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chimpanzee-behaviour-ethogram

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dorsal, put ventral, scoop infant, solicit riding, solicit companion, and grasp and push shoulders. Cf. go ahead. See video. dermatophytosis Infections skin disease on face of chimpanzee. See video. D 54 dig descend Climb down from elevated site: Includes six patterns: descend by brachiation, descend tree trunk feet first, descend tree trunk head first, leap down, slide down vertically and descend orthograde feet first. Cf. fall. See video. D descend by brachiating While progressing by brachiation, supportive bough gradually bends down until ground can be reached with feet. Also in bonobos. See video. descend orthograde feet first Grasp branch below with feet to move to another tree below. See video. descend tree trunk feet first Climb down backwards. Seen in bonobos. See videos. descend tree trunk head first

Climb down forwards. Seen in bonobos. See video. detach See kidnap. detain Hold body part of partner such as leg in order to continue social play, etc. while partner tries to escape. Mother restricts infants movement e.g., by grabbing its legs when it approaches possibly dangerous object such as adult male. See video. detour Deliberately take roundabout route. When chimpanzee seeks to avoid dominant rival, or when shortest route is difficult, e.g. gap in tree or crossing river. Kanos Detour and Pass with detour for bonobos of Wamba. See video. diarrhea Loose stools often seen after eating fruits of Pycnanthus or Toddalia. Also when being attacked by another, or traveling near territorial boundary. See sickness. die Lie motionless and breathless. Dead body usually found on ground in bush

along animal track. See death. See Fig. 8. dig Make hole in ground using hands or stick. dig for army ant with hand/stick 55 D Fig. 8 Die: Infant of Wakasunga died of disease dig for army ant with hand/stick Chimpanzees of Bossou dig up army ants nest by hand but also use digging stick. Absent at Mahale. dig for underground storage organ by hand Dig up root such as Aeschynomene sp. tree by hand while sitting or standing quadrupedally. Chimpanzees of Tongo dig up tubers as substitute for drinking. Cassava is dug up and eaten at Bossou. See video. dig for underground storage organ with tools Dig up underground storage organs with tool, reported only from Ugalla. Absent at Mahale. dig for water by hand Dig well

in wet ground of dry streambed using one or both hands to get drinking water. At least five individuals of M group do this. Behavior common at Semliki. Bonobos of Lilungu and Wamba do not dig for water. See also remove objects from water surface. 5. dig for water with stick Use tool to dig wet ground of dry streambed to get water. Only one juvenile female of Mahale seen to do this. Absent in bonobos of Wamba. 9. 5 dip ant directly dig stingless bees nest with stick Customary at Gashaka. Also present at Bwindi. Absent at Mahale. D dig termite nest with stick Dig into termite nest with stout stick. Known from Ndoki and Goualougo. Absent at Mahale. dip ant and wipe Ant-dip, Ant-dip-wipe. Dip wand

of vegetation into massed driver ants, and when ants climb wand, remove ants with sweeping movement of loosely flexed hand and pop ants into mouth. Custom at Gombe, Assirik, Gashaka, and Goualougo. Present at Bossou. Likely to occur at Kalinzu, Fongoli and Ngotto Forest. Absent at Mahale. See Fig. 9. dip ant directly Whiten et al. Ant-dip-single. Chimps dip short stick into masses of driver ants and when ants climb stick, quickly mouth them off stick. Custom at Fig. 9 Dip ant and wipe: Adult female, Skosha, of Gombe dips and wipes Dorylus ants dip fluid 57 D Fig. 9 Bossou, Ta and Goualougo. Present at Gombe and Nimba. Absent at Mahale. See video. dip fluid Whiten et al. Fluid dip: Use probe to extract fluids such as

honey. Some immature chimpanzees use sticks for drinking water in tree hollows. See also leaf-sponge. Customary at Gombe, Mahale, Kibale, Lop, Assirik and Ta. Present at Goualougo. 8. See video. dip fruit wadge into water Chimpanzees of Ta dip fruits and their wadges of Sacoglottis gabonensis into water apparently to suck out most of sweet juice. Absent at Mahale. 58 display as contest dip hand and lick water Immature chimpanzee immerses hand into stream or crevice of tree and lick water from fingers, knuckles, or back of hand. 8. See video. D directed scratch See scratch self. disarm See confiscate. discard Abandon object. See video. discard fruit skin with mouth Remove fruit skin with movements of lips, teeth and tongue, then abandon it by mouth. See video. disperse

seed Excrete feces away from place where eaten fruits originated, so chimpanzees unintentionally disperse seeds. See swallow seed. See Fig. 10. Fig. 10 Disperse seed: Seeds of fruit recovered from chimpanzee feces display as contest When male does charging display, more dominant male rushes up and displays against him. May elicit displays by more dominant males, and chain display away 59 may continue up to five individuals. If final displayer is alpha male, display contest ends. 7. See video. display away Goodall: After interaction with another, usually an attack, the chimp departs in a charging display. Seen in bonobos. display, charging Plooij Charging display. Goodall s description of Charging display is modified here. May move in slow rhythmic gait, run at moderate speed or very fast. Display patterns

include scratch dead leaves, drag branch, shake branch and sway woody vegetation, bristle, compress lips, slap, stamp, slap-stamp, flail, throw, drum, rake, swagger, bipedal and pant-hoot. May be bipedal, quadrupedal, or tripedal. Typically male display though some females do it also. Goodall discriminated Non-vocal and Vocal displays. The former when displayer makes no calls, tends to be directed toward another individual, and may be followed by attack. It is typical component of male dominance rivalry. Vocal display is accompanied by pant-hoot and typically not directed toward any other chimpanzee. Common during reunions, food excitement and when individual has been frustrated in obtaining desired goal. At Gombe, charging display is sometimes followed by beat chest, but not at Mahale. Bonobos charging display is accompanied by charge, stamp, and sometimes

by drag branch. See video. display past Goodall: 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 regularly hit human observers during charging displays. Seen in bonobos. 5. display, rain Goodall Rain display or Rain dance. At Mahale, two types of displays occur at onset or during heavy rain or storms. Not directed at other individuals. One is a noisy, active display accompanied by pant-hoot, charge, run bipedal, stamp etc. Other is prolonged, slow tempo display, of which main components are sway woody vegetation, shake branch, run bipedal, climb rapidly, brachiate and leap. Customary at Gombe, Mahale, Kibale, and habitual at Ta and Budongo. At start of storms

or heavy rain, adult male bonobos may run few meters while calling loudly, but no other activities such as those of chimpanzees occur. 3. See video. display, streambed Charging display done in water course. Goodalls Stream bed displays resemble rain displays but occur in streambeds. Goodalls Waterfall display probably should be included here. At Mahale, main components of streambed display are run, leap, drag branch, throw splash, D 0 dominant shake branch, swing and sway woody vegetation. Absent in bonobos. 5. See video. display toward Goodall: Display directed toward one or more others. Seen in bonobos. See video. D distance scratch See scratch self distantly. distract Shift anothers attention away from what it tries to do. For example, mother grooms or plays with infant to divert its attention

from sucking from nipples during weaning. If infant is male and throws temper tantrums in response to his mothers refusal, she sometimes allows him to copulate with her. Goodalls Distract. Also in bonobos of Wamba. See video. distress call Includes hoo, staccato call, whimper, and whimper-scream. 3. See video. Dja Study area of Pan troglodytes troglodytes in Dja Biosphere Reserve of southeast Cameroon. Research done by Deblauwe and colleagues from 2002 to 2005. dominant Higher-ranking of two individuals. See also pant-grunt. See Fig. 11. Fig. 11 Dominant: Subordinate male showed temper tantrum when dominant male approached dorso-ventral embrace 1 dorso-ventral embrace See mount. drag and circle Older individual holds arm or hand of playmate in mouth and drags other in social play. Performer may circle around tree. 5.

See video. drag branch Goodall Branch dragging: A branch broken from a tree, or one lying on the ground is dragged by one hand during a charging display Finally, branch is typically thrown forward. Widely known across Africa: Ndoki, Bossou, Ta, Lop, Gombe, Mahale, Kibale, Budongo and Goualougo. In bonobos of Wamba, branch-dragging displays have many more behavioral elements than those of chimpanzees. Late adolescent male bonobos use display to tease and to challenge males higher in rank. Bonobos sometimes drag branch while walking without stamping or vocalizing. Bonobos of Lomako also drag branch. See video. drag by hand Goodall: Drag by hand along ground individual, branch, palm frond, large food object, etc. Shown in aggression, threat, play, and maternal solicitude. Plooijs Drag. Bonobos of Wamba drag only

branches. drag by mouth Drag while holding object, animal or another individual in mouth. Cf. pull with mouth. 3. drag carcass by hand Drag large carcass by hand along ground, moving forward. Simple transport or element of charging display. 5. See video. drag carcass by mouth Drag large carcass forward by mouth along ground. Simple transport. 5. drag dry leaves Walk forward fast quadrupedally with head down and shoulders hunched, while pushing dry leaves with hands and feet, producing distinctive sound. Adult males do in charging displays. Adolescent female, Ivana, teased newcomer female, Qanat, by displays including this. Done by juvenile as solicitation of play. Cf. push leaf-pile. 7. See video. drag other by hand Goodall Drag other. Drag by arm and leg in aggression or play. Mother

often drags older infant when infant delays departing with her. D 2 drip Adult male may abuse infant by dragging it forcibly along path. Kanos Drag other and Pikas Grab-push-pull for bonobos. See video. D drag other by mouth Drag another individual while holding hand, arm, foot, leg, nape or back in mouth in play. Usually move backwards. 3. See video. See also video hold body part in mouth. drag to kill Adult male, Musa, once ran and dragged with one hand large red colobus monkey that he had captured, so that monkey was hit and injured by woody vegetation and choked by woody vines, causing death. Tactic prevented monkey from biting chimpanzee predator. 8. See videos. drape Goodall Drape: Drape bunch of fruits, leaves, or skin of

colobus over shoulders, nape or head. Category 15. See video. drink Goodall: Drinking of water or other liquids. This includes drinking directly by leaning over and sucking from the water source in a quadrupedal or sitting posture; dripping water from the fingers into the mouth; dipping fingers into water and licking them; drinking with or licking back of hand, licking drops of water from surfaces such as leaves, hair, and so on See also dig for water, keep water in mouth, leaf-sponge, leaf-spoon, and remove objects from water surface. Bonobos rarely drink from running or standing water. See video. See also videos leaf-spoonand remove objects from water surface. drink from hole in tree Put hand, fingers, or leaf sponges into hollow of tree, then lick water extracted. Customary

at Gombe, Mahale and Bossou. 5. See video. drink from lake Chimpanzees of lakeside groups, B, K, M, N, and L groups, drink from Lake Tanganyika. Some afraid of high waves and do not do so. Category 1. See video. drink from stream Chimpanzees of Mahale mostly depend on streams and rivers for drinking water. See video. drip Juvenile male, Xmas, once dribbled water repeatedly from back of hand at pool. Solo play with water. 9. See video. drop 3 drop Let object fall, occasionally deliberately onto another individual or human observer. See video. drop bark bits Remove by hand without obvious purpose outer bark little by little and drop. Shown by nervous individual or when waiting for companion engaged in long bout of sleep, grooming, or fishing

for ants or termites, etc. See also kill time. drop branch Drop branch from overhead to threaten human observer below, sometimes hitting target. Some adult males forcibly kick dead branches, detaching them, so that they fall. Occasionally seen at Mahale when chimpanzees were not well habituated. Seen in female chimpanzee in Gabon. Ingmanson described Object dropping, e.g. branch in bonobos of Wamba. See video. drop infant Mother drops older infant when mother tired during long bout of travel. Includes shake rump, shrug, lower head and shoulder, reject-sit, push, lower rump, etc. 5. See video. See also video lower rump. drop self Release branch and let self drop to lower vegetation or ground without leaping. Common practice when leaving tree. Youngsters in play repeat sequence of climbing and dropping.

Occasionally fall occurs accidentally when branch breaks. Cf. leap down and fall. See video. drum Component of charging display. Goodall: Hit and/or kick tree trunks, especially those with buttresses. Often they grab the buttress with their hands and stamp on it with their 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. Plooij Drum. Mahale chimpanzees do same. Drumming

patterns vary individually. For example, one male stands erect and beats walls of research buildings with only one palm, another does so with two palms, third one with one fist, fourth one kicks with one foot, and fifth one pounds with both feet. See also kick and slap. D 4 dunk face Drumming reported from Kibale and Ta. Bonobos drum on buttresses, most often immediately after onset of traveling. Cf. beat chest. See video. D drum belly See slap self. dunk Immerse object in water without releasing it. Juvenile male, Michio, once dunked fruit in puddle, then sucked it. Adult male, Musa, dunked colobus skin in stream apparently to clean it. Wadge dipping observed at Ta. Absent in bonobos of Wamba. leaf-sponge is kind of dunking. See wash.

5. dunk face Juveniles repeatedly submerge face in river in play. See video. E E eat Remove foodstuff from substrate directly with mouth or by hand, bite, chew, and swallow it. Sometimes, process food or wadge before swallowing. Goodalls Feed. Kanos Feed. See video. eat algae Enter stream and eat algae, standing quadrupedal in water up to 30 cm deep. Adolescent immigrant female, Sally, only few times seen to eat. Likely customary in groups inhabiting higher elevations of Mahale. Also recorded at Bossou and Odzala National Park. See videos. eat ant Ants are eaten everywhere, but species eaten vary from place to place. For example, army ants are not eaten at Mahale, but at many other sites. Bonobos of Wamba do not eat ants, but Lilungu bonobos eat

adult and larvae of Tetraponera. Humans in the tropics eat ants. See eat Camponotus ant, eat Crematogaster ant, and eat Oecophylla ant. See video. See also videos brush away from branch, brush away from self and fish for carpenter ants. eat beetle larva Ingest larvae of Coleoptera. Occasionally seen at Mahale. Regularly eaten at Bossou. Likely to be eaten at Ngogo. See video. eat blossom Ingestion techniques resemble those for eating leaves. Eaten by bonobos. See video. 5 eat fruit, pulp eat Camponotus ant Eat carpenter ants by cracking open grass stem or by using tools. 7. See video. E eat carcass Scavenge. Eat carcass of mammals, such as red colobus, red-tailed monkey, bushbuck, blue duiker, etc., not killed by self or ones companions. Scavenging occasionally seen at

Gombe and Mahale, but virtually absent at Ngogo. Absent in bonobos of Wamba. 3. eat Crematogaster ant Eat Crematogaster ants by splitting open grass stem or dry branch with teeth, breaking off large branch with hands or occasionally hands and feet. See break branch, break branch with foot and brush away from branch. 7. See video. eat Dorylus ant Dorylus ants are available, but never eaten at Mahale. See also dig for army ant with hand/stick, dip ant and wipe and dip ant directly. See video. eat egg Eggs of birds such as francolin and guinea fowl eaten by chimpanzees of Mahale and Gombe. Gombes chimpanzees readily ate domestic fowl eggs when offered. Some Mahale chimpanzees also ate them. 5. See video. eat eye mucus Eat discharge from

eyes. Rare. 9. See video. eat feces Goodall Coprophagy. At Mahale, eating feces occurs rarely. Adult female, Chausiku, picked out and ate undigested seeds of Saba comorensis from her feces. Gombe apes picked undigested meat from feces. Assirik chimpanzees picked out baobab seeds from feces. Semliki apes defecated directly into their own hand and raised feces to mouth to reingest seeds of Saba comorensis. Bonobos of Wamba reingested seeds of Dialium. 8. eat fruit, inner skin Scrape and ingest inner skin of fruit such as Saba comorensis. See video. eat fruit, pulp Pick fruits in tree, from shrubs or herbs on ground, or pick up fallen fruits from ground, then, open up fruit shell with hands or mouth, and eat gall 7 ingest contents. See video. See also

video spit seed. eat gall Gall of insects inhabiting parts of plants such as Milicia. 5. See video. eat honey Insert finger or probe into honeycomb and eat honey of honeybees, or gnaw with incisors to widen entrance of nest of stingless bees. Honey eating widespread in chimpanzees. Bonobos of Lilungu eat honey of stingless bees. eat infant Cannibalism. Chimpanzees of Mahale occasionally eat infants. Known also from Budongo, Gombe, Ngogo and Ta. Absent in bonobos. See Fig. 12. Fig. 12 Eat infant: Alpha male, Ntologi, ate infant of newly-immigrated female eat insect Chimpanzees of Mahale eat ants, honey bees, stingless bees, termites, and beetle larvae. See also eat ant and eat termite for details. eat invertebrate See eat insect. Chimpanzees eat only insects, but bonobos eat earthworms.

E 8 eat phloem eat leaf Put leaves into mouth and ingest them. Bring mouth to leaves or bring plucked leaves to mouth. Important cropping technique is pull through. See video. See also videos pull through with mouth and store. E eat meat Ingest teeth, bones, bone marrow, brain, entrails, stomach and intestinal contents, and skin, in addition to muscle. Eating meat unusually prolonged, sometimes for more than 3 h. Prey include red colobus, blue, red-tailed, and vervet monkeys, yellow baboon, greater galago, infant chimpanzee, blue duiker, bushbuck, bushpig, giant rat, rock hyrax, civet, mongoose, sun squirrel, francolin, guinea fowl, chicken, weaver bird, and black kite. Meat-eating occurs throughout chimpanzee range. After Ueharas review, black and white colobus, white-cheeked mangabey and red duiker and banded mongoose recorded as

prey. Bonobos eat meat of blue, bay, and black-fronted duikers, flying squirrel, fruit bat, galago, red-tailed monkey, black mangabey, Wolfs guenon, rodent, birds, etc. Recorded at Wamba, Lomako, Lilungu, and Lui Kotale, but less often and extensively. See also hunt. See video. eat nasal mucus Eat nasal mucus after picking nose or sniffling. Adult male often eats nasal mucus of his partner during social grooming. See also pick nose, probe nasal passage, sneeze and sniffle. See video. See also video sneeze. eat Oecophylla ant Grab weaver ants nest with one hand, peel it apart on ground, quickly eating ants. 7. See video. eat petiole Eat only petiole of Ipomoea rubens, Cordia millenii, Myrianthus arboreus, Vernonia amygdalina, etc. Category 35. See video. eat phloem Eat phloem or inner bark

rich in sugar and protein by scraping inner surface of outer bark with incisors. Eaten widely by humans. See Fig. 13. eat pith 9 E Fig. 13 Eat phloem: Inner bark of Brachystegia bussei teeth-marked by chimpanzees of K group eat pith Eat soft, juicy piths of herbs such as elephant grass, gingers, ginger lilies, and Marantochloa, herb vines such as Ipomoea, woody vines such as Landolphia and Saba, and shrubs such as Vernonia by removing outer surface with teeth. Pith of Megaphrynium eaten by both bonobos and humans at Wamba. See video. See also video peel with teeth. eat resin Scrape resin of Terminalia mollis with incisor or pick large lump of resin of Albizia glaberrima with fingers. Bonobos and African pastoralists eat resin. See video. eat

rock Nibble or lick rock. See lick rock. Category. eat root Dig ground to reach and eat live root of tree of Aeschynomene sp. Cf. dig. See video. eat seed Crack seed pod with teeth, chew and ingest seeds of plants such as Pterocarpus, Diplorhynchus, Baphia, Parkia, etc. Bonobos eat Pterocarpus tinctorius. Category 12. See video. See also video beg. eat semen Females, more often than males, eat semen ejaculated in copulation. 7. See video. 70 E elbow clamp eat termite Topple tower of termite mound with hands and pick up soldiers or winged reproductive forms of termites. Fishing for soldiers of Pseudacanthotermes recorded only for K group chimpanzees. Fishing for soldiers of Macrotermes has been seen in chimpanzees of B group. Widely known throughout chimpanzee range. See

fish for termite and shake wet arm to catch termite. Bonobos of Lilungu eat termites of different genera from those eaten by chimpanzees by inserting fingers into hole opened manually. Termites eaten widely by tropical people. See videos. eat termite soil Pick up and ingest small pieces of soil from termite towers of Pseudacanthotermes. Humans and bonobos eat termite soil. See video. eat vertebrate See eat meat. eat vomit See reingest vomit. eat with foot Pick up and put food into mouth with foot. Infant male, Caesar, plucked several ripe fruits of Garcinia with foot and ingested them. 9. See video. eat wood See eat xylem. eat xylem Eat dry dead wood of Pycnanthus angolensis, Ficus vallis-choudae, Garcinia huillensis, etc. Unknown elsewhere. Category. See video. Ebo Forest Study

area of Pan troglodytes vellerosus in Ebo Forest, Cameroon. Research organized by B.J. Morgan and E.E. Abwe from 2005 to present. ejaculate Ejection of seminal fluid from penis. Males begin to ejaculate at 9 years old at Mahale. Semen often eaten by both sexes. See eat semen. See video. elbow clamp See clamp elbow. emasculate 71 emasculate Remove testicles, occasionally in extreme male-male competition. Evidence from Kanyawara and Ta, as well as from captivity. Some male bonobos of Wamba lack testicles but cause unknown. 3. embrace full Goodall Ventro-ventral embrace: Two individuals face each other and each puts one or both arms around the other. van Hooffs Embrace, Plooijs Embrace full. Contexts include greeting, reassurance, alliance, and reconciliation. Embrace full by adult males may be accompanied by thrust.

In bonobos, embrace full posture seen only in genito-genital rubbing by females and ventro-ventral copulation between males and females, so both embrace-full in erect posture and embrace-full by adult males are lacking. 3. See video. See also video console. embrace half Plooij Embrace half. Goodalls Arm round: Puts one arm around another as in a half embrace. Often occurs when one individual pant-grunts to another. Adult male puts his arm around another male while walking and pant hooting. Occurs in copulatory solicitation in bonobos. Category 1. See video. emigrate Adolescent female disperses from natal unit group and enters another group for reproduction, at about 11 years old. Parous females rarely leave resident group. See also transfer. enter hole Enter cavity, such as burrow of aardvark in exploration or

play in juveniles. See video. erect hair See bristle. erect penis Plooij PEN. Goodalls Penile erection. Occurs during sexual and food excitement. See also adduct penis. Kanos Penile erection for bonobos. See video. See also video open thighs. escape Leave area when more dominant individual targets present in party. For example, when estrous female moves to another sex partner, she departs while often looking back or glancing at other adult males to monitor possible attack. 3. E 72 extend hand estrous cycle See swelling of sexual skin. estrus Female copulation confined mostly to period of maximal swelling of sexual skin, which includes day of ovulation. See swelling of sexual skin. See Fig. 14. E Fig. 14 Estrus: Sexual skin with maximal swelling expel Whiten et al. Expel/stir. Nishidas

Expelling stick. Push stout stick into hole of tree or other cavity and stir it violently side to side or back and force with arm. Mahale chimpanzees try to expel rock hyraxes from cavities in rocks. Customary at Gombe, Mahale and Ta. 3. See video. extend Extend arm or leg to another individual. See video. extend arm as ladder Nishida: Caregiver extends arm upwards in tree, to allow infant to climb up easily, by making ladder with arms. See also aid in locomotion and extend leg as ladder. Absent in bonobos of Wamba. 7. See video. extend hand Goodall Extend hand. van Hooff 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 about midway between pronation and supination. The hand may be bent at the wrist so that its back is turned to the extend hand and put knuckle on ground 73 partner with the fingers bent or fully stretched Goodalls Extend hand limited to gesture with wrist and fingers extended and palm up or down. Goodall described varying contexts in begging, as reassurance to out-of-reach subordinate, and when submissive chimpanzee seeks reassurance after aggression, or solicits support from third party. In mother-infant interactions, extend hand is contact-seeking. For bonobos, extend hand seen only as extend hand, palm downward or extend hand, palm upward below in food begging or mother seeking contact with offspring. See video. See also video

extend. extend hand and put knuckle on ground Put arm forward and place knuckle on ground. Often performed to solicit grooming as in extend hand to hold tree. Seen in bonobos. extend hand, palm downward Goodall Arm stretch: The arm or arms are extended toward another. The palm of the hand usually faces down. Seen in courtship or greeting. Also as solicitation of play. Mother may invite infant to riding on her or to go ahead. van Hooff Stretch over, Plooij Extend hand, palm downward, Pika Reach, Nishida Arm-stretch. Seen in bonobos. See video. extend hand, palm sideways Arm extended to another with elbow bent, palm sideways, and fingers straightened or flexed, occasionally accompanied by panting. Greeting and when mother seeks to retrieve infant from rough social play.

Function may be reassurance, appeasement, or reconciliation. 7. See video. extend hand, palm upward Plooij Extend hand, palm upward. Bipedal or quadrupedal. Used to solicit play, to beg, when mother solicits infant to ride on her belly or back, or to pant-grunt to dominant individual. Once, juvenile female, Flavia reached left arm to TN while standing bipedally. Seen in bonobos. See video. See also video pant-grunt with bent elbow. extend hand to beg Plooij Beg with hand: Stretch hand to possessors hand, mouth or food. Goodalls Beg hand-to-food, hand-to-hand and hand-to-mouth. Kanos Food beg: hand to hand and hand to mouth. Cf. mouth for begging. See video. extend hand to hold tree Reach arm forward and grasp tree trunk. Often performed to solicit grooming. Seen in bonobos. extend

leg Reach leg instead of hand to another individual, with similar function to extend hand. Shown especially when performing extend hand E 74 eyes open difficult, e.g. in tree or when performer holds infant and food. Seen in bonobos. See video. extend leg as ladder Mother extends leg to help infant to come to her. See also aid in locomotion and extend arm as ladder. See video. E extended grunt See grunt, extended. extract Sanz and Morgan Gathering an item or fluid with a tool for the purpose of extracting it from a location that is difficult to access manually. Includes fish, dip, pick out bone marrow, etc. 3. eyes closed See close eyes. eyes open See open eyes. F F face close See peer. fall Drop by

accident from higher level to lower level. Cf. drop self and leap down. See video. fall over backward Topple backward from seated position, arms extended. Thud caused by hitting dry leaves; sometimes after being pushed. Signals play solicitation as attention getter. Weanling uses to attract mothers attention. Cf. somersault, backwards and headstand. 5. See video. family Chimpanzee family consists of mother and her immature offspring and adult sons. If daughters do not emigrate, it may become group of three or more generations. See Fig. 15. Fig. 15 Family: Old adult female, Wabunengwa, and her adolescent daughter and infant son 75 7 fish fantasy play See play, imaginary. farsightedness See presbyopia. fart Break wind. See video. F fashion Behavior learned socially, shared by at least most members of at

least one age or sex class, and transmitted horizontally within one generation. See also culture and tradition. feed See eat. female solicit See solicit copulation. fend Goodall: 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, e.g., fend against being tickled. See also clamp elbow and cover nipple. Also in bonobos. See video. fight Mutual attack. See attack. See video. Filabanga Study area of Pan troglodytes schweinfurthii in western Tanzania. Research done from 19 5 to 19

by T. Kano and colleagues. fill mouth with food Stuff food by hand into mouth. See video. finger in mouth See push finger into mouth and take finger in mouth. finger wrestling See wrestle with fingers. fireman slide See slide down vertically. fish Insert flexible strip of bark, vine, twig, grass etc. into nest of social insects, withdraw and pick off insects with lips, teeth or tongue. Absent in bonobos fish for army ant 77 of Wamba, Lomako, and Lui Kotale. Category 13. See video. fish for army ant See dip ant and wipe and dip ant directly. fish for carpenter ant Insert fishing probe such as peeled bark, vine, twig, modified branch, midrib of leaf, or split wood into entrance of wood-boring carpenter antsnest, withdraw probe laden

with soldiers, and take them into mouth with lips and tongue. Usually arboreal and aseasonal, sometimes continues for more than 3 h. Customary at Assirik, Lop, and Gashaka. Chimpanzees of Kasakela group at Gombe do not fish for carpenter ants, but those of Mitumba do. 5. See video. See also videos fish, supplant, take, use tool and wait turn. fish for termite Resembles ant fishing. Fishing for the termites of Pseudacanthotermes spiniger at Mahale only in K group. Fishing for Macrotermes only in B group F 78 flap Collins 1985). Fishing for termites not seen in M group, apparently because Macrotermes is absent in range. Recorded for all subspecies of chimpanzees; Mt. Assirik, Bossou, Fongoli, Gashaka, Campo, Ndoki, Goualougo and Belinga, in addition to Gombe. Not recorded for

bonobos. Bantu people fish for termites with more elaborate techniques. See videos. F fission and fusion Chimpanzee unit groups repeatedly split into small parties, then later rejoin into larger ones, depending largely on abundance and distribution of food supply. Also bonobo unit groups and human hunting bands. fist See hit. fist ground See hit ground with fist. fixed stare See stare fixedly. flabby bottom Goodall: Loose and wrinkled sexual swelling during the final stage of detumescence. flail Brandish arm or branch for threat display, solicitation of play, or solo play, or brandish and beat animal to ground in order to kill it. See also flail arm or flail long object. See video. flail arm Brandish or bring down arm as threat display or play solicitation. 2. See video.

flail long object Brandish branch or other long object. Goodall: Brandish branch in hand and waves this weapon at an opponent. Plooijs FLL. Kortlandts Tree-swish. At Mahale flailing at another rarely done by adult male. Flailing at human observer more rarely done by adolescents. Adult male of Mahale caught, flailed and hit to death monkey or bird against ground. Cf. shake branch. Branch-flailing is element of branch-dragging among bonobos of Wamba. 2. See video. flap Goodall: downward slapping movement of the hand, usually repeated several times, in the direction of another individual. Often seen in flee 79 female squabbles Plooijs Flap. Absent in bonobos of Wamba. 5. flee Goodall Flight: rapid progression away from an alarming or dangerous stimulus Also seen in chasing play. Elements include scream, run,

climb, jump, and descend. van Hooffs Flight. Kanos Flee. See video. flee after startle Goodall Startle flee: 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. Also in bonobos. See video. flee from colobus male Adult male colobus occasionally descends to ground and chases chimpanzees away. Both adult males and females fearful of colobus male and run away. Also chimpanzees of Gombe, but those of Ta do not flee from colobus. 7. See video. flight See flee. flinch See bend away and parry. flip lip Goodall

Lip flip: The upper lip is rolled up and back towards the nose. Usually relaxed situation Plooijs FLI. Function of everted lip facial expression unknown, but may reflect boredom. Absent in bonobos. 5. See video. flop Lie supine and kick, rake, and wriggle on ground in pile of dry leaves, like flopping fish on line. Solo play of juvenile male, Xmas, perhaps idiosyncratic to him. See video. fluid-dip See dip fluid. fly whisk See whisk fly with leafy stick. fold leaf Tonooka Leaf-folding: Tear leaf or leaves with hand or mouth, stuff into mouth, chew them repeatedly, take them out of mouth and hold between index and middle fingers, soak them in water in hollow, pick up and suck water from them. Final shape is not like sponge,

but folded leaves. Cf. leaf-sponge. Only at Bossou? Absent at Mahale. See Fig. 17. F 80 fondle penis F Fig. 17 Fold leaf: Leaf folded by Bossou chimpanzee follow Trail after, trying to maintain close proximity to another. Goodalls Follow. Plooijs Follow. Kanos Follow. See videos. follow in contact Follow another with hands placed on back or shoulder of leader. This occurs when follower feels uneasy, is in stressful situation, enlists support from leader, or is interested in sexual swelling of leader. See video. follow specific female Newly-immigrated female chooses, follows, and forms affiliative relationship with one particular resident female by grooming her and caring for her infant. Also in immigrant female bonobos. See video. fondle penis See fumble penis. Fongoli 81 Fongoli Study area of Pan troglodytes

verus in southeastern Senegal. Research organized by J. Pruetz from 2001 to present. food grunt See grunt, food. food-pound on other See pound food on other. food-pound on wood See pound food on wood. food scream See scream, food. food sharing See share food. food transfer See share food. friendship Long-term association with mutual dependence. Cf. coalition. Past-prime male, Kalunde and past-prime female, Nkombo, traveled and rested most of time together, and she supported Kalunde when he was challenged for alpha status by adult male. See video. fumble clitoris Adult or adolescent female repeatedly touches own clitoris. Males occasionally grasp clitoris of young females. Rarely seen. 8. See video. fumble nipple Manipulate own nipples with thumb or finger for self-reassurance or self-stimulation. Done by few chimpanzees. Occurs when

frightened or nervous when dominant individuals closeby. Usually thumb only or thumb and index finger are used. Adult male, Alofu, dexterously alternates between right and left nipple when anxious and pant-grunting. When using right hand, simultaneously fumbles right nipple with thumb and left nipple with little finger, but when he using left hand, fumbles left nipple with thumb and right nipple with little finger. Nipples are massaged by circular movement of thumb and finger. Captive adolescent male bonobo did same in selfreassurance. 8. See video. F 82 funny face fumble penis Male occasionally fumbles his penis by hand, leading to penile erection. At Mahale, never culminates into ejaculation. For bonobos of Wamba, only one case of fumbling penis, which did not lead to ejaculation. 7. See video. fumble

penis with foot Rub penis up and down with own heel, toe or sole. Few juvenile, adolescent and adult males do so, without ejaculation. Cf. shake penis. See video. F funny face Adult male, Alofu, shows unusual facial expression to estrous females and adult males. Function unknown, but as he showed funny face to female unwilling to follow him despite frequent solicitation for consortship, function may be to reassure. See video. G G gallop Goodall: Fastest run of the chimps. van Hooffs Gallop. Pikas Gallop?. See video. gang attack See attack concertedly. Gashaka-Gumti Study area of Pan troglodytes vellerosus within Gashaka-Gumti National Park, Nigeria. Research organised by Sommer from 2002 to present. gaze Includes stare fixedly, peer, and watch. See video. genital inspection See inspect genitals. genito-genital contact

See rub genitals. geophagy See eat termite soil. gestation period Average = 229.4 days, range = 203244 days. G-G rub See rub genitals. give Possessor hands over food to another who may or may not have begged for it. Food given includes meat, fruit, leafy branch, and dead wood. Size of given food not necessarily large. Rare at Mahale. Unknown in bonobos of Wamba. 8. See video. T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 8, Springer 2010 83 84 grapple give birth Childbirth usually occurs in bed at night. See also travel alone after childbirth. glance Goodall: looks toward another for less than 2 s. Subordinate chimpanzee looks only briefly at food, estrous female, or other object of competition when dominant

male present. Cf. look, peer, stare fixedly, and watch. Kanos Glance for bonobos. See video. G glottal cramps See choke in tantrum. glove Wear cylindrical bark ring on lower arm. Female infant, Imani, did this once. See video. gnaw See mouth. go ahead Travel ahead of party, e.g. mother goes ahead of infant. Cf. depart together. See video. Gombe Study area of Pan troglodytes schweinfurthii in Gombe National Park, Tanzania. Research organized by Jane Goodall, continuing from 19 0 to present. Goualougo Goualougo Triangle, study area of Pan troglodytes troglodytes in NouabaleNdoki National Park, Republic of Congo. Research organized by C. Sanz and D. Morgan from 1999 to present. grab Goodall: 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 in aggression, grooming and play. Bonobos grab in same contexts. See video. grab and shake Grab anothers body part and repeatedly shake it. Threatening but also playful. Also, as solicitation of play. 7. See video. grapple Goodall: When two individuals go into a clinch with arms locked and heads bowed, and sometimes roll over and over. Individuals may be grasp 85 fighting or playing. Not uncommon when adult females fight one another. Bonobos do same. See video. grasp Grasp object such as branch to carry it or to support body. Infant may grasp mothers nipple during rest. See video. grasp and heave Grasp partners shoulders with both hands, fall on back, kick upwards

to partners belly, pushing partner forward and away. 7. See video. grasp and push shoulders Infant, Mitsue, in bipedal posture, grasps hair of mothers shoulders and pushes them upward, so mother departs with Mitsue on her back. Soliciting departure likely originated as intention movement of dorsal travel. 9. See video. grasp hand Goodall Hold hand. Dominant individual grasps hand extended by subordinate for reassurance, but usually without shaking motion. Also in hand-wrestle play. Absent in bonobos of Wamba. 5. See video. greet Friendly reunion as Goodalls Greeting behavior: Typical friendly behavior includes bob, crouch, touch, kiss, embrace, groom, present, mount, inspect genitals, grasp hand, etc. Vocalizations associated with greeting behavior are pant-grunt, pant-hoot, grunt, pant-bark, scream, etc. In bonobos, genital contact behavior and peer may be comparable to

greeting, in addition to groom, present, mount, inspect, and grasp hand, but the other greeting elements listed above are lacking. 2. See video. grimace See grin. grin Goodall: Facial expression with the corners of lips are drawn back, exposing the lower, or both upper and lower teeth. Includes van Hooffs Silent bared-teeth display. Often shown by chimpanzee threatened by another, when copulating, begging for foods, or simply watching dominant chimpanzee. May accompany male courtship. Facial expression sometimes accompanies vocalizations such as scream and squeal in copulation, and may reflect fear or frustration; but also reflect pleasure during copulation. An adult male, Fanana, occasionally shows grin and food screams during feeding on fruits such as lemon or Saba fruit or termites, which seems to express pleasure. See video. G

8 groom by hand grin-full-closed Goodall Full closed grin: Both upper and lower teeth are revealed by the horizontal retraction of upper and lower lips, and the teeth are closed, or almost closed. Plooijs Grin-full-closed. Kanos Grin for bonobos of Wamba. See video. grin-full-open Goodall Full open grin: Both upper and lower teeth are revealed by horizontal retraction of upper and lower lips, and the teeth are partially or widely open. Plooijs Grin-full-open. Grin for bonobos at Wamba. See video. G grin-low-closed Goodall Low closed grin: Only the lower teeth are revealed, and the teeth are closed, or almost closed. Absent in bonobos?. See video. grin-low-open Goodall Low open grin: Only the lower teeth are revealed, and the teeth are partially or widely open. Absent in bonobos? 3.

See video. groin pocket See transport in groin pocket or hold object in groin pocket. groom Goodall 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 reclining posture. Grooming functions include appeasement, reassurance, reconciliation, and hygiene. See video. groom-branch-clasp Groom mutually like groom-hand-clasp, but non-grooming hand is not clasped, but held overhead on branch. Seen at Gombe and Mahale. Whiten et al.s Branch-clasp. Also recorded for bonobos

of Lomako. See video. groom briefly Groom another individual briefly in greeting or appeasement. See video. groom by hand Groom another with one or both hands. Seem to groom important companions such as relatives and allies with two hands, but others with one hand and often leisurely. See video. See also videos groom with mouth and smack lip. groom carcass 87 groom carcass Groom carcass or even skin of animal such as colobus monkey or leopard. Function unknown, but shown by most adult members of M group. See also care alloparentally for another species. 7. See video. groom, dyadic Two chimpanzees, either one or both grooming. Groom unilaterally, groom mutually, or groom reciprocally. See video. groom ground See groom object or substrate. groom-hand-clasp McGrew and Tutins Grooming-hand-clasp. Kind

of mutual grooming at Mahale, Kibale, Lop, Budongo, Ta and Semliki, but not at Gombe or Bossou. In captivity, done by chimpanzees at Yerkes Primate Research Center. Seen in bonobos of Wamba, Lomako and Lui Kotale. 5. See video. See also video groom-hand-clasp unilaterally. groom-hand-clasp unilaterally One grooming partner assumes typical hand clasp posture, but does not groom companion; instead rests or grooms ground, although being groomed. 9. See video. groom leaf Goodall 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. At Mahale, chimpanzee puts small louse in folded leaf and kills it. Louse egg is also crushed during leaf-grooming. Customary at Gombe, Mahale, Kibale and present at Budongo, but not seen at Ta and Bossou, nor in bonobos of Wamba. See also inspect leaf and squash leaf. 5. See video. groom mutually Goodall Mutual grooming. Two chimpanzees groom each other simultaneously. Common in chimpanzees, but occurs in Wamba bonobos only during role changes of social grooming. Also rare at Lomako. Humans may not groom mutually. 3. See video. groom object or substrate Groom soil, leaves on ground, tree trunk, or moss on rock, usually with one hand. Solitary or social grooming, even during grooming-hand-clasp. Seen G 88 grunt, aha in only

two young adult males of Mahale in 1992, but done also by three adult or adolescent females in 1995. See video. groom, polyadic Simultaneously, three or more individuals groom in tandem or mutually. See Nakamura. See video. groom reciprocally Goodall Reciprocal grooming. Two chimpanzees alternately groom each other. Most common type of social grooming among adults. Common in chimpanzees and bonobos. See video. G groom self Goodall Self groom, Plooijs Self-groom, van Hooffs Autogroom. Kanos Self-groom for bonobos. See video. groom unilaterally Goodall Unilateral grooming. Groom another without reciprocation. Kanos Unilateral grooming for bonobos. See video. groom with mouth Use lips and teeth to remove items when grooming, especially from face. Also in bonobos. See video. groom wound Groom wound of another individual. Cf. inspect wound. 3. See

video. See also video inspect wound. grope Put arm into tree hole and feel for something to eat. Cf probe, probe with finger. 3. See video. ground bed or cushion See make ground bed/cushion. group Make any association of individuals, temporary or long-term. grunt Various utterances including grunt, aha, grunt in bed, grunt, extended, grunt, food and pant-grunt. grunt, aha Food grunts uttered loudly when walking hastily in group to known food patch. Probably absent in bonobos of Wamba. 5. See video. grunt, extended 89 grunt, extended Goodall: A long drawn out sound usually composed of two syllables- ehmmmmm- heard during rest sessions, significance not clear. Mahale chimpanzees often discovered because of this sound uttered in response to passers-by or appearance of human beings. Category 35. See video.

grunt, food Uttered during or just before eating, especially during first minutes. Cf. aha-grunt. Bonobo give similar grunt. See video. grunt in bed Goodall 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. Same as Boeschs Night call? Not heard in bonobos of Wamba. 5. G H H habituated No overt response to stimuli, after many repeated presentations that earlier elicited some reaction. Chimpanzees of M group now act normally in human presence although they fled before habituation succeeded. See Fig. 18. Fig. 18 Habituated: Chimpanzees of M group are followed by researchers without any apparent influence on their behavior hair erection See bristle. 90 hammer nut with

stone 91 hammer nut with stone Use stone to hammer nut on stone anvil or wood anvil: Customary at Bossou and Ta; Ebo; also at Nimba. At Bossou, only stone anvil used. At Ebo, stone hammer used in tree. Absent at Mahale. See video. hammer nut with wood Use wooden club to hammer nut on wooden or stone anvil. Customary only at Ta. Absent at Mahale. See Fig. 19. H Fig. 19 Hammer nut with wood: Drawn from photo of chimpanzees of Ta hand clap See clap hand. hand-clasp See groom-hand-clasp. hand-rubbing See rub hand/feet. hand support See transport with hand support. 92 hang tripedal handedness Preference for use of right or left hand. Handedness for tool use was found on individual basis, but not found population-level among

wild chimpanzees. handicap self Older individual inhibits rough acts and shows gentle restraint in social play, especially with youngsters, e.g. lie supine, lie prone, somersault, flee, fall over backward, etc. See video. handwrestle See wrestle with fingers. H hang Suspended freely by one or both hands from tree branch. Neither foot touches substrate. Kanos Suspend in bonobos. See video. hang and spin Hang from horizontal branch with one hand, and revolve repeatedly. Small infants and show as solo play. Sometimes, hanging immature plays with, or solicits play from, individual resting on ground. See video. hang and stamp Hang by one or both hands from large bough and stamp on ground in solo or social play. May repeat bouncing up and down like trampoline. Infants, juveniles and adolescents enjoy

this. See video. hang in sloth position Goodall Sloth position: Hanging underneath a branch using all hands and feet or a combination of any three. Usually part of locomotor play in infants. At Mahale done as solo play by immature chimpanzees. Rarely locomote in this position. Bonobos not only hang, but also travel in this posture. See video. hang object See suspend. hang-stand Hang from branch while one or both feet touch substrate below. Temporary posture changes to another posture or locomotion, or sometimes brief resting posture. Also in bonobos. 2. See video. hang tripedal Hang while gripping branches with feet and one hand. Head and trunk more or less erect. Other hand used to eat fruits or young leaves at ends of high hang upside-down by feet

93 branches, or extended to the playmate below in social play. Also in bonobos. See video. hang upside-down by feet Hang by feet from branch. Rare in infant and juvenile social and solo play and play invitation. Kanos Upside-down suspension for bonobos. See video. hang upside-down by hands Grip branch with hands, flex legs and hang upside down. Infants and juveniles in social and solo play and play invitation. 3. See video. See also video play with object. hang with legs pitterpat Hang from horizontal branch and shake legs up and down, e.g., in response to play partner on ground who tries to grasp leg of suspended individual, or to solicit play from partner on ground. See video. hang-wrestle Two arboreal chimpanzees hang by one hand, and each

tries to grab, push, or pull other with other hand. Making playmate fall seems to be aim of most interactions. 3. See video. harass Subordinate chimpanzee threatens dominant one. Adangs Tease. At Mahale older juvenile and adolescent males pester and threaten adult females by shake branch, slap, hit, throw branch, charge, etc. until females finally pant-grunt to them. Cf. tease. Bonobo adolescent males at Wamba pester adult males with threatening gestures, such as bipedal swagger or branch-dragging, and so dominate him. Termed as adolescent harassment by Kano. See video. head down hip up See stand with head down, bottom up. head tip See tip head. headstand Stand on ones head to solicit play and in social play. Done by mature and immature individuals. 7. See video. See also

video circle quadrupedal. heel kick See kick heel. heel-tap See tap heel. H 94 hit and run herd Complex of male behavior to control travel of cycling female, so that she follows him. Includes waiting such as sit, glance, wait for companion, reverse, and look back, threatening such as stare fixedly, shake branch, club, and scratch dry leaves, and aggression such as hit and kick. Alpha male pulled female during herding when she did not follow him. Thus, herding maintains consortship and possessiveness. Cf. consort and lead. Bonobos show similar behavior but without aggressive components. 3. See video. H herd in coalition Watts Coalitionary mate guarding: pairs or trios of top-ranking males engaged in cooperative aggression to prevent estrous females from mating with other males, but tolerated each

others mating activities. May be demography-dependent male mating strategy in unit group that has unusually many adult and adolescent males. Not seen at Mahale except for few hours. 5. hesitate Uncertain to do something, for example, repeatedly advance to and retreat from targeted individual. Stimuli include aggressive posture or facial expression of targeted individual, presence of others nearby, or close proximity of observers or dangerous animal such as warthog. 3. See video. hiccup Involuntary contraction of diaphragm produces distinctive sound. Sounds like humans. See video. hide Goodall: 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 and Ta. Unhabituated bonobos do same thing. Cf. detour. 2. hit Bring down an

arm from above and strike partner, animal or insect, etc. with fist or knuckles. van Hooffs and Goodalls Hit includes striking with palm, here called slap. Hit either in play or in attack. Kanos Hit and Pikas Punch in bonobos. See video. hit and run Hit or touch potential playmate and flee to solicit play. Category 18. See video. hit bush bipedal 95 hit bush bipedal Stand bipedal and hit bush with one or both arms, in charging display, threat or soliciting play. See video. hit ground with fist Hit ground with fist. Middle-aged mother, Juno, standing on all fours, hit ground with fist, watching her infant nearby. Infant did not respond and Juno grinned. Middle-aged female, Opal, hit ground and pant-grunted when alpha male approached. Juvenile male,

Xmas, hit ground with fist as solo play. Adolescent male, Cadmus, did so with fists alternately few meters from large grooming clique. Function probably attention-getting. See video. hit toward Threatening gesture. Goodall: When an animal is threatening it will make a hitting movement with the back of its hand. This gesture is usually accompanied by a soft bark. It may be directed toward chimps, other primates, snakes, large insects, birds, and so forth. Cf. raise arm quickly and shake arm. 5. hit with long branch See club. hold body part in mouth Hold by mouth body part such as fingers, hand, toes, foot, nape, limb or back of companion, usually in play. 5. See video. hold down See press down. hold finger in mouth See hold body part

in mouth. hold foot in mouth See hold body part in mouth. hold genitals Goodall Hold genitals. Adult grabs or touches scrotum or penis of another as reassurance. Often occurs when adult male mounted by another male; mounted reaches back between his thighs and holds scrotum of mounter. When adult female pant-grunts to adult male, she may hold his scrotum. Absent in bonobos of Wamba. 5. hold hand See grasp hand. H 9 hoot hold hand in mouth See hold body part in mouth. hold head or face Support anothers face or head with hand while grooming it with other hand or with lips and teeth. Seen in bonobos. 2. See video. hold object in groin pocket Keep object in fold of thigh and groin. See also transport

in groin pocket. 5. See video. H hold object in mouth Keep object between jaws. Small infant often keeps twig or leaf in mouth and nips off pieces in solo play. Small stone also held. See also solicit play with object in mouth and transport in mouth. See video. hold object in neck pocket Keep object in fold between lower jaw and flexed upper shoulder. 5. See video. hold object on head Keep object on head. Adult female, Opal, held her dead infant on head. See transport on head or nape. 3. See video. hold toe in mouth See hold body part in mouth. hold up chin or head See raise others chin. hoo Goodall: 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 etc. Mahale infants hoo when begging for food, when wanting to return to mother after separation, and when seeking to mate with estrous female. Mothers also hoo when older infant does not return to her, often while playing overhead when she wants to leave. Hoo expresses frustration. Cf. whimper. Absent in bonobos? 5. See video. hook branch Sugiyama Branch-hauling, Whiten et al.s Branch-hook: Use branch to hook another branch. Present only at Bossou. Absent at Mahale. See Fig. 20. hoot Goodall: Roar pant hoot. Continuous low-pitched calls given only by highly aroused individuals, rarely by females. Always accompanied by a charging display. 5. hoot

face 97 H Fig. 20 Hook branch: Chimpanzees of Bossou hooked branch overhead with stick hoot face Goodall Hoot face: pouted trumpeted lips that go with pant-hooting. The mouth may open completely or partially. hop bipedal on spot Nishida Jump bipedal. Juvenile male in courtship jumps in place in front of estrous female, to elicit her to present, or in front of adult female to harass her. Some juveniles jump up and down while pant-grunting vigorously in front of dominant male. Cf. leap bipedal and hop quadrupedal on spot. See video. hop quadrupedal on spot Hop repeatedly on four limbs on ground or on mothers back as solo play. Infant may hop while pant-grunting. Courtship by juvenile male. Also hop on branch, vegetation, bed, etc. as trampolining. Cf.

leap quadrupedal. 5. See video. hug Caretaker embraces infant with arms. Used for consolation or intention movement of mounting. Kanos Protective embrace in bonobos. Cf. rush to embrace. See video. hunch Goodall: Hunching shoulders in contexts of aggression, courtship and greeting. Nearly always accompanied by hair erection. Plooijs Hunch. 3. hunch and sit Goodall Sitting hunch: while sitting, raises shoulders, arms held out from the body either to the side or in front. At Mahale, sitting hunch 98 huu courtship often accompanied by stamping ground or shaking branch. Bonobo males Sitting erect may be homologous. 3. See video. hunch bipedal Goodall: Chimp stands on ones feet, shoulders hunched up, arms held slightly out and away from the body. If he sways from foot to foot, known as bipedal

swagger. Also in Kanka Sili. Cf. swagger bipedal. Absent in bonobos. See video. H hunch over Goodall: Similar to bipedal hunch, but arms held forward over the back of another chimp. de Waals Hunch over and Kanos Bluff over for bonobos. See video. hunch quadrupedal Goodall: Stand, walk or run on all fours with rounded back and head pulled in between shoulders. Seen in adult males at Mahale. Also in bonobos. See video. hunt Behavioral complex of stalk, pursue, capture, and kill large birds or small to medium-sized mammals. Include monitor, climb, stalk, run, chase, leap on, grab, flail, drag, drag to kill, throw, bite, pull, etc. Known throughout chimpanzee range. Bonobos hunt less often than chimpanzees. See also eat meat. See video. hunt with tool Thrust stick

to expel or to skewer prey from tree hole or rocky cavity at Mahale and Fongoli. Prey species are squirrel, hyrax, and bushbaby. See also expel. See video. hurl self Throw oneself at another individual. Play pattern employed in tree. 8. See video. huu Goodall:.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. Sound often helps researcher to find otherwise silent, hidden chimpanzee in bush. Absent in bonobos? 5. I ignore Adult female does not respond to harassment by immature male, so avoids escalation of harassment. Mother does not approach weanling despite its request to suckle, ride, etc. Adult male

does not respond to others pant-grunt. Goodalls Ignore includes both ignore and snub. Kanos Ignore for bonobos. Cf. snub. See videos. I imaginary play See play, imaginary. imitate Goodall Imitation When one chimp, after watching the behavior of another, then does the same. See video. immature Age-class consisting of infancy, juvenility and adolescence. immerse See dunk. immigrate Adolescent female leaves natal group to enter neighboring group for reproduction. Immigrant usually welcomed by males, but threatened by adult and adolescent females of new group. Juveniles inspect immigrant. See transfer. See video. immobilize Two or more attackers restrain single mature chimpanzee, using hold. Tactic used in lethal coalitionary attack on victim. Behavior includes hold, pull, bite. Recorded or suspected to occur at Gombe, Mahale, and Kanyawara. Cf. press down. 5.

99 100 infant incest Copulate with close relatives. Mother and infant son copulate during weaning period at Mahale. Mating between mother and adult son once observed. Mother and adolescent son or brother and sister mating not seen at Mahale, although juvenile male once tried to mate with mother but was refused. At Gombe, copulation between mother and mature son, and brother and sister rarely seen and alpha male produced offspring with mother. See avoid incest. 5. See video. index-hit See squash ectoparasite on arm. infant Youngster from birth to weaning. 04 years old. See Fig. 21. I Fig. 21 Infant: Three-year-old female infanticide 101 infanticide See kill infant and eat infant. infantile bark See bark, infantile. insect-pound See pound insect. inspect Investigate object by peer, gaze, sniff, touch,

climb, etc. Cf. monitor. See video. inspect fruit Travel through concentrated patch of fruit trees, apparently to check quality of crop, even long before fruit ripens. Includes walk, watch, touch fruit, sniff, and bite. 5. See video. inspect genitals Goodall Inspect:.male or female touches the vaginal opening of a female and sniffs finger, or sniffs with nose directly. Both hands used to pull apart the lips of vagina. Plooijs INS. Nishidas Genital inspection. Recoded from Ta. Adult male bonobos do not inspect female genitalia, but juveniles do. 3. See video. See also video in clip leaf by mouth. inspect leaf Whiten et al. Leaf-inspect: Inspect ectoparasite on leaf. Component of leaf-grooming. During social and self grooming, take leaf, bring to lips, fold leaf with finger and thumb, open

leaf and often inspect something on leaf. Twice confirmed to be louse or its egg. Customary at Budongo and Mahale and present at Gombe. Pattern at Budongo may be cultural variant. inspect self Inspect own body-part, e.g. genitals. May be combined with groom self. 3. See video. inspect wound Show keen interest in own or others wound. May approach another to peer, touch, lick, mouth, or groom wound. Category 3. See video. interact with human After habituation, some chimpanzees of Mahale began to communicate with humans by extend hand, touch, pull, slap, push, throw stone or rock, throw branch, club ground, etc. Some agonistic and others playful. Cf. play with another animal. I 102 intervene interfere Prevent another from engaging in self-maintenance or social behavior. Female weanling, Aqua,

interfered with consort grooming her mother. Interference in dominance interactions is intervene. Bonobos interfere in sexual and grooming interactions. See video. interfere fishing Infant often reaches hand to hand or fishing probe of mother or older siblings who is fishing for ants, trying to snatch probe or ants. This act much interferes with performers activity. Mother often allows infant to take probe. 7. See video. I interfere in copulation Weaning infant, juvenile or adolescent male, often offspring of mating female, runs to push itself between mating pair, waves arms, or touches or pushes at copulating male, while emitting whimper-scream or loud scream. Juvenile male may interfere with elder brothers mating. Dominant adult male may interfere in others mating by bark, threat, or direct charge. Adult female may squeeze

herself between mating pair. Dominant adult male bonobos interfere in mating of adult females, which Kano called Sexual interference. See video. interfere play Third party prevents ongoing social play by rushing at player, sometimes stopping play. Mother may prohibit rough alloparent from continuing play with her infant. Intruder may solicit play in various ways, take over play partner and engage in social play. 3. See video. See also video kick. inter-group behavior Behavior related to presence or absence of neighboring unit groups. Generally, antagonistic. Groups usually avoid one another, but larger party of unit group occasionally intrudes territory of another group and kills some of them, in particular infants. However, young females in estrus are allowed to enter neighboring group. See immigrate, intrude, kill adult male, kill infant,

patrol, respond to neighboring unit group and transfer. intervene Exert influence on agonistic interaction such as quarrel between two or more others by approach, threat or aggression. Goodalls Interference: Intervention in which a third individual prevents or tries to prevent an interaction between two others. Intervener may be partial or impartial. Also in bonobos. Includes interfere and support. See video. intervene to separate 103 intervene to separate Alpha male charges at rival male and third male who are grooming each other or sitting together, apparently to prevent formation or development of coalition between them. de Waals Separating intervention. See also Nishida and Hosaka. Absent in bonobos, but dominant individual thrust torso, or arm between grooming pair to take over grooming. See video. intimidate See threaten. intrude Wrangham Deep

incursion. Patrol crosses group boundary and invades territory of neighboring group to attack its members or to explore for resources such as food or mates. Recorded at Gombe, Mahale, Kanyawara, Ngogo, and Ta. 3. investigation probe See probe. investigatory probe See probe. invitation slap See slap in invitation. invitation stamp See stamp in invitation. invite See solicit. I J J join play Small infant rushes into vigorous play between seniors and interferes with them by approach, touch, slap, hit, kick, wedge, hug, mouth, etc. Infant rarely allowed to join. Adult, mother, adolescent or juvenile also joins play group of immatures and often succeeds in playing with one or two of them. See video. jump See leap and hop. juvenile Weaned youngster follows mother in daily ranging most of

time. 58 years old. See Fig. 22. Fig. 22 Juvenile: Eight-year-old male 104 K Kabogo Study area of Pan troglodytes schweinfurthii in western Tanzania. Research done from 19 1 to 19 3 by K. Imanishi and colleagues. Kahuzi Study area of Pan troglodytes schweinfurthii in Kahuzi-Biega National Park, Democratic Republic of Congo. Research organized by J. Yamagiwa from 1991 to present. Kalinzu Study area of Pan troglodytes schweinfurthii in Kalinzu Forest Reserve, Uganda. Research organized by C. Hashimoto and T. Furuichi, continuing from 1995 to present. Kanka Sili Study area of Pan troglodytes verus near Kindia in Guinea. Pioneer studies done by A. Kortlandt in the early 19 0s and his colleagues in 19 819 9. Kanyawara Study area of Pan troglodytes schweinfurthii in Kibale National Park, Uganda.

Research organized by R. Wrangham, continuing from 1987 to present. Kasakati Study area of Pan troglodytes schweinfurthii in western Tanzania. Research done from 19 3 to 19 7 by J. Itani, K. Izawa, M. Kawabe, and A. Suzuki. Kasoje Northwestern part of Mahale Mountains National Park, with headquarters of research and ecotourism. 105 K 10 kidnap keep water in mouth Drink from stream and sometimes retain water in mouth for more than 10 s before swallowing. Function unknown. See video. kick Goodall: Make contact with an objective with one or both feet. Kicking is a forward, sideways or backward movement, different from a stamp which is always downward. In aggression and play. See also drum. Cf. stamp. Many types of kicking described for bonobos by Kano. See video.

kick back Goodall: Walking individual kicks backwards towards a following youngster Seen in bonobos. See video. kick backward quadrupedal Kick backward to buttress, tree trunk, wall, etc. while standing on four limbs. Goodalls Drum/Kick. See video. K kick bipedal Kick buttress or tree trunk with foot while standing bipedal. See also drum. Absent in bonobos? 5. See video. kick buttress Kick buttress or alternative such as tree trunk and metal wall. Done mainly by adult males. Demonstrate own geographic position. See also drum. 5. See video. kick heel Goodall Heel kicking: resting individual raises one leg and thumps its heel down on a youngster who is playing around him or her. Also at Mahale. Goodalls other type is 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 infants rump. Absent in bonobos. 5. See video. kick other Kick another in fight and play. Seen in bonobos. Pikas Kick. See video. kick up Lie supine and kick into air object, such as carcass. If carcass, may be conflict behavior. See video. kidnap Nishida Detach: Take infant from mother and keep it under alloparents control for as long as alloparent wishes. Hard for infant to return kill 107 to mother if taken far away from her. Goodalls Kidnap: Borrow by consent. When an infant is carried away from the mother by a sibling or by another chimp. See also care alloparentally. Seen in bonobos. See video. kill Fatal attack to another. 3. kill

adolescent male At Gombe, two cases of between-group fatal attack of adolescent males seen or suspected. kill adult female Adult females killed within group at Ngogo and between groups at Gombe. Adult female attacked between groups but rescued by researchers at Mahale. 5. kill adult male At Mahale, ex-alpha male probably killed within group and several adult males between groups. One case of within-group attack on young adult male, considered to be ostracism. Killing adult male within group recorded or suspected at Gombe, Budongo and Ngogo and killing of adult males between groups at Gombe, Kanyawara, Ngogo and Kalinzu. 5. See Fig. 23. Fig. 23 Kill adult male: Ex-alpha male, Ntologi, suspected to have been killed by group members, given so many wounds on body K 108 kill

time kill another species Kill another species for eating or teasing by flail, knock down, throw, drag to kill, hit, bite, etc. See video. kill infant Infanticide. Infants killed at Mahale, with most eaten by killers and others: between-group 3 times and within group 5 times. Betweengroup infanticide recorded from Budongo, Gombe, and Ngogo. Within-group infanticide recorded at Gombe and Kanyawara, and suspected at Ta. Most infanticide was carried out by adult males, but occasionally by adult females. No infanticide in bonobos of Wamba. See also eat conspecific. See Fig. 24. K Fig. 24 Kill infant: Alpha male, Ntologi, killed and ate infant kill juvenile Between-group fatal attack on juvenile male reported from Ngogo and suspected at Gombe. Absent at Mahale. kill time Behavior when waiting for long

time, e.g. when close companion sleeps for long bout, sits, grooms, fishes for ants or termites, etc. Includes yawn, flip lip, drop bark bits, scratch self, groom object or substrate, etc. 3. See video. kiss 109 kiss Goodall divides Kiss into two forms: pout-kiss and open-mouth kiss. Both types rarely seen in bonobos, but absent in adult bonobos at Wamba. See video. kiss with open mouth Plooij OMK. Goodall 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 others upper lip and nose, the other has his mouth over the lower lip and chin of the first Open-mouth kiss often occurs in reunion, reconciliation, or social

excitement, especially, among adult males, accompanied by panting. Kanos Open mouth kiss directed only to infants and juveniles by older bonobos, and not between adults. 3. See video. kiss with pout face Goodall Pout-kiss: 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. Mother kisses infant in head or face for self-reassurance after agitated interaction with other. Even colobus carcass kissed. Absent in bonobos. See video. kiss with tongue Tongues pressed each other during kissing. Juvenile female and male, Imani and Oscar, showed this French kiss for 7 s as reconciliation after quarrel. 9. See video. knock with both arms Hold, lift, and knock object such as colobus

monkey against substrate with both arms. Kills prey and simultaneously intimidates followers of meat-holder. Also seen at Ta. 8. See video. knock with one arm Hold, lift, and knock object against substrate with one arm. Adolescent male, Cadmus, knocked colobus carcass with one hand, holding branch with other. Kills prey and simultaneously intimidates followers of meat-holder. Also seen at Ta. 8. See video. See also video knock down with both arms. knuckle-knock See rap. knuckle walk See walk quadrupedal on knuckles. K L laugh See play-pant. lead Mature male or female, and especially adolescent male, leads sexual partner into undergrowth or higher in tree to avoid interference by more dominant males. See also consort. Cf. herd. Seen in bonobos. See video. L leaf-clip See clip leaf. leaf cushion

See make leaf cushion. leaf-dab See dab leaf. leaf-fold See fold leaf. leaf-groom See groom leaf. leaf-inspect See inspect leaf. leaf-midrib spoon Juvenile male, Xmas, and infant female, Liz, removed leaf-blade by clipping leaf and used midrib for licking water. Rare solo play. Cf. leaf-spoon. See video. leaf-mop See mop ants with leaves. leaf-napkin See wipe with detached object. 110 leaf-pile pull 111 leaf-pile pull See pull leaf-pile. leaf-pile push See push leaf-pile. leaf-sponge Remove leaves from tree or shrub, dip into water, then put into mouth, and compress. Result is kind of absorbant sponge, which is soaked into tree-hole or stream, then sucked to drink water. Whiten et al. s Leaf-sponge. Chimpanzees of Tongo used moss for leaf sponges. Customary at Gombe, Mahale, Kibale, Budongo, Bossou, Ta,

and Goualougo. Young bonobos of Lomako also use moss sponges. 7. See video. leaf-spoon Put leaf into stream and suck, pour or lick water from it. Cf. leaf-midrib spoon. See video. leaf-squash See squash leaf. leaf strip See strip lea. leaf-swallow See swallow leaf. lean Incline torso to rest against tree trunk, bough, rock, or another. Kanos Reclining sit. See video. lean forward Flex trunk to expose back for grooming. Seen in bonobos. leap Jump propelled by feet. Plooijs JUM. Kanos Leap and Pikas Jump in bonobos. leap between trees Leap horizontally from tree to tree. Done by youngsters rather than adults, during play and when many food trees grow close together. See video. leap between trees with object Leap horizontally from tree to tree, while carrying object

on shoulders. See video. L 112 leave leap bipedal Adult male stands upright, swings body forward and back, to gain momentum, then jumps from rock to rock in upright posture when crossing stream on stepping stones. Absent in bonobos of Wamba. leap bipedal with squared shoulders Adult male with arms half-raised jumps bipedally up and down repeatedly facing another male doing same. Threatening gesture in adult males and females. 7. See video. leap down Goodall Vertical leap: Leap vertically from one structure to another. Leap from tree down to vegetation or ground during play, at start of departure, or when chased. Infant leaps down from mothers back to ground. Cf. fall and drop self. Kanos Vertical leap for bonobos. See video. leap down and wait As mother travels,

older infant leaps down from mothers back, runs ahead, climbs shrub or fallen log, waits for her arrival, then leaps on her back, and resumes ride. Infant play, repeated again and again. 7. L leap into lap Goodall: An infant jumps into the lap of a sitting chimp, during greeting. Seen at Mahale, but not in greeting. leap on Spring onto another in play and aggression. Infant may leap from mothers back to older playmates back in traveling. Kanos Jump on similar for bonobos, but jumper likely to mount. See video. leap quadrupedal Usual way of crossing stream via stepping stones. Before leaping, chimpanzees swing body back and forth to gain momentum in quadrupedal posture. Cf. hop quadrupedal on spot. Bonobos commonly leap on all fours arboreally and

on ground. See video. leap up Jump upward suddenly from sitting to grasp overhead branch or hanging playmate. Done by infant without anything overhead as solo play. Cf. leap up in surprise. 7. See video. leap up in surprise While standing bipedal, may leap up when surprised. Category 38. See video. leave Travel away from another with no parting signals. Plooijs LVE. Cf. depart. See video. leave and climb 113 leave and climb Infant leaves mothers back and climbs tree. 3. See video. leave to protest Weanling protests to mother by leaving and keeping distance from her, usually with whimper or whimper-scream. See also monitor mother. Cf. fall over backward. See video. leg cradle See cradle with leg. leopard See respond to leopard. lever open Whiten et al.