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BEHAVIOUR  - Editorial Comment

Editorial Comment

(Editorial Overview Text Replicated on Overall Species page - Pan paniscus - Bonobo)

Female bonobos carry their infants mainly on their belly (ventrally) initially; after about two years of age it is carried mainly on her back, and it is carried during travelling to about 3-4 years of age. Bonobos probably nurse their offspring to about four years. Mothers keep their infant in reach at all times when they first start moving off their mother at about six months of age. They play with their infants during rest periods and are generally indulgent, and they protect their offspring against other bonobos. The bond between females and their daughters is broken in late juvenile/early adolescence, but the bond with male offspring continues, even into adulthood. When the inter-birth interval is relatively short, a female may care for, carry and suckle two dependent offspring at the same time. Other bonobos, including older male offspring and even unrelated adult males, sometimes show alloparenting, being very caring and affectionate towards infants, including playing with them, carrying them, and sharing nests and food with them, although mothers generally limit the distance the male can take the infant away from her. On one occasion when a female gave birth when her previous infant was only a year old, an adult male often cared for the older infant; on another occasion, a youngster orphaned at under six years of age (too young for independent survival) was cared for by two males.

Further information on reproduction is provided in Pan paniscus - Bonobo - Life Stages (Literature Reports)

(References are available in detailed literature reports below)

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Parental Behaviour

Source Information

  • A female carries her infant ventrally most of the time until it is two years old, after which it is regularly carried on her back, but may still be carried ventrally in a crisis to four years. (B577.7.w7)
  • Initially when the infant starts to move off its mother at about six months old, she keeps it in reach at all times; if it tries to go further away she uses a hand to bar its way and carries it again. (B577.7.w7)
  • The female regularly carries her infant while travelling for 3-4 years. To initiate this, she vocalises, then walks up to 6m before standing still with one foot slightly lifted, sole raised behind her, until the juvenile runs and jumps onto her back. In a tree, she waits in an ascending or descending posture. She may extend a hand towards her offspring if it is close by. (B577.7.w7)
  • Females often play with their infants during rest periods. A female may tickle, grab at the infant with her hands, and play-bite. Also, in a day nest, lie on her back and raise it up with her hands and feet (airplane). She rarely plays with her offspring once it becomes a juvenile. (B577.7.w7)
  • A females start grooming her infant once it is about 5 - 6 months old; the amount of grooming increases with the offspring's age. (B577.7.w7)
  • Females probably nurse their offspring to about four years. (B577.7.w7)
  • Females are indulgent towards their offspring, not threatening or punishing them and, with rare exceptions, not taking food from them. (B577.7.w7)
  • A female will protect her offspring from aggression shown by another adult, even if the aggressive adult outranks the female. (B577.7.w7)
  • Females have been seen carrying two offspring at one time, the younger on her belly and the older on her back, and have been seen suckling two offspring at one time. (J552.4.w1)
  • Females often move rhythmically and rock their infant. Several types of play have been recorded from mother to offspring including: shaking her head from side to side, suspending the infant by its hand or foot, rhythmically shaking the branch the infant is on, grasping the infant by the hand and swinging it, rubbing the infant with her sexual skin, holding the infant between her legs and shaking it and play panting at the infant. Play behaviours by the infant on her which she permits include climbing on her, jumping onto her, hanging from her, hanging from her and swinging, touching, grasping or rubbing her sexual skin, and kicking. It was noted that "infants looked as though they regarded their mothers as a jungle gym." Females sometimes interfere with play fights between their own offspring and another bonobo. (J576.31.w3)
  • Mothers may support their adolescent or young adult sons in agonistic interactions with other bonobos; this may lead to the son rising in rank and becoming dominant over older males (including young adults becoming dominant over prime males). (J576.33.w1)
  • Females carried infants and juveniles on their belly or back. While the female was feeding, an older infant or juvenile often left its mother to play be itself, but when the female left the tree she would generally carry her offspring; she would stretch out a hand, or stop while standing on all four limbs, back towards the juvenile, for it to climb on her back. If a juvenile was left behind by a female while she descended, it would usually cry out and the female would wait for it to catch up, although in one instance a female was seen returning to get her offspring. Sometimes females would share food with juveniles who were capable of getting the food item in question (Croton haumanianus fruit) themselves. (J552.4.w1)
  • A female may appear severely depressed by the death of an infant and may carry the infant. (B577.7.w7)
  • The relationship with female offspring is broken sometime during the daughter's adolescence, but that with sons continues. During adolescence and even adulthood, a son usually forages with his mother, and they engage in grooming. Even adult males sometimes show begging behaviour towards their mother. Females show less maternal care as their offspring get older, but sometimes intervene with warning vocalisations or even physically attacking if an adolescent or adult son is attacked. (B577.7.w7)
  • Daughters show increased distance from their mother during foraging from about 6-7 years, but remain in the same foraging party. At about nine years of age, daughters emigrate to other unit groups (communities. Males at 6-7 years still remain close to their mothers and return to them once travel begins; as adults, they still remain in the same foraging party as their mothers. (B594.2.w2d)
Alloparenting
  • Males show generalised paternalism, being very caring and affectionate towards infants, including sharing nests and food. (B580.5.w5)
    • Males cannot know whether any particular infant is or is not their own offspring. (B580.5.w5)
  • Alloparenting behaviour may be displayed by an adolescent male towards a younger sibling. Adolescent and adult males show tolerance towards their younger (infant or juvenile) siblings and often groom them. One adolescent male, Tawashi, would frequently approach his mother, peer into her face then take his infant sibling away, carrying her on a walk, and even playing (tickling, holding up by the arms, embracing and open-mouth kissing all over. This adolescent male also showed alloparenting when the infant died, carrying her, placing the corpse in a tree fork near his nest for the night and grooming it briefly the following morning. (B577.7.w7)
  • Alloparenting occurs by unrelated males: the male will approach the female and stare at her infant while extending his hand; the infant may jump on his back or cling to his stomach, and the male leaves with the infant. The mother will allow only a distance of about 5m and if the male goes further away, or the infant whimpers, she will hurry after to retrieve it. (B577.7.w7)
  • Strong attachments of juveniles to males were seen, including a juvenile going into the day nest of a male and staying for an hour, with reciprocal grooming occurring for 20 minutes. Another time, an older juvenile foraged near one male for about two hours, and once in the yoku a party of an adult male and an older juvenile was seen. (J552.4.w1)
  • In observations at Wamba, adult and adolescent males showed a variety of behaviours towards infants, including embracing, sham-biting, holding the infant between his legs and abdomen while gazing at its face, often with pelvic thrusting, dandling (sitting the infant on his legs and rocking it repeatedly with the legs), head-bobbing, shaking the head from side to side while embracing the infant, touching with the mouth (on the head, shoulder, loin, back, mouth or genitals), tickling with the foot, grasping, placing on abdomen, lifting (lying on his back, placing the infant on his abdomen then grasping it with his hands and/or feet and lifting it over his abdomen), wrestling, grooming and play panting. Infants showed the following behaviours towards allofathers: touching the penis, getting on the male (either climbing up him or jumping on), kicking, making a play face, play pant, and pant. (J576.31.w3)
  • At Wamba, short than usual interbirth intervals sometimes resulted in a female having more than one dependent and nursing offspring at one time. In a case in which the interbirth interval was only one year, the older offspring was often cared for by an adult male. (J552.19.w2)
  • When a female at Wamba gave birth when her previous offspring was only 3.5 years old (compared to the more usual six years), a special relationship developed between the older infant and five of the mid- to high-ranking males, who proceeded to share much of the care normally provided by the mother for infants of this age. These males performed behaviours often seen performed to younger maternal siblings by males: playing, grooming, carrying, huddling, protection from other group members, and food sharing. (J564.20.w1)
  • Based on mitochondrial DNA sequencing, the probable (deceased) mother of a juvenile male, seen alone (orphaned) at about 5.5-6 years old, was determined. Records indicated he had been orphaned at under four years of age. None of the group showed any aggressive behaviour towards him and two adult males (not including his older brother) were observed taking care of him - moving with, feeding and grooming him in a similar way to mothers with their offspring; these behaviours may have enabled him to survive. (J576.37.w1)
  • Captive data: In a group of bonobos at San Diego Zoo (adult male, unrelated adult female, two female offspring of five and three years of age), the male sometimes carried the infant either ventrally or dorsally, as well as demonstrating other parental behaviours such as retrieving the infant when she was "hurt", patted her, and held her to prevent her from leaving. Parental behaviours by the male were less frequent than those by the female. The female carried the infant ventrally (88% of the time) or dorsally (in the wild, infants over one year of age are mainly carried dorsally), retrieved the infant when the infant was "hurt" or "endangered", held the infant, preventing it leaving her or interacting with the other individuals, "kissed" the infant (touching the infant's face with pouted, protruding lips). On one occasion, in response to "pout face" and "hou hou" verbalisation from the infant sitting before her, the female got up, sucked water into her mouth from a drinking spigot, walked back, sat down facing the infant, leaned forward and puckered her lower lip, allowing the infant to suck water from her mouth; this was repeated a further three times. (J576.20.w2)
    • Occasionally the male carried the infant, ventrally or dorsally, and was seen to retrieve it when it was "hurt", hold it to prevent it leaving, and to pat the infant. These behaviours were similar to those shown by the mother, but much less frequent. (J576.20.w2)
  • Captive data: A juvenile female, Kichele, was first seen to snatch her infant brother, Ke Ke, from her mother when the infant was five months old. Within a week, she was allowed to take him for a minute or two at a time, always approaching and grooming Diatou before taking or trying to take the infant. Later (at eleven months of age) Keke was seen back-riding both adults and occasionally Kickele. It was notice that as the infant grew older, caretaking duties were shared more by bonobos other than his mother. (N4.23.w3)
  • Captive data: Data from comparative studies at Yerkes indicate that mothers show closer and more continuous attention to their infants than do Pan troglodytes mothers. (B594.2.w2d)
  • Captive data: A comparative study considered that female bonobos were more likely to leave very young infants than Pan troglodytes females were, but that for older infants chimpanzees were more likely than bonobos to break contact and leave their infant. Bonobos show more, shorter nursing bouts than do chimpanzees, and showed higher rejection and nipple rejection (preventing or breaking nursing by the infant); hypotheses for this were more active regulation of infant nursing to regulate postpartum ovulation, or restriction of nursing at times when it would interfere with other activities. (J564.64.w1)
  • Captive data: An instance of infant theft was observed in the group at Twycross Zoo. The dominant female, Diatou, was found holding a newborn infant. However, she had been showing regular sexual cycles in the weeks prior to the birth, while the lowest ranking female, Banya, appeared weak, had heavy, irregular vaginal bleeding, and showed an irregular swelling cycle in the weeks after the birth. Genetic testing later confirmed that Banya was the mother. (J577.77.w2)

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Authors & Referees

Authors

Dr Debra Bourne MA VetMB PhD MRCVS (V.w5)

Referee

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