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chimpanzee-behaviour-ethogram

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Lever open: Use stick to enlarge entrance. Customary at Gombe, Ta, Lop, and Goualougo. Absent at Mahale. lick Plooij LIK. Repeatedly stroke with tongue. See video. lick lips Lick away items on lips. See video. lick rock Lick rocks at few sites along shore of Lake Tanganyika or in riverbed of large rivers. More often in dry than wet season. May continue an hour. At Mahale, seen in chimpanzees of B, K, M and L groups. Nutritional significance unknown. Rocks licked not salty to human taste. Not seen at Gombe. Category. See video. lick snot See eat nasal mucus. lick wood Licks de-barked surface of dead trees such as Pycnanthus angolensis, Ficus capensis, Ficus vallis-choudae, Garcinia huillensis, etc., with extended upward laps of tongue. Surface exudes some sugar

alcohol, which might be nutritionally meaningful. Not seen at Gombe. Category. See video. lick wound Lick wound apparently to clean it, directly with tongue or touch wound and then lick fingers. Sometimes done to others, often by infants or juveniles. Reported from Gombe, Mahale and Ta. Seen in bonobos, who also lick menstrual blood. See video. L 114 lie with back to another lie Recline on ground or other horizontal substrate such as stout branch. Plooijs LIE. Kanos Lie for bonobos. See video. lie and hug Nishida: Caretaker lies supine on ground while embracing infant to chest. Seen in bonobos. See video. lie and watch Adult or adolescent lie on ground and watch playing juveniles or infants to solicit play. 7. See video. lie lateral Lie on side.

Common resting or sleeping posture on ground. See video. lie on other Lie on others belly, etc. in play. Larger individual may lie supine on smaller play partner, immobilizing it during wrestling. Cf. step on in which smaller playmate rides on older reclining companion. Cf. press down in which larger individual stands quadrupedal, sits, or lies on stomach. 7. See video. L lie prone Lie on belly with legs and arms flexed. Assumed occasionally when waiting for play partner to come after solicitation of play. Also posture assumed by estrous or pregnant females near term. See video. lie-sit Typical sitting or reclining posture of estrous female, apparently forced by very swollen sexual skin. See videos. lie supine Lie on back with legs not spread. Most common sleeping posture

in bed or on ground. Also, as solicitation of play, in which playmate may step on him/her. See video. lie supine and shake arms and legs Lie supine and shake arms and legs quickly and irregularly. Solo play of adolescent males, Cadmus and Primus, juvenile male, Xmas, and infant female, Xantip. Occasionally used as solicitation of play, as juvenile male, Michio, did so on back of adult male. See video. lie supine with legs apart Lie on back with legs spread. Often occurs when male solicits another to groom scrotum. See video. lie with back to another Pattern of soliciting grooming of back. See also solicit grooming. Seen in bonobos. See video. lie with legs crossed 115 lie with legs crossed Lie supine with raised legs crossed. Resting

posture. 3. See video. lift Raise up heavy object by hands. Seen in bonobos. See video. lift and drop Raise up heavy log or thick woody vine by hand, then drop it on ground to make loud, thumping sound. Display by young adolescent males, Primus and Orion, and adult male, Pim, as courtship, mild threat, or solicitation of play. See video. lift rock Adult male lifts rock with hands, then usually throws into water to make noisy splash, apparently heightening effect of charging display. See throw splash. No rocks in habitat of Wamba bonobos. See video. Lilungu Study area of Pan paniscus in Equateur Province of DRC. Research done by Sabater Pi and colleagues from 1988 to 1990. limp Walk lame because of wounded limb. See video. lion

See respond to lion. lip flip See flip lip. lip-smack See smack lip. listen Orient head to source of sound or calls, while stopping on-going activities, such as groomimg or eating. Cf. search for conspecific. See video. locomote Includes walk, crutch, retreat, limp, stamp trot, run, gallop, somersault, pirouette, climb, leap, brachiate, sway and move, swing and grasp, fall, descend, slide down, etc. locomotion See locomote. locomotion play See play. L 11 look between thighs locomotor behavior See locomote. locomotor play See play. locomotory aid See aid in locomotion. Lomako Study area of Pan paniscus in Equateur Province of DRC. Research initiated by A and N Badrian from 1974 and taken over by many researchers after 1980s. lone play See play solo. longevity Longest lifespan. About 50 years

at Mahale. L look around Look side to side or back and forth, often accompanied by straightening back and tilting head, in order to monitor surroundings. Cf. scan and search for conspecific. See video. look at water Sit or stand quadrupedally and look at waters surface, either to watch object in water, or to view reflection of self-image. Cf. nod quadrupedal. See also play with water. 8. See video. See also video drink from stream. look back Groomee turns to look at groomer who has stopped grooming, and groomer immediately resumes grooming. Thus, turning round is request to resume grooming. Walking chimpanzee looks back to see if companion is following. e.g. when adult male leads sexual partner during consortship, or other chimpanzee is going to charge. Also seen

in bonobos of Wamba. See video. look between thighs Quadrupedal individual lowers head and looks at another through spread thighs to solicit play. Mother uses to capture infant as it continues to play when mother seeks to depart. Two-year-old female, Athena, looked at following observer between thighs at rear of group travel. Estrous young adult female presented to adolescent male, but he did not copulate. Then female looked at him through thighs in order to monitor his further response. 8. See video. look up 117 look up Sitting or standing individual raises up face to monitor arboreal stimuli, e.g. fruit, colobus monkey, or others. See video. Lop Study area of Pan troglodytes troglodytes in Lop Faunal Reserve of Gabon. Research organized by C.E.G. Tutin and M. Fernandez from

1984 to 1993. loser support See support subordinate. lost call See whimper-scream. lost child See search for conspecific. louse Ectoparasites of two genera, Pediculus and Pthiris, parasitizing chimpanzees. See remove lice. See Fig. 25. L Fig. 25 Louse: Louse of Pediculus schaeffi recovered from chimpanzee of M group 118 Luo Scientific Reserve lower arm Groomee lowers arm, e.g., when grooming-hand-clasp ends, signaling intention to change role or posture. Groomer uses to request groomee to change role in bonobos. 3. lower head Two individuals face each other in social grooming, and one flexes neck to offer top of head for grooming. Also in bonobos. 2. See video. lower head and shoulder Mother lowers head and shoulders to drop off infant forward. See also drop infant. See video. lower leg

Resting or lying groomee lowers raised leg, apparently stimulating groomer to groom another body-part. Seen in bonobos. See video. lower rump Mother lowers rump in order to drop infant. Sometimes, almost stands bipedally but with one hand on ground. See drop infant. 7. See video. L Lui Kotale Study area of Pan paniscus in Salonga National Park, DRC. Research organized by G. Hohmann and B. Fruth from 2000 to present. Lukuru Study area of Pan paniscus in Lukuru Forest, DRC. Research organized by J. Thompson from 1992 to present. lunge See leap on. Luo Scientific Reserve See Wamba. M Mahale Research area of Pan troglodytes schweinfurthii, used as synonym of Kasoje area, and abbreviation of Mahale Mountains and Mahale Mountains National Park. Research organized by T. Nishida from

19 5 to present. make bed Goodall Nest. Make platform for sleep or rest, by bending branches over each other. Bed preferable to nest because major function is for sleeping, not rearing. Includes bend, transport, sit, and stand bipedal. See video. make cushion Bend or break one or few shrubs or other vegetation, then sit or lie on them above ground. Used for daytime rest or nap. Cushion is less complex construction than bed. Weanling may make cushion in order to be suckled. Customary at Mahale. Bonobos of Wamba and Lomako do the same. See video. make day bed Goodall Day nest. Bed made in daytime. Simpler in construction and made more quickly than night bed. Bonobos make day beds. See also make cushion. See video. make ground

bed or cushion Make bed or cushion on ground for daytime rest, by bending over few shrubs or grasses. Mahale chimpanzees make no ground beds for overnight use, but those of Nimba sometimes spend night on ground bed probably because of lack of predators, or as mate-guarding. Also, ground beds seem to be used for overnight sleeping at Kalinzu, with few large carnivores. See video. 119 M 120 marrow-pick make leaf cushion Hirata et al.s Leaf cushion: Use leaves as cushions to sit on wet ground. Whiten et al.s Seat vegetation. Different from simple cushion in that materials for leaf cushion is cut off from plant. Habitual at Ta, present at Bossou, Goualougo and Kibale. Absent at Mahale. See Fig. 2. Fig. 2 Make leaf cushion: Leaves left

by chimpanzee at Bossou M make night bed Goodall Night nest. Plooijs Nestbuilding. Evening bed-making is common characteristic of great apes. Durable sometimes for more than months 2. See video. make sound Produce audible sound by exerting force on external object. Includes clip leaf, drum, rap, slap, splash water, stamp, etc. Cf. vocalize. make tool Modify natural object, so that it serves more effectively. At Mahale, this is limited to a few patterns. For example, make probe for fishing for carpenter ants: peel bark and cut it to appropriate size, use twig by removing leaves and small branch, remove leaf-blade and use mid-rib, etc. Also bend branches or shrubs for bed-making and courtship. Many other tool-making behavioral patterns seen at other sites. See video. male invite See open

thighs. marrow-pick See pick out bone marrow. mass excitement 121 mass excitement Nishida; Moris Booming situation; Reynolds and Reynolds Carnival; Goodalls Social excitement. Simultaneous, noisy display, when many chimpanzees gather at one place, such as new food source. massage shoulder When trouble started and screaming began, adult female, Xtina, massaged shoulder of female, Ruby, with left hand, while both pant-hooting. Kind of reassurance? 9. See video. masturbate Includes fumble penis for males and fumble clitoris for females. 8. mate See copulate. mate guarding See herd. maternal behavior See care maternally. maternal leave See travel alone after childbirth. mature See adult. meat-eating See eat meat. meat-sharing See share food. medicate self Includes lick wound, lie, remove lice, remove sand flea, push peri-anogenital region with finger, and probe nasal passage.

Swallow leaf and chew Vernonia piths may be internal selfmedication. menstrual cycle See swelling of sexual skin. misunderstand Misconstrue others intention, e.g. flinch from accidentally dropped stick. Adolescent female, Ai, tried to escape when adult female, Xtina, approached to groom her. Ai misunderstood that Xtina would attack her. 3. See video. M 122 mount mock bite See mouth. monitor Investigate changes in environment or social interactions. Includes watch, look up, stand bipedal, walk bipedal, sniff, etc. See video. monitor monkeys Chimpanzees in hunting mood sit on ground, scan canopy, and walk little by little, perhaps to look for immature monkeys and to check general movements. Includes walk, run, climb, watch, look up etc. See video. See also video hunt. monitor mother Infant being weaned by rejecting mother runs

screaming from her to few meters away, then repeatedly glances at mother to see if she will allow suckling, or cradling. Includes whimper, scream, whimper-scream, leave to protest, run, look back, glance, and throw temper tantrum. Also in bonobos. 2. See video. M mop ant Collect Camponotus ants with hair on back of hand, usually when many wood-boring ants emerge from nest and move about on surface of tree trunk. Small infant may use palmar side of hand. See video. mop ant with leaves Wipe away Camponotus ants with crumpled leaves. Observed only once for K group female, Chausiku. moralistic aggression See aggress morally. mother-offspring relations Mother and offspring spend most time together throughout infancy and juvenility. Since son remains in natal group throughout life, mother-son relationship lasts

until either of them dies. By contrast, since daughter usually emigrates from natal group and never returns, mother-daughter relationship terminates abruptly around 11 years of daughters age. Moukalaba Study area of Pan troglodytes troglodytes in Moukalaba-Doudou National Park, Gabon. Research organized by J. Yamagiwa from 2003 to present. mount Embrace another with arms from behind. Goodall: Part of the ventral surface of the mounter is in contact with part of the dorsal surface of mount in copulation 123 the other, and the mounter leans forward over the other, usually grasping him/her. She also mentions that the mounted individual may reach back to touch the genital area of the mounter with a hand, or the mounter may raise his/her own foot to the scrotum of the other. Goodall lists

as synonyms: Dorso-ventral embrace, Mounting embrace or Mount-embrace. Perhaps, Kortlandts Sham copulation. Plooijs MOU, de Waals Mount. Males and females mount. Mounting occurs, especially among adult males, during high social excitement such as reunion of rival males, in which each rival male mounts adult male or female, and both participants often scream or pant-scream. Usually bipedal with pelvic thrusts. Appears to function as self-reassurance, reassurance, recruitment of coalition partners, etc. In bonobos, mounting occurs in similar contexts such as appeasement and easing of tension, but posture differs, so that mounted individual usually stands quadrupedal. See video. mount in copulation Goodall Copulation mount: Male places both hands firmly on the females back or sides, during which he may open mouth kiss her. accompanied by intromission and thrusting movements of

the pelvis. See video. mount, misdirected van Hooff Disoriented mounting: Thrusting towards the head, face, shoulder or any other part except the rear. See video. mouth van Hooff Gnaw: Keep month wide open without the retraction of lips and press the teeth to the back, shoulder or other body part without biting. Reassurance often accompanied by panting. Similar pattern used to tickle with mouth pressed to shoulder, chest, belly, etc. Nishidas Mouthing or Mouth contact. Plooijs Mouthing. Goodalls Biting includes mouth. Goodalls Play biting occurs in social play. See also play bite. Cf. bite. Kanos Mock bite for bonobos. See video. mouth and shake Stand quadruped, mouth and shake another continuously, in social play. mouth for begging Beg for food by putting lips to lips or hand of

feeding other. Goodalls Beg mouth-to-mouth. Plooijs Beg with mouth. Kanos Food beg: mouth to hand and mouth to mouth. Cf. extend hand to beg. See video. move Dislodge object such as rock, heavy branch, etc. by push, pull, lift, stamp, etc. Pikas Move in bonobos. See video. M 124 muzzle-rubbing mummify Mother sometimes keeps dead infant for months, so it may be mummified if death occurs in dry season. Also reported from Bossou. See Fig. 27. Fig. 27 Mummify: Adult female, Wakasunga, continued to carry corpse of her infant until it was mostly bare bones M mummy See mummify. mutual grooming See groom mutually. muzzle-rubbing See rub muzzle. N nasal probe See probe nasal passage. Ndakan Study area of Pan troglodytes troglodytes in Dzanga-Sangha region of Central African

Republic. Research done by J.M. Fay in 1988. Ndoki Study area of Pan troglodytes troglodytes in Nouabale-Ndoki National Park, Republic of Congo. Research organized by S. Kuroda from 1989 to 1992. neck pocket See transport in neck pocket and hold object in neck pocket. N neglect See snub. nest See bed and make bed. nest-build See make bed. nest-grunt See grunt in bed. nest play See play in bed. Ngogo Study area of Pan troglodytes schweinfurthii in Kibale National Park, Uganda. Research organized by J.C. Mitani and D.P. Watts, continuing from 1994 to present. Pioneer study done by M.P. Ghiglieri in 197 1978 and in 1981. 125 12 nod with play face Ngotto Forest Study area of Pan troglodytes troglodytes in Central African Republic. Research organized by R.S.

Fouts from 2000 to present. night nest See make night bed. Nimba Study area of Pan troglodytes verus in Nimba Mountains Biosphere Reserve, in Guinea and Cote dIvoire. Research done intermittently from 1991 to present by Y. Sugiyama and colleagues, especially K. Koops and T. Humle. nipple contact See contact with nipple. nipple press See fumble nipple. nod and mouth When seated, holds playmate in lap, or as it stands quadrupedal, repeatedly brings mouth to body of playmate, in nodding motion. Usually elicits play pants from recipient. Adult male hugs another from back and repeatedly puts open mouth to latters back for selfreassurance. See video. N nod to water surface Nod up and down repeatedly while standing quadrupedal and watching reflection of self on waters surface. Few individuals

at Mahale such as Cadmus, Michio, Jiddah and Totzy do so. Cf. look at water. See video. nod to water surface and mouth water Adult female, Totzy, nods to waters surface. She repeatedly takes water into mouth when face down and spits it out when face up. Rare water-contact play. See video. nod with body part in mouth Juvenile male, Xmas, sits and nods repeatedly with hand of infant playmate in mouth. Rare social play pattern. See video. nod with object in mouth Juvenile female, Jiddah, puts pebble from stream into mouth, retains it and nods repeatedly. Idiosyncratic solo play. See video. nod with play face Nod up and down repeatedly, with play face, while quadrupedally standing or approaching playmate. Solicitation of play. 8. See video. nut cracking

127 nut cracking See hammer nut. nut-hammer See hammer nut. nuzzle Goodall Nuzzle: The young infant, until about 3 months old, moves its head from side to side and up and down against the mothers body when searching for the nipple Termed rooting behavior in other mammalian species. Kanos Nuzzle in bonobos. N O object play See play with object. Odzala Study area of Pan troglodytes troglodytes in Odzala National Park, Republic of Congo. Research done by C. Devos and colleagues in 2001. offer arm Dominant individual extends arm for subordinate to mouth, to be reassured, or to be groomed. Absent in bonobos of Wamba. 5. See video. offer back Present back to solicit grooming of it. 2. See video. O Okorobiko Study area of Pan troglodytes troglodytes

at Mt. Okorobiko of Rio Muni, Equatorial Guinea. Research done by C. Jones and J. Sabater Pi from 19 to 19 8. old age See senescence. open eyes Plooij Eyes open. See video. open mouth See play face and yawn. Some individuals open mouth when showing relaxed face. open mouth kiss See kiss with open mouth. open thighs Goodall Male invite: male sits with thighs splayed and penile erection, looking toward a female in estrus. Sometimes his hair is erect 128 orphan 129 Male courtship display. Similar pattern in bonobos. See video. orphan Dependent offspring who has survived mothers death. Usually accompanies adult males, particular adult female, or sibling. See adopt. ostracize Most members of unit group punish or expel individual from group by repeated, concerted attack. Reported

first by Nishida et al. See kill adult male and attack concertedly. 5. See Fig. 28. Fig. 28 Ostracize: Ex-alpha male, Ntologi, spent about 8 months alone after being ostracized O P palmigrade walk See walk quadrupedal on palms. pant Goodall: various calls which are linked by audible inhalations and which may be described as vocalized pant. Non-vocal panting sometimes occurs when chimpanzees are grooming each other, during greeting, etc. Sometimes, when the mouth is closed, only breathing sounds are heard. The best indication of panting is the quick rhythmic movements of the body that accompany it. Bonobos pant in play, but not in other contexts? 3. See video. pant-bark Goodall: Series of bark-like sounds joined by voiced inhalations. See also bark. Status unknown in bonobos. 3. See

video. P pant-grunt Goodall: 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 Hooffs Rapid OhOh. At Mahale, pant-grunting adult females may present to adult males, who may mount her and pelvic thrust. In response to pant-grunting from young adult and adolescent males, alpha male may jump on or attack them without being appeased. See also bob and bow. Pant-grunt absent in bonobo. See video. pant-grunt with bent elbow Pant-grunt while crouched quadrupedally with elbow flexed. Often, part of pant-grunt complex performed by older juvenile and adolescent males at Mahale, although adult females also do so. Cf. bow. Category 35.

See video. 130 pant-hoot 131 pant-hoot Goodall: 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 Higher-ranking males pant-hoot more often than lower-ranking ones. Chimpanzees advertise their presence and numerical strength to rival groups. At Mahale adult males also pant hoot when agonistic confrontations are continuing, in which pant hoots function as vocal threat. Pant-hoots have four stages: introduction, build-up, climax and let-down, but females lack climax and let-down stages. Elements of pant-hoot calls vary between Mahale

and Gombe. Exact bonobo version of pant-hoot absent, but Kanos Waa call, or Okayasus Hoot may be homologous to pant-hoot. Bonobo Waa call emitted upon arrival at feeding site, during feeding and resting, and at sunset after making night beds. See video. See also videos drum, pat and pull rock to roll. pant in copulation Goodall Copulation pant: Rather hoarse panting given by some males during copulation. In its loudest form may be confused with laughing. At Mahale rarely heard and seems idiosyncratic. pant-scream See pant-grunt. panting-laugh See play-pant. paraplay See solicit play and rebuff play. parry van Hooffs 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. Goodalls Startle flinch: When a chimpanzee is startled by a sudden movement nearby 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, Goodalls Startle reaction; P 132 patrol Plooijs Startle. Bonobos of Wamba duck head in similar context. See video. See also video splash water. party Sugiyama Party. Nishida Subgroup. Temporary subgroup of unit group or community, in fission-fusion. Size of parties changes seasonally, dependent on distribution and quantity of major fruit. See unit group. pass Walk past another, without body contact. See walk quadrupedal. Goodalls Pass. Kanos Pass without body contact for bonobos. See video. pass under Go under low horizontal obstacle such as fallen log.

Infant may walk quadrupedal under adult walking or standing, perhaps for amusement. Juvenile female, Imani, once passed through between legs of human. See video. pass with body contact Pass by another and make unnecessary body contact, apparently intentionally. Two young adult males and three juvenile females at Mahale did so to human observer in early 2000s. Significance unknown. Kanos Pass with body contact for bonobos. See video. P pat Plooij Pat. Goodall: 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. It has a calming effect. Also directed to infants head or back. Shown in play by juvenile bonobos of

Wamba. See video. patrol Nishida Scout. Goodall: Party of males moves deliberately and silently in the peripheral part of community range. Elements include sniff, stalk, stare fixedly, attack, and flee. Also from Budongo, Kanyawara, Ngogo and Ta. Bonobo males silently approach adjacent group, but only after hearing latter vocalize. Kanos Scout. 3. See Fig. 29. peel with hand 133 Fig. 29 Patrol: Adult males of M group patrolled and invaded K groups range at Myako Valley. Two adult males lead travel peel with hand Pull tip of outer layer of stalks of grasses, herbs and woody vines by hand while holding stalk with other. Supplementary movement of peel with teeth. Also in bonobos. See video. peel with teeth Pull with teeth tip of outer bark while holding stalk

in hands, thus removing hard outer layer of stalks of grasses, herbs and woody vines to eat pith. Peeled bark used as probe for ant-fishing. Also in bonobos. See video. peep Look through hole or slit. Cf. peer. See video. peer Look intently into another individuals face from few cm. distance, usually in quadrupedal stance. Goodall: when another eats, leaf-grooms, investigates or licks wound. In addition, at Mahale, when trying to remove sand fleas. Plooijs Put face close. Peering usually does not elicit food sharing. Cf. glance, look, stare fixedly, watch, and sniff mouth. Bonobos show same pattern. See video. P 134 pick out nutmeat peer together Up to 11 individuals gather, encircle, and peer at targeted individual who attracts attention. Peered-at has new wound, or is removing

ectoparasites. Recorded also at Bossou. 3. See video. pelvic thrust See thrust. penile adduction See adduct penis. penile erection See erect penis. perforate Chimpanzees of Goualougo reopen termite exit/entry holes on the surface of epigeal nests after unsuccessfully attempting to open the holes by hand. Perforating twig tools are held with a precision grip as the tip is pressed into the surface of the mound. The end of the tool is used to push through the soil replaced by termites to seal the nest. Absent at Mahale. See video. pestle-pound See pound pestle. Petit Loango Study area of Pan troglodytes troglodytes in Petit Loango Faunal Reserve, Gabon. Research done by J. Yamagiwa and colleagues from 1994 to 1998. P pick ear Remove ear wax with finger. See sickness.

3. pick nose Remove nasal mucus with finger and usually eat it. See eat nasal mucus and sickness. See video. See also video eat nasal mucus. pick out bone marrow Marrow-pick: Put stout probe into cracked bone and extract bone marrow with it. Customary only at Ta. Present at Goualougo. Absent at Mahale. pick out nutmeat Chimpanzees of Bossou pick out residual contents of nutshell with probe after cracking open oil palm fruit and eat contents. Present at Goualougo. Absent at Mahale. See Fig. 30. pick out pulp 135 Fig. 30 Pick out nutmeat: Drawn from photo of Bossou chimpanzee pick out pulp Pick out with forefinger pulp from large fruit such as Saba comorensis and eat it. See video. pick up Take object such as food by

hand from substrate such as ground. Cf. lift. See video. See also video search for object. pick up and release While sitting, grasp and lift objects, such as dry leaves, sand, pebbles, etc. and release in air. Solo play in infants. 8. See video. pick up discarded food Nishida and Turner Food retrieval. Pick up food left by others. Infant often picks up food left by mother and ingests it, thus gradually learning what to eat. See video. pick up lice See remove lice. pile dry leaves Spread arms to right and left sides and bring together dry leaves in front of body by adduction of arms. Produces distinctive rustling sound. Solo play by youngsters. 7. See video. pin down See press down. P 13 play face pinch

aggressively Goodall Pinch. Plooijs Pinch. Squeeze skin between thumb and index finger. Idiosyncracy of male chimp at Gombe. Absent at Mahale. Absent in bonobos. pinch clitoris Pinch clitoris of juvenile or infant female as tickling. See video. pirouette Goodall: progresses in a series of tight circles, moving on all fours or on their feet. Type of locomotor play. Seen in older infants, juveniles and young adolescents at Mahale. Juvenile female, Ivana, once pirouetted in order to solicit play from small infant, but usually done as solo play. Probably absent in bonobos of Wamba, perhaps different from Pikas Ice skating. See video. P play Goodall divided play into lone play and social play. She split play into four categories, locomotor play, or van Hooffs Gymnastic, nest play, object play,

self play. She added that, There is a facial expression connected with play, the play face, a type of locomotion that is seen only in the play context, the play walk, and a vocalization, laughing. Goodall listed as elements of each play type: locomotor play: dangle, gambol, swing, pirouette, and somersault. nest play: make bed, leap, and climb. object play: pick up, throw, rub, rotate fruit, drag, drape, flail, and move. self play: grab, dangle, slap, and tickle. Social play included play bite, push head into ventral, chase, circle, drag, wrestle with fingers, hang-wrestle, kick heel, kick, kick back, pinch, play walk, pull, push, step on, rub, slap, spar, tickle, pull object from opposing sides, wrestle, etc. in addition to most patterns used in lone play. See video.

play bite Goodall Play bite: Contact made on the partners body with teeth. Sometimes one of the players gets a firm grip on a limb and holds on for minutes at a time Infant, Teddy, bit mothers supra-orbital region, etc. softly as play. See mouth, as play bite is mouth in play. 2. See video. play face van Hooff Relaxed open-mouth display. Goodall: Only slight retraction of the lip corners. Jaws may be closed or open Occurs during play. Also in bonobos. See video. play face, full 137 play face, full Goodall Full play face: Upper and lower teeth show as the play gets rougher and tickling more vigorous Plooijs Play face full. Also in bonobos. See video. play face, half Plooij, Goodall: Low play face: Only the

lower teeth show. Status unknown in bonobos. 3. See video. play fight See wrestle. play, imaginary Solo play, apparently with imaginary companion or tool. Multiparous adult female lay supine with all limbs raised and play-panted as if playing aeroplane with infant who had been dead for few months. Adolescent male ran round and round tree while play-panting as if chasing, or fleeing from playmate. Adolescent male of Kanyawara carried log for over 4 h, treating it as infant. Rarely observed. See also Gomez and Martin-Andrade. 8. play in bed Goodall Arboreal nest play. Also in bonobos. See video. play invitation See solicit play. play-pant Plooijs Laugh. Goodall Laugh: Somewhat resembles human laughter, heard when chimpanzees are playing socially. As the play becomes rougher, or involves more tickling, there

is more laughter Chimpanzees rarely emit play-pant during solo play. In social play, tickled or chased chimpanzee emits most play-pants. Bonobos also play-pant. See video. See also video hang with legs pitterpat. play, parallel Two or more infants play simultaneously in vicinity, but without interacting with one another. See video. play, rough and tumble Most physical type of play, usually between youngsters. Paquettes Rough play. Wrestling and often locomotor play. Elements include chase, run, climb, leap, grab, tag, wrestle, push, pull, mouth, play bite, drag, stamp, slap, thrust, play-pant, scream, play face, grin, etc. See video. P 138 play with sand play self See play solo. play socially Goodall Social play. Play with conspecific. Usually in twos, but sometimes in 35. See also play. See video. play socially

with object Goodall: Social object play. Social play that includes chasing individual who has object such as fruit, branch, stone, animal skin, etc. and simultaneously pulling object from opposing sides. At Mahale, youngster may also press leafy branch to playmate while wrestling. Ingmansons Tool use in play for bonobos. See video. See also video discard. play solo Goodall Self play. Mendoza-Granados and Sommers Solitary play. Play alone. Cf. play socially. See video. See also video enter hole. play start See solicit play with object in mouth. play walk Goodall: The chimp walks with a rounded-back, its head slightly bent down and pulled back between the shoulders while it takes small stilted steps. See also tilt head. Plooijs Play walk. Seen in bonobos. See videos. P play with another

animal Play with animal of another species. Includes wrestle and chase-flee play. At Gombe chimpanzee youngsters play with olive baboons. At Mahale chimpanzees from older infancy to adolescence seem to solicit human observers to play with them by hit and run, somersault, lift and drop, etc. Bonobos of Wamba play with red-colobus monkeys. Cf. care alloparentally for another species and trifle with. play with object Goodall Object play. Solo play, handling object such as fruit or stone. See also play, play socially with object and toy. See video. play with sand Sand apparently stimulates youngsters of M group to lone or social play. They roll over sand or somersault on it. Repeatedly grab, raise and release handfuls of sand in sitting or quadrupedal posture, appearing to enjoy movement

and sound of sand. Not reported from elsewhere. 7. See video. play with urine 139 play with urine Infant touches with hand urine passed by another chimp sitting overhead. 7. See video. play with water Solo play including splash water, look at water, stir water, move rock, nod quadrupedal etc. 7. See video. poke Nishida: Tickle, by prodding repeatedly with fingertips, with one or more extended fingers to infants ventrum. Adult also poke dominant individuals hand rhythmically for reassurance. Goodall mentioned that Sometimes an individual may poke another chimp who has ignored a solicitation for grooming. Plooijs Poke. Seen in bonobos. See video. poke in social scratch See scratch socially, poke type. poke with play face Poke once another individual with one finger, while showing play face. Type

of soliciting play. 7. See video. police Type of impartial intervention by adult males. Dominant males rush to place of commotion and run past quarreling parties, in order to stop quarrel. Boehms control role by alpha male at Gombe. Category 3. See videos. possessive behavior See herd. post-conflict behavior See aggress, redirected, appease, console and reconcile. pound Sanz and Morgan: percussion of tool onto another object or substrate. Pounding beehive customary at Goualougo, and likely to occur at Bwindi. Absent at Mahale. pound food on object Whiten et al.: Food-pound onto substrate, e.g. stone, root, etc. Customary at Gombe and habitual at Ta. Absent at Mahale. pound food on wood Whiten et al. Food-pound onto wood: smash food. Customary at Bossou, Ta and Gombe. Absent at Mahale. pound

insect Whiten et al Insect-pound: Use object to mash insect. Present at Bossou. Absent at Mahale. P 140 present with limbs flexed pound pestle Yamakoshi and Sugiyama Pestle pound: Bossou chimpanzee climbs to crown of oil palm and mashes apical growth tip of crown with mature petiole detached from oil palm. Unique to Bossou. See video. pound resin Whiten et al. Resin-pound: extract resin by pounding. Present at Bossou. Absent at Mahale. pout Plooij Pout face. Goodall Pout face: Lip corners pushed forward, mouth may be slightly open. Lips together, except at the front where they may funnel out somewhat. This is seen with the hoo call and hoo whimper. Seen in bonobo. See video. pout face See pout. pout kiss See kiss with pout face. predation on

chimpanzees See respond to predator. pregnancy Females in last stage of pregnancy often lie prone, rest, and travel alone. Pregnant female often recognizable from large belly. Cf. travel alone after childbirth. See gestation period. P presbyopia Farsightedness from aging presumed from greater distance old chimpanzee maintains from body part that he/she is grooming. See video. present for grooming See solicit grooming. present with limbs extended Normal quadrupedal stance with hindquarters directed to dominant chimpanzee. Goodalls Present, also Rump-turn. Plooijs PRE. van Hooffs Mount-present. Kanos Dorso-ventral presenting for bonobos. Bonobo females also show ventro-ventral courtship gesture. See video. present with limbs flexed van Hooff Crouch-present. Quadrupedal posture with limbs flexed, and hindquarters turned towards another. Differs from crouch, in which hindquarters not directed to partner. See video. press 141

press Force contact with shoulder, side or hip against another. Function unclear, but may initiate social play or aggression. 7. press down Restrain another under body to immobilize partner against substrate. Common in social play, such as wrestle and sometimes aggression. Cf. immobilize. See video. press neck with lower arm Press neck of wrestling partner with lower arm. See video. press nipple See fumble nipple. press object on Press branch, herb, etc. against playmate, thus indirectly applying force. Adolescent and juvenile males, e.g. Cadmus and Michio, used in play. See video. press teeth against back Adult male mounts another male, and mounter presses upper row of teeth against mounteds back, while mounter embraces other, thrusts pelvis, and has grin-full-open. Intensified form of mouth. Mount accompanied by nod and

mouth. Functions as reassurance and coalition formation, and sometimes during copulation. Rarely, juvenile male presses teeth against objects, such as mass of dry leaves. Absent in bonobos of Wamba. See video. probe Use vine, stem, bark, grass, twig, or stick to touch object. Goodalls Investigation probe. Infant chimpanzees appear innately inclined to push investigatory probe into tree hole, long before starting to fish for arboreal ants. Juvenile sometimes inserts probe inside of large fruit such as Voacanga to try to obtain juice. Customary at Gombe, Mahale, Bossou, and Ta. Present at Goualougo and Kibale. Probing for bees is customary at Ta and present at Mahale. Absent in bonobos of Wamba. See video. See also video probe with finger. probe nasal passage Adult male, Kalunde, inserted probe into nostril

to stimulate sneeze, which discharged much nasal mucus. He showed this behavior for decade. Only one other member of M group, Maggie, female adolescent, did so. See video. probe with finger Infant inserts finger into small tree hole, etc., to investigate contents of cavity. Cf. grope. 3. See video. P 142 pull each other process food Change raw material of food for ingestion by removing inedible parts, rendering into manageable bits with hand and teeth, etc. Include break branch with use of foot, break open by pushing chin, pull through with hand, pull through with mouth, twist, etc. prop anvil Whiten et al. Anvil-prop: Insert smaller stone under anvil to level its working surface. Seen multiple times at Bossou. Unique to Bossou. protect Goodall Protect. Caretaker protects infant

from rain, storm, danger, or social harassment. Also in bonobos, except for protection from rain. See video. protest When attacked, threatened, or stolen from, resolutely display dissatisfaction with bark, stare fixedly, scream, and chase to dominant attacker, especially if supporter present, then attacker may flee. Supporter of victim may join in or remain calm. Cf. retaliate. 3. See video. protrude tongue Small infant sticks out tongue. Function unclear. 8. See video. pucker cheek Suck in cheeks. Function unknown. 8. See video. P pull Grasp and tug at object or other by flexing arms. Often in feeding, play, grooming, or agonism. Alpha male of M group pulled estrous female unwilling to follow him. Estrous female, Ako, pulled juvenile grooming adult male so vacating place for her to copulate with

him. Plooijs Pull. Kanos and Pikas Pull for bonobos. See video. pull down Stretch arm to reach upper branch of shrub, often while standing bipedal, and pull branch downward. Then, sit and eat fruits or leaves while gripping branch with hand. Mother or playmate pulls down infant from overhead. Also in bonobos. See video. pull each other Two individuals simultaneously pull from opposite sides. Social play rarely observed. Two adolescent chimpanzees once pulled each other at stream, one standing in water and other standing on stepping stone. 9. See video. pull face to face 143 pull face to face Pull hand or other body part of another, to face each other. Occurs when chimpanzee wants to groom others venter. Also in bonobos. 2. pull head with hands Pull

head of another with hands, often in grooming. Also in bonobos. 2. See video. pull leaf-pile Walk backwards quadrupedally while raking dry leaves with hands. Locomotor-object play. Cf. push leaf-pile. See video. pull object from opposing sides Tug of war. Two individuals simultaneously pull object competitively from opposite sides. Component of social play. Also in bonobos. Cf. pull each other. See videos. See also video play socially with object. pull out Pull forcefully with hands, often countered by pushing feet against ground. Chimpanzee tries to uproot stalk of banana, elephant grass, or African ginger from earth. Also, standing quadrupedal and pulling up heavy rotten wood with arms. Also in bonobos. See video. pull rock to roll Adult or adolescent male pulls rock along substrate with hand while walking

or running, as part of charging display. Immature pulls or rakes rock as solo play while squatting. Same movement to pull up heavy wood with arms. 7. See video. pull through in courtship Nishida Stem pull-through. Pull leafy branch of shrub or clump of grass stems through hand by rapid upward movement of forearm. Then immediately release stem, producing conspicuous sound, while seated and watching estrous female. Only two of oldest adult males, Musa and Ntologi, did this courtship. Exaggerated form of pull through with hand in eating context. One adult male tore off shoot of stem from shrub in courtship. Customary at Bossou. pull through with hand Upward or outward sweeping movement of thumb and index finger or palmar surface of hand along leafy branch or leafy

herb stem. Common food-processing technique for leaves, vine, herbs or blossoms. Leaf-stripping technique also used to make tool for fishing carpenter ants. See fish. Gombe chimpanzees use in ant dipping P 144 push and pull sponge. Seen in bonobos. Category 12. See video. See also videos eat leaf and store. pull through with mouth Hold twig with hand and pull it between lips or teeth. Common foodprocessing technique for soft leaves and blossoms of trees, vines, and herbs. Used to make tool for fishing carpenter ants. 7. See video. pull with mouth Mouth and pull body part of another, in social play. If body dragged extensively, is drag other by mouth. 7. See video. punch See hit. puncture Dig underground termite nest with stout stick. Stand bipedal, push

stout stick with foot, while holding stick with hands. Customary at Goualougo and recorded at Ndoki. Absent at Mahale. See video. P push Extend arm or foot to make forceful contact in aggression, grooming, infantile courtship, social play, and play solicitation. To groom another, push part of back, so that groomee turns torso. Groomer stimulates groomee to change posture after extensive grooming of body-part. Mother pushes infant to descend from her back to ground. Mother gives infants back or rump swift push with hand or foot to make it walk ahead. Juvenile male sometimes pushes back of estrous female to solicit copulation. Kanos and Pikas Push in bonobos. See video. push ahead Mother drops off older infant from back or belly and encourages it to go ahead by

pushing or pulling so as to travel comfortably and secure infants security. Goodall: A mother pushes her infant ahead of her during weaning to prevent dorsal travel. Often she does this with little slaps of her hand on its rump and this is apparently perceived by the infant as playful. Whether or not playful slaps is unclear at Mahale. Absent in bonobos of Wamba. 5. See video. push and pull sponge Whiten et al. Sponge push-pull. Put leaf-sponge into hole of tree and pull it out with stick. At Mahale, two juveniles used stick to pick out leaf-sponge from tree hole. Present also at Gombe and Ta. push away 145 push away Shove away infant with palmar side of hand forcibly, or back side of hand gently. Goodall

Push: Shove away from another individual with one or both hands, and Goodalls 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. Adult male shoves away older infant from mother when its approach hinders grooming. Kanos Push away for bonobos. 2. See video. push backward Push rock backward with hand while running forward in riverbed. Element of charging display of some adult male chimpanzees of M group. Not in bonobos. See video. push down Adult male, Masudi, runs while pushing down short plants, such as grasses and shrubs one after another, in policing. See video. push finger into mouth Nishida: Insert one or more fingers/toes into the mouth of another

individual. Goodalls Finger in mouth includes both push finger into mouth and take finger in mouth. Appeases or reassures another individual, or calms excited adult male. Mother uses to stop infants threat-like vocalization. Not in bonobos, except when infants or juveniles beg for food. See video. push forward Juvenile female, Maggie, playfully and repeatedly pushes hard-shelled fruit forward on ground. Adult male, Alofu, repeatedly pushes stone forward, facing consorted female, Abi, to threaten that if she flees he will throw stone at her. See video. push head into ventral Social play with wrestling: Youngster pushes crown of head into playmates chest or abdomen. Goodalls Butt. Category 35. See video. push leaf-pile Nishida and Wallauer: Walk forward quadrupedally while pushing dry leaves with both hands. Locomotory-object play done mostly

by juveniles. Adults and adolescents use to threaten other. Cf. drag dry leaves with feet and pull leaf-pile. See video. push object into Stuff leaves, leaf sponges, or some other objects into tree hole. Occurs when drinking water by leaf sponge, inspecting inside of cavity, etc. Cf. probe. See video. P 14 put rump to rump push peri-anogenital region with finger Knead peri-anogenital region with extended fingers to extract hardened feces, helping softer feces inside to be passed. 8. See video. push/pull swing Mother or play partner pushes and/or pulls back and forth with hand youngster or vertical woody vine to which it clings. 8. See video. push shoulders See grasp and push shoulders. push to shoulder Push object against shoulder with hand. Function unknown. 8. See video.

put dorsal Nishida and Goodall Take dorsal. When infant not on mother, she grasps infants arm and brings to her shoulder to place onto her back. Cf. scoop infant, put ventral, and solicit riding. Also in bonobos. See video. put dorsal from ventral Goodall Take dorsal. Caretaker transfers infant from belly to back with scooping upward movement of hand. Cf. put ventral from dorsal. Also known for bonobos. See video. put face close See peer. P put face to Put face to carcass, apparently enjoying feel of soft hair. Done by adolescent male, Cadmus. Rare behavior. See video. put heel on back Adolescent male, Cadmus, put right heel on back of adult female, Totzy, in response to her pant-grunting to him. See video. put mouth into water Immerse

mouth in stream but without drinking. Water-contact play of juvenile chimpanzees in quadruped or lying posture. See video. put rump to rump Goodall Rump-to-rump: In response to a submissive present, the higher ranking chimp turns his rump to the other and they touch bottoms. Only seen in a few males. Very rare at Mahale and in few males and females. Cf. rub genital. Common in bonobos. 8. See video. put ventral 147 put ventral Goodall Take ventral: Mother reaches to infant, grasps, and presses to a ventral position. Cf. put dorsal, scoop infant and solicit riding. Kanos Take ventral in bonobos. See video. put ventral from dorsal Mother transfers infant from dorsal to ventral position, when entering dense bush or meeting possible danger, such as unknown humans. Cf.

put dorsal from ventral. See video. P Q quadrupedal hunch See hunch quadrupedal. quadrupedal jump See hop quadrupedal on spot and leap quadrupedal. quadrupedal run See run quadrupedal. quadrupedal transport See transport quadrupedal. quadrupedal walk See walk quadrupedal on knuckles. Q 148 R rain cover See cover from rain. rain dance See display, rain. raise Elevate limb while lying supine or sitting on ground. Also in bonobos. See video. raise and hold leg Raise leg and hold with one or both hands while lying supine. Pattern of grooming solicitation and also in resting. In grooming and resting in bonobos. See video. raise arm bipedal Extend arm and direct palm to companion. Intention movement to embrace or pull individual overhead during play or stressful confrontation. In stress, usually done

by grinning adult male. In play, usually done by youngster with play face. Also, used to grasp fruit in tree, or to climb tree. Infant does without anything overhead when requesting mother to allow access to nipple or to seek rescue. Signal for help probably originated from intention movement to seek dorsal ride on mother. See video. See also video approach and withdraw. raise arm quickly Goodall Arm threat: The arm is raised in a quick jerky movement and the fingers flexed slightly. This is usually accompanied by head tipping and cough threat or waa bark. Plooijs arm raise. Cf. shake arm. Adolescent male, Cadmus, raises arm quickly while standing bipedally for courtship as well as intimidation. Also in bonobos of Wamba. See video. raise arm slowly Extend

arm overhead, seeking to solicit grooming. See also solicit grooming. Also in bonobos. See video. T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 9, Springer 2010 149 R 150 rapid ohoh raise arm to hold branch Raise arm and grasp overhead branch or liana to solicit grooming. See also solicit grooming. Also in bonobos. See video. raise arm with elbow bent Extend arm overhead with bent elbow, to elicit grooming. If accompanied by self-scratching of armpit may stimulate grooming of armpit. Invitation for grooming-hand-clasp often complied with by mirror-image gesture from other, leading to grooming-hand-clasp. Also used when mother solicits infant to ride on her belly or back at onset of travel. See also solicit grooming. Also in bonobos. See video. See

also video solicit grooming raise leg while lying Lie supine or sideways and raise leg, soliciting grooming of thigh, scrotum, or leg. Probably same as Goodalls ventro-ventral present: When the sitting subordinate faces the dominant, leans back, raises one foot and rotates thighs laterally. Also in bonobos. See video. See also video groom raise others arm Elevate anothers arm. Groomer seeks to change groomees posture, so as to change body part to be groomed, and for groomers convenience. Also in bonobos. See video. raise others chin Elevate or lift up anothers lower jaw with hand, to groom exposed neck or jaw. Also in bonobos. 7. See video. See also video groom unilaterally raise others leg Same function as raise others arm. See video. R rake Rake or pull

dry leaves or sand toward self with one or both hands, while lying, sitting or quadrupedal. Solo play of youngsters. Used also as solicitation of play. Plooijs Rake. See also pull leaf-pile. 7. See video. See also video sway woody vegetation. rake-hunch See scratch dead leaves. rap Nishida Thump: Courtship pattern of males and females; sit and hit stem of shrub, branch, or ground with knuckles. Customary at Ta and Mahale. Habitual at Gombe. 8. rapid ohoh See pant-grunt. reach 151 reach See extend. reach hand See extend hand. reach leg See extend leg. reach wrist toward Goodall Wrist towards. Reach flexed wrist and back of hand to another. Cf. extend hand which is limited to gesture with wrist and fingers extended. At Mahale, subordinate behavior by some

adolescents and adult females when pant-grunting to adult males, or threatened by dominant females. Also when prohibiting youngster from approaching infant. See video. reassurance suck See suck in reassurance. reassure Response to submission or fearful expressions. Shown mostly by higher-ranking individuals. Goodalls Reassurance includes pat, touch, embrace, kiss, and groom. Behavioral patterns exist in bonobos, but not in reassurance. 3. See video. rebuff play Ignore play solicitation from another. Cf. solicit play. Cf. Hayakis Paraplay, which is unsuccessful attempt to initiate play with another. See video. reciprocal grooming See groom reciprocally. reconcile de Waal Reconciliation. Post-conflict behavioral patterns between combatants accompanying body contact, such as approach, touch, embrace, mount, kiss with open mouth and groom. Reported from Gombe, Mahale and Ta. Bonobos of Wamba reconcile by approach, touch,

mount, groom, rub genitals, and put rump to rump. See video. recover See retrieve infant. recruit support See solicit support. R 152 release infant to fall redirection of aggression See aggress, redirected. regulate direction Mother interferes in traveling of older infant by pulling or pushing until it changes direction of travel as she wants. Also, adult male forces estrous female to travel as he wants by threat and attack. See herd. Cf. depart together. 3. See video. reingest vomit Consume own vomit. Goodall Reingesting vomit: A sick chimp may vomit into one hand, then eat the vomit. Not seen in bonobos at Wamba. 5. See video. reject Turn down partners request, e.g. to groom, care alloparentally, mate, suckle, play, transport, etc. See video. reject infant Goodall Rejection of

infant by mother: Maternal behavior preventing her infant from having access to the nipple, climbing dorsal or ventral, sharing her food or sleeping in her bed, when the infant is being weaned. Behavioral patterns include cover nipple, fend, turn around, reject-move, reject-move, shrug, push ahead, reject-sit, turn around, fend, clamp elbow, and reject-move and threaten. See also wean. See video. reject-move Goodall: Mother avoids infant attempting to suckle, ride on her or beg food by moving away. May occur in bonobos. See video. R reject-sit Goodall: 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 bonobos. 5.

See video. relaxed face Goodall Relaxed face. Mouth closed and eyes open in normal circumstances. Kanos Relaxed face. See video. relaxed open mouth display See play face. release infant to fall Juvenile grasping hand of infant playmate released it, and let infant fall to ground. Seen once. 9. See video. remove 153 remove Divest object from self. Pick up with thumb and forefinger small objects, such as sticky seeds of grasses or beggars lice from body surface, in grooming. Also in bonobos. 7. See video. remove lice Groomer pulls from hair with hand and removes louse egg, or puts teeth directly to hair to pull off louse imago. See also scrape. See video. remove objects from water surface When streams begin to dry up, remove dry leaves, stones,

or other debris, to search for good drinking pool. With digging, important technique to obtain water, pick up. At least five chimpanzees at Mahale do this. Cf. dig for water. 8. See video. remove sand flea Prise open toe and toe nail with both hands and suck out sand flea eggs. 8. See video. rescue Mother or allomother rushes to scene of attack, retrieves youngster. See video. resin-pound See pound resin. respond to baboon Upon encountering yellow baboons, bark at, chase, or charge them. Chimpanzees of Mahale recently began to prey on yellow baboons. 7. See video. respond to dead animal Encounter intact dead bodies of red colobus monkey, red-tailed monkey, bushpig, bushbuck, aardvark, giant rat, leopard, etc. Approach and stare fixedly at carcasses of large-sized animals. Do

not eat old carcass, but rarely eat fresh one, which they had not killed. Responses to fresh adult leopard carcass included inspect, watch, sniff, slap, hit, bite, groom, lift and drop, drag, pull, throw at. See also eat carcass. 8. See video. respond to dead chimpanzee Mother carries dead infant for weeks to months. Other chimpanzees approach, stare fixedly, sniff and whisk fly with arms from corpse. May react to dead adult group mate by emitting wraa calls. May beat, bite, mutilate or eat dead chimpanzee. See transport corpse of infant. Cf. abuse carcass. See video. R 154 respond to lion respond to leopard React to leopards wood-sawing call by emitting wraa, bark or pant-hoot. Youngsters approach adults for reassurance. Chimpanzees killed leopard cub at Mahale and Ta.

See video. See Fig. 31. R Fig. 31 Respond to leopard: Adult male Bakali slapped leopard cub respond to lion React to lions roaring at night by emitting fear calls then pant-hoot. Also, chimpanzees shifted daytime range to higher altitudes. Chimpanzees escaped to tree canopy and emitted wraa calls when chased by lions in savanna woodland of Ugalla and Assirik. respond to neighboring unit group 155 respond to neighboring unit group Upon hearing calls from neighboring group, chimpanzees respond by pant-hooting and charging displays. They embrace each other and listen. If party is few in members, they silently retreat to center of territory. See video. respond to predator Chimpanzees stalked, attacked, killed and eaten by predators such as lions, leopards and humans. In west and central Africa, chimpanzees

are objects of bushmeat trade. At Mahale at least six chimpanzees were taken by lions, but no record of predation by leopard. At Ta several chimpanzees killed by leopard. No record of predation on chimpanzees by bird of prey at Mahale. See respond to lion and respond to leopard. rest Goodall Rest. Relax and remain immobile, sit, lean or lie. Kanos Resting. See video. restrain Prevent partner from continuing ongoing behavior to its or partners advantage. Cf. interfere. See video. restrict See detain. retaliate Counter-attack, that is return attack to initial attacker. Goodall: Aggressive response directed toward the original aggressor by a chimp who has been threatened or attacked. In bonobos, retaliation occurs more often when females are attacked than when males are targeted. Kano included as retaliation,

dominants response to challenging subordinate. Cf. protest. See video. retreat Walk backwards quadrupedally. Subordinate pant-grunts to dominant, and occasionally walks backwards fearfully in front of him. Adult male walks backwards, facing estrous or semi-estrous female, to confirm her following him. Also in bonobos. Cf. creep. See video. retreat bipedal Walk backwards bipedally. Juvenile does rarely, in order to solicit play from partner following it. 9. See video. retrieve infant Nishida Recover. Mother returns to infant left alone or with others, touches, grasps or transports it, in order to depart, groom, suckle, etc. R 15 reunite Occurs after infant emits distress calls, when danger threatens, or when mother wants to depart. Also in bonobos. See video. return infant Alloparent returns infant to mother, when infant emits distress calls. Absent

in bonobos of Wamba. See Fig. 32. R Fig. 32 Return infant: Adolescent female, Wakilufya, returned infant to its mother reunite Individuals meeting after separation show friendly or aggressive behavior. reverse 157 reverse Turn and travel in opposite direction. See video. ride bipedal Infant stands bipedal on mothers back, to reach branch overhead or just before leaping down to ground. 7. See video. ride clinging Goodall: Infant clings to mothers arm or leg as she travels. Also in bonobos. See video. ride dangling Infant hangs below mother with one or both hands only. Kanos Dangling riding for bonobos. See video. ride dangling and touch dry leaves One- to two year-old infant hangs below mother with hand and touches dry leaves on ground, apparently enjoying the sound produced. Solo

play in dry season at Mahale. 8. See video. ride dorsal Infant approaches, puts hands on mothers buttocks, and climbs up to ride on her back, or occasionally, another individual. Goodall discriminates three infant dorsal positions: Jockey, Lying and Quadrupedal. Kanos Dorsal travel for bonobos. See video. ride jockey Goodall: 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 others back, or gripping around the others side. Kanos Dorsal travel: jockey style for bonobos. See video. ride onto See step on. ride prone Infant travels by lying prone on mothers back. Kanos Dorsal travel: cling style for bonobos. See video. ride quadrupedal Goodall: When an infant stands on the back of another.

Kanos Dorsal travel: quadrupedal style for bonobos. See videos. ride supine Infant travels by lying supine on mothers back, occasionally and briefly. 8. See video. ride ventral Goodall Ventral ride: The infant is transported as he clings to the mothers belly, gripping hair between flexed fingers and toes.There are R 158 rub dorsum to conspecific following variations: Extended, Dangle, Arm or Leg-cling. Kanos Ventral riding for bonobos. See video. ride ventral with limb extended Goodall Ventral riding-extended: Ventral infant grips mothers hair with four limbs extended, so its back almost brushes ground as she travels. Seen in bonobos. See video. rinse Shake object sideways repeatedly in water. See wash. 9. rock back and forth Move torso with rhythmic, alternating forward and backward movements while sitting or standing bipedal,

in threat. Also at onset of charging display. Seen also during leap bipedal. Plooijs Rock. See video. rock side to side Goodall Rock: Slight or vigorous side to side movements of the body when the chimp is sitting.Rocking occurs when a male is working up to a charging display. At Mahale, rocking prelude to charging display and when nervous. Bonobo rocking occurs only in courtship, not as warming up for charging display. See video. roll Goodall Roll: A chimpanzee may hit another and roll over him during aggression or play. Bonobos use in aggression. See video. rooting See nuzzle. R rotate fruit Juvenile male, Michio, lies supine on ground, extends legs upward and rotates big fruit such as Voacanga lutescens with his feet. Solo object play. See video.

rough and tumble play See play, rough and tumble. rub dorsum Lean against vertical tree trunk and rub wet shoulders or back against it, often holding arms upward, after heavy rain. May lie supine on rock, ground, grass, horizontal bough, or day-bed to rub against surface. Recorded at Gombe. Also in bonobos of Wamba. 5. See video. rub dorsum to conspecific Juvenile may lean against back of adult male to solicit play. Rare at Mahale. Idiosyncratic?. See video. rub genitals 159 rub genitals Female rubs genitals against another female. At Mahale rarely seen, and limited to pairs in which one participant is juvenile or adolescent. At Bossou more frequent and ventro-dorsal, and put rump to position seen. Bonobo females often show species-specific G-G -rubbing as reassurance. See video.

rub genitals to substrate Female rubs genitals against substrate such as ground or tree trunk. 9. See video. rub hand or foot Corp et al. Hand-rubbing. Rub hand or foot against branch of tree, stalk of shrub, rock, or ground when eating juicy fruit of lemon or sticky fruit such as Saba. After touching feces or other dirty object, immediately rub soiled hand or foot against ground. Cf. brush away from branch. 7. See video. rub hand with hand Rub hands together briskly and repeatedly, e.g. after removal of ectoparasite from body. 7. See video. rub muzzle Corp et al. Muzzle-rub. Rub muzzle stained with fruit juice against shrub, grass stalk, slender vine, bark of tree, or stone, to wipe off mess. Differs from wipe with detached object

because body-part put to part of the environment, not tool to body. At Mahale, only one adult female did this, after eating seeds of Parkia filicoidea several times before 1990s. Became common in 1998; by 2004, 29 chimps of M group did so when eating juicy fruit of lemon or sticky fruit of Saba comorensis and Voacanga lutescens. Goodall omitted but occurs at Gombe. See video. rub object to body Move quickly object such as leafy branch along body, back and forth. Adult male, Pim, rubbed large dead bird on nape and head while pressing it firmly. Juvenile female, Athena, lay on her back and rubbed her belly with smooth stone. 9. See video. rummage Search for target object by removing or separating other objects. See video. rump

to rump See put rump to rump. rump turn See present with limbs extended. R 1 0 rush to embrace run Rapid locomotory gait with which body momentarily off ground in each stride cycle. See video. run away See flee. run bipedal Goodall Bipedal run. Chimpanzees run on feet only during charging display. Adult male, Masudi, once ran on feet over ground swarming with Dorylus ants, to avoid being bitten. Also in bonobos. See video. run quadrupedal Goodall Quadrupedal run. See video. run tripedal Run using feet and arm while other arm carries object. See drag branch. Kanos Tripedal run. rush to embrace Frightened youngster dashes into arms of older individual, usually mother, often followed by sucking. 3. R S safari See consort. Salonga See Lui Kotale. sand

See play with sand. sand flea Jigger. Common ectoparasite that often burrows under toenails and lays eggs, annoying human, chimpanzee and baboon. See also remove sand flea and medicate self. scan Look in all directions seeking something. Lost juvenile often does so seeking mother after climbing tree. Adults do when looking for others. See also search for conspecific. Also in bonobos. Cf. monitor. scatter dry leaves Rake, shift to left or right, around, or throw forward dry leaves with hand. Infantile expression of dissatisfaction or protest, or solo play. Cf. scratch dry leaves, which adults show as prelude to charging display. 7. See video. scavenge See eat carcass. scoop algae Scoop algae from water using wand of vegetation. Customary at Bossou, and at least one adult male at

Odzala, but not elsewhere. Cf. eat algae. See video. 1 1 S 1 2 scratch self scoop infant Goodall: When the mother is about to travel and the infant is on the 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. Cf. put dorsal, put ventral and solicit riding. Plooijs Scoop. Kanos Scoop for bonobos. See video. scramble for food Compete for precious food such as meat by agonistic gestures and calls. See video. scrape Scrape lice eggs through hair with incisors. Also resins of trees such as Terminalia mollis. See also eat resin. Category 37. See video. scratch Rake nails of partly flexed fingers over surface. Goodalls

Scratch. Plooijs Scratch. Seen as Self-scratch in bonobos. See videos. scratch aggressively Scratch another to inflict wound. Goodalls Aggressive scratch: A chimpanzee may scratch another during a fight. Probably also in bonobos. S scratch dry leaves Scratch or rake with one or both hands dead leaves piled on ground, as initiation of pant-hooting or charging display, as solicitation of grooming, as solicitation of play, or as courtship. Done by infants as solo play. Maybe same as Goodalls Scrub or Rake. Cf. scatter dry leaves. Plooijs Rake. Bonobos rake before dragging Haumania vine in display. 5. See video. scratch self Goodall 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. Goodalls Scratch

self directly: ---during social grooming the groomee often scratches a body part that is then groomed by the partner. Occurs when frustrated and uneasy. Plooijs Self-scratch. Kanos Self-scratch for bonobos. See video. scratch self directly 1 3 scratch self directly See scratch self. scratch self signalling Goodall 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. Bonobos use at start of grooming. See video. scratch socially Nishida: Scratch or Rub. Nakamura et al. Social

scratch. Scratch vigorously skin of anothers back or arm at beginning or middle of grooming bout, usually with one hand, making audible sound. Absent at Gombe as tradition but seen in one male and two females in 2001. Bonobos of Wamba occasionally scratch another with first and second fingers in grooming. Category. See video. scratch socially, poke type When scratching another at Mahale, few rarely poke with straight fingers. Customary among chimpanzees of Ngogo. 9. See video. scratch socially, stroke type When scratching another at Mahale, usually stroke gently with bent fingers. See video. scream Goodall:.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. See also squeal in copulation

and whimper-scream. Kano divided bonobo scream calls into Scream and Screech. See video. See also video monitor mother. scream, food Adult male, Fanana, screams when eating prized food, such as ripe fruit of lemon, Saba or Garcinia, especially at beginning of feeding bout. Absent in Goodall. Bonobos Food peep or Food grunt may be homologous. scrub See scratch dry leaves. scrub pelt Rub colobus skin against rock in streambed, apparently to soften it. Done by one adult male. See wash. S 1 4 self-suckle search for conspecific Look for unseen others, in several ways: sit and listen, walk around, then return to original place, sniff ground, in particular, at path intersections and footprints, or climb tree and scan surroundings, frequently done by lost, whimpering juvenile. Mother also looks

for unweaned infant. Mature male looks for estrous female. Behavior includes: scan, walk, run, climb, grin, listen, whimper, whimper-scream, scream, pant-hoot, sniff, look back, etc. Absent in bonobos of Wamba. 5. See video. search for object Search for object, such as food, rock to throw, sleeping tree, etc. Includes: scan, walk, climb, move, touch, pull, stare fixedly, sniff, etc. 3. See video. seat-stick Branch used to avoid sharp thorns when sitting on kapok tree. Reported from Tenkere and Goualougo. Absent at Mahale. See also stepping-stick. seat vegetation See make leaf cushion. seed dispersal See disperse seed. self-clasp See clasp self. self-groom See groom self. self-inspect See inspect self. S self-medication See medicate self. self play See play solo. self-scratch See scratch self. self-slap See slap self. self-suckle See

suck self. self-tickle 1 5 self-tickle See tickle self with object. semen Whitish rubbery liquid containing sperm ejected by penis. See ejaculate. See Fig. 33. Fig. 33 Semen: Semen drips from females bottom after copulation Semliki Study area of Pan troglodytes schweinfurthii within Toro-Semliki Wildlife Reserve, Uganda. Research organized by K. Hunt from 199 to present. senescence Symptoms of old age are presbyopia, cataract, greying hair on head, bent and balding back, worn teeth, loss of teeth, lack of estrus, slow movement, avoidance of climbing tall trees, reduced range, etc. See video. separate Party splits into two or more smaller parties or subgroups. See video. separating intervention See intervene to separate. sexual skin Anogenital region of females. See swelling of sexual skin. See Fig. 34. S 1 shake

branch Fig. 34 Sexual skin: Sexual skin often bitten by adult males, so deforming shape shake arm, abduct Sit and shake horizontally-held arm towards another, in mild threat, often accompanied by cough bark. Cf. raise arm quickly and hit toward. 5. See video. shake arm, adduct Sit or stand bipedal and shake horizontally-held arm at another, to come closer or solicit play. Probably originated as intention movement of grab, grasp or hug. Juvenile male, Xmas, once shook hand above waters surface while looking down at his reflection. Sometimes in threat, if shaken quickly. See video. shake back and forth See sway woody vegetation. S shake back and forth with four limbs Cling to and shake tree trunk forward and backward using pushing force of feet and pulling force

of arms. Threatening gesture. Infants and young juveniles do as solo play. 7. See video. shake body See shiver. shake branch Goodall Branch shake: A branch is shaken with quick jerky movements of the arm, slight or vigorous. The chimp may be sitting or shake branch up and down with feet 1 7 standing. Plooijs Branch. Nishidas Branch-shake: Courtship pattern of males and females: Sit while shaking horizontally or vertically stem of shrub, branch of tree, or woody vine. Branch-shaking signals request to another to come closer or to depart together. Contexts include courtship, continuation of consortship, enticement to travel together, threat, and solicitation of play. Cf. sway woody vegetation in which chimp bipedally shakes tree trunk, etc. back and force forcibly with both arms. Cf. also flail

long object. Kanos Branch-shaking and Ingmansons Branch waving for bonobos of Wamba. Hohmann and Fruths Branch shake for bonobos of Lomako. See video. shake branch up and down with feet Stand on horizontal branch, woody vine, or fallen log on ground, and shake it up and down with one or both feet, in threat display or solo play. Weanling does as part of temper tantrum, namely, reaction to lack of response of mother or another, as attention-getting. Reported only at Mahale. See video. shake detached branch Hold detached, leafy branch in hand or foot and shaken horizontally in courtship, done by several juvenile males. shake face side to side Amuse self by watching reflection on waters surface. Juvenile male, Xmas, vigorously shook face side to side. Cf. nod

quadrupedal. 9. See video. shake face with object in mouth Hold object such as leaf-sponge in mouth and vigorously shake face side to side in stream. Done as solo play by adolescent male, Cadmus, only. See video. shake hand See shake arm, abduct or shake arm, adduct. shake hand side to side quickly Shake hand from side to side quickly, while pant-grunting. Conflict behavior shown by subordinate chimpanzee to dominant male. See video. shake hands See grasp hand. S 1 8 share bed shake head Shake vigorously head side to side while trying to spray about water or sand atop head. Immatures shake head during play, such as play walk. Cf. tilt head and nod quadrupedal. 5. See video. shake long object irregularly Adolescent male, Cadmus, sits and

irregularly shakes long dry grass, both vertically and horizontally, as threat to human observer. Category 9. See video. shake object up and down Stand and shake up and down bush or dead tree by hand, in threat or social play. See video. shake off While seated or standing, forcefully shake off play partner from body contact. Throw down opponent by hand in wrestling as in sumo. See video. shake other Old female, Calliope, grasped knee of Fanana and shook it back and force in greeting. Adult female, Xtina, grasped shoulder of another adult female, Ruby, and shook it back and forth when agitated. Adolescent male, Cadmus, grasped and shook juvenile males heel as play. Idiosyncratic?. See video. shake penis Adult male grasps and shakes vertically anothers penis, while

sometimes panting. Rare at Mahale. Function as reassurance? Cf. fumble penis. See video. shake rock up and down with feet Stand quadrupedal on rock and bounce up and down on it with feet. Solo play pattern of water-contact. 8. See video. S shake rump Mother shakes lower back from side to side, causing infant to descend to ground. See drop infant. 5. See video. shake wet arm to catch termite Catch winged termites by shaking arms wet with rain. Technique employed for termites of Pseudacanthotermes. 9. share bed Mother allows infant offspring to sleep in same bed. Cf. allow. See Fig. 35. share food 1 9 Fig. 35 Share bed: share food Allow another to take food under control of owner. Same as Feistner and McGrews Transfer of

food. Most common between mothers and infants. Meat is most common food to be shared among adults. Meat sharing among unrelated adults occurs at Gombe, Mahale, Ta, and Ngogo. Fruit sharing occurs at Gombe, Mahale, Budongo, Bossou, Ngogo and Ta. Large fruits such as Treculia africana and papaya often shared. Sugar cane sharing occurs at Mahale. Food share occurs in bonobos at Wamba and Lomako. See video. shelter from rain Take cover from rain e.g., by going under canopy of big tree. Includes walk and sit. Cf. cover from rain. shiver Chimps shiver when cold or wet. show off Display by dominant individual in aggression between individuals meeting after long separation. Includes bristle, charge, display, charging, hunch, slap, stamp, splash water, swagger bipedal, attack, etc. In reunion, bonobos

show display, charging in aggression. Cf. display as contest. S 170 sit prone shrub-bend See bend shrub. shrug Goodall: 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. Plooijs Shrug. Common at Mahale. See also drop infant. Cf. lower head and shoulder. Absent in bonobos of Wamba. 5. See video. sickness Symptoms of sickness include cough, diarrhea, lethargy, lie, pick ear, pick nose, sneeze, sniffle, vomit, runny nose, unstable walking, etc. See video. silent bared-teeth display See grin. sit Plooij SIT. Goodall: Rump on ground, branch, etc., body more or less vertical. Arms sometimes crossed. Kanos Sit. See videos. sit and lean Sit and extend both arms backwards and