

Before Embarking on a Necropsy

Personnel and Training

Interdisciplinary Studies

Access to Material for Postmortem Examination

Postmortem Procedures

Equipment

Recording of Findings

Performance

External Examination and Sampling

Internal Examination and Sampling

Description of Lesions

Forensic Postmortem Examinations

Postmortem Changes and Determination of Time of Death/PMI

Causes of Death Diagnosis

Chapter 7. Methods of Investigation – Sampling and Laboratory Tests

Abstract

Introduction

Sampling

Methods

Environmental Testing

Quality Control

Recording and Reporting of Findings

Personnel and Equipment

Safety in Laboratory Work

Storage of Samples and Reference Collections

Chapter 8. Nonspecific Pathology

Abstract

Introduction

General Considerations in Pathology, Not Related to Organ Systems

Age-Related Changes

Developmental Abnormalities

Genetic Anomalies

Behavioural Pathology

Aggression

Effects of Capture, Transportation and Captivity

Welfare

Pain

Neoplasia

Poisoning

Chapter 9. Skin and Integument

Abstract

Introduction

Biology of the Skin

Investigative Methods

Diseases and Pathology of the Skin – General

Noninfectious Causes

Infectious Causes

Ectoparasites (By Ian Redmond)

Diseases of Uncertain Aetiology

Conclusions

Chapter 10. Respiratory and Cardiovascular Systems

Abstract

Introduction

Respiratory System

Lung Parasites (By Ian Redmond)

Investigation of the Respiratory Tract

Cardiovascular System

Investigation of the Cardiovascular System

Cardiovascular Disease

Research on Heart Disease in Great Apes, Including Gorillas

Pathogenesis of Cardiovascular Disease

Other Pathology of the Vascular System

Chapter 11. Alimentary Tract and Associated Organs

Abstract

Introduction

Oral Cavity

Dental Disease and Pathology

Some Specific Conditions and Their Pathogenesis

The Gastrointestinal Tract

Investigation of the Liver

Pathology and Pathogenesis of Hepatic Disorders

Investigation of the Biliary System

Investigation of the Exocrine Pancreas

Chapter 12. Lymphoreticular and Haemopoietic Systems and Allergic Conditions

Abstract

Introduction

The Lymphoreticular System

Immunity

Immunological Disorders

Hypersensitivities and Allergies

Immunodeficiency (by Geoffrey Pearson and John E. Cooper)

Blood and Bone Marrow

Blood Parasites (by Ian Redmond and John E. Cooper)

Neoplasia

Chapter 13. Urinary and Reproductive Systems

Abstract

Introduction

Urinary System

Reproductive System

Chapter 14. Musculoskeletal System

Abstract

Introduction

Movement Disorders

The Pathology of Bone

The Importance of the Skeleton

Skull

Vertebral Column

Joints

Investigation of Bones

Skeletal Disease and Pathology

Fractures and Their Repair

Specific Diagnosis of Diseases of the Skeleton

Muscle

Connective Tissue

Adipose Tissue

Chapter 15. Nervous System and Special Senses

Abstract

Introduction

Psychological Studies on Gorillas

Studies on the Brain of Gorillas

Investigation of the Nervous System

Pathology and Diseases of the Nervous System

The Special Senses

Chapter 16. Endocrinological and Associated Conditions

Abstract

Introduction

The Endocrine System

Sleep

Stress

Chapter 17. Field Studies in Pathology and Health Monitoring

Abstract

Introduction

Pre-release Health Considerations for Gorillas

Quarantine

Setting-Up Long-Term Health Monitoring Systems for Mountain Gorillas Using a One Health Approach

Cryptosporidium, Giardia and Helminths of Habituated Nyakagezi Mountain Gorilla Group in Mgahinga Gorilla National Park

Working With Great Apes During an Ebola Outbreak

Chapter 18. Legal Considerations

Abstract

Introduction

General

Free-Living Gorillas

Gorillas in Captivity

Other Relevant Law

Chapter 19. Pathology, Health and Conservation – The Way Forward

Abstract

Introduction

Knowledge of the Genus and Its Biology

Threats to Survival

The Implications and Importance of Studies on Pathology and Health

Advances in Health Care

Collaboration and Sharing of Information

A Greater Role for Africa

Involvement of African Academic Institutions

Scientific Advances

Human Health

Attention to Welfare

Continued Search for Gorilla Material

Greater Dissemination of Information

Broadening the Holistic Approach

Concluding Remarks

Part II: A Catalogue of Preserved Materials

Introduction to the Catalogue

Scope and Method of Compilation

Layout

Gorilla History, Taxonomy and Synonymy

Hanno’s gorillai

Etymological Considerations

Behavioural Considerations

Correspondents

Catalogue of Preserved Gorilla Materials

Argentine Republic

Australia

Austria

Belgium

Brazil

Bulgaria

Canada

Central African Republic

Channel Islands

Chile

Cuba

Czech Republic

Denmark

Egypt

Finland

France

Germany

Hungary

India

Ireland

Israel

Italy

Japan

Kenya

Kuwait

Morocco

Netherlands

New Zealand

Norway

Poland

Portugal

Romania

Russian Federation

Rwanda

Slovenia

South Africa

Spain

Sweden

Switzerland

Uganda

United Kingdom

United States of America

Uruguay

Zimbabwe

Appendix 1. Glossary of Terms

Introduction

List of Terms and Meanings

Appendix 2. Protocols and Reports

Introduction

Postmortem Examination of Gorillas – Some Basic Rules

Nonhuman Primate Postmortem Examination

Cardiac Examination for Great Apes (and Other Primates In Which Cardiac Disease Is Present)

Postmortem Examination of Nonhuman Primate Fetuses and Neonates

Postmortem Examination of the Air Sacs of Orangutans and Other Nonhuman Primates

Gorilla Postmortem Examination

Cardiac Examination for Apes and Other Primates

Postmortem Examination of Primate Fetuses, Neonates and Placentas

Postmortem Examination of the Air Sacs of Gorillas and Other Apes

CNS/Musculoskeletal/Other

Guide to the Nonhuman Primate Postmortem Examination Tips for Tissue Collection During the Necropsy Examination

Great Ape Tag Placental Examination

Postmortem Examination of the Air Sacs of Apes

GAHP Recommended Cardiac Necropsy Prosection Guide

Ape Cardiac Necropsy Protocol

Decision Point

GAHP Recommended Cardiac Trimming Protocol for Pathologists

GAHP Cardiac Necropsy Check Sheet (Fillable PDF) – Pathologists: Please Submit With Report, Slide Key and Images

Skeletal Examination Form

Appendix 3. Field Pathology

Introduction

Appendix 4. Hazards, Including Zoonoses

Introduction

Appendix 5. Case Studies – Museums and Zoological Collections

Introduction

Studies on the Skeleton of ‘Guy’ at the NHM

Studies on Three Skeletons in the Osman Hill Collection

The Research Proposal

Initial Observations on the Specimens

Sampling

Processing the Samples for Histological Examination

Processing of Samples for DNA Testing

Results

Acknowledgements

Student Projects, Bristol, United Kingdom, 2015

Project 1: A Review of the Data Relating to the Animal Management and Welfare of Gorillas at Bristol Zoo Gardens From 1930 to 2015

Project 2: The Development of Osteoarthritis in Captive Primates

Acknowledgements

Appendix 6. Scientific Names of Species and Taxa Mentioned in Text

References and Further Reading

References

Appendix CA1. Use of Collections and Handling of Biological Material

Introduction

The Needs Pertaining to Museums and Collections

The Rise and Fall of Collections

The Need for Pathology-Based Studies

Working in Collaboration With Museums

Making Maximum Use of Material in Museums

Handling Preserved Material

Appendix CA2. Retrieval, Preparation and Storage of Skeletal and Other Material

Introduction

Skeleton

Soft Tissue Preservation

Skins

The Recovery and Preparation of Skeletal Remains

Invited Contribution – Skeletal Remains: Recovery, Preparation and Preservation

Purposes of Recovery and Preservation of Skeletons

Recovery of Skeletons From Fleshed Animals

Preparation, Preservation and Storage of Recovered Skeletons

Factors Affecting the Preservation of Material

Comments on Postmortem Change

Index

Copyright

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Dedication

This book is dedicated to those of all backgrounds, beliefs and races who have contributed to our knowledge and understanding of gorillas, and thereby enhanced the health, welfare and conservation of these endangered species. We remember particularly those who have died during the course of their work.

Thank you very much

Merci beaucoup

Asante sana

Murakoze cyane

Quotations

Read not to contradict and confute; nor to believe and take for granted; nor to find talk and discourse; but to weigh and consider.

Sir Francis Bacon (1561–1626)

Almost all the wise world is little else in nature but parasites and sub-parasites.

Ben Jonson (1606)

Foreword

Louise Leakey, Research Professor, Stony Brook University, Turkana Basin Institute

Emmanuel de Merode, Director, Virunga National Park/ Parc National des Virunga

This is a book about gorillas, those close relatives of human beings which are confined to the continent of Africa, that have become emblematic of the remarkable conservation and sustainable development effort, and yet whose future survival remains largely uncertain.

Gorillas have only been known to science for a little over 150 years, whereas other great apes, such as the chimpanzee and orangutan, were recognised several centuries ago.

During the first 100 years, knowledge of the biology of gorillas progressed from meticulous studies on their anatomy (often comparing their morphology with that of other extant and extinct hominins), followed by research on behaviour – both in the wild and in captivity. Only over the past few decades has serious attention been paid to the factors that influence the survival of gorillas in the wild.

It was Dr Louis (LSB) Leakey who first advocated that a greater understanding of gorillas (and other great apes) and their relevance to the evolution of Homo sapiens required detailed fieldwork. Encouraged by the early studies of Carl Akeley and the seminal research of George Schaller, Louis Leakey recruited the late Dian Fossey, who subsequently played an instrumental role in bringing the plight of the species to the world’s attention. Alongside the achievements of these scientists, the work of Central African conservation practitioners has played the critical role in the survival of the species, albeit less recognised and celebrated. The outcome has seen some remarkable successes, such as the dramatic increase in mountain gorilla populations since their catastrophic low point in 1985. The circumstances of this achievement, which include the Rwandan genocide and the horrors of four civil wars in the eastern Democratic Republic of Congo, make it one of the great miracles of modern conservation. But this success has come at a tragic price, such as the 140 of Virunga National Park’s rangers who have lost their lives while protecting these species and their ecosystems.

The conservation effort has had multiple outcomes, of which the most important has been the contribution of gorilla populations across their range, in supporting the livelihoods of the people living in and around their habitats. The combined efforts of African and international conservationists has seen the growth of a remarkable industry generating tens of millions in revenue for post-conflict economies and offering quality employment for a new generation of young professionals from rural communities. Yet this opportunity carries with it enormous risks. While gorilla viewing plays a determining role in their conservation, tourism also implies the close proximity of human visitors to gorilla groups, exposing them to the greatest of all threats, disease. It is comforting to see the depth of understanding contained within this work that complements the substantial efforts by veterinarians and rangers from the various park services and highly committed partners, such as the Mountain Gorilla Veterinary Project, who provide the data, knowledge and experience that are the foundation of good science.

In this book, John Cooper and Gordon Hull bring together much of the available information about the pathology of gorillas – that is, how infectious and non-infectious diseases (including injuries) manifest themselves as well as how gorillas respond – and the influence of such complex interactions on the health, welfare and conservation of these threatened creatures. The chapters of the book are written mainly by John Cooper, a specialist wildlife veterinary pathologist, supported by contributions from Margaret Cooper, Ian Redmond, Jane Hopper, Gladys Kalema-Zikusoka, Jenny Jaffe, Celsus Sente and Ogeto Mwebi. The text is supplemented with a unique catalogue of information as to the whereabouts internationally of skulls, skeletons and skins, fluid-preserved specimens, casts, histological sections and samples, together with images and clinical, postmortem and laboratory records. Such data will be of great value to primatologists, veterinarians, evolutionary biologists and osteologists, as well as to field conservationists.

We welcome this book, not only because it is an important contribution to gorilla and great ape biology but also because it should help in much-needed attempts to save these magnificent animals from extinction. It also will assist those who are involved in human medicine, not least of all because large numbers of free-living gorillas have died in West Africa from Ebolavirus infection in recent years; a proper understanding of the epidemiology of this distressing, highly fatal, disease requires knowledge of wildlife that may either harbour or transmit the causal organism.

Living and working in Africa, as we both do, we appreciate the emphasis throughout the text on the importance, in studies on gorillas and other wildlife, of involving local people, ranging from veterinarians and scientists to trackers and village leaders. Only with such an environmentally sensitive approach, coupled with interdisciplinary scientific research, will the vital work of saving and conserving the two species of gorilla prove a success.

January 2016

Preface

John E. Cooper and Gordon Hull, London, England

The more you know, the more you know you don’t know.

Aristotle 384–322 BC

Why a book about the pathology of gorillas? And why is it accompanied by a catalogue of materials and resources? The rationale is straightforward. We know so little about the pathology of gorillas. Yet the answers to some of the most pressing questions posed by conservationists may lie hidden in existing collections of material, which can provide valuable data on pathology and diseases. What can be learned? Where can collections be found? This publication is intended to answer these questions and play a part in rectifying the current dearth of information.

The two species of gorilla that exist in Africa today are, like all the world’s great apes, threatened by persecution, habitat destruction, illegal trade and diseases.

It is conventional wisdom that attempts to halt the decline of any species require an understanding of the biology of that taxon and, based on this knowledge, the application of appropriate conservation measures. An aspect of the biology that has tended to be overlooked in the past is the susceptibility of a species to infectious and noninfectious diseases, how it responds to such ‘insults’ and, insofar as pathogens are concerned, the intricacies of host-parasite relationships. In the case of the genus Gorilla, these matters have only recently attracted serious scientific attention, and much remains to be learned.

The genesis of the book goes back to 1994 when, following the evacuation of the senior author and his wife (the Coopers) from the fighting and genocide in Rwanda, they embarked upon a study of skeletal and dental pathology of the mountain gorilla (Gorilla beringei) in the collections of the Royal College of Surgeons of England and the National Museums of Kenya (Cooper and Cooper, 2007, 2008). This led to a project involving the gross examination of material from both species of gorilla in museums and collections in Belgium, France, Germany, the Netherlands, Kenya, Rwanda, South Africa and Uganda; this was accompanied by appeals for information to colleagues who may have had access to uncatalogued skeletal material. Those studies revealed that the gorilla skeletal material from collections in Europe and Africa showed a similar range of pathological changes to those reported by Lovell (1990a) in her review of great ape skeletons in the United States.

It soon became apparent that not only skeletons but also other material could provide valuable data on the pathology and diseases of gorillas. As Part II: A Catalogue of Preserved Materials, amply illustrates, various specimens are to be found in the world’s scientific collections and many are available for investigation, provided that appropriate arrangements are made. Most of the material is from gorillas that died, or were killed, decades ago and they therefore provide valuable reference (baseline) data. Gross and microscopic studies can, increasingly, be supplemented with imaging and DNA and other molecular studies.

While some of these collections have been extensively investigated, many have not, and studies on them by medical scientists are few and far between.

On their return to Rwanda after the cessation of hostilities, the Coopers resumed their research on gorilla material there. Some specimens had been destroyed or damaged, and gorilla bodies and tissues that had been left for safe keeping at the Laboratoire Nationale de Rubilizi in Kigali prior to the outbreak of the war were not to be found.

The compilation of this book and its accompanying catalogue are essentially the result of the belief by the two authors that there is a need a) to collate, as far as possible, published and unpublished data about the pathology of Gorilla spp. and to attempt to relate this information to the health, welfare and conservation of gorillas in the wild and in captivity and b) to produce a global list of skeletal and other material resources. Our aim to produce a global list has certainly been achieved. The catalogue that comprises Part II of this book, is, we believe, the first such database of its kind and owes its existence to years of dedicated work by Gordon Hull.

Our awareness of how gorillas respond to infectious and noninfectious conditions, including injuries, has advanced over the past three decades, largely driven by proponents of high quality zoo animal medicine and by far-sighted field projects. It remains a fact, though, that in contrast to what we know about humans, domesticated species and laboratory animals, there is still a paucity of properly collated, scientifically assessed, information about the causes and pathogenesis of diseases in gorillas. The need to have access to such data has, however, never been greater. Gorillas are a threatened species and both infectious and noninfectious diseases contribute to their ill health and death. In the case of some infectious conditions, such as Ebolavirus disease (EVD), whole populations are at risk and there are profound health implications for humans.

In his book Listellany (2014), John Rentoul claimed that ‘Lists are the future of journalism, the internet and the world. Also they are the past’. Veterinary and medical pathologists love lists, however – of differential diagnoses, for example – but Part II: A Catalogue of Preserved Materials, is already a list and an additional one, detailing all the reports of pathological changes in gorillas, would not make interesting, or necessarily useful, reading. Instead, therefore, the approach in Part I, Gorilla Pathology and Health, is an applied and practically orientated one, with an emphasis on how pathology affects the individual animal, its group and its species.

At this stage our definition of the term ‘pathology’ needs to be explained. The etymology is clear from its Greek roots – pathos, suffering/disease and logos, reason/plan. In this book we use the term ‘pathology’ to mean the science of the study of disease, with particular reference to its causes, its development and its appearance. In common (lay) parlance ‘pathology’ tends to be linked with death, carcases and postmortem examination and not, as it should, to the investigation also of clinical disease and the application of laboratory tests. This book intentionally incorporates the latter as well as the former.

The authors are two British enthusiasts with a shared interest in gorillas but rather different backgrounds.

John Cooper (JEC) is a member of a husband and wife team, from the United Kingdom. He and his wife Margaret (a lawyer, who is a contributor in her own right to this book) have travelled widely and lectured together in many countries. They have spent nearly 20 years living overseas, mainly in Africa, the Middle East and the Caribbean. The Coopers worked with mountain gorillas in Rwanda from 1993 to 1995 (see below) and they continue to study these and other primates. The Coopers are now based in Britain but are also engaged in voluntary work with wildlife, domesticated animals and communities in East Africa. JEC trained as a veterinary surgeon (veterinarian), has had a lifelong interest in natural history and is now a specialist veterinary (comparative) pathologist with particular involvement in studies on wildlife and so-called ‘exotic’ species, tropical diseases and comparative medicine. His attraction to what is now termed ‘One Health’ is in part because, as a veterinary student, he was taught by medical lecturers: his books on physiology, histology and pathology were primarily concerned with Homo sapiens, not conventional domesticated animals. It was, therefore, perhaps providential that he should spend 13 years of his professional life at the Royal College of Surgeons of England (RCS) as Senior Lecturer in Comparative Pathology. The RCS had a ‘Hunterian’ approach to comparative medicine at least 250 years before the current mantra of ‘One Health’ became so popular.

In the early 1970s, at the request of the renowned palaeontologist Dr Louis S.B. Leakey, John served as honorary veterinary surgeon for the National Primate Research Centre (now the Institute of Primate Research) in Kenya. Later, from 1993 to 1995, he was the Director of the Centre Vétérinaire des Volcans/Volcano Veterinary Center (CVV/VVC) in Rwanda, responsible for the health of the mountain gorillas.

Gordon Hull (GH) is an amateur naturalist with a deep interest in primates and certain other mammals. He has specialised in the study of gorillas over many years, during which time he has amassed a great deal of technical and historical information about specimens in zoos, museums, and other institutions throughout the world. Although unaffiliated, he has been, and remains, an assiduous and effective researcher, able to elicit excellent responses and cooperation from professional scientists and lay persons alike. Gordon is a member of the Gorilla Pathology Study Group (GPSG) and was a coauthor with John Cooper and Gladys Kalema-Zikusoka of the paper ‘Diseases and Pathology of the Genus Gorilla: The Need for a Database of Material and Resources’, that was presented at an International Scientific Conference in Nairobi, Kenya, in September 2013, and which arguably set the scene for the production of this book and its accompanying catalogue.

The VVC, where the Coopers were based in the 1990s, was established in 1986 at the request of Dian Fossey, with moral and financial support from Mrs Ruth Morris Keesling, and administered by the Morris Animal Foundation (MAF). Dr James Foster established the VVC and served as its Director from 1986 to 1988 (see picture below). Several of the subsequent Directors are referred to in this book but the Coopers would like to pay particular tribute to their friend the late Barkley Hastings. Barkley enthusiastically contributed to RCS/ZSL clinicopathological seminars in his London days, then played a pivotal role in initiating research at the VVC that ranged from extensive and enterprising studies on protozoal and helminth parasites to morphometrics on dead and immobilised animals and serology, microbiology, haematology and blood biochemistry.

A picture by a local artist, Sinanenye J.M.V., on paper and cardboard. It depicts the first CVV/VVC veterinarian, James Foster, treating a mountain gorilla on Sabinyo in the Parc National des Volcans/Volcanoes National Park, Rwanda.

Following the metamorphosis of the CVV/VVC into an independent body, the Mountain Gorilla Veterinary Project (MGVP), now often termed ‘Gorilla Doctors’, it was not infrequently suggested that whereas the VVC was originally a ‘mostly clinical programme’ (Cranfield and Minnis, 2007), the MGVP had changed it into ‘a holistic conservation medicine programme’. One feature published in 2006 in an eminent American veterinary journal even stated that ‘During its early years, the project’s (VVC’s) efforts were small in scope, limited to up to four medical interventions a year and the occasional necropsy to determine how a gorilla died’. This is all very misleading. It is not only an unjustifiable criticism of MAF’s innovative and humane management of a unique project but totally overlooks the sterling work of Barkley Hastings and others who, under the most difficult of circumstances (social unrest, sometimes war, no mobile telephones or emails, no running water, often working in three languages) managed between 1986 and 1996 to carry out much significant work in such fields as parasitology and cytology. In the 6 months prior to the outbreak of the 1994 genocide, the VVC was embarking on a whole range of progressive initiatives with the aim of providing a veterinary service, including laboratory investigations, for the whole of the Parc National des Volcans and intended to assist various kinds of wildlife as well as local people.

Gorillas are only found in Africa and their long-term survival in the wild depends on those who live and work there. Throughout this book, therefore, there is an emphasis on the need to involve African scientists (veterinarians, pathologists, primatologists and biologists), as well as African institutions, in studies on the health and pathology of gorillas and in locating and conserving specimens within that continent. This approach is coupled with championing the need for tissues and other samples from gorillas to be retained within the species’ range states, where access can be readily provided to local students and researchers, rather than overseas. This is in keeping with the spirit of the 1992 Convention on Biological Diversity (CBD), which deals with the fair and equitable sharing of genetic resources (see Chapter 18, Legal considerations). The essence of that Convention is that scientists from western (richer) countries should be taking their expertise to poorer nations, not removing material from range states so that it can be ‘safely’ processed and archived in North America or Europe. In respect of specimens from mountain gorillas, the importance of this approach to African countries was emphasised at the Population and Habitat Viability Assessment held in Uganda in December 1997 (Werikhe et al., 1998) and discussed further at a C.L. Davis Foundation Symposium in 1999, also in Uganda. At the latter the attendees had been told of plans to establish a database/reference collection of mountain gorilla material in Colorado, USA (described in a brochure a year later as ‘the Biological Resource Center. Here, mountain gorilla samples can be stored in one central facility. These samples are available to scientists worldwide whose studies focus on mountain gorilla health issues’). The C.L. Davis Foundation Symposium in Uganda resolved that records and specimens concerning mountain gorillas should be duplicated and one set kept in the host country (Rwanda, Uganda and DRC), but whether this was done routinely in the succeeding years is not clear.

For some time prior to the completion of this book, reference had been made in various quarters to ‘a biobank inventory’ of gorilla material held by MGVP. Some limited information about this, made available to us courtesy of Gorilla Doctors, is to be found in Part II, A Catalogue of Preserved Materials. However, it should be said that earlier enquiries about its whereabouts and contents largely proved unfruitful. This was not only disappointing but raised the question of how modestly funded African scientists, from whose homelands all this gorilla material originated, could hope to gain access to such resources.

The vital role of local Africans in studies on gorillas was, of course, recognised long ago. Early hunters, explorers and biologists would not have succeeded in their quest to find gorillas without the help of village people who had knowledge of the animals and the terrain. This was exemplified by the life of Reuben (sometimes spelt Rueben or Ruben) Rwanzagire in Uganda: the finding of his last resting place by a group from Makerere University was recounted by Cooper and Cooper (2001a).

In more recent years Africans have played an increasingly major role in the science of conservation and healthcare of gorillas. Their task is not always easy. A letter published in Veterinary Times (Cooper and Cooper, 2004) paid tribute to African veterinarians who had worked with mountain gorillas and, mindful of history, concluded ‘They are a constant source of support to the expatriates with whom they work, not least because of their local standing and their facility with vernacular languages: however, unlike the expatriate vets, they cannot depart when the working environment becomes dangerous’.

Over the succeeding 20 years the situation has certainly improved insofar as involvement of local people is concerned; but for as long as most of the funding for gorilla projects comes from the United States and Europe, there will be a tendency for expatriates to take the lead over policy and for Africans to have to follow. Gorilla organisations stress that they are ‘international’ but in reality ‘internationalism’ means ‘existing or carried on between nations; agreed on by many nations’ (Oxford English Dictionary), not just having members of staff who come from other countries.

One initiative to involve African colleagues more fully in their own right was the formation of the Gorilla Pathology Study Group (GPSG) in 2008. The GPSG receives no specific funding but small grants are attracted for training purposes. Thus, for example, in November 2011 the GPSG organised workshops on primate pathology in both Kenya and Uganda.

Apart from their intrinsic importance, as endangered species in precarious habitats, gorillas are of considerable relevance to the health and wellbeing of humans – extending from being natural ‘models’ of cardiovascular and osteoarthritic conditions to their (regrettable) susceptibility to Ebolavirus infection. Working with live gorillas in the wild or in captivity provides opportunities to learn about the role of disease in these species and the concurrent collection, collation and publication of data helps to fill gaps in our knowledge of relevant epidemiology and pathology in both Gorilla and Homo.

The background to Part II: A Catalogue of Preserved Materials, is detailed in its Introduction. As far as Part I of the book is concerned, when the project started it was hoped that the chapters would include reference to the vast majority of published papers about pathogenesis of known diseases of gorillas, together with a comprehensive (meta) analysis of causes of death and pathological findings in gorillas from the majority of the world’s zoos that keep this species. Neither has proved possible. The number of references relating directly to gorillas, or relevant to them, is enormous and continuously growing. In addition, though, it has been disappointing to note how many scientists representing zoo collections have declined to share basic information, or even failed to respond to our polite request for collaboration. This is puzzling. Such a refusal to cooperate not only denies others, including Africans from range countries, the chance to see, perhaps to use, samples, but also dents the claim of modern zoos to be contributing to conservation, research and education – the three tenets of the European Zoos Directive.

How different this is from the kindness we have received from the majority of our correspondents, and indeed, the experiences of the British (medical) pathologist GBD Scott, who wrote in his 1992 book: ‘… Dr Kurt Benirschke, the then Director of Research at the San Diego Zoo, placed the Zoo’s records of autopsies on primates at my disposal … he gave me every encouragement …’.

Alas, attempts at collaboration over gorillas and their conservation appear often to engender competition and possessiveness rather than openness. Two veterinary colleagues who have generously contributed material for this book are worth quoting:

Problems only arise when people don’t share information.

Sharon Redrobe, UK

I personally feel no one should have ownership over gorilla-related information and you are fighting the good fight getting the information out there…

Yvonne Schulmann, USA

Another unexpected difficulty that we have encountered is that the accuracy of diagnosis is not always taken seriously by those who hold specimens. A senior curator at a prestigious London museum told one of us (JEC) that providing accurate catalogue descriptions of lesions of gorilla skulls was not essential ‘because someone else will come along later and make corrections’! Such an attitude does not augur well for the scientific documentation of material that, as the species face increasing pressure in the wild, becomes more and more important in terms of providing valuable background data. Perhaps the answer to the question of unreliable diagnoses is for museums to restrict their catalogue entries to descriptions, supplemented whenever possible with photographs, and to leave attempts at interpretation of any morphological abnormalities or distinct lesions to veterinary, medical or dental pathologists.

This book is written in British English. This not only expresses the background of the authors, but also reflects the fact that many people working with gorillas in their range states use European terminology and syntax.

The word ‘native’ (or its equivalent in French or German) was employed routinely in earlier reports when Europeans and other expatriates were referring to local African people but the term is never used in this book (including the catalogue) except when it forms part of a quotation or citation.

Abbreviations that cause confusion are excluded from this work, in particular ‘cbc’ which is often used, without explanation, in North American texts. It means ‘complete blood count’ but in many other parts of the world different terms are employed, especially ‘total blood count’.

The word ‘quite’ is avoided throughout the text, intentionally, except where it is part of a quotation. In British English ‘quite’ generally means ‘fairly; to a certain extent or degree’ whereas in American English it usually equates with ‘very; to the utmost or most absolute extent or degree’. In a scientific text such a difference in connotation could be dangerous!

A few terms need specific explanation. In this book the term ‘free-living’ is used to describe gorillas that are living in the wild. ‘Free-ranging’ is favoured by some authors, especially American, but that term is used rather differently in Britain and other parts of the world.

The international nature of studies on endangered species means that the metric system should be used. This is the case in all gorilla range states. Standard international units (Système International d’Unités, SI) were introduced in 1960 as the result of an initiative that started in 1948. Animals should be weighed in grams (or kilograms) and measured in millimetres, centimetres or metres. However, the reader should be aware that in the United States and a few other countries, ‘conventional’ units are still routinely employed, and be prepared to request conversion or to do so oneself. An excellent explanation of the SI system was given by Douglas (1977), whose short paper concluded with the following words ‘Data from one laboratory can only be compared with those from another laboratory if identical methods are used’. That statement, and the clear description of the system by Douglas, are as true today as they were nearly 40 years ago.

The need also to use international dates was well illustrated in Rwanda where a British veterinarian’s field notes (using the European system) were put on to a computer by an American colleague using North American (USA) dates; subsequent analysis of them by a third person resulted in considerable confusion because one of the days cited (3/6/94, translated into 3rd June 1994) was when the Rwandan civil war was in full force and there were no veterinarians in the field!

In this book reference is made to both recent and previous (older) literature. There is an increasing tendency for journals and editors to insist on the inclusion in manuscripts only of the former, suggesting that this is more relevant and reliable. Such is a mistake. In a book such as this it is important that the references quoted reflect the history and development of the subject, not just ‘new’, ‘cutting edge’ publications. An appreciation of how our understanding of gorilla pathology and health has evolved helps put current work in context and elucidates where there are still significant gaps in our knowledge. Our friend Andy Richford, who guided us in the formulation of this book, advised us against including too much ‘grey literature’: in other words, material that is anecdotal or based upon an individual’s observations and deduction, rather than properly executed scientific study (see elsewhere regarding ‘hypothesis-based’ research). This was sound advice but we have not been able to follow it in toto. Many significant reports on gorillas and their health, especially in the earlier days, were the result of meticulous observation and record keeping by trackers of free-living gorillas or keepers of those in captivity. More recently too, there has been an abundance of relevant published case reports and observations in this book in various journals and magazines. These should not be overlooked.

Part I of this book is intended to be a monograph on the pathology (in its broadest sense) of gorillas and relevant implications for the health of these animals, both in the wild and in captivity. It is therefore particularly, but not exclusively, aimed at primatologists, veterinarians, biologists, osteologists and conservationists – especially African colleagues with very limited resources.

In order to save space, the full English names of the different species and subspecies of gorilla are not used in the following text. The two species are referred to broadly as ‘lowland’ (or ‘gorilla’) and ‘mountain’ (or ‘beringei’) except where it is necessary to distinguish them more specifically. The scientific names of all species mentioned in the text are listed in Appendix 6: Scientific Names of Species.

In photomicrographs, the stain used was haematoxylin and eosin (H&E), generally Cole’s, unless otherwise indicated. Except where specifically stated, the magnification of photomicrographs is either ×100 (low power) or 400 (high power).

There is an ancient proverb that states: I hear ~~ I forget, I see ~~ I remember, I do ~~ I understand. In this book, therefore, the reader will find not just information about the pathogenesis of disease in the genus Gorilla but also practical guidance, coupled with appropriate references, as to how to investigate animals, how to select samples and how to examine them. In other words, how to do it.

We have also, intentionally, included in the book some vignettes – brief, hopefully evocative, descriptions of incidents, mainly based on the Coopers’ experiences in Rwanda, that help illustrate the pleasures and the challenges of work with wildlife in Africa.

To conclude, the gorilla has attracted a great deal of scientific interest since its discovery and recognition as a separate genus, Gorilla, in the mid-19th century – less than 200 years ago. Nevertheless, it took nearly a century for an authoritative textbook to be published detailing the anatomy of gorillas. Notwithstanding the appearance in recent years of a substantial number of papers on infectious diseases of the genus, remarkably little is yet known about the gross and microscopic pathology of gorillas and how they respond to infectious and noninfectious insults. Our joint publication is intended to play a part in rectifying the situation. The centenary volume of The Pathological Society of Great Britain and Ireland in 2006 had as its title ‘Understanding Disease’ – that is the main aim of this work.

April 2016

Acknowledgements

John E. Cooper and Gordon Hull

Nia zikiwa moja, kilicho mbali huja

(Swahili: When minds are one, what is far comes near)

The successful compilation of this book would not have been possible without the help, advice and support of many people. Panegyrics, lofty writings in praise of a person, are not popular nowadays but it is important that those who have helped us in the immediate or distant past are acknowledged.

First, we both (JEC and GH) thank Louise Leakey and Emmanuel de Merode for writing the Foreword. They are a husband and wife team who have not only contributed much to Africa, which is their home, but also have many links with gorillas and their conservation. Louise’s grandfather, Dr LSB (Louis) Leakey is extolled in Chapter 1: The genus Gorilla – Morphology, Anatomy and the Path to Pathology. In addition to his contributions to palaeontology (and those of his wife, Mary, and other members of the family), Louis Leakey recruited Dian Fossey to study mountain gorillas in Central Africa.

When the Coopers first lived in Kenya, from 1969 to 1973, JEC served as (honorary) veterinary surgeon to Louis’s Primate Centre at Tigoni. Following Louis’s death in 1972 JEC and Peter Holt (thanked elsewhere in these pages) continued to tend the monkeys and to support the Centre until its future was assured. Ultimately it metamorphosed into the National Museums of Kenya’s Institute of Primate Research (IPR) (see Chapter 2: The Growth of Studies on Primate Pathology).

It is an honour to use Figs. 1.4 and 13.5, together with the front cover picture, that were taken in Rwanda by the late Robert (Bob) Campbell and were kindly provided by Heather Campbell, his widow and a friend and former colleague from the Coopers’ days in Kenya. Other images are supplied by Margaret E. Cooper or John E. Cooper except for those specifically credited beneath the figure or attributed below:

• Cambridge Veterinary School/Kate Hughes, Fernando Constantino-Casas, Michael Day (Figs. 2.5, 9.1, 9.2, 9.4, 9.6, 10.1, 10.2, 10.3, 10.4, 11.5, 13.1, 14.5, 14.6, 15.3, 16.3)

• Paddy Mannion (Figs. 6.2, 11.2 and 14.1)

• David Perpiñán (Figs. 10.7–10.9)

• Sam Young (Fig. 15.4)

• Carina Phillips (Figs. A.5.14–5.16)

• Michelle Burrows (Fig. A.5.17).

These images have greatly enhanced the book and complemented our own.

We express our sincere appreciation to Melissa Read (freelance Project Manager assisting the Elsevier Production team in Oxford) who, during the final stages of completing this book, has dealt with two idiosyncratic authors and their queries with much calmness, kindness and humour. There are many who, throughout the gestation of this publication, have given us wise counsel. We particularly thank Colin Groves for his interest in – and support for – our project and for sending us interesting images. Despite his many commitments he has always responded promptly to our email messages and furnished us with sound advice, especially but not exclusively about taxonomy. In his own inimitable way, early in this venture, he suggested ‘For the benefit of those who have been asleep over the past 30 years, you might explain that the family Hominidae used to be restricted to humans only, but now includes great apes’. In the same vein we are grateful to David Chivers and Roger Short for their advice and support.

We thank our contributors: Margaret Cooper, Ian Redmond, Jane Hopper, Gladys Kalema-Zikusoka, Jenny Jaffe, Ogeto Mwebi, and Celsus Sente; the ‘teams’ who worked with us on (1) ‘Guy’ at the NHM, Brian Livingstone, Paolo Viscardi, Keith Maybury, Allen Goodship, Richard Sabin and Roberto Portela Miguez, (2) skeletons in the Osman Hill Collection at the RCS, Carina Phillips, Martyn Cooke and Paul Budgen and (3) the student projects at Bristol Zoo Gardens, Sophia Keen, Dermot McInerney, Michelle Barrows, Christoph Schwitzer, Bryan Carroll, Kate Robson-Brown, Jonathan Musgrave and Allen Goodship. Mick Carman and Don Cousins also provided helpful information relating to our study of ‘Guy’.

Sally Dowsett, family friend and long-term supporter of our projects in Africa, typed drafts and for some years served as honorary secretary to the Gorilla Pathology Study Group. Pauline Muirhead kindly gave similar assistance while Gemma Saunders and Jenny Cocking cheerfully prepared laminates.

Many people kindly agreed to read (review) parts of this book before publication and/or provided helpful comments on content and orthography. We thank (arranged alphabetically) David Alderton, Ekane Humphrey Anoah (Wildlife Law Expert of the Last Great Ape Organisation, Cameroon), Helen Chatterjee, Vanessa Cooper, Max Cooper, Sarah Cooper, Chris Daborn, Jeremy Dearling, Ellie Devenish Nelson, Scott Dillon, Ofir Drori, Keith Hardy, Mamun Jeneby, Peter Kertesz, Arthur Kemoli, Brian Livingstone, Keith Maybury, Jaimie Morris, Jonathan Musgrave, Emily Neep, Howard Nelson, David Ojigo, Caroline Pond, Ian Redmond, Sharon Redrobe, Jenny Rees Davies, Rob Shave, Nick Short, Victoria Strong, Simon King, and David Williams.

Many others who kindly helped are cited as a ‘personal communication’ in the text or acknowledged in the catalogue. In addition, literature, contacts and/or helpful suggestions were received at various times from Wilbur Amand, Frances Barr, Donald M. Broom, Debra C. Bourne, Tom Butynski, Héctor Sanz Cabañes, Sarah Chapman, Christopher P. Conlon, Chris Furley, Hayley Murphy, Rick Murphy, Navarro Serra, Norma Chapman, Andrew Dixson, James Hassell, the late Barkley Hastings, Richard Jakob-Hoff, Jesús Maria Pérez Jiménez, Ian Keymer, Sascha Knauf, Felix Lankester, Michael Marks, Shelly Masi, Nelly Ménard, Dick Montali, Torsten Morner, Stuart Nixon, John Bosco Nizeyi (JBN), Richard J. de Norman, Marc Nussbaumer, Mark Rose, Bruce Rothschild, Peter W Scott, Angelique Todd, Anna and Steve Tolan, Lydia Tong, Tonino Van Wonterghem, Roberta Wallace, David Warrell, Michael Woodford and Sam Young.

Copies of postmortem reports were made available to us by many kind colleagues, including Rachael Liebmann and other medical pathologists.

Colleagues in the Gorilla Pathology Study Group (GPSG) have not only contributed to this book but also collaborated enthusiastically in relevant primate workshops in Africa. We are indebted to the late Professor Phillip V. Tobias, the eminent South African palaeontologist, GPSG Patron, for his encouragement, and we wish to record his role, for nearly half a century, in promoting the involvement of Africans in studies on the mountain gorilla and in helping to facilitate access to gorilla skulls at ‘Wits’.

Dr Sam Thompson, the lifeblood of the C. L. Davis Foundation in the United States, died in 2014. JEC remembers well his kindness and support, especially in respect of training programmes for veterinary pathologists in East Africa and the opportunity for JEC to record lectures for teaching purposes.

JEC is grateful to former Volcano Veterinary Center (Centre Vétérinaire des Volcans) (VVC)/Mountain Gorilla Veterinary Project (MGVP) veterinarians who responded personally to his letters, in particular Jonathan Sleeman, Antoine Mudakikwa and Ken Cameron. Officially sanctioned, summarised, information about MGVP findings was kindly furnished by Linda Lowenstine and she and Karen A. Terio provided the most recent versions of their general ape necropsy form, the GAHP recommended cardiac necropsy (prosection) guide and the guide for pathologists.

More than 1000 people have contributed to Part II: A Catalogue of Preserved Materials. We would like to thank them collectively here, since it has not been possible to mention everyone by name. Museum curators and collection managers, zoo vets, university academics, secretaries and others have selflessly dealt with many time-consuming enquiries (particularly from GH), despite already being under ample pressure from their everyday duties. Without their cheerful collaboration, this book would never have come about. In addition, GH would like to thank Spartaco Gippoliti for his help in contacting Italian collections, and Colin Groves, Rebecca Jabbour, Andrew Kitchener, Peter Holt, James L. Newman and Esteban Sarmiento for kindly reviewing the text for the Introduction to Part II. Paolo Viscardi provided some useful text (Tyranny of the test tube) for the Introduction. GH would also like to thank the experts in linguistics and the classics who have contributed in no small measure to the Introduction: their enthusiastic willingness to help with a project lying outside their normal sphere of activity is both exemplary and laudable.

We are grateful to Rob Shave for information about the International Primate Heart Project. The Coopers are indebted to Terry B. Kensler, Collection Manager, Laboratory of Primate Morphology, Caribbean Primate Research Center, Puerto Rico, for her welcome and for sharing information about skeletal methods so freely.

Both John and Margaret Cooper thank Professor Richard Griffiths for permission to use the DICE (Durrell Institute of Conservation and Ecology, University of Kent) address for this publication. Since DICE’s inception, it has trained hundreds of postgraduate students, from over 80 countries, and the Coopers are proud to have been associated with it, as Honorary Research Fellows, for many years.

JEC owes a great deal to his many mentors, both as a schoolboy and in later years. He particularly appreciates the training and guidance he received in respect of:

• natural history from Major Maxwell Knight (former M15 agent, the prototype for ‘M’ in the James Bond books), Mr Gerald Durrell (animal collector extraordinaire and Founder of the Jersey Zoo/Durrell Wildlife Conservation Trust/‘Durrell’ in the (British) Channel Islands), Mr Henry Berman (teacher extraordinaire). These and others with a love of the countryside enabled him to apply lessons about the natural world learned in the heaths and woodlands of England to wild places in Africa and elsewhere.

• osteology and its links with primatology from Dr Louis S.B. Leakey and Professor Phillip Tobias.

• odontology methods from Professor Loma Miles, Professor David Poswillo and Dr Caroline Grigson at the RCS and Professor Arthur Kemoli in Nairobi.

• comparative pathology (creatures great and small) from Dr Edward Elkan, Professor Peer Zwart, Professor Norman Ashton, Professor Gerry Slavin, Drs Ariela Pomerance, Mike Bennett, Ashley Price, Rodney Finlayson and R.N. T-W. Fiennes.

JEC also acknowledges the help of former colleagues in respect of primates and other matters at the University of the West Indies (UWI) – Paluri Murti, Rahul Naidu, Haytham Al-Bayaty, Lee Koma, Ravi Seebaransingh, Rod Suepaul and Richard Spence.

JEC owes a particular debt of gratitude to his teachers over the years, especially those who taught him medical and veterinary pathology at Bristol University and tropical veterinary pathology at Edinburgh University.

From 1978 to 1991, when JEC was Senior Lecturer in Comparative Pathology at the RCS, working with, and teaching, both medical and dental graduates, he was able to practise ‘One Health’ on a daily basis. He also received much support from RCS staff members Nina Wedderburn, Martyn Cooke, Liz Allen, John Turk, Ian Lyle, Derek Manning, Ashley Miles, Bari Logan, Ken Applebee, Alan Graham and Brian Eaton and Professor AJE Cave (who was publishing material on gorillas before JEC was born). He received encouragement from former Presidents such as Sir Alan Parks and Sir Reginald Murley. The many veterinary students who ‘saw practice’ at the RCS helped JEC with numerous relevant projects.

During his time as Guest Professor in Berlin, at the Institut für Zoo und Wildtierforschung/Institute for Zoo Biology and Wildlife (IZW), JEC and his wife received much support from the Director, Professor Reinhold R. Hofmann, an old friend from Kenya, and a warm welcome from colleagues there. Amongst those who contributed notably to studies on gorillas was Dr Roland Frey who performed a skilled dissection of a gorilla’s hand that was transported by car, with all necessary permits, across the length of Western Europe!

Over the years very many people have helped nurture the Coopers’ studies on skeletal lesions in gorillas.

In Rwanda, before the outbreak of genocide, they included staff of the VVC, notably Innocent Kiragi. Afterwards, following the Coopers’ evacuation to England in 1994 prior to returning to the VVC, Elizabeth Allen, John Turk and Martyn Cooke at the RCS provided facilities and support. The Coopers’ strong links with Jersey Zoo/Durrell Wildlife Conservation Trust/‘Durrell’, ranging from helping run their Summer Schools to JEC’s period as consultant veterinary pathologist to the Trust, greatly helped in this respect too. The support at Jersey Zoo of Gerald and Lee Durrell, Jeremy Mallinson, John Hartley, Nick Le Q. Blampied, Tony Allchurch, Richard Johnstone-Scott and other Trust members was much appreciated.

Following their departure from the VVC in Rwanda in 1995, the Coopers extended their skeletal research by visiting numerous museums and collections in different parts of the world. Those institutions where they worked — and the people who offered them a warm welcome — are too many to mention individually but are amongst the establishments listed in Part II: A Catalogue of Preserved Materials. The Coopers are grateful to staff of the Medical School of Witwatersrand, South Africa, for their support and assistance with mountain gorilla specimens, especially Professor B Kramer who gave permission for the examination of the material and Peter Dawson, Mary-Ann Costello, Llewellyn Sinclair and Peter Montja for the preparation and radiographing.

Most recently we have received help and been given access to material (see also Part II: A Catalogue of Preserved Materials): at the Muséum National d’Histoire Naturelle, Paris (Jacques Cuisin); Booth Museum (John A. Cooper); Royal Zoological Society of Scotland (RZSS) (Simon Girling, Lynda Burrill, Roslin Talbot and Jo Elliott); the Zoological Society of London (Edmund Flach and Tom Kearns); Bristol Zoo Gardens (Michelle Barrows, Allen Goodship, Rowena Killick, Lynsey Bugg, Sarah Gedman, Ryan Walker and Sam Matthews). Moses Cooper, probably the youngest contemporaneous gorilla researcher (he was born in 2006), has for 2 years ably assisted his grandfather in skeletal studies.

Over the years colleagues at Cambridge University Department of Veterinary Medicine have been generous with their time and facilities. These include Michael Herrtage, Fernando Constantino-Casas, Kate Hughes, Paddy Mannion, Chrissie Willers, Anna Dussek, Lucy Webb, Vicky O’Mahony, Mathew Rhodes, Madeline Fordham, Rayna Skoyles, Scott Dillon, Louise Grimson and Paul Tonks. Three colleagues at Cambridge have helped with translation of papers – Emilie Cloup (merci!), Heidi Radke and Angelika Rupp (danke!) and Carolina Arenas (gracias!).

The cordial welcome and generosity provided by Michael Day at the School of Veterinary Sciences, Langford, University of Bristol, has been particularly appreciated.

The authors are grateful to the librarians at the RCVS (Clare Boulton), Linnean Society (Lynda Brooks and Gina Douglas), National Museums of Kenya (Asha Owano) and Institute of Primate Research (Grace Mathani) for their help and use of their reading and literature-searching facilities.

By contrast, the keepers of some collections and databases either failed to reply to our polite requests or declined to help. Those attitudes only serve to accentuate the kind, generous, responses of many others. Two examples illustrate this:

Always unrestricted access to the bones

Ogeto Mwebi, National Museums of Kenya

And, last but not least, please remember that the doors of the collections stay open for further investigations!

Jacques Cuisin, Muséum National d’Histoire Naturelle.

Such is the spirit of true scientific collaboration.

Having read widely in order to compile this book, we realise more than ever the debt owed to earlier pioneers, most of whom we never met but who contributed to a greater understanding of gorillas. For instance, in giving the Coopers a copy of her 1991 book, Penelope Bodrey-Sanders opened our eyes even wider in respect of the achievements of Carl Akeley. As she put it: Nearly three quarters of a century ago Akeley became a pioneer in African conservation through his art and love of gorillas. Stimulated directly or indirectly by him, many others have over the years accepted the challenge and moral obligation to fight on behalf of Africa’s wildlife. The mountain gorilla is Akeley’s living monument, and those that follow his lead remain his most enduring legacy. There are other, often forgotten, pioneers – for example, Kenneth Carr, hunter and explorer, whose papers and photographs were passed to us by Edward Le Conte, and Jean-Marie Eugène Derscheid the Belgian zoologist who became the initial director of Africa’s first national park. During the Second World War Derscheid served in the Belgian Resistance and helped Allied soldiers and airmen escape from occupied Europe. He was shot as a spy by the Nazis on 13 March 1944, 5 days after JEC was born.

While the origins of the catalogue, go back to GH’s absorbing interest in gorillas that began in the 1960s, the book has a much more recent origin – the Coopers’ time (1993–95) with the Morris Animal Foundation’s (MAF’s) Volcano Veterinary Center (Centre Vétérinaire des Volcans) in Rwanda. More is said about this in the Preface, but this is an opportunity to pay tribute to many affected by the outbreak of violence (the ‘Rwandan genocide’) in April 1994 and its aftermath. Some Rwandans who helped the VVC or collected samples for the project were killed. Others survived but with terrible legacies. Etched in the Coopers’ memories forever are the words of one tracker who, in faltering English, wrote: Do not forget we. They will, indeed, not forget him nor many, many others.

The Coopers particularly remember those who did so much to keep them safe and to help them escape from the fighting and from Rwanda. The United Nations troops, so maligned by many and so small in numbers, must never be forgotten. Those from Ghana and Bangladesh who served in Kinigi were the Coopers’ friends, not just their peacekeepers. Some were never to return to their home countries. Three other blue-hatted UN soldiers, amidst the carnage going on around them, with only a pistol apiece, helped 169 evacuees, including the Coopers, to cross the border to relative safety in Zaire (now DRC). Those who criticise the United Nations usually do so from their comfortable office and forget Dag Hammarskjöld’s words: The UN was not created to take mankind to heaven, but to save humanity from hell.

In the context of those days, John and Margaret Cooper particularly wish to acknowledge the support of the staff of the Centre Vétérinaire des Volcans before, during and following the genocide in 1994 and to recognise in these pages the friendship and courage of JEC’s Rwandan counterpart. Other thanks and poignant memories, can only be summarised here: the safe haven provided (when they had crossed into Zaire) by Alyette de Munck; support and friendship from Nicole Merlo; the Belgian government plane to Burundi; the flight with the US Marines to Kenya (Coopers’ nyumba ya pili – second home); the Institute of Primate Research who provided a secure temporary home for them and JEC’s counterpart; the good-humoured comradeship of Garry Richardson with whom JEC returned to Rwanda very