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Chimpanzee Behavior in the Wild Toshisada Nishida Koichiro Zamma Takahisa Matsusaka Agumi Inaba William C. McGrew Chimpanzee Behavior in the Wild An Audio-Visual Encyclopedia Toshisada Nishida Executive Director Japan Monkey Centre 2 Kanrin, Inuyama Aichi 484-0081, Japan [email protected] Takahisa Matsusaka Assistant Professor Faculty of Health and Well-being Kansai University 1-11-1 Kaorigaoka-cho Sakai-ku, Sakai Osaka 590-8515, Japan [email protected] Koichiro Zamma Researcher Great Ape Research Institute Hayashibara Biochemical Laboratories, Inc. 952-2 Nu, Tamano Okayama 70 -031, Japan [email protected] Agumi Inaba Researcher Japan Monkey Centre 2 Kanrin, Inuyama Aichi 484-0081, Japan [email protected] William C. McGrew Professor Department of Biological Anthropology University of Cambridge Fitzwilliam Street Cambridge CB2 1QH, U.K. [email protected] ISBN 978-4-431-53894-3 e-ISBN 978-4-431-53895-0 DOI 10.1007/978-4-431-53895-0 Springer Tokyo Dordrecht Heidelberg London New York Library of Congress Control Number: 2010932131 Springer

2010 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Cover Spine: An adult male, Alofu, having descended to the ground from his night bed, yawns in the early morning. Front cover, from the top: An adult female, Xtina, eating a lemon. An adult male, Alofu, having descended to the ground

from his night bed, yawns in the early morning. The Sinsiba swamp, frequently visited by M-group chimpanzees. Back cover: A juvenile female, Xantip, watches her mother, Xtina, removing a sand flea from her toe. DVDs All photos on the DVD disks and the indexes in the DVDs were taken at Mahale by Koichiro Zamma. Printed on acid-free paper Springer is part of Springer Science+Business Media Preface Where We Stand Field workersscientists of animal behavior in naturehave long been fascinated by wild chimpanzees. A person who once has studied wild chimpanzees will be eager to observe them again. A person who has studied them twice will make every effort to continue the study, unless prevented from doing so. In short, behavioral primatology is addictive! Many people, among them Jane

Goodall, Richard Wrangham, and I, do not regret that they have dedicated their whole lives to the study of wild chimpanzees. This is because the apes behavior is always challenging: chimpanzees are cheerful, charming, playful, curious, beautiful, easygoing, generous, tolerant, and trustworthy most of the time, but also are cautious, cunning, ugly, violent, ferocious, bloodthirsty, greedy, and disloyal at other times. We human beings share both the light and dark sides with our closest living relatives. For decades, we have documented huge across-population variation in behavior, as well as within-population variation. Cultural biology, as proposed 0 years ago by Kinji Imanishi, recently has flourished. However, until now there has been no extensive glossary with illustrations, upon which systematic behavioral comparison could be based. In addition to fellow field

workers, we hope that zoo keepers, laboratory researchers and technicians, veterinarians, conservationists, eco-tourism managers, educators and students now will be able to widen and deepen the scope of their activities by making use of the descriptions and audio-visual images in this volume. We hope that laypersons also will find this an enjoyable read at home on the sofa! The team of authors began to use videography in 1999 and launched this editing project in 2003, and we have invested most of our time in it over the last 3 years. After much time and effort spent, the result is perhaps the first attempt to compile a comprehensive ethogram of a single species of mammal. This is a cooperative product in the truest sense of the word. If any

of us had been absent, then we would Preface not have been able to complete this work. Roles were allocated to film, copy, playback, select, and edit, and to edit, interpret, name, describe, reference, and synthesize. The resulting collaborative efforts have been discussed and refined 20 times over in the process. We hope that this book and its accompanying DVD will help to conserve these irreplaceable creatures forever. Toshisada Nishida Contents Color Plates. 1 Introduction. 7 Methods. 11 Study Sites of Chimpanzees and Bonobos. 13 Catalogue. 15 Remarks. 25 Glossary. 27 Discussion. 209 References. 213 Appendix. 229 Color Plates 1. A juvenile female, Xantip, watches her mother, Xtina, removing a sand flea from her toe. 2. An adult male, Primus, ingests a mature leaf while chewing meat. 3.

An adult female and two juvenile females licking a rock. 4. Grooming party of six adult males. 5. An adult female, Gwekulo, resting in the bush with her adopted juvenile daughter, Puffy. A juvenile female, Mitsue, looking at her own image in a stream. 7. An adult male and two adult females grooming on a fallen log. 8. A newly-immigrated female, Ua, who did not descend to the ground when human observers were present. 9. An adolescent male, Caesar, self-grooming. 10. A juvenile female, Mitsue. T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 1, Springer 2010 1 Introduction This book is an audio-visual ethogram that lists, describes, and defines, with video clips, all the known behavioral patterns of wild chimpanzees. As far

as we know, this is the first attempt to do this for any mammalian species on earth. The behavioral patterns listed have been recorded mostly for the M-group chimpanzees of the Mahale Mountains National Park, Tanzania, where we have been doing research since 19 5. However, we have made a great effort to add patterns described from other study-sites known to us only through articles, books and videos. This book is based on a previous ethogram that had neither video clips nor illustrations, but the glossary has been much expanded. Since 1999, videographic recording often has been used at Mahale for research on various topics, particularly social interactions, although earlier cinematographic records were occasionally made from 1971 onwards. We have compiled video clips for almost all of the

behavioral patterns listed. From 1999 to 2009, new long-term study sites have been established, and many articles and books dealing with the behavioral patterns of chimpanzees have appeared. Research on gestures in captivity has also been published. With this new material, it is now easier to recognize possible differences across sites than was the case in 1999. It is well-known that chimpanzees show much local, as well as age, sex, and idiosyncratic, variation in behavior. What implications does the compilation of the audio-visual ethogram have for these issues? Our books purpose is to stimulate and to facilitate ape behavioral studies by addressing the questions below. In particular, this publication pays attention to some behavioral variations that have been neglected so far, and we hope that this will encourage

more detailed comparative and developmental studies. First, we address the question of what kind of behavioral patterns may have been present in the last common ancestor of Pan and Homo. This is a crucial question if we wish to reconstruct the behavior and ecology of early hominids such as the australopithecines, because they may have had similar patterns. T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 2, Springer 2010 7 8 Introduction Second, it is often said that among animals only human beings depend much more on learning than on instinct. However, the extent to which chimpanzees depend on learning, especially social learning, is not well understood. A simple behavioral pattern such as social scratch not only is absent at some sites

but also way of scratching varies from site to site in its details. Audio-visual representations help to persuade skeptics of the richness of this variation. Third, how do chimpanzees acquire behavioral patterns as they mature? In relation to the second question above, we study the development of behavioral patterns. Some behavioral patterns are learned from the mother, but others from playmates, as with humans. Only recently has it been shown that social customs such as the grooming-hand-clasp develop more slowly than do ant-fishing techniques. Audio-visual data make it easier to compare changes in the behavioral patterns of individuals as they mature. Fourth, we study individuality and idiosyncracy in behavior. Little attention has been paid to this aspect of behavior, with a few notable exceptions. Idiosyncratic behavioral patterns are

important because they are the seeds or origins of local culture. Fifth, in which domains are local behavioral differences most remarkable? This should be addressed if we want to infer the evolutionary origins of human culture. The extent of local differences has not been well-investigated, except for tool use, partly because most study-sites of chimpanzees lack a detailed ethogram. As new research sites have been opened, new behavioral patterns have been reported one after another, such as digging in the ground with implements or penetrating it with perforating tools. In order to know the most variable domains, however, we need for the full range of behavioral patterns to be illuminated, despite the continuing attractiveness of tool use. It may be that social behavior shows more local and individual

differences than does tool using. Sixth, heretofore unknown sex differences in behavioral patterns may emerge as more detailed observations are made. For example, through video-recording, we have noticed that immature females tend to play more in trees than do their male counterparts, and when females play on the ground, they tend to perform more somersaults than do males. Seventh, this book also may contribute to the further testing of the gestural origin hypothesis of human language origins. Gestures appear to be integral to spoken language, because they accompany the speech of people who are blind from birth. Observations of spontaneous behavior in captivity show that chimpanzees and bonobos use brachio-manual gestures more flexibly across contexts than they do facial expressions and vocalizations. Gestures seem less closely tied to

particular emotions, hence they possess more adaptable functional potential. Pollick and de Waal advocate the gestural origins of language Introduction 9 hypothesis because gestures are evolutionarily younger, and thus likely to be under greater cortical control than are facial or vocal signals. Few studies of hand preference in gestural communications have been done, although studies of lateralization of tool use are many. Plentiful examples of gestural communications shown in this book might stimulate research on the symbolic connotations of ape gestures and on hand preference in gestures. Last but not least, we expect that our efforts at exhaustive listing of behavioral patterns will merge in future with research on the chimpanzee genome. Genomic research on chimpanzees, although successful, is still far from explaining the genetic basis of behavioral

patterns. We anticipate that this ethogram will contribute to the synthesis of the chimpanzee genome study. The audio-visual ethogram will be useful not only in facilitating more precise comparisons between sites, but also in providing students and ordinary people with knowledge and materials that will stimulate them to conserve great apes. Listing the behavioral patterns of a local population of chimpanzees in principle includes: behavioral patterns of the LCA of Pan and Homo; patterns common to the genus Pan, namely chimpanzees and bonobos, but not to Homo; patterns common to the chimpanzee Pan troglodytes, but not the bonobo Pan paniscus; patterns common to eastern and central, but not western and Nigerian chimpanzees, in view of the recent cladistic analyses of culture; patterns unique to the eastern chimpanzees, P.

t. schweinfurthii; patterns unique to the Mahale population; customary patterns unique to many individuals of M group chimpanzees; habitual patterns limited to a few individuals of M group; rare patterns observed only in a single individual. The class of behavioral category outlined above to which each behavioral pattern belongs will not be conclusively clear until extensive research and comparison is done across many groups of local populations. However, a preliminary estimate can be made for some of the best-known patterns; for example, few would deny that groom was in the behavioral repertoire of the LCA. In such cases, we put, in the column termed Category after each behavioral patterns description. Studies of the behavioral patterns of Homo sapiens and bonobo have been published, although to a lesser extent

than studies on chimpanzees. Therefore, behavioral comparisons between chimpanzee, bonobo and human can be made to some extent. Many authors have pointed out behavioral similarities and differences between human and chimpanzee. Some behavioral patterns are common to both human and chimpanzee but not bonobo, as well as patterns common to both human and bonobo but not chimpanzee. 10 Introduction These behavioral patterns are either those of the LCA of Pan and Homo, which either chimpanzee or bonobo has lost, or patterns that were lacking in the LCA but evolved in both man and only one species of Pan in parallel after the split of chimpanzee and bonobo. In such cases, we tentatively show 3, the status of which remains ambiguous. The behavioral patterns of the LCA more likely

are to be found in Categories, or than in others. Preliminary comparisons have already suggested interesting contrasts between chimpanzee and bonobo. If we scrutinize the minute details of the behavioral patterns of these sibling species, we may find many more differences between them. As less is known of the behavior of the two chimpanzee subspecies of central Africa, it is presently impossible to delineate Category above. However, recent progress at Goualougo is encouraging. Also, an increasing amount of information has emerged on West African chimpanzees, especially those of the Ta Forest, Bossou, and Fongoli, in addition to older data from Kanka Sili and Assirik, we can tentatively regard the common patterns of eastern and western chimpanzees as candidates for behavior common to the species Pan troglodytes. Since most

chimpanzees kept in European zoos are of the west African subspecies, behavior of captive chimpanzees provide much information on this subspecies. Behavioral patterns belonging to Categories 5, or 7 are most likely to be cultural, but their status remains ambiguous, until it is determined by careful comparison whether the environmental factors that might explain the local differences are present or absent. Methods In compiling an audio-visual ethogram of Mahale chimpanzees, we mostly followed principles used by Nishida et al. First, we tried to make the ethogram maximally inclusive. Second, we distinguished functional definitions from anatomical ones. Elements of the latter should be provided for all functionally-defined behavioral patterns. In order to attain this goal, we adopted the format of putting verbs in front of nouns or adjectives for

the description of behavioral patterns. For example, we used hunch bipedal or clip leaf instead of bipedal hunch or leaf-clip. Moreover, in this version, we used only a single noun when it was more appropriate. For example, we listed toy instead of play toy because toy is enough to show that play is involved. Third, differently from the previous version, in this paper we included as many video clips as possible in the DVD. If we had no video clips for a particular behavior pattern, then we made effort to add illustrations of the behavioral pattern in the form of photographs, video-frames, or drawings, so that researchers at other study sites can verify whether the patterns that they see are the same or different from the ones described

by us. In order to classify behavioral patterns hierarchically and to describe social relationships, both anatomical and functional terms are needed. This books major aim is to list the names of all the behavioral elements in terms of anatomical details, but we also need functional terms in order to denote behavior that is meaningful in the natural environment. For example, approach is a functional term. Although it is simply walk or run from the anatomical perspective, when one chimpanzee comes closer to another, some interaction will likely occur between them. Therefore, we must have a term approach in addition to walk, and so on. Moreover, from the perspective of cognitive psychology, we need functional terms. For example, a chimpanzee who detours illustrates an important aspect of social cognition,

in that the chimpanzee predicts from experience an associates future behavior and changes her route accordingly, although detour may be nothing but glance plus walk. Deceive is a much more difficult functional term, as it entails an inference by the observer about the apes intentions. Deception may not be linked to any particular motor pattern, but if we ignore this term we will fail to capture an important part of the cognitive abilities of the chimpanzee. T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 3, Springer 2010 11 12 Methods Some behavior occurs as a sequence of different behavioral patterns or behavioral complexes; the charging display is an example of such a sequence. Some terms designate several morphologically distinct behavioral patterns that

occur in similar contexts and have similar functions, although they may not occur simultaneously. Thus, we divide behavior patterns into four types: Type A denotes simple, anatomical terms such as walk bipedal; Type B is for complex anatomical terms such as wrestle; Type C is for simple functional terms such as approach; and Type D is for complex functional terms such as abuse. We designate the applicable Type for each term. We have employed as many terms as possible from previous works, in particular Goodall, van Hooff, Bygott, Berdecio and Nash, Plooij, Kano and Call and Tomasello, as well as Nishida et al. Many new terms, such as groom self are introduced here, but we do not seek to persuade readers to use neologisms. If we were to

follow conventional usage and list terms by alphabetical order, then groom and selfgroom would be placed many pages apart from each other although they designate the same motor pattern. To avoid such inconsistency, conventional terms were rearranged so that the action was emphasized, and this policy of neologism was consistent with the verb first principle outlined above. Three-letter representations of behavioral patterns are useful in recording activities quickly and for displaying them economically in illustrations. Accordingly, we borrowed from Plooij when the appropriate abbreviation was available, but because Plooijs ethogram concerned mostly mother-infant behaviors, most of our abbreviations were new. Goodalls glossary descriptions were reproduced in the explanation of terms, if the behavioral patterns were the same as those seen at Mahale. The category numbers of used in

the glossary below follow the principles explained above in the Introduction. Study Sites of Chimpanzees and Bonobos Local names of the major study-sites referred in the Glossary are listed below with main reference sources. Shorter-term study sites such as Bai Hokou, Beni, Campo, Filabanga, Kabogo, Kanka-Sili, Kasakati, Moukalaba, Ndoki, Odzala, Petit Loango, and Tongo are described in the glossary. Chimpanzees P. t. verus Assirik: Niokolo-Koba National Park, Senegal Bossou: Guinea Fongoli: Senegal Ta: Ta National Park, Cote dIvoire Tenkere: Sierra Leone P. t. vellerosus Ebo: Ebo Forest, Cameroon Gashaka: Gashaka Gumti National Park, Nigeria P. t. troglodytes Goualougo: Nouabale-Ndoki National Park, Republic of Congo Lop: Lop Reserve, Gabon T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 4, Springer 2010 13 14 Study

Sites of Chimpanzees and Bonobos P. t. schweinfurthii Budongo: Budongo Forest Reserve, Uganda Bwindi: Bwindi Impenetrable National Park, Uganda Gombe: Gombe National Park, Tanzania Kahuzi: Kahuzi-Biega National Park, Democratic Republic of Congo Kalinzu: Kalinzu Forest Reserve, Uganda Kanyawara: Kibale National Park, Uganda Mahale: Mahale Mountains National Park, Tanzania Ngogo: Kibale National Park, Uganda Semliki: Toro-Semliki Wildlife Reserve, Uganda Ugalla: Ugalla Forest Reserve, Tanzania Bonobo Lilungu: Lomako: Lomako Forest Reserve Lui Kotale: Salonga National Park Lukuru: Wamba: Luo Scientific Reserve Yalosidi: Catalogue Classification of Behavior Patterns 1. Social interaction 2. Feeding and foraging 3. Expressive behavior 4. Positional behavior 5. Object manipulation. Culture 7. Others 1. Social interaction 1-1. 1-2. 1-3. 1-4. Agonism 1-1-1. Aggression 1-1-2. Threat and display 1-1-3. Submission and defense 1-1-4. Reassurance and appeasement Sexual behavior

1-2-1. Copulation 1-2-2. Courtship 1-2-3. Others Maternal behavior 1-3-1. Maternal/alloparental care and infant behavior 1-3-2. Infant transport 1-3-3. Weaning Grooming 1-4-1. Social grooming and self-grooming 1-4-2. Grooming posture and grooming request T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 5, Springer 2010 15 1 Catalogue 1-5. 1-. 1-7. 1-8. Social/solo play and play invitation Inter-group relationships Inter-species relationships Other social interaction 2. Feeding and foraging 2-1. Eat 2-2. Techniques 2-3. Food sharing 2-4. Hunting 3. Expressive behavior 3-1. Vocalization 3-2. Facial expression 3-3. Gesture 4. Positional behavior 4-1. Locomotion 4-2. Posture 5. Object manipulation 5-1. Hold and transport object 5-2. Tool-use to feed 5-3. Tool-use in threat and aggression 5-4. Tool-use in other contexts. Culture 7. Others 7-1. Physiological behavior 7-2. Self-directed

behavior 7-3. Movement and grouping 7-4. Inspection 7-5. Maintenance 1-1-1. Agonism: Aggression abuse abuse carcass aggress aggress in sexual frustration aggress morally aggress, redirected attack attack concertedly attack human battle bite brush aside chase club club another compress lips drag by hand drag other by hand emasculate fight grab grapple hit immobilize interact with human kick kick other kill kill adolescent male kill adult female kill adult male kill infant kill juvenile leap on ostracize pinch aggressively press press down push push away retaliate Classification of Behavior Patterns 17 roll scratch aggressively show off slap other spar stamp other support support dominant support older support subordinate swing and kick 1-1-2. Agonism: Threat and display aggress bark bark, infantile beat chest break branch break tree charge clap hand compress

lips cough bark display as contest display away display past display toward display, charging display, rain display, streambed drag by hand drag carcass by hand drag dry leaves drum flail flail arm gaze harass hit bush bipedal hit toward hoot hunch hunch and sit hunch bipedal hunch over ignore intervene intervene to separate kick backward quadrupedal kick bipedal kick buttress knock down with both arms knock down with one arm leap bipedal with squared shoulders lift and drop police protest pull rock to roll push backward push down push forward raise arm quickly rock back and forth rock side to side scramble for food scratch dry leaves shake arm, abduct shake back and forth with four limbs shake branch shake branch up and down with feet shake long

object irregularly shake object up and down show off slap buttress or tree trunk slap ground slap self slap-stamp slap wall snub stamp stamp bipedal stamp quadrupedal stamp trot stare fixedly strip leaf supplant swagger bipedal sway woody vegetation swing swing and kick tease threaten throw throw at throw at animate object throw dry leaves throw sand tip head waa bark 1-1-3. Agonism: Submission and defense appease bend away bob bob bipedal bow crouch dab escape flee glance greet grin hesitate hop bipedal on spot hop quadrupedal on spot pant-grunt pant-grunt with bent elbow parry present with limbs extended present with limbs flexed scream shake hand side to side quickly shake other struggle free throw dry leaves vacate 1-1-4. Agonism: Reassurance and appeasement appease bend and stretch knee

confiscate console embrace full embrace half extend hand grasp hand greet hold genitals hug kiss kiss with open mouth kiss with pout face kiss with tongue massage shoulder mount mount, misdirected mouth nod and mouth offer arm pat poke press teeth against back push finger into mouth put heel on back put rump to rump reach wrist toward reassure reconcile shake penis solicit reassurance contact solicit support suck in reassurance take finger in mouth thrust thrust bipedal thrust, misdirected touch touch scrotum 1-2-1. Sexual behavior: Copulation copulate copulate between group copulate dorso-ventral copulate ventroventral dart incest interfere in copulation mount in copulation pant in copulation squeal in copulation thrust 18 Catalogue 1-2-2. Sexual behavior: Courtship bend shrub bend shrub in courtship clip leaf by hand clip leaf by

mouth club ground dance orthograde hop bipedal on spot hop quadrupedal on spot hunch hunch and sit hunch bipedal lift and drop open thighs present with limbs extended present with limbs flexed pull through in courtship push raise arm quickly rap scratch dry leaves shake branch shake detached branch slap branch solicit copulation stamp bipedal stand bipedal strip leaf swagger bipedal swagger on knuckles sway woody vegetation tap heel throw branch thrust bipedal touch 1-2-3. Sexual behavior: Others adduct penis adolescent infertility adolescent swelling aggress in sexual frustration avoid incest check penile erection consort eat semen ejaculate erect penis estrus flabby bottom fumble clitoris fumble penis fumble penis with foot herd herd in coalition inspect genitals lead masturbate rub genitals rub genitals to substrate semen sexual skin shake

penis swelling of sexual skin thrust in vacuum 1-3-1. Maternal behavior: Maternal/alloparental care and infant behavior abandon adopt aid in locomotion balance bounce foot/feet up and down rhythmically care alloparentally care alloparentally for another species care maternally change nipple clamp elbow cling cradle with hand cradle with leg extend arm as ladder extend leg as ladder gestation period give birth grasp hug kidnap kiss with pout face lie and hug mother-offspring relations mummify nuzzle orphan pat pregnancy protect rescue respond to dead chimpanzee retrieve infant return infant rush to embrace scan share bed share food spring suck suck in reassurance suckle travel alone after childbirth wedge 1-3-2. Maternal behavior: Infant transport crutch grasp and push shoulders put dorsal put dorsal from ventral put ventral put ventral from dorsal

ride bipedal ride clinging ride dangling ride dangling and touch dry leaves ride dorsal ride jockey ride prone ride quadrupedal ride supine ride ventral ride ventral with limb extended scoop infant solicit riding step up on leg touch transport corpse of infant transport two offspring transport with hand support transport with thigh support 1-3-3. Maternal behavior: Weaning approach and withdraw brush aside contact with nipple cover nipple distract drop infant fall over backward fend ignore interfere in copulation leave to protest lower head and shoulder lower rump monitor mother push ahead push away reject reject infant reject-move reject-sit scatter dry leaves shake rump shrug stand bipedal stare fixedly with head down and bottom up throw temper tantrum turn around wean whimper whimper-scream Classification of Behavior Patterns 19 1-4-1.

Grooming: Social grooming and self-grooming allow clack teeth crush gaze groom groom briefly groom by hand groom carcass groom leaf groom mutually groom object or substrate groom reciprocally groom self groom unilaterally groom with mouth groom wound groom, dyadic groom, polyadic groom-branch-clasp groom-hand-clasp groom-hand-clasp unilaterally inspect leaf presbyopia remove remove lice remove sand flea scrape scratch scratch socially scratch socially, poke type scratch socially, stroke type smack lip sputter squash ectoparasite on arm squash leaf 1-4-2. Grooming: Grooming posture and grooming request grab groom-branch-clasp groom-hand-clasp groom-hand-clasp unilaterally hold head or face lean forward lie supine with legs apart lie with back to another look back lower arm lower head lower leg offer arm offer back pull pull face to face pull head with hands push raise and hold

leg raise arm slowly raise arm to hold branch raise arm with elbow bent raise leg while lying raise others arm raise others chin raise others leg scratch dry leaves scratch self signalling sit and turn back sit behind individual sit face to face sit sideways solicit grooming solicit grooming turn stare fixedly with head down and bottom up touch turn face upward turn up lip 1-5. Social/solo play and play invitation backstroke balance beat sole with palm bend shrub bite and pull hairs bounce foot/feet up and down rhythmically chase circle orthograde circle orthograde off ground circle quadrupedal club other collect dry leaves contact watercover self drag and circle drag by hand drag by mouth drag dry leaves drag other by hand drag other by mouth drape

drip drop drop bark bits drop self dunk face enter into hole fend flail flail arm flee flop glove grab grab and shake grapple grasp and heave grasp hand handicap self hang and spin hang and stamp hang in sloth position hang upside-down by feet hang upside-down by hands hang with legs pitterpat hang-wrestle headstand hit and run hit bush bipedal hit ground with fist hold body part in mouth hop quadrupedal on spot hurl self interact with human interfere play join play kick kick back kick heel kick other kill time kick up leap down and wait leap on leap up lie and watch lie on other lie supine lie supine and shake arms and legs lift and drop look at water look between thighs mount, misdirected

mouth mouth and shake nod and mouth nod to water surface nod to water surface and mouth water nod with body part in mouth nod with object in mouth nod with play face pass under pick up and release pile dry leaves pinch clitoris pirouette play play bite play face play face, full play face, half play, imaginary play in bed play-pant play, parallel play, rough and tumble play socially play socially with object play solo play walk play with another animal play with object play with sand play with urine play with water poke poke with play face press press 20 Catalogue down press neck with lower arm press object on pull pull down pull each other pull leaf-pile pull object from opposing sides pull rock to

roll pull with mouth push push forward push head into ventral push leaf-pile push/ pull swing put face to put mouth into water rake rebuff play release infant to fall ride dangling and touch dry leaves roll rotate fruit rub dorsum to conspecific rub object to body scatter dry leaves scratch dry leaves shake branch up and down with feet shake face side to side shake face with object in mouth shake head shake object up and down shake off shake other shake rock up and down with feet sit on slap in invitation slap other slap self slide down boulder solicit play solicit play with object solicit play with object in mouth somersault somersault, backward somersault, backward with dry leaves somersault, forward somersault, side spar spit water

splash water stamp stamp in invitation stamp other stamp quadrupedal stamp water stand with head down, bottom up step on stir water struggle free suspend and shake up and down swing swing above swing and kick swing forward and upward tag tease throw throw at throw at animate object throw dry leaves throw sand throw stone or rock thrust thrust in vacuum thrust, misdirected tickle tickle self with object tilt head toy trample travel and play trifle with walk in sloth position walk quadrupedal on backs of hands wrestle wrestle bipedal wrestle with fingers 1-. Inter-group relationships attack concertedly battle copulate between group emigrate follow specific female immigrate inter-group behavior intrude kill adult male kill infant patrol respond to neighboring unit group stalk territory transfer unit group visit

1-7. Inter-species relationships abuse carcass attack human bark care alloparentally for another species catch with hand clap hand club ground drop branch flee from colobus male groom carcass habituated hide interact with human kill another species play with another animal respond to baboon respond to dead animal respond to leopard respond to lion respond to predator throw at throw at animate object trifle with whisk fly with arm whisk fly with leafy stick wraa 1-8. Other social interaction alliance coalition conflict behavior deceive friendship imitate interfere interfere copulation interfere fishing interfere play misunderstand reject restrain snatch steal stroke teach turn face away wait turn 2-1. Feeding and foraging: Eat chew disperse seed drink drink from hole in tree drink from lake drink from stream eat eat algae eat

ant eat beetle larva eat blossom eat Camponotus ant eat carcass eat Crematogaster ant eat Dorylus ant eat egg eat eye mucus eat feces eat fruit, inner skin eat fruit, pulp eat gall eat honey eat Classification of Behavior Patterns 21 infant eat insect eat leaf eat meat eat nasal mucus eat Oecophylla ant eat petiole eat phloem eat pith eat resin eat rock eat root eat seed eat semen eat termite eat termite soil eat xylem grunt, food lick lick lips lick rock lick wood lick wound reingest vomit scream, food suck 2-2. Feeding and foraging: Techniques break bone at joint break branch break branch with foot break open by pushing chin break tree brush away from branch brush away from self catch with hands dig dig

for underground storage organ by hand dig for water by hand dip fruit wadge into water dip hand and lick water discard fruit skin with mouth dunk eat with foot fill mouth with food grope handedness keep water in mouth mop ant peel with hand peel with teeth pick out pulp pick up discarded food process food pull down pull out pull through with hand pull through with mouth remove objects from water surface rinse scrape scrub pelt shake wet arm to catch termites sniff fruit spit juice spit seed squeeze stand bipedal store swallow leaf swallow seed tear throw fruit against rock touch fruit transport food twist wadge wadge by adding leaf wadge without adding leaf wash wash colobus skin 2-3. Feeding and foraging: Food sharing allow

beg clamp elbow extend hand to beg fend give mouth for begging peer push away share food take turn around twist 2-4. Feeding and foraging: Hunting chase drag to kill dunk eat meat expel hunt hunt with tool knock with both arms knock with one arm monitor monkeys scrub pelt share food stalk twist wash wash colobus skin 3-1. Expressive behavior: Vocalization bark bark, infantile choke in tantrum clack teeth cough bark distress call grunt grunt, aha grunt, extended grunt, food grunt in bed hoo hoot huu mass excitement pant pant-bark pant-grunt pant-hoot pant in copulation play-pant scream scream, food smack lip sputter squeal in copulation staccato call vocalize waa bark whimper whimper-scream wraa 3-2. Expressive behavior: Facial expression compress lips flip lip funny face grin grin-full-closed grin-full-open

grin-low-closed grin-low-open hoot face play face play face, full play face, half pout protrude tongue pucker cheek relaxed face sneer 3-3. Expressive behavior: Gesture beat chest bend and release bend and stretch knee bend away bob bob bipedal bow clap hand clip leaf by hand clip leaf by mouth dab extend extend hand extend hand and put knuckle on ground extend hand to beg 22 Catalogue extend hand to hold tree extend hand, palm downward extend hand, palm sideways extend hand, palm upward extend leg fall over backward flap hit ground with fist hit toward lower arm lower head lower leg make sound offer arm open thighs pull through in courtship raise and hold leg raise arm bipedal raise arm quickly raise arm slowly raise arm to hold

branch raise arm with elbow bent raise leg while lying rap reach wrist toward scratch dry leaves scratch self signalling shake arm, abduct shake arm, adduct shake back and forth with four limbs shake branch shake branch up and down with feet shake detached branch slap slap branch slap buttress or tree trunk slap ground slap in invitation slap self slap-stamp slap wall solicit companion solicit copulation solicit grooming solicit play solicit reassurance contact solicit riding solicit support stamp stamp bipedal stamp in invitation stamp quadrupedal strip leaf throw temper tantrum tip head 4-1. Positional behavior: Locomotion backstroke brachiate climb climb cliff climb vertical climb vertical, extended elbow crutch descend descend by brachiating descend orthograde feet first descend tree trunk feet first descend tree trunk head first drop

self fall gallop hop bipedal on spot hop quadrupedal on spot leap leap between trees leap between trees with object leap bipedal leap bipedal with squared shoulders leap down leap quadrupedal leap up limp locomote pirouette play walk retreat retreat bipedal run run bipedal run quadrupedal run tripedal slide down vertically somersault somersault, backward somersault, forward somersault, side stamp trot stumble sway and move swing swing and grasp swing forward and upward tumble wade walk bipedal walk in sloth position walk quadrupedal on backs of hands walk quadrupedal on knuckles walk quadrupedal on palms walk tripedal 4-2. Positional behavior: Posture cling cross cross arms cross arms on head cross legs crouch dangle groom-branch-clasp groom-hand-clasp hang hang in sloth position hang tripedal hang upside-down by feet hang upside-down by

hands hang-stand headstand hunch hunch and sit hunch bipedal hunch quadrupedal lean lean forward lie lie and hug lie lateral lie prone lie-sit lie supine lie supine with legs apart lie with legs crossed raise raise and hold leg raise arm to hold branch raise leg while lying sit sit and lean sit face to face sit prone squat stand bipedal stand quadrupedal stand quadrupedal heel up stand with head down, bottom up 5-1. Object manipulation: Hold and transport object discard drag by hand drag carcass by hand drag carcass by mouth grasp hold object in groin pocket hold object in mouth hold object in neck pocket hold object on head leap between trees with object lift lift and drop lift rock move pick up suspend transport transport

bipedal transport corpse of infant transport food transport in foot transport in groin pocket transport in Classification of Behavior Patterns 23 hand transport in mouth transport in neck pocket transport on back transport on head or nape transport on shoulder transport quadrupedal 5-2. Object manipulation: Tool-use to feed brush-tip fishing probe clip leaf for fishing probe dig for army ant with hand/stick dig for underground storage organ with tools dig for water with stick dig stingless bees nest with stick dig termite nest with stick dip ant and wipe dip ant directly dip fluid expel extract fish fish for carpenter ant fish for termite fold leaf hammer nut with stone hammer nut with wood hunt with tool leaf-midrib spoon leaf-sponge leaf-spoon lever open make tool mop ant with

leaves perforate pick out bone marrow pick out nutmeat pound pound food on object pound food on wood pound insect pound pestle pound resin prop anvil puncture push and pull sponge push object into scoop algae tool use tool 5-3. Object manipulation: Tool-use in threat and aggression club club ground club other drag branch drop drop branch flail long object throw at inanimate object throw branch throw splash throw stone or rock tool use tool 5-4. Object manipulation: Tool-use in other contexts bed bend branch clip leaf by hand clip leaf by mouth comb cover from rain dab leaf drape groom leaf hook branch make bed make cushion make day bed make ground bed or cushion make leaf cushion make night bed make tool play socially with object

play with object probe probe nasal passage push object into push to shoulder seat-stick sniff with tool solicit play with object solicit play with object in mouth squash leaf stepping-stick tickle self with object tool tool composite tool set toy use tool whisk fly with leafy twig wipe with detached object. Culture bend and release bend shrub in courtship clip leaf by hand clip leaf by mouth comb cover from rain culture dab leaf dig for army ant with hand/stick dig for underground storage organ by hand dig for underground storage organ with tools dig stingless bees nest with stick dig termite nest with stick dip ant and wipe dip ant directly dip fruit wadge into water eat algae eat ant fashion fish for carpenter ant fish for

termite fold leaf groom leafgroom-hand-clasp hammer nut with stone hammer nut with wood inspect leaf leaf-sponge lever open lick rock lick wood make leaf cushion perforate pick out bone marrow pick out nutmeat pound food on object pound food on wood pound insect pound pestle pound resin prop anvil pull leaf-pile puncture rub muzzle scoop algae scratch socially, poke type scratch socially, stroke type sputter squash ectoparasite on arm squash ectoparasite on palm squash leaf strip leaf tap heel throw fruit against rock throw splash tradition whisk fly with leafy stick wipe with detached object 24 Catalogue 7-1. Physiological behavior adduct penis bristle close eyes cough death defecate defecate, prone defecate quadrupedal defecate, sitting diarrhea die ejaculate erect penis fart hiccup open eyes parry senescence shiver sleep sneeze

sniffle urinate urinate, prone urinate quadrupedal urinate, sitting vomit wriggle yawn 7-2. Self-directed behavior bite self clasp self fumble clitoris fumble nipple fumble penis fumble penis with foot inspect self lick wound masturbate medicate self pick ear pick nose probe nasal passage push peri-anogenital region with finger scratch self slap self suck self suck thumb suck toe tickle self with object 7-3. Movement and grouping approach approach and withdraw arise arrive avoid avoid fallen log bend away chase core area creep dart depart depart together detain detour family fission and fusion flee flee after startle follow follow in contact follow specific female go ahead group herd hesitate hide lead leap up in surprise leap into lap leave leave and climb party pass pass under pass with body contact

regulate direction rest reunite reverse scan search for conspecific separate solicit companion stalk stand quadrupedal heel up step over supplant territory travel travel alone after childbirth vacate wade wait for companion 7-4. Inspection crowd gaze glance grope inspect inspect fruit inspect genitals inspect leaf inspect self inspect wound listen look around look at water look back look between thighs look up monitor monitor monkeys monitor mother patrol peep peer peer together probe probe with finger respond to dead animal respond to dead chimpanzee rummage scan search for conspecific search for object sniff sniff finger sniff fruit sniff mouth sniff with tool stand bipedal stare fixedly tilt head touch fruit touch with foot vigilance watch 7-5. Maintenance brush away from self catch with hand comb cover from rain dab

leaf eat eye mucus lick wound medicate self pick ear pick nose probe nasal passage push peri-anogenital region with finger rub dorsum rub hand or foot rub hand with hand rub muzzle scratch self shake head shelter from rain sickness whisk fly with arm whisk fly with leafy stick wipe with detached object Remarks 1. Principle of text descriptions in the catalogue goes as follows: Start with first-person singular verb whenever possible Assume chimpanzee as subject of verb Omit articles for conciseness Omit verb to be Eliminate redundancy, e.g. one hand to hand 2. Use of symbols, etc. Behavioral patterns listed in this book are bracketed by Local behavioral patterns that have been recorded at other sites, but not at Mahale, are shown in bold italics and are

not categorized In addition to the behavioral catalogue, we include some basic terms such as Infant and place names of study sites, so readers can use this book as a dictionary. Such terms are discriminated from behavioral patterns by using bold font Synonyms are in italics T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0, Springer 2010 25 Glossary A A abandon Nishida: Alloparent leaves infant charge and departs without returning it to its mother. Also occurs in bonobos. Category 2. See Fig. 1. Fig. 1 Abandon: Five-year-old female, Blondy, left infant charge 10 m away from its mother abuse Nishida: Older animal apparently mistreats an infant by brush aside, push away, roll, kick, push, stamp, hit, pull, pull foot, press legs, etc.

Mother may abuse her infant. Also seen in bonobos of Wamba. See video. abuse carcass Behavior directed to dead animal without eating it after chimpanzees have killed it. Apparently not directly related to consumption of meat. T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 7, Springer 2010 27 28 A adolescent swelling Include slap, stamp, kick, knock with both arms, knock with one arm, throw, throw branch, bite, suspend, shake object up and down, and sit on. Colobus and leopard carcasses were abused at Mahale. However, chimpanzees also hug, kiss with pout face, groom, lie on other, and even share day bed. Cf. respond to dead animal, trifle with, and care alloparentally for another species. 7. See video. adduct penis Goodall: Erect

penis spasmodically jerks up and down. See also erect penis. See video. adolescent Sexually mature but socially immature. Males 915 years old, Females 913 years old. See Fig. 2. Fig. 2 Adolescent: Ten-year-old male of K group, Masisa adolescent infertility Adolescent females usually do not conceive although they copulate often. adolescent sterility See adolescent infertility. adolescent swelling Goodall Adolescent swelling: The first tiny swellings of the sex skin initially involve only the labia. These gradually become larger and more regular over the next 3 years or so until the first adult-sized swelling attracts the attention of the adult males See Fig. 3. adopt 29 A Fig. 3 Adolescent swelling: Nine-year-old female, Abi adopt Long-continuing alloparental care of orphaned infant by single alloparent, sometimes lasting more than a year.

At Mahale, at least five older adolescent or young adult nulliparous females seen to hug, groom, transport, protect, play, share food, and share bed with orphan under 5 years old. Long-term adoption by adult males was reported from Ta. In bonobos, care of 2-year-old infant by newly immigrated female seen for one week. See Fig. 4. Fig. 4 Adopt: Ten-year-old female, Tula, adopted 3-year-old female, Maggie, and cared for her for more than 2 years 30 A adult aha-grunt Sexually and socially mature. Males 1 years old, Females 14 years old. See Fig. 5. Fig. 5 Adult: Eighteen-year-old male, Nsaba aeroplane See balance. aggress attack or threaten. See video. See also video aggress, redirected. aggress in redirection See aggress, redirected. aggress in sexual frustration Aggress to sexual partner

in response to refusal to copulate. 5. See video. aggress morally de Waal Moralistic aggression. Chimpanzee retaliates against companion after betrayal. When alpha male supported adult female against his subordinate male ally during competition for meat, latter chased alpha male. aggress, redirected Aggress to another individual after being aggressed against by more dominant individual. Goodalls Redirection of aggression. Kanos Redirection. See video. aggressive scratch See scratch aggressively. aha-grunt See grunt, aha. aid in combat 31 aid in combat See support. aid in locomotion Locomotory aid: Mother or other individual assists its charge by extending arm or leg and retrieving infant stranded in tree, e.g., if it cannot cross gap. Cf. extend arm as ladder: allomother who is below infants mother extends arm upwards to mother to facilitate infants

return to her. See videos. See also video fall. aimed throw See throw at. algae-scoop See scoop algae. alliance Short-term and expedient mutual support against common rival. Cf. coalition. See video. alloparental care See care alloparentally. allow Individual A permits individual B to approach, make contact with A and perform act of concern to A. For example, A allows B to suck As wound, to groom A, to take food from As control, to kidnap As infant, etc. May include sit, stand quadrupedal, watch, etc. See video. alpha male Most dominant male of unit group or community. All other individuals pant-grunt to him. See video. ant-dip-single See dip ants directly. ant-dip-wipe See dip ants and wipe. ant-fish See fish for carpenter ants. anvil-prop See prop anvil. appease Goodall:

Make submissive gestures directed toward a dominant individual after aggression, or in an attempt to prevent aggression. Includes pant-grunt, touch, extend hand, kiss, embrace half, present with limbs extended, present with limbs flexed, groom briefly, etc. Bonobos at Wamba appease by touch, embrace A 32 A attack half, present, groom briefly, etc. approach Goodall: Individual moves toward another. Plooijs Approach. van Hooff divided approach into smooth and hesitant. Includes walk quadrupedal, walk bipedal, and run. Cf. leave. See video. approach and withdraw Infant, especially weanling, repeatedly approaches and withdraws from mother often with whimper when wanting to suck from mother. Estrous female may also do so when soliciting mating from male. Cf. leave to protest and monitor mother. 5. See video. arise Shift from reclining posture to sitting

or standing. Rising by groomee stimulates groomer to resume grooming. 2. See video. arm ladder See extend arm as ladder. arm round See embrace half. arm stretch See extend hand, palm downward. arm swing See brachiate. arm threat See raise arm quickly, shake arm and hit toward. arrive Goodall: Chimp arrives and joins the group. Often accompanied by greeting. See video. Assirik Study area of Pan troglodytes verus in Niokolo-Koba National Park, Senegal. Research done by W.C. McGrew and C.E.G. Tutin from 197 to 1979. attack Goodall Attack: Aggressive physical contact between two, or more, individuals. Includes charge, push, hit, grab, scratch, pinch, slap, drag, kick, stamp, and bite. Cf. battle. Also in bonobos. See video. attack concertedly 33 attack concertedly Nishida et al. Gang attack. Two or

more chimpanzees simultaneously attack one or two others. Occurs both within and between groups. See also ostracize. Also in bonobos of Wamba. See video. attack human Wild chimpanzees occasionally attack human infants and juveniles and kill and eat them. Recorded from Gombe and near Budongo. Habituated adult male chimpanzees occasionally push, slap or kick human observers as part of charging displays. Non-habituated adult female once bit two adult male humans fiercely when her infant was being attacked by M group males at Mahale. See also interact with human, play with another animal and eat meat. 5. auto-groom See groom self. avoid Locomote away from another. Plooijs Avoid, Goodalls Avoidance, van Hooffs Avoidance. See detour, creep and hide. Kanos Avoidance for bonobos. See video. avoid fallen log When fallen

log blocks path, infant moves onto log, then resumes ride on her back after mother passes under log. 3. See video. avoid incest Does not mate with close relatives such as parent, sibling or offspring. Incest avoidance known from many mammals and birds. See incest. A B B baboon See respond to baboon. backstroke Solo locomotor play. Infant or young juvenile slides supine downwards, on slope covered with dry leaves, head first while kicking ground with feet, while keeping both arms ahead. Performed by juvenile male, Michio, and infant female, Athena. 9. See video. Bai Hokou Study area of Pan troglodytes troglodytes in Dzanga-Sangha region of Central African Republic. Research done by RW Carroll in 19881989. balance Nishida Aeroplane, Plooijs Balance: Mother or adult/ adolescent alloparent lying supine

holds infant and lifts infant above ground by one to four of its limbs. Caretaker may gently bounce infant up and down with foot or feet. Caretaker may grasp one or both of infants hands. Similar behavior by bonobo and gorilla mothers. See video. bang See throw fruit against rock. bare teeth See grin. bark Goodall: Loud, sharp sounds, usually given in long sequences with much variation in pitch. Functions to protest to another individual of same or different species. Probably corresponds to bonobos Bark or Wa! call. See also cough bark, pantbark, waa bark, and wraa. See video. 34 bark eating 35 bark eating See eat phloem. bark, infantile High-pitched bark emitted by infants. 3. See video. battle Direct intergroup agonistic interactions, in which both sides contain

multiple males. Two groups concertedly attack each other. 3. beat chest Hit chest with fists. Idiosyncratic element of male charging display at Gombe. Cf. slap self. Absent at Mahale. beat ground with stick See club ground. beat sole with palm Lie supine and repeatedly and simultaneously beat each sole with palm of corresponding left or right hand. Play solicitation signal and also solo play shown by adolescent male, Cadmus. 9. See video. beat with fist See hit. bed Goodall Nest: Platform or bed for sleeping or resting. See also make day bed, make night bed, grunt in bed, and play in bed. beg Goodall: Beg for food, toy, or any desirable object from the possessor. Includes extend hand to beg and mouth for begging. Begging is often accompanied

by whimpering, and if unsuccessful, the beggar may even throw a tantrum. Kanos Food beg for bonobos. See video. See also video reject-move. beg with hand See extend hand to beg. beg with mouth See mouth for begging. bend and release Whiten et al. Branch din: Bend and release saplings, so noisy movement alerts others. Customary only at Lop. Recorded among Lomako bonobos during inter-group encounters. Absent at Mahale. B 3 B bipedal swagger bend and stretch knee Sit orthograde, bend and stretch knee while extending arm palm down. Shown by adult male, Fanana. Idiosyncratic greeting response to adult female. See video. bend away Goodall: With elbow and wrist flexed, arm drawn close to body, the chimp leans slightly away from a passing higher ranking animal. This is

a submissive gesture. van Hooffs Flinch/shrink. Plooijs Bend away. Also seen in bonobos of Wamba. 2. bend branch Bend attached branch when making bed or cushion. See make bed and make cushion. See video. bend shrub Bend shrub, grass, bamboo or herb with one or both hands. In making day-bed or in courtship on ground, done in sitting or quadrupedal posture. In charging display done bipedally. Done also by immatures as solo play. See video. bend shrub in courtship Nishida Shrub-bend courtship. Male courtship pattern at Mahale. While in sitting or quadrupedal posture, push down or pull stem of shrub, bamboo grass, or herb such as ginger, and put one foot on plant; repeat many times same series of actions, thus making crude ground-cushion or bed. Usually followed

by stamp or rap ground, or rarely by hop quadrupedal on spot. Customary among males at Mahale. Seen also as courtship at Ta. Absent in bonobos of Wamba. See video. Beni Study area of Pan troglodytes schweinfurthii in eastern province of DRC. Research done by A. Kortlandt in 19 0, 19 3 and 19 4. bipedal hunch See hunch bipedal. bipedal jump See leap bipedal and hop bipedal on spot. bipedal run See run bipedal. bipedal sex dance See dance bipedal. bipedal swagger See swagger bipedal. bipedal transport 37 bipedal transport See transport bipedal. bipedal walk See walk bipedal. birth See give birth. bite Use teeth to apply force to object. Includes Goodalls Bite: nipping or cutting into the anatomy of another individual by pressing the teeth on

the skin and closing the jaws hard. Plooijs Bite. Goodall did not differentiate from mouth. Cf. mouth and play bite. Bonobo females bite more often than males. Male bonobos do not bite females. See video. bite and pull hairs Player bites some hairs of partner and pulls them with teeth. Infant male, Ichiro, once bit and pulled, and another time sucked, hairs of adult male. Juvenile male, Xmas, bit and pulled hair of back of infant in play. Playful behavior. 9. See video. bite self Bite own palm, wrist, or arm. Pattern expressed in temper tantrum, in response to stressful situation. See video. bluff over See hunch over. bob Goodall: The body bobs up and down as elbows are flexed and straightened. Shown typically by adolescent males but

also by adolescent females, juvenile males and adult males when a high-ranking individual passes. Sometimes as the dominant recipient of the gesture moves on, the bobber backs away in front of the dominant animal. This may provoke an aggressive response. Bobbing is accompanied by pant-grunts which may become rather frenzied pant-screams. Plooijs Bob. van Hooffs Squat bob. Absent in bonobos. See video. bob bipedal Like bob, but standing on feet only. See video. boredom Goodall Boredom. See kill time. B 38 B branch slap Bossou Study area of Pan troglodytes verus in southwestern part of Guinea. Research organised by Y. Sugiyama and T. Matsuzawa, continuing from 197 to present. Pioneer studies done by A. Kortlandt in the early 19 0s and his colleagues in 19 819 9. bounce

foot/feet up and down rhythmically Lie supine, flex and extend leg rhythmically, thus bounce foot up and down. Element of balance. See video. bow Plooij Bow: Similar to bob in context. Deep flexion of legs but arms extended forward, so that head is lower than hips. Absent in bonobos, but see Pika. Category 3. See video. brachiate Goodall Brachiation. Hanging from branch, swings hand-over-hand to move to terminal branch of tree to eat or to descend to ground. Plooijs Brachiate. Same as Kanos Brachiation and Susman et al.s Armswinging. Cf. swing. See video. branch break See break branch. branch-clasp See groom-branch-clasp. branch din See din branch. branch drag See drag branch. branch-flail See flail long object. branch-haul See hook branch. branch-hook See hook branch. branch shake See shake

branch. branch slap See slap branch. break bone at joint 39 break bone at joint Hold and snap limb bone at joint with both hands. Seen in eat meat. Cf. twist. 7. See video. break branch Chimpanzee detaches branch and throws or drags it in display on ground, or eats from it comfortably above ground. In feeding, chimpanzee breaks off terminal branch laden with fruits or young leaves, carries it to safe place, such as big bough or position closer to tree trunk, and eats from branch. Many large food trees therefore modified into different shapes. Recorded for P. t. troglodytes of Gabon. Seen in bonobos in feeding and before branch-dragging display. 2. See video. break branch with foot Before eating Crematogaster ants or Xylocopa bees, hold both

ends of thick, dry branch by hand and use foot to push forcibly against it, in order to break it in two. Also reported from Goualougo. 7. See video. break open by pushing chin Hold large fruit with fairly soft shell such as ripe Saba comorensis or lemon in mouth and push chin with hand or wrist in order to break open and eat contents. break tree Break tree trunk by leaping and pushing against it, or pushing and pulling with hand repeatedly. Kortlandts Break tree. Element of charging display by adult male. leap, push, pull. Also when eating pith from small tree. Also in bonobos. See video. break wind See fart. bristle Goodall Hair erection: The hairs stand on end. Plooijs Hair erection or pilo erection. Bonobos

bristle, but less so. See video. brush aside Mother sweeps off infant her back by hand in refusing to carry it. Male sweeps away by hand infant who wedges itself between copulating pair. Kanos Rebuff for bonobos. brush away from branch Brush repeatedly with hand dry branch to wipe away ants on it. Used to remove Crematogaster ants, when eating their pupae, larvae, or eggs. See video. B 40 B Bwindi brush away from self Rubs away ants with hand which cling to its own body. See video. brush-tip fishing probe Modified probe used for termite fishing. Removes the leaf from the distal end of the tool with his/her mouth. One end of the tool is then pulled sideways through partially closed teeth several times to fray the

end of the herb into a brush tip. Some individuals were also observed biting the end of the tool with their incisors to pull apart the fibers. First described as brush-stick by Sugiyama for chimpanzees of Campo. Absent at Mahale. See video. Budongo Study area of Pan troglodytes schweinfurthii in Budongo Forest Reserve, Uganda. Research organized by Reynolds, continuing from 1990 to present. Pioneer studies done by and F. Reynolds in 19 219 3, by Y. Sugiyama in 19 19 9, and A. Suzuki in 19 91970. bump See pass with body contact. butt See push head into ventral. buttress-beat See drum. Bwindi Study area of Pan troglodytes schweinfurthii in Bwindi Impenetrable National Park, Uganda. Research organised by C. Stanford from 199 to 2005. C C call See

vocalize. Campo Study area of Pan troglodytes troglodytes within Campo Animal Reserve, Cameroon. Research done by Y. Sugiyama in 19841985. cannibalism See eat infant. care alloparentally Nishida Alloparental behavior: Class of behavioral patterns similar to maternal care, shown to infants by individuals other than mother, including hug, groom, transport, mouth, lie and hug, put dorsal, pat, touch, balance, protect, scratch, share food, take finger in mouth, etc. Similar behavior in bonobos. See video. See also video retrieve infant. care alloparentally for another species Alloparental behavior directed to member of another species. Adolescent female carried dead leopard cub to day bed, hugging and grooming it. Cf. trifle with. 9. See Fig. 41 42 carry C Fig. Care alloparentally for another species: Juvenile female carried dead leopard cub into tree

and groomed it care maternally Complex of nurturing behavior shown by mothers to their offspring. Consists of hug, suckle, transport, groom, protect, play socially, share food, share bed, reject, support, etc. No juvenile offspring is suckled, transported, or allowed to share bed. Adolescent offspring still groomed by mother but usually on reciprocal basis, and supported occasionally during fighting with another individual. Mother grooms adult offspring reciprocally. Bonobo mother supports her adult son even against his rival. See video. carry See transport. catch with hand 43 catch with hand Seize flying insect such as fly by quick sideways jerking movement of hand, then open palm to inspect it. Also in bonobos. grab is used for other contexts. Cf. whisk fly with arm. See video. catch with hands Clutch food

thrown by human. When chimpanzees of K group were given bananas and sugarcane, some chimpanzees of both sexes skillful at catching piece of sugar cane with both hands, even when in tree. Behavior developed without training, in response to human throwing to chimpanzees. change nipple During sucking, infant changes nipple from right to left or vice versa. Plooijs Change nipple. See video. charge Goodall: Fast run directed toward another individual. Bonobos charge too. See video. charging display See display, charging. chase Goodall Chase. Run after fleeing individual, seeking to grab it in aggression, play or hunt. Chaser usually dominant to chased, but not always. Chasing play between adults is slower. Goodall Sometimes, a subordinate individual, screaming or perhaps uttering waa barks, chases after his/her aggressor as the latter

charges away See aggress morally. See also circle orthograde or circle quadrupedal. Bonobo chasing is more ritualized, as chaser often gets ahead of fleeing individual. Kano refers to Chase play for chase in play context. See video. check penile erection Estrous female approaches male in order to investigate whether he is ready to copulate. Nishida: Erection check. 9. See video. chest-beating See slap self. chew Plooij Chew. Press item repeatedly between upper and lower teeth. Also occur in bonobos. See video. childhood See Juvenile. Childhood should be applied only to human children, who are not suckled by mother but yet cannot obtain food enough to subsist by themselves. C 44 C clasp self choke in tantrum Goodall Glottal cramps: During loud and prolonged tantrum screaming, the chimpanzee may

seem to choke and only hoarse squeaks or rasping sounds can be heard. See also throw temper tantrum. Absent in bonobos? 5. circle orthograde Occurs in similar context as circle quadrupedal. Individual moves round and round tree bipedally or tripedally, while holding tree trunk with hand and keeping torso upright. See also circle quadrupedal. 5. See video. circle orthograde off ground Hold tree trunk by hand while revolving around it with feet off ground. Solo play or solicit play. 9. See video. circle quadrupedal In social play, two or more individuals may chase round and round tree, another individual, or without any pivot-point. One or both of players may transport leafy twig in mouth. Chased one may take turn to chase, and vice versa. Occasionally occurs as solo

play. Often accompanied by play-pant at Mahale and Gombe. Seen in bonobos of Wamba. See also circle orthograde. 2. See videos. clack teeth Goodall Teeth-clack: Mouth rhythmically opened and closed during grooming. The teeth are clapped together each time the mouth is closed. Usually the lips are quite tightly drawn over the teeth when the mouth is open. Occurs during vigorous grooming. See also sputter, and smack lip. Not seen in bonobos of Wamba. 5. See video. clamp elbow Goodall Elbow clamp: When small infant pesters mother for food, she may clamp it to her breast with both elbows and continue her meal over its head. See also fend. Cf. cover nipple. 5. clap hand Koops and Matsuzawa Hand clapping. Unhabituated adult female of Nimba Mts. once observed

to clap hands as part of threatening gesture against human observers. Only record of hand clapping by wild chimpanzees, although known in captivity. Absent at Mahale. clasp self Goodall Self-clasp: Sit or lie with arms tightly clasped around self. in a relaxed manner during resting. Not seen at Mahale, nor in bonobos of Wamba. Infants of both species of Pan in captivity show this gesture. clear nasal passage 45 clear nasal passage See probe nasal passage. climb Plooij Climb. Includes not only climb vertical and climb vertical, extended elbow, but also climbing by walk quadrupedal on palms on slanting trunk or bough of big tree. Goodalls Vertical climb and Kanos Vertical climb up for bonobos probably includes both climb vertical and climb vertical, extended elbow. See video. climb

cliff Similar to human rock-climbing without any artificial climbing equipment. 3. See video. climb down See descend. climb vertical Hunt et al.s Flexed-elbow vertical climb. Climb woody vine or small tree by alternately using palmar surfaces of four limbs. See video. climb vertical, extended elbow Hunt et al. Extended-elbow vertical climbing. Extend arms around trunk of large tree and propel upwards by simultaneous thrusting or walking movements of feet. See video. cling Goodall: Gripping a tree trunk or branch with hands or hands and feet. An infant clings to mothers or others belly, back, arm or leg. Also in bonobos. See video. clip leaf by hand Tear leaf by hand, separating midrib and leaf-blade. At Mahale, adolescent male, Cadmus, used to solicit mating. Customary at Budongo, and habitual

at Ta and Kibale. See video. clip leaf by mouth Nishida Leaf clip: Pull leaf repeatedly between lips or teeth with one hand, producing conspicuous sound that attracts attention from prospective sex partner. Among most common courtship displays at Mahale. Act of frustration or play at Bossou and prelude to make drumming-enacted frustration at Ta. Thus, contexts differ from place to place, e.g., nuances of behavior differ between Mahale and Ta. Customary at Budongo and Ngogo. Reported as habitual solicitation of social play for Lomako bonobos. See also use midrib for fishing probe. See video. C 4 C collect dry leaves clip leaf for fishing probe Use midrib of leaf as fishing probe after removing leaf-blade by leafclipping with mouth or hand. Customary in fishing for carpenter ants

at Mahale and termite-fishing at Assirik. See video. close eyes Close eyes: in sleep, as reflex when frightened, and when seemingly in bliss. Adult male, Alofu, lies on ground with eyes closed, apparently in bliss, after stuffing many sweet fruits in his mouth. Plooijs Eyes Closed. See video. club Strike target with long, thick stick in overarm movement. Directed to conspecific, another species, or inanimate object. van Hooff and Kortlandt and Kooij described this pattern. Seen rarely in a few individuals at Mahale. Habitual at Ta and Gombe. Present at Bossou and Assirik. Absent in bonobos. 8. See video. club ground Strike ground with long, thick stick to solicit copulation, by two juvenile males. Adult male, Fanana, clubs ground to coerce estrous female to follow him during consortship.

Juvenile male clubs the ground to threaten another juvenile during play or to threaten newly-immigrated female and human observer. Several adult males, Lukaja, Fanana, and Dogura, club ground or wall of house as part of charging display. See video. club other Club another individual with stick. Juvenile males, Michio and Cadmus, club another in solicitation of play. Adult male, Toshibo, twice hit another adult male with stout stick after flailing it during fighting. 8. See video. See also video fight coalition Long-term and strategic mutual support against common rival. Bonding made by pair such as Pinky and her babysitter, Gwekulo, and Opal and her adult daughter, Ruby. Cf. alliance. See also attack concertedly. See video. coalitionary attack See attack concertedly. coalitionary mate guarding See herd in coalition. collect

dry leaves Infant or juvenile reclines or sits and collects dry leaves by raking, pulling or grabbing and hugs them in solo play. See video. comb 47 comb Whiten et al. Comb: Use stem to comb through hair. Present at Budongo and Goualougo. Absent at Mahale. community See unit group. compress lips Goodall Compressed lips: The lips are pressed tightly together so that the upper lip is bunched up and protrudes beyond the point where the lips meet. Seen during displays and attacks. Plooijs Compressed lips. Absent in bonobos. 3. See video. confiscate To prevent aggression, take away potentially dangerous object such as rock or stout branch from another who is irritated. At Arnhem Zoo, adult female confiscated stone from adult male. De Waal s Disarm. At Mahale,

however, only mothers took away stone or stick from adolescent male or juvenile offspring. 3. See video. conflict behavior Situational acts, when two or more contradictory drives are stimulated in actor, such as groom carcass, kick up, shake hand side by side quickly, etc. consolation See console. console de Waal and van Roosmalen Consolation. Third party touches, hugs, grooms or licks victim attacked and wounded by another. Consolation reported from Gombe, Mahale and Ta. See lick wound. Kanos Conciliating behavior by bonobos. See video. consort Goodall Consortship: An exclusive sexual relationship between adult male and female.the female is taken to a peripheral part of the community range away from other rival males. McGinniss Consortship, Tutins Safari behavior. Consortship seen for Ta chimpanzees. At Mahale, consortship is mating strategy

of non-alpha males. See also lead. Cf. herd. Not seen in bonobos of Wamba. See Fig. 7. C 48 copulate between group C Fig. 7 Consort: Adult male, Kamemanfu and young estrous female, Wakapala of K group, in consort visited Myako camp to eat sugarcane when main party was absent contact water Contact with water includes: stir water, look at water, drink, play with water, splash water, throw splash, etc. See also wade. 7. contact with nipple Weanling puts mouth to mothers nipple without sucking. Final stage of weaning. See video. coprophagy See eat feces. copulate Intromission and pelvic thrusting between male and estrous female. Goodalls Copulation, Plooijs Copulate. See video. See also video in bend shrub in courtship, check penile erection, dart, erect penis, press teeth against

back, and wedge. copulate between group Young, and less often prime, estrous females visit neighboring unit group and copulate at Gombe, Mahale and Ta. Bonobo unit groups occasionally mingle peacefully, and copulation by members of different groups occurs at Wamba and Lomako. Adult copulate dorso-ventral 49 males do not interfere with such copulations. copulate dorso-ventral Male mounts female from rear. Major copulation pattern of mature males and females in chimpanzees and bonobos. See video. See also video interfere in copulation. copulate ventro-ventral Male mounts female from front. Occurs rarely in adolescent females and immature male chimpanzees. More common in adolescent females and mature and immature male bonobos. copulation pant See pant in copulation. copulation scream See squeal in copulation. copulatory dart See dart. copulatory squeal See squeal in

copulation. core area Area used intensively by individual chimpanzee although also used by other members of unit group. Females compete one another for good core area. Cf. territory. cough Goodall:When chimps have colds and coughs they sneeze and cough, sounding like humans. See sickness. Bonobos also cough. See video. cough bark Goodall: Soft bark. A grunt-like sound uttered through slightly open mouth directed by higher ranking chimpanzees to subordinates. Indicates mild annoyance. It functions as a mild threat. Threat call of one syllable. Also in bonobos. See video. cough-threat See cough bark. counter-attack See retaliate. courtship display See solicit copulation. cover breast See cover nipple. C 50 cross legs cover from rain Cover body with leafy twig to avoid rain. Present at Goualougo. Also seen for bonobos of

Wamba. Cf. shelter from rain. Absent at Mahale. C cover nipple Plooij Cover nipple. Goodalls Cover breast: Mother prevents access to breast by placing her arm across her nipple.She may also lie with breasts pressed to the ground or branch. Part of mothers weaning of infant. Absent in bonobos of Wamba. See video. cover self Cover self with dry leaves or sand. Solo play of infants. Category 8. See video. cradle with hand Mother supports newborn infant with arms. Goodalls Cradle. Cf. transport with hand support. Kanos Cradle for bonobos of Wamba. cradle with leg Mother hangs from tree branch supports infant clinging to her belly with one thigh flexed, so that babys body is pressed against her. Cf. transport with thigh support. Plooijs Cradle. Seen in bonobos.

creep Walk stealthily. Goodall states that.a fearful subordinate tries to creep from the vicinity of a particular higher ranking individual. The subordinate proceeds with extreme caution, moving slowly while keeping a constant sharp watch on the other. Bonobos of Wamba show similar pattern. See video. cross Flex one or both limbs to rest on opposite limb. See video. cross arms Flex arms across chest. See video. cross arms on head Flex arms on head. 7. See video. cross legs Flex one leg to rest on opposite leg, which is supported by substrate. See video. crouch 51 crouch van Hooff and Plooij Crouch. Goodalls Crouch: Quadrupedal posture with limbs flexed, hindquarters not turned towards another animal. Goodall states that:.Often occurs in greeting when it is usually accompanied by pant

grunts or during submission after aggression. Kanos Crouch and Present in agonistic interaction for bonobo. See video. crouch-present See present with limbs flexed. crowd Many chimpanzees gather to peer at individual or gaze at object. Occurs when chimpanzee tries to extract thorn from foot, inspects seriously injured skin, carries newly-dead baby, possesses prized food such as meat, or an unusual object in environment that stimulates curiosity. When crowded, chimpanzees appear to allow closer contact with each other than usual. No adult bonobos crowd. See video. crush Press small arthropod such as louse between finger-tips. Absent in bonobos. 8. See video. crutch Goodall: A method of progressing, usually when the chimp is going downhill, during which the arms are used as crutches. With both hands on the ground, the

legs and body are swung forward through the arms. Plooijs crutch. At Mahale, also occurs when a mother carries newborn on belly, accompanied by transport with thigh support. In bonobos, crutch used only as last step of a long walk or as one-step movement while eating or resting. See video. cry See whimper-scream. culture Behavior learned socially after birth, shared by at least most members of one age/sex class in group. See also tradition and fashion. cushion See make cushion. C D D dab Goodall: Rapid hitting movement in which the backs of the flexed fingers are directed toward a more dominant individual, typically an adult male. Only a few individuals, mostly adolescents, have been observed to dab. Usually these individuals dabbed several times in succession, and the

gesture was mostly directed toward the face of the other. Plooijs Dab. At Mahale, seen only in few adolescents and juveniles when pant-grunting to adult males. Absent in bonobos of Wamba. dab leaf Whiten et al. Leaf-dab: Dab leaf on wound and examine it. Customary at Kibale, present at Gombe, Goualougo and Ta. Absent at Mahale. dance orthograde Nishida Bipedal sex dance: Stand bipedal while raising arm or performing branch-shaking display. Accompanied by scratch self. Done by estrous female irritated by apparent indifference of adult male to females sexual advance. Only two adult females, Wakilufya and Ako, of M group showed this type of courtship, but de Waal mentioned similar pattern. Similar pattern in bonobos of Wamba. dangle Plooij Dangle. Goodall: Infant hangs, under, from the side of,

or below mother or another individual with one or both hands. Does not grip with feet. May also dangle from a branch See hang. Seen in bonobos of Wamba. See also ride dangling. See video. dart Estrous female runs quickly few meters after copulation. Often accompanied by copulatory squeal and grin, then, female often lies down for few minutes. Nishidas Copulatory dart. Absent in bonobos. 5. See video. 52 day nest 53 day nest See make day bed. death Chimpanzees die of diseases, predation, mothers death, fall from tree, killing by conspecifics, injuries, etc. See also die. deceive Mislead others to own advantage by not showing posture, gesture, or facial expression usually done in that context, mislead others to own advantage by showing posture, gesture, or facial expression

that is normally inappropriate or irrelevant in that context. Reported from Gombe and captivity. See video. deep incursion See intrude. defecate Excrete feces. See also diarrhea and push peri-anogenital region with finger. See video. See also video push perianogenital region with finger. defecate, prone Lie prone and defecate, when chimpanzee has diarrhea. Category 38. See video. defecate quadrupedal Stand on four limbs and defecate. Common posture. defecate, sitting Defecate while crouching with weight on legs only. Common posture of defecation. See video. defecate with hand support See push peri-anogenital region with finger. depart Begin to travel, increasing distance from starting point. See video. depart together Mother approaches infant to initiate travel together. Older infants and juveniles often precede mothers, who then may follow them. See also pull, put