|Male at Apenheul Primate Park|
The bonobo (/ /,; Pan paniscus), also historically called the pygmy chimpanzee and, less often, the dwarf or gracile chimpanzee, is an endangered great ape. It is one of the two species making up the genus Pan, the other being the common chimpanzee (Pan troglodytes). Although bonobos are not a subspecies of chimpanzee (Pan troglodytes), but rather a distinct species in their own right, both species are sometimes referred to collectively using the generalized term chimpanzees, or chimps. Taxonomically, the members of the chimpanzee/bonobo subtribe Panina (composed entirely by the genus Pan) are collectively termed panins.
The bonobo is distinguished by relatively long legs, pink lips, dark face, 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 frugivorous and inhabits primary and secondary forests, including seasonally inundated swamp forests. Because of political instability in the region and the timidity of bonobos, 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. As 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 are 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. Bonobos typically live 40 years in captivity; their lifespan in the wild is unknown, but it is almost certainly much shorter.
Formerly the bonobo was known as the "pygmy chimpanzee", despite the bonobo having a similar body size to the common chimpanzee. The name "pygmy" was given by the German zoologist Ernst Schwarz in 1929, who classified the species on the basis of a previously mislabeled bonobo cranium, noting its diminutive size compared to chimpanzee skulls.
The name "bonobo" first appeared in 1954, when Austrian zoologist Eduard Paul Tratz and German biologist Heinz Heck proposed it as a new and separate generic term for pygmy chimpanzees. The name is thought to derive from a misspelling on a shipping crate from the town of Bolobo on the Congo River near the location from which the first bonobo specimens were collected in the 1920s.
The bonobo was first recognised as a distinct taxon in 1928 by German anatomist Ernst Schwarz, based on a skull in the Tervuren Museum in Belgium which had previously been classified as a juvenile chimpanzee (Pan troglodytes). Schwarz published his findings in 1929, classifying the bonobo as a subspecies of chimpanzee, Pan satyrus paniscus. In 1933, American anatomist Harold Coolidge elevated it to species status. Major behavioural differences between bonobos and chimpanzees were first discussed in detail by Tratz and Heck in the early 1950s. American psychologist and primatologist Robert Yerkes was also one of the first to notice major behavioural differences.
The exact timing of the Pan–Homo last common ancestor is contentious, but DNA comparison suggests continual interbreeding between ancestral Pan and Homo groups, post-divergence, until about 4 million years ago. DNA evidence suggests the bonobo and common chimpanzee species diverged approximately 890,000–860,000 years ago due to separation of these two populations possibly due to acidification and the spread of savannas at this time. Currently, these two species are separated by the Congo River, which had existed well before the divergence date, though ancestral Pan may have dispersed across the river using corridors which no longer exist. The first Pan fossils were reported in 2005 from the Middle Pleistocene (after the bonobo–chimp split) of Kenya, alongside early Homo fossils.
According to A. Zihlman, bonobo body proportions closely resemble those of Australopithecus, leading evolutionary biologist Jeremy Griffith to suggest that bonobos may be a living example of our distant human ancestors. According to Australian anthropologists Gary Clark and Maciej Henneberg, human ancestors went through a bonobo-like phase featuring reduced aggression and associated anatomical changes, exemplified in Ardipithecus ramidus.
The first official publication of the sequencing and assembly of the bonobo genome was released in June 2012. The genome of a female bonobo from Leipzig Zoo 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.
Genetics and genomics
|NCBI genome ID|
|Genome size||2,869.21 Mb|
|Number of chromosomes||24 pairs|
|Year of completion||2012, 2021|
Relationships of bonobos to humans and other apes can be determined by comparing their genes or whole genomes. While the first bonobo genome was published in 2012, a high-quality reference genome became only available in 2021. The overall nucleotide divergence between chimpanzee and bonobo based on the latter is 0.421 ± 0.086% for autosomes and 0.311 ± 0.060% for the X chromosome. The reference genome predicts 22,366 full-length protein-coding genes and 9,066 noncoding genes, although cDNA sequencing confirmed only 20,478 protein-coding and 36,880 noncoding bonobo genes, similar to the number of genes annotated in the human genome. Overall, 206 and 1,576 protein-coding genes are part of gene families that contracted or expanded in the bonobo genome compared to the human genome, respectively, that is, these genes were lost or gained in the bonobo genome compared to humans.
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 broadly overlap in body size. Adult female bonobos are somewhat smaller than adult males. Body mass ranges from 34 to 60 kg (75 to 132 lb) with an average weight of 45 kilograms (99 lb) in males against an average of 33 kg (73 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). Male bonobos average 119 cm (3.90 ft) when standing upright, compared to 111 centimetres (3.64 ft) in females. 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 parting. 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 the common chimpanzee does. 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 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. Bonobos warn each other of danger less efficiently than chimpanzees in the same situation.
Bonobos are unique among nonhuman apes for a lack of male dominance and relatively high social status of females, due to the latter forming long-lasting, powerful alliances among each other. Although a male bonobo is dominant to a female in a dyadic interaction, depending on the community, socially-bonded females may be co-dominant with males or dominant over them. At the top of the hierarchy is a coalition of high-ranking females and males typically headed by an old, experienced matriarch who acts as the decision-maker and leader of the group. Female bonobos typically earn their rank through age, rather than physical intimidation, and top-ranking females will protect immigrant females from male harassment. While bonobos are often called matriarchal, in many communities there is an alpha male who will associate closely with the high-ranking females in the group. He is co-dominant or at least the second most dominant group member after the alpha female, and helps her in curbing the aggressive impulses of adolescent males, has priority access to feeding sites alongside the females, and resident females may show preference for him as a mate. He alerts the group to any possible threats, protects the group from predators such as pythons and leopards, and in some cases, plays a decisive role in deciding where the group travels to, and where they feed. While female bonobos initiate group travels in most cases, males do appear to play a role in chasing off predators which threaten the group. 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, and although the son of a high ranking female may outrank a lower female, rank plays a less prominent role than in other primate societies. Relationships between different communities are often positive and affiliative, and bonobos are not a territorial species. Bonobos will also share food with others, even unrelated strangers. Bonobos exhibit paedomorphism (retaining infantile physical characteristics and behaviours), which greatly inhibits aggression and enables unfamiliar bonobos to freely mingle and cooperate with each other.
Males engage in lengthy friendships with females and, in turn, female bonobos prefer to associate with and mate with males who are respectful and easygoing around them. Because female bonobos can use alliances to rebuff coercive and domineering males and select males at their own leisure, they show preference for males who are not aggressive towards them.
Aging bonobos lose their playful streak and become noticeably more irritable in old age. Both sexes have a similar level of aggressiveness.
Bonobos live in a male philopatric society where the females immigrate to new communities while males remain in their natal troop. However, it is not entirely unheard of for males to occasionally transfer into new groups.
Alliances between males are poorly developed in most bonobo communities, while females will form alliances with each other and alliances between males and females occur, including multisex hunting parties. There is a confirmed case of a grown male bonobo adopting his orphaned infant brother.
A mother bonobo will also support her grown son in conflicts with other males and help him secure better ties with other females, enhancing her chance of gaining grandchildren from him. She will even take measures such as physical intervention to prevent other males from breeding with certain females she wants her son to mate with. Although mothers play a role in aiding their sons, and the hierarchy among males is largely reflected by their mother's social status, some motherless males will still successfully dominate some males who do have mothers.
Bonobos are not known to kill each other, and are generally less violent than chimpanzees, yet aggression still manifests itself in this species. Although males are unable to dominate females, with more chivalrous males enjoying more success in attaining high rank and fathering large amounts of young, rank among males is often violently enforced and the alpha status heavily coveted. Male bonobos are known to attack each other and inflict serious injuries such as missing digits, damaged eyes and torn ears. Some of these injuries may also occur when a male threatens the high ranking females and is injured by them, as the larger male is swarmed and outnumbered by these female mobs.
Due to the promiscuous mating behavior of female bonobos, a male cannot be sure which offspring are his. As a result, the entirety of parental care in bonobos is assumed by the mothers. However, bonobos are not as promiscuous as chimpanzees and slightly polygamous tendencies occur, with high-ranking males enjoying greater reproductive success than low-ranking males. Unlike chimpanzees, where any male can coerce a female into mating with him, female bonobos enjoy greater sexual preferences and can rebuff undesirable males, an advantage of female-female bonding, and actively seek out higher-ranking males.
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.
Female bonobos more often than not secure feeding privileges and feed before males do, although they are rarely successful in one-on-one confrontations with males, a female bonobo with several allies supporting her has extremely high success in monopolizing food sources. Different communities favour different prey. In some communities females exclusively hunt and have a preference for rodents, in others both sexes hunt, and will target monkeys.
In captive settings, females exhibit extreme food-based aggression towards males, and forge coalitions against them to monopolize specific food items, often going as far as to mutilate any males who fail to heed their warning.
In wild settings, however, female bonobos will quietly ask males for food if they had gotten it first, instead of forcibly confiscating it, suggesting sex-based hierarchy roles are less rigid than in captive colonies.
Female bonobos are known to lead hunts on duikers and successfully defend their bounty from marauding males in the wild. They are more tolerant of younger males pestering them yet exhibit heightened aggression towards older males.
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 tongue kissing. Bonobos and humans are the only primates to typically engage in face-to-face genital sex, although a pair of western gorillas has been photographed in this position.
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.
More often than the males, female bonobos engage in mutual genital-rubbing 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. 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 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 analogue of tribadism, engaged in by some 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. As 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". 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 engage in various forms of male–male genital behavior. The most common form of male–male mounting is similar to that of a heterosexual mounting: one of the males sits "passively on his back [with] the other male thrusting on him", with the penises rubbing together due to both males' erections. In another, rarer form of genital rubbing, which is the non-human analogue of frotting, engaged in by some human males, 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) often occurs to express reconciliation between two males after a conflict, when they stand back-to-back and rub their scrotal sacs together, but such behavior also occurs outside agonistic contexts: Kitamura (1989) observed rump–rump contacts between adult males following sexual solicitation behaviors similar to those between female bonobos prior to GG-rubbing. Takayoshi Kano observed similar practices among bonobos in the natural habitat. Tongue kissing, oral sex, and genital massaging have also been recorded among male bonobos.
Bonobo reproductive rates are no higher than those of the common chimpanzee. However, female bonobo oestrus periods are longer. During oestrus, females undergo a swelling of the perineal tissue lasting 10 to 20 days. 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. Mothers will help their sons get more matings from females in oestrus. Adult male bonobos have sex with infants, although without penetration.
Infanticide, while well documented in chimpanzees, is apparently absent in bonobo society. The highly sexual nature of bonobo society and the fact that there is little competition over mates means that many males and females are mating with each other, in contrast to the one dominant male chimpanzee that fathers most of the offspring in a group. The strategy of bonobo females mating with many males may be a counterstrategy to infanticide because it confuses paternity. If male bonobos cannot distinguish their own offspring from others, the incentive for infanticide essentially disappears. This is a reproductive strategy that seems specific to bonobos; infanticide is observed in all other great apes except orangutans.
Observations in the wild indicate that the males among the related common chimpanzee communities are 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.
While bonobos are more peaceful than chimpanzees, it is not true that they are unaggressive. In the wild, among males, bonobos are half as aggressive as chimpanzees, while female bonobos are more aggressive than female chimpanzees. Both bonobos and chimpanzees exhibit physical aggression more than 100 times as often as humans do.
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. Bonobos have more grey matter volume in the right anterior insula, right dorsal amygdala, hypothalamus, and right dorsomedial prefrontal cortex, all of which are regions assumed to be vital for feeling empathy, sensing distress in others and feeling anxiety. They also have a thick connection between the amygdala, an important area that can spark aggression, and the ventral anterior cingulate cortex, which has been shown to help control impulses in humans. This thicker connection may make them better at 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." Although male bonobos have not yet been seen to practice infanticide, there is documented incident in captivity involving a dominant female abducting an infant from a lower-ranking female, treating the infant roughly and denying it the chance to suckle. During the kidnapping, the infant's mother was clearly distressed and tried to retrieve her infant. Had the zookeepers not intervened, the infant almost certainly would have died from dehydration. This suggests female bonobos can have hostile rivalries with each other and a propensity to carry out infanticide.
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.
The bonobo is an omnivorous frugivore; 57% of its diet is fruit, but this is supplemented 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 dead infant was described in 2008. A 2016 paper reported two more instances of infant cannibalism, although it was not confirmed if infanticide was involved.
Cognitive comparisons to chimpanzees
In 2020 the first whole-genome comparison between chimpanzees and bonobos was published and shows genomic aspects that may underlie or have resulted from their divergence and behavioral differences, including selection for genes related to diet and hormones. A 2010 study found that "female bonobos displayed a larger range of tool use behaviours than males, a pattern previously described for chimpanzees but not for other great apes". This finding was affirmed by the results of another 2010 study which also found that "bonobos were more skilled at solving tasks related to theory of mind or an understanding of social causality, while chimpanzees were more skilled at tasks requiring the use of tools and an understanding of physical causality".
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. The communication system of wild bonobos includes a characteristic that was earlier only known in humans: bonobos use the same call to mean different things in different situations, and the other bonobos have to take the context into account when determining the meaning. 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 which humans theoretically accord to all persons (See great ape personhood). In the 1990s, Kanzi was taught to make and use simple stone tools. This resulted from a study undertaken by researchers Kathy Schick and Nicholas Toth, and later Gary Garufi. The researchers wanted to know if Kanzi possessed the cognitive and biomechanical abilities required to make and use stone tools. Though Kanzi was able to form flakes, he did not create them in same way as humans, who hold the core in one hand and knap 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.
Distribution and habitat
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. 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.
In the Congo tropical rainforest, the very great majority of plants need animals to reproduce and disperse their seeds. Bonobos are the second largest frugivorous animals in this region, after elephants. During its life, each bonobo will ingest and disperse nine tons of seeds, from more than 91 species of lianas, grass, trees and shrubs. These seeds will travel 24 hours in the bonobo digestive tract, which will transfer them over several kilometers (mean 1.3 km; max: 4.5 km), far from their parents, where they will be deposited intact in their feces. These dispersed seeds remain viable, germinate better and more quickly than unpassed seeds. For those seeds, diplochory with dung-beetles (Scarabaeidae) improves post-dispersal survival. Certain plants such as Dialium may even be dependent on bonobos to activate the germination of their seeds, characterized by tegumentary dormancy. The first parameters of the effectiveness of seed dispersal by bonobos are present. Behavior of the bonobo could affect the population structure of plants whose seeds they disperse. The majority of these zoochorous plants cannot recruit without dispersal and the homogeneous spatial structure of the trees suggests a direct link with their dispersal agent. Few species could replace bonobos in terms of seed dispersal services, just as bonobos could not replace elephants. There is little functional redundancy between frugivorous mammals of the Congo, which face severe human hunting pressures and local extinction. The defaunation of the forests, leading to the empty forest syndrome, is critical in conservation biology. The disappearance of the bonobos, which disperse seeds of 40% of the tree species in these forests, or 11.6 million individual seeds during the life of each bonobo, will have consequences for the conservation of the Congo rainforest.
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 still rely on local and community involvement. The issue of parks versus people is salient in the Cuvette Centrale, in 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 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.
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. This model, implemented mainly through DRC organizations and local communities, has helped bring about agreements to protect over 50,000 square miles (130,000 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".
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.
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 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
- Bonobo Conservation Initiative
- Chimpanzee genome project
- Claudine André
- Great ape personhood
- Great Ape Project
- List of apes – notable individual nonhuman apes
- Lola ya Bonobo
- Koba, a fictional bonobo and antagonist of the Planet of the Apes reboot series
- Gynecocracy, among people, 'women's government over women and men' or 'women's social supremacy'
- Groves CP (2005). Wilson DE, Reeder DM (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. p. 183. ISBN 0-801-88221-4. OCLC 62265494.
- Fruth B, Hickey JR, André C, Furuichi T, Hart J, Hart T, Kuehl H, Maisels F, Nackoney J, Reinartz G, Sop T, Thompson J, Williamson EA (2016) [errata version of 2016 assessment]. "Pan paniscus". IUCN Red List of Threatened Species. 2016: e.T15932A102331567.
- de Waal F, Lanting F (1997). Bonobo: The Forgotten Ape. University of California Press. ISBN 978-0-520-20535-2.
- Angier N (September 10, 2016). "Beware the Bonds of Female Bonobos". The New York Times. Retrieved September 10, 2016.
- Muehlenbein MP (2015). Basics in Human Evolution. Elsevier Science. pp. 114–115. ISBN 9780128026526.
- Diogo R, Molnar JL, Wood B (April 2017). "Bonobo anatomy reveals stasis and mosaicism in chimpanzee evolution, and supports bonobos as the most appropriate extant model for the common ancestor of chimpanzees and humans". Scientific Reports. 7 (1): 608. Bibcode:2017NatSR...7..608D. doi:10.1038/s41598-017-00548-3. PMC 5428693. PMID 28377592.
- Beaune, David (November 2012). "The Ecological Role of the Bonobo. Seed Dispersal Service in Congo Forests". Research Gate. Retrieved May 27, 2021.
- Rowe N (1996). Pictural Guide to the Living Primates. West Hampton: Pogonias Press. ISBN 0-9648825-1-5.
- Herzfeld C (2007). "L'invention du bonobo" (PDF). Bulletin d'Histoire et d'Épistémologie des Sciences de la Vie (in French). 14 (2): 139–162. doi:10.3917/bhesv.142.0139. Retrieved 21 December 2011.
- Savage-Rumbaugh S, Lewin R (1994). Kanzi: the ape at the brink of the human mind. John Wiley & Sons. p. 97. ISBN 978-0-385-40332-0.
- de Waal F (2005). Our Inner Ape. Riverhead Books. ISBN 978-1-57322-312-6.
- Schwarz E (April 1, 1929). "Das Vorkommen des Schimpansen auf den linken Kongo-Ufer" (PDF). Revue de Zoologie et de Botanique Africaines. 16: 425–426.
- Coolidge Jr HJ (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.
- de Waal FB (2002). Tree of Origin: What Primate Behavior Can Tell Us About Human Social Evolution. Harvard University Press. p. 51. ISBN 978-0674010048.
- Savage-Rumbaugh S (21 June 2010). "Bonobos have a secret". New Scientist. 206 (2765): 48. Bibcode:2010NewSc.206...48S. doi:10.1016/S0262-4079(10)61507-2. Retrieved 21 December 2011.
- Takahata N, Satta Y, Klein J (October 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.
- Waterson RH, Lander ES, Wilson RK, et al. (Chimpanzee Sequencing Analysis Consortium) (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.
- Patterson N, Richter DJ, Gnerre S, Lander ES, Reich D (June 2006). "Genetic evidence for complex speciation of humans and chimpanzees". Nature. 441 (7097): 1103–8. Bibcode:2006Natur.441.1103P. doi:10.1038/nature04789. PMID 16710306. S2CID 2325560.
- Won YJ, Hey J (February 2005). "Divergence population genetics of chimpanzees". Molecular Biology and Evolution. 22 (2): 297–307. doi:10.1093/molbev/msi017. PMID 15483319.
- McBrearty S, Jablonski NG (September 2005). "First fossil chimpanzee". Nature. 437 (7055): 105–8. Bibcode:2005Natur.437..105M. doi:10.1038/nature04008. PMID 16136135. S2CID 4423286.
- Zihlman AL, Cronin JE, Cramer DL, Sarich VM (October 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. S2CID 4252525.
- Griffith J (2013). Freedom Book 1. Part 8:4G. WTM Publishing & Communications. ISBN 978-1-74129-011-0.
- Clark G, Henneberg M (2015). "The life history of Ardipithecus ramidus: A heterochronic model of sexual and social maturation". Anthropological Review. 78 (2): 109–132. doi:10.1515/anre-2015-0009.
- Prüfer K, Munch K, Hellmann I, Akagi K, Miller JR, Walenz B, et al. (June 2012). "The bonobo genome compared with the chimpanzee and human genomes". Nature. 486 (7404): 527–31. Bibcode:2012Natur.486..527P. doi:10.1038/nature11128. PMC 3498939. PMID 22722832.
- Karow J (2008-05-13). "Neandertal, bonobo genomes may shed light on human evolution; MPI, 454 preparing drafts". In Sequence. Genome Web. Retrieved 2011-12-08.
- Prüfer, Kay; Munch, Kasper; Hellmann, Ines; Akagi, Keiko; Miller, Jason R.; Walenz, Brian; Koren, Sergey; Sutton, Granger; Kodira, Chinnappa; Winer, Roger; Knight, James R. (June 2012). "The bonobo genome compared with the chimpanzee and human genomes". Nature. 486 (7404): 527–531. Bibcode:2012Natur.486..527P. doi:10.1038/nature11128. ISSN 0028-0836. PMC 3498939. PMID 22722832.
- Mao, Yafei; Catacchio, Claudia R.; Hillier, LaDeana W.; Porubsky, David; Li, Ruiyang; Sulovari, Arvis; Fernandes, Jason D.; Montinaro, Francesco; Gordon, David S.; Storer, Jessica M.; Haukness, Marina (2021-05-05). "A high-quality bonobo genome refines the analysis of hominid evolution". Nature. 594 (7861): 77–81. Bibcode:2021Natur.594...77M. doi:10.1038/s41586-021-03519-x. ISSN 1476-4687. PMC 8172381. PMID 33953399.
- Kingdon, Jonathan (2013). Mammals of Africa: Volume II. Bloomsbury Publishing. p. 69.
- Scholz MN, D'Août K, Bobbert MF, Aerts P (September 2006). "Vertical jumping performance of bonobo (Pan paniscus) suggests superior muscle properties". Proceedings. Biological Sciences. 273 (1598): 2177–84. doi:10.1098/rspb.2006.3568. PMC 1635523. PMID 16901837.
- "Bonobo videos, photos and facts – Pan paniscus". ARKive. Archived from the original on 2012-08-25. Retrieved 2012-08-15.
- Burnie D, Wilson DE, eds. (2005). Animal: The Definitive Visual Guide to the World's Wildlife. DK Adult. ISBN 0789477645.
- Novak RM (1999). Walker's Mammals of the World (6th ed.). Baltimore: Johns Hopkins University Press. ISBN 0-8018-5789-9.
- Coolidge, Shea (1982). "External body dimensions of Pan paniscus and Pan troglodytes chimpanzees". Primates. 23 (2): 245–251. doi:10.1007/BF02381164. S2CID 27818900.
- Doran DM (May 1993). "Comparative locomotor behavior of chimpanzees and bonobos: the influence of morphology on locomotion". American Journal of Physical Anthropology. 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 (May 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): 353–61. doi:10.1111/j.0021-8782.2004.00292.x. PMC 1571309. PMID 15198700.
- Wiessner PW, Wiessner P, Schiefenhövel W (1996). Food and the status quest : an interdisciplinary perspective. Providence: Berghahn Books. p. 50. ISBN 1571818715.
...twenty-two mature community members (eight males, fourteen females)could be identified using facial features...
- Shea BT (November 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 LR, Sutherland MR (January 1996). "Paradox of peramorphic paedomorphosis: heterochrony and human evolution". American Journal of Physical Anthropology. 99 (1): 17–42. doi:10.1002/ajpa.1330990102. PMID 8928718.
- de Waal F (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. 20 (4): 131–42. doi:10.1002/evan.20308. PMID 22038769. S2CID 17830996.
- Stanford CB (1998). "The Social Behavior of Chimpanzees and Bonobos: Empirical Evidence and Shifting Assumptions". Current Anthropology. 39 (4): 399–420. doi:10.1086/204757. S2CID 8452514.
- Wilson ML, Boesch C, Fruth B, Furuichi T, Gilby IC, Hashimoto C, et al. (September 2014). "Lethal aggression in Pan is better explained by adaptive strategies than human impacts". Nature. 513 (7518): 414–7. Bibcode:2014Natur.513..414W. doi:10.1038/nature13727. PMID 25230664. S2CID 4449515.
- "In the wild, chimpanzees are more motivated to cooperate than bonobos". phys.org. Retrieved 2020-06-24.
- White, Frances J.; Wood, Kimberley D. (August 2007). "Female feeding priority in bonobos, Pan paniscus, and the question of female dominance". American Journal of Primatology. 69 (8): 837–850. doi:10.1002/ajp.20387. ISSN 0275-2565. PMID 17358018. S2CID 17628292.
- Paoli T, Palagi E, Tarli SM (May 2006). "Reevaluation of dominance hierarchy in bonobos (Pan paniscus)". American Journal of Physical Anthropology. 130 (1): 116–22. doi:10.1002/ajpa.20345. PMID 16353224.
- Surbeck M, Hohmann G (November 2013). "Intersexual dominance relationships and the influence of leverage on the outcome of conflicts in wild bonobos (Pan paniscus)". Behavioral Ecology and Sociobiology. 67 (11): 1767–80. doi:10.1007/s00265-013-1584-8. S2CID 15709567. Lay summary – ScienceDaily.
- Pallardy R (2012-05-21). "The Scandalous Social Lives of Bonobos". Saving Earth | Encyclopedia Britannica. Retrieved 2021-01-13.
- Sivasubramanian S. "Feminist bonobos are taking a stand against male aggression". Topics. Retrieved 2021-03-12.
- "Zoo Story". Milwaukee Magazine. 2007-08-13. Retrieved 2021-05-07.
- Raffaele P. "The Smart and Swinging Bonobo". Smithsonian Magazine. Retrieved 2020-09-07.
- Corredor-Ospina, Nicolas; Kreyer, Melodie; Rossi, Giulia; Hohmann, Gottfried; Fruth, Barbara (2021-07-01). "First report of a leopard (Panthera pardus)–bonobo (Pan paniscus) encounter at the LuiKotale study site, Democratic Republic of the Congo". Primates. 62 (4): 555–562. doi:10.1007/s10329-021-00897-8. ISSN 1610-7365. PMC 8225524. PMID 33950405.
- White F (1996). "Comparative socio-ecology of Pan paniscus". In McGrew WC, Marchant LF, Nishida T (eds.). Great ape societies. Cambridge, England: Cambridge Univ Press. pp. 29–41. ISBN 0521555361.
- Nicholls H (17 March 2016). "Do bonobos really spend all their time having sex?". BBC.
- Furuichi T (July 2011). "Female contributions to the peaceful nature of bonobo society". Evolutionary Anthropology. 20 (4): 131–42. doi:10.1002/evan.20308. PMID 22038769. S2CID 17830996.
- Hare B, Kwetuenda S (March 2010). "Bonobos voluntarily share their own food with others". Current Biology. 20 (5): R230-1. doi:10.1016/j.cub.2009.12.038. PMID 20219170. S2CID 28319610.
- Wobber V, Wrangham R, Hare B (February 2010). "Bonobos exhibit delayed development of social behavior and cognition relative to chimpanzees". Current Biology. 20 (3): 226–30. doi:10.1016/j.cub.2009.11.070. PMID 20116251. S2CID 3398517.
- Callaway E. "Peter Pan ways make bonobos the most amiable of apes". New Scientist. Retrieved 2021-03-14.
- Surbeck M, Deschner T, Schubert G, Weltring A, Hohmann G (March 2012). "Mate competition, testosterone and intersexual relationships in bonobos, Pan paniscus". Animal Behaviour. 83 (3): 659–69. doi:10.1016/j.anbehav.2011.12.010. S2CID 53198728. Lay summary – Max-Planck-Gesellschaft.
- Hogenboom M. "First personality test shows that female apes are irritable". www.bbc.com. Retrieved 2021-03-12.
- Furuichi T, Idani GI, Ihobe H, Hashimoto C, Tashiro Y, Sakamaki T, Mulavwa MN, Yangozene K, Kuroda S (2012), Kappeler PM, Watts DP (eds.), "Long-Term Studies on Wild Bonobos at Wamba, Luo Scientific Reserve, D. R. Congo: Towards the Understanding of Female Life History in a Male-Philopatric Species", Long-Term Field Studies of Primates, Berlin, Heidelberg: Springer, pp. 413–433, doi:10.1007/978-3-642-22514-7_18, ISBN 978-3-642-22514-7, retrieved 2021-01-13
- Jirik K. "LibGuides: Bonobo (Pan paniscus) Fact Sheet: Behavior & Ecology". ielc.libguides.com. Retrieved 2021-01-13.
- Surbeck, Martin; Hohmann, Gottfried (2018). "Affiliations, aggressions and an adoption: Male–male relationships in wild bonobos". Oxford Scholarship. doi:10.1093/oso/9780198728511.003.0003. ISBN 978-0-19-872851-1.
- Surbeck M, Mundry R, Hohmann G (February 2011). "Mothers matter! Maternal support, dominance status and mating success in male bonobos (Pan paniscus)". Proceedings. Biological Sciences. 278 (1705): 590–8. doi:10.1098/rspb.2010.1572. PMC 3025686. PMID 20810444. Lay summary – phys.org.
- Sample I (2019-05-20). "Pushy bonobo mothers help sons find sexual partners, scientists find". The Guardian. Retrieved 2021-01-13.
- "The Momma's Boy Strategy: Why Bonobo Males Tend Not To Form Coalitions". Traditions of Conflict. Retrieved 2021-05-07.
- Tokuyama N, Toda K, Poiret ML, Iyokango B, Bakaa B, Ishizuka S (March 2021). "Two wild female bonobos adopted infants from a different social group at Wamba". Scientific Reports. 11 (1): 4967. Bibcode:2021NatSR..11.4967T. doi:10.1038/s41598-021-83667-2. PMC 7973529. PMID 33737517. Lay summary – phys.org.
- Tokuyama N, Toda K, Poiret ML, Iyokango B, Bakaa B, Ishizuka S (March 2021). "Two wild female bonobos adopted infants from a different social group at Wamba". Scientific Reports. 11 (1): 4967. Bibcode:2021NatSR..11.4967T. doi:10.1038/s41598-021-83667-2. PMC 7973529. PMID 33737517.
- Clint E (2017-10-09). "Faux-nobo: "Naked Bonobo" demolishes myth of sexy, egalitarian bonobos". Incredulous. Retrieved 2020-12-18.
- Cawthon Lang KA (December 2010). "Bonobo (Pan paniscus) behavior". Primate Factsheets. University of Wisconsin. Archived from the original on 2016-04-12.
- "16 The Absence of Sexual Coercion in Bonobos", Sexual Coercion in Primates and Humans, Harvard University Press, pp. 410–423, 2009-12-31, doi:10.4159/9780674054349-016, ISBN 978-0-674-05434-9
- White FJ, Wood KD (August 2007). "Female feeding priority in bonobos, Pan paniscus, and the question of female dominance". American Journal of Primatology. 69 (8): 837–50. doi:10.1002/ajp.20387. PMID 17358018. S2CID 17628292.
- Samuni, Liran; Wegdell, Franziska; Surbeck, Martin (2020-09-01). Weigel, Detlef; Van de Waal, Erica; Van de Waal, Erica (eds.). "Behavioural diversity of bonobo prey preference as a potential cultural trait". eLife. 9: e59191. doi:10.7554/eLife.59191. ISSN 2050-084X. PMC 7462605. PMID 32869740.
- Nicholls H. "Do bonobos really spend all their time having sex?". www.bbc.com. Retrieved 2020-08-08.
- Jones N (2018-04-05). "Bonobos Spied Sharing a Feast". SAPIENS. Retrieved 2020-08-08.
- Wakefield ML, Hickmott AJ, Brand CM, Takaoka IY, Meador LM, Waller MT, White FJ (2019). "New Observations of Meat Eating and Sharing in Wild Bonobos (Pan paniscus) at Iyema, Lomako Forest Reserve, Democratic Republic of the Congo". Folia Primatologica; International Journal of Primatology. 90 (3): 179–189. doi:10.1159/000496026. PMID 30889597. S2CID 84183655. Lay summary – The Leakey Foundation.
- "Aggression topics". University of New Hampshire. Archived from the original on 20 March 2005.
- Manson JH, Perry S, Parish AR (1997). "Nonconceptive Sexual Behavior in Bonobos and Capuchins". International Journal of Primatology. 18 (5): 767–86. doi:10.1023/A:1026395829818. S2CID 3032455.
- Nguyen TC (2008-02-13). "Gorillas Caught in Very Human Act". Live Science.
- de Waal FB (March 1995). "Bonobo sex and society" (PDF). Scientific American. 272 (3): 82–8. Bibcode:1995SciAm.272c..82W. doi:10.1038/scientificamerican0395-82. PMID 7871411. Archived from the original (PDF) on 27 January 2012. Retrieved 21 December 2011.
- Balcombe JP (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 N (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 SB (April 2006). "Perineal swelling, intermenstrual cycle, and female sexual behavior in bonobos (Pan paniscus)". American Journal of Primatology. 68 (4): 333–47. doi:10.1002/ajp.20228. PMID 16534808. S2CID 25823290.
- Hohmann G, Fruth B (July 2000). "Use and function of genital contacts among female bonobos". Animal Behaviour. 60 (1): 107–120. doi:10.1006/anbe.2000.1451. PMID 10924210. S2CID 39702173.
- "Courtney Laird, "Social Organization"". Bio.davidson.edu. 2004. Archived from the original on 2011-05-19. Retrieved 2009-07-03.
- de Waal FB (2001). "Bonobos and Fig Leaves". The ape and the sushi master : cultural reflections by a primatologist. Basic Books. ISBN 978-84-493-1325-7.
- Kitamura K (August 1989). "Genito-Genital Contacts in the Pygmy Chimpanzees (Pan paniscus)". African Study Monographs. 10 (2): 49–67. doi:10.14989/68052. hdl:2433/68052.
- Hogenboom M (6 February 2015). "Are there any homosexual animals?". BBC.
- Ihobe H, Furuichi T (1994). "Variation in Male Relationships in Bonobos and Chimpanzees". Behaviour. 130 (3–4): 211–228. doi:10.1163/156853994x00532. ISSN 0005-7959.
- Williams A, Myers P (2004). "Pan paniscus". Animal Diversity Web. Retrieved 6 January 2012.
- de Waal FB (1990). "Sociosexual behavior used for tension regulation in all age and sex combinations among bonobos". In Feierman JR (ed.). Pedophilia: Biosocial Dimensions. New York: Springer. pp. 378–393. ISBN 9781461396840.
- Small MF (1 June 1992). "Casual Sex Play Common Among Bonobos". Discover.
Even juveniles participate by rubbing their genital areas against adults, although ethologists don't think that males actually insert their penises into juvenile females.
- Gottfried H, Vigilant L, Mundry R, Behringer V, Surbeck M (May 2019). "Aggression by male bonobos against immature individuals does not fit with predictions of infanticide". Aggressive Behavior. 45 (3): 300–309. doi:10.1002/ab.21819. PMID 30710459. S2CID 73440844.
- Hare B, Wobber V, Wrangham R (March 2012). "The self-domestication hypothesis: evolution of bonobo psychology is due to selection against aggression". Animal Behaviour. 83 (3): 573–585. doi:10.1016/j.anbehav.2011.12.007. ISSN 0003-3472. S2CID 3415520.
- Beaudrot LH, Kahlenberg SM, Marshall AJ (September 2009). "Why male orangutans do not kill infants". Behavioral Ecology and Sociobiology. 63 (11): 1549–1562. doi:10.1007/s00265-009-0827-1. PMC 2728907. PMID 19701484.
- Sharp PM, Shaw GM, Hahn BH (April 2005). "Simian immunodeficiency virus infection of chimpanzees". Journal of Virology. 79 (7): 3891–902. doi:10.1128/jvi.79.7.3891-3902.2005. PMC 1061584. PMID 15767392.
- "Chimpanzee behavior: Killer instincts". The Economist. June 24, 2010. Retrieved 2011-12-08.
- Wrangham R (2019). The Goodness Paradox: The Strange Relationship Between Virtue and Violence in Human Evolution. Knopf Doubleday Publishing Group. pp. 19–20. ISBN 978-1-101-97019-5.
- Raffaele P (November 2006). "The Smart and Swinging Bonobo". Smithsonian Magazine.
- Caswell JL, Mallick S, Richter DJ, Neubauer J, Schirmer C, Gnerre S, Reich D (April 2008). McVean G (ed.). "Analysis of chimpanzee history based on genome sequence alignments". PLOS Genetics. 4 (4): e1000057. doi:10.1371/journal.pgen.1000057. PMC 2278377. PMID 18421364.
- White FJ, Wrangham RW (1988). "Feeding competition and patch size in the chimpanzee species Pan paniscus and Pan troglodytes". Behaviour. 105 (1/2): 148–164. doi:10.1163/156853988X00494. JSTOR 4534684. S2CID 18285801.
- Rilling JK, Scholz J, Preuss TM, Glasser MF, Errangi BK, Behrens TE (April 2012). "Differences between chimpanzees and bonobos in neural systems supporting social cognition". Social Cognitive and Affective Neuroscience. 7 (4): 369–79. doi:10.1093/scan/nsr017. PMC 3324566. PMID 21467047.
- Davidson RJ, Putnam KM, Larson CL (July 2000). "Dysfunction in the neural circuitry of emotion regulation--a possible prelude to violence". Science. 289 (5479): 591–4. Bibcode:2000Sci...289..591D. doi:10.1126/science.289.5479.591. PMID 10915615.
- Pezawas L, Meyer-Lindenberg A, Drabant EM, Verchinski BA, Munoz KE, Kolachana BS, et al. (June 2005). "5-HTTLPR polymorphism impacts human cingulate-amygdala interactions: a genetic susceptibility mechanism for depression". Nature Neuroscience. 8 (6): 828–34. doi:10.1038/nn1463. PMID 15880108. S2CID 1864631.
- Vastag B (11 April 2011). "Brain differences may explain varying behavior of bonobos and chimpanzees". The Washington Post. Retrieved 26 December 2012.
- de Waal F (August 8, 2007). "Bonobos, Left & Right". Skeptic.
- Clint, Edward (2017-10-09). "Faux-nobo: "Naked Bonobo" demolishes myth of sexy, egalitarian bonobos". Incredulous. Retrieved 2021-06-03.
- Surbeck M, Fowler A, Deimel C, Hohmann G (February 2009). "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. S2CID 32622605.; Surbeck M, Hohmann G (October 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. S2CID 6708310.
- Lang KC (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. 33 (2): 247–250. doi:10.1007/BF02382754. S2CID 10063791.
- Rafert J, Vineberg EO (1997). "Bonobo Nutrition – relation of captive diet to wild diet" (PDF). Archived from the original (PDF) on 2012-04-25." Bonobo Husbandry Manual, American Association of Zoos and Aquariums
- Parker I (2007-07-30). "Swingers". Our Far-Flung Correspondents. The New Yorker. Retrieved 2011-12-08.
- de Waal F (2009-10-18). "Was "Ardi" a Liberal?". The Huffington Post. Retrieved 2009-10-18.
- Walker M (2010-02-01). "Wild bonobo mother ape eats own infant in DR Congo". BBC News.
- Callaway E (1 February 2010). "Hippy apes caught cannibalising their young". New Scientist. Retrieved 18 April 2013.
- Tokuyama, Nahoko, Deborah Lynn Moore, Kirsty Emma Graham, Albert Lokasola, and Takeshi Furuichi. "Cases of maternal cannibalism in wild bonobos (Pan paniscus) from two different field sites, Wamba and Kokolopori, Democratic Republic of the Congo." Primates 58, no. 1 (2017): 7-12.
- Herrmann E, Hare B, Call J, Tomasello M (August 2010). "Differences in the cognitive skills of bonobos and chimpanzees". PLOS ONE. 5 (8): e12438. Bibcode:2010PLoSO...512438H. doi:10.1371/journal.pone.0012438. PMC 2929188. PMID 20806062. Available under CC BY 4.0.
- Kovalaskas S, Rilling JK, Lindo J (March 2021). "Comparative analyses of the Pan lineage reveal selection on gene pathways associated with diet and sociality in bonobos". Genes, Brain, and Behavior. 20 (3): e12715. doi:10.1111/gbb.12715. PMID 33200560. S2CID 226988471. Lay summary – phys.org.
- Gruber T, Clay Z, Zuberbühler K (1 December 2010). "A comparison of bonobo and chimpanzee tool use: evidence for a female bias in the Pan lineage" (PDF). Animal Behaviour. 80 (6): 1023–1033. doi:10.1016/j.anbehav.2010.09.005. S2CID 14923158.
- Dielenberg, Robert A. "The comparative psychology of human uniqueness: A cognitive behavioral review." (2013).
- Anderson, James R., and Gordon G. Gallup. "Mirror self-recognition: a review and critique of attempts to promote and engineer self-recognition in primates." Primates 56, no. 4 (2015): 317-326.
- "Bonobos at the Columbus Zoo". Columbus Zoo. Archived from the original on 2006-05-02. Retrieved 2006-08-01.
- Webb J (August 4, 2015). "Bonobo squeaks hint at earlier speech evolution". BBC News. Retrieved 2015-08-05.
- "Meet our Great Apes: Kanzi". Great Ape Trust. 2007. Archived from the original on 2008-06-30. Retrieved 2008-09-28.
- Raffaele P (2006). "Speaking Bonobo". Smithsonian. Retrieved 2008-09-28.
- Schick KD, Toth N, Garufi G, Savage-Rumbaugh ES, Rumbaugh D, Sevcik R (1999). "Continuing Investigations into the Stone Tool-making and Tool-using Capabilities of a Bonobo (Pan paniscus)". Journal of Archaeological Science. 26 (7): 821–832. doi:10.1006/jasc.1998.0350.
- Palagi E, Paoli T, Tarli SB (January 2004). "Reconciliation and consolation in captive bonobos (Pan paniscus)". American Journal of Primatology. 62 (1): 15–30. doi:10.1002/ajp.20000. PMID 14752810. S2CID 22452710.
- 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. PMC 3818457. PMID 24223924.
- Beale B (2003). "Where Did Laughter Come From?". ABC Science Online. Retrieved 2008-08-10.
- Dawkins R (2004). "Chimpanzees". The Ancestor's Tale. Houghton Mifflin. ISBN 978-1-155-16265-2.
- Quammen D. "The Left Bank Ape". The New Age of Exploration, 2013. National Geographic. Retrieved 28 February 2013.
- Beaune D, Bretagnolle F, Bollache L, Hohmann G, Surbeck M, Fruth B (January 2013). "Seed dispersal strategies and the threat of defaunation in a Congo forest". Biodiversity and Conservation. 22 (1): 225–238. doi:10.1007/s10531-012-0416-x. ISSN 0960-3115. S2CID 17600509.
- Beaune D, Bretagnolle F, Bollache L, Bourson C, Hohmann G, Fruth B (September 2013). "Ecological services performed by the bonobo (Pan paniscus): seed dispersal effectiveness in tropical forest". Journal of Tropical Ecology. 29 (5): 367–380. doi:10.1017/s0266467413000515. ISSN 0266-4674. S2CID 86540021.
- Beaune D, Bollache L, Bretagnolle F, Fruth B (2012-08-29). "Dung beetles are critical in preventing post-dispersal seed removal by rodents in Congo rain forest". Journal of Tropical Ecology. 28 (5): 507–510. doi:10.1017/s0266467412000466. ISSN 0266-4674. S2CID 86529605.
- Beaune D, Bretagnolle F, Bollache L, Hohmann G, Surbeck M, Bourson C, Fruth B (April 2013). "The bonobo-dialium positive interactions: seed dispersal mutualism". American Journal of Primatology. 75 (4): 394–403. doi:10.1002/ajp.22121. PMID 23307414. S2CID 21817223.
- Beaune D (2015-02-27). "What would happen to the trees and lianas if apes disappeared?". Oryx. 49 (3): 442–446. doi:10.1017/s0030605314000878. ISSN 0030-6053. S2CID 84253241.
- Reid J (2006-06-15). "Parks and people, not parks vs. people". San Francisco Chronicle.
- "Bonobo and large mammal survey". Zoological Society of Milwaukee. Archived from the original on May 6, 2012.
- Curwood S, Parish A (July 2006). "The Make Love, Not War Species". Living on Earth. National Public Radio.
- Hart T (2012-07-27). "Searching for Bonobo in Congo". Bonoboincongo.com. Retrieved 2012-12-26.
- "Lola Ya Bonobo (Bonobo Heaven)]". Lolayabonobo.wildlifedirect.org. Archived from the original on 2016-11-24. Retrieved 2012-12-26.
- "Bonobo Reintroduction in the Democratic Republic of Congo" (PDF). friendsofbonobos.org. November 2009. Archived from the original (PDF) on 2010-12-18. Retrieved 2012-12-26.
- "Bonobo and Congo Biodiversity Initiative". Zoological Society of Milwaukee. Archived from the original on 10 February 2012.
- Chapin M (December 2004). "Vision for a Sustainable World" (PDF). WORLDWATCH magazine.
- de Waal F, Lanting F (1997). Bonobo: The Forgotten Ape. University of California Press. ISBN 0-520-20535-9.
- Takayoshi K (1992). The Last Ape: Pygmy Chimpanzee Behavior and Ecology. Stanford, CA: Stanford University Press.
- Savage-Rumbaugh S, Lewin R (1994). Kanzi: The Ape at the Brink of the Human Mind. John Wiley. ISBN 0-471-58591-2.
- Woods V (2010). Bonobo Handshake. Gotham Books. ISBN 978-1-59240-546-6.
- Sandin J (2007). Bonobos: Encounters in Empathy. Zoological Society of Milwaukee & The Foundation for Wildlife Conservation, Inc. ISBN 978-0-9794151-0-4.
- de Waal F (2013). The Bonobo and the Atheist. Norton. ISBN 978-0393073775.
- de Waal F (1995). "Bonobo: Sex & Society". Scientific American. 272 (3): 82–88. Bibcode:1995SciAm.272c..82W. doi:10.1038/scientificamerican0395-82. PMID 7871411.
- DeBartolo A (11 June 1998). "The Bonobo: 'Newest' apes are teaching us about ourselves". Chicago Tribune. Archived from the original on 2003-02-11.
- Schweller K (July 2012). "Apes with Apps". IEEE Spectrum Magazine. 49 (7): 38–45. doi:10.1109/MSPEC.2012.6221081. S2CID 22556649.
- Madrigal A (11 June 2014). "Brian the Mentally Ill Bonobo, and How He Healed". The Atlantic.
- Parker I (30 July 2007). "Swingers". The New Yorker.
- Bechard D (February 2014). "Viral Conservation". The Solutions Journal.
- Fischer A, Prüfer K, Good JM, Halbwax M, Wiebe V, André C, et al. (29 June 2011). Joly E (ed.). "Bonobos fall within the genomic variation of chimpanzees". PLOS ONE. 6 (6): e21605. Bibcode:2011PLoSO...621605F. doi:10.1371/journal.pone.0021605. PMC 3126833. PMID 21747915.
- Zsurka G, Kudina T, Peeva V, Hallmann K, Elger CE, Khrapko K, Kunz WS (September 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. PMC 2942848. PMID 20813043.
- Wildman DE, Uddin M, Liu G, Grossman LI, Goodman M (June 2003). "Implications of natural selection in shaping 99.4% nonsynonymous DNA identity between humans and chimpanzees: enlarging genus Homo". Proceedings of the National Academy of Sciences of the United States of America. 100 (12): 7181–8. Bibcode:2003PNAS..100.7181W. doi:10.1073/pnas.1232172100. PMC 165850. PMID 12766228.
|Wikimedia Commons has media related to Pan paniscus.|
|Wikispecies has information related to Bonobo.|
- Evolution: Why Sex?
- Bonobos: Wildlife summary from the African Wildlife Foundation
- Primate Info Net Pan paniscus Factsheet
- Susan Savage-Rumbaugh: Apes that write, start fires and play Pac-Man – Ted.com
- WWF (World Wide Fund for Nature / World Wildlife Fund) – Bonobo species profile
- San Diego Zoo Library: Bonobo, Pan paniscus
- View the panPan1 genome assembly in the UCSC Genome Browser.