Bonobo
Bonobo[1] | |
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Species: | P. paniscus |
Pan paniscus | |
Bonobo distribution | |
The Bonobo (English pronunciation: /bəˈnoʊboʊ/[3][4] /ˈbɒnɵboʊ/[5]), Pan paniscus, previously called the Pygmy Chimpanzee and less often, the Dwarf or Gracile Chimpanzee,[6] is a great ape and one of the two species making up the genus Pan. The other species in genus Pan 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, while Pan paniscus is usually referred to as the Bonobo.
The Bonobo is endangered and is found in the wild only in the Democratic Republic of the Congo. Along with the Common Chimpanzee, the Bonobo is the closest extant relative to humans. Since the two species are not proficient swimmers, it is possible that the formation of the Congo River 1.5–2 million years ago led to the speciation of the Bonobo. They live south of the river, and thereby were separated from the ancestors of the Common Chimpanzee, which live north of the river.[7]
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.[8]
The species is distinguished by relatively long legs, pink lips, dark face and tail-tuft through adulthood, and parted long hair on their head. Although, it had been reported that Bonobos were a matriarchal species, more recent research on wild bonobos and accounts reported in the Smithsonian magazine indicate that Bonobo troops are led by a dominant alpha male who plays an active role in defending the troop from predators:
"The alpha male is usually in charge," Ihomi says. The alpha male determines where the troop eats and sleeps and when it moves, and he is the first to defend the troop from leopards and pythons.
In captivity, Bonobos were perceived to be a matriarchal species that commonly engages in casual sexual activity, as well as significant homosexual contact.[10][11] The limited field research on bonobos in the wild reveals that these behaviors may be exaggerated by captivity, as well as by food provisioning by researchers in the field.[10]
This primate is mainly frugivorous, but supplements its diet with leaves and meat from small vertebrates, such as flying squirrels and duikers,[12] and invertebrates.[13] In some instances, bonobos have been shown to consume lower-order primates.[14][15] Some claim that Bonobos have also been known to practice cannibalism in captivity, a claim disputed by others.[10][11]
Contents[hide] · 1 Name |
Name
Common name
The name Bonobo first appeared in 1954, when Edward Tratz and Heinz Heck proposed it as a new and separate generic term for pygmy chimpanzees. The term has been reported variously[citation needed] as being a word for "chimpanzee" or "ancestor" in a Bantu language. Another suggestion for the derivation of the name is that the name is a misspelling of the name of the town of Bolobo on the Congo River, which has been associated with the collection of chimps in the 1920s.[16]
Taxonomy
The scientific name for the Bonobo is Pan paniscus. Initial genetic studies characterised their DNA as being as much as 98% (99.4 in one study) identical to that of Homo sapiens.[17] Later studies showed that chimpanzees are more closely related to humans than to gorillas.[18] The most recent genetic analyses (published in 2006) of chimpanzee and human genetic similarity come from whole genome comparisons and have shown that the differences between the two species are more complex, both in extent and character, than the historical 98% figure suggests [19] .
In the seminal Nature paper reporting on initial genome comparisons, researchers identified thirty-five million single-nucleotide changes, five million insertion or deletion events, and a number of chromosomal rearrangements which constituted the genetic differences between chimpanzees and humans, covering 98% of the same genes.[20] While many of these analyses have been performed on the Common Chimpanzee rather than the Bonobo, the differences between the two chimpanzee species are unlikely to be substantial enough to affect the Pan-Homo comparative data significantly.
There still is controversy, however. Scientists such as Jared Diamond in The Third Chimpanzee, and Morris Goodman [21] of Wayne State University in Detroit argue 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 Rule 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 other species closely related to humans, including Australopithecus. It should be noted that ideas such as this are considered far outside the mainstream; the current line between homo and non-homo is drawn about 2 million years ago, and chimpanzee and human ancestry only converges about 7 million years ago, nearly three times earlier. It also should be noted that those promoting chimpanzees as "third humans" are mostly chimpanzee researchers, with an obvious bias.
Recent DNA evidence[when?] suggests the Bonobo and Common Chimpanzee species effectively separated from each other less than one million years ago.[22][23] The chimpanzee 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.
Physical characteristics
Bonobo
The Bonobo is sometimes considered to be more gracile than the Common Chimpanzee, and females are somewhat smaller than males. Its head is 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. The Bonobo walks upright approximately 25% of the time during ground locomotion. Its quadrupedal ground locomotion generally is characterized by forelimb 'palm walking', similar to orangutans and in contrast to the predominant use of knuckles as characteristic of gorillas and the Common Chimpanzees. These physical characteristics and its posture, give the Bonobo an appearance more closely resembling humans than that of the Common Chimpanzee (see: bipedal Bonobos). 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.
The controversial idea that Bonobos are more attractive and human-like than chimpanzees has added to their popularity and a devoted following among some people.[10]
Psychological characteristics
Frans de Waal, one of the world's most respected and popular primatologists, states that the Bonobo is capable of altruism, compassion, empathy, kindness, patience, and sensitivity. How peaceful bonobos are has been disputed by some, but in general scientists agree with these assessments and the fact remains that thus far there are no confirmed observations of lethal aggression among bonobos, either in the wild or in captivity.
Observations in the wild indicate that the males among the related Common Chimpanzee communities are extraordinarily hostile to males from outside of the community. Parties of males 'patrol' for the unfortunate neighbouring males who might be traveling alone, and attack those single males, often killing them. This does not appear to be the behavior of the Bonobo males or females in their own communities, where they 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 resolve conflicts with sexual contact (hence the "make love - not war" characterization for the species). Bonobos live in different areas from the more aggressive Common Chimpanzee. Neither of the species swims, their respective ranges being separated by the great Congo River with Bonobos living south of the river and Chimpanzees living north of the river. 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.
The popular image of the Bonobo as a peaceful ape does not always apply to captive populations. Accounts exist of Bonobos confined in zoos mutilating one another and engaging in bullying. These incidents may be due to the practice in zoos of separating mothers and sons, which is contrary to their social organization in the wild. 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" as well as writing that "when first writing about their behavior, 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". The immature state of Bonobo research in the wild, compared to that of the Common Chimpanzee means that lethal aggression between Bonobos could still be discovered.
Hohmann and Surbeck published in 2008 that Bonobos sometimes do hunt monkey species. Having observed a group of bonobos in Salonga National Park they witnessed five incidents in five years where Bonobos preyed on monkeys. Their research indicates it was deliberate hunting, where a group of Bonobos would coordinate their actions—contrary to their normal hunting behaviour, which is quite solitary and less purposeful. In three occasions the hunt was successful and infant monkeys were captured, once a redtail monkey and twice a Cercopithecus wolfi. The spoils, however, were distributed quite peacefully among the members of the group.[15][24]
Social behavior
Bonobo fishing for termites
In the past, Females were considered to have a higher social status and Bonobo were thought to have a matriarchal culture.[citation needed], however, recent field work and observation indicates that Bonobo troops are led by an alpha male although females are not subordinate to all adult males as in Chimpanzees.[25] Aggressive encounters between males and females are rare, and males are tolerant of infants and juveniles. A male's status is derived from the status of his mother. The mother-son bond often stays strong and continues throughout life. While social hierarchies do exist, rank does not play so prominent a role as it does in other primate societies.
Bonobo party size tends to be variable since the groups exhibit a fission-fusion pattern. A community of approximately one hundred will split into small groups during the day while looking for food, and then come back together to sleep. They sleep in trees in nests that they construct.
Sexual social behavior
Sexual intercourse plays a major role in bonobo society observed in captivity, being used as what some scientists perceive as a greeting, a means of conflict resolution, and post-conflict 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, tongue kissing, and oral sex[26] (although a pair of Western Gorillas have been photographed performing face to face genital sex[27]). In scientific literature, the female-female behavior of touching genitals together is often referred to as GG rubbing or genital-genital rubbing.
The sexual activity happens within the immediate family as well as outside it. Bonobos do not form permanent 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 intercourse between mothers and their adult sons; some observers believe these pairings are taboo. 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 allowing for peaceful feeding.[28]
Bonobo males occasionally engage in various forms of male-male genital behavior (frot).[29][30] In one form, two males hang from a tree limb face-to-face while "penis fencing".[31][32] Frot also may occur when two males rub their penises together while in face-to-face position. A special form of frot called "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.
Bonobo females also engage in female-female genital behavior, possibly to bond socially with each other, thus forming a female nucleus of Bonobo society. The bonding among females allows them to dominate Bonobo society—although male Bonobos are individually stronger, they cannot stand alone against a united group of females.[32] Adolescent females often leave their native community to join another community. Sexual bonding with other females establishes the new females as members of the group. This migration mixes the Bonobo gene pools, providing genetic diversity.
Bonobo reproductive rates are not any higher than that of the Common Chimpanzee. Female Bonobos carry and nurse their young for five years and can give birth every five to six years. Compared to Common Chimpanzees, Bonobo females resume the genital swelling cycle much sooner after giving birth, allowing them to rejoin the sexual activities of their society. Also, Bonobo females who are sterile or too young to reproduce still engage in sexual activity.
Craig Stanford, an American primatologist, has challenged the claim that Bonobos are more sexually active than Common Chimpanzees.[clarification needed] Stanford compared existing data on Common Chimpanzees and Bonobos in the natural habitat and found that female Common Chimpanzees copulated at least as often as female Bonobos, while he recorded that male chimpanzees copulated more than male Bonobos.[33] His comparison excluded same-sex sexual contacts, however, which are very common in Bonobos. De Waal's book on Bonobos includes interviews with field workers and relies especially on the studies by Takayoshi Kano, the only scientist to have worked for more than two decades with wild Bonobos.[34] Kano describes frequent sexual encounters of all kinds among the bonobos of Wamba, hence there is no reason to believe that such encounters are limited to captive bonobos.
Closeness to humans
Bonobos are capable of passing the mirror-recognition test for self-awareness. They communicate primarily through vocal means, although the meanings of their vocalizations are not currently known. However, most humans do understand their facial expressions[17] 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[35] and he has comprehension of around 3,000 spoken English words.[36] Some, such as philosopher and bioethicist Peter Singer, argue that these results qualify them for the "rights to survival and life," rights that humans theoretically accord to all persons.
There are instances in which non-human primates have been reported to have expressed joy. One study analyzed and recorded sounds made by human babies and Bonobos when they were tickled.[37] It found although the Bonobo's laugh was a higher frequency, the laugh followed a similar spectrographic pattern to human babies.[37]
Habitat
Around 10,000 Bonobos are found only south of the Congo River and north of the Kasai River (a tributary of the Congo),[38] in the humid forests of the Democratic Republic of Congo of central Africa. They are an endangered species, due both to habitat loss and hunting for bushmeat, the latter activity having increased dramatically during the current civil war due to the presence of heavily armed militias even in remote "protected" areas such as Salonga National Park. Today, at most several thousand Bonobos remain. This is part of a more general trend of ape extinction.
Conservation efforts
Since 1996, the first and second Congo wars in the Democratic Republic of Congo (DRC) have had a major impact on both the Bonobo and human population. Bonobos are in danger of being hunted to extinction. The key to Bonobo conservation efforts is balancing these issues.
As the Bonobo's habitat is shared with people, the ultimate success of conservation efforts will rely on local and community involvement. The issue of parks vs. people,[39] is very cogent in the Cuvette Centrale, the Bonobo's range. There is strong local and broad-based Congolese resistance to establishing national parks, as indigenous communities often have 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 recent surveys[when?] indicate that the Bonobo, the African Forest Elephant, and other species have been severely devastated by poachers and the thriving bushmeat trade. In contrast to this, there are areas where the Bonobo and biodiversity still thrive without any established parks, due to the indigenous beliefs and taboos against killing Bonobos.
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 twenty 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 (BCBI). 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, as of 2007[update], 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 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."[40]
This initiative has been gaining momentum and greater international recognition and it recently[when?] has gained greater support through Conservation International, the Global Conservation Fund, United States Fish and Wildlife Service's Great Ape Conservation Fund, and the United Nations' Great Apes Survival Project.
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 in order to find ways to help protect these apes.
· Develop anti-poaching measures to help save apes, forest elephants, and other endangered animals in Congo's Salonga National Park, a U.N. 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. As of 2007[update], 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.[41]
The Congo is setting aside more than 11,000 square miles (28,000 km2) of rain forest 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 rain forest, the U.S.-based Bonobo Conservation Initiative. The area amounts to just over 1 percent 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 thirty years, though surveys have been hard to carry out in war-ravaged central Congo. Estimates range from 60,000 to fewer than 5,000 living, according to the World Wildlife Fund.
The Sankuru reserve also contains Okapi, closely related to the Giraffe, that also is native to Congo, elephants, and at least ten other primate species.
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 non scientific or ecological sites have created various groups to collect donations to help with the conservation of this species.
See also
· Homosexual behavior in animals
· List of apes – notable individual apes
· Polygamy
References
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2. ^ 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. Downloaded on 4 January 2009. Listed as Endangered (EN A4cd v3.1)
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4. ^ "dictionary.com". Dictionary.reference.com. http://dictionary.reference.com/browse/bonobo. Retrieved 2009-07-03.
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6. ^ Frans de Waal, Frans Lanting (1997) Bonobo: The Forgotten Ape, University of California Press
7. ^ Caswell JL, Mallick S, Richter DJ, et al (April 2008). "Analysis of chimpanzee history based on genome sequence alignments". PLoS Genet. 4 (4): e1000057. doi:10.1371/journal.pgen.1000057. PMID 18421364.
8. ^ See Herzfeld, Chris (2007), « L’invention du bonobo », Bulletin d’histoire et d’épistémologie des sciences de la vie, volume 14, numéro 2, 2007.
9. ^ http://www.smithsonianmag.com/science-nature/bonobo.html?c=y&page=4
10. ^ a b c d Parker, Ian (2009-01-07). "Our Far-Flung Correspondents: Swingers: Reporting & Essays". The New Yorker. http://www.newyorker.com/reporting/2007/07/30/070730fa_fact_parker. Retrieved 2009-07-03.
11. ^ a b de Waal, Frans (2009-10-18). "Was "Ardi" a Liberal?". The Huffington Post. http://www.huffingtonpost.com/frans-de-waal/was-ardi-perhaps-liberal_b_325201.html. Retrieved 2009-10-18.
12. ^ Ihobe H (April 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. http://www.springerlink.com/content/h7145027g60n708l/.
13. ^ 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
14. ^ 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. http://www3.interscience.wiley.com/journal/121543963/abstract.
15. ^ a b Surbeck M, Hohmann G (14 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. http://www.current-biology.com/content/article/abstract?uid=PIIS0960982208011172.
16. ^ Sue Savage-Rumbaugh & Roger Lewin (1994). Kanzi: the ape at the brink of the human mind. John Wiley & Sons. p. 97 isbn=0471585912.
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27. ^ Tuan C. Nguyen, "Gorillas Caught in Very Human Act" in Live Science (February 13, 2008)
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29. ^ Frans de Waal, "Bonobo Sex and Society" in Scientific American (March 1995), p. 82ff
30. ^ "Courtney Laird, "Social Organization"". Bio.davidson.edu. 2004. http://www.bio.davidson.edu/people/vecase/Behavior/Spring2004/laird/Social%20Organization.htm. Retrieved 2009-07-03.
31. ^ Frans B. M. de Waal (2001). "Bonobos and Fig Leaves". The ape and the sushi master : cultural reflections by a primatologist. Basic Books.
32. ^ a b de Waal, Frans B.M. (March 1995). "Bonobo Sex and Society". Scientific American 272 (3). http://primate.uchicago.edu/DeWaalSciAm.doc.
33. ^ Stanford, C. B. (1998). "The social behavior of chimpanzees and bonobos". Current Anthropology 39: 399–407. doi:10.1086/204757.
34. ^ Kano, Takayoshi (1992). The Last Ape: Pygmy Chimpanzee Behavior and Ecology. Stanford University Press.
35. ^ "Meet our Great Apes: Kanzi". http://www.greatapetrust.org/bonobo/meet/kanzi.php. Retrieved 2008-09-28.
36. ^ Raffaele, P. (2006). "Speaking Bonobo". Smithsonian. http://www.smithsonianmag.com/science-nature/10022981.html. Retrieved 2008-09-28.
37. ^ a b Beale, B. (2003). "Where Did Laughter Come From?". ABC Science Online. http://dsc.discovery.com/news/afp/20031006/laughter.html. Retrieved 2008-08-10.
38. ^ Dawkins, Richard (2004). "Chimpanzees". The Ancestor's Tale. Houghton Mifflin.
39. ^ Reid, John, Parks and people, not parks vs. people, San Francisco Chronicle, June 15, 2006
40. ^ "The Make Love, Not War Species," Living on Earth, (July 2006), National Public Radio
41. ^ Chapin, Mac, (November/December 2004), Vision for a Sustainable World, WORLDWATCH magazine
Further reading
Books
· 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)
· de Waal, Frans, Our Inner Ape, Riverhead Books, 2005. ISBN 1-57322-312-3
· 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)
Articles
· de Waal, Frans, "Bonobo: Sex & Society", Scientific American, 1995
· de Waal, Frans, "Bonobos, Left & Right", Skeptic, August 8, 2007.
· DeBartolo, Anthony. "The Bonobo: 'Newest' apes are teaching us about ourselves", Chicago Tribune June 11, 1998.
· Q&A on bonobos on a field research site from Congo
· ARKive - BBC images and movies of the bonobo (Pan paniscus)
· Bonobos: Wildlife summary from the African Wildlife Foundation
· Chimpanzee and Human Behaviors
· Bonobos: The Left-Bank Chimpanzees
· The Great Ape Trust: Studying the Bonobo
· NPR's Science Friday piece on bonobos
· Primate Info Net Pan paniscus Factsheet
· U.S. Fish & Wildlife Service Species Profile
· "The Last Great Ape", an episode of Nova.
· Image: bonobos genito-genital rubbing
· Susan Savage-Rumbaugh: Apes that write, start fires and play Pac-Man - Ted.com
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