AnthropologicalScience 107 (2), 141-188, 1999

Ethogram and Ethnography of Mahale Chimpanzees

Toshisada Nishida1, Takayoshi Kano2, Jane Goodall3, William C. McGrew44, and Michio Nakamura1

1 Sub-Departmentof Anthropology,Graduate Schoolof Science,Kyoto University, Kyoto 2 Primate Research Institute, Kyoto University,Inuyama 3 Jane GoodallInstitute, Silver Spring, U.S.A. 4 Departments of Sociology,Gerontology and Anthropology,and of Zoology, Miami University, Oxford, U.S.A.

(Submitted January 4, 1998; Review sent March 4, 1999; Accepted April 26, 1999)

Abstract This paper aims to compile an exhaustive list of the behavioral patterns exhibited by the chimpanzees of the Mahale Mountains National Park, Tanzania. The compilation is based on the glossary compiled by Goodall (1989), but a substantial numbers of new terms have been added. Thus, we list 316 simple anatomical terms, 81 complex anatomical terms, 37 simple functional terms, and 81 complex functional terms, in addition to 116 synonyms. The behavioral patterns are divided into eight categories on the basis of degree of universality: (1) commonly seen in both Homo and two species of Pan, (1?) commonly seen in Homo and only one species of Pan, (2) patterns common to the genus Pan but not to Homo, (3) patterns common to the chimpanzee Pan troglodytes but not the bonobo Pan paniscus, (4) patterns common to eastern (P.t. schweinfurthii) and central (P.t. troglodytes) but not western (P.t. verus) chimpanzees, (5) patterns unique to the eastern chimpanzees, P.t. schweinfurthii, (6) patterns unique to the population of Mahale, (7) patterns unique to many individuals (at least most members of an age/sex class) of M group chimpanzees, (8) patterns limited to a single (idiosyncrasy) or a few individuals of M group. It is most likely that the behavior patterns of the last common ancestor of Homo and Pan are found in Categories 1 and 1? and less likely in Categories 2 and 3. It is possible that behavior patterns belonging to Categories 5, 6 or 7 are cultures.

Keywords: ethogram, chimpanzee, Mahale Mountains, behavior, culture

Corresponding author: Toshisada Nishida Sub-Department of Anthropology, Faculty of Science, Kyoto University Kitashirakawa-Oiwakecho, Sakyo, Kyoto, Japan TEL:+81-75-753-4084 FAX: +81-75-753-4098 E-mail: [email protected] 142 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M .

Introduction This article aims to create an ethogram, that is, to list and describe all the behavioral patterns that have been recorded for the chimpanzees of the Mahale Mountains National Park, Tanzania. Previous such attempts only achieved a preliminary level of classification (Nishida, 1970; Mori, 1982). It is well known that chimpanzees show a great deal of local, as well as age, sex, and idiosyncratic, differences in behavior (for example, Goodall, 1973; Nishida , 1987; McGrew 1998). In which domains are the local behavioral differences most remarkable? This is an important question to be addressed if we want to solve the origins of human culture. The extent of local differences has not been well elucidated , partly because most of the study sites of chimpanzees lack a detailed ethogram. Gombe is an important exception: Goodall and her colleagues were pioneers in preparing a detailed glossary of the behavior of Gombe chimpanzees (Goodall, 1968, 1986, 1989; Bygott, 1974; Plooij, 1984). Consequently, it is relatively easy to compile a list of behavioral patterns of a local population by consulting the Gombe ethogram. A list of the behavioral patterns of a local population of the chimpanzees may in principle include (1) behavioral patterns of our last common ancestor of Pan and Homo, (2) patterns common to the genus Pan, namely chimpanzees and bonobos, but not to Homo, (3) patterns common to the chimpanzee Pan troglodytes , but not the bonobo Pan paniscus, (4) patterns common to eastern (P.t. schweinfurthii) and central (P.t. troglodytes), but not western (P.t. verus) chimpanzees, in view of the recent DNA cladistics (Morin and others, 1994), (5) patterns unique to the eastern chimpanzees, P. schweinfurthii, (6) patterns unique to the population of Mahale, (7) patterns unique to many individuals (at least most members of an age/sex class) of M group chimpanzees , (8) patterns limited to a single (idiosyncrasy) or a few individuals of M group. In addition, there are behavior patterns that we have found in both man and chimpanzees but not bonobos, as well as patterns common to both man and bonobos but not chimpanzees. These behavior patterns are either those of the last common ancestors of Pan and Homo, which either chimpanzee or bonobo has lost, or patterns that were lacking in the last common ancestor but evolved in both man and only one species of Pan in parallel after the split of chimpanzees and bonobos. The last possibility is less likely from the viewpoint of the parsimonious principle. Thus , we add Category 1? for these patterns whose status remain ambiguous. It is most likely that the behavior patterns of our last common ancestor are found in Categories 1 and 1? and less likely in Categories 2 and 3. Since little of the behavior of the chimpanzees of central Africa is known (Kuroda , 1998), it is impossible to delineate the Category (4) above . However, since there has been an increasing amount of information on west African chimpanzees, in particular those of the Tai Forest (for example, Boesch and Boesch, 1999), Bossou (Sugiyama, 1998; Matsuzawa and Yamakoshi, 1996), and Bossou and Kanka Sili' (Kortlandt and Bresser, 1963; Kortlandt and Kooij, 1963; Albrecht and Dunnett, 1971) , we can Ethogram of chimpanzees 143 tentatively regard the common patterns of eastern (Gombe and Mahale) and western (Tai) chimpanzees as candidates of behavior common to the species Pan troglodytes. Studies of the behavior patterns of humans (for example, Eibl-Eibesfeldt, 1972; McGrew, 1972; Morris, 1977) and bonobos (for example, Kano, 1980, 1992, 1998; Kuroda, 1980, 1984; Susman and others, 1980; de Waal, 1988) have been published, although to a lesser extent than studies on chimpanzees. Therefore, behavioral comparisons between chimpanzees, bonobos and humans can be made to some extent. Many authors have pointed out behavioral similarities and differences between humans and chimpanzees. Preliminary comparisons have already suggested interesting contrasts between chimpanzees and bonobos (for example, Mori, 1983; Nishida and Hiraiwa-Hasegawa , 1987; de Waal and Lanting, 1997). If we scrutinize the minute details of the behavioral patterns of these species, we might find many more differences between the two species. We need such comparisons in order to reconstruct the behavior of the last common ancestor of Homo and Pan. The class of behavioral category outlined above that each behavioral pattern belongs to cannot be elucidated before extensive research and comparison is made across many groups of local populations. However, a preliminary estimate can be made for some of the best known patterns. For example, no one would deny that "grooming" would be among the behavioral patterns of the last common ancestor. In such cases, we put "Category 1" , for example, in the last part of each behavioral pattern. Behavior patterns belonging to Categories 5, 6 or 7 are most likely to be cultures, but the 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 ethogram of Mahale chimpanzees, we tried to incorporate some new principles. 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 form 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". Third , we made every effort to list the available references that had illustrations of behavioral patterns in the form of photographs, video frames, or drawings so that researchers at other study sites could verify whether the patterns they see are the same or different from the ones described by us. In order to classify behavioral patterns hierarchically and describe social relationships, both "anatomical" and functional terms are needed. This paper's major aim is to list names for all the behavioral elements in terms of anatomical details, but we also need functional terms in order to describe behavior that is meaningful in the natural habitat. For example, "approach" is a functional term. Although it is simply or 144 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. 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 , and so on. Moreover, from the perspective of cognitive psychology, we need functional terms. For example, a chimpanzee sometimes "detours", namely takes a long way round, avoiding, for example, an ill-tempered male who is sitting on a path. Detour is also simply or . However, the term "detour" illustrates an important aspect of social cognition in that the chimpanzee predicts the associate's future behavior from experience and changes the travel route accordingly. 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 is described by simple, anatomical terms such as "walk bipedal", Type B by complex anatomical terms such as "wrestle" , Type C by simple functional terms such as "approach", and Type D by complex functional terms such as "abuse". We designate the applicable Type in each term. We have employed as many terms as possible from previous works, in particular Goodall (1968, 1986, 1989) and van Hooff (1973). A useful list of behavior patterns among captive chimpanzees by Berdesio and Nash (1981) was also consulted, although their terms were not employed here because their study subjects were limited to infants and juveniles. Many new terms are introduced here, but we do not intend to persuade readers to use neologisms. If we follow conventional usage, "groom" and "self-groom", for example, would be placed many pages apart from each other even though they designate the same action. To avoid such inconsistency, conventional terms are rearranged so that "action" is emphasized. Therefore, the policy of the neologism is consistent with the "verb first" principle mentioned above. Three-letter representations of behavioral patterns are useful in recording activities quickly and describing them in charts. Accordingly, we borrow from Plooij (1984) when the appropriate abbreviation is available, but because Plooij's (1984) ethogram concerned mostly mother-infant behaviors, we have had to coin most of the abbreviations. Goodall's (1989) glossary descriptions are reproduced in the explanation of terms if the behavioral patterns are not different from those observed at Mahale. The ethogram category numbers used in the Ethogram below are the same as used in the third paragraph in the Introduction above.

Ethogram A Abandon (ABD)(Type D): Nishida(1983a): Alloparent leaves her/his infant charge and departs without returning it to its mother. (Therefore, the infant must return to its mother on its own accord or be retrieved.) Also occursin bonobos (Kano, unpublished).Category 2. Abuse (ABU) (Type D): Nishida(1983a): Older animalapparently mistreatsan infant by , , , ,, , ,, , , and so on. Similar behavior is seen in the bonobos of Wamba (Kano, unpublished). Category 1. Adduct, penis (ADP) (Type A): Goodall (1989): "Erect penis spasmodically jerks." Category 1. Adopt (ADP) (Type D): Long-continuing alloparental care of an orphaned infant by a single alloparent. May continue for over one year. At Mahale late adolescent or young adult nulliparous females have been seen in more than five cases to , , , , , , and , (with) a particular orphan under 5 years of age. In the bonobo, adoption of a two-year-old infant by a newly immigrated female for one week has been observed (Kano, 1998). Category 1? Aeroplane (AER): See Balance. Aggress (AGG) (Type D): Attack or Threaten. Category 1. Aggress in redirection (ARE) (Type D): Aggress against another (usually subordinate) individual after being aggressed against by a more dominant individual. Goodall's (1989) Redirection of aggression. Kano's (1998) Redirection. Category 1. Aggress morally (AGM) (Type D): de Waal's (1991) Moralistic aggression. Chimpanzee may retaliate upon one's companion for his/her betrayal. When an alpha male supported an adult female against his subordinate male ally during the competition for meat, the latter became angry and chased the alpha male (Nishida, 1994). Category 1? Aha-grunt: See Grunt, aha. Alloparental care: See Care alloparentally. Appease (APS) (Type D): Goodall (1989): "Make submissive gestures directed toward a dominant individual after aggression, or in an attempt to prevent aggression." Includes , , , , , , and so on. The bonobos of Wamba appease by , , , , and so on (Kano, unpublished). Category 1. Approach (APP) (Type C): Goodall (1989): "Individual moves toward another." Plooij's approaching (APP). Van Hooff (1973) divided approach into "smooth" and "hesitant." Includes , , and . Cf. Leave. Category 1. Arm ladder: See Extend arm as ladder. Arm threat: See Raise arm quickly. Arrive (ARV) (Type C): Goodall (1989): "The chimp arrives and joins the group." Category 1. Attack (ATT) (Type D): Goodall's (1989) Attack: "Aggressive physical contact between two, or more, individuals." Includes , , , , , , , , , , and . Similar behavior patterns exit in bonobos (Kano, 1998). Category 1. Attack concertedly (ATC) (Type D): Nishida and others' (1995) Gang attack. Two or more chimpanzees simultaneously attack one or two chimpanzees. This occurs both within (for example, Takahata, 1985; Nishida and Hiraiwa-Hasegawa, 1985; Nishida, 1994; Nishida and others 1995; Goodall, 1986, 1992) and between groups (Goodall, 1986; Nishida, 1979; Boesch and Boesch, 1999). Gang attack is seen in the bonobos of Wamba (Kano, unpublished). Category 1. Auto-groom: See Groom self. Avoid (AVD) (Type D): Locomotion away from a partner. Plooij's (1984) Avoid (AVO), Goodall's (1989) Avoidance, van Hooff's (1973) Avoidance. See Detour, Creep and Hide. Kano's (1998) Avoidance for the bonobo. Category 1.

B Balance (BAL) (Type A): Nishida's (1983a) aeroplane, Plooij's (1984) Balance (AER): Mother or adult/ adolescent alloparent lying supine holds the infant in a ventro-ventral position and lifts the prone infant from the ground by its four limbs. Caretaker bounces the infant gently up and down with its feet, a pattern strikingly similar to that of humans. The caretaker grasps one or both of the infant's hands. Similar behavior is done by bonobo mothers (Kano, 1998). See Figure 4 of Nishida (1983a) and de Waal (1995, p.64) for photograph. Category 1. Bang: See Throw fruit at. Bark (BAR) (Type A): Goodall (1989): "Loud, sharp sounds, usually given in long sequences with much variation in pitch." It functions to protest against another individual of the same or different species (for example, baboons). This probably corresponds to the bonobo's Bark (Okayasu, 1991) or Wa! call (Kano, 1998). Category 2? Beat ground with stick: See Club. 146 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M.

Beat with fist: See Hit. Beg (BEG) (Type D): Goodall (1989): "Beg for food, toy, or any desirable object from the possessor. Begging is often accompanied by whimpering, and if unsuccessful, the beggar may even throw a tantrum." Kano's (1998) Food beg for the bonobo. Category 1. Beg with hand (BWH) (Type A): Plooij's (1984) Beg with hand (BWH): Beg by stretching hand to the possessor's hand, mouth or food. Goodall's (1989) Beg hand-to-food, hand-to-hand and hand-to-mouth. Kano's (1998) Food beg: hand to hand and hand to mouth. Category 1. Beg with mouth (BWM) (Type A): Beg for food by putting lips to the lips or hand of feeding possessors. Goodall's (1989) Beg mouth-to-mouth. Plooij's (1984) Beg with mouth (BWM). See Figure 14. 1 of Nishida (1990) for photo. Kano's (1998) Food beg: mouth to hand and mouth to mouth. Category 1. Bend away (BEN) (Type A): Goodall (1989): "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 Hooff's (1973) Flinch/shrink. Also seen in the bonobos of Wamba (Kano, 1998). Category 2. Bend shrub (BES) (Type B): Nishida's (1997) Shrub-bend, Male courtship pattern: While sitting, push down the stem of a shrub, grass, or herb such as ginger, and put one foot on the plant, and repeat many times the same series of actions, thus apparently making a crude ground cushion or bed . Usually followed by stamping or thumping the ground. Not seen in the bonobos of Wamba. Category 7. Bipedal hunch: See Hunch bipedal. Bipedal jump: See Leap bipedal. Bipedal run: See Run bipedal. Bipedal sex dance: See Dance bipedal. Bipedal swagger: See Swagger bipedal. Bipedal transport: See Transport bipedal. Bipedal walk: See Walk bipedal. Bite (BIT) (Type A): Goodall's (1989) Bite: "nipping or cutting into the anatomy of another individual by pressing the teeth on the skin and closing the jaws hard." Plooij's (1984) bite (BIT): Goodall (1989) did not differentiate from "mouth." In bonobos females bite more often than males (Kano , 1998). Male bonobo does not bite female (de Waal and Lanting, 1997). Category 1. Bob (BOB) (Type A): Goodall (1989): "The body bobs up and down as elbows are flexed and straightened. Shown typically by adolescent males but also by adolescent females and juvenile 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 ." Plooij's (1984) Bob (BOB). Van Hooff's (1973) Squat bob. Absent in bonobos. Category 3. Bow (BOW) (Type A): Plooij's (1984) Bow (BOW): Similar to Bob in the context. However, this is deep flexion of legs but arms flexed forward, so that head is lower than hips. Absent in bonobos. Category 3. Brachiate (BRA) (Type A): Goodall's (1989) Brachiation: "Hanging from branch , animal moves by swinginghand-over-hand." Also, Goodall's (1989) Swing: "During brachiation in the trees ... when the body is swung through space between one support and the next." Plooij's (1984) Brachiate (BRA) . Same as Kano's (1998) Brachiation and Susman and others' (1980) Armswinging. See Figure 5 of Goodall (1968) for illustration. Category 1-2. Branch break: See Break branch. Branch drag: See Drag branch. Branch shake: See Shake branch. Break branch (BRB) (Type A): Chimpanzee breaks off a branch in order to throw or drag it in display on the ground (see Drag branch), or eat from it comfortably above the ground. In feeding, chimpanzee breaks off a terminal branch laden with fruits or young leaves, carries it to a safe place such as a big bough or position closer to the tree trunk, and eats from the branch. By such manipulations, many large food trees have been modified into different shapes . Recorded for P. t. troglodytes of Gabon (Takenoshita and others, 1998). Seen among the bonobo in feeding context and before branch-dragging display (Kano, 1998). Category 1 or 2. Break tree (BRT) (Type B): Break a tree trunk by leaping and pushing against the trunk of a sapling. Kortlandt's (1967) Break tree. Element of charging display by adult male. , , . Seen also in bonobos (Kano, unpublished). Category 2. Bristle (BRS) (Type A): Goodall's (1989) Hair erection: "The hairs stand on end." Plooij's (1984) Hair Ethogram of chimpanzees 147 erection (HAI). Bonobos also bristle, but to a lesser extent (Kano, 1998). Category 1 or 2. Brush aside (BRU) (Type A): Mother sweeps off her infant from her back with her hand in refusing to carry it. Male partner sweeps away with his hand infant who squeezes between copulating pair. Kano's (1998) Rebuff for the bonobo. Category 1. Bump (BUM) (Type B)*: While walking along, chimpanzee bangs its body (for example, shoulder) against another (occasionally human observer), apparently intentionally. Likely absent in bonobos. Category 7 or 8?

C Cannibalism: See Eat chimpanzee. Care alloparentally (CAL) (Type D): Nishida's (1983a) Alloparental behavior: Class of behavioral patterns similar to maternal care, shown to infants by individuals other than the mother, including , , , , , , , , , , , , , and so on. See Nishida (1983a) for photographs . Similar behavior has been seen in the bonobo (Kano, 1998). Category 1. Care maternally (CAM) (Type D): Complex of nurturing behavior shown by mothers to their offspring. Consists of , , , , , , , , , , and so on. Bonobo mother supports her adult son even against his rival (Kano, 1992). Category 1. Carry: See Transport. Catch with both hands (CAB) (Type A): Clutch airborne food thrown by a human. When chimpanzees of K group were given bananas and sugarcane, some chimpanzees both sexes became skillful at catching a piece of sugar cane with both hands even while sitting in a tree. This pattern of behavior developed without training, in response to human throwing to chimpanzees. Category 8. Catch with one hand (CAT) (Type A): Seize a flying insect such as a fly by a quick sideways jerking movement of a hand, then open the palm to inspect the insect. Seen also in bonobos (Kano, unpublished). "Grab" is used for other contexts. Category 2. Charge (CHG) (Type C): Goodall (1989): "A fast run directed toward another individual." The bonobo's charge is accompanied by stamping, and sometimes by branch-dragging (Kano, 1998). Category 2. Charging display: See Display, charging. Chase (CHA) (Type C): Goodall's (1989) Chase. Run after a fleeing individual, in order to grab it in aggression, play or hunt. Usually the chaser is dominant to the chased, but this is not always the case. Goodall (1989) commented that "sometimes, a subordinate individual, screaming or perhaps uttering waa-barks, chases after his/her aggressor as the latter charges away" (see Aggress morally). During play, two or more individuals may chase round and round a tree or another individual. The bonobo's chase is more ritualized, as the chaser often gets ahead of the fleeing individual (Kano, 1998). Kano (1998) refers to "Chase play" for chase in the play context. Category 1. Chew (CHE) (Type A): Plooij's (1984) Chew (CHE). Press repeatedly between upper and lower teeth. Also occur in bonobos. Category 1. Choke in tantrum (CHT) (Type A): Goodall's (1989) Glottal cramps: "During loud and prolonged tantrum screaming, the chimpanzee may seem to choke and only hoarse squeaks or rasping sounds can be heard." Absent in bonobos? Category 3-5. Clack teeth (CTE) (Type A): Goodall's (1989) 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." Not seen in the bonobos of Wamba (Kano, 1998). Category 3-5. Clasp self (CLS) (Type A): Goodall's (1989) Self-clasp: "Sit or lie with arms tightly clasped around self..." Observed at Gombe as "in a relaxed manner during resting." Not seen at Mahale? Not seen in the bonobo of Wamba (Kano, 1998). However, infants of both species of Pan captured by humans show this gesture. Category 1-2? Climb (CLI) (Type D): Plooij's (1984) Climb (CLI). Includes not only CLV and CLE, but also chimpanzees climb by walking quadrupedal palmigrade on a slanting trunk or bough of a big tree (same as WQP). Goodall's (1989) Vertical climb ("The chimp climbs up a trunk, branch, pole, or other vertical structure") and Kano's (1998) "Vertical climb up" for the bonobo probably includes both CLV and CLE. Category 1. 148 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M.

Climb down: See Descend. Climb vertical (CLV) (Type A): Hunt and others' (1996) "Flexed-elbow vertical climb". Climb a woody vine or small tree by alternately using palmar surfaces of four limbs. See Figure 3 of Nishida (1983a). Category 1. Climb vertical, extended elbow (CLE) (Type A): Hunt and others' (1996) "Extended-elbow vertical climbing". Extend arms around trunk of a huge tree and propel upwards by the simultaneous kicking or walking movements of feet. Category 1. Cling (CLN) (Type A): Goodall (1989): "Gripping a tree trunk or branch with hands or hands and feet. An infant clings to mother's belly, back, arm or leg..." Also seen in bonobos (Kano, unpublished). See Photograph 5 in Nishida (1968). Category 2. Clip leaf (CLL) (Type B): Nishida's (1980b) Leaf clip: Pull a leaf repeatedly between lips or teeth with one hand, producing a conspicuous sound that attracts attention from the prospective sex partner. One of the most common courtship display patterns in Mahale chimpanzees. Also occurs as an act of frustration or play at Bossou (Sugiyama, 1981) and as a prelude to male drumming behavior or an act of frustration at Tai (Boesch, 1995). Thus, the contexts in which this occurs differ from place to place. Moreover, the detail of the behavior pattern differs between Mahale and Tai. See Figure 38-2 of Nishida (1987) for drawing. Category 7. Close eyes (EYC) (Type A): Chimpanzees close eyes: (1) in sleep, (2) as a reflex when they are frightened, and (3) when they seem to be in bliss. An adult male (Alofu) lies on the ground with eyes closed apparently in bliss, after stuffing a great amount of sweet fruit in his mouth. Plooij's (1984) Eyes Closed (EYC). Category 1. Club (CLB) (Type A): Strike target with a long (usually more than 1m), thick stick in overthrow movement. Van Hooff (1973) and Kortlandt and Kooij (1963) described this pattern. Seen only rarely and displayed by a few individuals at Mahale. Absent in bonobos (Kano, unpublished). Category 1?. Club another (CLA) (Type A): Club another individual with a stick. Adult male (Toshibo) twice hit another adult male with a stout stick after flailing it. Category 8. Club ground (CLG) (Type A): Strike the ground with a long, thick stick to solicit copulation, seen in two juvenile males. Adult male (Fanana) clubs the ground in order to entice an estrous female to follow him during consortship. Juvenile male clubs the ground to threaten another juvenile during play. Several adult males (Lukaja, Fanana, Dogura) club the ground or wall of house as a component of charging display. Category 8? Compress lips (COM) (Type A): Goodall's (1989) 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." Plooij's (1984) Compressed lips (COM). Absent in bonobos. Category 3. Console (CSL) (Type D): Third party touches, grooms or licks a victim attacked and wounded by another. See Lick wound. Kano's (1998) "Conciliating behavior" by the bonobo. Category 1. Consort (CST) (Type D): Goodall's (1989) 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." McGinnis's (1973) consortship, Tutin's (1974) Safari behavior. Consortship has been deduced for the chimpanzees of Tai from a genetic study (Gagneux and others 1999). Not seen in the bonobos of Wamba (Kano, 1998). Category 3. Coprophagy: See Eat feces. Copulate (COP) (Type A): Intromission and pelvic thrusting between male and estrous female. Goodall's (1989) Copulation, Plooij's (1984) Copulate.Category 1. Copulate between group (CPG) (Type D): Young, and less often prime, estrous females may visit a neighboring unit group and mate at Gombe (Goodall, 1986), Mahale (Nishida and others, 1985) and Tai (Gagneux and others, 1999). Bonobo unit groups occasionally mingle peacefully, and mating between members of different groups occurs at Wamba (Idani, 1990) and Lomako (Fruth , cited in de Waal and Lanting, 1997). Adult males do not interfere with such copulations (Kano, 1998). Category 1. Copulate dorso-ventral (CDV) (Type A): Male mounts the rear of the female. Major copulating pattern among mature males and females of both chimpanzees and bonobos. See Figure 11-1 of McGrew and others (1996) for photograph. Category 1. Copulate ventro-ventral (CVV) (Type A): Male mounts the front of the female. Occurs rarely between adolescent females and immature male chimpanzees. More common between adolescent females and mature and immature male bonobos (Kano, 1998). Category 1. Ethogram of chimpanzees 149

Copulatory dart: See Dart. Copulatory squeal: See Squeal in copulation. Cough (COH) (Type A): Goodall (1989): "When chimps have colds and coughs they sneeze and cough, sounding like humans." Bonobos also cough (Kano, unpublished). Category 1. Cough-threat (COT) (Type A): Goodall (1989): "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 uttered by bonobos (Kano, unpublished). Category 2. Counter-attack: See Retaliate. Courtship display: See Solicit copulation. Cover breast: See Cover nipple. Cover nipple (CON) (Type A): Plooij's (1984) Cover nipple (CON). Goodall's (1989) 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 mother's weaning of her infant. Not recorded for the bonobos of Wamba (Kano, 1998). Category 3. Cradle with hand (CDH) (Type A): Mother supports her newborn infant with both arms. Goodall's (1989) Cradle. Kano's (1998) Cradle. for the bonobo of Wamba. Category 1. Cradle with leg (CDL) (Type A): Mother hanging from a tree branch supports her infant clinging to her belly with one thigh flexed so that the baby's body is pressed against her. Plooij's (1984) Cradle (CRA). Seen in bonobos (Kano, unpublished). Category 2. Creep (CRE) (Type D): Walk stealthily. Goodall (1989) 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 a similar pattern (Kano, 1998). Category 1-2? Cross (CRS) (Type B): Flex one or both limbs to rest on opposite limb. Category 1. Cross arms (CRA) (Type A): Flex arms across chest. Category 1. Cross legs (CRL) (Type A): Flex one leg to rest on opposite leg, which is supported on substrate. Category 1. Crouch (CRO) (Type A): Van Hooff's (1973) and Plooij's (1984) Crouch (CRO). Goodall's (1989) 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..." Kano's (1998) Crouch and Present in agonistic interaction for the bonobo. Category 2. Crouch-present: See Present with limbs flexed. Crowd (CRW) (Type D): Many chimpanzees gather to peer at an individual or gaze at an object. Occurs when a chimpanzee tries to pick out a thorn from its foot, inspects its seriously damaged skin, carries a baby that has just died, has precious food such as meat, or an unusual object in the environment that stimulates chimpanzees' curiosity. When crowded, chimpanzees appear to allow closer contact with each other than usual. No adult bonobos crowd (Kano, unpublished). Category 1? Crush (CRH) (Type A): Press louse between finger-tips of both hands. Absent in bonobos. Category 8? Crutch (CRU) (Type A): Goodall (1989): "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..." Plooij's (1984) crutch (CRU). At Mahale, also occurs when a mother carries her newborn on her belly, accompanied by . In bonobos, crutch is used only as the last step of the long walk or as a one-step movement while eating or resting (Kano, 1998). See Figures 2c and 2d of Goodall (1968) for illustration. Category 2. Cry: See Whimper-Scream. Cushion: See Make a cushion.

D Dab (DAB) (Type A): Goodall (1989): "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." At Mahale, seen only in a few adolescents and juveniles. Absent in the bonobos of Wamba (Kano, 1998). Category 8? Dance bipedal (DCB) (Type B): Nishida's (1997) Bipedal sex dance: Stand on one's feet while raising the 150 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. arms or performing the branch-shaking display. Only one adult female (Wakiluhya) of M group showed this type of courtship, but de Waal (1982) mentioned similar pattern. Similar pattern displayed by the bonobos of Wamba. Category 8. Dangle (DAN) (Type A): Plooij's (1984) Dangle (DAN). Goodall (1989): "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..." Seen in the bonobos of Wamba (Kano, 1998). See Figures 19 a, and 19b of Goodall (1968). Category 2. Dart (DAR) (Type D): Estrous female runs quickly a few meters after copulation. Often accompanied by copulatory squeal and grin, then, the female often lies down for a few minutes. Nishida's (1997) Copulatory dart. Absent in bonobos (Kano, unpublished). Category 3-5. Deceive (DCV) (Type D): (1) mislead other individuals for one's own advantage by not showing posture, gesture, or facial expression usually done in that context, (2) mislead other individuals for one's own advantage by expressing posture, gesture, or facial expression that is irrelevant in that context (Nishida, 1990, 1998). Reported from Gombe (Goodall, 1971) and captivity (de Waal, 1982). Likely to be Category 1. Depart (DPT) (Type C): Begin to move. From the rest to travel phase. Category 1. Descend (DSC) (Type D): Climb down from an elevated site: Includes 5 patterns: DBR, DQF, DQH , Leap down and Slide down. Category 1. Descend by brachiating (DBR) (Type A): While the chimpanzee progresses by brachiation, the bough gradually bends down until the ground can be reached with the feet. Known for the bonobo (Kano, 1998). Likely to be Category 1. Descend quadrupedal feet first (DQF) (Type A): Climb down backwards. Seen in bonobos (Kano , unpublished). Category 1. Descend quadrupedal head first (DQH) (Type A): Climb down forwards. Seen in bonobos (Kano, unpublished). Category 2. Detach (DTC) (Type D): Nishida (1983a): Take infant from its mother. Keeps the infant under the alloparent's control for as long as the alloparent wants, since it is hard for the infant to return to the mother on its own if it is taken away far from the mother. Goodall's (1989) Kidnap: "(Borrow by consent) When an infant is carried away from the mother by a sibling or by another chimp." See Figures 2 and 3 of Nishida (1983a) for photograph. Known for the bonobos of Wamba (Kano, 1998). Category 1-2. Detour (DET) (Type C): Deliberately take a roundabout route. Occurs when a chimpanzee wants to avoid a dominant rival, or when a chimpanzee finds the shortest route to be difficult, for example, gap in a tree or crossing a river. Kano's (1998) Detour and Pass with detour for the bonobos of Wamba . Category 1. Dig with hand (DGH) (Type A): Dig into wet ground of a dry stream with hand to get drinking water. More than 4 individuals of M group did this. The bonobos of Lilungu and Wamba dig for earthworms and mushrooms, not for water (Bermejo and others, 1994; Kano, 1998). See also Remove leaves. Category 8. Dig with stick (DGS) (Type A): Dig with tool wet ground of a dry stream to obtain water . Only one juvenile female was seen to do this. Absent in the bonobos of Wamba (Kano, unpublished). Category 8. Display away (DSA) (Type C): Goodall (1989): "After interaction with another, usually an attack , the chimp departs in a charging display." Seen in bonobos (Kano , unpublished). Category 2? Display, charging (CHD) (Type D): Plooij's (1984) charging display (CHD) . Goodall (1989): "The chimp may move in a slow rhythmic" cantering gait, "run at a moderate speed or very fast. Display patterns include , , , , , , , , , and occasionally ... The charging display may also incorporate , or . It can be bipedal , quadrupedal, or tripedal. This is typically a male display though females do it also." Goodall (1989) also discriminates Non-vocal and Vocal displays. The former, "when the displayer makes no calls, is most likely to be directed toward another individual, and may be followed by an attack. It is a typical component of male dominance rivalry." The latter "is accompanied by pant hoots, and typically is not directed toward any other chimpanzee. It is common during reunions, food excitement and when an individual has been frustrated in obtaining a desired goal..." At Mahale, the charging display is never accompanied by , but often by and as well as the behavior above mentioned. The bonobo's charging display has only two patterns: and (Kano, 1998). Category 2. Display past (DSP) (Type D): Goodall (1989): "When a chimp, who is clearly directing a display toward another, passes close, and may hit or kick the other in passing." In 1995 at Mahale , two adult males Ethogram of chimpanzees 151 regularly hit human observers during their charging displays. Seen in bonobos. Category 8? Display, rain (DSR) (Type D): Goodall's (1989) Rain display or Rain dance. At Mahale, two types of displays are associated with the onset or during heavy rain or storms. One is a noisy, active display accompanied by , , , . and so on. Another is prolonged, slow tempo displays, of which the main components are , , , and . Although at the start of storms or heavy rain, several adult male bonobos may run a few meters while calling loudly, no other activities such as those of chimpanzees occur (Kano, 1998). Category 2? Display, streambed (DSS) (Type D): Goodall's (1989) Streambed displays are like rain displays that occur in a streambed. Goodall's (1989) Waterfall display should probably be included here. At Mahale, the main components of the streambed display are , , , and . Absent in bonobos (Kano, unpublished). Category 3-5. Display toward (DST) (Type D): Goodall (1989): "Display directed toward one or more others." Seen in bonobos (Kano, unpublished). Category 2. Distance scratch: See Scratch self distantly. Distract (DTR) (Type D): Goodall's (1989) Distract. Mother grooms infant to divert its attention from sucking from nipples during the weaning period. If the infant is male and he throws temper tantrums in response to his mother's refusal, she sometimes allows him to copulate with her. The bonobos of Wamba display the same behavior pattern (Kano, 1998). Category 1-2. Drag (DRA) (Type B): Goodall (1989): Drag along the ground another individual, branch, palm frond, large food object. Shown in aggression, threat, play, and maternal solicitude. Plooij's (1984) Drag (DRA). The bonobos of Wamba drag only as branch-dragging (Kano, 1998). Category 1. Drag branch (DRB) (Type A): Goodall's (1989) Branch dragging: "A branch broken from a tree, or one lying on the ground is dragged by one hand during a charging display..." Widely known across populations from east to central Africa (Kuroda, 1998), to west Africa (Sugiyama, 1998). Among the bonobos of Wamba, branch-dragging displays (Ingmanson, 1996; Kano, 1998) consist of many more behavioral elements than those of chimpanzees. Late adolescent male bonobos use this display in order to tease and surpass older males in rank. Bonobos sometimes drag a branch while walking without stamping or vocalizing. See Kano (1990, p.62) for photograph. Category 2. Drag other (DRS) (Type A): Goodall's (1989) Drag other. Drag by arm and leg of another individual. Occurs during aggression and play. Mother often drags her older infant when the latter does not agree to go with her. Kano's (1998) Drag other for the bonobo. Category 1. Drag to kill (DRK) (Type A): Adult male chimpanzee (Musa) once ran and dragged with one hand a large red colobus monkey that he had captured, so that the monkey was hit and injured by woody vegetation and choked by woody vines. Tactic prevented the monkey from biting the chimpanzee predator and eventually killed thevictim. Category 8? Drape (DRP) (Type A): Goodall (1989): "Bunch of fruit or leaves, piece of cloth, and so on. is draped over shoulders, neck - or even over the head. Thus adorned, the chimp performs other patterns of lone play." At Mahale, a juvenile female (Tula) once draped cardboard and a cooking pan over her shoulder and head. Adult female (Ako) once draped "necklace" -like colobus skin around her neck (McGrew and Marchant, 1998). Category 8? Drink (DRI) (Type A): Goodall (1989): "Drinking of water or other liquids. This includes drinking directly by leaning over and sucking from the water source; dripping water from the fingers into the mouth; licking drops of water from surfaces such as leaves, hair, and so on..." Chimpanzees sometimes keep water in the mouth, and transport it for up to 10 m, without drinking it. See Figure 11 of Nishida and Uehara (1983) for photograph. Bonobos rarely drink from running or standing water (Kano, 1998). Category 1. Drop (DRO) (Type B): Let something fall just above another individual or human observer . Category 2. Drop branch (DPB) (Type A): Drop a branch from above the ground to threaten human observer (occasionally another conspecific) below, so that sometimes the target is hit. Some adult males forcibly kick dead branches, and thus detach the branches so that they fall. Occasionally observed at Mahale when chimpanzees were not habituated well. Reported for a female chimpanzee of Gabon (Takenoshita and others, 1998). Ingmanson (1996) describes "Object dropping" for the bonobos of Wamba , which includes a branch. Category 2. Drum (DRM) (Type B): Component of the charging display. Goodall (1989): "Hit and/or kick tree trunks, especially those with buttresses. Often they grab the buttress with their hands and stamp on it with their 152 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M.

feet. They may kick backwards with their feet or hit the trunk or buttress with their hands. Often a particular" drumming tree "triggers a drumming display and the individuals of a traveling group are likely to drum one after the other as they pass. Usually drumming is accompanied by the pant-hoot call, but sometimes...drumming is without vocalization. Chimps at Gombe also pound on the walls of research buildings and on a barrel placed at the feeding area..." Mahale's chimpanzees do the same. Drumming patterns vary from one individual to another. For example, an individual male who is standing erect beats the walls of research buildings with only one palm, another does so with two palms, and the third one with one fist, while the fourth one kicks with one foot and the fifth does so with both feet. Drumming reported from Kibale and Tai (Arcadi and others, 1998). Bonobos also drum on buttresses, most often immediately after the onset of traveling (Kano, 1998). See Figure 3 of Nishida (1994) for photograph. Category 2. Drum belly (DMB) (Type A): Idiosyncratic performance shown by only one juvenile male. Five-year-old Cadmus slapped his belly with his right hand to make drum-like sounds, while hanging from a branch by his left arm. There were no other chimpanzees nearby. Perhaps this was play or was in response to the approach of a human observer. Absent in the bonobos of Wamba (Kano, unpublished). Category 8. Dunk (DUN) (Type B): Put an object into water without releasing it. Absent in the bonobos. Category 8. Dunk colobus skin (DUC) (Type A): Adult male (Musa) once dunked a red colobus skin into a stream apparently in order to clean it. See Wash. Category 8. Dunk face (DUF) (Type A): Juveniles dunk their faces into water repeatedly in playin the river. Category 8.

Eat (EAT) (Type B): Remove foodstuff (leaves and so on.) from the substrate , process, put into mouth, bite and chew, wadge and swallow it. Goodall's (1989) Feed. Kano's (1998) Feed. Category 1. Eat ants (EAA) (Type B): Eat Crematogaster ants by cracking a grass stem or dry branch with teeth (See Nishida and Uehara, 1983 for photograph), breaking off a large log with hands and occasionally hands and feet. Eating carpenter ants (Camponotus spp.) by cracking a grass stem or by using tools (fishing for ants). A nest of weaver ants (Oecophylla longinoda) is grabbed with one hand, and often peeled on the ground, with the ants being quickly eaten. Ants are eaten everywhere, but the species eaten vary from place to place (McGrew, 1992). The bonobos of Wamba do not eat ants, but those of Lilungu eat adult and larvae of Tetraponera (Bermejo and others, 1994). Humans in the tropical region eat ants (Bodenheimer, 1951). Category 1? Eat blossom (EAB) (Type B): Eating techniques resemble those for eating leaves. Eaten by bonobos (Kano, 1992). Category 1. Eat carcass (EAX) (Type B): Scavenge. Eat the carcass of mammals that have not been killed by oneself or one's companions. Scavenging has been seen at Gombe (Muller and others, 1995) and Mahale (Hasegawa and others, 1983; Nishida, 1994). Not recorded for the bonobo of Wamba. Category 1? Eat conspecific (EAC) (Type B): Cannibalism. Chimpanzees of Mahale occasionally eat infants (Hamai and others, 1992). Known also from Budongo (Suzuki, 1971), Gombe (Bygott , 1972; Goodall, 1986), Kibale (Mitani, unpublished) and Tai (Boesch and Boesch, 1999). See photographs 1-4 of Kawanaka (1981). Not recorded for the bonobo (Kano, 1992). Category 3. Eat feces (EAD) (Type B): Goodall's (1989) Coprophagy. At Mahale, eating one's feces has been seen only rarely. Adult female (Chausiku) picked out and ate undigested seeds of Saba florida from her feces. Gombe apes picked undigested meat from feces (Wrangham, 1977); Assirik chimpanzees picked out baobab seeds from feces (McGrew and others, 1988). Not observed for the bonobos of Wamba (Kano, 1998). Category 8. Eat fruit (EAF) (Type B): Crop fruits in a tree, from shrubs on the ground, or pick up fallen fruits from the ground, then, open the fruit shell with hands or teeth, and ingest the pulp. See Figure 4-1 of McGrew and others (1996) for photograph. Category 1. Eat honey (EHO) (Type B): Put stick into honeycomb and eat honey in the case of honeybees (Apis), and bite wide open the entrance of the nest with incisors in the case of stingless bees ( Trigona). Honey eating is widely known in the chimpanzee range (Tuttle, 1986). The bonobos of Lilungu eat honey of stingless bees (Bermejo and others, 1994). Category 1. Eat leaves (EAL) (Type B): Put leaves into the mouth and ingest them. Chimpanzee may bring mouth to the leaves or bring cropped leaves into the mouth. Important cropping technique is . Ethogram of chimpanzees 153

Category 1. Eat meat (EAM) (Type B): Meat eating consists of the ingestion of bones, teeth, bone marrow, brain, intestines, stomach and intestinal contents, and skin, in addition to muscles. Eating meat is unusually prolonged, sometimes for more than 3 hr. Prey includes red colobus, blue, red-tailed, and vervet monkeys, greater galago, infant chimpanzees (see Eat conspecific), blue duiker, bushbuck, bushpig, giant rat, rock hyrax, civet, francolin, guinea fowl, and weaver bird. Meat-eating has been confirmed throughout the chimpanzee range (Uehara, 1997). See Figure 9-2 of McGrew and others (1996) for photograph. Bonobos have also been recorded to eat meat in Wamba (Kano, 1992), Lomako (Hohman and Fruth, 1993) and Lilungu (Bermejo and others, 1994), but in much less often and extensively (Kano, 1998). See also Hunt. Category 1. Eat phloem (EAH) (Type B): Eat phloem or inner bark rich in sugar and protein by scraping outer bark with incisors. See Figure 1 of Nishida (1976) for photograph. Eaten widely by humans (Nishida, 1976). Category 1? Eat pith (EAP) (Type B): Eat soft, juicy piths of herbs such as elephant grass, gingers, ginger lilies and Marantochloa, and woody vines such as Landolphia and Saba by removing outer surface with teeth. See Figure 11.1 of Uehara (1990). Pith of Megaphrynium eaten by both bonobos and humans at Wamba (Kano, 1992). Category 1. Eat resin (EAR) (Type B): Scrape the resin of Terminalia mollis with frontal teeth or picking the large lump of the resin of Albizia glaberrima. See Figure 6 of Nishida and Uehara (1983) for photograph. Resin eaten by bonobos (Kano, unpublished). Category 1. Eat seeds (EAS) (Type B): Crack seed pod with teeth, and chew and ingest the seeds. Eaten by bonobos (Kano, 1992). Category 1-2. Eat termites (ETT) (Type B): Push over the tower of termite mounds with hands and pick up soldiers or winged reproductive forms of the termites of Pseudacanthotermes. See Figure X of Uehara (1982) for photograph. Fishing for soldiers of Macrotermes has been observed in the chimpanzees of B group (Nishida and Uehara, 1980, McGrew and Collins, 1985). Widely known throughout the chimpanzee range (Tuttle, 1986). The bonobos of Lilungu eat termites of different genera from those eaten by chimpanzees by introducing fingers into the hole that they had opened manually (Bermejo and others, 1994). Termites are widely eaten bytropical people (Bodenheimer, 1951). Category 1. Eat termite soil (ETS) (Type B): Pick up and ingest small pieces of the soil from termite towers of Pseudacanthotermes. Termite soil also eaten by the bonobo (Kano, 1998). Category 1. Eat wood (EAW) (Type B): Eat dry dead wood of, for example, Pycnanthus angolensis. See Nishida and Uehara (1983) for photograph. Not known in other localities. Category 6. Ejaculate (EJA) (Type A): Ejection of seminal fluid from penis. Males begin to ejaculate at 9 years old at Mahale. Semen is often eaten by both sexes. Category 1. Embrace full (EMF) (Type A): Goodall's (1989) Ventro-ventral embrace: "Two individuals face each other and each puts one or both arms around the other." Van Hooff's (1973) Embrace, Nishida's (1983a) Embrace, Plooij's (1984) Embrace full (EMF). Embrace full by adult males may be accompanied by . In the bonobo, the embrace full posture is seen only in genito-genital rubbing by females and ventro-ventral copulation between males and females (Kano, 1998), and so both embrace full in erect posture and embrace-full by adult males are lacking. Category 3? Embrace half (EMH) (Type A): Plooij's (1984) Embrace half (EMH). Goodall's (1989) Arm round: "Chimp puts one arm around another as in a half embrace ." Often occurs when one individual pant-grunts to another. An adult male may put his arm around another (usually dominant) male while walking with pant hoots. Seen in copulatory solicitation in bonobos (Kano, unpublished). Likely to be Category 1. Erect hair: See Bristle. Erect penis (PEN) (Type A): Plooij's (1984) PEN. Goodall's (1989) Penile erection. Occurs during sexual and food excitement. Kano's (1998) Penile erection for the bonobo. Category 1. Extend (EXT) (Type B): Extend arm or leg to another individual. Category 1. Extend arm as ladder (EXA) (Type C): Nishida (1983a): Alloparent extends arm up to the mother above in a tree, in order to allow her infant to climb up easily by making a ladder with her arms. See Figure 8 of Nishida (1983a) for photograph. Absent in bonobos. Category 8? Extend hand (EXH) (Type B): Nishida's (1983a) Reach hand. Goodall's (1989) Extend hand. Van Hooff's (1973) Hold out hand: "A variety of forms were observed. Most common is a form in which the actor, either sitting or standing, extends its arm roughly horizontally towards a fellow. The arm is in a position 154 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. about midway between pronation and supination (that is with the thumb up). The hand may be bent at the wrist so that its back is turned to the partner with the fingers bent or fully stretched..." Goodall's (1989) "Extend hand" is limited to the posture with wrist and fingers extended and palm up or down. Goodall (1989) pointed out that the behavioral pattern is seen during begging, as a reassurance gesture towards a subordinate who is out of reach, and when a submissive chimpanzee seeks reassurance after aggression. In mother-infant interactions, extend hand is a contact-seeking behavior. For the bonobo, extend hand is only seen in the form of EAD or EAU below, and in the context of food begging and mother seeking contact with her offspring (Kano, 1998). Category 1. Extend hand and hold tree (EHT) (Type A): Reach one arm forward and grasp the trunk of a tree. This is often performed to solicit being groomed. Seen in bonobos. Category 2. Extend hand and put knuckle on ground (EKG) (Type A): Reach one arm forward and put knuckle on the ground instead of holding the tree trunk. Often performed to solicit being groomed as in EHT . Seen in bonobos (Kano, unpublished). Category 2. Extend hand, palm downward (EXD) (Type A): Van Hooff's (1973) Stretch over, Plooij's (1984) Extend hand, palm downward (EHD). Goodall's (1989) Arm stretch: "The arm or arms are extended toward another. The palm of the hand usually faces down. Seen in courtship or greeting." Nishida's (1997) Arm-stretch. See Figure 4 of Hiraiwa-Hasegawa (1989). Seen in bonobos (Kano, unpublished). Category 2? Extend hand, palm upward (EXU) (Type A): Plooij's (1984) Extend hand, palm upward (EHU) . Often used in begging. Seen in bonobos (Kano, 1998). Category 1. Extend leg (EXL) (Type A): Reach leg instead of hand to another individual, in similar function to REH. This is particularly shown when performing REH is difficult, for example , in a tree or when the performer holds an infant and food. Seen in bonobos. Category 2. Extended grunt: See Grunt, extended. Eyes closed: See Close eyes. Eyes open: See Open eyes.

Face close: See Peer. F Feed: See Eat. Fend (FEN) (Type D): Goodall (1989): "Most often seen when a mother keeps her child away with her hand or foot when she is being pestered for a share of her food, when her infant tries to suckle during weaning, and so on. But adults sometimes fend off individuals who are begging from them." Not limited to food but any case of deflecting reach of another. Bonobos show similar behavior (Kano, 1998). Category 1. Fight (FIG) (Type D): Attack each other. See attack. Category 1. Finger wrestle: See Wrestle with fingers. Fish (FIS) (Type B): Insert a strip of bark, vine, twig, grass and so on. into the nest of ants or termites, withdraw and pick off insects with lips, teeth and tongue. Absent in the bonobos of Wamba . Category 3? Fish for carpenter ants (FIA) (Type B): Insert a fishing probe such as peeled bark, unmodified vine, branch, modified branch, midrib of leaf, or scraped wood into the entrance of wood-boring carpenter ants (Camponotus spp.) and withdraw the probe laden with the soldier ants and lick them off with lips and tongues. Usually an arboreal activity that can continue for more than 3 hours at a time, it is performed throughout the year (Nishida and Hiraiwa, 1982). See Plates 1-6 of Nishida (1973) for photographs . The chimpanzees of Gombe do not fish for carpenter ants, but dip for driver ants (McGrew, 1974). Category 6. Fish for termites (FIT) (Type B): Resembles ant fishing. Fishing for the termites of Pseudacanthotermes spongier has been observed for only K group chimpanzees (Uehara 1982). Fishing for Macrotermes only for B group chimpanzees (Nishida and Uehara, 1980; Uehara, 1982; McGrew and Collins, 1985). Fishing for termites has not been seen for M group chimpanzees apparently because of the lack of Macrotermes species. Recorded from both west (McGrew and others, 1979; Humle, 1999) and central (Sugiyama, 1985; Suzuki and others, 1995, McGrew and Rogers, 1983) Africa in addition to Gombe. Not recorded for bonobos. African Bantu people fish for termites with more elaborate techniques (Bodenheimer , 1951). Category 1? Fixed stare: See Stare fixedly. Flail (FLL) (Type A): Goodall (1989): Brandish a branch in one hand and "waves this weapon at an opponent." Plooij's (1984) FLL. Kortlandt's (1967) Tree-swish. At Mahale flailing against another Ethogram of chimpanzees 155

individual is rarely done by an adult male. Flailing against a human observer is more rarely done by an adolescent male and female. Branch-flailing appears as an element of branch-dragging among the bonobos of Wamba (Kano, 1998). Category 1-2. Flap (FLP) (Type A): Goodall (1989): "...downward slapping movement of the hand , usually repeated several times, in the direction of another individual. Often seen in female squabbles..." Not recorded for the bonobos of Wamba (Kano, 1998). Category 3-5. Flee (FLE) (Type C): Goodall's (1989) Flight (Run away): "A rapid progression away from an alarming or dangerous stimulus..." Elements include , , , , and . Van Hooff's (1973) Flight. Kano's (1998) Flee. Category 1. Flight: See Flee. Flip lip (FLI) (Type A): Goodall's (1989) Lip flip: "The upper lip is rolled up and back towards the nose. Usually relaxed situation..." Plooij's (1984) FLI. The function of this facial expression is unknown. Not seen in bonobos (Kano,unpublished). Category 3-5. Follow (FOL) (Type C): Walk after, trying to maintain close proximity (but not physical contact) to another individual. Goodall's (1989) Follow. Plooij's (1984) Follow. See Figure 6 of Nishida (1994) for photo. Kano's (1998) Follow. Category 1. Follow specific female (FSF) (Type C): Newly-immigrated females tend to choose, follow and form affiliative relationship with, one particular resident female by grooming her and caring for her infant (Nsihida, 1989). Immigrant female bonobos do the same (Furuichi, 1989; Idani, 1991). Category 2. Food grunt: See Grunt, food. Food scream: See Scream, food. Fumble with clitoris (FWC) (Type A): An adult or adolescent female repeatedly touches own clitoris. Rarely seen. Category 1-5. Fumble with nipple (FWN) (Type A): Self-reassurance or self-stimulation performed by a few chimpanzees (one male and 3 females in 1997), by manipulation of the nipples with thumb and fingers. Occurs when they are frightened or become nervous in the proximity of dominant individuals. Usually thumb only or thumb and index finger are used. One male, Alofu, alternates the right and left nipple very skillfully during anxiety and pant-grunting. When using the right hand, he fumbles the right nipple with his thumb and the left nipple with his little finger simultaneously, and when he uses the left hand, he fumbles the left nipple with his thumb and the right nipple with the little finger. Thus, the nipples are massaged by the circular movement of the thumb and finger. A captive adolescent male bonobo did the same as an act of self-reassurance (deWaal and Lanting, 1997). Category 8. Fumble with penis (FWP) (Type A): Male occasionally fumbles with his penis, which leads to penile erection. At Mahale, this never culminates into ejaculation. For the bonobo of Wamba, there was only one case observed of fumbling with penis, and it did not lead to ejaculation (Kano, 1998). Category 8?

GGallop (GLP) (Type A): Goodall (1989): "Fastest run of the chimps." Van Hooff's (1973) Gallop. See Figure 7b of Goodall (1968) for illustration. Category 2. Gang attack: See Attack. Give (GIV) (Type C): Hand over food by the owner to another who may or may not have been begging. Rare at Mahale. This has never been observed among the bonobos of Wamba (Kano, 1998). Category 8? Glance (GLN) (Type A): Goodall (1989): "The chimp looks toward another for less than 2 seconds." Subordinate chimpanzee may watch only briefly food, estrous female or other object of competition when there is a dominant male. Kano's (1998) Glance for the bonobo. Category 1. Glottal cramps: See Choke in tantrum. Grab (GRA) (Type A): Goodall (1989): "When one individual roughly seizes another with one or both hands. This is an aggressive gesture. It is usually followed by a sequence of aggressive interactions, -and may escalate into fight." Grab occurs not only in aggression, but also in grooming and play. Bonobos grab in the same context (Kano, 1998). Category 1. Grapple (GRP) (Type A): Goodall (1989): "When two individuals go into a clinch with arms locked and heads bowed, and sometimes roll over and over. Individuals may be fighting or playing. Not uncommon when adult females fight one another." Bonobos do the same (Kano, 1998). Category 1. Grasp hand (GSH) (Type A): A dominant individual grasps the hand extended to him by a subordinate for reassurance, but usually, there is no shaking motion. Absent in bonobos of Wamba (Kano, 156 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. unpublished). Category 3-5? Greet (GRE) (Type D): Elements of friendly reunion that are the same as Goodall's (1989) Greeting Behavior: "Typical friendly behavior includes , , , , , , , , , , and so on. Vocalization associated with greeting behavior are , , , , , and so on." In bonobos, genital contact behavior and may be comparable to greeting, in addition to , , , , and , but the other GRE elements described above are lacking (Kano, 1998). Category 1-2. Grimace: See Grin. Grin (GIN) (Type B): Goodall (1989): Facial expression with "the corners of lips are drawn back, exposing the lower, or both upper and lower teeth (or teeth and gums). Category 2. Grin-full-closed (GFC) (Type A): Goodall's (1989) Full closed grin: "Both upper and lower teeth (and gums) are revealed by the horizontal retraction of upper and lower lips, and the teeth are closed, or almost closed". Kano's (1998) Grin for the bonobo of Wamba. Category 2. Grin-full-open (GFO) (Type A): Goodall's (1989) Full open grin: "Both upper and lower teeth (and gums) are revealed by horizontal retraction of upper and lower lips, and the teeth are partially or widely open ." See Figure 3 of Nishida (1970) for photograph. Seen for the bonobo at Wamba (Kano, 1998). Category2. Grin-low-closed (GLC) (Type A): Goodall's (1989) Low closed grin: "Only the lower teeth (sometimes with gums) are revealed, and the teeth are closed, or almost closed." See Figure 7 of Nishida (1970) for photo. Absent in bonobos? Category 3? Grin-low-open (GLO) (Type A): Goodall's (1989) Low open grin: "Only the lower teeth (sometimes with gums) are revealed, and the teeth are partially or widely open." See Figure 6 of Nishida (1994) for photograph. Absent in bonobos? Category 3? Groin pocket: See Transport in groin pocket. Groom (GRM) (Type B): Goodall's (1989) Grooming behavior: "...use both hands, pushing the hair back with the thumb or index finger of one hand and holding it back while picking at the exposed skin with the nail of the thumb or index finger of the other. The chimpanzee can also use one hand , parting the hair in the same way and holding it back with the lower lip". Grooming may occur in bipedal,quadrupedal , sitting, or lying posture. Function of grooming includes appeasement, reassurance and reconciliation in addition to elimination of ectoparasites. See Figure 6 of Nishida (1983a) for photograph. Category 1. Groom ground (GRG) (Type A): Groom the soil, leaves on the ground, or moss on a rock, usually with one hand. Solitary or social grooming, even during the grooming-hand-clasp. See Figure 5 of Nishida (1994) for photo. Seen in only two young adult males of Mahale in 1992, but done also by 3 adult or adolescent females in 1995. Category 8. Groom-hand-clasp (GHC) (Type A): McGrew and Tutin's (1978) Grooming-hand-clasp. Pattern of mutual grooming seen at Mahale, Kibale, Lope, Budongo and Tai, but not at Gombe or Bossou. In captivity, the same gesture done by 7 chimpanzees at Yerkes Primate Research Center (de Waal and Seres, 1997). Similar pattern is seen for the bonobo of Wamba (Kano, 1998). Category 6. Groom-hand-clasp unilaterally (GHU) (Type A): One grooming partner holds hands with the partner in a typical hand clasp posture, but does not actually groom companion; instead rests or just grooms the ground, although he is being groomed. See Figure 5 of Nishida (1994) for photograph. Category 8. Groom in mutual arm raise (GNH) (Type A): Mutually groom like GRH, but hands not engaged in grooming are not clasped, but hold branches. This hand posture is seen at Gombe as well as at Mahale. See Figure X of McGrew and Tutin (1978) for photograph. Category 3-5. Groom leaf (GRL) (Type B): Goodall's (1989) leaf-grooming: "A leaf may be suddenly picked, seemingly at random. Holding the leaves in one hand, the chimp makes grooming movements with both thumbs, often pushing his lower lip to the leaf surface as in social grooming. Other chimps often crowd around to look. After a few minutes the leaves are discarded... It seems to be a form of displacement or redirection." Function is unclear. Not seen at Tai and Bossou, nor in the repertoire of the bonobo of Wamba (Kano, 1998). Category 4 or 5. Groom mutually (GMU) (Type A): Goodall's (1989) Mutual grooming. Two chimpanzees groom each other simultaneously. Mutual grooming is common in the west African subspecies (Boesch and Boesch, 1999), but occurs in bonobos only during role changes of social grooming (Kano, 1998). See Figures 8-9 of McGrew and others (1996) for photograph. Category 2-3? Groom reciprocally (GRR) (Type A): Goodall's (1989) Reciprocal grooming. Two chimpanzees groom Ethogram of chimpanzees 157 each other alternately. Most common type of social grooming among adults. Also, common among the west African subspecies (Boesch and Boesch, 1999) and the bonobo (Kano, 1998). Category 1. Groom self (GRS) (Type A): Goodall's (1989) Self groom, Plooij's (1984) GRS, van Hooff's (1973) Autogroom. See Figure 2 of Nishida (1983a) for photograph. Kano's (1998) Self-groom for the bonobo. Category 1. Groom unilaterally (GRU) (Type C): Goodall's (1989) Unilateral grooming. An individual grooms another without reciprocation. See Figure 6 of Nishida (1983a) for photo. Kano's (1998) Unilateral grooming. Category 1. Groom with hand (GRH) (Type A): Groom other with one or both hands. It seems that chimpanzees groom their important companions such as relatives and allies with two hands, and others with one hand and often leisurely. Category 1. Groom with mouth (GMO) (Type A): Pick up something with lips and teeth when grooming, in particular, the face. Similar behavior pattern seen in bonobos (Kano, unpublished). Category 2? Group (GOU) (Type D): Chimpanzees form a social unit called "unit group" or "community" that consists of plural adult males, plural adult females andimmature individuals (Nishida, 1968; Goodall, 1973; Boesch and Boesch, 1999). Adult females usually outnumber adult males. Shows fission and fusion grouping pattern. Inter-group relationship is antagonistic. Bonobos have similar social unit (Kano, 1982). Category 2. Grunt (GNT): See GNA, GNB, GNE, GNF and Pant-grunt. Grunt, Aha (GNA) (Type A): Food grunts uttered loudly when walking hastily in a group towards the known food patch (Goodall, 1986). Probably absent in the bonobos of Wamba (Kano, unpublished). Category 3-5. Grunt, in bed (GNB) (Type A): Goodall's (1989) Nest grunt: "Distinctive double grunt given when an individual begins to look around for a suitable nest site, during nest making, or as he settles down for the night." Not heard in the bonobos of Wamba (Kano, unpublished). Category 3-5. Grunt, extended (GNE) (Type A): Goodall (1989): "A long drawn out sound usually composed of two syllables-ehmmmmm-heard during rest sessions, significance not clear." At Mahale chimpanzees are often "discovered" because of this sound uttered in response to the passing-by or appearance of human beings. Category 3-5. Grunt, food (GNF) (Type A): Uttered during or just before eating food, especially during the first minutes. (Cf. Aha-grunt). Similar grunt occurs in bonobos (Kano, unpublished). Category 2?

Hair erection: See Bristle. H Hand support: See Transport with hand support. Handwrestle: See Wrestle with fingers. Hang (HAN) (Type A): Hang by one or both hands from a tree branch (Cf. Brachiate). Neither foot touches the substratum. Kano's (1998) Suspend for the bonobo.Category 1. Hang in sloth position (HSP) (Type A): Goodall's (1989) Sloth position: "Hanging underneath a branch using all hands and feet or a combination of any three. Usually part of locomotor play in infants." Seen also as a part of self-play by immature chimpanzees at Mahale. The bonobo not only hangs, but also travels in the posture, which Kano (1998) calls "Sloth-walk".See Figure 7a of Goodall (1968). Category 2. Hang-stand (HST) (Type A): Hang from the branch while one or both feet touch the substratum below. Usually assumed as a temporary posture changing to another posture or locomotion, and sometimes as a brief resting posture. Also occurs in bonobos (Kano, unpublished). Category 1-2. Hang tripedal (HAT) (Type A): Hang while gripping the branches with two feet and one hand. The head and trunk is maintained more or less erect. The other hand is used for eating fruits or young leaves at the ends of high branches. Occurs also in bonobos. Category 2. Hang upside-down (HUD) (Type A): Hang with two feet from a branch. Rarely seen in play among infants and juveniles. Kano's (1998) Upside-down suspension for the bonobo. Category 2? Head tip: See Tip head. Heel kicking: See Kick heel. Herd (HER) (Type D): Complex of male behavior to control the travel of a cycling female, so that she follows him. Includes waiting such as , , , and , threatening such as , , , and aggression such as and . Thus, 158 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. herding is all the activities to maintain consortship and possessiveness. Similar behavior is seen in bonobos, but aggressive components (hit and kick) and are lacking. Category 3? Hide (HID) (Type D): Goodall (1989): "Unhabituated chimps may hide behind a tree trunk, or pull a thickly foliated branch in front of them when a human approaches". Similar behavior seen at Mahale (Nishida, unpublished) and Tai (Boesch and Boesch, 1999). Unhabituated bonobos do the same thing (Kano, 1998). (Cf. Detour). Category 1. Hit (HIT) (Type A): Bring down an arm from above and strike the partner with the fist. Van Hooff's (1973) and Goodall's (1989) "Hit" includes the case of striking with the palm, which is called "Slap" in this paper. is seen either during play or as a component of an attack (Goodall, 1989). Kano's (1998) Hit for the bonobo. Category 1. Hit conspecific or other species with objects: See Club. Hold chin/head up (HLU) (Type A): Lift chin or head of another up with one hand in order to groom jaw or neck. Also occurs in bonobos (Kano, unpublished). Category 1. Hold face (HLF) (Type A): Support the face or head of another with one hand while grooming the face or head with the other hand. Seen in bonobos (Kano, unpublished). Category 1. Hold genitals (HLG) (Type A): "Goodall's (1989) Hold genitals. Adult usually grabs or touches the scrotum or penis of a male as reassurance. Often occurs when an adult male is mounted by another male, the mounted reaches his hand between his thighs and holds the scrotum of the mounter. When an adult female pant-grunts to an adult male she sometimes holds his scrotum. Never seen in the bonobos of Wamba (Kano, 1998). Category 3-5. Hold hand (HLN) (Type A): Goodall (1989): "The most commonly seen form of holding... This occurs when a higher ranking individual grasps the stretched out hand of a lower ranking chimp. A type of hand shaking has been observed when the subordinate holds the hand, given in reassurance , of the higher ranking animal." Not seen in these contexts in bonobos (Kano, 1998). Category 3? Hold head (HLH) (Type A): Hold head of another with both hands in order to groom the face with lips and teeth. Occurs also in bonobos (Kano, unpublished). Category 2? Hold in mouth (HLM) (Type A): Have an object between jaws. Category 1. Hoo (H00) (Type A): Goodall (1989): "A single syllable soft whimper. A single hoo may be uttered several times in succession. It is the typical sound given by an infant which cannot reach the nipple and wants to reestablish contact with the mother and so on." At Mahale, an infant hoos when it begs for food , when it wants to return to its mother after separation, and even when it wants to mate with an estrous female. A mother also hoos when her older infant does not come back to her, often while playing above the ground when she wants to leave. Thus, hoo expresses frustration (Cf. Whimper). Absent in bonobos? Category 3-5. Hoot (HOT) (Type A): Goodall (1989): "Roar pant hoot. Continuous low-pitched calls given only by highly aroused individuals, rarely by females. Always accompanied by a charging display." Category 3-5. Hug (HUG) (Type A): Caretaker embraces an infant with both arms. Also, intention movement of mounting. See Figure 8 of Nishida (1970) and Figure 3 of Nishida (1983b). Kano's (1998) Protective embrace for the bonobo. Category 1. Hunch (HUC) (Type B): Goodall (1989): Hunching of shoulders in the "contexts of aggression, courtship and greeting". "Nearly always accompanied by hair erection." Absent in bonobos? Category 3? Hunch and sit (HUS) (Type A): Goodall's (1989) Sitting hunch: "The chimp, while sitting, raises shoulders, arms held out from the body either to the side or in front." At Mahale , sitting hunch courtship is often accompanied by stamping the ground or branch-shaking. Bonobo male's "Sitting erect" (Kano, 1998) might be homologous to HUS. Category 2? Hunch bipedal (HUB) (Type A): Goodall (1989): "Chimp stands on one's feet, shoulders hunched up, arms held slightly out and away from the body. If he sways from foot to foot, known as bipedal swagger". Known also from West Africa (Plate 31 of Albrecht and Dunnett, 1971). Absent in bonobos? Category 3? Hunch over (HUO) (Type A): Goodall (1989): "Similar to bipedal hunch, but arms held forward over the back of another chimp." De Waal's (1988) Hunch over. Kano's (1998) Bluff over for the bonobo. Category 2. Hunch quadrupedal (HUQ) (Type A): Goodall (1989): "Stand, walk or run on all fours with rounded back and head pulled in between shoulders." Seen among adult males at Mahale. Also occurs in bonobos (Kano, unpublished). Category 2. Hunt (HUN) (Type D): Behavioral complex of stalking, pursuing, capturing, and killing large birds (for Ethogram of chimpanzees 159 example, guinea fowl) or small to medium-sized mammals. Include , , , , , , , , , , , , , and so on. Known throughout the chimpanzee range (Boesch and Boesch, 1989; Uehara, 1997). Bonobos hunt less often than chimpanzees (Kano, 1992). Category 1. Huu (HUU) (Type A): Goodall (1989): "...very like the hoo whimper, but is usually higher pitched and the huu does not show the pouted lips characteristic of hoo. It is made when a chimpanzee suddenly hears or sees a strange object or sound." This sound often helps a researcher to "discover" an otherwise silent, hidden chimpanzee in the bush. Absent in bonobos? Category 3-5?

I Ignore (IGN) (Type C): Not respond to stimuli, as a result of being habituated, that usually elicit some responses. For example, chimpanzees of M group now act normally in human presence although they would run away before habituation succeeded. Goodall's (1989) Ignore: "When an individual, sometimes dominant, does not respond to the gesture in any manner apparent to the human observer." Kano's (1998) Ignore for the bonobo. Cf. Snub. Category 1. Imaginary play: See Play, imaginary. Infanticide: See Kill. Inspect (INS) (Type D): Investigate the environment including conspecifics by , , , and so on. Category 1. Inspect genitals (ING) (Type B): Goodall's (1989) Inspect: "When a male or female touches the vaginal opening of a female and sniffs finger, or sniffs with nose directly. Both hands may be used to pull apart the lips of vagina." See Figure 11 of Nishida (1970) for photograph. Plooij's (1984) INS: Also directed to self. By contrast, no adult male bonobos inspect female genitals, although juveniles do (Kano , 1998). Category 2? Interfere (ITF) (Type D): Goodall's (1989) Interference: A pattern of intervention (see Intervene) in which "a third individual prevents or tries to prevent an interaction between two others. This is most often seen in the sexual context..." Bonobos also interfere in sexual and grooming interactions (Kano, 1998). Category 1. Interfere copulation (ITC) (Type D): Weaning infant (sometimes even adolescent male), often the offspring of the mating female, runs to push itself between the mating pair, waves arms, or touch or push at a male who is copulating, while emitting whimper/scream or loud scream (Tutiu, 1979; Nishida, 1997). A juvenile male may do the same thing to his elder brother. An (often dominant but not always) adult male may interfere mating between a female and a male by barking, threatening, or charging directly . An adult female (often in estrus) may squeeze herself into a mating pair. Dominant adult males of the bonobo interfere with mating involving adult females, which Kano (1998) calls "Sexual interference". Category 2. Interfere to separate (ITS) (Type D): Charging display performed by an alpha male towards a rival male and a third male who are grooming each other, apparently in order to prevent the formation or development of the coalitionary relationships between them. De Waal's (1982) Separating intervention. See also Nishida and Hosaka (1996). Absent in bonobos, but dominant individual (females more often than males) may thrust oneself, or insert one's arm, into a grooming pair to take over grooming (Kano, unpublished). Category 3. Intervene (ITV) (Type D): Exert influence on social interaction of two or more conspecifics by threat or aggression. Intervener may be partial or impartial. Also occurs in bonobos. Includes Interfere and Support. Category 1. Invitation slap: See Slap in invitation. Invitation stamp: See Stamp in invitation.

Jump: See Leap. J

K Kick (KCK) (Type B): Goodall (1989): "Make contact with an objective (usually another chimp) with one or both feet. Kicking is a forward, sideways or backward movement, different from a stamp which is always downward..." See Figure 3 of Nishida (1994). Many types of kicking are described for the bonobo 160 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. by Kano (1998). Category 1. Kick back (KCB) (Type A): Goodall (1989): "Walking individual kicks backwards towards a following youngster..." The same behavior is seen for the bonobo (Kano, 1998). Category 2? Kick backward quadrupedal (KCQ) (Type A): Kick backward buttress/tree trunk/wall, while standing on all fours. Goodall's (1989) Drum/Kick. Category 2-3? Kick bipedal (KCP) (Type A): Kick buttress/tree trutik/wall with one foot while upright. Goodall's (1989) Drum/Kick. Absent in bonobos? Category 3-5. Kick heel (KCH) (Type A): Goodall's (1989) Heel kicking: "When a resting individual (often an adolescent or adult) raises one leg and thumps its heel down on a youngster who is playing around him or her." Seen in the chimpanzees of Mahale. However, Goodall's other type ("An older male, when lying on back holding an infant in a ventro-ventral embrace, kicks down gently one heel after the other , on the infant's rump") is not seen in Mahale. Not seen for the bonobo (Kano, 1998). Category 3-5. Kick social (KCS) (Type C): Kick other individual in fight and play. Seen in bonobos. Category 1. Kidnap: See Detach (cf. Care alloparentally). Kill (KIL) (Type D): Fatal attack to another chimpanzee. At Mahale only infants were observed to have been killed and most of them were eaten by killers and other chimpanzees. Infanticide has also been recorded from Budongo, Gombe, Kibale and Tai (see Eat conspecific). Adult chimpanzees were suspected to have been killed within a group (Nishida, 1996) and between groups (Nishida and others, 1985). Killing adults have been recorded at Gombe (Goodall, 1986) and Budongo (Whiten, pers. comm.) and suspected at Kibale (Wrangham and Peterson, 1996). Category 1? Kiss (KIS) (Type B): Goodall (1989) divides Kiss into two forms: pout-kiss and open-mouth kiss. Both of these types are seen among the bonobo, but rare. Adult bonobos at Wamba do not kiss (Kano, 1998). Category 1. Kiss in pout face (KPO) (Type A): Goodall (1989): "The lips are slightly pouted and pressed against, or briefly laid against, the body, face, or limbs of another. Pout kissing often occurs in greeting, submissive and reassurance contexts." Absent in bonobos (Kano, unpublished). Category 3. Kiss with open mouth (KOM) (Type A): Plooij's (1984) OMK. Goodall's (1989) Open mouth kiss: "The open mouth is pressed to body or mouth of another. If two chimps open-mouth kiss each other on the face one often presses his mouth over the other's upper lip and nose, the other has his mouth over the lower lip and chin of the first..." Open-mouth kissing often occurs during reunion and social excitement , in particular, among adult males, accompanied by panting. See Figure 5 and 6 of Nishida (1970) for photograph. Kano's (1998) Open mouth kiss is directed only to infants and juveniles by older bonobos, and never seen between adults. Category 2 or 3. Knuckle walk: See Walk quadrupedal with knuckle.

L Lap into (LAP) (Type A): Goodall (1989): "An infant jumps into the lap of a sitting chimp, during greeting." Seen at Mahale, but not in greeting. Category 8? Laugh: See Play-pant. Lead (LED) (Type D): Mature male or female, and especially adolescent male, leads sexual partner into undergrowth or higher in a tree in order to avoid interference by more dominant males. See Nishida (1997). Seen in bonobos (Kano, unpublished). Category 2? Leaf-clip (CLP): See Clip leaf. Lean (LEA) (Type A): Incline torso to rest against a tree trunk, bough, rock or another individual. Kano's (1998) Reclining sit. Category 1. Lean forward (LEF) (Type A): Flex the trunk in order to expose the back for grooming. Seen also in bonobos (Kano, unpublished). Category 1-2. Leap (LEP) (Type B): Jump propelled by both feet. Plooij's (1984) JUM. Kano's (1998) Leap for the bonobo. See Figure 6 of Goodall (1968) for illustration. Category 1. Leap bipedal (LPB) (Type A): Adult males occasionally stand upright, swing their body back and forth in order to gain momentum, and then jump from a rock to another in upright posture when crossing a stream by using stepping stones. Absent in bonobos of Wamba (Kano, unpublished). See Figure 3 of Goodall (1968) for illustration. Category 3? Leap bipedal on the spot (LBS) (Type A): Juvenile male jumps on his feet in front of an estrous female in courtship, to elicit her to "present". See Nishida (1997). Juveniles also jump up and down while Ethogram of chimpanzees 161 pant-grunting vigorously in front of a dominant male. Category 8? Leap down (LPD) (Type A): Leap from a tree down to vegetation or ground. Similar behavior is known for the bonobo of Wamba (Kano, 1998). Category 1. Leap on (LPO) (Type C): Spring on another individual in aggression. Kano's (1998) Jump on is similar for the bonobo, but then the jumper is likely to mount. Category 2-3? Leap quadrupedal (LPQ) (Type A): Usual pattern of crossing a stream via stepping stones . Before leaping, chimpanzees swing their body back and forth in order to gain momentum in a quadrupedal posture. Bonobo commonly leaps on all fours above and on the ground (Kano, 1998). See Figure 3 of Goodall (1968) for illustration. Category 2. Leap vertical (LPV) (Type A): Goodall's (1989) Vertical leap: "Leap vertically from one structure to another." Seen at Mahale when a chimpanzee leaps from a tree crown to another crown below. Kanos (1998) Vertical leap for the bonobo. Category 2. Leave (LVE) (TypeC): Move away from another individual. Plooij's (1984) LVE. (cf. DEP), Category 1. Leg cradle: See Cradle with leg. Lick (LIK) (Type A): Plooij's (1984) LIK. Repeatedly stroke with tongue. Category 1. Lick rock (LIR) (Type B): Lick the rocks at limited sites along the shore of Lake Tanganyika or in the riverbed of large rivers. Happens more often in the dry season than wet (Nishida, 1980a), and sometimes continues as long as one hour. Confirmed for the chimpanzees of B, K, M and L groups (unpublished data). Nutritional significance is unknown. Rocks licked are not salty to a human observer. See Figure 3 of Nishida (1980a) and Figure 12 of Nishida and Uehara (1983) for photo. Not observed at Gombe (Goodall, 1986). Category 6. Lick sneeze (LSN) (Type B): Insert finger into nostril and lick it. Seen in bonobos (Kano , unpublished). Category 1. Lick wood (LIU) (Type B): Licks bark-free surface of dead trees such as Pycnanthus angolensis, Ficus capensis, Garcinia huillensis and so on., with extended upward movements of tongue. Nutritional significance is unknown. Not observed at Gombe. Category 6. Lick wound (LIW) (Type B): Lick a wound apparently to clean it, directly with tongue or touch wound and then lick finger (s). Sometimes done socially, often by infants or juveniles. Also seen in the bonobo, who even lick menstrual blood (Kano, 1998). Category 1. Lie (LIE) (Type B): Plooij's (1984) LIE. Kano's (1998) Lie for the bonobo. Category 1. Lie-hug (LIH) (Type A): Nishida (1983a): Caretaker lies supine on the ground while embracing an infant to the chest. Seen in bonobos (Kano, unpublished). Category 1. Lie lateral (LID) (Type A): Lie on side. Common resting or sleeping posture on the ground. Category 1. Lie prone (LIP) (Type A): Lie on belly. Posture is often assumed by estrous females or pregnant females near term. See Figure 5 of Nishida (1983a) for photograph. Category 1. Lie-sit (LSI) (Type A): Typical sitting or lying posture of an estrous female, apparently forced to assume by extremely swollen sexual skin. Category 3? Lie supine (LIS) (Type A): Lie on back with legs not spread. Most common sleeping posture in the bed or on the ground. See Figure 4 of Nishida (1983a) for photograph. Category 1. Lie supine and legs apart (LIL) (Type A): Lie on back with legs spread. Often occurs when male solicits another to groom his scrotum. Category 1? Lie with back to another (LIB) (Type C): Pattern of soliciting grooming of one's back. Seen in bonobos (Kano, unpublished). Category 2. Lie with legs crossed (LIX) (Type A): Lie supine with raised legs crossed. Resting posture. Category 1? Lift (LFT) (Type A): Hold heavy object up with both arms. Seen in bonobos. Category 1. Lift and drop (LFD) (Type B): Lift heavy log or thick woody vine with both hands and drop it on the ground to make a loud sound. Display done by two young adolescent males (Primus, Orion) as a courtship or mild threat. Category 8. Lift rock (LFR) (Type A): Adult male lifts rock with both hands. (This leads to throwing it into water to make splashing sounds, apparently to heighten the effect of the charging display). See Throw splash. No rock in the habitat of Wamba bonobos. Category 6-7? Limp (LMP) (Type A): Walk lame because of wounded limb. Category 1. Lip flip: See Flip lip. Lip-smack: See Smack lip. Locomotor behavior: See Travel 162 Nishida T., Kano T., Goodall J., McGrew W. C., and Nakamura M.

Locomotor play: See Play. Look back (LOB) (Type A): Groomee looks back at the groomer who has just stopped grooming, and the latter immediately resumes grooming. Thus, turning round is a request to resume grooming. A walking chimpanzee looks back in order to monitor whether the companion is following. This often occurs when an adult male leads his sexual partner during consortship. Also occurs in the bonobos of Wamba in the similar contexts (Kano, 1998). Category 1. Look up (LOU) (Type A): While sitting or standing, a chimpanzee raises the face up in order to monitor arboreal stimuli, for example, fruit, colobus monkey, or other conspecifics. Category 1. Lost call: See Whimper-scream. Lower arm (LWA) (Type A): Groomee lowers arm, for example, when the grooming-hand-clasp stops, signaling intention to change role or posture. Used by groomer to request groomee to change role among bonobos (Kano, unpublished). Category 2? Lower head (LWH) (Type A): Two individuals face each other in social grooming, and one partner flexes the neck to offer the top of head for grooming. Occurs also in bonobos (Kano, unpublished). Category 1-2. Lower leg (LWL) (Type A): Resting or lying groomee lowers raised leg, which apparently stimulates the groomer to groom another body part. Similar patterns is seen in bonobos (Kano, unpublished). Category 2?

M Make a bed (MBD) (Type B): Goodall's (1989) Nest. Construct a platform for sleep or rest by bending branches over each other. "Bed" is preferable to nest because the major function is not for rearing but for sleeping. Includes , , , and . Category 2. Make a cushion (MCU) (Type B): Bend or break one or a few shrubs or other vegetation and sit or lie on them. Above or on the ground. Used for daytime rest or nap. A bed is a more complex construction than a cushion. Bonobos of Wamba make cushions above or on the ground (Kano, 1998). Category 2-5. Make a day bed (MDB) (Type B): Goodall's (1989) Day nest. Day bed is made much more quickly than night bed. Bonobos also make day beds (Kano, 1998). Category 2. Make a night bed (MNB) (Type A): Goodall's (1989) Night nest. Plooij's (1984) Nestbuilding (NES). Category 2. Male invite: See Open thighs. Masturbate: Goodall's (1989) Masturbation. See Fumble with penis for males and Fumble with clitoris for females. Maternal behavior: See Care maternally. Maternal leave: See Travel alone during childbirth. Monitor (MNT) (Type D): Investigate the changes in the environment or social interactions. See Figure 5 of Nishida (1983a) for photograph. Category 1? Monitor fruits (MNF) (Type D): Travel through a concentrated patch of fruit trees and apparently check the good or poor crop even long before the fruit ripens. Includes , , , , and . Category 3-5? Monitor monkeys (MNC) (Type D): When in the "hunting mood", chimpanzees sit on the ground , look up, and move little by little, perhaps in order to look for immature monkeys and to check their movements in general. Includes , , , , and so on. Category 3? Monitor mother (MNM) (Type D): When an infant is being weaned by a rejecting mother, it runs screaming from her, stay a few meters away and repeatedly glances to her to see if she will accept it again (namely, allow it to suckle, or to put ventral/dorsal). Includes , , , , , , and . Also seen in bonobos (Kano, unpublished). Category 1 or 2. Mop ants (MOP) (Type A): Take Camponotus ants with the back of a hand. This often occurs when many wood-boring arts emerge from the nest and move about on the surface of the trunk. See Nishida and Hiraiwa (1982). Category 6? Mount (MOU) (Type A): Embrace another with both arms from behind. Goodall (1989): "Part of the ventral surface of the mounter is in contact with part of the dorsal surface of the other, and the mounter leans forward over the other, usually grasping him/her". She also mentions that "the mounted individual (male or female) may reach back to touch the genital area of the mounter with a hand, (or, rarely, the foot), or the mounter may raise his/her own foot to the scrotum of the other." Goodall (1989) lists as synonyms: "Dorso-ventral embrace", "Mounting embrace" or "Mount-embrace". Perhaps, Kortlandt's Ethogram of chimpanzees 162

(1964) Sham copulation. Plooij's (1984) MOU, de Waal's (1982) Mount. Both male and female can mount. Mounting occurs especially among adult males during high social excitement such as the reunion of rival males in which each rival male mounts an adult male or female, and both participants often scream or pant-scream. Mounting is usually accompanied by pelvic thrusts. Mounting appears to function as self-reassurance, reassurance, recruitment of coalition partners, and so on. Among the bonobos , mounting occurs in similar contexts such as appeasement and easing of tension, but posture differs , such that the mounted usually stands on all fours (Kano, 1998). Category 2. Mount in copulation (MOC) (Type A): Goodall's (1989) Copulation mount: "The male places both hands firmly on the female's back or sides, during which he may open mouth kiss her... accompanied by intromission and thrusting movements of the pelvis." Category 1. Mount, misdirected (MOM) (Type A): Van Hooff's (1973) Maldirected mounting: "Thrusting towards the head, face, shoulder or any other part except the rear." Category 3? Mouth (MOT) (Type A): Van Hooff's (1973) Gnaw: "Keep mouth wide open without the retraction of lips and press the teeth to the back, shoulder or other body part without biting". Reassurance behavior often accompanied by panting. Similar pattern is used when tickling with mouth. The mouth is pressed to the shoulder, chest, belly and any other part of the body. Nishida's (1983a) Mouthing or Mouth contact. Plooij's (1984) Mouthing (SAB). Goodall's (1989) Biting includes Mouthing. Goodall's (1989) Play biting occurs in the contexts of social play. See Figure 5 of Nishida (1983a) for photo. Kano's (1998) Mock bite for the bonobo. Category 1. Mumble (MUM) (Type A): Sounds of lips and tongues while grooming. Category 2-3?

N Neck pocket: See Transport in neck-pocket. Nest-build: See Make a bed. Nest-grunt: See Grunt in bed. Nest play: See Play in bed. Nuzzle (NZL) (Type A): Goodall's Nuzzle: "The young infant, until about 3 months old, moves its head from side to side and up to and down against the mother's body when searching for the nipple ..." Also called rooting behavior in other mammalian species. Kano's (1998) Nuzzle for the bonobo. Category 1.

Object play: See Play with object. Offer arm (OFA) (Type C): Dominant individual extends arm to another to allow the latter to mouth the arm to be reassured. Not seen in the bonobos of Wamba (Kano, unpublished). Category 3-5. Open eyes (EYO) (Type A): Plooij's (1984) Eyes open (EYO). Category 1. Open mouth kiss: See Kiss with open mouth. Open thighs (OPT) (Type A): Goodall's Male invite: "The male sits with thighs splayed and penile erection, looking toward a female in estrus. Sometimes his hair is erect..." Male courtship display pattern. Similar pattern is shown by the bonobo (Kano, 1998). Category 2.

P Palmigrade walk: See Walk quadrupedal palmigrade. Pant (PAN) (Type B): Goodall (1989): "...various calls which are linked by audible inhalations and which may be described as vocalized pant. Non-vocal panting sometimes occurs whenchimpanzees are grooming each other, during greeting, and so on. Sometimes, when the mouth isclosed, only breathing sounds are heard. The best indication of panting is the quick rhythmic movements of the body that accompany it." Pant occurs in bonobos in play, but unknown in other contexts? Category 2-3? Pant-bark (PBA) (Type A): Goodall (1989): "Series of bark-like sounds joined by voiced inhalations." Status unknown in bonobos. Category 2-3? Pant-grunt (PGR) (Type A): Goodall (1989): "A series of soft or loud grunts functioning as a token of respect given during greeting by submissive chimpanzees and during submissive interactions... A highly fearful individual may utter frenzied pant barks that may be labeled pant-screams." Van Hooff's(1973) Rapid OhOh. At Mahale, while pant-grunting adult females may to adultmales, who may mount the females and show thrusting. In response to pant-grunting by young adult and adolescent males, alpha males may jump on or attack them without being appeased. See Figure 2 of Nishida (1994) for

O 164 Nishida T., Kano T., Goodall J., McGrew W. C., and Nakamura M.

photo. Pant-grunt is completely lacking in the bonobo (Nishida and Hiraiwa-Hasegawa, 1987).Category 3. Pant-grunt with bent elbow (PGE) (Type A): Pant-grunt while crouched quadrupedally with elbow flexed. Pattern in pant-grunt complex performed by older juvenile and adolescent males at Mahale. Category 3-5. Pant-hoot (PHT) (Type A): Goodall (1989): Series of long calls "which are often contact calls between distant groups or calls given by chimpanzees at night, from their nests, when they are within earshot of another group... give pant-hoots when they arrive at a food source, cross a ridge, or face into a new valley. A subordinate greeting a dominant often gives pant hoots and pant grunts..." At Mahale adult males also pant hoot when agonistic confrontations are continuing, in which pant hoots may function as a vocal threat. See Figures 4, 5 and 6 of Nishida (1983b) for photograph. Pant-hoots may be divided into four stages: build-up, pause, climax and end, but females lack the climax and end stages (Mitani and Nishida, 1993). Exact bonobo edition of pant-hoot call is absent (Nishida, personal observation), but, Kano's (1998) Waa call, or Okayasu's (1991) Hoot may be homologous to the pant-hoot. The bonobo Waa call is emitted at the time of arrival in the feeding ground, during feeding, during resting and at sunset after making the night beds (Kano, 1998). Category 2. Pant-scream: See Pant-grunt. Parry (PAR) (Type A): Van Hooff's (1973) Parry: "One or both arms are raised. The forearm is kept in a roughly horizontal position over or in front of the head, thus shielding it off from possible beats from a fellow." Goodall's (1989) Startle flinch: "When a chimpanzee is startled by a sudden movement nearby (such as a low-flying bird, large insect, unexpected gesture of human or even another chimp, and so on.) he/she will immediately duck his head and fling one or both arms across his/her face, or if very startled , throw both hands in the air..." Also, corresponds to Goodall's (1968) Startle reaction. The bonobos of Wamba may duck their heads in the similar context (Kano, unpublished). Category 1. Pass (PAS) (Type C): Walks past another, without body contact. . Goodall's (1989) Pass. Kano's (1998) Pass without body contact for the bonobo. Category 1. Pass with body contact (PAB) (Type B): Pass by another while making body contact (not with hand or foot) although the performer can pass without doing so. Two young adult males and three juvenile females at Mahale did so to a human observer. Significance of the behavior is unknown. Kano's (1998) Pass with body contact for the bonobo. Category 2. Pat (PAT) (Type A): Plooij's (1984) Pat. Goodall (1989): "Repeated brief contacts , with the palmar surface of the hand, on the body of another, usually on the head, back or hand. This is a reassurance gesture directed towards a submissive individual by a more dominant individual's behavior. It has a calming effect." Not shown by adult, but by juvenile, bonobos of Wamba in play (Kano, 1998). Category 1? Patrol (PTR) (Type D): Nishida's (1979) Scout. Goodall (1989): "Party of males (occasionally accompanied by a female, usually in estrus) moves deliberately and silently in the peripheral part of community range". Elements include , , , , , and . Reported also from Budongo (Newton-Fisher, 1998), Kibale (Wrangham and Peterson , 1996) and Tai (Boesch and Boesch, 1999). Bonobo males similarly approach an adjacent group silently, however, only after they have heard the latter vocalize, and so Kano (1998) calls it "Scout". Category 1 or 2. Peel with hand (PEH) (Type A): Pull the tip of the outer layer of stalks of grasses, herbs and woody vines with one hand while holding it with another. Supplementary movement of PET. Similar behavior is seen in the bonobo (Kano, 1998). Category 1. Peel with teeth (PET) (Type A): Pull with teeth the tip of the outer bark while holding the stalk with both hands, thus remove the hard outer layer of stalks of grasses (sugar cane and elephant grass) , herbs (ginger and Marantochloa) and woody vines (Saba and Landolphia) in order to eat pith. Similar pattern is seen in the bonobo (Kano, 1998). Category 1. Peer (PER) (Type A): Look intently into another individual's face from a few cms. distance usually in a quadrupedal stance. Goodall (1989) saw peering when another individual eats, leaf-grooms, or investigates or licks a wound. Plooij's (1984) Put face close. Peering does not usually elicit food sharing. See Figure la of Nishida (1983a) for photograph. Bonobos show the same pattern (Idani, 1995; Kano, 1998). Category 1. Pelvic thrust: See Thrust. Penile adduction: See Adduct, penis. Penile erection: See Erect penis. Pick up (PIU) (Type A): Pick up with the thumb and forefinger small objects such as sticky seeds of grasses or beggar's lice from the body surface in grooming. Also seen in bonobos. Category 1. Ethogram of chimpanzees 165

Pirouette (PIR) (Type A): Goodall (1989): "The chimp (usually a youngster) progresses in a series of tight circles, moving on all fours or on their feet. A type of locomotor play". See Figure 7f of Goodall (1968) for illustration. Seen among juveniles at Mahale. Probably absent in the bonobos of Wamba (Kano, unpublished). Category 3?

Play (PLY) (Type B): Goodall (1989) divided play into lone play and social play. Further, she divided lone play into 4 categories, (a) locomotor play, or van Hooff's (1973) Gymnastics, (b) nest play, (c) object play, and (d) self play. She added that, "There is a facial expression connected with play, the play face, a type of locomotion that is seen only in the play context, the play walk, and a vocalization, laughing." Goodall (1989) listed as elements of each play type as follows: (a) locomotor play: , , , , , and . (b) nest play: , , and . (c) object play: , , , , , , , and . (d) self play: , , , and . Goodall's social play included , , , , , , , , ,

, , , , , and so on. Category 1.

Play-bite (PLB) (Type A): Goodall's (1989) play bite: "Contact made on the partner's body with teeth.

Sometimes one of the players gets a firm grip on a limb and holds on for minutes at a time..." See Mouth, since play-bite is displayed in the play contexts. See Figure 4.9 of Hasegawa (1990) for photograph. Category 1 or 2. Play face: See Show place face.

Play, imaginary (PLI) (Type D): Engaged in self-play, but apparently with imaginary companion or tool.

A multiparous adult female lying on her back with all limbs up was seen to play-pant as if she was playing "aeroplane" with her infant who had been dead for a few months (Nishida , unpublished). Hayaki (1985) recorded an adolescent male running around a tree repeatedly while play-panting loudly as if he was chasing, or fleeing from a playmate. Rarely observed. Category 8?

Play in bed (PLN) (Type B): Goodall's (1989) Nest play above. Seen also among the bonobo (Kano,

1998). Category 2.

Play in rough and tumble (PRT) (Type B): Most physical type of play usually between youngsters . Comprises wrestling and often locomotor play. Elements include , , , ,

, , , , , , , , , , ,

, , , and so on. Category 1.

Play-pant (PLP) (Type A): Plooij's (1984) Laugh (LAC). Goodall's (1989) Laugh: "Somewhat resembles human laughter, heard when chimpanzees are playing socially. As the play becomes rougher, or involves more tickling, there is more laughter..." Bonobos also play-pant (de Waal, 1988; Okayasu , 1991; Kano, 1998). Category 1.

Play self (PLS) (Type B): Goodall's (1989) Self play. Kano's (1998) Lone (Self) play. Category 1.

Play socially with object (PSO) (Type B): Goodall's (1989) Social object. Social play including chasing an individual who has an object such as a fruit, branch, stone and animal skin and so on . Ingmanson's (1996) Tool use in play for the bonobo. Category 1 or 2.

Play walk (PWL) (Type A): Goodall (1989): "The chimp walks with a rounded-back, its head slightly bent down and pulled back between the shoulders while it takes small stilted steps•c" Seen also in bonobos

(Kano, unpublished). Category 2. Play with object (PLO) (Type B): Goodall's (1989) Object play above. Category 1.

Play with water (PLW) (Type B): Type of self play including , , and so on. Category 1-7.

Poke (POK) (Type A): Nishida (1983a): Tickling, by prodding repeatedly with fingertip contacts with one or several extended fingers to an infant's ventrum while lying on its back. Goodall (1989) mentioned that "Sometimes an individual may poke another chimp who has ignored a solicitation for grooming ." Poking in this context has not been seen at Mahale. See Figure la of Nishida (1983a) for photograph. Seen in bonobos. Category 1.

Pout (POU) (Type A): Plooij's (1984) PTF. Goodall's (1987) Pout face: "Lip corners pushed forward, mouth may be slightly open. Lips together, except at the front where they may funnel out somewhat. This is seen with the hoo call and hoo whimper". Bonobos have a similar facial expression (Kano , 1998). Category 1.

Pout face: See Pout.

Pout kiss: See Kiss in pout face. 166 Nishida T., Kano T., Goodall J., McGrew W. C., and Nakamura M.

Presentwith limbs extended(PRE) (TypeA): Normal quadrupedal stancewith hindquartersdirected to dominant chimpanzee. Goodal1's (1989) Present.Goodall mentions Rump-turn as a synonym . Plooij' s (1984) PRE. VanHooff' s (1973) Mount-present.Kano' s (1998)Dorso-ventral presenting for the bonobo. In addition,bonobo females have ventro-ventralcourtship gesture. Category 2. Presentwith limbs flexed(PRF) (TypeA): van Hooff (1973)'s Crouch-present.Quadrupedal posturewith limbs flexed,and hindquartersturned towards another individual.See Figure 7 of Nishida (1979) for photograph.Differs with Crouch in which hindquartersare not directedto the partner.Category 3. Press (PRS) (TypeA): Pushone s shoulder,side or hip againstanother individual. Function unclear , but sometimes initiatesocial play. Category 3-8? Press teeth againstback (PTB) (TypeA): Adult male mounts another male, and the mounter often presseshis upper row of teethagainst the back of the mounted male, while the mounter embraces the latter,performs a pelvicthrusts and shows a wide open grin.Intensified form of . Not seen in the bonobos of Wamba. Category 3-6. Probe (PRB) (TypeA): Use a vine,twig or stickto touchsome seen or unseen objects.Goodall's (1989) Investigationprobe. Infantchimpanzees appear to have an innatetendency to push an investigation probe into any hole of a tree long before they fishfor arborealants. Not been seen among the bonobos of Wamba (Kano, 1998).Category 3? Protect (PTC) (TypeD): Goodall' s (1989) Protect.Caretaker protect an infantfrom rain,storm, dangerous situation(for example, gaps in canopy) or socialharassment (for example , malecharging displays).Simar patternis seen in the bonobo, exceptfor protectionfrom rain(Kano , 1998). Category 1. Pull (PUL) (TypeA): Grasp and tug at an objector otherindividual by flexingarms. Occurs frequently duringfeeding, play, grooming and agonism.Kano' s Pullfor the bonobo. See Figure l ofNishida (1983a). Category 1. Pull down (PLD)(TypeA): Stretcharm to reach upper branch of shrub,often whilestanding on both feet,and pullthe branch downward. Followed by sittingand eatingfruits or leaves whilegripping the branch with one hand. Also seen in bonobos (Kano, unpublished).Category 1. Pull face to face (PLF)(Type C): Pullhand or otherbody part of anotherindividual , so thatthey face each other.Occurs when chimpanzee wants to groom the frontalpart of the other.Also seen in bonobos (Kano, unpublished).Category 1-2. Pull from opposing sidessimultaneously (POS)(TypeB): Tug of war. Two chimpanzees simultaneously pull an object (infant,branch, carcassand so on.)competitivelyfrom opposing sides. Seen also in bonobos. Category 1. Pullhead with both hands (PLH)(TypeA): Pullhead of anotherwith both hands,of ten in grooming.Also seen in bonobos (Kano, unpublished).Category 1-2. Pull out (PUO)(TypeA): Pullfor cibly with both hands, oftencountered by pushingmovement of both feet on the ground.Occurs when chimpanzee triesto draw stalksof banana, elephantgrasses, or African gingersfrom the soil.Also seen in bonobos. Category 1. Pullthrough (PTH)(TypeA): Upward sweeping movement of thumb and index fingeror palmar surface of hand along a leafybranch or leafyherb stem. This is a food-processingtechnique for leaves, vines, herbs or blossoms (forexample, Erythrinaabyssinica) while holding a twig with one hand and pulling it through between lips/teeth.Also used by Gombe chimpanzees in ant dipping(McGrew , 1974).Seenin bonobos (Kano, unpublished).Category 1-2. Pull through stem (PTS)(TypeA): Nishida's (1997) Stem pull-through(courtship). Pull the leafybranch of a shrub or clump of grassstems throughthe hand by a rapidupward movement of the forearm, then, the stem is immediatelyreleased. Produces a conspicuoussound, while seated and watching an estrous female.Only two of the oldestadult males, Musa and Ntologi,did thiscourtship. An exaggerated form of pullingthrough in the eatingcontext. One adultmale (Fanana)toreoff the shootof a stem from a shrub as a part of courtship.Category 8. Push (PUS)(TypeA): Exert forceby extendingarms in contactduring aggression and grooming.In order to groom another,push part of the back (forexample, shoulder)of anotherso thatthe latterwill turn its back.Groomer stimulatesthe groomee to change postureafter extensive grooming of a particularpart of the body. Kano's (1998) Push for the bonobo. Category 1. Push ahead (PUA)(Type C): Goodall (1989):"A mother pushes her infantahead of her duringweaning to preventdorsal travel. Often she does thiswith littleslaps of her hand on itsrump and thisisa pparently perceivedby the infantasplayful". PUA isnot seen in the bonobos of Wamba (Kano, 1998).Category 3-5. Ethogram of chimpanzees 167

Push away (PAW) (Type A): Goodall's (1989) Push: "Shove away from anotherindividual with one or both hands...", and Goodall's (1989) Push away: "When a mother pushes her child away when it approaches to suckle or ride on her; also occurs during begging when adults are pushed away". Kano's (1998) Push away for the bonobo. Category 1-2. Push backward (PUB) (Type A): Push a rock backward with one hand while running on the river bottom. Element of the charging display of some adult male chimpanzees of M group. Not seen in bonobos (Kano, unpublished). Category 3-8. Push finger into mouth (PFM) (Type A): Nishida (1983a): "insert one or morefingers/toes into the mouth of another individual." Goodall's (1989) Finger in mouth includes both PFM andTFM. This appeases or reassures another (usually, more dominant) individual, or calms an excited adultmale. Not seen among the bonobo except when infants or juveniles are begging for food (Kano, 1998).Category 2. Push forward (PUF) (Type A): Juvenile female (Maggy) played by pushing repeatedl a hard shelled fruit forward along the ground. Category 8. Put dorsal (PTD) (Type A): Nishida's (1983a) Put dorsal, Goodall's (1989) Takedorsal: Caretaker places infant on back, either over the shoulder, often with both hands, or with ascooping upward movement of the hand when the infant was on the belly. Similar behavior is known for thebonobo (Kano, 1998). Category 1. Put face close: See Peer. Put ventral (PTV) (Type A): Goodall's (1989) Take ventral: "Mother reaches toinfant, grasps, and presses to a ventral position." Kano's (1998) Take ventral for the bonobo.Category 1.

Q Quadrupedal hunch: See Hunch quadrupedal. Quadrupedal jump: See Leap quadrupedal. Quadrupedal run: See Run quadrupedal. Quadrupedal transport: See Transport quadrupedal. Quadrupedal walk: See Walk quadrupedal with knuckle.

R Raise (RAI) (Type B): Elevate limb while lying supine or sitting on the ground.Seen in bonobos (Kano, unpublished). Category 1-2. Raise and hold leg (RAH) (Type A): Raise a leg and hold it with one or both hands while lying supine, often when being groomed. Seen in bonobos (Kano, unpublished). Category 2. Raise arm quickly (RAQ) (Type A): Goodall's (1989) Arm threat: "The arm (whole or forearm only) is raised in a quick jerky movement and the fingers flexed slightly. This is usually accompanie by head tipping and cough threat or waa bark." Plooij's (1984) arm raise (ARA). Similar behavior is seen in bonobos (Kano, unpublished). Category 2? Raise arm slowly (RAS) (Type A): Extend arm overhead, in seeking to solicitgrooming. Seen in bonobos (Kano, unpublished). Category 2. Raise arm to hold branch (RAB) (Type A): Raise an arm and grasp a branch or liana. Solicit grooming. Seen in bonobos (Kano, unpublished). Category 2. Raise arm with elbow bent (RAE) (Type A): Extend arm overhead with bent elbow. Elicit grooming-hand-clasp at Mahale. Invitation of grooming-hand-clasp is often complied with by the same arm gesture, and results in grooming-hand-clasp. RAE is seen in bonobos. Category 2. Raise both arms, while bipedal (RTE) (Type B): Palms are directed to the companion. Intention movement to embrace an individual above the ground during play or stressful confrontation. In play, the performer is a youngster who shows play face, and in stress, the performer is usually an adult male who grimaces and the partner is above the ground. Category 3-6. Raise leg and hold foot with hand while seated (RHL) (Type A): Pattern of grooming solicitation. Seen in grooming and resting in bonobos (Kano, unpublished). Category 2. Raise leg while supine (RAL) (Type A): Lie supine and raise one leg. This solicits grooming of thigh or leg. Probably the same as Goodall's (1989) ventro-ventral present: "When the sittingsubordinate faces the dominant, leans back, raises one foot and rotates thighs laterally". Seen in bonobos (Kano, unpublished). Category 2. Raise other's arm (ROA) (Type A): Raise or lift other's arm. Groomer's initiativ to change the groomee's 168 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. posture, so as to change the body part to be groomed, and sometimes for groomer's convenience. Seen also in bonobos (Kano, unpublished). Category 2. Raise other's leg (ROL) (Type A): Same as Raise other arm (ROA) in function.Category 2. Rake (RAK) (Type A): Goodall (1989): "The sweeping of ground vegetation withstraight arm movements while showing a quadrupedal hunch. Usually this occurs prior to a chargingdisplay." Pattern is probably the same as the "Scratching/scrubbing dry leaves" by Mahale's adult males. Bonob rakes before dragging a Haumania vine in display (Kano, 1998). Category 2? Rap (RAP) (Type A): Nishida's (1997) Thump: Courtship pattern of both males and females: Sit and hit stem of shrub or branch or the ground with the knuckles of one hand. Category 7-8. Reach: See Extend. Reach hand: See Extend hand. Reach leg: See Extend leg. Reassurance suck: See Suck in reassurance. Reassure (REA) (Type D): Goodall (1989): "Response to submission... Includes , , , , and ." Kano (1998) states that although these behavioral patterns exist in the bonobo, they do not occur in reassurance, so, there is no reassurancebehavior in bonobos. Category 3? Reconcile (RCL) (Type D): Post-conflict behavioral patterns between combatants,such as , , , , and . The bonobos of Wambareconcile by , , , , , and (Kano, unpublished). See also de Waal (1982, 1989). Category 1. Redirection of aggression: See Aggress in redirection. Reject (RJT) (Type D): Goodall's (1989) Rejection of infant by mother: Maternal behavior "preventing her infant from (1) having access to the nipple, (2) climbing dorsal or ventral, (3) sharing her food or (4) sleeping in her bed", when the infant is being weaned. Behavioral patterns include (1) ", , , , (2) , , , , (3) , , , and and (4) " . Category 1. Reject-move (RJM) (Type C): Goodall (1989): "Mother avoids infant attempting to suckle, rideon her or beg food by moving away." Likely to occur in bonobos (Kano, unpublished).Category 2-5? Reject-sit (RJS) (Type C): Goodall (1989): "Mother sits when the infant is riding on her back and leans slightly back, so the infant can no longer ride. She may sit down the moment the infant climbs onto her back, or during travel..." Clear example unknown for the bonobo (Kano, 1998). Category 3-5? Relaxed face (RLF) (Type A): Goodall's Relaxed face. Mouth closed and eyes open in normal circumstances. Kano's (1998) Relaxed face. Category 1. Remove leaves (RML) (Type C): When streams begin to dry up, chimpanzees remove dry leaves and search for good drinking pool. With digging, this is an important technique to obtain water , . Displayed by at least five chimpanzees of Mahale. Category 3-8. Rest (RES) (Type B): Goodall's (1989) Rest: Remain immobile, or . Kano's (1998) Resting . Category 1. Retaliate (RTL) (Type D): Counter-attack, namely attack returned to the initialattacker . Goodall (1989): "Aggressive response directed toward the original aggressor by a chimp who has been threatened or attacked." Among the bonobos, retaliation occurs more often when females are attacked than when males are (Kano, 1998). Kano (1998) also called retaliation by a dominant individual to a challenging subordinate "punish attack". Category 1. Retreat (RET) (Type A): Walk backwards on all fours. When a subordinate pant-grunts to a dominant, the subordinate occasionally walks backwards in fearful way. Seen in bonobos. Cf. Creep. Category 2. Retrieve (RTR) (Type D): Nishida (1983a): Mother approaches and takes her infant away from the alloparent, when the infant begins to emit distress calls. Seen in bonobos (Kano, unpublished). Category 1. Return infant (RTI) (Type D): Alloparent brings back infant to its mother when the infant begins to emit distress calls. See Figure 8 of Nishida (1983a). Not seen in the bonobos of Wamba (Kano , unpublished). Cateogory 1. Reunite (REU) (Type D): Individuals meet after a separation. In such occasions, both friendly behavior (see Greet) and aggressive behavior (see Show off) occur. Category 1. Ride clinging (RDC) (Type A): Goodall (1989): "Infant clings to mother's arm or leg as she travels." Seen in bonobos (Kano, unpublished). Category 2. Ride dangling (RDA) (Type A): Goodall (1989): "Clings below mother with one or both hands only." Ethogram of chimpanzees 169

Kano's (1998) Dangling riding for the bonobo. Category 2. Ride dorsal (RDD) (Type A): Goodall's (1989) Dorsal travel: "When an infant rides on .the back of another. Three infant dorsal positions: Jockey, Prone, Quadrupedal and Supine." Kano's (1998) Dorsal travel for the bonobo. Category 1. Ride extended (RDE) (Type A): Goodall (1989): "Infant in a ventral position grips mother's hair with hands and feet but has arms and legs extended so that its back almost brushes the ground as she travels ." Seen in bonobos. Category 2. Ride jockey (RDJ) (Type A): Goodall (1989): "When an infant sits on the back of another in a straight upright position. Its legs may be drawn up, with the feet on the other's back, orgripping around the other's side." Kano's (1998) "Dorsal travel: jockey style" for the bonobo. Category 2. Ride prone (RDP) (Type A): Infant lies prone on the back of mother. Kano's (1998) "Dorsal travel: cling style" for the bonobo. See Figures 2 and 3 of Nishida (1983a) for photograph. Category 1. Ride quadrupedal (RDQ) (Type A): Goodall (1989): "When an infant stands on the back of another ." Kano's (1998) "Dorsal travel: quadrupedal style" for the bonobo. Category 2. Ride supine (RDS) (Type A): Infant lies supine on the back of mother. Occurs onl occasionally and briefly. Category 3-8. Ride ventral (RDV) (Type A): Goodall's (1989) Ventral ride: "The infant is transported as he clings to the mother's belly, gripping hair between flexed fingers and toes...There are followingvariations: Extended , Dangle, Arm or Leg-cling." Kano's (1998) Ventral riding for the bonobo. Category 1. Rinse (RIN) (Type A): Hold and shake object sideways repeatedly in the water. See Wash. Category 8. Rise (RIS) (Type A): Shift from lying posture to sitting or standing posture . Rising by a groomee stimulates the groomer who has stopped grooming to re-start. Category 2. Rock (ROC) (Type A): Goodall's (1989) Rocking: "Slight or vigorous side to sidemovements of the body when the chimp is sitting... Rocking occurs when a male is working up to a charging display..." At Mahale, rocking occurs not only before a charging display, but also when some individuals become nervous. Rocking occurs only in the context of courtship among the bonobo, not as warming up for a charging display (Kano, 1998). Category 2? Roll (RLL) (Type A): Goodall's (1989) Roll: "A chimpanzee may hit another and roll over him during aggression or play." Seen in bonobos in aggression (Kano, unpublished). Category1 or 2. Rough and tumble play: See Play in rough and tumble. Rub dorsum (RBD) (Type A): Lean against a straight tree trunk and rub shoulders or back against it, while often holding arms upward. See Figure 15 of Goodall (1968). May lie supine on a rock, the ground, grasses, horizontal bough, or day bed to rub themselves. See Figure 4 of Nishida (1980a). Seen in the bonobos of Wamba (Kano, unpublished). Category 2. Rub muzzle (RBM) (Type A): Rub muzzle stained with fruit juice against a shrub, grass stalk, slender vine, or bark of tree or stone in order to wipe off the mess. This is different from "wipe" because the chimpanzee puts the body part to a "part of the environment", not the tool to the body (Wipe) . This has been seen for only one adult female (Chausiku) who did so several times, in particular, after eating the seeds of Parkia filicoidea. Not described in Goodall (1989) but occurs at Gombe (McGrew, unpublished). Category 8. Run (RUN) (Type B): Locomote quickly. Category 1. Run bipedal (RUB) (Type A): Goodall's (1989) Bipedal run. Chimpanzees run on both feet during a charging display. Adult male (MA) once ran on both feet over ground swarming with Dorillus ants, apparently in order to avoid being bitten. See Plate 33 of Albrecht and Dunnett (1971) for photograph. Bonobos also run as bipeds (Kano, 1998). Category 1. Run quadrupedal (RUQ) (Type A): Goodall's (1989) Quadrupedal run. Category 2? Run tripedal (RUT) (Type A): Run using feet and one arm while other arm carriesobject. Sometimes, this is the same as Drag branch. Kano's (1998) Tripedal run. Category 2?

S Scan (SCN) (Type D): Pattern of Search: lost juvenile climbs a tree and looks around in all directions in order to find its mother. Adults do this too when looking for others. Bonobos show similar behavior (Kano , unpublished). Cf. Monitor. Category 2. Scavenge: See Eat carcass. Scoop (SCP) (Type A): Goodall(1989): "When the mother is about to travel and the infant is on the 170 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. ground near her rump she reaches back and pushes the child up onto her back with a backward and upward movement, using the palm of her hand and fingers." Kano's (1998) Scoop for the bonobo. Category 1. Scratch (SCR) (Type B): Rake partly flexed finger nails over surface. Goodall's (1989) Scratch. Seen as self-scratch in bonobos (Kano, 1998). Category 1. Scratch aggressively (SCA) (Type A): Scratch another animal in order to inflict wound. Goodall's (1989) Aggressive scratch: "A chimpanzee may scratch another during a fight." Probably occurs also in bonobos (Kano, unpublished). Category 1? Scratch dead leaves (SCL) (Type A): Scratch or rake with one or both hands the dead leaves that have been piled up on the ground, as an initiation of the charging display. Maybe the same behavior as Goodall's (1989) Scrub ("A component of charging displays. The chimp takes a bunch of vegetation, burlap nag and so on, and moves his hand in a semi-circular motion from side to side. Often in preparation for a charging display") or Rake. Absent in the bonobos of Wamba (Kano, unpublished). Category 3-5. Scratch self (SCE) (Type A): Goodall's (1989) Self-scratch: "Nails drawn deliberately across skin. In self-grooming, scratching is directed against the direction of hair growth and is followed by grooming of the scratched part... During social grooming the groomee often scratches a body part that is then groomed by the partner." Also occurs when a chimpanzee is frustrated, and feel uneasy. See Figure 12 of Nishida (1970) for photograph. Kano's (1998) Self-scratch for the bonobo. Category 1. Scratch self distantly (SCD) (Type C): Kind of Scratch self. Goodall's (1989) Distance scratch: "A grooming session sometimes initiated when one chimp sits, holds an overhead branch, and makes deliberate scratching movements while staring at the desired grooming partner who may be several meters away. Loud frustrated scratching may serve as a signal to a reluctant female who is being led away on a consortship, indicating that the patience of the male is wearing thin." Occasionally seen in bonobos in the beginning of grooming (Kano, unpublished). Category 2? Scratch socially (SCS) (Type A): Nishida's (1983a) Scratch or Rub: Scratch the skin of other's back vigorously during the beginning or middle of a grooming bout, usually with one hand. Making audible sound. Pattern is absent at Gombe. Nakamura and others' (1999) Social scratch. The bonobos of Wamba occasionally scratch another individual with the first and second fingers in grooming (Kano, 1998). Category 6. Scream (SRM) (Type A): Goodall (1989): "...high pitched and loud, almost always given in a series. Usually heard in contexts of aggression and general social excitement by highly stressed, fearful, frustrated, or excited individual..." Kano (1998) divided the bonobo scream calls into two, Scream and Screech. Category 1. Scream, food (SCF) (Type A): Uttered by some adult males when eating a prized food, such as the ripe fruit of Saba or Garcinia, especially during the beginning of the bout. Not mentioned in Goodall (1989). Bonobo's Food peep (Okayasu, 1991) or Food grunt (Kano, 1998) might be homologousto FSC. Category 2? Scrub pelt (SCC) (Type A): Scrub colobus skin against a rock in the streambed, apparently in order to soften it. Idiosyncratic to one adult male (Nishida, 1993). See Wash. Category 8. Search (SEA) (Type D): Look for unseen chimpanzees, by several methods: (1) sit and listen, (2) walk around, often returning to the original place, (3) sniff the ground, in particular, path intersections and footprints, and (4) climb tree and scan the environment, frequently employed by a lost juvenile who is whimpering. Not recorded in the bonobos of Wamba. Category 1?-7. Self-clasp: See Clasp self. Self-groom: See Groom self. Self-scratch: See Scratch self. Separating intervention: See Interfere to separate. Shake branch (SHB) (Type A): Goodall's (1989) Branch shaking: "A branch is shaken with quick jerky movements of the arm, slight or vigorous. The chimp may be sitting or standing..." Plooij's (1984) Branching (BRN). Nishida's (1997) Branch-shake: Courtship pattern of males and females: Sit while shaking horizontally stem of a shrub, branch of a tree, or a woody vine. Branch-shaking can be understood as a request for a partner to come closer or to leave together. Contexts include courtship, continuation of consortship (herding technique), and enticement to travel together. Kano's (1998) Branch-shaking and Ingmanson's (1996) Branch waving for the bonobos of Wamba. Category 2. Shake detached branch (SHR) (Type A): Detached, leafy branch is held in one hand or foot and shaken horizontally in courtship by several juvenile males. Category 8. Ethogram of chimpanzees 171

Shake hands: See Grasp hand. Shake penis (SHP) (Type A): Adult male grasps and shakes the penis of another vertically, while panting. Rare at Mahale. Category 8. Share food (SHF) (Type D): Allow another individual to take food that is under the control of the owner (for example, in the mouth or hand, on the lap, or in proximity). Same as Feistner and McGrew's (1989) Transfer of food. Meat is the most common food to be shared among adults. Food sharing is most common between mothers and infants. "Food share" (Kuroda, 1984; Kano, 1998) occurs in bonobos a Wamba and at Lomako (Hohmann and Fruth, 1993). See Figures 14a, b, and c of Nishida (1970) and Figure 8-8 of McGrew and others (1996) for photograph. Category 1. Show off (SHO) (Type D): Aggressive behavior that occurs between individuals meeting after a separation. Includes , , , , , and attacks. In reunion, bonobos show as aggressive elements (Kano, unpublished). Category 2. Show play face (PFA) (Type B): Goodall (1989): "Only slight retraction of the lip corners. Jaws may be closed or open... Occurs during play." Play face is seen among the bonobos (Kano, 1998). Category 1. Show play face full (PFF) (Type A): Goodall's (1989) Full play face: "Upper and lower teeth show as the play gets rougher and tickling more vigorous..." Plooij's (1984) Place face full (PFF). Seen in bonobos (Kano, unpublished). Category 1. Show play face half (PFH) (Type A): Plooij's (1984). Goodall's (1989) Low play face: "Only the lower teeth show." See Figure 7 of Nishida (1983a). Status unknown in bonobos. Category 1-3. Shrug (SHG) (Type A): Goodall (1989): "Mother raises one shoulder slightly and lowers the other. Signals the infant to climb off its dorsal position. Gesture occurs during weaning and is a sign of rejection." Not confirmed for the bonobo (Kano, 1998). Category 3-5. Sit (SIT) (Type A): Plooij's (1984) SIT. Goodall (1989): "Rump on ground, branch, and so on, body more or less vertical." See Figures 14a, c, d, and g of Goodall (1968) for illustration. Kano's (1998) Sit. Category 1. Sit and lean (SIL) (Type A): Sit and extend both arms backwards and lean against them. Not confirmed for the bonobos. Category 1-3? Sit and turn back (SIB) (Type C): Pattern of soliciting grooming. Seen in bonobos. Category 2. Sit behind individual (SIH) (Type C): Change position in order to groom back of companion. Seen in bonobos (Kano, unpublished). Category 2. Sit face to face (SIF) (Type C): Express the intention of social grooming or seeking shared food. Similar behavior is seen in bonobos (Kano, unpublished). Category 1-2. Sit prone (SIP) (Type A): Slouch or bend forward, extend both arms, and put hands on the ground. Often when being groomed. Seen in bonobos (Kano, unpublished). Category 2. Sit sideways (SIS) (Type C): Pattern of solicitation of grooming. Seen in bonobos, too. Category 2. Sitting hunch: See Hunch and sit. Slap (SLA) (Type B): Plooij (1984) 's SLA. Goodall (1989): "A hitting down gesture when the palmar surface of the hand is brought into contact with the objective..." Bonobos slap in aggression, charging displays, drumming and social play (Kano, 1998). Category 1. Slap buttress/tree trunk (SLB)(Type A): Performed with one hand or alternate hands during charging display. Goodall's (1989) Drum. Included in Kano's (1998) Drum for the bonobo. Category 2. Slap ground (SLG) (Type A): Done with one hand or alternate hands during a charging display. May slap ground with pant-hoot in response to pant hoot calls from a distance. Recorded from Beni (Kortlandt, 1964). Bonobo slaps the ground while staring at the opponent (Kano, 1998), so the context is more narrow. Category 2. Slap in invitation (SLI) (Type D): Nishida's (1983a) Invitation slap. Repeatedly slaps a rock or ground with one hand in order to entice youngster into play. Elements include: , , . Absent in the bonobos of Wamba. Category 3-5? Slap other (SLO) (Type A): Done during play, charging display or fighting. Category 1. Slap-stamp (SLS) (Type A): Element of charging display. Goodall (1989): "Slapping with hands and stamping with feet together." Not seen in bonobos of Wamba (Kano, 1998). Category 3-5. Slap wall (SLW) (Type B): Slap wall of metal house with one or both hands or alternate hands, while standing on both feet or on all fours. Performed regularly by M group adult males at Mahale and by Kasakela males at Gombe. See Figure 3 of Nishida (1994) for photograph. Category 7. 172 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M.

Slide down (SLD) (Type A): Goodall's (1989) Fireman slide: "A chimp may slide down a vertical or diagonal pole or tree in an upright or crouched position. All four limbs are in contact with the pole or tree." Kano's (1998) Vertical slide. Category 1. Sloth position: See Hang in sloth position. Smack lip (SLP) (Type A): Goodall's (1989) Lip smack: "Occurs during grooming. Mouth slightly opened and closed rhythmically. As the mouth is opened, there is a smacking sound when the tongue is drawn across the palate." Plooij's (1984) LIP. Similar behavior was recognized for the bonobo, but it is unknown whether it is homologous to SLP or not (Kano, 1998). Category 2 or 3. Smell: SeeSniff. Snatch (SNA) (Type C): Take forcibly an object such as food from another individual without consent. Adult males snatch food, meat in particular, from individuals of any age-sex class, mothers from immature offspring, and juvenile sons from mothers. In bonobos, SNA is mostly done by juveniles (Kano, 1998).Category 1. Sneer (SNR) (Type A): Goodall (1989): "The upper lip is retracted, sometimes on one side more than the other, to expose part of the upper teeth. In the wild the context is associated with fear of a human or a human associated situation. Only afew chimps sneer." At Mahale also only afew individuals sneer. Category 8. Sneeze (SNZ) (Type A): Goodall (1989): "Chimps sneeze in the same way as humans." Kano's (1998) for the bonobo. Category 1. Sneeze with stick (SNS) (Type A): Push stick into nostril to stimulate sneeze,which discharges a large amount of nasal mucus. Only one adult male showed this type of tool-use. See Nishida and Nakamura (1993). Category 8. Sniff(SFF) (Type A): Plooij's (1984)SFI. Goodall's (1989) Sniff. Put nose close to something in the environment, such as the ground, vegetation, tree trunk, fruit, food wadge , feces, urine and so on. apparently to obtain information on which chimpanzees were there or what kind of animal was there . Holding in the hand and sniffing a large hard-shelled fruits such as Saba florida and Voacanga lutescens is an important way to judge their ripeness. Chimpanzee often sniffs the ground after being left behind by companions, or sniffs the ground when arriving at a place where strange chimpanzees had been immediately before. In bonobos, sniffing is a predominantly juvenile behavior (Kano, 1998). Category 1. Sniff finger(SFI) (Type A): Plooij's (1984)SFI. Goodall (1989): "One finger , usually index finger, is carefully smelled after being used to investigate some object. The most usual context is during inspection of the genital area... of a female." Shown also by juvenile bonobos (Kano, 1998). Category 1. Sniff with tool (SFT) (Type A): Goodall's (1989) Investigation probe: "A stick or twig is used by a chimp for touching some object..." Not observed among the bonobos of Wamba (Kano, 1998). Category 3-5. Snub (SNB) (Type C): Apparently deliberate refusal to look at a conspecific that is approaching closely to seek contact. At Mahale, an alpha male, Ntologi, did not look at the beta male, Nsaba even if Nsaba continuously followed Ntologi apparently seeking to groom him. Apparently a kind of intimidation signaling that the alpha male does not wish to interact with the beta. (Cf. Ignore). Category 7-8? Solicit copulation (SOC) (Type D): Courtship. Show gesture and posture directed to an estrousfemale by a male or vice versa that normally leads to copulation. Goodall (1989) includes only male behavior toward an estrous female. Goodall lists , , , , , , and , , , and . At Mahale, , , , , and so on. also occur (Nishida, 1997). Among the bonobos of Wamba , , , , , and have been recorded as courtship display for copulation (Kano, 1998). Category 2. Solicit grooming (SOG) (Type D): Van Hooff's (1974) Groom-present . Goodall's (1989) Present for (solicit) grooming: "The rump, back, or bowed head are the most usual body parts presented to a chosen grooming partner at the start of a session..." Alter the position of limbs, head or body part tofacilitate the continuation of grooming. Self-scratching is the most important of 4 patterns of solicitation for grooming in the bonobos of Wamba (Kano, 1998). Category 2. Solicit grooming turn (SGT) (Type D): Signal to ask partner to take turn of grooming: (1) stop grooming and release the partner's body, (2) stop grooming, and scratch or groom self, (3) stop grooming and present the back or other body part to the partner. Similar behavior is shown among the bonobo (Kano, 1998). Category 2. Ethogram of chimpanzees 173

Solicit play (SOP)(Type D): Invite another individual to play. Consists of , , , , , , , and others. Bonobos invite play by approach with play face, hitting lightly or pulling the hand of partner (Kano, 1998). Category 2. Solicit reassurance contact (SOR)(Type D): Aggressed-against or frightened infant runs to its mother, whimpers with extended hand or throws temper tantrum. Aggressed-against adults seek contact with a third party or with the aggressor by extending a hand, and so on. Goodall (1989) mentioned, but did not name, this behavioral pattern. Kano's (1998) Request for appeasement. Aggressed-againstbonobo female pesters the aggressorfemale and peers at her persistently so that the aggressor invites the victim into GG-rubbing (Kano, 1992). Category 1-2. Solicit support (SOS) (Type D): Request the support of third party when one is being threatened or attacked. Includes patterns such as , , , , , , , , and so on. At Wamba, bonobo solicits only mother for support (Kano , unpublished). Category 1. Somersault (SOM) (Type A): Goodall (1989): Roll "head over heels". Usually repeated many times. At

Mahale, 3-4 year old infants often engage in this pattern of self play when the whole group travels a long distance. See Figure 7e of Goodall (1968) for illustration. Absent in the bonobos of Wamba. Category 1?

Spar (SPA) (Type A): At the beginning of fight, both parties stand bipedally and flail one or both of their arms in hitting movement, with or without screaming. Usually do not really hit. Similar movement is shown during play. Bonobo juveniles hit each other during play (Kano, 1998). Category 2.

Spit seed (SPS) (Type A): Chimpanzees occasionally spit large seeds such as those of Garcinia huillensis and Voacanga lutescens at the spot where they are eating. Seen in bonobos. Category 1.

Squat (SQT)(Type A): Hunt and others' (1996): "The body weight is borne solely by the feet/foot, both hip and knee are strongly flexed." Seen in bonobos. Cf. Sit. Category 1. Squeal in copulation (SCO) (Type A): High-pitched, scream-like sound accompanied by a grin , emitted by an estrous female at the final stage of copulation. Goodall's (1989) Copulation scream (previously called a squeal). See Figure 11-1 of McGrew and others (1996) for photograph. Similar call is emitted by bonobos (Kano, 1998). Category 2.

Squeeze (SQZ) (Type A): Hold object such as fruit and pith in mouth and extract juices by firm pressing of lower and upper palate, tongue and lips. Sometimes, a hand is pressed against lower jaw to facilitate extraction. See also Wadge. Seen in bonobos. Category 2.

Stamp (STA) (Type B): Goodall (1989): "Forceful downward kick of one foot, or alternate feet... (with sole making contact)." She also states that as a component of charging displays that "The chimp stamps his feet one after the other as he runs along. One foot is often stamped harder than the other. Astanding chimp may stamp alternatefeet... A sitting chimp may stamp several times with one foot." VanHooff's

(1974) Stamp. See Figures 13a, and b of Nishida (1970) for the photograph of stamping on the buttress, and Figure 4 of Nishida (1994) for the photograph of stamping on the colobus skin. Bonobos stamp while running, but not while sitting (Kano, 1998). Category 2.

Stamp bipedal (STB) (Type A): Stamp one's sole on the ground, buttress, tree trunk, wall of house or another individual while upright. Component of charging displays. Some adolescent males stamp with alternate foot on the ground in courtship display. Bonobos of Wamba stamp bipedal in courtship, but not in charging displays. Category 2-3?

Stamp in invitation (STI) (Type A): Nishida's (1983a) Invitation stamp. Mainly youngsters, lightly stamp on the ground with one foot while watching potential playmate, in invitation to social play or alloparental care. May be bipedal or quadrupedal. Absent in bonobos of Wamba. Category 3-7.

Stamp other (STS)(Type A): Goodall (1989): "Stamping on a victim is a major component of within-species fighting, and can result in severe bruising and contusions." Seen in bonobos. Category 2.

Stamp quadrupedal (STQ) (Type A): Stamp on all fours on the ground, buttress, tree trunk, and wall of house or another individual. Component of charging displays and so on. Absent in bonobos? Category 3?

Stamp trot (STT)(Type A): van Hooff (1973): "The animal walks or trots along while stamping heavily with its hind feet on the ground. Often this is done in a rhythmical manner in that one foot is placed down gently and the other with force. The head is mostly kept rather low and tucked back between the shoulders which are pulled upwards. The head may make low amplitude vertical rocking movements•@ " Kano's (1998) Stamping run for the bonobo. Category 2.

Stand bipedal (SDB) (Type A): Plooij's (1984) STB. Stand upright. Stance assumed when watching a 174 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. distant object or taking food from tall shrubs such as Ficus urceolaris. (cf. WAB). See Figure 4 of Nishida (1970) for photograph. Bipedal posture occurs in courtship, when mixed with other movement such as , , , and . Commonly seen in bonobos. Category 1. Stand in head down, bottom up (SHD) (Type A): Nishida's (1983a) Head down hip up: A postural pattern with tickling by mouth. "The mother or caretaker stands on all fours, with elbows flexed, and putting its mouth against the infant's body on the ground. The open mouth may be pressed against any part of the infant's body, but especially its belly, back, head, neck, foot or arms." Similar to "Fore-limb crouch" (Figure 8B of Hunt and others, 1996). Absent in bonobos? Category 3? Stand quadrupedal (SDQ) (Type A): Stop walking in order to scan the environment or wait for another and so on.Plooij's (1984) STQ. Commonly seen in bonobos. Category 2. Stand quadrupedal heel up (SQH) (Type A): When waiting for a companion, or monitoring the environment, performer stands still with heel up and toes on the ground. Kano's (1998) Waiting posture for the bonobo. Category 2. Stare fixedly (STF) (Type A): Goodall's (1989) Fixed stare: "The chimp stares intently at the individual with whom it is interacting..." At Mahale, groomee often stares fixedly at groomer during intense social grooming. Other contexts include threat, predation, and monitoring. Kano's (1998) Stare for the bonobo, who stare mutually during mating. Category 1. Startle flee (SFL) (Type A): Goodall (1989): "When a resting chimp is suddenly startled by an unexplained sound he or she may run to climb the nearest tree, or at least hold the trunk ready to climb. One chimp in a resting group who behaves thus will trigger the same reaction in most or all of the others." Same pattern is seen among the bonobos (Kano, 1998). Category 2. Startle flinch: See Parry. Steal (STL) (Type C): Remove an object surreptitiously, without the owner's knowledge. This is a part of Goodall's (1989) Steal, which includes both Snatch and Steal. Seen in bonobos (Kano, unpublished). Category 1. Stem pull-through: See Pull through Stir water (STW) (Type A): Stir water vigorously with one hand in a pool of a river. Juveniles do this when playing by themselves. Category 8. Store (STO) (Type A): Keep leaves in one hand and begin to chew only after collecting a handful of them. This often occurs when chimpanzees eat the hard leaves of Ficus exasperata. Category 3-5. Strip leaves: See Pull through. Stroke (STR) (Type A): Stroke theface of another with fingers and palm. Rare behavior of unknown function. Category 8. Suck (SKK) (Type A): Take nipple in mouth and suck. Goodall's (1989) Suckle (sic). Kano's (1998) Suckle (sic) for the bonobo. Category 1. Suck in reassurance (SKR) (Type C): Goodall's (1989) Reassurance suckle (sic): "A brief suck when the infant has been frightened or hurt". Not observed in the bonobo (Kano, 1998). Category 3? Suck self (SKS)(Type A): Suck from own nipples. Rarely, a female who has lost her newborn infant sucks from her own nipples. Matsumoto-Oda's (1997) Self suckle. Category 8. Suck thumb (SKT)(Type A): Infant sucks its own thumb. Category 1? Suckle (SUC)(Type A): Mother allows her infant to suck from her nipples. Goodall's (1989) Suckle. Kano's (1998) Suckle for the bonobo. Category 1. Supplant (SPL)(Type D): Drive off another chimpanzee from a food patch by , , or . Adult males usually supplant adult females and other adult males. In bonobos, adult males are supplanted by adult females, or other adult males (Kano, 1998). Category 1-2. Support (SUP) (Type D): Intervene in a fight and side with either of the conflicting parties. Rare in bonobos. Category 1. Support dominant (SUD) (Type D): Intervene in a fight and side with dominant party (cf. SUS). De Waal's (1982) Winner support. Category 1? Support older (SUO) (Type D): Nishida and Hosaka (1996): Intervene in a fight and side with older party . Often an intervention tactic by dominant male in fight between young adult and past-prime adult males. Category1? Support subordinate (SUS) (Type D): Intervene in a fight and side with a subordinate party. De Waal's (1982) Loser support. (cf. SUD). Category 1? Swagger bipedal (SWB)(Type A): Goodall's (1989) Bipedal swagger: "In an upright or semi-upright Ethogram of chimpanzees 175 posture the chimp sways, often rhythmically, from one foot to the other. The animal may remain in one spot to swagger or move forward during the swagger. The arms are normally held out from the body, the shoulders hunched up. It is often used during aggression, greeting, and courtship." Van Hooff's (1973) Sway walk. Plooij's (1984) Bipedal swagger (BIS). At Mahale, this is often performed as the separating intervention just before a charging display. Similar behavior occurs in the bonobos of Wamba (Kano , 1998). Category 2. Swallow leaves (SWL) (Type A): Wrangham and Nishida's (1983) Leaf-swallow. Swallow leaves of particular species of trees and herbs slowly and without chewing, usually in the early morning. So far, the leaves of Aneilema spp., Aspilia mossambicensis, Commelina spp., Ficus exasperata, Lippia pulicata and Trema orientalis have been seen to be swallowed at Mahale. See Figure 1 of Nishida (1994) for photograph. Known also from Tai (Boesch, 1995), Kibale, Bossou and other localities (Huffman, 1997) in addition to Gombe and Mahale. Species of swallowed leaves differ from place to place. Likely to be a culture universal in chimpanzees. Category 3? Sway and move (SWM) (Type A): Arboreal movement: Climb high up, sway the tree trunk back and forth using the body weight, and catch and pull the terminal branch of another tree when the original tree is bent. Kano's (1998) Sway tree/branch for the bonobo. Category 2. Sway woody vegetation (SWV) (Type A): Component of charging display. Usually standing bipedally, vigorously sway back and forth with both hands on the trunk of small tree, large horizontal branch or fallen trunk of a tree. Probably the same as Goodall's (1989) Branch sway: "The chimp vigorously sways a growing branch usually when standing on one'sfeet. During a charging display, a male may pause and sway branches." Plooij's (1984) Branch-sway. Absent in the bonobos of Wamba (Kano, 1998). Category 3-5. Swing (SWN) (Type A): Hang from woody vine by arms and swing back and forth. Pattern of charging display. If the performer changes from one support to another, it is called Brachiate. Thus, Goodall's (1989) Swing is equivalent to brachiation. Kano's (1998) Swing for the bonobo. Category 1. Swing and drum (SWD) (Type B): Hang from woody vine, swing, and kick buttress of tree forcibly with feet, taking advantage of the impetus of swinging movement. Absent in bonobos of Wamba (Kano, unpublished). Category 3?

Tag (TAG)(Type B): Play tag. Alternate chase and flee. See Nishida (1983a). Includes , , and so on. Seen in bonobos (Kano, unpublished). Category 1. Take (TAK) (Type C): Goodall (1989): "When a begging chimpanzee takes food with the owner's knowledge and presumed consent." See Figures 14a, b, c, d of Nishida (1970) for photograph. Cf. Snatch and Steal. Seenin bonobos (Kano, 1992). Category 1. Take finger in mouth (TFM) (Type A): Nishida (1983a), Goodall (1989): "reach for the hand/foot of another and place part of hand/food or fingers/toes in its own mouth." Like PFM (Push finger into mouth), this is performed as a reassurance behavior. TFM is seen much less often than PFM at Mahale. Not seen for the bonobos of Wamba (Kano, 1998). Category 3-5. Teach (TCH) (Type C): At Mahale, teaching as defined by Caro and Hauser (1992) consists only of discouragement (Nishida, 1987). Chimpanzee mother or allomother rarely removes and discards a leaf held in the mouth by an infant if the leaf is not in the diet of the group (Nishida, 1983a; Goodall, 1986). Doubtful that this is systematic teaching. Teaching is not recorded in the bonobos of Wamba. Category 3-8. Tease (TEA)(Type D): Adang's (1984) Tease. Mahale adolescent males pester and threaten adult females by , , and , , and so on until femalespant-grunt to them. Adolescent or juveniles throw branch at another adolescent or juvenile apparently to tease him/her. Once an adolescent female with a monkey tail "tried to fish" for a juvenile male: She carried a colobus tail and climbed a tree with it and dangle it from above, and whenever the male jumped fo it, she withdrew it so that he could not reach it. After several trials, the male finally got the tail, and the female grimaced, suggesting that she was just trifling with the male, without any intent of giving the tail. Bonobo adolescent male at Wamba pesters a particular adult male, shows threatening gesture such as bipedal swagger or branch-dragging, and as a result may dominate him, which is termed as "adolescent harassment" by Kano (1998). Category 1 or 2. Temper tantrum: See Throw temper tantrum. Thigh support: See Transport with thigh support. Threaten (THR)(Type B): Intention movement or preparatory gesture of aggression. Goodall's (1989) 176 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M.

Threat:"•@ a repertoire of gestures to elicit submissive behavior in the individual the gestures are directed toward... The repertoire includes , , , , , , , , , , , , , , , and ". Kano (1998) lists similar threatening: "stare", "head-tip", "wrist-shake", "slap", "branch-sway", "arm-raise", "threatening approach", and "pseudo-charge". Category 1-2.

Throw (THO) (Type B):Plooij's (1984) THO. Goodall (1989): "Objects, such as stones , rocks, branches, sticks, handful of grass, and so on. may be thrown under arm or overarm..." Bonobos also throw , but underarm only (Kano, 1998). Category 1.

Throw at (THA) (Type B): Goodall (1989): "When an object is aimed at a specific objective..." Goodall also uses the term "aimed throwing". Adult males of Mahale sometimes throw with one arm a heavy stick or rock against the wall of a metal house to make a loud noise. Absent in bonobos (Kano, unpublished). See also Throw splash. Category 3-5?

Throw branch (THB)(Type A): Branch or stick is thrown in an upright posture with one arm . (1) Element of a charging display by both males andfemales, and (2) Courtship pattern to an estrousfemale by juvenile/older infant males. Older juvenile male once threw a stick as long as 1 m from a distance of 8 m while he was pant-grunting at an adult male. Bonobo throw branch forward at the end of the branch-dragging display (Kano, 1998). Category 2. Throw dry leaves (THD) (Type A): In the dry season, male chimpanzees of Mahale run , pick up a pile of dry leaves from the ground, and throw it with one hand at the climax of the charging display . Not used in courtship. Absent in bonobos of Wamba. Category 7.

Throw sand (THS) (Type A): Pattern of the male charging display. One adult male (Fanana) throws sand with one hand while running bipedally. Within the M group's range the ground is rarely covered with sand.

Category 8. Throw splash (THW) (Type A): Special case of Throw stone/rock. Fully adult male of M group lifts and throws a heavy rock into a stream (aimedthrowing), which produces loud splash that intimidates others

(see Nishida, 1993). Category 7. Throw stone/rock (TSR)(Type A): Element of the charging display. Not used the courtship. Bonobos of

Wamba throw only branches (Kano, 1998). See Throw splash. Category 3? Throw temper tantrum (TTT)(Type B): Goodall (1989): "The chimpanzee screams loudly , and may leap up, fling arms above his/her head and then slap them onto the ground or beat the ground with his hands . He may hold himself to the ground on his face, hug a tree, or himself. He may rush off , tumbling over and over, still screaming. The screaming often results in glottal cramps..." Typically occurs amonginfants during weaningconflict, when motherrejects sucking,food sharing, transport, grooming and soon., however, even adults sometimes show in response to rejection by dominant individuals . At Mahale, an extreme case was that of an infant hanging and swinging withfeet as if it were about tofall headfirst from high above the ground. By temper tantrums, an infant not only protests, but appears to monitor how much it can still garner the care of a rejecting mother (see Monitor mother). Similar behavior is seen among the bonobos (Kano, 1998). Category 1. Thrust (THU) (Type A): Plooij's (1984) THU. Goodall's (1989) Pelvic thrust: "Rhythmic back and forth movements of the pelvis... Occurs during copulationand...in reassurance mounting . Thrusting commonly occurs in play between infants." Two adult males embrace ventro-ventrally and bipedally in the context of reassurance or reconciliation, and one of the males may thrust against another. Adultfemale also occasionally mounts another female and shows a pelvic thrust. Bonobo males thrust like chimpanzees, but the female thrusts right and left in GG-rubbing (Kano, 1998). Category 1.

Thrust bipedal(TBI)(Type A): Stand upright on one's feet and thrust in non-mating context . Courtship shown by a few adult and adolescent males. Category 8.

Thump: See Rap. Tickle (TIC) (Type B): Plooij's (1984) TIC. Goodall (1989): "The chimp puts one or both hands on the body of the partner, usually between the neck and shoulder or in the groin, and makes tickling movements with the fingers." See Figure 7 of Nishida (1983a) for photograph. Bonobos may touch, push lightly , hold, or grasp an infant or a juvenile in social play, which may be equivalent to chimpanzee tickling (Kano,

1998). Category 1. Tickle with hand: See Tickle. Tickle with mouth: See Mouth. Ethogram of chimpanzees 177

Tip head (TPH)(Type A): Goodall's (1989) Head tip: "---a threatening gesture. Head is jerked very slightly backwards, at the same time chin is raised. The performer faces the individual being threatened and the gesture is usually accompanied by a cough bark and often an arm raise." Seen in the bonobo (Kano, 1998). Category 2. Tool use: See Use tool. Touch (TOU)(Type A): van Hooff's (1973) Touch. Plooji's (1984) TOU. Nishida's (1983a) Touch. Goodall (1989): "A chimpanzee reaches out with a hand (or occasionally foot) and touches another, with fingers (or toes), by laying the whole palmar surface of the hand on the other's body..." Touch signals reassurance to a subordinate from a dominant, and appeasement to a dominant from asubordinate . At Mahale, some estrous females approach and touch males on the shoulder, and so on, to solicit copulation. To touch the head of another is sometimes a signal to the groomee by the groomer to change posture. Parts frequently touched are mouth, chin, head, hand, shoulder, and genitals. The mouth was most often touched by an adult male (Kalunde) in order to appease the then alpha male (Fanana). See Figure 9 of Nishida (1970) for photograph, in which an adult male touches the mouth of a juvenile female and lets her bite his hand in order to reassure her. See also , , , and . Adult bonobos touch each other much less since Kano (1998) mentions that touching is predominantly byinfants and juveniles. Category 1. Touch fruit (TOF)(Type A): Touching hard-shelled fruit is an important way to get information about the ripeness of the fruit, in addition to. Absent in the bonobos of Wamba. Category 3-6. Touch scrotum (TOS) (Type A): A pattern of reassurance behavior. When mounting occurs, the mounted male often reaches back to touch the genital area of the mounter with a hand, while screaming loudly. While pant-grunting to an adult male, adult female often reaches to touch the scrotum. Kortlandt's (1964) Caress another's scrotum for the chimpanzees ofBeni. Recorded for humans in New Guinea (Eibl-Eibesfeldt, 1972). Category 1-5. Transfer (TRS)(Type B): Permanent migration from one unit group to another (Nishida and Kawanaka, 1972). Females normally transfer from their natal group to another during adolescence (Nishida and others, 1990). No sexually mature males have been known to transfer at Mahale. Thus, chimpanzees are "patrilocal" (Nishida , 1979). This is also the case for Gombe (Goodall, 1986) and Tai (Boesch and Boesch, 1999) and the bonobos of Wamba (Kano, 1992). Rarely, mother with juvenile offspring also transfers (Nishida and others, 1985; Takahata and Takahata, 1989).Cf. Visit. Category 1. Transport (TRP) (Type B): Nishida (1983a): Carry infant on the belly or on the back on all fours. Carry food between lips, in hand or foot, on the back or head, in the groin pocket or neck pocket, or drag food by one hand. Bonobos also transport food similarly (Kano, 1998). During play, juvenile chimpanzees often transport objects such as twig, fruit, ant-nest and so on. McGrew (1992) called the latter Play-start. See Object play. Category 1-2. Transport bipedal (TRB)(Type A): When carrying a lot of food, an individual often walks on both feet . Bonobos often carry sugar cane bipedally (Kano, 1998). See photo 4 and 7 of Nishida (1968). Category 1. Transport corpse (TRC) (Type B): Mother carries the dead body of her infant by hand, neck or groin pocket, or on her back. May persist up to 3 months, and the corpse becomes a mummy. Known from west (Matsuzawa, 1997) and central (Kuroda, 1998) Africa in addition to Gombe and Mahale. See Figure 22 of Goodall (1968). Seen in bonobos (Kano, 1992). Category 2. Transport food (TRF) (Type B): Transport of food occurs when: (1) individual (in particular, youngster) wants to follow a companion who has already eaten and departed, (2) individual (in particular, adult male) wants to eat in a safe position in a tree, and (3) individual wants to eat in the shade. See Ride for transport of infants. Bonobos also carry food (Kano, 1998). Category 1. Transport in groin-pocket (TGP) (Type A): Goodall's (1989) Groin pocket. Kano's (1998) Groin pocket for the bonobo. Category 2. Transport in mouth (TRM)(Type A): Hold and carry an object in mouth (between teeth). Chimpanzees hold vines or twigs as materials of fishing tools in mouth and transport to fishing sites. Hold food such as leafy twigs or fruit in mouth during transportation. Bonobos carry objects in the mouth (Kano, 1998). Category 1. Transport in neck-pocket (TNP)(Type A): Goodall's (1989) neck-pocket, Plooij's (1984) ONP. Rare in the bonobo (Kano, 1998). Category 2. Transport on head (TOH)(Type A): Plooij's (1984) OOH. One mother (Chausiku) regularly carried her newborn on her head. Category 8. 178 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M.

Transport quadrupedal (TRQ) (Type B): Transport object with one hand, in mouth, neck pocket, on shoulder or on head, while walking on all fours. Category 2. Transport two offspring (TTO)(Type B): Mother simultaneously carries juvenile offspring on her back andinfant offspring on her belly, or rarely carries both of them on her back. More likely to occur in dangerous situations such as crossing a large river. Bonobo mothers seem to carry two offspring more often than the chimpanzee mother, because the birth interval is shorter (Kano, 1998). Category 1. Transport with hand support (TRH)(Type A): Goodall's (1989) Hand support. Mother supports infant's back with one hand. Aninfant newborn orenfeebled by wound/disease is typically carried in this way. Kano's (1998) Hand support for the bonobo. Category 1 or 2. Transport with thigh support (TRT) (Type A): Goodall's (1989) Thigh support: "Method of supporting infant especially newborn, where the mother walks or runs with her thighs flexed, thus providing support for the infant's back. Usually the mother moves with rounded back and takes short steps." Kano's (1998) Thigh support for the bonobo. Category 2. Travel (TRV) (Type D): Goodall (1989): "Moving between A and B." Include , , , , , , , , and . Kano's (1998) Travel for the bonobo. See also Susman and others (1980) for bonobo locomotor patterns. Category 1. Travel alone during childbirth (TRA)(Type D): Maternal leave. Days or weeks after parturition, a mother with a newborn baby tends to spend time alone, apart from other group members, probably avoiding harassment by them. At Wamba, mother bonobo is often accompanied by one or two males directly after she has given birth (Kano, 1998). Category 1 or 2. Tug of war: See Pull from opposing sides simultaneously. Turn around (TUR)(Type A): Turn one's back to an approaching individual (for example, to beg). Goodall's (1989) Turn away: "Mother may turn from her infant to prevent suckling or begging. Adults may also turn their backs on beggars." Kano's (1998) Turn back for the bonobo. Category 1. Turn face away (TFA)(Type A): Mother turns her face away from her infant who is begging. Kano's (1998) Turn away for the bonobo among which TFA is often seen in a similar situation. Category 1. Turn face downward (TFD)(Type A): Drop one's head as a signal to request grooming of the head. Seen in bonobos. Category 1-2. Turn face upward (TFU)(Type A): Incline head backward. Groomee often does this when being groomed on the face. Seen in bonobos. Category 1-2. Turn round: See Look back. Twist (TWI)(Type A): Wrench. When a begging chimpanzee finds it difficult to detach a piece of meat by pulling from a carcass possessed by another chimpanzee, twists the bone so that a piece of carcass may be detached. Not mentioned in Goodall (1989). Category 8?

U Urinate (URN)(Type A): Category 1. Use tool (UST)(Type D): Extract an object from the outer environment and use it with certain purpose. Mahale chimpanzees' repertoire includes , , , , , , , , , , , and . Great local variation exists in tool-use (see McGrew, 1992 and Sugiyama, 1997 for review). Tool using repertoire of bonobos is limited: , , and at Wamba (Ingmanson, 1996; Kano, 1998). Category 1-8.

V Ventral riding: See Ride ventral. Ventro-ventral embrace: See Embrace full. Vertical climb: See Climb vertical. Vertical leap: See Leap vertical. Visit (VST) (Type D): Temporary migration from one unit group to another. Adolescent and young nulliparous adult females may visit one of the neighboring groups when they are in estrus, and return to their natal group when their sexual skin shrinks. Reported from Gombe (Goodall, 1986) and Tai (Boesch and Boesch, 1999), in addition to Mahale (Nishida and others, 1990). Different from Transfer, which is permanent. Category 1-2. Ethogram of chimpanzees 179

Vomit (VOM)(Type A): Plooij's (1984) KOT. Category 1.W

Waa bark (WAA) (Type A): Goodall (1989): "Loud, sharp sound given in a variety of agonistic contexts. It is usually accompanied by an arm threat or more vigorous gestures." Category 3? Wade (WDE)(Type A): When crossing a river, chimpanzees avoid contact with water and use stepping stones if available. If not, they walk quadrupedal in the river so long as the river is shallow. Category 1-7? Wadge (WDG)(Type A): Goodall's (1989) Wadge: Extract juice by chewing and compressing sticky fruits, their seeds and skin with leaves, or meat with leaves "between the lower lip andteeth", or "between tongue and palate" . Honey, eggs and even semen may be wadged. After juice is extracted, the remaining inedible parts are spat out. Fruits frequently wadged at Mahale include Canthium crassum, Harungana madagascariensis, Parinari curatellifolia, Pseudospondiasmicrocarpa,Psychotria peduncularis, Syzigium guineense, Uapaca kirkiana and Uapaca nitida. Bonobos also wadge (Kano, 1998). Category 2. Wait (WIT)(Type D): Goodall (1989): "An individual sets off, then looks back at a companion. If the latter is not following, the first stops to wait..." Ntologi (the alpha male) often waited for other adult males to follow. Also at the ant-fishing and drinking sites which are limited in number. Bonobos wait in similar contexts (Kano, 1998). Category 1. Waiting posture: See Stand quadrupedal heel up. Walk bipedal (WAB)(Type A): Goodall's (1989) Bipedal walk. Plooij's (1984) WAB. Walk upright when chimpanzees have lots of food in the arms, walk on muddy terrain, or initiate a charging display. Susman and other's (1980) Bipedal walk for the bonobo. See Figure 2 of Goodall (1968) and de Waal (1995, p.64) for illustration or photograph. Category 1. Walk quadrupedal palmigrade (WQP)(Type A): Quadrupedal walk with open palm on the slender horizontal branch. Susman and others' (1980) Palmigrade quadrupedalism. Category 2. Walk quadrupedal with knuckle (WQK)(Type A): Goodall's (1989) Knuckle walking, Plooij's (1984) WAQ. Quadrupedal walk with knuckles on the ground or large horizontal bough. See Figure 6 of Nishida (1994). Kano's (1998) Walk. Category 2. Walk tripedal (WQT)(Type A): Walk with object in one hand. Category 2. Wash (WAS)(Type B): Seen only as "Wash the colobus skin" below. Wash means the whole process of ,, , and . Category 8. Wash colobus skin (WAC) (Type B): Old adult male (Musa) once dunked a colobus pelt into running water, kept it immersed for a while, rinsed it in the water, brought it out, stamped on one's feet it on the rock repeatedly and repeated the whole process several times. After that, Musa nibbled the skin (Nishida, 1993). See Figure 4 of Nishida (1994). At least 6 bonobos of Lilungu wash aquatic herbs and/ or earthworms in the water before consuming them (Bermejo and others, 1994). Category 8. Watch (WAT) (Type A): Goodall (1989): "Gaze intently at what another is doing. If the face of the watching individual is very close to the other this is described as peering" (cf. Peer). Van Hooff's (1973) Watch. Kano's (1998) Watch for the bonobo. Category 1. Wean (WEA) (Type D): Goodall's (1989) Wean. Patterns of maternal rejection, encouraging the independence of the infant from the mother: such as preventing the infant from sucking, riding on her or sharing night bed. Kano's Wean for the bonobo. Category 1. Whimper (WHP) (Type A): Goodall (1989): "A whole series of soft, low pitched sounds, rising and falling in pitch, which may lead to crying and screaming...". Whimper by a male infant sometimes elicits presentation by an estrous female, thus serving as a courtship display (Nishida, 1997). Similar sound is emitted by bonobos (Kano, 1998). Category 2. Whimper-scream (WHS)(Type A): Goodall's (1989) Cry: "A combination of loud whimpers and tantrum screaming. This is commonly heard when an infant is separated from mother". At Mahale, weaning infants, especially those weaned when unusually young (for example, 3 years old), whimper-scream continuously almost all day throughout their mothers' estrous periods. Bonobo's lost call seems to be distinct from that of chimpanzees in acoustic nature and in that both mature and immature individuals emit it (Kano, 1998). Category 2. Wipe (WIP)(Type A): Goodall (1989): "Use of leaves, straw, paper, and so on to wipe dirt (feces, urine, mud, ejaculate, and so on) from self or others..." Wiping is very rare at Mahale, with only a handful of cases of rump wiping with a leaf. Goodall (1989) stated that the most common form at Gombe is penis wiping by a male after copulation, but this has never been seen among Mahale chimpanzees. Wiping dirt 180 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M. from others has also not been observed (Cf. Rub with a stalk/vine). Among the bonobos of Wamba , wiping has been seen only three times (Ingmanson, 1996). Category 8. Wraaa (WRA) (Type A): Goodall (1989): "---a long drawn out, pure sounding call, neither high or low pitched... Given by chimpanzees when confronted with new or disturbing objects in the environment - a human before they were habituated to humans, sometimes buffalo, python, dead bush pigs, and so on..." Category 3-5. Wrestle (WRE) (Type B): Goodall (1989): "Two or more youngsters grab hold of each other and often roll over as they bite, tickle, kick and so on without losing contact." (Cf. Rough and tumble play) . Kano's (1998) Wrestling play for the bonobo. Category 1. Wrestle with fingers (WFI)(Type A): Van Hooff's (1973) Handwrestle. Goodall (1989): "Gentle fondling, holding, squeezing, tickling of one hand (occasionally foot) of intended play partner. This is how a mature individual typically seeks to initiate play with another adult. It may lead to a vigorous bout of social play." At Mahale, this is used to initiate social play among adult males and between adult males and females, and is the most common pattern of social play overall. Not seen in the bonobo (Kano, 1998). Category 3.

Y Yawn (YAW) (Type A): Goodall's (1989) Yawning. Plooij's (1984) YAW . Kano's (1998) Yawn for the bonobo. Category 1.

Behavior Patterns Listed in Goodall (1989), But Not Seen at Mahale Eat vomit (EAV): Goodall's (1989) Reingest vomit: "A sick chimp may vomit into one hand, then eat the vomit". Not recorded for the bonobos at Wamba (Kano , 1998). Elbow clamp (ECL): Goodall (1989): "When small infant pesters mother for food, she may clamp it to her breast with both elbows and continue her meal over its head." Pinch (PIN): Goodall's (1989) Pinch. Not recorded for the bonobo (Kano, 1998). Throw fruit at (THF): The chimpanzee of Gombe throw a hard-shelled fruit against a rock or the trunk of a tree to crack it open (McGrew and others, 1999). This was termed Bang (BAN) by Plooij (1984).

Acknowledgments We thank Tanzania Commission for Science and Technology , Serengeti Wildlife Research Institute, and Tanzania National Parks for permission to do the field research , and Mahale Mountains National Park and Mahale MountainsWildlife Research Centre for logistic support. Our gratitude goes to H.Y. Kayumbo, C. Mlay, G. Sabuni, E. Massawe, B.C. Mwasaga, A.K. Seki and their staff. We are indebted to our field assistants, in particular, R. Kitopeni, H. Rashidi, M.B. Kasagula, K. Athumani, M. Matumla, M. Hawazi and M. Hamisi. Video tapes taken by various video teams were most valuable for detailed comparison of the behavior patterns. We are much indebted to Miho Nakamura and T. Asou of ANC Production, M.Mori and M. Matsuya of the East Company, and K. Sugimoto of the University of the Air for providing film rushes. We thank two anonymous referees for their constructive comments. Field research was supported by funds from the Monbusho International Scientific Research Program (#07041138 to T. Nishida). Analysis o the behavior based on videotapes and photos was funded by a Monbusho Grant-in-Aid for Basic Scientific Research(#08454278 to T. Nishida) and Grant-in-Aid for COE Research (#10CE2005 too. Takenaka).

REFERENCES Albrecht H. and Dunnett S.C. (1971) Chimpanzees in West Africa. Piper Verlag , Ethogram of chimpanzees 181

Munich. Adang O. M. J. (1984) Teasing in young chimpanzees. Behaviour, vol. 88, pp. 98-122. Arcadi A. C., Daniel R., and Boesch C. (1998) Buttress drumming by wild chimpanzees: Temporal patterning, phrase integration into loud calls, and preliminary evidence for individual distinctiveness. Primates, vol.39, pp.505-518. Berdecio S. and Nash L. T. (1981) Chimpanzee visual communication. Arizona State University, Anthropological Research Papers No.26. Bermejo M., Illera G., and Sabater Pi J. (1994) Animals and mushrooms consumed by bonobos (Pan paniscus): New records from Lilungu (Ikela), Zaire. International Journal of Primatology, vol.15, pp.879-898. Bodenheimer F. S.(1951) Insects as human food. Dr. W. Junk, Publishers, Hague. Boesch C.(1995) Innovation in wild chimpanzees (Pan troglodytes). International Journal of Primatology, vol.16, pp.1-16. Boesch C. and Boesch H. (1989) Hunting behavior of wild chimpanzees in the Tai National Park. American Journal of Physical Anthropology, vol.78, pp.547-573. Boesch C. and Boesch H.(1999) The chimpanzees of the Tai Forest, Cambridge University Press, Cambridge (in press). Bygott J. D.(1971) Cannibalism among wild chimpanzees. Nature, vol.238, pp.410-411. Bygott J. D.(1974) Agonistic behaviour and dominance in wild chimpanzees. Ph. D. thesis, University of Cambridge, Cambridge. Caro T. M. and Hauser M.D.(1992) Is there teaching in nonhuman animals? Quarterly Review of Biology, vol.67, pp.151-174. Eibl-Eibesfeldt I.(1972) Love and hate. The natural history of behavior patterns. Holt, Rinehart and Winston, New York. Feistner A. T. C. and McGrew W. C.(1989) Food-sharing in primates: A critical review. In Seth P. K. and Seth S. eds., "Perspectives in primate biology vol. 3", Today & Tomorrow's, New Delhi, pp.21-36. Furuichi T. (1989) Social interactions and the life history of female Pan paniscus in Wamba, Zaire. International Journal of Primatology, vol. 10, pp.173-197. Gagneux P., Boesch C., and Woodruff D. S.(1999) Female reproductive strategies, paternity and community structure in wild chimpanzees. Animal Behaviour, vol.57, pp.19-32. Goodall J.(1968) The behavior of free-living chimpanzees in the Gombe Stream Reserve. Animal Behaviour Monograph, vol.1, pp.161-311. Goodall J.(1971) In the shadow of man. Collins, London. Goodall J.(1973) Cultural elements in a chimpanzee community. In Menzel E. W. ed., "Precultural primate behavior" , S. Karger, Basel, pp. 144-184. Goodall J.(1986) The chimpanzees of Gombe. Harvard University Press, Cambridge. Goodall J.(1989) Glossary of chimpanzee behaviors. Jane Goodall Institute, Tucson. Goodall J.(1992) Unusual violence in the overthrow of an alpha male chimpanzee at Gombe. In Nishida T., McGrew W. C., Marler P., Pickford M., and de Waal F. B. M. 182 Nishida T., Kano T., Goodall J., McGrew W.C., and Nakamura M.

eds.,"Topics in primatology 1. Human origins", University of Tokyo Press, Tokyo, pp. 131-142. Hamai M., Nishida T., Takasaki H., and Turner L. A.(1992) New records of within- group infanticide and cannibalism in wild chimpanzees. Primates, vol.33, pp.151-162. Hasegawa T.(1990) Sex differences in ranging patterns. In Nishida T. ed.,"The chimpanzees of the Mahale Mountains", University of Tokyo Press, Tokyo, pp.99-114 . Hasegawa T., Hiraiwa-Hasegawa M., Nishida T., and Takasaki H.(1983) New evidence of scavenging behavior of wild chimpanzees. Current Anthropology, vol.24 , pp. 231-232. Hayaki H.(1985) Social play of juvenile and adolescent chimpanzees in the Mahale Mountains National Park, Tanzania. Primates, vol. 26, pp.343-360. Hiraiwa-Hasegawa M.(1989) Sex differences in behavioral development at Mahale. In Heltne P.G. and Marquardt L. A. eds.,"Understanding chimpanzees" , Harvard University Press, Cambridge, pp.104-115. van Hooff J. A. R. A. M.(1973) A structural analysis of the social behavior of a semi- captive group of chimpanzees. In von Cranach M. and Vine I. eds., "Social communication and movement", Academic Press, London, pp.75-162. Hohmann G. and Fruth B.(1993) Field observations on meat sharing among bonobos (Pan paniscus). Folia Primatologica, vol.60, pp.225-229. Huffman M. A.(1997) Current evidence for self-medication in Primates: A multidisciplinary perspective. Yearbook of Physical Anthropology, vol. 40, pp.171-200. Humle T.(1999) New record of fishing for termites (Macrotermes) by the chimpanzees of Bossou (Pan troglodytes verus), Guinea. Pan Africa News, vol.6, pp.3-4. Hunt K. D., Cant J. G., Gebo D. L., Rose M. D., Walker S.E., and Youlatos D. (1996) Standardized descriptions of primate locomotor and postural modes . Primates, vol.37, pp.363-387. Idani G. (1990) Relations between unit-groups of bonobos at Wamba , Zaire: Encounters and temporary fusions. African Study Monographs, vol.11, pp.153-186. Idani G.(1991) Social relationships between immigrant and resident bonobo (Pan paniscus) females at Wamba. Folia Primatologica, vol. 57, pp.83-95 . Idani G.(1995) Function of peering behavior among bonobos (Pan paniscus) at Wamba . Primates, vol.36, pp.377-383. Ingmanson E. J.(1996) Tool-using behavior in wild Pan paniscus: Social and ecological considerations. In Russon A.E., Bard K.A., and Parker S.T. eds., "Reaching into thought", Cambridge University Press, Cambridge, pp. 190-210. Kano T.(1980) Social behavior of wild pygmy chimpanzees (Pan paniscus) of Wamba: A preliminary report. Journal of Human Evolution, vol.9, pp.243-260. Kano T.(1982) The social group of pygmy chimpanzees (Pan paniscus) of Wamba . Primates, vol.23, pp.171-188. Kano T. (1990) The bonobo's peaceable kingdom. Natural History , November, pp. 62-71. Ethogram of chimpanzees 183

Kano T.(1992) The last ape. Stanford University Press, Stanford. Kano T.(1998) A preliminary glossary of bonobo behaviors at Wamba. In Nishida T. ed ., "Comparative study of the beh avior of the genus Pan by compiling video ethogram", Nissho Printer, Kyoto, pp.39-81. Kawanaka K.(1981) Infanticide and cannibalism in chimpanzees, with special reference to the newly observed case in the Mahale Mountains. African Study Monographs, vol. 1, pp.69-99. Kortlandt A.(1964) Experimentation with forest-dwelling chimpanzees in the Congo. The verbatim text of the 16mm film. Kortlandt A.(1967) Experimentation with chimpanzees in the wild. In Satrck D., Schneider R., and Kuhn H.-J. eds., "Neue Ergebnisse der Primatologie", Fischer, Stuttgart , pp. 208-244. Kortlandt A. and Bresser M.(1963) Experimentation with forest-dwelling chimpanzees in the Congo, 1963. 8mm Film. Kortlandt A. and Kooij M.(1963) Protohominid behavior in primates. Symposia of Zoological Society of London, No.10, pp.61-87. Kuroda S.(1980) Social behavior of the pygmy chimpanzees. Primates, vol. 21, pp. 181-197. Kuroda S.(1984) Interaction over food among pygmy chimpanzees. In Susman R. L. ed., "The Pygmy Chimpanzee" , Plenum Press, New York, pp.301-324. Kuroda S.(1998) Preliminary ethogram of tschego chimpanzees. In Nishida T. ed., "Comparative study of the behavior of the genus Pan by compiling video ethogram". Nissho Printer, Kyoto. pp.82-91. Matsumoto-Oda A.(1997) Self-suckling behavior by a wild chimpanzee. Folia Primatologica, vol.68, pp.342-343. Matsuzawa T.(1997) The death of an infant chimpanzee at Bossou, Guinea. Pan Africa News, vol.4, pp.4-6. Matsuzawa T. and Yamakoshi G.(1996) Comparison of chimpanzee material culture between. Bossou and Nimba, West Africa. In Russon A. E., Bard, K. A., and Parker S.T. eds.,"Reaching into thought", Cambridge University Press, Cambridge, pp. 211-232. McGinnis P.R. (1973) Patterns of sexual behavior in a community of free-living chimpanzees. Ph. D. thesis, University of Cambridge. McGrew W. C.(1972) An ethological study of children's behavior. Academic Press, New York. McGrew W. C. (1974) Tool use by wild chimpanzees in feeding upon driver ants. Journal of Human Evolution, vol.3, pp.501-508. McGrew W. C. (1992) Chimpanzee material culture: Implications for human evolution. Cambridge University Press, Cambridge. McGrew W.C. (1998) Culture in nonhuman primate? Annual Review of Anthropology, vol.27, pp.301-328. 184 Nishida T., Kano T., Goodall J., McGrew W. C., and Nakamura M.

McGrew W. C., Baldwin P.J., and Tutin C.E. G.(1988) Diet of wild chimpanzees (Pan troglodytes verus) at Mt. Assirik, Senegal. I. Composition. American Journal of Primatology, vol.16, pp.213-226. McGrew W. C. and Collins D. A.(1985) Tool use by wild chimpanzees (Pan troglodytes) to obtain termites (Macrotermes herus) in the Mahale Mountains, Tanzania. American Journal of Primatology, vol.9, pp.47-62. McGrew W.C. and Marchant F. L.(1998) Chimpanzee wears a knotted skin "necklace". Pan Africa News, vol.5, pp.8-9. McGrew W. C., Marchant F.L., and Nishida T.(1996) Great Ape Societies. Cambridge University Press, Cambridge, Mass. McGrew W. C., Marchant F. L., Wrangham R.W., and Klein H. (1999) Manual laterality in anvil use: Wild chimpanzees cracking Strychnos fruits. Laterality 4: 79-87. McGrew W. C. and Rogers M. E.(1983) Chimpanzees, tools and termites: new records from Gabon. American Journal of Primatology, vol.5, pp.171-174. McGrew W. C. and Tutin C. E. G.(1978) Evidence for a social custom in wild chimpanzees? Man (n. s.), vol.13, pp.234-251. McGrew W. C., Tutin, C.E. G., and Baldwin P.J.(1979) Chimpanzees, tools and termites: cross-cultural comparison of Senegal, Tanzania and Rio Muni. Man (n. s.), vol.14, pp. 185-214. Mitani J. and Nishida T. (1993) Contexts and social correlates of long-distance calling by male chimpanzees. Animal Behaviour, vol.45, pp.735-746. Mori A.(1982) An ethological study on chimpanzees at the artificial feeding place in the Mahale Mountains, Tanzania----with special reference to the booming situation. Primates, vol.23, pp.45-65. Mori A.(1983) Comparison of the communicative vocalizations and behaviors of group ranging in eastern gorillas, chimpanzees and pygmy chimpanzees. Primates , vol.24, pp.486-500. Morin P. A., Moore J. J., Chakraborty R., Jin L., Goodall J., and Woodruff D. S.(1994) Kin selection, social structure, gene flow, and the evolution of chimpanzees. Science, vol.265, pp.1193-1201. Morris D.(1977) Man watching. A field guide to human behaviour. Elsevier Publishing Projects Ltd., Oxford. Muller M. N., Mpongo E., Stanford C. B., and Boehm C.(1995) A note on scavenging by wild chimpanzees. Folia Primatologica, vol. 65, pp.43-47. Nakamura M., McGrew W. C., Marchant L.F., and Nishida T.(1999) Social scratch. (submitted) Newton-Fisher N. (1998) The socio-ecology of Budongo's chimps. Budongo Forest Project, vol. 1, no.2, p.2. Nishida T.(1968) The social group of wild chimpanzees in the Mahali Mountains . Primates, vol.9, pp.167-224. Nishida T. (1970) Social behavior and relationship among wild chimpanzees of the Ethogram of chimpanzees 185

Mahali Mountains. Primates, vol.11, pp.47-87. Nishida T.(1973) The ant-gathering behaviour by the use of tools among wild chimpanzees of the Mahali Mountains. Journal of Human Evolution, vol.2, pp.357-370. Nishida T.(1976) The bark-eating habits in Primates, with special reference to their status in the diet of wild chimpanzees. Folia Primatologica, vol.25, pp.277-287. Nishida T.(1979) The social structure of chimpanzees of the Mahale Mountains. In Hamburg D. A. and McCown E. R. eds.,"The great apes", Benjamin/Cummings, Menlo Park, pp.72-121. Nishida T.(1980a) Local differences in responses to water among wild chimpanzees. Folia Primatologica, vol.33, pp.189-209. Nishida T.(1980b) The leaf-clipping display: a newly-discovered expressive gesture in wild chimpanzees. Journal of Human Evolution, vol.9, pp.117-128. Nishida T.(1983a) Alloparental behavior in wild chimpanzees of the Mahale Mountains, Tanzania. Folia Primatologica, vol.41, pp.1-33. Nishida T.(1983b) Alpha status and agonistic alliance in wild chimpanzees (Pan troglodytes schweinfurthii). Primates, vol.24, pp.318-336. Nishida T.(1987) Local traditions and cultural transmission. In Smuts B. B., Cheney L., Seyfarth R. M., Wrangham R. W., and Struhsaker T. T. eds., "Primate societies", University of Chicago Press, Chicago, pp.462-474. Nishida T.(1989) Social interactions between resident and immigrant female chimpanzees. In Heltne P. G. and Marquardt L. eds.,"Understanding chimpanzees", Cambridge University Press, Cambridge, Mass., pp.68-89. Nishida T.(ed.)(1990) The chimpanzee of the Mahali Mountains. University of Tokyo Press, Tokyo. Nishida T.(1993) Chimpanzee are always new to me. In Cavalieri P. and Singer P. eds., "Great ape project" , Fourth Estate, London, pp.24-26. Nishida T.(1994) Review of recent findings on Mahale chimpanzees: implications and future research directions. In Wrangham R.W., McGrew W. C., de Waal F. B. M., and Heltne P. eds., "Chimpanzee cultures", Harvard University Press, Cambridge, MA., pp.373-396. Nishida T.(1996) The death of Ntologi. The unparalleled leader of M group. Pan Africa News, vol.3, no.1, p.4. Nishida T.(1997) Sexual behavior of adult male chimpanzees of the Mahale Mountains National Park, Tanzania. Primates, vol.38, pp.379-398. Nishida T.(1998) Deceptive tactic by an adult male to snatch a dead infant from a mother. Pan Africa News, vol.5, pp.13-15. Nishida T. and Hiraiwa M. (1982) Natural history of a tool-using behavior by wild chimpanzees in feeding upon wood-boring ants. Journal of Human Evolution, vol.11, pp.73-99. Nishida T. and Hiraiwa-Hasegawa M.(1985) Responses to a stranger mother-son pair in the wild chimpanzee: A case report. Primates, vol.26, pp.1-13. 186 Nishida T., Kano T., Goodall J., McGrew W. C., and Nakamura M.

Nishida T. and Hiraiwa-Hasegawa M.(1987) Chimpanzees and bonobos: Cooperative relationships among males. In Smuts B.B., Cheney D. L., Seyfarth R.M., Wrangham R. W., and Struhsaker T. T., eds.,"Primate societies", The University of Chicago Press , Chicago, pp.165-177. Nishida T., Hiraiwa-Hasegawa M., Hasegawa T., and Takahata Y.(1985) Group extinction and female transfer in wild chimpanzees in the Mahale National Park, Tanzania. Zeitschrift fur Tierpsychologie, vol.67, pp.284-301. Nishida T. and Hosaka K.(1996) Coalition strategies among adult male chimpanzees of the Mahale Mountains, Tanzania. In McGrew W.C., Marchant L. F., and Nishida T. eds., "Great ape societies", Cambridge University Press, Cambridge, pp.114-134. Nishida T., Hosaka K., Nakamura M., and Hamai M.(1995) A within-group gang attack on a young adult male chimpanzee: Ostracium of an ill-mannered member? Primate , vol.36, pp.207-211. Nishida T. and Kawanaka K.(1972) Inter-unit-group relationships among wild chimpanzees of the Mahali Mountains. Kyoto University African Studies , vol.7, pp. 131-169. Nishida T. and Nakamura M.(1993) Chimpanzee tool use to clear a blocked nasal passage. Folia Primatologica, vol.61, pp.218-220. Nishida T. and Nakamura M.(1998) Ethogram for Mahale chimpanzees: A preliminary draft. In Nishida T. ed.,"Comparative study of the behavior of the genus Pan by compiling video ethogram". Nissho Printer, Kyoto, pp.5-38. Nishida T., Takasaki H., and Takahata Y.(1990) Demography and reproductive profiles. In Nishida T. ed."The chimpanzees of the Mahale Mountains", University of Tokyo Press, Tokyo, pp.63-97. Nishida T. and Uehara S.(1980) Chimpanzees, tools , and termites: Another example from Tanzania. Current Anthropology, vol.21, pp.671-672. Nishida T. and Uehara S.(1983) Natural diet of chimpanzees (Pan troglodytes schweinfurthii): Long-term record from the Mahale Mountains, Tanzania. African Study Monographs, vol.3, pp.109-130. Nishida T. and Turner L. A.(1996) Food transfer between mother and infant chimpanzees of the Mahale Mountains National Park, Tanzania. International Journal of Primatology, vol.17, pp.947-968. Okayasu N.(1991) Vocal communication and its sociological interpretation of wild bonobos in Wamba, Zaire. In Ehara A., Kimura T., Takenaka O., and Iwamoto M. eds.,"Primatology today", Elsevier Science Publishers, pp.239-240 . Plooij F. X.(1984) The behavioral development of free-living chimpanzee babies and infants. Ablex Publishing Corporation, Norwood, New Jersey. Sugiyama Y.(1981) Observations on the population dynamics and behavior of wild chimpanzees at Bossou, Guinea, in 1979-1980. Primates, vol.26, pp.361-374. Sugiyama Y.(1985) The brush-sticks of chimpanzees found in southwest Cameroon and their cultural characteristics. Primates, vol.26, pp.361-374. Ethogram of chimpanzees 187

Sugiyama Y.(1997) Social tradition and the use of tool-composites by wild chimpanzees. Evolutionary Anthropology, vol. 6, pp.23-27. Sugiyama Y.(1998) Local variation of tool-using repertoire in wild chimpanzees . In Nishida T. ed."Comparative study of the behavior of the genus Pan by compiling video ethogram". Nissho Printer, Kyoto, pp.82-91 Susman R. L.(1984) The pygmy chimpanzee: Evolutionary biology and behavior. Plenum Press, New York. Susman R. L., Badrian N.L., and Badrian A. J. (1980) Locomotor behavior of Pan paniscus in Zaire. American Journal of Physical Anthropology, vol. 53, pp.69-80. Suzuki A. (1971) Carnivority and cannibalism observed among forest-living chimpanzees. Journal of Anthropological Society of Nippon, vol. 79, pp. 30-48. Suzuki S., Nishihara T., and Kuroda S.(1995) Tool-set for termite fishing by chimpanzees in the Ndoki Forest, Congo. Behaviour, vol. 132, pp.219-235. Takahata H. and Takahata Y.(1989) Inter-unit group transfer of an immature male of the common chimpanzee and his social interactions in the non-natal group. African Study Monographs, vol.9, pp.209-220. Takahata Y.(1985) Adult male chimpanzees kill and eat a male newborn infant: Newly observed intragroup infanticide and cannibalism in Mahale National Park, Tanzania. Folia Primatologica, vol. 44, pp.161-170. Takenoshita Y., Yamagiwa J., and Nishida T.(1998) Branch-dropping display by a female chimpanzee (Pan troglodytes troglodytes) of Petit Loango, Gabon. Pan. Africa News, vol. 5, pp.16-17. Tutin C.E. G.(1974) Exceptions to promiscuity in a feral chimpanzee community. Proceedings of the 5th Congress of the International Primatological Society, Nagoya, Japan, pp.28-29. Tutin C. E. G.(1979) Responses of chimpanzees to copulation, with special reference to interference by immature individuals. Animal Behaviour, vol. 27, pp.845-854. Tuttle R. H.(1986) Apes of the world. Noyes Publications, Park Ridge. Uehara S.(1982) Seasonal changes in the techniques employed by wild chimpanzees in the Mahale Mountains, Tanzania, to feed on termites (Pseudacanthotermes spiniger). Folia Primatologica, vol. 37, pp.44-76. Uehara S.(1990) A preliminary report on age differences in plant-feeding behaviors among adult and adolescent males. In Nishida T. ed., "The chimpanzees of the Mahale Mountains", University of Tokyo Press, pp. 223-236. Uehara S. (1997) Predation on mammals by the chimpanzee. Primates, vol. 38, pp. 193-214. de Waal F. B. M. (1982) Chimpanzee politics. Jonathan Cape, London. de Waal F. B. M. (1988) The communicative repertoire of captive bonobos (Pan paniscus), compared to that of chimpanzees. Behaviour, vol. 106, pp.183-251. de Waal F. B.M. (1989) Peace making among primates. Harvard University Press, Cambridge, Mass. 188 Nishida T., Kano T., Goodall J., McGrew W. C., and Nakamura M. de Waal F. B. M. (1991) The chimpanzee's sense of social regularity and its relation to the human sense of justice. American Behavioral Scientist, vol. 34, pp. 335-349. de Waal F. B.M. (1995) Bonobo sex and society. Scientific American, vol.272, pp.58-64. de Waal F. B.M. and Lanting F. (1997) Bonobo. The forgotten ape. University of California Press, Berkeley. de Waal F. B. M. and Seres M. (1997) Propagation of handclasp grooming among captive chimpanzees. American Journal of Primatology, vol. 43, pp.339-346. Wrangham R.W. (1977) Feeding behavior of chimpanzees in Gombe National Park, Tanzania. In Clutton-Brock T.H. ed., "Primate ecology", Academic Press, London, pp. 503-538. Wrangham R.W. and Nishida T. (1983) Aspilia spp. leaves. A puzzle in the feeding behavior of wild chimpanzees. Primates, vol. 24, pp.276-282. Wrangham R.W. and Peterson D. (1996) Demonic males. Houghton Mifflin Company, Boston.

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