Bonobos (Pan paniscus) live in the forests located centrally in the Democratic Republic of the Congo (formerly Zaire). Bonobo habitat lies in the Congo Basin. This area is located south of an arc formed by the Congo River (formerly the Zaire River) and its headwater, the Lualaba River, and north of the Kasai Rver.
Biogeographic Regions: ethiopian (Native )
- Kano, T. 1982. The social group of pygmy chimpanzees (Pan paniscus) of Wamba. Primates, 23/2: 171-188.
- Kano, T. 1983. An ecological study of the pygmy chimpanzees (Pan paniscus) of Yalosidi, Republic of Zaire. International Journal of Primatology, 4/1: 1-31.
- Kano, T. 1992. The Last Ape: Pygmy Chimpanzee Behavior and Ecology. Stanford, CA: Stanford University Press.
Contrary to the implication of one of its common names, "pygmy chimpanzee," this species is not particularly diminutive when compared to common chimpanzees (Pan troglodytes). The "pygmy" modifier may instead refer to its location: it lives in an area inhabited by people often referred to as such.
Unlike its closest cousins (common chimpanzees), bonobos are not divided into subspecies. Bonobos are apes about two-thirds the size of humans, with dark hair covering their bodies. The hair is generally longer than in common chimpanzees, and is particularly noticeable on the cheeks, which are relatively hairless in P. troglodytes. The portions of body not covered with hair (i.e. mid-face, hands, feet) are darkly colored throughout life. This contrasts with common chimpanzees, which have lighter skin, particularly during the younger years.
Bonobos are primarily knuckle-walkers, although at times they walk bipedally and do so more frequently than P. troglodytes. Bonobos have longer extremities, particularly hind legs, as compared to common chimpanzees. Although sexual dimorphism exists with males around 30% heavier (37 to 61 kg, 45 kg average) than females (27 to 38 kg, 33.2 kg average), bonobos are less sexually dimorphic than many primates, and skeletons are nearly the same size. Average height is 119 cm for males and 111 cm for femals. Average cranial capacity is 350 cubic centimeters.
Range mass: 27 to 61 kg.
Average mass: 39 kg.
Range length: 104 to 124 cm.
Average length: 115 cm.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: male larger
- Boesch, C. 2002. Behavioural diversity in Pan . Pp. 1-8 in C Boesch, G Hohmann, M Linda, eds. Behavioural Diversity in Chimpanzees and Bonobos. Cambridge, UK: The Press Cyndicate of the University of Cambridge.
- Jungers, W., R. Susman. 1984. Body size and skeletal allometry in African apes. Pp. 131-177 in R Susman, ed. The Pygmy Chimpanzee: Evolutionary Biology and Behavior. New York, NY: Plenum Press.
- Zihlman, A. 1984. Body build and tissue composition in Pan paniscus and Pan troglodytes, with comparisons to other hominoids. Pp. 179-200 in R Susman, ed. The Pygmy Chimpanzee: Evolutionary Biology and Behavior. New York, NY: Plenum Press.
Within the Congo Basin, bonobos inhabit several vegetation types. The area generally is classified as tropical rainforest; however, local agriculture and areas reverted to forest from agriculture (“young” and “aged secondary forest”) are intermingled. Species composition, height, and density of trees are different in each, yet all are utilized by bonobos. In addition to the forested areas, swamp forests opening into marsh-grassland areas occur, which are also utilized. Foraging occurs in each type of habitat, while sleeping occurs in forested areas. Some bonobo populations may have a preference to sleep in relatively small (15 to 30 m tall) trees, particularly those found in secondary growth forests.
Range elevation: 299 to 479 m.
Habitat Regions: tropical ; terrestrial
Terrestrial Biomes: forest ; rainforest
- Fruth, B., G. Hohmann. 1993. Ecological and behavioral aspects of nest building in wild bonobos (Pan paniscus). Ethology, 94: 113-126.
- Uehara, S. 1988. Grouping patterns of wild pygmy chimpanzees (Pan pansicus) observed at a marsh grassland amidst the tropical rain forest of Yalosidi, Republic of Zaire. Primates, 29/1: 41-52.
- Uehara, S. 1990. Utilization patterns of a marsh grassland within the tropical rain forest by the bonobos (Pan paniscus) of Yalosidi, Republic of Zaire. Primates, 31/3: 311-322.
Habitat and Ecology
Fruit comprises the largest portion of the diet of P. paniscus, although bonobos incorporate a wide variety of other food items into their diet. Plant parts consumed include fruit, nuts, stems, shoots, pith, leaves, roots, tubers and flowers. Mushrooms are also occasionally consumed. Invertebrates form a small proportion of the diet and include termites, grubs, and worms. On rare occasions, bonobos have been known to eat meat. They have been directly observed eating flying squirrels (Anomalurus sp.), duiker (Cephalophus dorsalis and Cephalophus nigrifrons), and bats (Eidolon sp.).
Animal Foods: mammals; eggs; insects; terrestrial worms
Plant Foods: leaves; roots and tubers; wood, bark, or stems; seeds, grains, and nuts; fruit; flowers
Other Foods: fungus
Primary Diet: herbivore (Frugivore )
- Badrian, N., R. Malenky. 1984. Feeding ecology of Pan paniscus in the Lomako Forest, Zaire. Pp. 275-299 in R Susman, ed. The Pygmy Chimpanzee: Evolutionary Biology and Behavior. New York, NY: Plenum Press.
- Kano, T., M. Mulavwa. 1984. Feeding ecology of the pygmy chimpanzees (Pan paniscus) of Wamba. Pp. 233-274 in R Susman, ed. The Pygmy Chimpanzee: Evolutionary Biology and Behavior. New York, NY: Plenum Press.
The quantity of fruit consumed by bonobos suggests that they may play a role in dispersal of the species eaten.
Ecosystem Impact: disperses seeds
Species Used as Host:
- non known
- none known
- none known
The only verified predators of bonobos are humans. Although the hunting of bonobos is illegal, poaching is still common. It has been speculated that leopards and pythons, known to prey on common chimpanzees, may also feed on bonobos.
- humans (Homo sapiens)
- Van Krunkelsven, E. 2001. Density estimation of bonobos (Pan paniscus) in Salonga National Park, Congo. Biological Conservation, 99: 387-391.
Life History and Behavior
Bonobos communicate in a variety of ways. Females have a scream, but males bark, grunt, and pant-hoot. A bark may indicate alarm, whereas other vocalizations may indicate aggression, excitement, satisfaction, etc. The separate types of calls are used in multiple contexts, and cannot be thought of as "words".
In addition to this vocal communication, tactile communication is important. Social rank is communicated by GG rubbing, mounting, or rump contact. (See behavior section.) Other forms of tactile communication are seen between mothers and their offspring, and between rivals.
Visual communication also occurs. Bonobos often "peer" at another individual. This behavior indicates interest in the activity of the "peered at" individual. Peering may occur when oneother bonobo has a food item that is wanted, or it may be included in the courtship behavior of a male.
Communication Channels: visual ; tactile ; acoustic ; chemical
Perception Channels: visual ; tactile ; acoustic ; chemical
Some researchers have suggested that one difference between humans and chimpanzees is that only humans voluntarily share their food with others. Hare and Kwetuenda (2010) experimentally investigated voluntary food sharing in unrelated Bonobos. In their experiments one individual (the "subject") was given the choice of either monopolizing food or actively sharing it with another individual (the "recipient") by releasing the other individual from an adjoining room. This experimental design eliminated relatedness and harassment as motivating factors for sharing. The researchers found that subjects showed a significant preference to open the potential recipient’s door rather than another door to an empty room. Subjects released recipients to co-feed for the majority of the total feeding time during each trial. Compared to control trials in which subjects were presented with one empty room and another room with additional food but no other Bonobo, subjects opened the door to a room with a potential food recipient more quickly than they opened the door to a room with additional food. In a limited set of additional control trials, two test subjects more often released a recipient into a room with food rather than into an empty room, suggesting the possibility of striking altruism. Results from this set of experiments suggest that our own species’ propensity for voluntary food sharing may not be unique among the apes. (Hare and Kwetuenda 2010)
Limited information exists on bonobo longevity, and there has been no ongoing study that lasted longer than the expected bonobo lifespan. The longest semi-continuous study of bonobos began at Wamba in 1976. At that time, the age of each individual was estimated, and from extrapolation, a female that died in 1993 was in the 45 to 50 year age range when she died. This would make the lifespan of these animals comparable to that of common chimpanzees.
Status: wild: 50 (high) years.
Status: captivity: 35.0 years.
Status: captivity: 48.0 years.
Status: wild: 20.0 years.
- Furuichi, T. 1989. Social interactions and the life history of female Pan paniscus in Wamba, Zaire. International Journal of Primatology, 10/3: 173-197.
Lifespan, longevity, and ageing
Bonobos are polygynandrous. Females may be approached by and copulate with, any male in the group except their sons. However, the mating system may be confused by the use of sexual activity in these animals as part of social bond formation.
Mating System: polygynandrous (promiscuous)
Basic life history traits of bonobos are under-researched. Some of the seminal studies of this species have noted that “bonobos have not yet been studied long enough to provide data on age at sexual maturity or birth interval” (Nishida and Hiraiwa-Hasegawa, 1987), the most frequently researched “Wamba and Lomako study populations lack long-term demographic data” (Thompson-Handler et al., 1984), and “information on the demography of wild bonobos is very limited compared to that for chimpanzees” (Furuichi et al., 1998).
Female bonobos undergo estrus, marked by distinctive swelling of the perineal tissue lasting 10 to 20 days. Matings are concentrated during the time of maximal swelling. Breeding occurs throughout the year. Postpartum amenorrhea lasts less than one year in bonobos. A female may resume external signs of estrus (i.e. swelling) within a year of giving birth. At this point, copulation may resume, although these copulations do not lead to conception, indicating that the female is probably not fertile. During this period, she continues to lactate until her offspring is weaned at around 4 years. The average interbirth interval is 4.6 years (4.8 if one only includes live births). Therefore, lactation may suppress ovulation, but not the outward signs of estrus. As no study has lasted longer than a bonobo lifespan, total number of offspring per female is unknown. However, at Wamba, many adult females had four offspring during the 20 year study length.
Adult female bonobos have an estrus period that is marked externally by physical changes in their genitalia. During this time, males of the group approache the female, displaying their erect penises. Females are generally receptive, and will move toward a male to allow copulation. There is no clear pattern of mate choice: females are courted by many males of the group during estrus, with the exception of their sons. Because of this, paternity is generally unknown to both partners.
Breeding interval: Breeding occurs nearly all the time in this species, however, a female may produce one offspring approximately every 5 years.
Breeding season: Bonobos have no marked breeding season.
Average number of offspring: 1.
Average gestation period: 240 days.
Average weaning age: 48 months.
Range time to independence: 7 to 9 years.
Range age at sexual or reproductive maturity (female): 13 to 15 years.
Range age at sexual or reproductive maturity (male): 13 to 15 years.
Key Reproductive Features: iteroparous ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization ; viviparous
Average birth mass: 1331 g.
Average gestation period: 232 days.
Average number of offspring: 1.
Information is limited on parental investment. However, bonobos are highly social mammals and live around 15 years before achieving full adult status. During this time, the mother provides most of the parenting, although the males may contribute indirectly (i.e. in alerting the group of danger, sharing food, and possibly helping to protect young).
Bonobo babies are born relatively helpless. They are dependent on mothers’ milk and cling to their mother for several months. Parental care is provided by the mother, as paternity is generally unclear. Weaning is a gradual process, and is usually commenced by the time the offspring is 4 years of age. Throughout the weaning process, mothers generally have their offspring feed by their side, allowing them to observe the feeding process and food choice, rather than providing them with food directly. Weaning may be enforced by a mother’s refusal to allow a juvenile into her nest, thereby encouraging it to build a nest of its own.
As adults, male bonobos typically remain in their natal social group, so they have contact with their mothers throughout her remaining years. Female offspring leave their natal group during late adolescence, so they do not maintain contact with their mothers in adulthood.
Parental Investment: altricial ; pre-fertilization (Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male, Female); pre-independence (Protecting: Male, Female); post-independence association with parents; extended period of juvenile learning; inherits maternal/paternal territory; maternal position in the dominance hierarchy affects status of young
- Dahl, J. 1986. Cyclic perineal swelling during the intermenstrual intervals of captive female pygmy chimpanzees (Pan paniscus). Journal of Human Evolution, 15: 369-385.
- Fruth, B., G. Hohmann. 1993. Ecological and behavioral aspects of nest building in wild bonobos (Pan paniscus). Ethology, 94: 113-126.
- Furuichi, T. 1987. Sexual swelling, receptivity, and grouping of wild pygmy chimpanzee females at Wamba, Zaire. Primates, 23/3: 309-318.
- Furuichi, T., G. Idani, H. Ihobe, S. Kuroda, K. Kitamura, A. Mori, T. Enomoto, N. Okayasu, C. Hashimoto, T. Kano. 1998. Population dynamics of wild bonobos (Pan paniscus) at Wamba. International Journal of Primatology, 19/6: 1029-1043.
- Horn, A. 1980. Some observations on the ecology of the bonobo chimpanzee (Pan paniscus, Schwarz 1929) Near Lake Tumba, Zaire. Folia primatologica, 34: 145-169.
- Kano, T. 1992. The Last Ape: Pygmy Chimpanzee Behavior and Ecology. Stanford, CA: Stanford University Press.
- Savage-Rumbaugh, E., B. Wilkerson. 1978. Socio-sexual behavior in Pan paniscus and Pan troglodytes: A comparative study. Journal of Human Evolution, 7: 327-344.
Molecular Biology and Genetics
Barcode data: Pan paniscus
Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.
See the BOLD taxonomy browser for more complete information about this specimen and other sequences.
-- end --
Download FASTA File
Statistics of barcoding coverage: Pan paniscus
Public Records: 27
Specimens with Barcodes: 36
Species With Barcodes: 1
Bonobos are an endangered species according to both IUCN and US Federal Endangered Species lists. The IUCN criteria project a 50% or greater reduction in their numbers within three generations, due to both exploitation and habitat destruction. Bonobos face “a very high risk of extinction in the wild in the near future” according to the IUCN Red List criteria. Civil war and its aftermath have hampered conservation efforts. Population estimates vary widely as conflict has limited the ability of researchers to work in the region. Estimates range from 5,000 to 17,000 individuals left.
US Federal List: endangered
CITES: appendix i; appendix ii
IUCN Red List of Threatened Species: endangered
IUCN Red List Assessment
Red List Category
Red List Criteria
- 1994Vulnerable(Groombridge 1994)
- 1990Vulnerable(IUCN 1990)
- 1988Vulnerable(IUCN Conservation Monitoring Centre 1988)
- 1986Vulnerable(IUCN Conservation Monitoring Centre 1986)
Date Listed: 10/19/1976
Lead Region: Foreign (Region 10)
Where Listed: Entire
Population location: Entire
Listing status: E
For most current information and documents related to the conservation status and management of Pan paniscus , see its USFWS Species Profile
Follow the link below for Table 1: population densities for Pan paniscus.
There are no substantive data concerning total numbers, although speculative estimates give numbers for a total population size between 29,500 (Myers Thompson 1997) and 50,000 (Dupain and Van Elsacker 2001). Recent surveys indicate that these numbers may still be underestimates. In any case, any number indicating total population size should be considered with the highest caution.
Commercial poaching has to be considered the most prominent threat. In some areas local taboos against bonobo hunting still exist, in others they are disintegrating due to changing cultural values associated with transient and immigrant human populations. Although commercial hunting is targeted at large-bodied ungulates and monkeys, the growing predominance of bushmeat commerce as an income-generating activity has led to increases in the number of commercial bushmeat hunters. These hunters, aided by military and local administration, are active in all areas, including those with legally protected status such as Salonga National Park. The importance of Salonga National Park as a significant reservoir of bonobos will be rapidly compromised if current hunting trends continue. Between 2003 and 2006, Hart et al. (2007) recorded evidence of hunting across the park in 51% of survey grids. Hunting pressure was considered to be high in the north and east of the park, and bonobo mortalities as a direct consequence of hunting were recorded. Bonobos are slow-breeding and thus particularly susceptible to loss caused directly by poaching or indirectly by snaring.
Infectious diseases are yet to be quantified but are undoubtedly an important threat to wild bonobo populations, and of particular concern in areas where bonobos live side-by-side with humans. The risk of transmission increases with increasing human population density as well as increasing proximity with wildlife.
In the context of the Congo Basin (Congo Basin Forest Partnership (CBFP) 2005), an international project for the protection of forests, DRC possesses three landscapes important for bonobo conservation: Lac Tele-Lac Tumba Landscape (Congo and DRC); Maringa-Lopori-Wamba Landscape (DRC); and Salonga-Lukenie-Sankuru Landscape (DRC).
Within these landscapes are areas with established protection status such as Salonga National Park (since 1970) and Luo Scientific Reserve (since 1990). However, significant portions of bonobo habitat such as the Lomami-Lualaba and the Lomela-Sankuru areas are not included in the landscapes defined by CBFP.
Due to the combined efforts of the ICCN (DRC’s national conservation authority) and international NGOs, two additional areas obtained official protected status in 2006: Faunal Reserve of Lomako-Yokokala (RFLY) (Maringa-Lopori-Wamba Landscape), and Tumba-Lediima Natural Reserve (RTL) (Lac Tele-Lac Tumba Landscape). Others are under consideration and protected only by local commitments: Yasa-Bososandja wildlife sanctuary (1998) (Myers Thompson 2001), and Kokolopori (reserve to be gazetted in 2007) (BCI 2007).
Although these areas still harbour sizeable numbers of bonobos, the threats cited above put all populations at risk irrespective of the conservation status of the area. Conservation education programmes are essential to help curb poaching and illegal trade, not only in areas adjacent to wild populations but also in urban centres such as Kinshasa or Kisangani, where the demand for bushmeat and pets is generated. Lola Ya Bonobo, a sanctuary for confiscated bonobos in Kinshasa, welcomes 15,000 visitors per annum, half of which are school children, who can influence local attitudes (ABC 2006).
In sum, for effective protection of bonobos, commercial hunting must be halted, intensification rather than expansion of local agriculture must be supported, and local industries must be actively persuaded to support rather than subvert conservation. Additional surveys are needed to better determine the species’ overall distribution and abundance.
Despite the fact that bonobos breed well in captivity, and captive propagation programmes exist in North American and European zoos, only conservation measures in situ can be considered useful attempts to contribute to the species’ survival in the wild.
It is listed on CITES Appendix I.
Relevance to Humans and Ecosystems
Bonobos may eat sugarcane that is being grown for profit. However, direct references to this being a problem for humans have not been encountered in the literature.
Bonobos, similar to common chimpanzees, carry many of the same diseases that can afflict humans, such as polio.
Negative Impacts: injures humans (carries human disease); crop pest
Bonobos and their sister species, common chimpanzees, are the closest relatives to Homo sapiens. They are an invaluable source of information in studying human origins and diseases.
Bonobos are endearing to humans as 'charasmatic megafauna' and may be useful in encouraging conservation for habitat preservation.
Bonobos continue to be a source of bush meat for human consumption, and although hunting bonobos has been legally outlawed, poaching continues.
Positive Impacts: food ; research and education
The bonobo (// or //) (Pan paniscus), formerly called the pygmy chimpanzee and less often, the dwarf or gracile chimpanzee, is an endangered great ape and one of the two species making up the genus Pan; the other is Pan troglodytes, or the common chimpanzee. Although the name "chimpanzee" is sometimes used to refer to both species together, it is usually understood as referring to the common chimpanzee, whereas Pan paniscus is usually referred to as the bonobo.
The bonobo is distinguished by relatively long legs, pink lips, dark face and tail-tuft through adulthood, and parted long hair on its head. The bonobo is found in a 500,000 km2 (190,000 sq mi) area of the Congo Basin in the Democratic Republic of the Congo, Central Africa. The species is omnivorous and inhabits primary and secondary forests, including seasonally inundated swamp forests. Political instability in the region and the timidity of bonobos has meant there has been relatively little field work done observing the species in its natural habitat.
Along with the common chimpanzee, the bonobo is the closest extant relative to humans. Because the two species are not proficient swimmers, the formation of the Congo River 1.5–2 million years ago possibly led to the speciation of the bonobo. Bonobos live south of the river, and thereby were separated from the ancestors of the common chimpanzee, which live north of the river. There is no concrete data on population numbers, but the estimate is between 29,500 and 50,000 individuals. The species is listed as Endangered on the IUCN Red List and is threatened by habitat destruction and human population growth and movement, though commercial poaching is the most prominent threat. They typically live 40 years in captivity, though their lifespan in the wild is unknown.
- 1 Etymology
- 2 Evolutionary history
- 3 Physical description
- 4 Behavior
- 5 Habitat and distribution
- 6 Conservation status
- 7 See also
- 8 Notes
- 9 References
- 10 Further reading
- 11 External links
Despite the alternative common name "pygmy chimpanzee", the bonobo is not especially diminutive when compared to the common chimpanzee. "Pygmy" may instead refer to the pygmy peoples who live in the same area. The name "bonobo" first appeared in 1954, when Eduard Paul Tratz and Heinz Heck proposed it as a new and separate generic term for pygmy chimpanzees. The name is thought to be a misspelling on a shipping crate from the town of Bolobo on the Congo River, which was associated with the collection of chimps in the 1920s. The term has also been reported as being a word for "ancestor" in an extinct Bantu language.
Fossils of Pan species were not described until 2005. Existing chimpanzee populations in West and Central Africa do not overlap with the major human fossil sites in East Africa. However, Pan fossils have now been reported from Kenya. This would indicate that both humans and members of the Pan clade were present in the East African Rift Valley during the Middle Pleistocene. According to A. Zihlman, bonobo body proportions closely resemble those of Australopithecus, leading evolutionary biologists like Jeremy Griffith to suggest that bonobos may be a living example of our distant human ancestors.
Taxonomy and phylogeny
German anatomist Ernst Schwarz is credited with having discovered the bonobo in 1928, based on his analysis of a skull in the Tervuren museum in Belgium that previously had been thought to have belonged to a juvenile chimpanzee. Schwarz published his findings in 1929. In 1933, American anatomist Harold Coolidge offered a more detailed description of the bonobo, and elevated it to species status. The American psychologist and primatologist Robert Yerkes was also one of the first scientists to notice major differences between bonobos and chimpanzees. These were first discussed in detail in a study by Eduard Paul Tratz and Heinz Heck published in the early 1950s.
|NCBI genome ID|
|Genome size||2,869.21 Mb|
|Number of chromosomes||24 pairs|
|Year of completion||2012|
The first official publication of the sequencing and assembly of the bonobo genome became publicly available in June 2012. It was deposited with the International Nucleotide Sequence Database Collaboration (DDBJ/EMBL/GenBank) under the EMBL accession number AJFE01000000 after a previous analysis by the National Human Genome Research Institute confirmed that the bonobo genome is about 0.4% divergent from the chimpanzee genome. In addition, Svante Pääbo's group at the Max Planck Institute for Evolutionary Anthropology is currently sequencing the genome of a female bonobo from the Leipzig zoo.
Initial genetic studies characterised the DNA of chimpanzees and bonobos as being as much as 98% (99.4% in one study) identical to that of Homo sapiens. Later studies showed that chimpanzees and bonobos are more closely related to humans than to gorillas. In the crucial Nature paper reporting on initial genome comparisons, researchers identified 35 million single-nucleotide changes, five million insertion or deletion events, and a number of chromosomal rearrangements which constituted the genetic differences between the two Pan species and humans, covering 98% of the same genes. While many of these analyses have been performed on the common chimpanzee rather than the bonobo, the differences between the two Pan species are unlikely to be substantial enough to affect the Pan-Homo comparison significantly.
There still is controversy, however. Scientists such as Jared Diamond in The Third Chimpanzee, and Morris Goodman of Wayne State University in Detroit suggest that the bonobo and common chimpanzee are so closely related to humans that their genus name also should be classified with the human genus Homo: Homo paniscus, Homo sylvestris, or Homo arboreus. An alternative philosophy suggests that the term Homo sapiens is the misnomer rather, and that humans should be reclassified as Pan sapiens, though this would violate the Principle of Priority, as Homo was named before Pan (1758 for the former, 1816 for the latter). In either case, a name change of the genus would have implications on the taxonomy of extinct species closely related to humans, including Australopithecus. The current line between Homo and non-Homo species is drawn about 2.5 million years ago, and chimpanzee and human ancestry converge only about 7 million years ago, nearly three times longer.
DNA evidence suggests the bonobo and common chimpanzee species effectively separated from each other fewer than one million years ago. The Pan line split from the last common ancestor shared with humans approximately six to seven million years ago. Because no species other than Homo sapiens has survived from the human line of that branching, both Pan species are the closest living relatives of humans and cladistically are equally close to humans. The recent genome data confirms the genetic equidistance.
The bonobo is commonly considered to be more gracile than the common chimpanzee. Although large male chimpanzees can exceed any bonobo in bulk and weight, the two species actually broadly overlap in body size. Adult female bonobos are somewhat smaller than adult males. Body mass in males ranges from 34 to 60 kg (75 to 132 lb), against an average of 30 kg (66 lb) in females. The total length of bonobos (from the nose to the rump while on all fours) is 70 to 83 cm (28 to 33 in). When adult bonobos and chimpanzees stand up on their legs, they can both attain a height of 115 cm (45 in). The bonobo's head is relatively smaller than that of the common chimpanzee with less prominent brow ridges above the eyes. It has a black face with pink lips, small ears, wide nostrils, and long hair on its head that forms a part. Females have slightly more prominent breasts, in contrast to the flat breasts of other female apes, although not so prominent as those of humans. The bonobo also has a slim upper body, narrow shoulders, thin neck, and long legs when compared to the common chimpanzee.
Bonobos are both terrestrial and arboreal. Most ground locomotion is characterized by quadrupedal knuckle walking. Bipedal walking has been recorded as less than 1% of terrestrial locomotion in the wild, a figure that decreased with habituation, while in captivity there is a wide variation. Bipedal walking in captivity, as a percentage of bipedal plus quadrupedal locomotion bouts, has been observed from 3.9% for spontaneous bouts to nearly 19% when abundant food is provided. These physical characteristics and its posture give the bonobo an appearance more closely resembling that of humans than that of the common chimpanzee. The bonobo also has highly individuated facial features, as humans do, so that one individual may look significantly different from another, a characteristic adapted for visual facial recognition in social interaction.
Multivariate analysis has shown bonobos are more neotenized than the common chimpanzee, taking into account such features as the proportionately long torso length of the bonobo. Other researchers challenged this conclusion.
Primatologist Frans de Waal states bonobos are capable of altruism, compassion, empathy, kindness, patience, and sensitivity, and described "bonobo society" as a "gynecocracy".[a] Primatologists who have studied bonobos in the wild, have documented a wide range of behaviors, including aggressive behavior and more cyclic sexual behavior similar to chimpanzees, even though the fact remains that bonobos show more sexual behavior in a greater variety of relationships. An analysis of female bonding among wild bonobos by Takeshi Furuichi stresses female sexuality and shows how female bonobos spend much more time in estrus than female chimpanzees. Some primatologists have argued that de Waal's data reflect only the behavior of captive bonobos, suggesting that wild bonobos show levels of aggression closer to what is found among chimpanzees. De Waal has responded that the contrast in temperament between bonobos and chimpanzees observed in captivity is meaningful, because it controls for the influence of environment. The two species behave quite differently even if kept under identical conditions. A 2014 study also found bonobos to be less aggressive than chimpanzees, particularly eastern chimpanzees. The authors argued that the relative peacefulness of western chimpanzees and bonobos was primarily due to ecological factors.
Most studies indicate that females have a higher social status in bonobo society. Aggressive encounters between males and females are rare, and males are tolerant of infants and juveniles. A male derives his status from the status of his mother. The mother–son bond often stays strong and continues throughout life. While social hierarchies do exist, rank plays a less prominent role than in other primate societies.
Because of the promiscuous mating behavior of female bonobos, there is a great deal of paternal uncertainty. If a male cannot be sure which offspring are his, he is less likely to invest any time or energy caring for them. It is because of this lack of certainty that the entirety of parental care in bonobos is assumed by the mothers.
Bonobo party size tends to vary because the groups exhibit a fission–fusion pattern. A community of approximately 100 will split into small groups during the day while looking for food, and then will come back together to sleep. They sleep in nests that they construct in trees.
Sexual activity generally plays a major role in bonobo society, being used as what some scientists perceive as a greeting, a means of forming social bonds, a means of conflict resolution, and postconflict reconciliation. Bonobos are the only non-human animal to have been observed engaging in all of the following sexual activities: face-to-face genital sex (though a pair of western gorillas has been photographed performing face-to-face genital sex), tongue kissing, and oral sex.
Bonobos do not form permanent monogamous sexual relationships with individual partners. They also do not seem to discriminate in their sexual behavior by sex or age, with the possible exception of abstaining from sexual activity between mothers and their adult sons. When bonobos come upon a new food source or feeding ground, the increased excitement will usually lead to communal sexual activity, presumably decreasing tension and encouraging peaceful feeding. This quality is also described by Dr. Susan Block as "The Bonobo Way" in her book of the same title "The Bonobo Way: The Evolution of Peace Through Pleasure"
Bonobo clitorises are larger and more externalized than in most mammals; while the weight of a young adolescent female bonobo "is maybe half" that of a human teenager, she has a clitoris that is "three times bigger than the human equivalent, and visible enough to waggle unmistakably as she walks". In scientific literature, the female–female behavior of bonobos pressing genitals together is often referred to as genito-genital (GG) rubbing, which is the non-human analog of tribadism, engaged in by human females. This sexual activity happens within the immediate female bonobo community and sometimes outside of it. Ethologist Jonathan Balcombe stated that female bonobos rub their clitorises together rapidly for ten to twenty seconds, and this behavior, "which may be repeated in rapid succession, is usually accompanied by grinding, shrieking, and clitoral engorgement"; he added that it is estimated that they engage in this practice "about once every two hours" on average. Because bonobos occasionally copulate face-to-face, "evolutionary biologist Marlene Zuk has suggested that the position of the clitoris in bonobos and some other primates has evolved to maximize stimulation during sexual intercourse". On the other hand, the frequency of face-to-face mating observed in zoos and sanctuaries is not reflected in the wild, and thus may be an artifact of captivity. The position of the clitoris may alternatively permit GG-rubbings, which has been hypothesized to function as a means for female bonobos to evaluate their intrasocial relationships.
Bonobo males occasionally engage in various forms of male–male genital behavior, which is the non-human analog of frotting, engaged in by human males. In one form, two bonobo males hang from a tree limb face-to-face while penis fencing. This also may occur when two males rub their penises together while in face-to-face position. Another form of genital interaction (rump rubbing) occurs to express reconciliation between two males after a conflict, when they stand back-to-back and rub their scrotal sacs together. Takayoshi Kano observed similar practices among bonobos in the natural habitat.
More often than the males, female bonobos engage in mutual genital behavior, possibly to bond socially with each other, thus forming a female nucleus of bonobo society. The bonding among females enables them to dominate most of the males. Although male bonobos are individually stronger, they cannot stand alone against a united group of females. Adolescent females often leave their native community to join another community. This migration mixes the bonobo gene pools, providing genetic diversity. Sexual bonding with other females establishes these new females as members of the group.
Bonobo reproductive rates are no higher than those of the common chimpanzee. During oestrus, females undergo a swelling of the perineal tissue lasting 10 to 20 days. Most matings occur during the maximum swelling. The gestation period is on average 240 days. Postpartum amenorrhea (absence of menstruation) lasts less than one year and a female may resume external signs of oestrus within a year of giving birth, though the female is probably not fertile at this point. Female bonobos carry and nurse their young for four years and give birth on average every 4.6 years. Compared to common chimpanzees, bonobo females resume the genital swelling cycle much sooner after giving birth, enabling them to rejoin the sexual activities of their society. Also, bonobo females which are sterile or too young to reproduce still engage in sexual activity. Adult male bonobos have sex with infants. Frans de Waal, an ethologist who has studied bonobos remarked, "A lot of the things we see, like pedophilia and homosexuality, may be leftovers that some now consider unacceptable in our particular society."
The bonobo is an omnivorous frugivore; 57% of its diet is fruit, but this is supplemented this with leaves, honey, eggs, meat from small vertebrates such as anomalures, flying squirrels and duikers, and invertebrates. In some instances, bonobos have been shown to consume lower-order primates. Some claim bonobos have also been known to practise cannibalism in captivity, a claim disputed by others. However, at least one confirmed report of cannibalism in the wild of a deceased infant was described in 2008.
Observations in the wild indicate that the males among the related common chimpanzee communities are extraordinarily hostile to males from outside the community. Parties of males 'patrol' for the neighboring males that might be traveling alone, and attack those single males, often killing them. This does not appear to be the behavior of bonobo males or females, which seem to prefer sexual contact over violent confrontation with outsiders. In fact, the Japanese scientists who have spent the most time working with wild bonobos describe the species as extraordinarily peaceful, and de Waal has documented how bonobos may often resolve conflicts with sexual contact (hence the "make love, not war" characterization for the species). Between groups, social mingling may occur, in which members of different communities have sex and groom each other, behavior which is unheard of among common chimpanzees. Conflict is still possible between rival groups of bonobos, but no official scientific reports of it exist. The ranges of bonobos and chimpanzees are separated by the Congo River, with bonobos living to the south of it, and chimpanzees to the north. It has been hypothesized that bonobos are able to live a more peaceful lifestyle in part because of an abundance of nutritious vegetation in their natural habitat, allowing them to travel and forage in large parties.
Recent studies show that there are significant brain differences between bonobos and chimps. The brain anatomy of bonobos has more developed and larger regions assumed to be vital for feeling empathy, sensing distress in others and feeling anxiety, which makes them less aggressive and more empathic than their close relatives. They also have a thick connection between the amygdala, an important area that can spark aggression, and the ventral anterior cingulate cortex, which helps control impulses. This thicker connection may make them better in regulating their emotional impulses and behavior.
Bonobo society is dominated by females, and severing the lifelong alliance between mothers and their male offspring may make them vulnerable to female aggression. De Waal has warned of the danger of romanticizing bonobos: "All animals are competitive by nature and cooperative only under specific circumstances" and that "when first writing about their behaviour, I spoke of 'sex for peace' precisely because bonobos had plenty of conflicts. There would obviously be no need for peacemaking if they lived in perfect harmony."
Surbeck and Hohmann showed in 2008 that bonobos sometimes do hunt monkey species. Five incidents were observed in a group of bonobos in Salonga National Park, which seemed to reflect deliberate cooperative hunting. On three occasions, the hunt was successful, and infant monkeys were captured and eaten.
Similarity to humans
Bonobos are capable of passing the mirror-recognition test for self-awareness, as are all great apes. They communicate primarily through vocal means, although the meanings of their vocalizations are not currently known. However, most humans do understand their facial expressions and some of their natural hand gestures, such as their invitation to play. Two bonobos at the Great Ape Trust, Kanzi and Panbanisha, have been taught how to communicate using a keyboard labeled with lexigrams (geometric symbols) and they can respond to spoken sentences. Kanzi's vocabulary consists of more than 500 English words, and he has comprehension of around 3,000 spoken English words. Kanzi is also known for learning by observing people trying to teach his mother; Kanzi started doing the tasks that his mother was taught just by watching, some of which his mother had failed to learn. Some, such as philosopher and bioethicist Peter Singer, argue that these results qualify them for "rights to survival and life" — rights that humans theoretically accord to all persons. (See great ape personhood.) Afterwards Kanzi was also taught how to use and create stone tools in 1990. Then within 3 years, three researchers- Kathy Schick, Nicholas Toth and Gary Garufi- wanted to test Kanzi's knapping skills. Though Kanzi was able to form flake technology, he didn't create it the way they expected. Unlike the way hominids did it where they held the core in one hand and knapped it with the other, Kanzi threw the cobble against a hard surface or against another cobble. This allowed him to produce a larger force to initiate a fracture as opposed to knapping it in his hands.
As in other great apes and humans, third party affiliation toward the victim – the affinitive contact made toward the recipient of an aggression by a group member other than the aggressor – is present in bonobos. A 2013 study  found that both the affiliation spontaneously offered by a bystander to the victim and the affiliation requested by the victim (solicited affiliation) can reduce the probability of further aggression by group members on the victim (this fact supporting the Victim-Protection Hypothesis). Yet, only spontaneous affiliation reduced victim anxiety – measured via self-scratching rates – thus suggesting not only that non solicited affiliation has a consolatory function but also that the spontaneous gesture – more than the protection itself – works in calming the distressed subject. The authors hypothesize that the victim may perceive the motivational autonomy of the bystander, who does not require an invitation to provide post-conflict affinitive contact. Moreover, spontaneous – but not solicited – third party affiliation was affected by the bond between consoler and victim (this supporting the Consolation Hypothesis). Importantly, spontaneous affiliation followed the empathic gradient described for humans, being mostly offered to kin, then friends, then acquaintances (these categories having been determined using affiliation rates between individuals). Hence, consolation in the bonobo may be an empathy-based phenomenon.
Instances in which non-human primates have expressed joy have been reported. One study analyzed and recorded sounds made by human infants and bonobos when they were tickled. Although the bonobos' laugh was at a higher frequency, the laugh was found to follow a spectrographic pattern similar to that of human babies.
Habitat and distribution
Bonobos are found only south of the Congo River and north of the Kasai River (a tributary of the Congo), in the humid forests of the Democratic Republic of Congo of central Africa. Ernst Schwarz's 1927 paper “Le Chimpanzé de la Rive Gauche du Congo”, announcing his discovery, has been read as an association between the Parisian Left Bank and the left bank of the Congo River; the bohemian culture in Paris, and an unconventional ape in the Congo.
The IUCN Red List classifies bonobos as an endangered species, with conservative population estimates ranging from 29,500 to 50,000 individuals. Major threats to bonobo populations include habitat loss and hunting for bushmeat, the latter activity having increased dramatically during the first and second Congo wars in the Democratic Republic of Congo due to the presence of heavily armed militias even in remote "protected" areas such as Salonga National Park. This is part of a more general trend of ape extinction.
As the bonobos' habitat is shared with people, the ultimate success of conservation efforts will rely on local and community involvement. The issue of parks versus people is salient in the Cuvette Centrale the bonobos' range. There is strong local and broad-based Congolese resistance to establishing national parks, as indigenous communities have often been driven from their forest homes by the establishment of parks. In Salonga National Park, the only national park in the bonobo habitat, there is no local involvement, and surveys undertaken since 2000 indicate the bonobo, the African forest elephant, and other species have been severely devastated by poachers and the thriving bushmeat trade. In contrast, areas exist where the bonobo and biodiversity still thrive without any established parks, due to the indigenous beliefs and taboos against killing bonobos.
The port town of Basankusu is situated on the Lulonga River, at the confluence of the Lopori and Maringa Rivers, in the north of the country, making it well placed to receive and transport local goods to the cities of Mbandaka and Kinshasa. With Basankusu being the last port of substance before the wilderness of the Lopori Basin and the Lomako River—–the bonobo heartland—conservation efforts for the bonobo use the town as a base.
In 1995, concern over declining numbers of bonobos in the wild led the Zoological Society of Milwaukee, in Milwaukee, Wisconsin, with contributions from bonobo scientists around the world, to publish the Action Plan for Pan paniscus: A Report on Free Ranging Populations and Proposals for their Preservation. The Action Plan compiles population data on bonobos from 20 years of research conducted at various sites throughout the bonobo's range. The plan identifies priority actions for bonobo conservation and serves as a reference for developing conservation programs for researchers, government officials, and donor agencies.
Acting on Action Plan recommendations, the ZSM developed the Bonobo and Congo Biodiversity Initiative. This program includes habitat and rain-forest preservation, training for Congolese nationals and conservation institutions, wildlife population assessment and monitoring, and education. The Zoological Society has conducted regional surveys within the range of the bonobo in conjunction with training Congolese researchers in survey methodology and biodiversity monitoring. The Zoological Society’s initial goal was to survey Salonga National Park to determine the conservation status of the bonobo within the park and to provide financial and technical assistance to strengthen park protection. As the project has developed, the Zoological Society has become more involved in helping the Congolese living in bonobo habitat. The Zoological Society has built schools, hired teachers, provided some medicines, and started an agriculture project to help the Congolese learn to grow crops and depend less on hunting wild animals.
During the wars in the 1990s, researchers and international non-governmental organizations (NGOs) were driven out of the bonobo habitat. In 2002, the Bonobo Conservation Initiative initiated the Bonobo Peace Forest Project supported by the Global Conservation Fund of Conservation International and in cooperation with national institutions, local NGOs, and local communities. The Peace Forest Project works with local communities to establish a linked constellation of community-based reserves, managed by local and indigenous people. Although there has been only limited support from international organizations, this model, implemented mainly through DRC organizations and local communities, has helped bring about agreements to protect over 5,000 square miles (13,000 km2) of the bonobo habitat. According to Dr. Amy Parish, the Bonobo Peace Forest "is going to be a model for conservation in the 21st century."
With grants from the United Nations, USAID, the U.S. Embassy, the World Wildlife Fund, and many other groups and individuals, the Zoological Society also has been working to:
- Survey the bonobo population and its habitat to find ways to help protect these apes
- Develop antipoaching measures to help save apes, forest elephants, and other endangered animals in Congo's Salonga National Park, a UN World Heritage site
- Provide training, literacy education, agricultural techniques, schools, equipment, and jobs for Congolese living near bonobo habitats so that they will have a vested interest in protecting the great apes – the ZSM started an agriculture project to help the Congolese learn to grow crops and depend less on hunting wild animals.
- Model small-scale conservation methods that can be used throughout Congo
Starting in 2003, the U.S. government allocated $54 million to the Congo Basin Forest Partnership. This significant investment has triggered the involvement of international NGOs to establish bases in the region and work to develop bonobo conservation programs. This initiative should improve the likelihood of bonobo survival, but its success still may depend upon building greater involvement and capability in local and indigenous communities.
The Congo is setting aside more than 11,000 square miles (28,000 km2) of rainforest to help protect the endangered bonobo, in this central African country. U.S. agencies, conservation groups, and the Congolese government have come together to set aside 11,803 square miles (30,570 km2) of tropical rainforest, the U.S.-based Bonobo Conservation Initiative. The area amounts to just over 1% of the vast Congo – but that means a park larger than the state of Massachusetts.
The bonobo population is believed to have declined sharply in the last 30 years, though surveys have been hard to carry out in war-ravaged central Congo. Estimates range from 60,000 to fewer than 50,000 living, according to the World Wildlife Fund.
In addition, concerned parties have addressed the crisis on several science and ecological websites. Organizations such as the World Wide Fund for Nature, the African Wildlife Foundation, and others, are trying to focus attention on the extreme risk to the species. Some have suggested that a reserve be established in a more stable part of Africa, or on an island in a place such as Indonesia. Awareness is ever increasing, and even nonscientific or ecological sites have created various groups to collect donations to help with the conservation of this species.
- Basankusu, DR Congo – base for bonobo research and conservation
- Chimpanzee genome project
- Claudine André
- Great ape personhood
- Great Ape Project
- Kanzi – a captive bonobo who uses language
- List of apes – notable individual apes
- Lola ya Bonobo
- Gynecocracy, among people, 'women's government over women and men' or 'women's social supremacy'
- Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M, eds. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. p. 183. OCLC 62265494. ISBN 0-801-88221-4.
- Fruth, B., Benishay, J.M., Bila-Isia, I., Coxe, S., Dupain, J., Furuichi, T., Hart, J., Hart, T., Hashimoto, C., Hohmann, G., Hurley, M., Ilambu, O., Mulavwa, M., Ndunda, M., Omasombo, V., Reinartz, G., Scherlis, J., Steel, L. & Thompson, J. (2008). Pan paniscus. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 4 January 2009.
- de Waal, Frans; Lanting, Frans (1997). Bonobo: The Forgotten Ape. University of California Press. ISBN 0-520-20535-9.
- Rowe, N. (1996) Pictural Guide to the Living Primates, Pogonias Press, East Hampton, ISBN 0-9648825-1-5.
- Williams, A.; Myers, P. (2004). "Pan paniscus". Animal Diversity Web. Retrieved 6 January 2012.
- Savage-Rumbaugh, Sue; Lewin, Roger (1994). Kanzi: the ape at the brink of the human mind. John Wiley & Sons. p. 97. ISBN 0-385-40332-1.
- de Waal, Frans (2005). Our Inner Ape. Riverhead Books. ISBN 1-57322-312-3.
- McBrearty, Sally; Jablonski, Nina G. (1 September 2005). "First fossil chimpanzee". Nature 437 (7055): 105–8. Bibcode:2005Natur.437..105M. doi:10.1038/nature04008. PMID 16136135.
- Zihlman AL; Cronin JE; Cramer DL; Sarich VM (1978). "Pygmy chimpanzee as a possible prototype for the common ancestor of humans, chimpanzees and gorillas". Nature 275 (5682): 744–6. Bibcode:1978Natur.275..744Z. doi:10.1038/275744a0. PMID 703839.
- Griffith, Jeremy (2013). Freedom Book 1. Part 8:4G. WTM Publishing & Communications. ISBN 978-1-74129-011-0.
- Schwarz, Ernst (April 1, 1929). "Das Vorkommen des Schimpansen auf den linken Kongo-Ufer". Revue de zoologie et de botanique africaines 16: 425–426.
- Coolidge, Harold Jefferson Jr. (July–September 1933). "Pan paniscus. Pigmy chimpanzee from south of the Congo river". American Journal of Physical Anthropology 18 (1): 1–59. doi:10.1002/ajpa.1330180113. Coolidge's paper contains a translation of Schwarz's earlier report.
- Herzfeld, Chris (2007). "L'invention du bonobo". Bulletin d'histoire et d'épistémologie des sciences de la vie (in French) 14 (2): 139–162. Retrieved 21 December 2011.
- Savage-Rumbaugh, Sue (21 June 2010). "Bonobos have a secret". New Scientist. Retrieved 21 December 2011.
- de Waal, Frans B. M. (2002). Tree of Origin: What Primate Behavior Can Tell Us About Human Social Evolution. Harvard University Press. p. 51. ISBN 0674010043.
- Prüfer, Kay et al. (2012) "The bonobo genome compared with the chimpanzee and human genomes". Nature doi:10.1038/nature11128 PMID 22722832
- Karow, Julia (2008-05-13). "Neandertal, bonobo genomes may shed light on human evolution; MPI, 454 preparing drafts". In Sequence. Genome Web. Retrieved 2011-12-08.
- "Colombus Zoo: Bonobo". Archived from the original on 2006-05-02. Retrieved 2006-08-01.
- Takahata, N.; Satta, Y.; Klein, J. (1995). "Divergence time and population size in the lineage leading to modern humans". Theoretical Population Biology 48 (2): 198–221. doi:10.1006/tpbi.1995.1026. PMID 7482371.
- The Chimpanzee Sequencing and Analysis Consortium (1 September 2005). "Initial sequence of the chimpanzee genome and comparison with the human genome". Nature 437 (7055): 69–87. Bibcode:2005Natur.437...69.. doi:10.1038/nature04072. PMID 16136131.
- Hecht, Jeff (19 May 2003). "Chimps are human, gene study implies". New Scientist. Retrieved 2011-12-08.
- Won, Yong-Jin; Hey, Jody (13 October 2004). "Divergence population genetics of chimpanzees". Molecular Biology & Evolution 22 (2): 297–307. doi:10.1093/molbev/msi017. PMID 15483319.
- Fischer, Anne; Wiebe, Victor; Pääbo, Svante; Przeworski, Molly (12 February 2004). "Evidence for a complex demographic history of chimpanzees". Molecular Biology & Evolution 21 (5): 799–808. doi:10.1093/molbev/msh083. PMID 14963091.
- Scholz, M. N.; d'Aout, K.; Bobbert, M. F.; Aerts, P. (2006). "Vertical jumping performance of bonobo (Pan paniscus) suggests superior muscle properties". Proceedings of the Royal Society B: Biological Sciences 273 (1598): 2177–2184. doi:10.1098/rspb.2006.3568. PMC 1635523. PMID 16901837.
- "Bonobo videos, photos and facts – Pan paniscus". ARKive. Retrieved 2012-08-15.
- Burnie D and Wilson DE (Eds.) (2005) Animal: The Definitive Visual Guide to the World's Wildlife. DK Adult. ISBN 0789477645
- Novak, R. M. (1999). Walker's Mammals of the World. 6th edition. Johns Hopkins University Press, Baltimore, ISBN 0-8018-5789-9
- Shumaker, Robert W.; Walkup, Kristina R. and Beck, Benjamin B. (2011) Animal Tool Behavior: The Use and Manufacture of Tools by Animals, JHU Press, ISBN 1421401282.
- Doran, D. M. (1993). "Comparative locomotor behavior of chimpanzees and bonobos: the influence of morphology on locomotion". Am J Phys Anthropol. 91 (1): 83–98. doi:10.1002/ajpa.1330910106. PMID 8512056.
- D’Août, K.; Vereecke, E.; Schoonaert, K.; De Clercq, D.; Van Elsacker, L.; Aerts, P. (2004). "Locomotion in bonobos (Pan paniscus): differences and similarities between bipedal and quadrupedal terrestrial walking, and a comparison with other locomotor modes". Journal of Anatomy 204 (5): 353–361. doi:10.1111/j.0021-8782.2004.00292.x. PMC 1571309. PMID 15198700.
- Wilson Wiessner, Pauline and Schiefenhövel, Wulf (1996). Food and the Status Quest: An Interdisciplinary Perspective. Berghahn Books. ISBN 1571818715. p. 50. "...twenty-two mature community members (eight males, fourteen females)could be identified using facial features...
- Shea, B.T. (1983). "Paedomorphosis and neoteny in the pygmy chimpanzee". Science 222 (4623): 521–2. Bibcode:1983Sci...222..521S. doi:10.1126/science.6623093. PMID 6623093.
- Godfrey L; Sutherland M. (1996). "Paradox of peramophic paedomorphosis: heterochrony and human evolution". Am J Phys Anthropol 99 (1): 17–42. doi:10.1002/ajpa.1330990102. PMID 8928718.
- de Waal, Frans (2013). The Bonobo and the Atheist: In Search of Humanism Among the Primates (1st ed.). W. W. Norton. p. 78. ISBN 978-0-393-07377-5.
- Furuichi, T (2011). "Female contributions to the peaceful nature of bonobo society". Evolutionary Anthropology: Issues, News, and Reviews 20 (4): 131–42. doi:10.1002/evan.20308. PMID 22038769.
- Stanford, C. B. (1998). "The Social Behavior of Chimpanzees and Bonobos: Empirical Evidence and Shifting Assumptions". Current Anthropology 39 (4): 399. doi:10.1086/204757.
- Wilson, M. L.; Boesch, C.; Fruth, B.; Furuichi, T.; Gilby, I. C.; Hashimoto, C.; Hobaiter, C. L.; Hohmann, G.; Itoh, N.; Koops, K.; Lloyd, J. N.; Matsuzawa, T.; Mitani, J. C.; Mjungu, D. C.; Morgan, D.; Muller, M. N.; Mundry, R.; Nakamura, M.; Pruetz, J.; Pusey, A. E.; Riedel, J.; Sanz, C.; Schel, A. M.; Simmons, N.; Waller, M.; Watts, D. P.; White, F.; Wittig, R. M.; Zuberbühler, K.; Wrangham, R. W. (2014). "Lethal aggression in Pan is better explained by adaptive strategies than human impacts". Nature 513 (7518): 414. Bibcode:2014Natur.513..414W. doi:10.1038/nature13727. PMID 25230664.
- White F. (1996) "Comparative socio-ecology of Pan paniscus", pp. 29–41 in: McGrew WC, Marchant LF, Nishida T (eds.) Great ape societies. Cambridge, England: Cambridge Univ Press, ISBN 0521555361.
- Cawthon Lang KA. (December 2010) Primate Factsheets: Bonobo (Pan paniscus) behavior. University of Wisconsin
- Aggression topics. University of New Hampshire
- Nguyen, Tuan C. (2008-02-13). "Gorillas Caught in Very Human Act". Live Science
- Manson, J.H.; Perry, S.; Parish, A.R. (1997). "Nonconceptive Sexual Behavior in Bonobos and Capuchins". International Journal of Primatology 18 (5): 767–86. doi:10.1023/A:1026395829818.
- de Waal, Frans B. M. (March 1995). "Bonobo Sex and Society" (PDF). Scientific American 272 (3): 58–64. Bibcode:1995SciAm.272c..82W. doi:10.1038/scientificamerican0395-82. PMID 7871411. Retrieved 21 December 2011.
- Block, Susan (2014) The Bonobo Way: The Evolution of Peace Through Pleasure Paperback. Gardner & Daughters Publishers. ISBN 0692323767
- Balcombe, Jonathan Peter (2011). The Exultant Ark: A Pictorial Tour of Animal Pleasure. University of California Press. p. 88. ISBN 978-0-520-26024-5. Retrieved 2012-11-22.
- Angier, Natalie (1999). Woman: An Intimate Geography. Houghton Mifflin Harcourt. p. 68. ISBN 978-0-395-69130-4. Retrieved 2012-11-22.
- Paoli, T.; Palagi, E.; Tacconi, G.; Tarli, S. B. (2006). "Perineal swelling, intermenstrual cycle, and female sexual behavior in bonobos (Pan paniscus)". American Journal of Primatology 68 (4): 333–347. doi:10.1002/ajp.20228. PMID 16534808.
- Hohmann, G.; Fruth, B. (2000). "Use and function of genital contacts among female bonobos". Animal Behaviour 60 (1): 107–120. doi:10.1006/anbe.2000.1451.
- "Courtney Laird, "Social Organization"". Bio.davidson.edu. 2004. Archived from the original on 2011-05-19. Retrieved 2009-07-03.
- Frans B. M. de Waal (2001). "Bonobos and Fig Leaves". The ape and the sushi master : cultural reflections by a primatologist. Basic Books. ISBN 84-493-1325-2.
- de Waal, F. B. M. (1990). "Sociosexual behavior used for tension regulation in all age and sex combinations among bonobos", pp. 378–393 in Pedophilia: Biosocial Dimensions, J. R. Feierman (ed.). Springer, New York. ISBN 9781461396840
- Casual Sex Play Common Among Bonobos. Discover, June 1992
- Paul M. Sharp; George M. Shaw; Beatrice H. Hahn (2005). "Simian Immunodeficiency Virus Infection of Chimpanzees". Journal of Virology 79 (7): 3891–3902. doi:10.1128/jvi.79.7.3891-3902.2005. PMID 15767392.
- Cawthorn Lang, Kristina (2011). "Bonobo: Pan paniscus". National Primate Research Center, University of Wisconsin – Madison.
- Ihobe H (1992). "Observations on the meat-eating behavior of wild bonobos (Pan paniscus) at Wamba, Republic of Zaire". Primates (journal) 33 (2): 247–250. doi:10.1007/BF02382754.
- Rafert, J. and E.O. Vineberg (1997). "Bonobo Nutrition – relation of captive diet to wild diet," Bonobo Husbandry Manual, American Association of Zoos and Aquariums
- Surbeck M; Fowler A; Deimel C; Hohmann G (2008). "Evidence for the consumption of arboreal, diurnal primates by bonobos (Pan paniscus)". American Journal of Primatology 71 (2): 171–4. doi:10.1002/ajp.20634. PMID 19058132.; Surbeck M; Hohmann G (2008). "Primate hunting by bonobos at LuiKotale, Salonga National Park". Current Biology 18 (19): R906–7. doi:10.1016/j.cub.2008.08.040. PMID 18957233.
- Parker, Ian (2007-07-30). "Swingers". Our Far-Flung Correspondents. The New Yorker. Retrieved 2011-12-08.
- de Waal, Frans (2009-10-18). "Was "Ardi" a Liberal?". The Huffington Post. Retrieved 2009-10-18.
- Walker, Matt (2010-02-01). "Wild bonobo mother ape eats own infant in DR Congo". BBC News.
- Hippy apes caught cannibalising their young. New Scientist (1 February 2010). Retrieved on 2013-04-18.
- "Chimpanzee behavior: Killer instincts". The Economist. June 24, 2010. Retrieved 2011-12-08.
- Raffaele, Paul (November 2006) The Smart and Swinging Bonobo. Smithsonian Magazine.
- Caswell JL; Mallick S; Richter DJ (2008). McVean, Gil, ed. "Analysis of Chimpanzee History Based on Genome Sequence Alignments". PLoS Genet. 4 (4): e1000057. doi:10.1371/journal.pgen.1000057. PMC 2278377. PMID 18421364.
- White, Frances J.; Wrangham, Richard W. (1988). "Feeding competition and patch size in the chimpanzee species Pan paniscus and Pan troglodytes". Behaviour 105: 148–164. doi:10.1163/156853988X00494. JSTOR 4534684.
- Brain differences may explain varying behavior of bonobos and chimpanzees. Washingtonpost.com (2011-04-12). Retrieved on 2012-12-26.
- de Waal, Frans (August 8, 2007). "Bonobos, Left & Right". Skeptic.
- Best, Steven (Spring 2009). "Minding the Animals: Ethology and the Obsolescence of Left Humanism". The International Journal of Inclusive Democracy 5 (2). Retrieved 2012-02-08.
- "Meet our Great Apes: Kanzi". Archived from the original on 2008-06-30. Retrieved 2008-09-28.
- Raffaele, P. (2006). "Speaking Bonobo". Smithsonian. Retrieved 2008-09-28.
- Schick, Kathy; Toth, Nicholas; Garufi, Gary; Savage-Rumbaugh, E.Sue; Rumbaugh, Duane; Sevcik, Rose (1999). "Continuing Investigations into the Stone Tool-making and Tool-using Capabilities of a Bonobo (Pan paniscus)". Journal of Archaeological Science 26 (7): 821. doi:10.1006/jasc.1998.0350.
- Palagi E; Paoli; Tarli (2004). "Reconciliation and consolation in captive bonobos (Pan paniscus)". Am J Primatol 62 (1): 15–30. doi:10.1002/ajp.20000. PMID 14752810.
- Palagi E; Norscia I (2013). "Bonobos Protect and Console Friends and Kin". PLoS ONE 8 (11): e79290. Bibcode:2013PLoSO...879290P. doi:10.1371/journal.pone.0079290. PMID 24223924.
- Beale, B. (2003). "Where Did Laughter Come From?". ABC Science Online. Retrieved 2008-08-10.
- Dawkins, Richard (2004). "Chimpanzees". The Ancestor's Tale. Houghton Mifflin. ISBN 1-155-16265-X.
- Quammen, David. "The Left Bank Ape". The New Age of Exploration, 2013. National Geographic. Retrieved 28 February 2013.
- Reid, John (2006-06-15). Parks and people, not parks vs. people, San Francisco Chronicle.
- Bonobo and large mammal survey. Zoological Society of Milwaukee.
- Hart, Therese (2012-07-27) Searching for Bonobo in Congo. Bonoboincongo.com. Retrieved on 2012-12-26.
- Lola Ya Bonobo (Bonobo Heaven). Lolayabonobo.wildlifedirect.org. Retrieved on 2012-12-26.
- Bonobo Reintroduction in the Democratic Republic of Congo. friendsofbonobos.org (November 2009) . Retrieved on 2012-12-26.
- "Bonobo and Congo Biodiversity Initiative". Zoological Society of Milwaukee.
- The Make Love, Not War Species, Living on Earth, (July 2006), National Public Radio
- Chapin, Mac, (November/December 2004), Vision for a Sustainable World, WORLDWATCH magazine
- de Waal, Frans, and Frans Lanting, Bonobo: The Forgotten Ape, University of California Press, 1997. ISBN 0-520-20535-9; ISBN 0-520-21651-2 (trade paperback)
- Kano, Takayoshi, The Last Ape: Pygmy Chimpanzee Behavior and Ecology, Stanford, CA: Stanford University Press, 1992.
- Savage-Rumbaugh, Sue, and Roger Lewin, Kanzi: The Ape at the Brink of the Human Mind, John Wiley, 1994. ISBN 0-471-58591-2; ISBN 0-471-15959-X (trade paperback)
- Woods, Vanessa, Bonobo Handshake, Gotham Books, 2010. ISBN 978-1-59240-546-6
- Sandin, Jo, Bonobos: Encounters in Empathy, Zoological Society of Milwaukee & The Foundation for Wildlife Conservation, Inc., 2007. ISBN 978-0-9794151-0-4
- de Waal, Frans, The Bonobo and the Atheist, Norton, 2013. ISBN 978-0393073775
- de Waal, Frans, "Bonobo: Sex & Society", Scientific American, 1995
- DeBartolo, Anthony. "The Bonobo: 'Newest' apes are teaching us about ourselves", Chicago Tribune June 11, 1998.
- Schweller, Ken, "Apes With Apps," IEEE Spectrum Magazine, July 2012.
- Madrigal, Alexis "Brian the Mentally Ill Bonobo, and How He Healed", The Atlantic, June 11, 2014.
- Parker, Ian "Swingers", The New Yorker, July 30, 2007.
- Fischer, Anne; Prüfer, Kay; Good, Jeffrey M.; Halbwax, Michel; Wiebe, Victor; André, Claudine; Atencia, Rebeca; Mugisha, Lawrence; Ptak, Susan E. et al. (29 June 2011). Joly, Etienne, ed. "Bonobos fall within the genomic variation of chimpanzees". PLoS ONE 6 (6): e21605. Bibcode:2011PLoSO...621605F. doi:10.1371/journal.pone.0021605.
- Zsurka, Gábor; Kudina, Tatiana; Peeva, Viktoriya; Hallmann, Kerstin; Elger, Christian E.; Elger, Konstantin; Khrapko, Konstantin; Kunz, Wolfram S. (2010). "Distinct patterns of mitochondrial genome diversity in bonobos (Pan paniscus) and humans". BMC Evolutionary Biology 10: 270. doi:10.1186/1471-2148-10-270.
- Wildman, Derek E.; Uddin, Monica; Liu, Guozhen; Grossman, Lawrence I.; Goodman, Morris (10 June 2003). "Implications of natural selection in shaping 99.4% nonsynonymous DNA identity between humans and chimpanzees: Enlarging genus Homo". PNAS 100 (12): 7181–7188. Bibcode:2003PNAS..100.7181W. doi:10.1073/pnas.1232172100. PMC 165850. PMID 12766228.
To request an improvement, please leave a comment on the page. Thank you!