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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
2
3
4
Sub-Departmentof Anthropology,Graduate Schoolof Science,Kyoto University,Kyoto
Primate Research Institute, Kyoto University,Inuyama
Jane GoodallInstitute, SilverSpring, U.S.A.
Departments of Sociology,Gerontologyand 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: nishida@jinrui.zool.kyoto-u.ac.jp
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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 <walk> or <move>
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 <walk>, 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 <walk> or <move>. 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):Alloparentleaves 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 <brushaside>,
<pushaway>, <turnupside down>, <kick>,<pushwithfoot>, <stamp on>, <pull>,<strikewithknuckles,
Ethogram
of chimpanzees
145
the back of the wrist or the fingertips>, <pull the foot>, <press its legs>, 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 <hug>, <groom>, <transport>, <protect>, <play>, <share food>, and
<share bed>, (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 <pant-grunt>, <touch>,
<kiss>, <embrace>, <present>, <brief grooming>, and so on. The bonobos of Wamba appease by
<touch>, <embrace-half>, <present>, <brief-groom>, 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 <walk on all fours>,
<walk bipedal>, and <run>. 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 <push>, <kick>, <hit>, <grab>, <slap>, <bite>, <drag>, <charge>, <stamp>,
<pinch>, and <scratch>. 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.
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and Nakamura
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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. <leap>, <push>, <pull>. 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 <hug>,
<groom>, <transport>, <mouth>, <lie-hug>, <put dorsal> , <pat>, <touch>, <balance>, <protect>,
<scratch>, <share food>, <push finger into mouth>, 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 <hug>, <suckle>, <transport>, <groom>, <protect>, <play with>, <share food>, <share bed>,
<reject>, <support>, 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.
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W.C.,
and Nakamura
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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.
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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 <cradle by leg>. 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
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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 <scrub>, <throw>, <drag branch>, <sway branch>, <slap>, <stamp>, <slap-stamp> , <flail>,
<drum>, <rake> and occasionally <beat chest>... The charging display may also incorporate <hair
erection>, <compressed lips and face> or <pant hoots>. 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 <beat chest> , but
often by <scratch leaves> and <bipedal swagger> as well as the behavior above mentioned. The bonobo's
charging display has only two patterns: <charge> and <drag branch> (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
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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 <pant-hoot>, <charge>, <run bipedal>, <stamp>. and so on. Another is prolonged, slow
tempo displays, of which the main components are <sway trunks of tree/shrub> , <run on one's feet>,
<climb tree rapidly>, <brachiate> and <leap>. 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 <run>, <jump>, <drag branch>, <throw splash> and
<sway and shake branch/woody vine>. 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
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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 <pull through>.
Ethogram
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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
<thrust>. 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
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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 <scream>, <run>, <climb>, <jump>, and <descend>. 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 <bob>, <crouch>, <touch>, <kiss>, <embrace>, <groom>,
<present>, <mount>, <inspect>, <hold hand>, and so on. Vocalization associated with greeting behavior
are <pant-grunt>, <pant-hoot>, <grunt>, <pant-bark>, <scream>, and so on." In bonobos, genital contact
behavior and <peer> may be comparable to greeting, in addition to <groom>, <present>, <mount>,
<inspect>, and <hold hand>, 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 <sit>, <glance>, <walk and stop>, <return> and <turn back> ,
threatening such as <gaze>, <shake branch>, <club>, and aggression such as <hit> and <kick>. 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 <club> 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. <Hit> 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 <monitor>, <climb>, <stalk>, <run> ,
<chase>, <lunge>, <grab>, <flail>, <drag>, <drag to kill>, <throw>, <bite>, <pull>, 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 <gaze>, <sniff>, <touch> ,
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 <bend>, <transport>, <sit>, and <stand bipedal>. 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 <walk>, <watch>, <touch>, <sniff>, and
<bite>. 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 <walk>, <move>, <climb>, <watch>, <look up> 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 <whimper> , <scream>,
<whimper-scream>, <run>, <turn round>, <glance>, and <temper tantrum>. 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
O
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 <present> 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
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 . <walk quadrupedal>. 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 <sniff>, <creep>, <maintain silence>, <stare>, <attack>, and
<flee>. 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
Pirouette
(PIR)
circles,
for
(Type
moving
on
illustration.
Seen
unpublished).
(PLY)
lone
play
a
type
<flail>,
(d)
play-bite
(d)
See
A multiparous
"aeroplane"
progresses
play".
absent
in
of
See
in
the
play
each
in
Figure
the
7f
a series
of
bonobos
an
chasing,
or
Play
in
1998).
on
a limb
grip
the
<slap>,
play
so
on.
and
and
contexts.
play,
the
a
bed
D):
female
infant
Engaged
lying
who
Play
in
<wrestle>,
from
on
had
in
self-play,
but
the
(Type
and
self
Play
socially
(Type
A):
is
social
back>,
minutes
Figure
4.9
with
up was
months
seen
to
(Nishida
has
(PWL)
(Type
and
pulled
back
play
with
object
Play
with
water
body
at
of
mouth
is
teeth.
See
Hasegawa
Mouth,
(1990)
for
imaginary
companion
play-pant
as
or
if she
was
Hayaki
play-panting
loudly
tool.
playing
(1985)
as
if
he
was
Seen
also
among
the
bonobo
(Kano,
type
of
play
include
usually
<chase>,
<stamp>,
between
<run>,
<slap>,
youngsters
<grab>,
<thrust>,
.
<tag>,
<leap>,
<climb>,
1.
(LAC).
playing
Bonobos
Goodall's
socially.
also
(1989)
As
play-pant
the
(de
Laugh:
play
"Somewhat
becomes
Waal,
resembles
rougher,
1988;
or
Okayasu
involves
, 1991;
Kano,
with
(1998)
Social
stone
and
Lone
object.
animal
(Self)
play.
Social
skin
and
Category
play
so
1.
including
chasing
on . Ingmanson's
an
(1996)
2.
"The
shoulders
Kano's
(1989)
branch,
1 or
(1989):
chimp
while
it
walks
takes
with
small
a rounded-back,
stilted
its
steps•c"
head
Seen
slightly
also
in
bent
bonobos
2.
(Type
B):
Goodall's
(Type
B):
Type
on.
Category
so
(1983a):
to
may
been
A):
slightly
the
a fruit,
the
play.
(1989)
of
self
at
play
including
above.
<touch
Category
and
"hit"
1.
water>,
<peer
into
water
1-7.
Tickling,
by
an infant's
ventrum
poke
another
chimp
seen
Object
play
Mahale.
See
prodding
while
who
Figure
lying
has
la
repeatedly
on its
ignored
of
with
fingertip
back.
Goodall
a solicitation
Nishida
(1983a)
contacts
(1989)
mentioned
for
grooming
for
with
one
that
." Poking
photograph.
Seen
in
1.
(Type
be
as
Self
Goodall's
Category
Nishida
not
B):
Goodall
and
A):
has
(POU)
such
Category
Category
Category
Laugh
(1989)
between
water>
context
seen
physical
Category
are
(Type
bonobo.
(PLO)
(Type
may
with
a time..."
8?
above.
Elements
<drag>,
on.
(1984)
Goodall's
A):
extended
fingers
an individual
bonobos.
Pout
the
(PLW)
<stir
(POK)
this
for
unpublished).
Play
in
so
laughter..."
object
use
or several
"Sometimes
play
Most
play.
<mouth>,
and
(PSO)
an
walk
Poke
while
Category
Nest
B):
chimpanzees
B):
object
Play
surface>,
repeatedly
observed.
(Type
Plooij's
more
(Type
with
who
(Kano,
play
<pinch>,
1.
Tool
down
tree
(1989)
<bite>,
when
there
in
a
locomotor
<grin>,
heard
(PLS)
individual
Goodall's
(PRT)
<pull>,
Category
Play
around
Rarely
often
<scream>,
tickling,
1998).
tumble
<push>,
laughter,
more
running
B):
and
(PLP)
human
, and
, <drape>,
Goodall's
partner's
for
apparently
her
back
with
all limbs
been
dead
for
a few
a playmate.
wrestling
pant>,
Play-pant
<dangle>,
<leap>
2.
rough
Comprises
<play
male
(PLN)
vocalization,
<drag>
<kick
(c)
play
1.
on
on
See
with
play:
<tickle>.
<kick>,
made
holds
play,
and
feet>,
divided
nest
a bed>,
one's
kick>,
she
(b)
locomotor
<make
Category
"Contact
walk,
(a)
with
<heel
and
bite:
play
play:
object
<dangle>,
play
nest
Further,
connected
follows:
(b)
wrestle>,
play.
face.
(Type
adolescent
Category
(Kano,
2.
place
fleeing
tight
(1968)
Wamba
, unpublished).
recorded
of
Goodall
of
Gymnastics,
the
as
<rotate
<wrestle>,
social
expression
type
<somersault>.
(1989)
and
(1973)
context,
play
<finger
in
play
Hooff's
is a facial
<grab>,
a firm
lone
van
<rub>,
<drag>,
displayed
1 or
adult
with
her
only
play:
into
or
"There
and
self
gets
<mouth>
(PLI)
seen
Goodall's
players
Show
imaginary
that,
<tickle>,
A):
Category
face:
a youngster)
locomotor
Probably
play
<throw>,
<chase>,
the
is
of
play,
elements
up>,
<spar>,
of
(usually
type
divided
is
as
<pick
(Type
one
Mahale.
<pirouette>,
<butt>,
<rub>,
photograph.
Play,
that
<move-away>.
Sometimes
at
added
listed
play:
(PLB)
chimp
A
locomotor
She
<swing>,
object
<play-walk>,
Play-bite
play.
(1989)
<bite>,
"The
feet.
(1989)
(a)
self
locomotion
and
included
Goodall
of
(c)
their
juveniles
categories,
<leap>,
<climb>.
(1989):
on
3?
Goodall
<gambol>,
Play
or
B):
4
and
laughing."
since
Goodall
among
(Type
into
play,
face,
A):
fours
Category
Play
object
all
165
of chimpanzees
Plooij's
open.
hoo
call
(1984)
Lips
and
1.
Pout
face:
See
Pout.
Pout
kiss:
See
Kiss
in
pout
face.
PTF.
together,
hoo
whimper".
Goodall's
except
at
Bonobos
(1987)
the
front
have
Pout
where
a
similar
face:
they
"Lip
may
facial
corners
funnel
expression
pushed
out
forward,
somewhat.
(Kano
This
,
1998).
166
Nishida
T.,
Kano
T., Goodall
J., McGrew
W. C., and Nakamura
M.
Presentwith limbs extended(PRE) (TypeA): Normal quadrupedal stancewithhindquartersdirectedto
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-ventralpresentingfor the bonobo.
In addition,bonobo females have ventro-ventral
courtshipgesture.Category 2.
Presentwithlimbs flexed(PRF) (TypeA): van Hooff (1973)'s Crouch-present.
Quadrupedal posturewith
limbs flexed,and hindquartersturnedtowards another individual.
See Figure 7 of Nishida (1979)for
photograph.Differswith Crouch in which hindquartersare not directedto the partner.Category 3.
Press (PRS) (TypeA): Pushone s shoulder,sideor hip againstanotherindividual.
Functionunclear, but
sometimes initiate
socialplay.Category 3-8?
Press teeth againstback (PTB) (TypeA): Adult male mounts another male, and the mounter often
presseshisupper row of teethagainst the back of the mounted male, while the mounter embraces the
latter,
performs a pelvicthrustsand shows a wide open grin.Intensified
form of <mouth> . Not seen inthe
bonobos of Wamba. Category 3-6.
Probe (PRB) (TypeA): Use a vine,twig or stickto touchsome seen or unseen objects.
Goodall's(1989)
Investigation
probe.Infantchimpanzees appear tohave an innatetendencyto push an investigation
probe
intoany hole of a treelong 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
(forexample, gaps in canopy) or socialharassment (forexample , malecharging
displays).
Simar patternisseen inthe bonobo, exceptfor protection
from 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 Pullforthe bonobo. See Figurel ofNishida (1983a).
Category 1.
Pulldown (PLD)(TypeA): Stretcharm to reach upper branch of shrub,often whilestandingon both
feet,and pullthe branch downward. Followed by sitting
and eatingfruitsor leaveswhilegrippingthe
branch with one hand. Also seen in bonobos (Kano, unpublished).
Category 1.
Pullface to face (PLF)(Type C): Pullhand or otherbody partof anotherindividual
, so thatthey face
each other.Occurs when chimpanzee wants to groom the frontalpartof the other.Also seen inbonobos
(Kano, unpublished).
Category 1-2.
Pullfrom opposing sidessimultaneously(POS)(TypeB): Tug of war . Two chimpanzees simultaneously
pullan object (infant,branch, carcassand so on.)competitivelyfrom opposing sides.Seen also in
bonobos.Category 1.
Pullhead withboth hands (PLH)(TypeA): Pullhead of anotherwithboth hands,often ingrooming.Also
seen in bonobos (Kano, unpublished).
Category 1-2.
Pullout (PUO)(TypeA): Pullforciblywith both hands, oftencounteredby pushingmovement of both
feeton the ground.Occurs when chimpanzee triesto draw stalksof banana, elephantgrasses,
or African
gingersfrom the soil.
Also seen inbonobos.Category 1.
Pullthrough (PTH)(TypeA): Upward sweeping movement of thumb and index fingeror palmar surface
of hand along a leafybranch or leafyherb stem. This isa food-processing
techniquefor leaves,vines,
herbs or blossoms(forexample, Erythrinaabyssinica)whileholdinga twig withone hand and pullingit
through between lips/teeth.
Also used by Gombe chimpanzees in ant dipping(McGrew , 1974).Seenin
bonobos (Kano, unpublished).
Category 1-2.
Pullthrough stem (PTS)(TypeA): Nishida'
s (1997)Stem pull-through
(courtship).
Pullthe leafybranch
of a shrub or clump of grassstems throughthe hand by a rapidupward movement of the forearm,then,
the stem isimmediatelyreleased.Produces a conspicuoussound,while seatedand watchingan estrous
female.Only two of the oldestadultmales,Musa and Ntologi,did thiscourtship.
An exaggeratedform
of pullingthroughinthe eatingcontext.
One adultmale (Fanana)toreoffthe shootof a stem from a shrub
as a part of courtship.
Category 8.
Push (PUS)(TypeA): Exert forceby extendingarms incontactduringaggressionand grooming.In order
to groom another,push partof the back (forexample, shoulder)of anotherso thatthe latterwillturnits
back.Groomer stimulates
the groomee to change postureafterextensivegrooming of a particular
partof
the body. Kano's (1998)Push for the bonobo. Category 1.
Push ahead (PUA)(Type C): Goodall (1989):"A mother pushesher infantahead of her duringweaning
to preventdorsaltravel.
Often she does thiswithlittle
slapsof her hand on itsrump and thisisapparently
perceivedby the infantasplayful".
PUA isnot seen inthe bonobos of Wamba (Kano, 1998).Category3-5.
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
Quadrupedal
Quadrupedal
Quadrupedal
Quadrupedal
hunch: See Hunch quadrupedal.
jump: See Leap quadrupedal.
run: See Run quadrupedal.
transport: See Transport 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 <pat>, <touch>,
<embrace>, <kiss>, and <groom>." 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 <approach> ,
<touch>, <embrace>, <mount>, <open-mouth-kiss> and <groom> . The bonobos of Wambareconcile by
<approach>, <touch>, <mount>, <groom>, <GG-rubbing> , and <rump-rump rubbing> (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) "<cover breast>,
<fend>, <turn away>, <reject-move>, (2) <reject-move>, <shrug>, <push ahead>, <reject-sit>, (3) <turn
away>, <fend>, <elbow-clamp>, and <reject-move> and (4) <threaten>" . 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 , <pick up>.
Displayed by at least five chimpanzees of Mahale. Category 3-8.
Rest (RES) (Type B): Goodall's (1989) Rest: Remain immobile, <sit> or <lie>. 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 <bipedal swagger>, <bristle>, <hunch>, <slap>, <stamp>, <charging displays> and
attacks. In reunion, bonobos show <charging display> 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: <slap ground>, <play face>,
<stand on all fours>. 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 <swagger bipedal>, <hunch and sit>, <shake branch>, <display
brachiation>, <male invite>, <arm stretch>, <penile erection> and <hair erection>, <gaze>, <glance
repeatedly>, and <approach with erect penis>. At Mahale , <leaf-clip>, <shrub bend>, <stem pullthrough>, <sex dance>, and so on. also occur (Nishida, 1997). Among the bonobos of Wamba , <bipedal
hunch>, <branch-shaking>, <rocking>, <male invite>, and <sitting hunch> 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
173
of chimpanzees
Solicit play (SOP)(Type D): Invite another individual to play. Consists of <approach>, <invitation
stamp>, <invitation slap>, <play face>, <hit lightly>, <hit and run>, <tickle>, <carry some object> 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 <extend arm>, <scream>, <approach>, <touch>, <mount>, <thrust>,
<embrace>, <kiss>, and so on. At Wamba, bonobo solicits only mother for support (Kano , unpublished).
Category 1.
Somersault
(SOM)
Mahale,
3-4
distance.
See
Spar
(SPA)
arms
in
seed
and
and
by
in
an
Squeeze
(SQZ)
of
and
extraction.
Stamp
See
(STA)
sole
making
his
feet
(1974)
and
after
may
Stamp.
Stamp
not
in
on
ground
the
care.
on
be
Stamp
the
Stamp
gently
shoulders
feet
and
the
and
(STQ)
which
(1998)
bipedal
other
are
Stamping
(SDB)
A):
while
can
solely
and
huillensis
1.
by
the
feet/foot,
by
a grin
scream
Similar
extract
both
is emitted
by
lower
, emitted
(previously
call
juices
against
of
one
the
by
firm
pressing
to
facilitate
jaw
the
with
of
feet...
"The
than
times
the
alternate
that
photograph
on
or
harder
several
for
foot,
displays
stamped
stamping
one
other.
skin.
stamps
Astanding
foot."
stamping
colobus
(with
chimp
VanHooff's
on
the
Bonobos
buttress,
stamp
while
2.
sole
on
of
charging
the
ground,
buttress,
displays.
Bonobos
of
result
tree
Some
Wamba
with
trunk,
wall
adolescent
stamp
pulled
(1973):
The
run
is
head
upwards.
The
for
the
A):
Plooij's
bonobos
of
males
bipedal
all
on
fours
in
house
or
stamp
courtship,
head
bonobo.
in
the
with
but
not
animal
may
STB.
so
walks
kept
make
is
Seen
low
major
in
bonobos.
or
trots
low
in
along
in
that
and
amplitude
or
stamp
alloparental
component
tree
Absent
manner
rather
a
buttress,
on.
lightly
play
3-7.
victim
ground,
and
social
Category
a
a rhythmical
mostly
Category
(1984)
Wamba.
youngsters,
to
contusions."
displays
"The
Mainly
invitation
on
and
done
is
of
bruising
charging
this
stamp.
in
"Stamping
on
of
Hooff
force.
in
severe
Stamp
Often
Invitation
playmate,
(1989):
in
A):
van
ground.
(1983a)
potential
Absent
Component
(Type
of
Nishida's
Goodall
(Type
A):
the
their
is shown
Garcinia
Copulation
charging
stamp
Category
watching
A):
individual.
on
of
Category
is pressed
kick
is often
may
display.
movement
those
photograph.
mouth
of
(1970)
one's
1?
of
2.
foot
Nishida
Component
quadrupedal.
(STT)(Type
hind
One
1998).
for
in
both
2.
accompanied
(1989)
a hand
a component
photograph
courtship
pith
or
1.
sound
downward
chimp
b of
Stamp
(Type
foot
or
and
as
At
a long
Category
one
Similar
is borne
Category
(1996)
Wamba.
flail
times.
travels
2-3?
(STS)(Type
another
trot
its
in
(STI)
fighting,
or
A):
Category
one
along.
the
upright.
others
hit.
bonobos.
weight
Sit.
Category
sitting
(Kano,
ground
quadrupedal
house
Kano's
sitting
bipedal
within-species
and
for
(Type
with
other
runs
A
13a,
while
invitation
May
Stamp
with
while
displays.
Stamp
he
body
of
Category
such
in
Goodall's
fruit
as
seeds
group
and
really
many
whole
bonobos
1998).
scream-like
"Forceful
that
not
Seen
Sometimes,
bonobos.
states
(1994)
(STB)
foot
charging
as
Figures
individual
alternate
also
other
Nishida
bipedal
another
in
as
lips.
do
large
Cf.
and
(1989):
alternatefeet...
See
but
Seen
"The
the
repeated
the
bipedally
(Kano,
copulation.
such
and
Goodall
She
the
4 of
running,
object
tongue
B):
stamp
Figure
Hold
Wadge.
contact)."
one
chimp
also
of
in
Usually
when
stand
eating.
bonobos.
McGrew
2.
A):
(Type
of
are
High-pitched,
stage
Absent
play
spit
they
(1996):
A):
final
play
Usually
during
in
heels".
self
parties
screaming.
Seen
Category
palate,
both
other
where
over
of
illustration.
fight,
others'
11-1
"head
pattern
occasionally
spot
(Type
Figure
(Type
each
and
the
1998).
upper
without
flexed."
at
See
(Kano,
lower
Hunt
(SCO)
a squeal).
or
this
for
of
hit
the
strongly
female
(1968)
Chimpanzees
at
copulation
bonobos
Stand
A):
A):
are
estrous
called
with
Roll
in
beginning
juveniles
(Type
(1989):
engage
Goodall
the
lutescens
knee
Squeal
of
movement,
(SPS)
Goodall
often
At
Bonobo
(SQT)(Type
hip
in
7e
A):
Voacanga
Squat
A):
infants
Figure
play.
Spit
old
(Type
hitting
during
(Type
year
while
tucked
vertical
trunk,
and
bonobos?
one
of
Category
2.
wall
Category
stamping
foot
heavily
is placed
back
rocking
of
3?
between
down
the
movements•@
"
watching
a
2.
Stand
upright.
Stance
assumed
when
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
<shake branch>, <thrust>, <jump>, and <stamp>. 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 <bark>, <push>, <arm
raise> or <shake-branch>. 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 <run>, <playpant>, <play-face> 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 <slap>, <hit>, and <throw branch>, <charge>, 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
Threat:"•@
a repertoire
toward...
The
<wraa
threatening:
(THO)
(Type
sticks,
handful
of
only
Throw
at
also
or
the
rock
See
against
also
Throw
Throw
of
the
a
infant
males.
juvenile/older
m
while
he
was
Throw
of
dry
in
courtship.
leaves
leaves
Throw
from
Absent
sand
with
one
(THS)
hand
Category
splash
throws
a heavy
Nishida,
throw
Throw
temper
tantrum
above
He
may
hold
himself
over,
,
stare>,
Kano
directed
<branch>,
<soft
(1998)
bark>
lists
"threatening
,
similar
approach",
such
as
overarm..."
stones
, rocks,
Bonobos
branches,
also
throw
, but
is aimed
at
sometimes
noise.
a specific
throw
Absent
in
objective..."
with
one
bonobos
Goodall
arm
a heavy
(Kano,
stick
unpublished).
an
(2)
once
upright
posture
Courtship
threw
a stick
as
throw
branch
Bonobo
with
pattern
one
to
long
as
arm
an
1 m
. (1)
Element
estrousfemale
from
forward
by
a distance
at
the
of
end
of
8
the
2.
dry
it
in
season,
with
male
one
hand
Category
chimpanzees
at
the
of
climax
of
Mahale
the
run
charging
, pick
display
up
a pile
. Not
used
7.
the
male
Within
the
M
charging
of
Throw
display.
group's
One
range
the
adult
male
ground
(Fanana)
is rarely
throws
covered
sand
with
sand.
still
to
extreme
case
from
high
much
it
among
above
the
Thrust
the
still
the
occurs
in
play
or
occasionally
female
Thrust
by
Thump:
a few
See
Tickle
the
grasp
the
(Type
B):
partner,
a tree,
results
M
group
lifts
intimidates
and
others
hanging
and
a
to
A):
Stand
infant
not
(see
may
as
only
rush
of
it
protests,
but
and
. At
soon.,
Mahale,
tofall
an
headfirst
to
Similar
.
over
amonginfants
appears
mother).
hands
, tumbling
occurs
about
leap
his
grooming
were
may
with
off
individuals
if
, and
ground
Typically
dominant
Monitor
Bonobos
loudly
the
transport,
withfeet
Goodall's
(1989)
copulationand...in
males
of
shows
may
monitor
behavior
how
is
seen
one's
Category
feet
Category
and
thrust
and
thrust
forth
commonly
bipedally
another.
males
back
. Thrusting
and
against
Bonobo
1998).
"Rhythmic
mounting
thrust
thrust.
(Kano,
on
thrust:
ventro-ventrally
males
a pelvic
upright
males.
Pelvic
reassurance
embrace
the
GG-rubbing
adolescent
cramps..."
by
courtship.
1.
one
in
an
beat
He
sharing,
rejection
the
screams
or
himself.
glottal
mother
THU.
adult
left
ground
or
swinging
rejecting
and
in
used
3?
chimpanzee
the
sucking,food
response
and
Plooij's
usually
fingers."
an
of
that
in
the
context
Adultfemale
like
also
chimpanzees,
but
1.
in
non-mating
context
. Courtship
8.
Rap.
(TIC)
of
hug
during
female
and
face,
in
Two
and
adult
his
Category
Occurs
bipedal(TBI)(Type
shown
male
splash
Not
Category
"The
onto
(1984)
infants."
right
(1989):
them
tantrums,
of
Plooij's
another
thrusts
adult
loud
display.
splash.
slap
temper
care
reconciliation,
mounts
Goodall
often
infant
By
pelvis...
between
reassurance
Fully
produces
charging
Throw
motherrejects
1998).
A):
the
then
on
show
an
(Kano,
the
See
screaming
the
(Type
of
ground
ground.
bonobos
movements
of
garner
(THU)
of
1998).
and
sometimes
that
stone/rock.
which
Element
B):
head
when
was
can
A):
(Kano,
The
adults
case
(aimedthrowing),
(TTT)(Type
screaming.
even
Special
7.
his/her
weaningconflict,
however,
or
the
of
stream
branches
arms
during
A):
a
Category
only
fling
with
or
object
and
male.
Wamba.
Pattern
(TSR)(Type
up,
body
grin>".
"Objects,
Mahale
is thrown
male
throw
bipedally.
into
1993).
Wamba
the
A):
stick
adult
In
of
(Type
rock
stone/rock
of
open
<fixed
"arm-raise",
arm
a loud
Category
A):
bonobos
(THW)
Throw
and
are
8.
Throw
(see
an
and
running
make
juvenile
1998).
ground,
(Type
while
an
of
andfemales,
at
(Type
in
"When
or
males
(Kano,
the
gestures
<throwat>
3-5?
Older
(THD)
full
(1989):
under
males
to
Branch
both
display
dry
the
<hunch>,
with
M.
1.
house
pant-grunting
branch-dragging
individual
"branch-sway",
thrown
Adult
a metal
A):
by
the
<hittoward>,
<scream
Goodall
be
(1989):
Category
display
and
THO.
may
Goodall
of
in
display>,
"slap",
Category
(THB)(Type
charging
on.
throwing".
splash.
branch
<charging
lip>
(1984)
so
B):
wall
behavior
and Nakamura
1-2.
1998).
"aimed
W.C.,
<armwave>,
"wrist-shake",
and
(Type
term
J., McGrew
submissive
<charge>,
B):Plooij's
(Kano,
elicit
<compressed
grass,
(THA)
uses
<slap>,
Category
Throw
Goodall
<headtip>,
"head-tip",
"pseudo-charge".
underarm
to
includes
call>,
"stare",
T.,
gestures
<stamp>,
bark>,
and
of
repertoire
<branch-wave>,
<waa
T., Kano
See
infant
Figure
or
1998).
Category
Tickle
with hand:
Tickle
with mouth:
(1984)
between
7 of
a juvenile
1.
See Tickle.
See Mouth.
TIC.
the
Nishida
in
social
Goodall
neck
(1983a)
play,
and
(1989):
shoulder
for
which
"The
or
photograph.
may
be
in
the
chimp
puts
one
groin,
and
makes
Bonobos
equivalent
may
to
or
touch,
chimpanzee
both
hands
tickling
push
on
the
movements
lightly
tickling
, hold,
(Kano,
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 <push finger into mouth>, <groom>, <inspect>, <pat> and
<push>. 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<sniff>. 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 <walk quadrupedal>,
<walk bipedal>, <walk tripedal>, <crutch>, <run>, <jump>, <somersault>, <climb>, <brachiate> and
<descend>. 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 <fish for ants>, <fish for termites>, <clip leaf>, <pull through
stem>, <bend shrub>, <drag branch>, <club>, <flail branch>, <throw rock>, <play start> , <sneeze
stick>, and <use a bed>. Great local variation exists in tool-use (see McGrew, 1992 and Sugiyama, 1997
for review). Tool using repertoire of bonobos is limited: <rain cover>, <drag branch>, <throw branch>
and <use a bed> at Wamba (Ingmanson, 1996; Kano, 1998). Category 1-8.
V
Ventral
riding:
Ventro-ventral
Vertical
Vertical
See Ride ventral.
embrace:
See Embrace
full.
climb: See Climb 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
<dunk>,<keep. immersed>, <rinse>, and <stamp>. 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).
Albrecht
REFERENCES
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 withingroup 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 semicaptive 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.
Handling editor: Saitou Naruya