The chimpanzee is one of our planet's best-loved and most instantly recognisable animals. Splitting from the human lineage between four and six million years ago, it is (along with its cousin, the bonobo) our closest living relative, sharing around 94% of our DNA. First encountered by Westerners in the seventeenth century, virtually nothing was known about chimpanzees in their natural environment until 1960, when Jane Goodall travelled to Gombe to live and work with them. Accessibly written, yet fully referenced and uncompromising in its accuracy and comprehensiveness, this book encapsulates everything we currently know about chimpanzees: from their discovery and why we study them, to their anatomy, physiology, genetics and culture. The text is beautifully illustrated and infused with examples and anecdotes drawn from the author's thirty years of primate observation, making this a perfect resource for students of biological anthropology and primatology as well as non-specialists interested in chimpanzees.

The history of chimpanzee scholarship has been one of generally gradual but occasionally sudden shifts in the perception of how closely related humans and chimpanzees are. Here I will try to bend that perspective a little bit to view chimpanzees more as a counterpoint to human uniqueness, taking care to point out how they differ from humans and how explaining those differences helps us to understand human evolution, including our frailties and failings. To draw out those insights, I will attempt an unhurried grand tour through the natural sciences, always with chimpanzees in the forefront, always contrasting their biology and behavior to that of humans. I intend this volume to be current and thoroughly grounded in the latest research, only one step from formal scholarship, but at the same time stripped of the jargon-heavy phraseology that weighs down many similar efforts. Where I use obscure terms, I will attempt to define them on the fly.
I see my audience as scholars from other fields, laypersons with a sincere interest in the behavior, biology, and history of primates, doctoral students in human evolution, human biology, and primatology, and colleagues in scholarly specialties neighboring my own. If successful, it will be a volume worth dipping into for scholars in any natural science field, even the chimpanzee expert. I hope a primatologist might turn to this volume if he or she wants to dabble in functional anatomy or endocrinology. While I realize this is probably too much to hope for, I sought to make the book engaging enough that a colleague or student might be drawn into exploring an area they had not anticipated investigating. Ideally, this volume would be an accessible version of the six-volume G.H. Bourne volume The Chimpanzee, published from 1969 to 1973.
As I worked, I found that sometimes the most thorough investigation of a subject – for example, balance, hearing, ape cognition, aspects of gross anatomy – is half a century old. In the course of discovering the chimpanzee, researchers sometimes got it right the very first time, in which case I cited the classic scholarship that shaped a particular issue, occasionally skipping a recent publication that added a minor flourish to the subject. I would like this volume to be something of the corrective to the tendency I have seen recently among younger scholars to cite the most recent publication on some subject, no matter how tangential, ignoring truly foundational work with which they ought to be familiar. Other times experts long labored under misapprehensions that have been corrected only recently, in which case I focused on the recent scholarship with only a nod to ancient errors. My shorthand way of summarizing my emphasis on studying micro- and macrobiology, as well as behavior, is that while the bricks are indeed interesting, they are most interesting when they help us understand the castle.
I began this project in a small way in 1996, when I had just begun to teach a class at Indiana that – in a modest way – surveyed the breadth of chimpanzee research. I called it Sister Species: Lessons from the Chimpanzee. The text for the class was a coursepack made up of review articles and a few specialist research articles. The readings that were most successful with students, I learned from student evaluations, were often those from Evolutionary Anthropology. These were professional-grade offerings, current and cite-able, but still accessible to committed neophytes. I have tried to work on that model in this book.
The first few times I taught Sister Species, the first reading in my coursepack was a brilliantly written news article by Claire Martin (1994), a short work that focused on chimpanzee and human cognitive similarities. Students loved it. As evocative as this work was, I was left feeling that the reading for my first lecture should have a slightly broader perspective and more robust scholarly citations. That same year, 1996, I started research on chimpanzees at the ToroSemliki Wildlife Reserve, Uganda. One day, problems with my vehicle left me stuck in a chimpanzee-less part of the reserve for two hot, humid days with nothing to do but sweat. During this time, I wrote a short exposition that quoted extensively from Martin's article but went on to cover a broader constellation of ideas that I hoped would best set the stage for my first lecture. It would have done its job well except that almost as soon as I finished it the intense heat fried my hard drive. I lost everything but a couple of paragraphs I had copied onto a floppy disk.
Anyone who has lost a piece of work like this knows how disheartening it can be, and how the

Foreword page ix

Richard Wrangham

Preface xi

Acknowledgments xv

1 Sister's Keeper: Humans and Chimpanzees 1

2 Wild Lesson: Why Study Animals in Nature? 9

3 A Most Surprising Creature: The Discovery of the Chimpanzee 20

4 Kin: The Chimpanzee's Place in Nature 40

5 Scratching Out a Living in an Unforgiving World: Habitat and Diet 61

6 Guts, Glorious Guts, Large Stomach, and Colon: Plant Chemistry, Fruit Ripening, Digestive Physiology, and Gut Anatomy 80

7 Thews, Sinews, and Bone: Chimpanzee Anatomy and Osteology 97

8 Arboreal Gathering, Terrestrial Traveling: Locomotion and Posture 119

9 Forged in Nature's Cauldron: Engineering the Chimpanzee 131

10 Up from the Protoape: The Evolution of the Chimpanzee 158

11 Building a Natural Wonder: Growth, Development, and Life History 188

12 The Source of Similarity: Chimpanzee Genetics 217

13 Making Your Way in the Great Wild World: Chimpanzee Senses 242

14 The Grim Reaper in the Forest Primeval: Wild Chimpanzee Diseases and Lessons for Healthy Living 252

15 Powering Life: Endocrinology and Physiology 274

16 Shelter from the Storm: Chimpanzee Mothering 298

17 Meat-Seeking Missiles: Chimpanzees as Hunters 312

18 The Mind of the Chimpanzee: Reasoning, Memory, and Emotion 323

19 The Brain of the Chimpanzee: The Mind's Motor 357

20 Tired Nature's Sweet Restorer: Chimpanzee Sleep 372

21 Chimpanzee Thought Transfer: Communication and Language 383

22 Ape Implements: Making and Using Tools 404

23 Wisdom of the Ages: Chimpanzee Culture 419

24 The Daily Grind: Within-Group Aggression 427

25 A Nation at War with Itself: Defending a Community of the Mind 437

26 The Sporting Chimpanzee: Dominance without Destruction 449

27 The Passion of Pan: Sex and Reproduction 458

28 Into the Light: Semliki Chimpanzees 473

29 The Other Sister, Bonobos: The Monkey Convergence Hypothesis 499

30 Sister Species: Lessons from the Chimpanzee 517

Appendix 1: Taxonomy of the Primates 536

Appendix 2: Professional Grade Chimpanzee Testable Hypotheses 553

Index 559

Color plates can be found between pages 268 and 269.

Kevin D. Hunt is Professor of Animal Behaviour and Anthropology in the Department of Anthropology at Indiana University, Bloomington. He is also Founder and Director of the Semliki Chimpanzee Project, which was established in 1996 to study and preserve the chimpanzees within the Toro-Semliki Wildlife Reserve. Broadly trained in various anthropological disciplines, much of Professor Hunt's published work has centred on functional morphology and what chimpanzee locomotion, posture and ecology can tell us about what led humans to diverge from apes, and what advantage bipedalism gave our chimpanzee-like ancestors roughly five million years ago.

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