Bonobo

The bonobo (/bəˈnoʊboʊ, ˈbɒnəboʊ/; Pan paniscus), also

historically called the pygmy chimpanzee and less often, the Bonobo[1]
dwarf or gracile chimpanzee,[3] is an endangered great ape and Temporal range:
one of the two species making up the genus Pan; the other being Early Pleistocene – Holocene
the common chimpanzee (Pan troglodytes).[4] Although bonobos ↓

PreЄ Є O S D C P T J K PgN
are not a subspecies of chimpanzee (Pan troglodytes), but rather a
distinct species in their own right, both species are sometimes
referred to collectively using the generalized term chimpanzees, or
chimps. Taxonomically, the members of the chimpanzee/bonobo
subtribe Panina (comprised entirely by the genus Pan) are
collectively termed panins.[5][6]

The bonobo is distinguished by relatively long legs, pink lips, dark

face, tail-tuft through adulthood, and parted long hair on its head.
The bonobo is found in a 500,000 km2 (190,000 sq mi) area of the
Congo Basin in the Democratic Republic of the Congo, Central
Africa. The species is omnivorous and inhabits primary and
secondary forests, including seasonally inundated swamp forests.
Because of political instability in the region and the timidity of Male at Apenheul Primate Park
bonobos, there has been relatively little field work done observing Conservation status
the species in its natural habitat.

Along with the common chimpanzee, the bonobo is the closest

extant relative to humans.[4] As the two species are not proficient
swimmers, the formation of the Congo River 1.5–2 million years Endangered (IUCN 3.1)[2]
ago possibly led to the speciation of the bonobo. Bonobos live
Scientific classification
south of the river, and thereby were separated from the ancestors
of the common chimpanzee, which live north of the river. There Kingdom: Animalia
are no concrete data on population numbers, but the estimate is
between 29,500 and 50,000 individuals. The species is listed as Phylum: Chordata
Endangered on the IUCN Red List and is threatened by habitat Class: Mammalia
destruction and human population growth and movement, though
commercial poaching is the most prominent threat. Bonobos Order: Primates
typically live 40 years in captivity; their lifespan in the wild is Suborder: Haplorhini
unknown, but it is almost certainly much shorter.[7]
Infraorder: Simiiformes
Family: Hominidae
Contents Subfamily: Homininae

Etymology Tribe: Hominini

Taxonomy Genus: Pan
Description Species: P. paniscus
Behavior Binomial name
General
Social behavior Pan paniscus
 Sociosexual behaviour Schwarz, 1929
Peacefulness
Diet
Similarity to humans
Distribution and habitat
Conservation status
See also
Notes
References
Further reading
Books Bonobo distribution
Articles
Journal articles
External links

Etymology
Despite the species' common name "pygmy chimpanzee", the bonobo is not especially diminutive when
compared to the common chimpanzee, with exception of its head. The appellative "pygmy" is attributable to
the species' namer, Ernst Schwarz, who classified the species on the basis of a previously mislabeled bonobo
cranium, noting its diminutive size compared to chimpanzee skulls.[8]

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.[9][10]

Taxonomy
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 chimp (Pan
troglodytes). Schwarz published his findings in 1929, classifying the bonobo as a subspecies of
chimp.[11][12] In 1933, American anatomist Harold Coolidge elevated it to species status.[12][13] Major
behavioural differences between bonobos and chimps were first discussed in detail by Tratz and Heck in the
early 1950s.[14] American psychologist and primatologist Robert Yerkes was also one of the first to notice
major behavioural differences.[15]

The first official publication of the sequencing and assembly of the bonobo genome was published in June
2012. The genome of a female bonobo from the Leipzig zoo was deposited with the International Nucleotide
Sequence Database Collaboration (DDBJ/EMBL/GenBank) under the EMBL accession number
AJFE01000000[16] 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.[17]

Bonobos and chimps are the two species which make up the genus Pan, and are the closest living relatives
to humans (Homo sapiens).[18][19]
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.[20] DNA evidence suggests the bonobo and common chimpanzee species diverged approximately
890,000–860,000 years ago due to separation of these two populations possibly due to acidification and the
spread of savannas at this time. Currently, these two species are separated by the Congo River, which had
existed well before the divergence date, though ancestral Pan may have dispersed across the river using
corridors which no longer exist.[21] The first Pan fossils were reported in 2005 from the Middle Pleistocene
(after the bonobo–chimp split) of Kenya, alongside early Homo fossils.[22]

According to A. Zihlman, bonobo body proportions closely resemble those of Australopithecus,[23] leading
evolutionary biologist Jeremy Griffith to suggest that bonobos may be a living example of our distant human
ancestors.[24] 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.[25]

Description
The bonobo is commonly considered to be more gracile than the Genomic information
common chimpanzee. Although large male chimpanzees can exceed
NCBI genome 10729 (https://www.
any bonobo in bulk and weight, the two species actually broadly
ID ncbi.nlm.nih.gov/ge
overlap in body size. Adult female bonobos are somewhat smaller
than adult males. Body mass in males ranges from 34 to 60 kg (75 to nome/?term=1072
132 lb), against an average of 30 kg (66 lb) in females. The total 9)
length of bonobos (from the nose to the rump while on all fours) is Ploidy diploid
70 to 83 cm (28 to 33 in).[26][27][28][29] When adult bonobos and
Genome size 2,869.21 Mb
chimpanzees stand up on their legs, they can both attain a height of
115 cm (45 in).[30] The bonobo's head is relatively smaller than that Number of 24 pairs
of the common chimpanzee with less prominent brow ridges above chromosomes
the eyes. It has a black face with pink lips, small ears, wide nostrils, Year of 2012
and long hair on its head that forms a parting. Females have slightly completion
more prominent breasts, in contrast to the flat breasts of other female
apes, although not so prominent as those of humans. The bonobo
also has a slim upper body, narrow shoulders, thin neck, and long legs when compared to the common
chimpanzee.

Bonobos are both terrestrial and arboreal. Most ground locomotion

Bonobos Kanzi (C) and Panbanisha
(R) with Sue Savage-Rumbaugh and
the outdoor symbols "keyboard"
characteristic adapted for visual facial recognition in social interaction.
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,[31] 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.[32] 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,[33] as humans do, so that one
individual may look significantly different from another, a
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.[34] Other researchers
challenged this conclusion.[35]

Behavior

General

Primatologist Frans de Waal states bonobos are capable of altruism, compassion, empathy, kindness,
patience, and sensitivity,[3] and described "bonobo society" as a "gynecocracy".[36][a] Primatologists who
have studied bonobos in the wild have documented a wide range of behaviors, including aggressive behavior
and more cyclic sexual behavior similar to chimpanzees, even though bonobos show more sexual behavior
in a greater variety of relationships. An analysis of female bonding among wild bonobos by Takeshi
Furuichi stresses female sexuality and shows how female bonobos spend much more time in estrus than
female chimpanzees.[37]

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.[38] 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.[39] Bonobos
warn each other of danger less efficiently than chimpanzees in the same situation.[40]

Social behavior

Many studies indicate that

females have a higher social
status in bonobo society.
Aggressive encounters
between males and females
are rare, and males are
tolerant of infants and
juveniles. A male derives his
status from the status of his
Bonobo searching for termites
mother.[41] The mother–son Bonobos are very social.
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.[42]

Due to the promiscuous mating behavior of female bonobos, a male cannot be sure which offspring are his.
As a result, the entirety of parental care in bonobos is assumed by the mothers.[43]

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.
Sociosexual behaviour

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.[44][4] Bonobos are the only non-human
animal to have been observed engaging in tongue kissing.[45]
Bonobos and humans are the only primates to typically engage in
face-to-face genital sex, although a pair of western gorillas has been
photographed in this position.[46]
Bonobos mating, Jacksonville Zoo
and Gardens.
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.[47]

Bonobo clitorises are larger and more externalized than in most mammals;[48] 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".[49] In scientific
literature, the female–female behavior of bonobos pressing genitals together is often referred to as genito-
genital (GG) rubbing,[47][50] which is the non-human analogue of tribadism, engaged in by some human
females. This sexual activity happens within the immediate female bonobo community and sometimes
outside of it. Ethologist Jonathan Balcombe stated that female bonobos rub their clitorises together rapidly
for ten to twenty seconds, and this behavior, "which may be repeated in rapid succession, is usually
accompanied by grinding, shrieking, and clitoral engorgement"; he added that it is estimated that they
engage in this practice "about once every two hours" on average.[48] As bonobos occasionally copulate face-
to-face, "evolutionary biologist Marlene Zuk has suggested that the position of the clitoris in bonobos and
some other primates has evolved to maximize stimulation during sexual intercourse".[48] 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.[51]

Bonobo males engage in various forms of male–male genital

behavior.[47][52] The most common form of male–male mounting is
similar to that of a heterosexual mounting: one of the males sits
"passively on his back [with] the other male thrusting on him", with
the penises rubbing together due to both males' erections.[3] In
another, rarer form of genital rubbing, which is the non-human
analogue of frotting, engaged in by some human males, two bonobo
males hang from a tree limb face-to-face while penis fencing.[47][53]
Group of bonobos
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.[54]
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.[55][3]
More often than the males, female bonobos engage in mutual genital behavior, possibly to bond socially
with each other, thus forming a female nucleus of bonobo society. The bonding among females enables them
to dominate most of the males.[47] Adolescent females often leave their native community to join another
community. This migration mixes the bonobo gene pools, providing genetic diversity. Sexual bonding with
other females establishes these new females as members of the group.

Bonobo reproductive rates are no higher than those of the common chimpanzee.[47] 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.[56] 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.[42] Adult male bonobos have sex with infants,[57] although without penetration.[58]

It is unknown how the bonobo avoids simian immunodeficiency virus (SIV) and its effects.[59]

Peacefulness

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.[60] This does not appear to be the behavior of bonobo
males or females, which seem to prefer sexual contact over violent
confrontation with outsiders.[4]

While bonobos are more peaceful than chimpanzees, it is not true

that they are unaggressive.[61] In the wild, among males, bonobos Bonobo (Pan paniscus) mother and
are half as aggressive as chimpanzees, while female bonobos are infant at Lola ya Bonobo
more aggressive than female chimpanzees.[61] Both bonobos and
chimpanzees exhibit physical aggression more than 100 times as
often as humans do.[61]

The ranges of bonobos and chimpanzees are separated by the Congo River, with bonobos living to the south
of it, and chimpanzees to the north.[62][63] 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.[64]

Recent studies show that there are significant brain differences between bonobos and chimps. The brain
anatomy of bonobos has more developed and larger regions assumed to be vital for feeling empathy, sensing
distress in others and feeling anxiety, which makes them less aggressive and more empathic than their close
relatives. They also have a thick connection between the amygdala, an important area that can spark
aggression, and the ventral anterior cingulate cortex, which helps control impulses. This thicker connection
may make them better at regulating their emotional impulses and behavior.[65]

Bonobo society is dominated by females, and severing the lifelong alliance between mothers and their male
offspring may make them vulnerable to female aggression.[4] 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."[66]

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.[67]

Diet

The bonobo is an omnivorous frugivore; 57% of its diet is fruit, but this is supplemented with leaves, honey,
eggs,[68] meat from small vertebrates such as anomalures, flying squirrels and duikers,[69] and
invertebrates.[70] In some instances, bonobos have been shown to consume lower-order primates.[67] Some
claim bonobos have also been known to practise cannibalism in captivity, a claim disputed by others.[71][72]
However, at least one confirmed report of cannibalism in the wild of a dead infant was described in
2008.[73][74]

Similarity to humans

Bonobos are capable of passing the mirror-recognition test for self-awareness,[75] as are all great apes. They
communicate primarily through vocal means, although the meanings of their vocalizations are not currently
known. However, most humans do understand their facial expressions[76] 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.[77] 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,[78] and he has comprehension of
around 3,000 spoken English words.[79] Kanzi is also known for learning by observing people trying to
teach his mother; Kanzi started doing the tasks that his mother was taught just by watching, some of which
his mother had failed to learn. Some, such as philosopher and bioethicist Peter Singer, argue that these
results qualify them for "rights to survival and life"—rights which humans theoretically accord to all persons
(See great ape personhood). In the 1990s, Kanzi was taught to make and use simple stone tools. This
resulted from a study undertaken by researchers Kathy Schick and Nicholas Toth, and later Gary Garufi. The
researchers wanted to know if Kanzi possessed the cognitive and biomechanical abilities required to make
and use stone tools. Though Kanzi was able to form flakes, he did not create them in same way as humans,
who hold the core in one hand and knap it with the other, Kanzi threw the cobble against a hard surface or
against another cobble. This allowed him to produce a larger force to initiate a fracture as opposed to
knapping it in his hands.[80]

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.[81] A 2013 study [82] found that both the affiliation spontaneously offered by a bystander to the
victim and the affiliation requested by the victim (solicited affiliation) can reduce the probability of further
aggression by group members on the victim (this fact supporting the Victim-Protection Hypothesis). Yet,
only spontaneous affiliation reduced victim anxiety—measured via self-scratching rates—thus suggesting
not only that non-solicited affiliation has a consolatory function but also that the spontaneous gesture—more
than the protection itself—works in calming the distressed subject. The authors hypothesize that the victim
may perceive the motivational autonomy of the bystander, who does not require an invitation to provide
post-conflict affinitive contact. Moreover, spontaneous—but not solicited—third party affiliation was
affected by the bond between consoler and victim (this supporting the Consolation Hypothesis). Importantly,
spontaneous affiliation followed the empathic gradient described for humans, being mostly offered to kin,
then friends, then acquaintances (these categories having been determined using affiliation rates between
individuals). Hence, consolation in the bonobo may be an empathy-based phenomenon.

Instances in which non-human primates have expressed joy have been reported. One study analyzed and
recorded sounds made by human infants and bonobos when they were tickled.[83] Although the bonobos'
laugh was at a higher frequency, the laugh was found to follow a spectrographic pattern similar to that of
human babies.[83]

Distribution and habitat

Bonobos are found only south of the Congo River and north of the Kasai River (a tributary of the
Congo),[84] 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.[85]

Conservation status
The IUCN Red List classifies bonobos as an endangered species, with conservative population estimates
ranging from 29,500 to 50,000 individuals.[2] Major threats to bonobo populations include habitat loss and
hunting for bushmeat, the latter activity having increased dramatically during the first and second Congo
wars in the Democratic Republic of Congo due to the presence of heavily armed militias even in remote
"protected" areas such as Salonga National Park. This is part of a more general trend of ape extinction.

As the bonobos' habitat is shared with people, the ultimate success of conservation efforts still rely on local
and community involvement. The issue of parks versus people[86] is salient in the Cuvette Centrale the
bonobos' range. There is strong local and broad-based Congolese resistance to establishing national parks, as
indigenous communities have often been driven from their forest homes by the establishment of parks. In
Salonga National Park, the only national park in the bonobo habitat, there is no local involvement, and
surveys undertaken since 2000 indicate the bonobo, the African forest elephant, and other species have been
devastated by poachers and the thriving bushmeat trade.[87] In contrast, areas exist where the bonobo and
biodiversity still thrive without any established parks, due to the indigenous beliefs and taboos against
killing bonobos.

During the wars in the 1990s, researchers and international non-governmental organizations (NGOs) were
driven out of the bonobo habitat. In 2002, the Bonobo Conservation Initiative initiated the Bonobo Peace
Forest Project supported by the Global Conservation Fund of Conservation International and in cooperation
with national institutions, local NGOs, and local communities. The Peace Forest Project works with local
communities to establish a linked constellation of community-based reserves, managed by local and
indigenous people. This model, implemented mainly through DRC organizations and local communities, has
helped bring about agreements to protect over 50,000 square miles (130,000 km2) of the bonobo habitat.
According to Dr. Amy Parish, the Bonobo Peace Forest "is going to be a model for conservation in the 21st
century".[88]

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[89] use the
town as a base.[90][91]

In 1995, concern over declining numbers of bonobos in the wild led the Zoological Society of Milwaukee,
in Milwaukee, Wisconsin, with contributions from bonobo scientists around the world, to publish the Action
Plan for Pan paniscus: A Report on Free Ranging Populations and Proposals for their Preservation. The
Action Plan compiles population data on bonobos from 20 years of research conducted at various sites
throughout the bonobo's range. The plan identifies priority actions for bonobo conservation and serves as a
reference for developing conservation programs for researchers, government officials, and donor agencies.

Acting on Action Plan recommendations, the ZSM developed the Bonobo and Congo Biodiversity Initiative.
This program includes habitat and rain-forest preservation, training for Congolese nationals and
conservation institutions, wildlife population assessment and monitoring, and education. The Zoological
Society has conducted regional surveys within the range of the bonobo in conjunction with training
Congolese researchers in survey methodology and biodiversity monitoring. The Zoological Society’s initial
goal was to survey Salonga National Park to determine the conservation status of the bonobo within the park
and to provide financial and technical assistance to strengthen park protection. As the project has developed,
the Zoological Society has become more involved in helping the Congolese living in bonobo habitat. The
Zoological Society has built schools, hired teachers, provided some medicines, and started an agriculture
project to help the Congolese learn to grow crops and depend less on hunting wild animals.[92]

With grants from the United Nations, USAID, the U.S. Embassy, the World Wildlife Fund, and many other
groups and individuals, the Zoological Society also has been working to:

Survey the bonobo population and its habitat to find ways to help protect these apes
Develop antipoaching measures to help save apes, forest elephants, and other endangered
animals in Congo's Salonga National Park, a UN World Heritage site
Provide training, literacy education, agricultural techniques, schools, equipment, and jobs for
Congolese living near bonobo habitats so that they will have a vested interest in protecting the
great apes – the ZSM started an agriculture project to help the Congolese learn to grow crops
and depend less on hunting wild animals.
Model small-scale conservation methods that can be used throughout Congo

Starting in 2003, the U.S. government allocated $54 million to the Congo Basin Forest Partnership. This
significant investment has triggered the involvement of international NGOs to establish bases in the region
and work to develop bonobo conservation programs. This initiative should improve the likelihood of bonobo
survival, but its success still may depend upon building greater involvement and capability in local and
indigenous communities.[93]

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.

See also
Basankusu, DR Congo – base for bonobo research and conservation
 Bonobo Conservation Initiative
Chimpanzee genome project
Claudine André
Great ape personhood
Great Ape Project
Kanzi – a captive bonobo who uses language
List of apes – notable individual nonhuman apes
Lola ya Bonobo

Notes
a. Gynecocracy, among people, 'women's government over women and men' or 'women's social
supremacy'

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paniscus)". Am J Primatol. 62 (1): 15–30. doi:10.1002/ajp.20000 (https://doi.org/10.1002%2Faj
p.20000). PMID 14752810 (https://pubmed.ncbi.nlm.nih.gov/14752810).
82. Palagi E; Norscia I (2013). "Bonobos Protect and Console Friends and Kin" (https://www.ncbi.n
lm.nih.gov/pmc/articles/PMC3818457). PLOS ONE. 8 (11): e79290.
Bibcode:2013PLoSO...879290P (https://ui.adsabs.harvard.edu/abs/2013PLoSO...879290P).
doi:10.1371/journal.pone.0079290 (https://doi.org/10.1371%2Fjournal.pone.0079290).
PMC 3818457 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3818457). PMID 24223924 (htt
ps://pubmed.ncbi.nlm.nih.gov/24223924).
83. Beale, B. (2003). "Where Did Laughter Come From?" (http://www.abc.net.au/science/articles/2
003/10/07/961420.htm). ABC Science Online. Retrieved 2008-08-10.
84. Dawkins, Richard (2004). "Chimpanzees". The Ancestor's Tale. Houghton Mifflin. ISBN 978-1-
155-16265-2.
85. Quammen, David. "The Left Bank Ape" (http://ngm.nationalgeographic.com/2013/03/bonobos/
quammen-text). The New Age of Exploration, 2013. National Geographic. Retrieved
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on/openforum/article/Parks-and-people-not-parks-vs-people-2533132.php), San Francisco
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87. Bonobo and large mammal survey (http://www.zoosociety.org/conservation/Bonobo/BCBI/Surv
ey.php) Archived (https://web.archive.org/web/20120506233748/http://www.zoosociety.org/con
servation/Bonobo/BCBI/Survey.php) May 6, 2012, at the Wayback Machine. Zoological Society
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0027#feature2), Living on Earth, (July 2006), National Public Radio
89. Hart, Therese (2012-07-27) Searching for Bonobo in Congo (http://www.bonoboincongo.com/t
he-bonobo). Bonoboincongo.com. Retrieved on 2012-12-26.
90. Lola Ya Bonobo (Bonobo Heaven) (http://lolayabonobo.wildlifedirect.org/) Archived (https://we
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depth/EP176A.pdf), WORLDWATCH magazine

Further reading

Books
de Waal, Frans, and Frans Lanting, Bonobo: The Forgotten Ape, University of California Press,
1997. ISBN 0-520-20535-9; ISBN 0-520-21651-2 (trade paperback)
 Kano, Takayoshi, The Last Ape: Pygmy Chimpanzee Behavior and Ecology, Stanford, CA:
Stanford University Press, 1992.
Savage-Rumbaugh, Sue, and Roger Lewin, Kanzi: The Ape at the Brink of the Human Mind,
John Wiley, 1994. ISBN 0-471-58591-2; ISBN 0-471-15959-X (trade paperback)
Woods, Vanessa, Bonobo Handshake, Gotham Books, 2010. ISBN 978-1-59240-546-6
Sandin, Jo, Bonobos: Encounters in Empathy, Zoological Society of Milwaukee & The
Foundation for Wildlife Conservation, Inc., 2007. ISBN 978-0-9794151-0-4
de Waal, Frans, The Bonobo and the Atheist, Norton, 2013. ISBN 978-0393073775

Articles
de Waal, Frans, "Bonobo: Sex & Society" (http://songweaver.com/info/bonobos.html), Scientific
American, 1995
DeBartolo, Anthony. "The Bonobo: 'Newest' apes are teaching us about ourselves" (http://ww
w.hydeparkmedia.com/bonobo.html), Chicago Tribune June 11, 1998.
Schweller, Ken, "Apes With Apps," (https://spectrum.ieee.org/computing/software/apes-with-ap
ps/) IEEE Spectrum Magazine, July 2012.
Madrigal, Alexis "Brian the Mentally Ill Bonobo, and How He Healed" (https://www.theatlantic.c
om/health/archive/2014/06/brian-the-mentally-ill-bonobo-and-how-he-healed/372596/), The
Atlantic, June 11, 2014.
Parker, Ian "Swingers" (http://www.newyorker.com/magazine/2007/07/30/swingers-2?currentP
age=all), The New Yorker, July 30, 2007.
Bechard, Deni "Viral Conservation (http://www.thesolutionsjournal.com/node/237106)" The
Solutions Journal, February 2014

Journal articles
Fischer, Anne; Prüfer, Kay; Good, Jeffrey M.; Halbwax, Michel; Wiebe, Victor; André, Claudine;
Atencia, Rebeca; Mugisha, Lawrence; Ptak, Susan E.; Pääbo, Svante (29 June 2011). Joly,
Etienne (ed.). "Bonobos fall within the genomic variation of chimpanzees" (https://www.ncbi.nl
m.nih.gov/pmc/articles/PMC3126833). PLoS ONE. 6 (6): e21605.
Bibcode:2011PLoSO...621605F (https://ui.adsabs.harvard.edu/abs/2011PLoSO...621605F).
doi:10.1371/journal.pone.0021605 (https://doi.org/10.1371%2Fjournal.pone.0021605).
PMC 3126833 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3126833). PMID 21747915 (htt
ps://pubmed.ncbi.nlm.nih.gov/21747915).
Zsurka, Gábor; Kudina, Tatiana; Peeva, Viktoriya; Hallmann, Kerstin; Elger, Christian E.; Elger,
Konstantin; Khrapko, Konstantin; Kunz, Wolfram S. (2010). "Distinct patterns of mitochondrial
genome diversity in bonobos (Pan paniscus) and humans" (https://www.ncbi.nlm.nih.gov/pmc/
articles/PMC2942848). BMC Evolutionary Biology. 10: 270. doi:10.1186/1471-2148-10-270 (htt
ps://doi.org/10.1186%2F1471-2148-10-270). PMC 2942848 (https://www.ncbi.nlm.nih.gov/pm
c/articles/PMC2942848). PMID 20813043 (https://pubmed.ncbi.nlm.nih.gov/20813043).
Wildman, Derek E.; Uddin, Monica; Liu, Guozhen; Grossman, Lawrence I.; Goodman, Morris
(10 June 2003). "Implications of natural selection in shaping 99.4% nonsynonymous DNA
identity between humans and chimpanzees: Enlarging genus Homo" (https://www.ncbi.nlm.nih.
gov/pmc/articles/PMC165850). PNAS. 100 (12): 7181–7188. Bibcode:2003PNAS..100.7181W
(https://ui.adsabs.harvard.edu/abs/2003PNAS..100.7181W). doi:10.1073/pnas.1232172100 (ht
tps://doi.org/10.1073%2Fpnas.1232172100). PMC 165850 (https://www.ncbi.nlm.nih.gov/pmc/
articles/PMC165850). PMID 12766228 (https://pubmed.ncbi.nlm.nih.gov/12766228).

External links
 ARKive – BBC images and movies of the bonobo (Pan paniscus) (https://web.archive.org/web/
20060321204742/http://www.arkive.org/species/GES/mammals/Pan_paniscus/)
Evolution: Why Sex? (https://www.pbs.org/wgbh/evolution/library/07/3/l_073_03.html)
Bonobos: Wildlife summary from the African Wildlife Foundation (http://www.awf.org/content/wi
ldlife/detail/bonobo)
Primate Info Net Pan paniscus Factsheet (http://pin.primate.wisc.edu/factsheets/entry/bonobo)
U.S. Fish & Wildlife Service Species Profile (https://web.archive.org/web/20060928044248/htt
p://ecos.fws.gov/species_profile/servlet/gov.doi.species_profile.servlets.SpeciesProfile?spcode
=A06E)
"The Last Great Ape" (https://www.pbs.org/wgbh/nova/bonobos/), an episode of Nova.
Susan Savage-Rumbaugh: Apes that write, start fires and play Pac-Man (http://www.ted.com/i
ndex.php/talks/susan_savage_rumbaugh_on_apes_that_write.html) – Ted.com
WWF (World Wide Fund for Nature / World Wildlife Fund) – Bonobo species profile (http://wwf.
panda.org/what_we_do/endangered_species/great_apes/bonobo/)
Encyclopedia of Life (http://eol.org/pages/326448/overview)
San Diego Zoo Library: Bonobo, Pan paniscus (https://web.archive.org/web/20130723063336/
http://library.sandiegozoo.org/factsheets/bonobo/bonobo.htm)
Human Timeline (Interactive) (http://humanorigins.si.edu/evidence/human-evolution-timeline-in
teractive) – Smithsonian, National Museum of Natural History (August 2016).
View the panPan1 (https://genome.ucsc.edu/cgi-bin/hgTracks?db=panPan1) genome
assembly in the UCSC Genome Browser.

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