Editing Bonobo (section)

==== Sociosexual behaviour ====
{{See also|Animal sexual behaviour#Genital-genital rubbing|Homosexual behavior in animals#Bonobos}}
[[File:Bonobo sexual behavior 1.jpg|thumb|Bonobos mating, [[Jacksonville Zoo and Gardens]].]]
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 [[reconciliation (democratic process)#In animals|postconflict reconciliation]].<ref>{{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]] }}</ref><ref name="NYT-20160910" /> Bonobos are the only non-human animal to have been observed engaging in [[tongue kiss]]ing.<ref name=Manson1997>{{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 }}</ref> Bonobos and [[human]]s are the only primates to typically engage in face-to-face genital sex, although a pair of [[western gorilla]]s has also been photographed in this position.<ref>{{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 }}</ref>

Bonobos do not form [[Monogamy in animals|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.<ref name="deWaal1995">{{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 }}</ref>

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.<ref name="deWaal1995" /> Adolescent females often leave their native community to join another community. This migration mixes the bonobo [[gene pool]]s, providing [[genetic diversity]]. Sexual bonding with other females establishes these new females as members of the group.

Bonobo [[clitoris]]es are larger and more externalized than in most mammals;<ref name="Balcombe 2011">{{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}}</ref> 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".<ref name="Angier">{{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 }}</ref> In scientific literature, the female–female behavior of bonobos pressing vulvas together is often referred to as [[Genital-genital rubbing|genito-genital (GG) rubbing]].<ref name="deWaal1995" /><ref>{{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 }}</ref> This sexual activity happens within the immediate female bonobo community and sometimes outside of it. [[Ethology|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.<ref name="Balcombe 2011" /> As bonobos occasionally copulate face-to-face,<ref>{{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}}</ref> "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".<ref name="Balcombe 2011" /> 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.<ref name="hohmannfruth2000">{{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 }}</ref>
[[File:Pan paniscus11.jpg|thumb|left|Group of bonobos]]
Bonobo males engage in various forms of male–male genital behavior.<ref name="deWaal1995" /><ref>{{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}}</ref> 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.<ref name="deWaal1997" /> In another, rarer form of genital rubbing, two bonobo males hang from a tree limb face-to-face while [[penis fencing#Other uses|penis fencing]].<ref name="deWaal1995" /><ref name=penisfencing>{{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}}</ref> 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.<ref>{{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 }}</ref> [[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.<ref>{{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}}</ref><ref name="deWaal1997" />

Wild females give birth for the first time at 13 or 14 years of age.<ref>{{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 }}</ref> Bonobo reproductive rates are no higher than those of the common chimpanzee.<ref name=deWaal1995 /> However, female bonobo oestrus periods are longer.<ref>{{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}}</ref> During [[oestrus]], females undergo [[Sexual swelling|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.<ref name="ADW">{{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 }}</ref> 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.<ref name="bbc.com" />

Adult male bonobos have sex with infants,<ref name="de Waal 1990">{{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}}</ref> although without penetration.<ref>{{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 }}</ref> 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.<ref name="de Waal 1990" /> They have also been shown to be sexually active even in the absence of any stimulation or learning from adults.<ref>{{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.}}</ref>

Infanticide, while well documented in [[chimpanzee]]s, is apparently absent in bonobo society.<ref name="Gottfried 300–309">{{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 }}</ref> Although infanticide has not been directly observed, there have been documented cases of both female<ref>{{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 }}</ref> and male<ref name="Questioning the “sexy” bonobo hype, part 2: a primatologist debunks Christopher Ryan - Incredulous">{{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 }}</ref> 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.<ref>{{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}}</ref>
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.<ref>{{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 }}</ref> 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.<ref name="Gottfried 300–309" /> This is a reproductive strategy that seems specific to bonobos; infanticide is observed in all other great apes except [[orangutan]]s.<ref>{{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 }}</ref> Bonobos engage in sexual activity numerous times a day.<ref>{{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 }}</ref>

It is unknown how the bonobo avoids [[simian immunodeficiency virus]] (SIV) and its effects.<ref>{{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 }}</ref>