Bonobo#Cognitive comparisons to chimpanzees

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.{{cite book | vauthors = Savage-Rumbaugh S, Lewin R |year=1994 |title=Kanzi: the ape at the brink of the human mind |url=https://archive.org/details/kanzi00sues |url-access=registration |publisher=John Wiley & Sons |isbn=978-0-385-40332-0 |page=[https://archive.org/details/kanzi00sues/page/97 97]}}{{cite book | vauthors = de Waal F |year=2005 |title=Our Inner Ape |publisher=Riverhead Books |isbn=978-1-57322-312-6}}

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.{{cite journal |vauthors=Schwarz E |date=April 1, 1929 |title=Das Vorkommen des Schimpansen auf den linken Kongo-Ufer |journal=Revue de Zoologie et de Botanique Africaines |volume=16 |pages=425–426 |url=http://www.metafro.be/primates/English_translation_of_Schwarz.pdf |access-date=April 4, 2015 |archive-date=July 19, 2021 |archive-url=https://web.archive.org/web/20210719053540/http://www.metafro.be/primates/English_translation_of_Schwarz.pdf |url-status=dead }}{{cite journal | vauthors = Coolidge Jr HJ |date=July–September 1933 |title=Pan paniscus. Pigmy chimpanzee from south of the Congo river |journal=American Journal of Physical Anthropology |volume=18 |issue=1 |pages=1–59 |doi=10.1002/ajpa.1330180113}} Coolidge's paper contains a translation of Schwarz's earlier report. In 1933, American anatomist Harold Coolidge elevated it to species status.{{cite journal |vauthors=Herzfeld C |year=2007 |title=L'invention du bonobo |language=fr |journal=Bulletin d'Histoire et d'Épistémologie des Sciences de la Vie |volume=14 |issue=2 |pages=139–162 |url=http://www.chrisherzfeld.com/userfiles/publications/BONOBO.pdf |access-date=21 December 2011 |doi=10.3917/bhesv.142.0139 |archive-date=12 December 2022 |archive-url=https://web.archive.org/web/20221212181134/http://www.chrisherzfeld.com/userfiles/publications/BONOBO.pdf |url-status=live }} Major behavioural differences between bonobos and chimpanzees were first discussed in detail by Tratz and Heck in the early 1950s.{{cite book | vauthors = de Waal FB |year=2002 |title=Tree of Origin: What Primate Behavior Can Tell Us About Human Social Evolution |isbn=978-0-674-01004-8 |publisher=Harvard University Press |page=51}} Unaware of any taxonomic distinction with the common chimpanzee, American psychologist and primatologist Robert Yerkes had already noticed an unexpected major behavioural difference in the 1920s.{{cite book |title=Our Inner Ape |first=Frans |last=de Waal |authorlink=Frans de Waal |page=30 |publisher=Penguin |year=2006 |isbn=978-1-59448-196-3 |url=https://books.google.com/books?id=lDP4TPccgC8C&dq=%22Robert+Yerkes%22+%22expressed+doubt+that+Prince+Chim+was+a+regular+chimp%22%22&pg=PA30 |access-date=2023-04-23 |archive-date=2023-07-30 |archive-url=https://web.archive.org/web/20230730121832/https://books.google.com/books?id=lDP4TPccgC8C&dq=%22Robert+Yerkes%22+%22expressed+doubt+that+Prince+Chim+was+a+regular+chimp%22%22&pg=PA30 |url-status=live }}

Bonobos and chimpanzees are the two species which make up the genus Pan, and are the closest living relatives to humans (Homo sapiens).{{cite journal | vauthors = Takahata N, Satta Y, Klein J | title = Divergence time and population size in the lineage leading to modern humans | journal = Theoretical Population Biology | volume = 48 | issue = 2 | pages = 198–221 | date = October 1995 | pmid = 7482371 | doi = 10.1006/tpbi.1995.1026 | doi-access = free | bibcode = 1995TPBio..48..198T }}{{cite journal | vauthors = Waterson RH, Lander ES, Wilson RK | collaboration = Chimpanzee Sequencing Analysis Consortium | title = Initial sequence of the chimpanzee genome and comparison with the human genome | journal = Nature | volume = 437 | issue = 7055 | pages = 69–87 | date = September 2005 | pmid = 16136131 | doi = 10.1038/nature04072 | doi-access = free | bibcode = 2005Natur.437...69.}}

According to studies published in 2017 by researchers at The George Washington University, bonobos, along with common chimpanzees, split from the human line about 8 million years ago; moreover, bonobos split from the common chimpanzee line about 2 million years ago.{{cite news |author=Staff |title=Bonobos May Resemble Humans More Than You Think - A GW researcher examined a great ape species' muscles and found they are more closely related to humans than common chimpanzees. |url=https://gwtoday.gwu.edu/bonobos-may-resemble-humans-more-you-think |date=5 May 2017 |work=The George Washington University |url-status=live |archive-url=https://archive.today/20230414135148/https://gwtoday.gwu.edu/bonobos-may-resemble-humans-more-you-think |archive-date=14 April 2023 |access-date=14 April 2023}}{{cite journal |last1=Diogo |first1=Rui |last2=Molnar |first2=Julia L. |last3=Wood |first3=Bernard |title=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 |journal=Scientific Reports |date=4 April 2017 |volume=7 |number=608 |page=608 |doi=10.1038/s41598-017-00548-3 |pmid=28377592 |pmc=5428693 |bibcode=2017NatSR...7..608D |s2cid=256924135 }}

Nonetheless, 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.{{cite journal | vauthors = Patterson N, Richter DJ, Gnerre S, Lander ES, Reich D | title = Genetic evidence for complex speciation of humans and chimpanzees | journal = Nature | volume = 441 | issue = 7097 | pages = 1103–8 | date = June 2006 | pmid = 16710306 | doi = 10.1038/nature04789 | s2cid = 2325560 | bibcode = 2006Natur.441.1103P }} DNA evidence suggests the bonobo and common chimpanzee species diverged approximately 890,000–860,000 years ago following separation of these two populations possibly because of 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.{{cite journal | vauthors = Won YJ, Hey J | title = Divergence population genetics of chimpanzees | journal = Molecular Biology and Evolution | volume = 22 | issue = 2 | pages = 297–307 | date = February 2005 | pmid = 15483319 | doi = 10.1093/molbev/msi017 | doi-access = free }} The first Pan fossils were reported in 2005 from the Middle Pleistocene (after the bonobo–chimpanzee split) of Kenya, alongside early Homo fossils.{{cite journal | vauthors = McBrearty S, Jablonski NG | title = First fossil chimpanzee | journal = Nature | volume = 437 | issue = 7055 | pages = 105–8 | date = September 2005 | pmid = 16136135 | doi = 10.1038/nature04008 | s2cid = 4423286 | bibcode = 2005Natur.437..105M }}

According to A. Zihlman, bonobo body proportions closely resemble those of Australopithecus,{{cite journal | vauthors = Zihlman AL, Cronin JE, Cramer DL, Sarich VM | title = Pygmy chimpanzee as a possible prototype for the common ancestor of humans, chimpanzees and gorillas | journal = Nature | volume = 275 | issue = 5682 | pages = 744–6 | date = October 1978 | pmid = 703839 | doi = 10.1038/275744a0 | bibcode = 1978Natur.275..744Z | s2cid = 4252525 }} leading evolutionary biologist Jeremy Griffith to suggest that bonobos may be a living example of our distant human ancestors.{{cite book |vauthors=Griffith J |author-link=Jeremy Griffith |year=2013 |title=Freedom Book 1 |chapter=Part 8:4H Humans’ development of integration through love-indoctrination and mate selection |publisher=WTM Publishing & Communications |isbn=978-1-74129-011-0 |chapter-url=http://www.worldtransformation.com/freedom-book1-integration-through-love-indoctrination/ |access-date=2013-03-28 |archive-date=2014-12-19 |archive-url=https://web.archive.org/web/20141219052145/http://www.worldtransformation.com/freedom-book1-integration-through-love-indoctrination/ |url-status=live }} 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.{{cite journal |doi=10.1515/anre-2015-0009 |title=The life history of Ardipithecus ramidus: A heterochronic model of sexual and social maturation |journal=Anthropological Review |volume=78 |issue=2 |pages=109–132|year=2015 | vauthors = Clark G, Henneberg M |doi-access=free }}

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{{cite journal | vauthors = Prüfer K, Munch K, Hellmann I, Akagi K, Miller JR, Walenz B, Koren S, Sutton G, Kodira C, Winer R, Knight JR, Mullikin JC, Meader SJ, Ponting CP, Lunter G, Higashino S, Hobolth A, Dutheil J, Karakoç E, Alkan C, Sajjadian S, Catacchio CR, Ventura M, Marques-Bonet T, Eichler EE, André C, Atencia R, Mugisha L, Junhold J, Patterson N, Siebauer M, Good JM, Fischer A, Ptak SE, Lachmann M, Symer DE, Mailund T, Schierup MH, Andrés AM, Kelso J, Pääbo S | display-authors = 6 | title = The bonobo genome compared with the chimpanzee and human genomes | journal = Nature | volume = 486 | issue = 7404 | pages = 527–31 | date = June 2012 | pmid = 22722832 | pmc = 3498939 | doi = 10.1038/nature11128 | bibcode = 2012Natur.486..527P }} 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.{{cite web |vauthors=Karow J |date=2008-05-13 |title=Neandertal, bonobo genomes may shed light on human evolution; MPI, 454 preparing drafts |publisher=Genome Web |work=In Sequence |url=http://www.genomeweb.com/sequencing/neandertal-bonobo-genomes-may-shed-light-human-evolution-mpi-454-preparing-draft |access-date=2011-12-08 |archive-date=2012-01-18 |archive-url=https://web.archive.org/web/20120118184736/http://www.genomeweb.com/sequencing/neandertal-bonobo-genomes-may-shed-light-human-evolution-mpi-454-preparing-draft |url-status=live }}

{{Infobox genome

| taxId = 10729

| ploidy = diploid

| chromosomes = 24 pairs

| size = 2,869.21 Mb

| year = 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,{{Cite journal|last1=Prüfer|first1=Kay|last2=Munch|first2=Kasper|last3=Hellmann|first3=Ines|last4=Akagi|first4=Keiko|last5=Miller|first5=Jason R.|last6=Walenz|first6=Brian|last7=Koren|first7=Sergey|last8=Sutton|first8=Granger|last9=Kodira|first9=Chinnappa|last10=Winer|first10=Roger|last11=Knight|first11=James R.|date=June 2012|title=The bonobo genome compared with the chimpanzee and human genomes|url= |journal=Nature|volume=486|issue=7404|pages=527–531|doi=10.1038/nature11128|pmc=3498939|pmid=22722832|bibcode=2012Natur.486..527P}} a high-quality reference genome became available only in 2021.{{Cite journal|last1=Mao|first1=Yafei|last2=Catacchio|first2=Claudia R.|last3=Hillier|first3=LaDeana W.|last4=Porubsky|first4=David|last5=Li|first5=Ruiyang|last6=Sulovari|first6=Arvis|last7=Fernandes|first7=Jason D.|last8=Montinaro|first8=Francesco|last9=Gordon|first9=David S.|last10=Storer|first10=Jessica M.|last11=Haukness|first11=Marina|date=2021-05-05|title=A high-quality bonobo genome refines the analysis of hominid evolution|journal=Nature|volume=594|issue=7861|pages=77–81|doi=10.1038/s41586-021-03519-x|pmid=33953399|pmc=8172381|bibcode=2021Natur.594...77M}} 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.

Researchers have found that both central (Pan troglodytes troglodytes) and eastern chimpanzees (Pan troglodytes schweinfurthii) share more genetic material with bonobos than other chimpanzee subspecies.{{Cite news |last=Owens |first=Brian |date=2016-10-27 |title=Chimps and bonobos interbred and exchanged genes |work=New Scientist |url=https://www.newscientist.com/article/2110682-chimps-and-bonobos-interbred-and-exchanged-genes/ |access-date=2022-04-06 |archive-date=2022-01-28 |archive-url=https://web.archive.org/web/20220128162354/https://www.newscientist.com/article/2110682-chimps-and-bonobos-interbred-and-exchanged-genes/ |url-status=live }} It is believed that genetic admixture has occurred at least two times within the past 550,000 years.{{Cite web |date=2022-04-06 |title=Ancient interbreeding between chimpanzees and bonobos |url=https://cnag.crg.eu/news/ancient-interbreeding-between-chimpanzees-and-bonobos |access-date=2022-04-06 |website=cnag.crg.eu |archive-date=2020-12-31 |archive-url=https://web.archive.org/web/20201231062259/https://www.cnag.crg.eu/news/ancient-interbreeding-between-chimpanzees-and-bonobos |url-status=dead }} In modern times hybridization between bonobos and chimpanzees in the wild is prevented as populations are allopatric and kept isolated on different sides of the Congo river.{{Cite web |last=Jewett |first=Katie |date=2017-04-25 |title=The Great Divide |url=https://www.biographic.com/the-great-divide/#:~:text=The%20Congo%20River%20separates%20the,this%2C%20the%20world's%20deepest%20river. |access-date=2022-04-06 |website=www.biographic.com |archive-date=2022-05-23 |archive-url=https://web.archive.org/web/20220523204316/https://www.biographic.com/the-great-divide/#:~:text=The%20Congo%20River%20separates%20the,this%2C%20the%20world's%20deepest%20river. |url-status=live }}

Within captivity, hybrids between bonobos and chimpanzees have been recorded. Between 1990 and 1992, five pregnancies were conceived and studied between a male bonobo and two female chimpanzees. The two initial pregnancies were aborted because of environmental stressors. The following three pregnancies however led to the birth of three hybrid offspring.{{Cite journal |last1=Vervaecke |first1=Hilde |last2=Elsacker |first2=L. van |date=1992 |title=Hybrids between common chimpanzees (Pan troglodytes) and pygmy chimpanzees (Pan paniscus) in captivity |journal=Mammalia |volume=56 |issue=4 |pages=667–669 |issn=1864-1547 |access-date=2023-12-01 |url=https://www.researchgate.net/publication/272356540 |archive-date=2024-03-09 |archive-url=https://web.archive.org/web/20240309175202/https://www.researchgate.net/publication/272356540_Hybrids_between_common_chimpanzees_Pan_troglodytes_and_pygmy_chimpanzees_Pan_paniscus_in_captivity |url-status=live }}

A bonobo and chimpanzee hybrid called Tiby was also featured in the 2017 Swedish film The Square.{{Cite book |last=Foundation |first=Arcus |title=Killing, Capture, Trade and Ape Conservation: Volume 4 |publisher=Cambridge University Press |date=2021-04-29 |isbn=9781108487948 |pages=111}} This same male bonobo and female chimpanzee had several offspring.Bonobo-chimpanzee Hybrids, Pan paniscus × Pan troglodytes, "EUGENE M. MCCARTHY, PHD GENETICS, ΦΒΚ", https://www.macroevolution.net/bonobo-chimpanzee-hybrids.htmlPan continuity: bonobo-chimpanzee hybrids., Hilde Vervaecke1,2, Jeroen Stevens1,2 Linda Van Elsacker 1,2, "1 University of Antwerp, Department of Biology, Universiteitsplein 1, B-2610 Wilrijk, Belgium, bisonobo@skynet.be.", "2 Royal Zoological Society of Antwerp, Centre for Research and Conservation, K. Astridplein 26, B-2018 Antwerp, Belgium.", https://lirias.kuleuven.be/retrieve/518289

File:Pan paniscus (female).jpg

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 {{convert|34|to|60|kg|lb|abbr=on}} with an average weight of {{Convert|45|kg|lb}} in males against an average of {{convert|33|kg|lb|abbr=on}} in females.{{Cite book|last=Kingdon|first=Jonathan|title=Mammals of Africa: Volume II|publisher=Bloomsbury Publishing|year=2013|pages=69}} The total length of bonobos (from the nose to the rump while on all fours) is {{convert|70|to|83|cm|in|abbr=on}}.{{cite journal | vauthors = Scholz MN, D'Août K, Bobbert MF, Aerts P | title = Vertical jumping performance of bonobo (Pan paniscus) suggests superior muscle properties | journal = Proceedings. Biological Sciences | volume = 273 | issue = 1598 | pages = 2177–84 | date = September 2006 | pmid = 16901837 | pmc = 1635523 | doi = 10.1098/rspb.2006.3568 }}{{cite web |url=http://www.arkive.org/bonobo/pan-paniscus/ |title=Bonobo videos, photos and facts – Pan paniscus |publisher=ARKive |access-date=2012-08-15 |archive-url=https://web.archive.org/web/20120825024053/http://www.arkive.org/bonobo/pan-paniscus/ |archive-date=2012-08-25 |url-status=dead }}{{cite book | veditors = Burnie D, Wilson DE | date = 2005 | title = Animal: The Definitive Visual Guide to the World's Wildlife | publisher = DK Adult | isbn = 0-7894-7764-5 }}{{cite book | vauthors = Novak RM | date = 1999 | title = Walker's Mammals of the World | edition = 6th | publisher = Johns Hopkins University Press | location = Baltimore | isbn = 0-8018-5789-9 }} Male bonobos average {{convert|119|cm|ft|abbr=on}} when standing upright, compared to {{Convert|111|cm|ft}} in females.{{Cite journal|last=Coolidge, Shea|title=External body dimensions of Pan paniscus and Pan troglodytes chimpanzees|url=https://www.scholars.northwestern.edu/en/publications/external-body-dimensions-of-pan-paniscus-and-pan-troglodytes-chim|journal=Primates|year=1982|volume=23|issue=2|pages=245–251|doi=10.1007/BF02381164|s2cid=27818900|access-date=2021-05-03|archive-date=2021-06-02|archive-url=https://web.archive.org/web/20210602215546/https://www.scholars.northwestern.edu/en/publications/external-body-dimensions-of-pan-paniscus-and-pan-troglodytes-chim|url-status=live}} 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.

File:Bonobos Kanzi and Panbanisha with Sue Savage-Rumbaugh.jpg (C) and Panbanisha (R) with Sue Savage-Rumbaugh and the outdoor symbols "keyboard"]]

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,{{cite journal | vauthors = Doran DM | title = Comparative locomotor behavior of chimpanzees and bonobos: the influence of morphology on locomotion | journal = American Journal of Physical Anthropology | volume = 91 | issue = 1 | pages = 83–98 | date = May 1993 | pmid = 8512056 | doi = 10.1002/ajpa.1330910106 }} 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.{{cite journal | vauthors = D'Août K, Vereecke E, Schoonaert K, De Clercq D, Van Elsacker L, Aerts P | title = Locomotion in bonobos (Pan paniscus): differences and similarities between bipedal and quadrupedal terrestrial walking, and a comparison with other locomotor modes | journal = Journal of Anatomy | volume = 204 | issue = 5 | pages = 353–61 | date = May 2004 | pmid = 15198700 | pmc = 1571309 | doi = 10.1111/j.0021-8782.2004.00292.x }} 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,{{cite book | vauthors = Wiessner PW, Wiessner P, Schiefenhövel W |title=Food and the status quest: an interdisciplinary perspective |date=1996 |location=Providence | publisher = Berghahn Books |isbn=1-57181-871-5 | url = https://books.google.com/books?id=Hp7dOJzVCeIC | page = 50 | quote = ...twenty-two mature community members (eight males, fourteen females)could be identified using 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.{{cite journal | vauthors = Shea BT | title = Paedomorphosis and neoteny in the pygmy chimpanzee | journal = Science | volume = 222 | issue = 4623 | pages = 521–2 | date = November 1983 | pmid = 6623093 | doi = 10.1126/science.6623093 | bibcode = 1983Sci...222..521S }} Other researchers challenged this conclusion.{{cite journal | vauthors = Godfrey LR, Sutherland MR | title = Paradox of peramorphic paedomorphosis: heterochrony and human evolution | journal = American Journal of Physical Anthropology | volume = 99 | issue = 1 | pages = 17–42 | date = January 1996 | pmid = 8928718 | doi = 10.1002/ajpa.1330990102 }}

Primatologist Frans de Waal states bonobos are capable of altruism, compassion, empathy, kindness, patience, and sensitivity,{{cite book|url=https://archive.org/details/bonoboforgottena0000waal|title=Bonobo: The Forgotten Ape|vauthors=de Waal F, Lanting F|publisher=University of California Press|year=1997|isbn=978-0-520-20535-2|url-access=registration}} and described "bonobo society" as a "gynecocracy" {{cite book | vauthors = de Waal F | title = The Bonobo and the Atheist: In Search of Humanism Among the Primates | publisher = W. W. Norton | edition = 1st | year = 2013 | isbn = 978-0-393-07377-5 | page = [https://archive.org/details/stormhasmanyeyes00lodg/page/78 78] | url = https://archive.org/details/stormhasmanyeyes00lodg/page/78 }}{{Efn|Gynecocracy among people, 'women's government over women and men' or 'women's social supremacy'}} (i.e. a matriarchy). 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.{{cite journal | vauthors = Furuichi T | title = Female contributions to the peaceful nature of bonobo society | journal = Evolutionary Anthropology | volume = 20 | issue = 4 | pages = 131–42 | year = 2011 | pmid = 22038769 | doi = 10.1002/evan.20308 | s2cid = 17830996 }}

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.{{Cite journal | doi = 10.1086/204757| title = The Social Behavior of Chimpanzees and Bonobos: Empirical Evidence and Shifting Assumptions| journal = Current Anthropology| volume = 39| issue = 4| pages = 399–420| year = 1998| vauthors = Stanford CB | s2cid = 8452514}} 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.{{cite journal | vauthors = Wilson ML, Boesch C, Fruth B, Furuichi T, Gilby IC, Hashimoto C, Hobaiter CL, Hohmann G, Itoh N, Koops K, Lloyd JN, Matsuzawa T, Mitani JC, Mjungu DC, Morgan D, Muller MN, Mundry R, Nakamura M, Pruetz J, Pusey AE, Riedel J, Sanz C, Schel AM, Simmons N, Waller M, Watts DP, White F, Wittig RM, Zuberbühler K, Wrangham RW | display-authors = 6 | title = Lethal aggression in Pan is better explained by adaptive strategies than human impacts | journal = Nature | volume = 513 | issue = 7518 | pages = 414–7 | date = September 2014 | pmid = 25230664 | doi = 10.1038/nature13727 | s2cid = 4449515 | bibcode = 2014Natur.513..414W | url = https://lib.dr.iastate.edu/anthr_pubs/13 | hdl = 10023/6258 | hdl-access = free | access-date = 2021-04-04 | archive-date = 2021-06-13 | archive-url = https://web.archive.org/web/20210613132947/https://lib.dr.iastate.edu/anthr_pubs/13/ | url-status = live }} Bonobos warn each other of danger less efficiently than chimpanzees in the same situation.{{Cite web|title=In the wild, chimpanzees are more motivated to cooperate than bonobos|url=https://phys.org/news/2020-06-wild-chimpanzees-cooperate-bonobos.html|access-date=2020-06-24|website=phys.org|archive-date=2023-03-13|archive-url=https://web.archive.org/web/20230313033221/https://phys.org/news/2020-06-wild-chimpanzees-cooperate-bonobos.html|url-status=live}}

Nonetheless, on 12 April 2024, biologists reported that bonobos behave more aggressively than thought earlier.{{cite news |last=Zimmer |first=Carl |authorlink=Carl Zimmer |title=No 'Hippie Ape': Bonobos Are Often Aggressive, Study Finds - Despite their peaceful reputation, bonobos act aggressively more often than their chimpanzee cousins, a new study found. |url=https://www.nytimes.com/2024/04/12/science/bonobo-chimpanzee-aggression.html |date=12 April 2024 |work=The New York Times |url-status=live |archive-url=https://archive.today/20240413053352/https://www.nytimes.com/2024/04/12/science/bonobo-chimpanzee-aggression.html |archive-date=13 April 2024 |access-date=13 April 2024}}{{cite journal |author=Mouginot, Maud |display-authors=et al. |title=Differences in expression of male aggression between wild bonobos and chimpanzees |date=12 April 2024 |journal=Cell |volume=34 |issue=8 |pages=1780–1785.e4 |doi=10.1016/j.cub.2024.02.071 |doi-access=free|pmid=38614078 |pmc=11167569 |bibcode=2024CBio...34.1780M }}

In a study published in February 2025, scientists determined that bonobos could tell when humans didn't know something.{{Cite journal |last1=Townrow |first1=Luke A. |last2=Krupenye |first2=Christopher |date=2025-02-11 |title=Bonobos point more for ignorant than knowledgeable social partners |journal=Proceedings of the National Academy of Sciences |volume=122 |issue=6 |pages=e2412450122 |doi=10.1073/pnas.2412450122 |pmc=11831142 |pmid=39899718|bibcode=2025PNAS..12212450T }} The findings advance researchers' proposal that like humans, chimpanzees and bonobos–humans’ closest evolutionary cousins–may also possess theory of mind.{{Cite web |last=Tamisiea |first=Jack |title=Bonobos Can Tell When a Human Doesn't Know Something |url=https://www.scientificamerican.com/article/bonobos-can-tell-when-a-human-doesnt-know-something/ |access-date=2025-02-25 |website=Scientific American |language=en}}

File:6 bonobos WHCalvin IMG 1341.jpg

File:A Bonobo at the San Diego Zoo "fishing" for termites.jpg

Bonobos are unusual among apes for their matriarchal social structure (extensive overlap between the male and female hierarchies leads some to refer to them as gender-balanced in their power structure). Bonobos do not have a defined territory and communities will travel over a wide range. Because of the nomadic nature of the females and evenly distributed food in their environment, males do not gain any obvious advantages by forming alliances with other males, or by defending a home range, as chimpanzees do. Female bonobos possess sharper canines than female chimpanzees, further fueling their status in the group.{{Cite book|chapter=The Other Sister, Bonobos: The Monkey Convergence Hypothesis|date=2020|title=Chimpanzee: Lessons from our Sister Species|pages=499–516|editor-last=Hunt|editor-first=Kevin D.|place=Cambridge|publisher=Cambridge University Press|doi=10.1017/9781316339916.029|isbn=978-1-107-11859-1|s2cid=236795730}} Although a male bonobo is dominant to a female in a dyadic interaction,{{Cite journal|last1=White|first1=Frances J.|last2=Wood|first2=Kimberley D.|date=August 2007|title=Female feeding priority in bonobos, Pan paniscus, and the question of female dominance|journal=American Journal of Primatology|volume=69|issue=8|pages=837–850|doi=10.1002/ajp.20387|pmid=17358018|s2cid=17628292}} depending on the community, socially-bonded females may be co-dominant with males{{cite journal |vauthors=Paoli T, Palagi E, Tarli SM |title=Reevaluation of dominance hierarchy in bonobos (Pan paniscus) |journal=American Journal of Physical Anthropology |volume=130 |issue=1 |pages=116–22 |date=May 2006 |pmid=16353224 |doi=10.1002/ajpa.20345}} or dominant over them, even to the extent that females can coerce reluctant males into mating with them.{{Cite web|title=What Bonobos Can Teach Us About Sexual Assault|url=https://www.theatlantic.com/ideas/archive/2018/10/are-men-wired-dominate-women-bonobos-suggest-not/571957/|access-date=2021-11-17|website=The Atlantic|date=3 October 2018|archive-date=2023-02-26|archive-url=https://web.archive.org/web/20230226031725/https://www.theatlantic.com/ideas/archive/2018/10/are-men-wired-dominate-women-bonobos-suggest-not/571957/|url-status=live}}{{cite journal | vauthors = Surbeck M, Hohmann G | title = Intersexual dominance relationships and the influence of leverage on the outcome of conflicts in wild bonobos (Pan paniscus). | journal = Behavioral Ecology and Sociobiology | date = November 2013 | volume = 67 | issue = 11 | pages = 1767–80 | doi = 10.1007/s00265-013-1584-8 | bibcode = 2013BEcoS..67.1767S | s2cid = 15709567}}

• {{cite press release |date=July 15, 2013 |title=Attractive and successful: In bonobos, attractive females are more likely to win conflicts against males |website=ScienceDaily |url=https://www.sciencedaily.com/releases/2013/07/130715105206.htm |access-date=April 5, 2021 |archive-date=February 11, 2021 |archive-url=https://web.archive.org/web/20210211205306/http://www.sciencedaily.com/releases/2013/07/130715105206.htm |url-status=live }}

At the top of the hierarchy is a coalition of high-ranking females and males typically headed by an old, experienced matriarch{{Cite web|vauthors=Pallardy R|date=2012-05-21|title=The Scandalous Social Lives of Bonobos|url=https://www.britannica.com/explore/savingearth/the-scandalous-social-lives-of-bonobos|access-date=2021-01-13|website=Saving Earth {{!}} Encyclopedia Britannica|archive-date=2022-11-12|archive-url=https://web.archive.org/web/20221112144759/https://www.britannica.com/explore/savingearth/the-scandalous-social-lives-of-bonobos|url-status=live}} who acts as the decision-maker and leader of the group. Female bonobos typically earn their rank through experience, age, and ability to forge alliances with other females in their group, rather than physical intimidation, and top-ranking females will protect immigrant females from male harassment.{{Cite web|vauthors=Sivasubramanian S|title=Feminist bonobos are taking a stand against male aggression|url=https://www.sbs.com.au/topics/science/nature/article/2016/07/27/feminist-bonobos-are-taking-stand-against-male-aggression|access-date=2021-03-12|website=Topics|archive-date=2023-02-26|archive-url=https://web.archive.org/web/20230226031727/https://www.sbs.com.au/topics/science/nature/article/2016/07/27/feminist-bonobos-are-taking-stand-against-male-aggression|url-status=dead}} While bonobos are often called matriarchal, and while every community is dominated by a female, some males will still obtain a high rank and act as coalitionary partners to the alpha female,{{Cite web|date=2007-08-13|title=Zoo Story|url=https://www.milwaukeemag.com/ZooStory/|access-date=2021-12-06|website=Milwaukee Magazine|archive-date=2023-02-26|archive-url=https://web.archive.org/web/20230226031723/https://www.milwaukeemag.com/ZooStory/|url-status=live}} often taking initiative in coordinating the group's movements. These males may outrank not only the other males in the group, but also many females.{{Cite web|date=2007-08-13|title=Zoo Story|url=https://www.milwaukeemag.com/ZooStory/|access-date=2021-05-07|website=Milwaukee Magazine|archive-date=2023-02-26|archive-url=https://web.archive.org/web/20230226031723/https://www.milwaukeemag.com/ZooStory/|url-status=live}} Certain males alert the group to any possible threats, protecting the group from predators such as pythons and leopards.{{Cite web|vauthors=Raffaele P|title=The Smart and Swinging Bonobo|url=https://www.smithsonianmag.com/science-nature/the-smart-and-swinging-bonobo-134784867/|access-date=2020-09-07|website=Smithsonian Magazine|archive-date=2023-03-30|archive-url=https://web.archive.org/web/20230330220205/https://www.smithsonianmag.com/science-nature/the-smart-and-swinging-bonobo-134784867/|url-status=live}}{{Cite journal|last1=Corredor-Ospina|first1=Nicolas|last2=Kreyer|first2=Melodie|last3=Rossi|first3=Giulia|last4=Hohmann|first4=Gottfried|last5=Fruth|first5=Barbara|date=2021-07-01|title=First report of a leopard (Panthera pardus)–bonobo (Pan paniscus) encounter at the LuiKotale study site, Democratic Republic of the Congo|journal=Primates|volume=62|issue=4|pages=555–562|doi=10.1007/s10329-021-00897-8|pmc=8225524|pmid=33950405}}

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.{{cite book | vauthors = White F | date = 1996 | chapter = Comparative socio-ecology of Pan paniscus | pages = 29–41 | veditors = McGrew WC, Marchant LF, Nishida T | title = Great ape societies | location = Cambridge, England | publisher = Cambridge Univ Press | isbn = 0-521-55536-1 }} 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.{{cite web | vauthors = Nicholls H | date = 17 March 2016 | url = http://www.bbc.com/earth/story/20160317-do-bonobos-really-spend-all-their-time-having-sex | title = Do bonobos really spend all their time having sex? | work = BBC | access-date = 11 October 2016 | archive-date = 7 October 2017 | archive-url = https://web.archive.org/web/20171007230355/http://www.bbc.com/earth/story/20160317-do-bonobos-really-spend-all-their-time-having-sex | url-status = live }} Relationships between different communities are often positive and affiliative, and bonobos are not a territorial species.{{cite journal | vauthors = Furuichi T | title = Female contributions to the peaceful nature of bonobo society | journal = Evolutionary Anthropology | volume = 20 | issue = 4 | pages = 131–42 | date = July 2011 | pmid = 22038769 | doi = 10.1002/evan.20308 | s2cid = 17830996 }} Bonobos will also share food with others, even unrelated strangers.{{cite journal | last1= Hare |first1=B |last2=Kwetuenda |first2=S | title = Bonobos voluntarily share their own food with others | journal = Current Biology | volume = 20 | issue = 5 | pages = R230-1 | date = March 2010 | pmid = 20219170 | doi = 10.1016/j.cub.2009.12.038 | s2cid = 28319610 | doi-access = free |bibcode=2010CBio...20.R230H }} Bonobos exhibit paedomorphism (retaining infantile physical characteristics and behaviours),{{cite journal | vauthors = Wobber V, Wrangham R, Hare B | title = Bonobos exhibit delayed development of social behavior and cognition relative to chimpanzees | journal = Current Biology | volume = 20 | issue = 3 | pages = 226–30 | date = February 2010 | pmid = 20116251 | doi = 10.1016/j.cub.2009.11.070 | s2cid = 3398517 | url = http://nrs.harvard.edu/urn-3:HUL.InstRepos:5270286 | doi-access = free | bibcode = 2010CBio...20..226W }} which greatly inhibits aggression and enables unfamiliar bonobos to freely mingle and cooperate with each other.{{Cite web|vauthors=Callaway E|title=Peter Pan ways make bonobos the most amiable of apes|url=https://www.newscientist.com/article/dn18443-peter-pan-ways-make-bonobos-the-most-amiable-of-apes/|access-date=2021-03-14|website=New Scientist|archive-date=2023-03-13|archive-url=https://web.archive.org/web/20230313035354/https://www.newscientist.com/article/dn18443-peter-pan-ways-make-bonobos-the-most-amiable-of-apes/|url-status=live}}

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.{{cite journal | vauthors = Surbeck M, Deschner T, Schubert G, Weltring A, Hohmann G | title = Mate competition, testosterone and intersexual relationships in bonobos, Pan paniscus | journal = Animal Behaviour | date = March 2012 | volume = 83 | issue = 3 | pages = 659–69 | doi = 10.1016/j.anbehav.2011.12.010 | s2cid = 53198728}}

• {{cite web |date=January 23, 2012 |title=Bonobos' unusual success story: Dominant males invest in friendly relationships with females |website=Max-Planck-Gesellschaft |url=https://www.mpg.de/4994603/Bonobos_testosterone |access-date=February 26, 2021 |archive-date=March 26, 2021 |archive-url=https://web.archive.org/web/20210326204542/https://www.mpg.de/4994603/Bonobos_testosterone |url-status=live }} Aging bonobos lose their playful streak and become noticeably more irritable in old age. Both sexes have a similar level of aggressiveness.{{Cite web|vauthors=Hogenboom M|title=First personality test shows that female apes are irritable|url=https://www.bbc.com/earth/story/20160622-why-female-apes-are-irritable|access-date=2021-03-12|website=www.bbc.com|archive-date=2021-03-31|archive-url=https://web.archive.org/web/20210331175649/http://www.bbc.com/earth/story/20160622-why-female-apes-are-irritable|url-status=live}} 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.{{Citation| vauthors = Furuichi T, Idani GI, Ihobe H, Hashimoto C, Tashiro Y, Sakamaki T, Mulavwa MN, Yangozene K, Kuroda S |title=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|date=2012|work=Long-Term Field Studies of Primates|pages=413–433| veditors = Kappeler PM, Watts DP |place=Berlin, Heidelberg|publisher=Springer|doi=10.1007/978-3-642-22514-7_18|isbn=978-3-642-22514-7}} Additionally, females with powerful mothers may remain in their natal clan.{{Cite web|title=Bonobo Alive {{!}} News|url=https://www.bonobo-alive.org/files/news.html|access-date=2021-11-21|website=www.bonobo-alive.org|archive-date=2023-03-13|archive-url=https://web.archive.org/web/20230313035350/https://www.bonobo-alive.org/files/news.html|url-status=dead}}

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.{{Cite web|vauthors=Jirik K|title=LibGuides: Bonobo (Pan paniscus) Fact Sheet: Behavior & Ecology|url=https://ielc.libguides.com/sdzg/factsheets/bonobo/behavior|access-date=2021-01-13|website=ielc.libguides.com|archive-date=2023-05-27|archive-url=https://web.archive.org/web/20230527233818/https://ielc.libguides.com/sdzg/factsheets/bonobo/behavior|url-status=live}} There is a confirmed case of a grown male bonobo adopting his orphaned infant brother.{{Cite journal |last1=Surbeck |first1=Martin |last2=Hohmann |first2=Gottfried |title=Affiliations, aggressions and an adoption: Male–male relationships in wild bonobos |journal=Oxford Scholarship |year=2018 |pages=35–46 |doi=10.1093/oso/9780198728511.003.0003|isbn=978-0-19-872851-1 }} 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.{{cite journal | vauthors = Surbeck M, Mundry R, Hohmann G | title = Mothers matter! Maternal support, dominance status and mating success in male bonobos (Pan paniscus) | journal = Proceedings. Biological Sciences | volume = 278 | issue = 1705 | pages = 590–8 | date = February 2011 | pmid = 20810444 | pmc = 3025686 | doi = 10.1098/rspb.2010.1572}}

• {{cite web |date=September 1, 2010 |title=High social status, maternal support play important role in mating success of male bonobos |website=Phys.org |url=https://phys.org/news/2010-09-high-social-status-maternal-important.html |access-date=September 28, 2020 |archive-date=October 28, 2020 |archive-url=https://web.archive.org/web/20201028185326/https://phys.org/news/2010-09-high-social-status-maternal-important.html |url-status=live }} 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.{{Cite web|vauthors=Sample I|date=2019-05-20|title=Pushy bonobo mothers help sons find sexual partners, scientists find|url=http://www.theguardian.com/science/2019/may/20/pushy-bonobo-mothers-help-sons-find-sexual-partners-scientists-find|access-date=2021-01-13|website=The Guardian|archive-date=2023-03-29|archive-url=https://web.archive.org/web/20230329024607/https://www.theguardian.com/science/2019/may/20/pushy-bonobo-mothers-help-sons-find-sexual-partners-scientists-find|url-status=live}} 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.{{Cite web|title=The Momma's Boy Strategy: Why Bonobo Males Tend Not To Form Coalitions|url=https://traditionsofconflict.com/blog/2018/12/18/the-mommas-boy-strategy-why-bonobo-males-tend-not-to-form-coalitions|access-date=2021-05-07|website=Traditions of Conflict|date=18 December 2018|archive-date=2021-05-10|archive-url=https://web.archive.org/web/20210510142819/https://traditionsofconflict.com/blog/2018/12/18/the-mommas-boy-strategy-why-bonobo-males-tend-not-to-form-coalitions|url-status=live}}

Female bonobos have also been observed fostering infants from outside their established community.{{cite journal |vauthors=Tokuyama N, Toda K, Poiret ML, Iyokango B, Bakaa B, Ishizuka S |title=Two wild female bonobos adopted infants from a different social group at Wamba |journal=Scientific Reports |volume=11 |issue=1 |pages=4967 |date=March 2021 |pmid=33737517 |pmc=7973529 |doi=10.1038/s41598-021-83667-2 |bibcode=2021NatSR..11.4967T}}

• {{cite web |date=March 18, 2021 |title=Female wild bonobos provide care for infants outside their social group |website=Phys.org |url=https://phys.org/news/2021-03-female-wild-bonobos-infants-social.html |access-date=March 19, 2021 |archive-date=March 19, 2021 |archive-url=https://web.archive.org/web/20210319065122/https://phys.org/news/2021-03-female-wild-bonobos-infants-social.html |url-status=live }}{{cite journal | vauthors = Tokuyama N, Toda K, Poiret ML, Iyokango B, Bakaa B, Ishizuka S | title = Two wild female bonobos adopted infants from a different social group at Wamba | journal = Scientific Reports | volume = 11 | issue = 1 | pages = 4967 | date = March 2021 | pmid = 33737517 | pmc = 7973529 | doi = 10.1038/s41598-021-83667-2 | bibcode = 2021NatSR..11.4967T }} Bonobos are not known to kill each other, and are generally less violent than chimpanzees, yet aggression still manifests itself in this species. Although female bonobos dominate males and selectively mate with males who do not exhibit aggression toward them, competition between the males themselves is intense and high-ranking males secure more matings than low-ranking ones.{{cite journal |vauthors=Stevens J, Vervaecke H, de Vries H, van Elsacker L |date=2 October 2007 |title=Sex Differences in the Steepness of Dominance Hierarchies in Captive Bonobo Groups |journal=International Journal of Primatology |volume=28 |issue= 6|pages=1417–1430 |doi=10.1007/s10764-007-9186-9 |s2cid=27240321 |url=https://www.researchgate.net/publication/227156283}} Indeed, the size difference between males and females is more pronounced in bonobos than it is in chimpanzees, as male bonobos do not form alliances and therefore have little incentive to hold back when fighting for access to females.{{cite book |vauthors=Hunt K |year=2020 |chapter=The Other Sister, Bonobos: The Monkey Convergence Hypothesis |title=Chimpanzee: Lessons from our Sister Species |pages=499–516 |location=Cambridge |publisher=Cambridge University Press |url=https://www.researchgate.net/publication/342897635 |doi=10.1017/9781316339916.029|s2cid=236795730 }} 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 a female mob.{{Cite web|vauthors=Clint E|date=2017-10-09|title=Faux-nobo: "Naked Bonobo" demolishes myth of sexy, egalitarian bonobos|website=Incredulous|url=https://skepticink.com/incredulous/2017/10/09/bonobo-myth-demolished/|access-date=2020-12-18|archive-date=2023-05-25|archive-url=https://web.archive.org/web/20230525223219/https://skepticink.com/incredulous/2017/10/09/bonobo-myth-demolished/|url-status=live}}

Because of 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.{{cite web | vauthors = Cawthon Lang KA | date = December 2010 | url = http://pin.primate.wisc.edu/factsheets/entry/bonobo/behav | work = Primate Factsheets | title = Bonobo (Pan paniscus) behavior | publisher = University of Wisconsin | archive-url = https://web.archive.org/web/20160412150955/http://pin.primate.wisc.edu/factsheets/entry/bonobo/behav | archive-date = 2016-04-12 }} 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.{{Citation|title=16 The Absence of Sexual Coercion in Bonobos|date=2009-12-31|work=Sexual Coercion in Primates and Humans|pages=410–423|publisher=Harvard University Press|doi=10.4159/9780674054349-016|isbn=978-0-674-05434-9}}

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, and 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.{{cite journal | vauthors = White FJ, Wood KD | title = Female feeding priority in bonobos, Pan paniscus, and the question of female dominance | journal = American Journal of Primatology | volume = 69 | issue = 8 | pages = 837–50 | date = August 2007 | pmid = 17358018 | doi = 10.1002/ajp.20387 | s2cid = 17628292 }} 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.{{Cite journal|last1=Samuni|first1=Liran|last2=Wegdell|first2=Franziska|last3=Surbeck|first3=Martin|date=2020-09-01|editor-last=Weigel|editor-first=Detlef|editor2-last=Van de Waal|editor2-first=Erica|editor3-last=Van de Waal|editor3-first=Erica|title=Behavioural diversity of bonobo prey preference as a potential cultural trait|journal=eLife|volume=9|pages=e59191|doi=10.7554/eLife.59191|pmid=32869740|pmc=7462605 |doi-access=free }} 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.{{Cite web|vauthors=Nicholls H|title=Do bonobos really spend all their time having sex?|url=https://www.bbc.com/earth/story/20160317-do-bonobos-really-spend-all-their-time-having-sex|access-date=2020-08-08|website=www.bbc.com|archive-date=2020-08-12|archive-url=https://web.archive.org/web/20200812160013/http://www.bbc.com/earth/story/20160317-do-bonobos-really-spend-all-their-time-having-sex|url-status=live}} 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.{{Cite web|vauthors=Jones N|date=2018-04-05|title=Bonobos Spied Sharing a Feast|url=https://www.sapiens.org/evolution/bonobos-meal-sharing/|access-date=2020-08-08|website=SAPIENS|archive-date=2020-06-24|archive-url=https://web.archive.org/web/20200624180404/https://www.sapiens.org/evolution/bonobos-meal-sharing/|url-status=live}} 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.{{cite journal | vauthors = Wakefield ML, Hickmott AJ, Brand CM, Takaoka IY, Meador LM, Waller MT, White FJ | title = New Observations of Meat Eating and Sharing in Wild Bonobos (Pan paniscus) at Iyema, Lomako Forest Reserve, Democratic Republic of the Congo | journal = Folia Primatologica; International Journal of Primatology | volume = 90 | issue = 3 | pages = 179–189 | date = 2019 | pmid = 30889597 | doi = 10.1159/000496026 | s2cid = 84183655}}

• {{cite web |author= |date=2019-05-01 |title=Bonobos Eat and Share Meat at Rates Similar to Chimpanzees |website=The Leakey Foundation |url=https://leakeyfoundation.org/bonobos-eat-and-share-meat-at-rates-similar-to-chimpanzees/ |access-date=2020-10-05 |archive-date=2020-10-08 |archive-url=https://web.archive.org/web/20201008081630/https://leakeyfoundation.org/bonobos-eat-and-share-meat-at-rates-similar-to-chimpanzees/ |url-status=live }}

In a study published in November 2023, scientists reported, for the first time, evidence that groups of primates, particularly bonobos, are capable of cooperating with each other.{{cite news |last=Zimmer |first=Carl |authorlink=Carl Zimmer |title=Scientists Find First Evidence That Groups of Apes Cooperate - Some bonobos are challenging the notion that humans are the only primates capable of group-to-group alliances. |url=https://www.nytimes.com/2023/11/16/science/bonobos-cooperation-study.html |date=16 November 2023 |work=The New York Times |url-status=live |archive-url=https://archive.today/20231116194259/https://www.nytimes.com/2023/11/16/science/bonobos-cooperation-study.html |archive-date=16 November 2023 |access-date=17 November 2023}}{{cite journal |author=Samuni, Liran |display-authors=et al. |title=Cooperation across social borders in bonobos |url=https://www.science.org/doi/10.1126/science.adg0844 |date=16 November 2023 |journal=Science |volume=382 |issue=6672 |pages=805–809 |doi=10.1126/science.adg0844 |pmid=37972165 |bibcode=2023Sci...382..805S |s2cid=265221570 |url-status=live |archive-url=https://archive.today/20231117125744/https://www.science.org/doi/10.1126/science.adg0844 |archive-date=17 November 2023 |access-date=17 November 2023}} Researchers observed unprecedented cooperation between two distinct bonobo groups in the Congo's Kokolopori Bonobo Reserve, Ekalakala and Kokoalongo, challenging traditional notions of ape societies. Over two years of observation, researchers witnessed 95 encounters between the groups. Contrary to expectations, these interactions resembled those within a single group. During these encounters, the bonobos engaged in behaviors such as grooming, food sharing, and collective defense against threats like snakes. Notably, the two groups, while displaying cooperative tendencies, maintained distinct identities, and there was no evidence of interbreeding or a blending of cultures. The cooperation observed was not arbitrary but evolved through individual bonds formed by exchanging favors and gifts. Some bonobos even formed alliances to target a third individual, demonstrating a nuanced social dynamic within the groups.

{{See also|Animal sexual behaviour#Genital-genital rubbing|Homosexual behavior in animals#Bonobos}}

File:Bonobo sexual behavior 1.jpg.]]

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.{{cite web | url = http://pubpages.unh.edu/~jel/512/aggression-01.html | archive-url = https://web.archive.org/web/20050320135050/http://pubpages.unh.edu/~jel/512/aggression-01.html | archive-date = 20 March 2005 | title = Aggression topics | publisher = University of New Hampshire }} Bonobos are the only non-human animal to have been observed engaging in tongue kissing.{{cite journal| vauthors = Manson JH, Perry S, Parish AR |year = 1997|title = Nonconceptive Sexual Behavior in Bonobos and Capuchins|journal = International Journal of Primatology|volume = 18|issue = 5|pages = 767–86 |doi = 10.1023/A:1026395829818|s2cid = 3032455 }} Bonobos and humans are the only primates to typically engage in face-to-face genital sex, although a pair of western gorillas has also been photographed in this position.{{cite web | vauthors = Nguyen TC | date = 2008-02-13 | url = http://www.livescience.com/2298-gorillas-caught-human-act.html | title = Gorillas Caught in Very Human Act | work = Live Science | access-date = 2011-10-01 | archive-date = 2012-07-04 | archive-url = https://web.archive.org/web/20120704074516/http://www.livescience.com/2298-gorillas-caught-human-act.html | url-status = live }}

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.{{cite journal | vauthors = de Waal FB | title = Bonobo sex and society | journal = Scientific American | volume = 272 | issue = 3 | pages = 82–8 | date = March 1995 | pmid = 7871411 | doi = 10.1038/scientificamerican0395-82 | url = http://www.biozentrum.uni-wuerzburg.de/uploads/media/Bonobo_sex_01.pdf | access-date = 21 December 2011 | url-status = dead | bibcode = 1995SciAm.272c..82W | archive-url = https://web.archive.org/web/20120127051545/http://www.biozentrum.uni-wuerzburg.de/uploads/media/Bonobo_sex_01.pdf | archive-date = 27 January 2012 | author-link = Frans de Waal }}

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;{{cite book | vauthors = Balcombe JP |author-link=Jonathan Balcombe |title=The Exultant Ark: A Pictorial Tour of Animal Pleasure |publisher=University of California Press |isbn=978-0-520-26024-5 |year=2011 |access-date=2012-11-22|page=88|url=https://books.google.com/books?id=tz9mSyTWh0oC&pg=PA88}} 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".{{cite book | vauthors = Angier N |author-link=Natalie Angier |title=Woman: An Intimate Geography |publisher=Houghton Mifflin Harcourt |isbn=978-0-395-69130-4 |year=1999 |access-date=2012-11-22|page=[https://archive.org/details/womanintimategeo00angi/page/68 68]|url=https://archive.org/details/womanintimategeo00angi|url-access=registration }} In scientific literature, the female–female behavior of bonobos pressing vulvas together is often referred to as genito-genital (GG) rubbing.{{cite journal | vauthors = Paoli T, Palagi E, Tacconi G, Tarli SB | title = Perineal swelling, intermenstrual cycle, and female sexual behavior in bonobos (Pan paniscus) | journal = American Journal of Primatology | volume = 68 | issue = 4 | pages = 333–47 | date = April 2006 | pmid = 16534808 | doi = 10.1002/ajp.20228 | s2cid = 25823290 }} 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,{{Cite book |last=Wrangham |first=Richard W. |url=https://books.google.com/books?id=IzBIHPeE45IC&pg=PA164 |title=Chimpanzee Cultures |date=1996 |publisher=Harvard University Press |isbn=978-0-674-11663-4 |language=en}} "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.{{cite journal | vauthors = Hohmann G, Fruth B | title = Use and function of genital contacts among female bonobos | journal = Animal Behaviour | volume = 60 | issue = 1 | pages = 107–120 | date = July 2000 | pmid = 10924210 | doi = 10.1006/anbe.2000.1451 | s2cid = 39702173 }}

File:Pan paniscus11.jpg

Bonobo males engage in various forms of male–male genital behavior.{{cite web|url=http://www.bio.davidson.edu/people/vecase/Behavior/Spring2004/laird/Social_Organization.htm |archive-url=https://web.archive.org/web/20110519005633/http://www.bio.davidson.edu/people/vecase/Behavior/Spring2004/laird/Social_Organization.htm |archive-date=2011-05-19 |title=Courtney Laird, "Social Organization" |publisher=Bio.davidson.edu |year=2004 |access-date=2009-07-03}} 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 because of both males' erections. In another, rarer form of genital rubbing, two bonobo males hang from a tree limb face-to-face while penis fencing.{{cite book| vauthors = de Waal FB |author-link= Frans de Waal| title=The ape and the sushi master: cultural reflections by a primatologist|publisher=Basic Books|chapter=Bonobos and Fig Leaves|year= 2001|isbn= 978-84-493-1325-7}} 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.{{cite journal | vauthors = Kitamura K |title=Genito-Genital Contacts in the Pygmy Chimpanzees (Pan paniscus) |journal=African Study Monographs |date=August 1989 |volume=10 |issue=2 |pages=49–67 |doi=10.14989/68052 |hdl=2433/68052 }} 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.{{cite web|website=BBC|url=http://www.bbc.com/earth/story/20150206-are-there-any-homosexual-animals|title=Are there any homosexual animals?|date=6 February 2015|vauthors=Hogenboom M|access-date=30 June 2017|archive-date=14 May 2021|archive-url=https://web.archive.org/web/20210514141924/http://www.bbc.com/earth/story/20150206-are-there-any-homosexual-animals|url-status=live}}

Wild females give birth for the first time at 13 or 14 years of age.{{Cite journal |last=De Waal |first=Frans B. M. |date=June 1, 2006 |title=Bonobo Sex and Society |url=https://www.scientificamerican.com/article/bonobo-sex-and-society-2006-06/ |journal=Scientific American |volume=16 |issue=2 |pages=14–21 |doi=10.1038/scientificamerican0606-14sp |access-date=June 26, 2022 |archive-date=January 4, 2023 |archive-url=https://web.archive.org/web/20230104143350/https://www.scientificamerican.com/article/bonobo-sex-and-society-2006-06/ |url-status=live }} Bonobo reproductive rates are no higher than those of the common chimpanzee. However, female bonobo oestrus periods are longer.{{Cite journal| vauthors = Ihobe H, Furuichi T |date=1994|title=Variation in Male Relationships in Bonobos and Chimpanzees|journal=Behaviour|volume=130|issue=3–4|pages=211–228|doi=10.1163/156853994x00532}} 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.{{cite web | url = http://animaldiversity.ummz.umich.edu/site/accounts/information/Pan_paniscus.html | vauthors = Williams A, Myers P | year = 2004 | title = Pan paniscus | publisher = Animal Diversity Web | access-date = 6 January 2012 | archive-date = 7 February 2011 | archive-url = https://web.archive.org/web/20110207001628/http://animaldiversity.ummz.umich.edu/site/accounts/information/Pan_paniscus.html | url-status = live }} 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,{{cite book | vauthors = de Waal FB | date = 1990 | chapter = Sociosexual behavior used for tension regulation in all age and sex combinations among bonobos | pages = 378–393 | title = Pedophilia: Biosocial Dimensions | veditors = Feierman JR | publisher = Springer | location = New York | isbn = 978-1-4613-9684-0}} although without penetration.{{cite web | vauthors = Small MF | date = 1 June 1992 | url = https://www.discovermagazine.com/mind/casual-sex-play-common-among-bonobos | title = Casual Sex Play Common Among Bonobos | work = Discover | quote = Even juveniles participate by rubbing their genital areas against adults, although ethologists don't think that males actually insert their penises into juvenile females. | access-date = 14 November 2020 | archive-date = 26 March 2023 | archive-url = https://web.archive.org/web/20230326180012/https://www.discovermagazine.com/mind/casual-sex-play-common-among-bonobos | url-status = live }} Adult females also have sex with infants, but less frequently. Infants are not passive participants. They quite often initiate contacts with both adult males and females, as well as with peers. They have also been shown to be sexually active even in the absence of any stimulation or learning from adults.{{Cite journal |last1=Woods |first1=Vanessa |last2=Hare |first2=Brian |date=2011-04-01 |title=Bonobo but not chimpanzee infants use socio-sexual contact with peers |journal=Primates |volume=52 |issue=2 |pages=111–116 |doi=10.1007/s10329-010-0229-z |pmid=21127940 |s2cid=7628992 |quote=Even as infants, bonobos use socio-sexual behavior, whereas the same behavior is completely absent in chimpanzee infants. Findings suggest that sociosexual behavior in infant bonobos serves functions during social feeding that is not closely tied to reproduction. Moreover, their sexual behavior development is not dependent on a gradual learning process facilitated by interactions with adults or by close observation of such behavior.}}

Infanticide, while well documented in chimpanzees, is apparently absent in bonobo society.{{cite journal | vauthors = Gottfried H, Vigilant L, Mundry R, Behringer V, Surbeck M | title = Aggression by male bonobos against immature individuals does not fit with predictions of infanticide | journal = Aggressive Behavior | volume = 45 | issue = 3 | pages = 300–309 | date = May 2019 | pmid = 30710459 | doi = 10.1002/ab.21819 | s2cid = 73440844 }} Although infanticide has not been directly observed, there have been documented cases of both female{{cite magazine | url=https://www.newyorker.com/magazine/2007/07/30/swingers | title=Swingers | magazine=The New Yorker | date=23 July 2007 | access-date=19 December 2018 | archive-date=28 March 2023 | archive-url=https://web.archive.org/web/20230328131137/https://www.newyorker.com/magazine/2007/07/30/swingers | url-status=live }} and male{{cite web | url=https://skepticink.com/incredulous/2014/12/29/questioning-sexy-bonobo-hype-part-2-primatologist-responds-christopher-ryan/ | title=Questioning the "sexy" bonobo hype, part 2: A primatologist debunks Christopher Ryan | date=29 December 2014 | access-date=1 July 2022 | archive-date=29 March 2023 | archive-url=https://web.archive.org/web/20230329043949/https://skepticink.com/incredulous/2014/12/29/questioning-sexy-bonobo-hype-part-2-primatologist-responds-christopher-ryan/ | url-status=live }} bonobos kidnapping infants, sometimes resulting in infants dying from dehydration. Although male bonobos have not yet been seen to practice infanticide, there is a 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.{{Cite web|last=Clint|first=Edward|date=2017-10-09|title=Faux-nobo: "Naked Bonobo" demolishes myth of sexy, egalitarian bonobos|url=https://skepticink.com/incredulous/2017/10/09/bonobo-myth-demolished/|access-date=2021-06-03|website=Incredulous|archive-date=2023-05-25|archive-url=https://web.archive.org/web/20230525223219/https://skepticink.com/incredulous/2017/10/09/bonobo-myth-demolished/|url-status=live}}

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.{{Cite journal| vauthors = Hare B, Wobber V, Wrangham R |date=March 2012|title=The self-domestication hypothesis: evolution of bonobo psychology is due to selection against aggression |journal=Animal Behaviour |volume=83 |issue=3 |pages=573–585 |doi=10.1016/j.anbehav.2011.12.007 |s2cid=3415520 }} 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.{{cite journal | vauthors = Beaudrot LH, Kahlenberg SM, Marshall AJ | title = Why male orangutans do not kill infants | journal = Behavioral Ecology and Sociobiology | volume = 63 | issue = 11 | pages = 1549–1562 | date = September 2009 | pmid = 19701484 | pmc = 2728907 | doi = 10.1007/s00265-009-0827-1 | bibcode = 2009BEcoS..63.1549B }} Bonobos engage in sexual activity numerous times a day.{{Cite web |title=Reproduction (Part of the Extreme Mammals exhibition.) | work=American Museum of Natural History |url=https://www.amnh.org/exhibitions/extreme-mammals/extreme-bodies/reproduction#:~:text=For%20the%20more%20promiscuous%20side,group%2C%20male%20and%20female%20alike |access-date=15 February 2024 |archive-date=15 February 2024 |archive-url=https://web.archive.org/web/20240215194504/https://www.amnh.org/exhibitions/extreme-mammals/extreme-bodies/reproduction#:~:text=For%20the%20more%20promiscuous%20side,group%2C%20male%20and%20female%20alike |url-status=live }}

It is unknown how the bonobo avoids simian immunodeficiency virus (SIV) and its effects.{{cite journal | vauthors = Sharp PM, Shaw GM, Hahn BH | title = Simian immunodeficiency virus infection of chimpanzees | journal = Journal of Virology | volume = 79 | issue = 7 | pages = 3891–902 | date = April 2005 | pmid = 15767392 | pmc = 1061584 | doi = 10.1128/jvi.79.7.3891-3902.2005 }}

File:Bonobo (Pan paniscus) at Lola Ya Bonobo - 3.JPG]]

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.{{cite news | date = June 24, 2010 | url = http://www.economist.com/node/16422404 | newspaper = The Economist | title = Chimpanzee behavior: Killer instincts | access-date = 2011-12-08 | archive-date = 2018-01-14 | archive-url = https://web.archive.org/web/20180114195507/http://www.economist.com/node/16422404 | url-status = live }} 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 more aggressive than chimpanzees, having higher rates of aggressive acts, about three times as much. Although, male chimpanzees are more likely to be aggressive to a lethal degree than male bonobos which are more likely to engage in more frequent, yet less intense squabbling. There is also more female to male aggression with bonobos than there is with chimpanzees.{{cite web | url=https://news.harvard.edu/gazette/story/2024/04/so-much-for-summers-of-love/#:~:text=Researchers%20observing%20bonobos%20and%20chimps,bonobos%20than%20among%20male%20chimps | title=Male bonobos fight three times as often as chimps, study finds | date=22 April 2024 }}{{cite journal |first1=Maud |last1=Mouginot |first2=Michael L. |last2=Wilson |first3=Nisarg |last3=Desai |first4=Martin |last4=Surbeck |year=2024 |title=Differences in expression of male aggression between wild bonobos and chimpanzees |journal=Current Biology |volume=34 |issue=8 |pages=1780–1785.e4|doi=10.1016/j.cub.2024.02.071 |pmid=38614078 |pmc=11167569 |bibcode=2024CBio...34.1780M }} Female bonobos are also more aggressive than female chimpanzees, in general. Both bonobos and chimpanzees exhibit physical aggression more than 100 times as often as humans do.{{cite book |vauthors=Wrangham R |title=The Goodness Paradox: The Strange Relationship Between Virtue and Violence in Human Evolution |date=2019 |publisher=Knopf Doubleday Publishing Group |isbn=978-1-101-97019-5 |pages=19–20 |url=https://books.google.com/books?id=pOi9DwAAQBAJ |access-date=2020-03-10 |archive-date=2024-03-09 |archive-url=https://web.archive.org/web/20240309175059/https://books.google.com/books?id=pOi9DwAAQBAJ |url-status=live }}

File:Grooming de Zoé.jpg

Although referred to as peaceful, bonobo aggression is not restricted to each other, and humans have also been attacked by bonobos, and suffered serious, albeit non-fatal, injuries.

Bonobos are far less violent than chimpanzees, though, as lethal aggression is essentially nonexistent among bonobos while being not infrequent among chimpanzees.

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.{{cite journal|year=1988|title=Feeding competition and patch size in the chimpanzee species Pan paniscus and Pan troglodytes|journal=Behaviour |volume=105 |issue=1/2 |pages=148–164 |doi=10.1163/156853988X00494 |jstor=4534684| vauthors = White FJ, Wrangham RW |s2cid=18285801}}

Recent studies show that there are significant brain differences between bonobos and chimpanzees. 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.{{cite journal | vauthors = Rilling JK, Scholz J, Preuss TM, Glasser MF, Errangi BK, Behrens TE | title = Differences between chimpanzees and bonobos in neural systems supporting social cognition | journal = Social Cognitive and Affective Neuroscience | volume = 7 | issue = 4 | pages = 369–79 | date = April 2012 | pmid = 21467047 | pmc = 3324566 | doi = 10.1093/scan/nsr017 }} 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.{{cite journal | vauthors = Davidson RJ, Putnam KM, Larson CL | title = Dysfunction in the neural circuitry of emotion regulation—a possible prelude to violence | journal = Science | volume = 289 | issue = 5479 | pages = 591–4 | date = July 2000 | pmid = 10915615 | doi = 10.1126/science.289.5479.591 | bibcode = 2000Sci...289..591D }}{{cite journal | vauthors = Pezawas L, Meyer-Lindenberg A, Drabant EM, Verchinski BA, Munoz KE, Kolachana BS, Egan MF, Mattay VS, Hariri AR, Weinberger DR | display-authors = 6 | title = 5-HTTLPR polymorphism impacts human cingulate-amygdala interactions: a genetic susceptibility mechanism for depression | journal = Nature Neuroscience | volume = 8 | issue = 6 | pages = 828–34 | date = June 2005 | pmid = 15880108 | doi = 10.1038/nn1463 | s2cid = 1864631 }} This thicker connection may make them better at regulating their emotional impulses and behavior.{{cite news | vauthors = Vastag B | date = 11 April 2011 | url = https://www.washingtonpost.com/national/brain-differences-may-explain-varying-behavior-of-bonobos-and-chimpanzees/2011/03/29/AFP2wUND_story.html | title = Brain differences may explain varying behavior of bonobos and chimpanzees | newspaper = The Washington Post | access-date = 26 December 2012 | archive-date = 14 February 2021 | archive-url = https://web.archive.org/web/20210214005916/https://www.washingtonpost.com/national/brain-differences-may-explain-varying-behavior-of-bonobos-and-chimpanzees/2011/03/29/AFP2wUND_story.html | url-status = live }}

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."{{cite web|vauthors=de Waal F|url=http://www.skeptic.com/eskeptic/07-08-08/|title=Bonobos, Left & Right|work=Skeptic |date=August 8, 2007|access-date=June 29, 2010|archive-date=November 21, 2016|archive-url=https://web.archive.org/web/20161121030108/http://www.skeptic.com/eskeptic/07-08-08/|url-status=live}}

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.

There is one inferred intraspecies killing in the wild,{{cite magazine |last=Parker |first=Ian |date=July 23, 2007 |title=Swingers |magazine=The New Yorker |issue=July 30, 2007 |url=https://www.newyorker.com/magazine/2007/07/30/swingers |url-access=limited |access-date=September 21, 2022 |archive-date=March 28, 2023 |archive-url=https://web.archive.org/web/20230328131137/https://www.newyorker.com/magazine/2007/07/30/swingers |url-status=live }} and a confirmed lethal attack in captivity.{{Cite web |title=Trauer um Bonobo Birogu |url=https://www.wuppertal.de/microsite/zoo/Aktuelles/20210115-trauer-um-bonobo-birogu.php |access-date=September 21, 2022 |website=Der Grüne Zoo Wuppertal |language=de |archive-date=May 26, 2022 |archive-url=https://web.archive.org/web/20220526204313/https://www.wuppertal.de/microsite/zoo/Aktuelles/20210115-trauer-um-bonobo-birogu.php |url-status=live }} In both cases, the attackers were female and the victims were male.

File:Les bonobos utilisent certaines feuilles aux vertus pharmacologique contre leurs parasites intestinaux.jpg

The bonobo is an omnivorous frugivore; 57% of its diet is fruit, but this is supplemented with leaves, honey, eggs,{{cite web|url=http://pin.primate.wisc.edu/factsheets/entry/bonobo|vauthors=Lang KC|title=Bonobo: Pan paniscus|year=2011|publisher=National Primate Research Center, University of Wisconsin – Madison|access-date=2005-11-08|archive-date=2006-03-01|archive-url=https://web.archive.org/web/20060301215912/http://pin.primate.wisc.edu/factsheets/entry/bonobo|url-status=live}} meat from small vertebrates such as anomalures, flying squirrels and duikers,{{cite journal | vauthors = Ihobe H |title=Observations on the meat-eating behavior of wild bonobos (Pan paniscus) at Wamba, Republic of Zaire |journal=Primates |volume=33 |issue=2 |pages=247–250|year=1992 |doi=10.1007/BF02382754|s2cid=10063791 }} and invertebrates.{{cite web | vauthors = Rafert J, Vineberg EO | year = 1997 | url = http://www.nagonline.net/HUSBANDRY/Diets_pdf/Bonobo_Nutrition.pdf | title = Bonobo Nutrition – relation of captive diet to wild diet | archive-url = https://web.archive.org/web/20120425232556/http://www.nagonline.net/HUSBANDRY/Diets_pdf/Bonobo_Nutrition.pdf | archive-date=2012-04-25}}" Bonobo Husbandry Manual, American Association of Zoos and Aquariums The truffle species Hysterangium bonobo is eaten by bonobos.{{cite journal |last1=Elliott |first1=Todd F. |last2=Georgiev |first2=Alexander V. |last3=Lokasola |first3=Albert Lotana |last4=Smith |first4=Matthew E. |title=Hysterangium bonobo: A newly described truffle species that is eaten by bonobos in the Democratic Republic of Congo |url=https://research.bangor.ac.uk/portal/files/35024619/Elliott_et_al._2020_Hysterangium_bonobo_REV2.pdf |journal=Mycologia |date=4 September 2020 |volume=112 |issue=6 |pages=1203–1211 |doi=10.1080/00275514.2020.1790234 |pmid=32886571 |s2cid=221503207 |access-date=7 July 2023 |archive-date=7 July 2023 |archive-url=https://web.archive.org/web/20230707162121/https://research.bangor.ac.uk/portal/files/35024619/Elliott_et_al._2020_Hysterangium_bonobo_REV2.pdf |url-status=live }} In some instances, bonobos have been shown to consume lower-order primates.{{cite journal | vauthors = Surbeck M, Fowler A, Deimel C, Hohmann G | title = Evidence for the consumption of arboreal, diurnal primates by bonobos (Pan paniscus) | journal = American Journal of Primatology | volume = 71 | issue = 2 | pages = 171–4 | date = February 2009 | pmid = 19058132 | doi = 10.1002/ajp.20634 | s2cid = 32622605 }}; {{cite journal | vauthors = Surbeck M, Hohmann G | title = Primate hunting by bonobos at LuiKotale, Salonga National Park | journal = Current Biology | volume = 18 | issue = 19 | pages = R906-7 | date = October 2008 | pmid = 18957233 | doi = 10.1016/j.cub.2008.08.040 | s2cid = 6708310 | doi-access = free | bibcode = 2008CBio...18.R906S }} Some claim bonobos have also been known to practise cannibalism in captivity, a claim disputed by others.{{cite web | vauthors = Parker I | url = http://www.newyorker.com/reporting/2007/07/30/070730fa_fact_parker?printable=true | work = Our Far-Flung Correspondents | title = Swingers | publisher = The New Yorker | date = 2007-07-30 | access-date = 2011-12-08 | archive-date = 2012-01-18 | archive-url = https://web.archive.org/web/20120118060731/http://www.newyorker.com/reporting/2007/07/30/070730fa_fact_parker?printable=true | url-status = live }}{{cite news |vauthors=de Waal F |url=http://www.huffingtonpost.com/frans-de-waal/was-ardi-perhaps-liberal_b_325201.html |title=Was "Ardi" a Liberal? |work=The Huffington Post |date=2009-10-18 |access-date=2009-10-18 |archive-date=2009-10-21 |archive-url=https://web.archive.org/web/20091021121234/http://www.huffingtonpost.com/frans-de-waal/was-ardi-perhaps-liberal_b_325201.html |url-status=live }} However, at least one confirmed report of cannibalism in the wild of a dead infant was described in 2008.{{cite news|url=http://news.bbc.co.uk/earth/hi/earth_news/newsid_8487000/8487138.stm|work=BBC News|title=Wild bonobo mother ape eats own infant in DR Congo|date=2010-02-01|vauthors=Walker M|access-date=2010-06-05|archive-date=2010-08-10|archive-url=https://web.archive.org/web/20100810211034/http://news.bbc.co.uk/earth/hi/earth_news/newsid_8487000/8487138.stm|url-status=live}}{{cite web | vauthors = Callaway E | url = https://www.newscientist.com/article/dn18454-hippy-apes-caught-cannibalising-their-young.html | title = Hippy apes caught cannibalising their young | work = New Scientist | date = 1 February 2010 | access-date = 18 April 2013 | archive-date = 1 August 2013 | archive-url = https://web.archive.org/web/20130801013918/http://www.newscientist.com/article/dn18454-hippy-apes-caught-cannibalising-their-young.html | url-status = live }} A 2016 paper reported two more instances of infant cannibalism, although it was not confirmed if infanticide was involved.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.

File:Composite image of male chimpanzee (left) and male bonobo (right) (CC BY version).png

File:Differences in the Cognitive Skills of Bonobos and Chimpanzees, 2010 study.png

In 2020, the first whole-genome comparison between chimpanzees and bonobos was published and showed genomic aspects that may underlie or have resulted from their divergence and behavioral differences, including selection for genes related to diet and hormones.{{cite journal | vauthors = Kovalaskas S, Rilling JK, Lindo J | title = Comparative analyses of the Pan lineage reveal selection on gene pathways associated with diet and sociality in bonobos | journal = Genes, Brain and Behavior | volume = 20 | issue = 3 | pages = e12715 | date = March 2021 | pmid = 33200560 | doi = 10.1111/gbb.12715 | s2cid = 226988471| doi-access = free }}

• {{cite web |date=December 16, 2020 |title=Whole genomes map pathways of chimpanzee and bonobo divergence |website=Phys.org |url=https://phys.org/news/2020-12-genomes-pathways-chimpanzee-bonobo-divergence.html |access-date=January 30, 2021 |archive-date=July 4, 2022 |archive-url=https://web.archive.org/web/20220704032737/https://phys.org/news/2020-12-genomes-pathways-chimpanzee-bonobo-divergence.html |url-status=live }} 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".{{cite journal |vauthors=Gruber T, Clay Z, Zuberbühler K |title=A comparison of bonobo and chimpanzee tool use: evidence for a female bias in the Pan lineage |journal=Animal Behaviour |date=1 December 2010 |volume=80 |issue=6 |pages=1023–1033 |doi=10.1016/j.anbehav.2010.09.005 |s2cid=14923158 |url=http://doc.rero.ch/record/278363/files/Gruber_T._-_A_comparison_of_bonobo_and_chimpanzee-20170117.pdf |access-date=17 April 2021 |archive-date=13 June 2023 |archive-url=https://web.archive.org/web/20230613195455/https://doc.rero.ch/record/278363/files/Gruber_T._-_A_comparison_of_bonobo_and_chimpanzee-20170117.pdf |url-status=live }} 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".{{cite journal | vauthors = Herrmann E, Hare B, Call J, Tomasello M | title = Differences in the cognitive skills of bonobos and chimpanzees | journal = PLOS ONE | volume = 5 | issue = 8 | pages = e12438 | date = August 2010 | pmid = 20806062 | pmc = 2929188 | doi = 10.1371/journal.pone.0012438 | bibcode = 2010PLoSO...512438H | doi-access = free }} File:CC-BY icon.svg Available under [https://creativecommons.org/licenses/by/4.0/ CC BY 4.0] {{Webarchive|url=https://web.archive.org/web/20171016050101/https://creativecommons.org/licenses/by/4.0/ |date=2017-10-16 }}. Bonobos have been found to be more risk-averse compared to chimpanzees, preferring immediate rather than delayed rewards when it comes to foraging. Bonobos also have a weaker spatial memory compared to chimpanzees, with adult bonobos performing comparably to juvenile chimpanzees.{{cite journal | vauthors = Hare B, Wobber V, Wrangham R | title = The self-domestication hypothesis: evolution of bonobo psychology is due to selection against aggression | journal = Genes, Brain and Behavior | volume = 83 | issue = 3 | pages = 573–585 | date = March 2012 | doi = 10.1016/j.anbehav.2011.12.007| s2cid = 3415520 }}

Bonobos are capable of passing the mirror-recognition test for self-awareness, as are all great apes.Dielenberg, R. A. (2013). "The comparative psychology of human uniqueness: A cognitive behavioral review", pp. 111–182 in R. G. Bednarik (Ed.), The psychology of human behavior. Nova Science Publishers. {{ISBN|1622579011}}{{cite journal | last1=Anderson | first1=James R. | last2=Gallup | first2=Gordon G. | title=Mirror self-recognition: a review and critique of attempts to promote and engineer self-recognition in primates | journal=Primates | volume=56 | issue=4 | date=2015 | pmid=26341947 | doi=10.1007/s10329-015-0488-9 | pages=317–326| hdl=2433/202619 | s2cid=254157787 | hdl-access=free }} They communicate primarily through vocal means, although the meanings of their vocalizations are not currently known. However, most humans do understand their facial expressions{{cite web | url = http://www.colszoo.org/animalareas/aforest/bonobo.html | work = Columbus Zoo | title = Bonobos at the Columbus Zoo | access-date = 2006-08-01 | archive-url = https://web.archive.org/web/20060502101116/http://www.colszoo.org/animalareas/aforest/bonobo.html | archive-date = 2006-05-02}} 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.{{cite news | vauthors = Webb J | date = August 4, 2015 | url = https://www.bbc.com/news/science-environment-33731444 | work = BBC News | title = Bonobo squeaks hint at earlier speech evolution | access-date = 2015-08-05 | archive-date = 2023-03-13 | archive-url = https://web.archive.org/web/20230313070243/https://www.bbc.com/news/science-environment-33731444 | url-status = live }}

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,{{cite web|url=http://www.greatapetrust.org/bonobo/meet/kanzi.php|title=Meet our Great Apes: Kanzi | work = Great Ape Trust | date = 2007 |access-date=2008-09-28 |archive-url = https://web.archive.org/web/20080630201101/http://www.greatapetrust.org/bonobo/meet/kanzi.php|archive-date = 2008-06-30}} and he has comprehension of around 3,000 spoken English words.{{cite magazine|vauthors=Raffaele P|year=2006|title=Speaking Bonobo|url=http://www.smithsonianmag.com/science-nature/10022981.html|magazine=Smithsonian|access-date=2008-09-28|archive-date=2013-12-16|archive-url=https://web.archive.org/web/20131216231356/http://www.smithsonianmag.com/science-nature/10022981.html|url-status=dead}}

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 the 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.{{cite journal| vauthors = Schick KD, Toth N, Garufi G, Savage-Rumbaugh ES, Rumbaugh D, Sevcik R |title=Continuing Investigations into the Stone Tool-making and Tool-using Capabilities of a Bonobo (Pan paniscus)|journal=Journal of Archaeological Science|volume=26|issue=7|pages=821–832|date=1999|doi=10.1006/jasc.1998.0350|bibcode=1999JArSc..26..821S }}

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.{{cite journal | vauthors = Palagi E, Paoli T, Tarli SB | title = Reconciliation and consolation in captive bonobos (Pan paniscus) | journal = American Journal of Primatology | volume = 62 | issue = 1 | pages = 15–30 | date = January 2004 | pmid = 14752810 | doi = 10.1002/ajp.20000 | s2cid = 22452710 }} A 2013 study{{cite journal | vauthors = Palagi E, Norscia I | title = Bonobos protect and console friends and kin | journal = PLOS ONE | volume = 8 | issue = 11 | pages = e79290 | year = 2013 | pmid = 24223924 | pmc = 3818457 | doi = 10.1371/journal.pone.0079290 | bibcode = 2013PLoSO...879290P | doi-access = free }} 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 bonobos have expressed joy have been reported. One study analyzed and recorded sounds made by human infants and bonobos when they were tickled.{{cite web|vauthors=Beale B|year=2003|title=Where Did Laughter Come From?|url=http://www.abc.net.au/science/articles/2003/10/07/961420.htm|publisher=ABC Science Online|access-date=2008-08-10|archive-date=2010-08-04|archive-url=https://web.archive.org/web/20100804050627/http://www.abc.net.au/science/articles/2003/10/07/961420.htm|url-status=live}} Although the bonobos' laugh was at a higher frequency, the laugh was found to follow a spectrographic pattern similar to that of human babies.

File:Ulrik, jeune de 1 ans avec sa mère Uma. LuiKotale, RDC.jpg

Bonobos are found only south of the Congo River and north of the Kasai River (a tributary of the Congo),{{cite book | vauthors = Dawkins R |author-link=Richard Dawkins |title=The Ancestor's Tale |year=2004 |publisher=Houghton Mifflin |chapter=Chimpanzees |isbn=978-1-155-16265-2}} 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.{{cite web| vauthors = Quammen D |title=The Left Bank Ape|url=http://ngm.nationalgeographic.com/2013/03/bonobos/quammen-text|archive-url=https://web.archive.org/web/20130221042436/http://ngm.nationalgeographic.com/2013/03/bonobos/quammen-text|url-status=dead|archive-date=February 21, 2013|work=The New Age of Exploration, 2013|publisher=National Geographic|access-date=28 February 2013}}

The ranges of bonobos and chimpanzees are separated by the Congo River, with bonobos living to its south and chimpanzees to the north.{{cite web | vauthors = Raffaele P | date = November 2006 | url = https://www.smithsonianmag.com/science-nature/the-smart-and-swinging-bonobo-134784867/ | title = The Smart and Swinging Bonobo | work = Smithsonian Magazine | access-date = 2018-01-23 | archive-date = 2023-03-30 | archive-url = https://web.archive.org/web/20230330220205/https://www.smithsonianmag.com/science-nature/the-smart-and-swinging-bonobo-134784867/ | url-status = live }}{{cite journal | vauthors = Caswell JL, Mallick S, Richter DJ, Neubauer J, Schirmer C, Gnerre S, Reich D | title = Analysis of chimpanzee history based on genome sequence alignments | journal = PLOS Genetics | volume = 4 | issue = 4 | pages = e1000057 | date = April 2008 | pmid = 18421364 | pmc = 2278377 | doi = 10.1371/journal.pgen.1000057 | veditors = McVean G | doi-access = free }}

File:Ida une jeune qui devra quitter son groupe à l'adolescence.jpg

In the Congo tropical rainforest, the very great majority of plants need animals to reproduce and disperse their seeds.{{Cite journal| vauthors = Beaune D, Bretagnolle F, Bollache L, Hohmann G, Surbeck M, Fruth B |date=January 2013|title=Seed dispersal strategies and the threat of defaunation in a Congo forest|journal=Biodiversity and Conservation|volume=22|issue=1|pages=225–238|doi=10.1007/s10531-012-0416-x|bibcode=2013BiCon..22..225B |s2cid=17600509}} Bonobos are the second largest frugivorous animals in this region, after elephants. It is estimated that 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 travel for about 24 hours in the bonobo digestive tract, which can 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, germinating better and more quickly than unpassed seeds.{{Cite journal| vauthors = Beaune D, Bretagnolle F, Bollache L, Bourson C, Hohmann G, Fruth B |date=September 2013|title=Ecological services performed by the bonobo (Pan paniscus): seed dispersal effectiveness in tropical forest |journal=Journal of Tropical Ecology|volume=29|issue=5|pages=367–380|doi=10.1017/s0266467413000515|s2cid=86540021}} For those seeds, diplochory with dung-beetles (Scarabaeidae) improves post-dispersal survival.{{Cite journal| vauthors = Beaune D, Bollache L, Bretagnolle F, Fruth B |date=2012-08-29|title=Dung beetles are critical in preventing post-dispersal seed removal by rodents in Congo rain forest |journal=Journal of Tropical Ecology|volume=28|issue=5|pages=507–510|doi=10.1017/s0266467412000466|s2cid=86529605}}

Certain plants such as Dialium may even be dependent on bonobos to activate the germination of their seeds, characterized by tegumentary dormancy.{{cite journal | vauthors = Beaune D, Bretagnolle F, Bollache L, Hohmann G, Surbeck M, Bourson C, Fruth B | title = The bonobo-dialium positive interactions: seed dispersal mutualism | journal = American Journal of Primatology | volume = 75 | issue = 4 | pages = 394–403 | date = April 2013 | pmid = 23307414 | doi = 10.1002/ajp.22121 | s2cid = 21817223 }} 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.{{Cite journal| vauthors = Beaune D |date=2015-02-27|title=What would happen to the trees and lianas if apes disappeared?|journal=Oryx|volume=49|issue=3|pages=442–446|doi=10.1017/s0030605314000878|s2cid=84253241|doi-access=free}} 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, would 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 many people, the ultimate success of conservation efforts still relies on local and community involvement. The issue of parks versus people{{cite web | vauthors = Reid J | date = 2006-06-15 | url = http://www.sfgate.com/opinion/openforum/article/Parks-and-people-not-parks-vs-people-2533132.php | title = Parks and people, not parks vs. people | work = San Francisco Chronicle | access-date = 2012-12-26 | archive-date = 2013-04-11 | archive-url = https://web.archive.org/web/20130411233300/http://www.sfgate.com/opinion/openforum/article/Parks-and-people-not-parks-vs-people-2533132.php | url-status = live }} is salient in the Cuvette Centrale, within the bonobos' range. There is strong local, and broad-based Congolese, resistance to establishing national parks, as indigenous communities have previously been driven from their forest homes by the forming of parks. In Salonga National Park (the only national park in bonobo habitat), there is no local involvement, and surveys undertaken since 2000 indicate the bonobo, the African forest elephant, the okapi, and other rare species have been devastated by poachers and the thriving bushmeat trade.{{cite web | url = http://www.zoosociety.org/conservation/Bonobo/BCBI/Survey.php | title = Bonobo and large mammal survey | archive-url = https://web.archive.org/web/20120506233748/http://www.zoosociety.org/conservation/Bonobo/BCBI/Survey.php | archive-date=May 6, 2012 | publisher = Zoological Society of Milwaukee }} In contrast, areas do exist where the bonobo and ecological biodiversity still thrive without any established park borders, because of the indigenous beliefs/taboos against killing bonobos and other animals.

During the wars in the 1990s, researchers and international non-governmental organizations (NGOs) were driven out of the bonobo habitat. In 2002, the Bonobo Conservation Initiative initiated the Bonobo Peace Forest Project (supported by the Global Conservation Fund of Conservation International), 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 {{convert|50,000|sqmi|km2}} of the bonobo habitat. According to Amy Parish, the Bonobo Peace Forest "is going to be a model for conservation in the 21st century".{{cite web | vauthors = Curwood S, Parish A | url = http://www.loe.org/shows/shows.htm?programID=06-P13-00027#feature2 | title = The Make Love, Not War Species | work = Living on Earth | date = July 2006 | publisher = National Public Radio | access-date = 2006-11-02 | archive-date = 2010-12-28 | archive-url = https://web.archive.org/web/20101228213230/http://www.loe.org/shows/shows.htm?programID=06-P13-00027#feature2 | url-status = live }}

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{{cite web | vauthors = Hart T | date = 2012-07-27 | url = http://www.bonoboincongo.com/the-bonobo | title = Searching for Bonobo in Congo | work = Bonoboincongo.com. | access-date = 2012-12-26 | archive-date = 2008-12-02 | archive-url = https://web.archive.org/web/20081202105930/http://www.bonoboincongo.com/the-bonobo/ | url-status = live }} use the town as a base.{{cite web | url = http://lolayabonobo.wildlifedirect.org/ | title = Lola Ya Bonobo (Bonobo Heaven) | archive-url = https://web.archive.org/web/20161124221827/http://lolayabonobo.wildlifedirect.org/ | archive-date = 2016-11-24 | work = Lolayabonobo.wildlifedirect.org. | access-date = 2012-12-26 }}{{cite web | url = http://www.friendsofbonobos.org/images/newsnov09.pdf | title = Bonobo Reintroduction in the Democratic Republic of Congo | archive-url = https://web.archive.org/web/20101218060615/http://www.friendsofbonobos.org/images/newsnov09.pdf | archive-date=2010-12-18 | work = friendsofbonobos.org | date = November 2009 | access-date = 2012-12-26 }}

In 1995, concern over declining numbers of bonobos in the wild led the Zoological Society of Milwaukee (ZSM), 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 ZSM has conducted regional surveys within the range of the bonobo in conjunction with training Congolese researchers in survey methodology and biodiversity monitoring. The ZSM'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 ZSM has become more involved in helping the Congolese living in bonobo habitat. They have 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.{{cite web | url = http://www.zoosociety.org/conservation/Bonobo/BCBI/ | archive-url = https://web.archive.org/web/20120210211110/http://www.zoosociety.org/conservation/Bonobo/BCBI/ | archive-date = 10 February 2012 | title = Bonobo and Congo Biodiversity Initiative | publisher = Zoological Society of Milwaukee}}

With grants from the United Nations, USAID, the U.S. Embassy, the World Wildlife Fund, and many other groups and individuals, the ZSM 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.{{cite web | vauthors = Chapin M | date = December 2004 | url = http://watha.org/in-depth/EP176A.pdf | title = Vision for a Sustainable World | work = WORLDWATCH magazine | access-date = 2012-12-26 | archive-date = 2021-07-19 | archive-url = https://web.archive.org/web/20210719061047/http://watha.org/in-depth/EP176A.pdf | url-status = dead }}

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.

World Bonobo Day is February 14 (Valentine's Day). This was established in 2017 by the African Wildlife Foundation.{{Cite web |url=https://www.awf.org/news/world-bonobo-day-protecting-our-closest-kin-hippie-chimps |title=Kibet R. World Bonobo Day: Protecting Our Closest Kin, the "Hippie Chimps" February 14, 2024 |access-date=February 15, 2024 |archive-date=February 15, 2024 |archive-url=https://web.archive.org/web/20240215041810/https://www.awf.org/news/world-bonobo-day-protecting-our-closest-kin-hippie-chimps |url-status=live }}

{{div col|colwidth=30em}}

• Basankusu, DR Congo – base for bonobo research and conservation
• Bonobo Conservation Initiative
• Chimpanzee genome project
• Claudine André
• Great ape personhood
• Great Ape Project
• International Primate Day
• Kanzi
• List of apes – notable individual nonhuman apes
• Lola ya Bonobo
• Koba, a fictional bonobo and antagonist of the Planet of the Apes reboot series

{{div col end}}

{{Notelist}}

{{Reflist}}

• {{cite book | vauthors = de Waal F, Lanting F | author-link1 = Frans de Waal | author-link2 = Frans Lanting | title = Bonobo: The Forgotten Ape | publisher = University of California Press | date = 1997 | isbn = 0-520-20535-9 }}
• {{cite book | vauthors = Takayoshi K | title = The Last Ape: Pygmy Chimpanzee Behavior and Ecology | location = Stanford, CA | publisher = Stanford University Press | date = 1992 }}
• {{cite book | vauthors = Savage-Rumbaugh S, Lewin R | author-link1 = Sue Savage-Rumbaugh | author-link2 = Roger Lewin | title = Kanzi: The Ape at the Brink of the Human Mind | publisher = John Wiley | date = 1994 | isbn = 0-471-58591-2}}
• {{cite book | vauthors = Woods V | author-link = Vanessa Woods | title = Bonobo Handshake | publisher = Gotham Books | date = 2010 | isbn = 978-1-59240-546-6}}
• {{cite book | vauthors = Sandin J | title = Bonobos: Encounters in Empathy | publisher = Zoological Society of Milwaukee & The Foundation for Wildlife Conservation, Inc. | date = 2007 | isbn = 978-0-9794151-0-4}}
• {{cite book | vauthors = de Waal F | author-link1 = Frans de Waal | title = The Bonobo and the Atheist | publisher = Norton | date = 2013 | isbn = 978-0-393-07377-5}}

• {{cite journal | vauthors = de Waal F | url = http://songweaver.com/info/bonobos.html | title = Bonobo: Sex & Society |journal= Scientific American | date = 1995 | volume = 272 | issue = 3 | pages = 82–88 | doi = 10.1038/scientificamerican0395-82 | pmid = 7871411 | bibcode = 1995SciAm.272c..82W }}
• {{cite journal | vauthors = DeBartolo A | url = http://www.hydeparkmedia.com/bonobo.html|url-status=usurped|archive-url=https://web.archive.org/web/20030211041752/http://www.hydeparkmedia.com/bonobo.html | title = The Bonobo: 'Newest' apes are teaching us about ourselves | journal = Chicago Tribune | date = 11 June 1998 | archive-date = 2003-02-11}}
• {{cite journal | vauthors = Schweller K | url = https://spectrum.ieee.org/apes-with-apps | title = Apes with Apps | journal = IEEE Spectrum Magazine | date = July 2012 | volume = 49 | issue = 7 | pages = 38–45 | doi = 10.1109/MSPEC.2012.6221081 | s2cid = 22556649 }}
• {{cite journal | vauthors = Madrigal A | url = https://www.theatlantic.com/health/archive/2014/06/brian-the-mentally-ill-bonobo-and-how-he-healed/372596/ | title = Brian the Mentally Ill Bonobo, and How He Healed | journal = The Atlantic | date = 11 June 2014 }}
• {{cite magazine | vauthors = Parker I | url = http://www.newyorker.com/magazine/2007/07/30/swingers-2?currentPage=all | title = Swingers | magazine = The New Yorker | date = 30 July 2007 }}
• {{cite journal | vauthors = Bechard D | url = http://www.thesolutionsjournal.com/node/237106 | title = Viral Conservation | journal = The Solutions Journal | date = February 2014 | access-date = 2016-07-13 | archive-date = 2016-03-26 | archive-url = https://web.archive.org/web/20160326142635/http://www.thesolutionsjournal.com/node/237106 | url-status = dead }}

• {{cite journal | vauthors = Fischer A, Prüfer K, Good JM, Halbwax M, Wiebe V, André C, Atencia R, Mugisha L, Ptak SE, Pääbo S | display-authors = 6 | title = Bonobos fall within the genomic variation of chimpanzees | journal = PLOS ONE | volume = 6 | issue = 6 | pages = e21605 | date = 29 June 2011 | pmid = 21747915 | pmc = 3126833 | doi = 10.1371/journal.pone.0021605 | editor-first = Etienne | bibcode = 2011PLoSO...621605F | editor-last = Joly | doi-access = free }}
• {{cite journal | vauthors = Zsurka G, Kudina T, Peeva V, Hallmann K, Elger CE, Khrapko K, Kunz WS | title = Distinct patterns of mitochondrial genome diversity in bonobos (Pan paniscus) and humans | journal = BMC Evolutionary Biology | volume = 10 | pages = 270 | date = September 2010 | issue = 1 | pmid = 20813043 | pmc = 2942848 | doi = 10.1186/1471-2148-10-270 | bibcode = 2010BMCEE..10..270Z | doi-access = free }}
• {{cite journal | vauthors = Wildman DE, Uddin M, Liu G, Grossman LI, Goodman M | title = Implications of natural selection in shaping 99.4% nonsynonymous DNA identity between humans and chimpanzees: enlarging genus Homo | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 100 | issue = 12 | pages = 7181–8 | date = June 2003 | pmid = 12766228 | pmc = 165850 | doi = 10.1073/pnas.1232172100 | bibcode = 2003PNAS..100.7181W | doi-access = free }}

{{Commons category}}

{{Wikispecies}}

• [https://www.pbs.org/wgbh/evolution/library/07/3/l_073_03.html Evolution: Why Sex?]
• [http://www.awf.org/content/wildlife/detail/bonobo Bonobos: Wildlife summary from the African Wildlife Foundation]
• [https://primate.wisc.edu/primate-info-net/pin-factsheets/pin-factsheet-bonobo/ Primate Info Net Pan paniscus Factsheet]
• [https://www.ted.com/talks/susan_savage_rumbaugh_the_gentle_genius_of_bonobos Susan Savage-Rumbaugh: The gentle genius of bonobos] – TED
• [https://wwf.panda.org/discover/knowledge_hub/endangered_species/great_apes/bonobo/ WWF (World Wide Fund for Nature / World Wildlife Fund) – Bonobo species profile]
• [https://web.archive.org/web/20130723063336/http://library.sandiegozoo.org/factsheets/bonobo/bonobo.htm San Diego Zoo Library: Bonobo, Pan paniscus]
• {{UCSC genomes|panPan1}}

{{Hominidae nav}}

{{Apes}}

{{Notable apes}}

{{Portal bar|Biology|Primates}}

{{Taxonbar|from=Q19537}}

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Category:Mammals described in 1929

Category:Primates of Africa

Category:Fauna of Central Africa

Category:Mammals of the Democratic Republic of the Congo

Category:Endemic fauna of the Democratic Republic of the Congo

Category:Tool-using mammals

Category:Extant Pleistocene first appearances

Category:Taxa named by Ernst Schwarz