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Bonobo Sex and Society
The behavior of a close relative challenges assumptions
about male supremacy in human evolution
Frans B. M. de Waal
B. M. de WAAL was trained as an ethologist in the European
tradition, receiving his Ph.D. from the University of Utrecht in
1977. After a six-year study of the chimpanzee colony at the Arnhem
Zoo, he moved to the U.S. in 1981 to work on other primate species,
including bonobos. He is now a research professor at the
Yerkes Regional Primate
Research Center in Atlanta and professor of psychology at
At a juncture in history during which women are seeking equality
with men, science arrives with a belated gift to the feminist
movement. Male-biased evolutionary scenarios – Man the Hunter, Man
the Toolmaker and so on – are being challenged by the discovery that
females play a central, perhaps even dominant, role in the social
life of one of our nearest relatives. In the past few years many
strands of knowledge have come together concerning a relatively
unknown ape with an unorthodox repertoire of behavior: the bonobo.
bonobo is one of the last large mammals to be found by science. The
creature was discovered in 1929 in a Belgian colonial museum, far
from its lush African habitat. A German anatomist, Ernst Schwarz,
was scrutinizing a skull that had been ascribed to a juvenile
chimpanzee because of its small size, when he realized that it
belonged to an adult. Schwarz declared that he had stumbled on a new
subspecies of chimpanzee. But soon the animal was assigned the
status of an entirely distinct species within the same genus as the
The bonobo was officially classified as Pan paniscus, or the
diminutive Pan. But I believe a different label might have been
selected had the discoverers known then what we know now. The old
taxonomic name of the chimpanzee, P. satyrus – which refers to the
myth of apes as lustful satyrs – would have been perfect for the
The species is best characterized as female-centered and
egalitarian and as one that substitutes sex for aggression. Whereas
in most other species sexual behavior is a fairly distinct category,
in the bonobo it is part and parcel of social relations – and not
just between males and females. Bonobos engage in sex in virtually
every partner combination (although such contact among close family
members may be suppressed). And sexual interactions occur more often
among bonobos than among other primates. Despite the frequency of
sex, the bonobo's rate of reproduction in the wild is about the same
as that of the chimpanzee. A female gives birth to a single infant
at intervals of between five and six years. So bonobos share at
least one very important characteristic with our own species,
namely, a partial separation between sex and reproduction.
A Near Relative
finding commands attention because the bonobo shares more than 98
percent of our genetic profile, making it as close to a human as,
say, a fox is to a dog. The split between the human line of ancestry
and the line of the chimpanzee and the bonobo is believed to have
occurred a mere eight million years ago. The subsequent divergence
of the chimpanzee and the bonobo lines came much later, perhaps
prompted by the chimpanzee's need to adapt to relatively open, dry
habitats [see "East Side Story: The Origin of Humankind," by Yves
Coppens; SCIENTIFIC AMERICAN, May 1994].
In contrast, bonobos probably never left the protection of the
trees. Their present range lies in humid forests south of the Zaire
River, where perhaps fewer than 10,000 bonobos survive. (Given the
species' slow rate of reproduction, the rapid destruction of its
tropical habitat and the political instability of central Africa,
there is reason for much concern about its future.)
If this evolutionary scenario of ecological continuity is true,
the bonobo may have undergone less transformation than either humans
or chimpanzees. It could most closely resemble the common ancestor
of all three modern species. Indeed, in the 1930s Harold J. Coolidge
– the American anatomist who gave the bonobo its eventual taxonomic
status – suggested that the animal might be most similar to the
primogenitor, since its anatomy is less specialized than is the
chimpanzee's. Bonobo body proportions have been compared with those
of the australopithecines, a form of prehuman. When the apes stand
or walk upright, they look as if they stepped straight out of an
artist's impression of early hominids.
Not too long ago the savanna baboon was regarded as the best
living model of the human ancestor. That primate is adapted to the
kinds of ecological conditions that prehumans may have faced after
descending from the trees. But in the late 1970s, chimpanzees, which
are much more closely related to humans, became the model of choice.
Traits that are observed in chimpanzees – including cooperative
hunting, food sharing, tool use, power politics and primitive
warfare – were absent or not as developed in baboons. In the
laboratory the apes have been able to learn sign language and to
recognize themselves in a mirror, a sign of self-awareness not yet
demonstrated in monkeys.
Although selecting the chimpanzee as the touchstone of hominid
evolution represented a great improvement, at least one aspect of
the former model did not need to be revised: male superiority
remained the natural state of affairs. In both baboons and
chimpanzees, males are conspicuously dominant over females; they
reign supremely and often brutally. It is highly unusual for a fully
grown male chimpanzee to be dominated by any female.
Enter the bonobo. Despite their common name – the pygmy
chimpanzee – bonobos cannot be distinguished from the chimpanzee by
size. Adult males of the smallest subspecies of chimpanzee weigh
some 43 kilograms (95 pounds) and females 33 kilograms (73 pounds),
about the same as bonobos. Although female bonobos are much smaller
than the males, they seem to rule.
In physique, a bonobo is as different from a chimpanzee as a
Concorde is from a Boeing 747. I do not wish to offend any
chimpanzees, but bonobos have more style. The bonobo, with its long
legs and small head atop narrow shoulders, has a more gracile build
than does a chimpanzee. Bonobo lips are reddish in a black face, the
ears small and the nostrils almost as wide as a gorilla's. These
primates also have a flatter, more open face with a higher forehead
than the chimpanzee's and – to top it all off – an attractive
coiffure with long, fine, black hair neatly parted in the middle.
Like chimpanzees, female bonobos nurse and carry around their
young for up to five years. By the age of seven the offspring reach
adolescence. Wild females give birth for the first time at 13 or 14
years of age, becoming full grown by about 15. A bonobo's longevity
is unknown, but judging by the chimpanzee it may be older than 40 in
the wild and close to 60 in captivity.
Fruit is central to the diets of both wild bonobos and
chimpanzees. The former supplement with more pith from herbaceous
plants, and the latter add meat. Although bonobos do eat
invertebrates and occasionally capture and eat small vertebrates,
including mammals, their diet seems to contain relatively little
animal protein. Unlike chimpanzees, they have not been observed to
Whereas chimpanzees use a rich array of strategies to obtain
foods – from cracking nuts with stone tools to fishing for ants and
termites with sticks – tool use in wild bonobos seems undeveloped.
(Captive bonobos use tools skillfully.) Apparently as intelligent as
chimpanzees, bonobos have, however, a far more sensitive
temperament. During World War II bombing of Hellabrun, Germany, the
bonobos in a nearby zoo all died of fright from the noise; the
chimpanzees were unaffected.
Bonobos are also imaginative in play. I have watched captive
bonobos engage in "blindman's buff." A bonobo covers her eyes with a
banana leaf or an arm or by sticking two fingers in her eyes. Thus
handicapped, she stumbles around on a climbing frame, bumping into
others or almost falling. She seems to be imposing a rule on
herself: "I cannot look until I lose my balance." Other apes and
monkeys also indulge in this game, but I have never seen it
performed with such dedication and concentration as by bonobos.
bonobos are incurably playful and like to make funny faces,
sometimes in long solitary pantomimes and at other times while
tickling one another. Bonobos are, however, more controlled in
expressing their emotions – whether it be joy, sorrow, excitement or
anger – than are the extroverted chimpanzees. Male chimpanzees often
engage in spectacular charging displays in which they show off their
strength: throwing rocks, breaking branches and uprooting small
trees in the process. They keep up these noisy performances for many
minutes, during which most other members of the group wisely stay
out of their way. Male bonobos, on the other hand, usually limit
displays to a brief run while dragging a few branches behind them.
Both primates signal emotions and intentions through facial
expressions and hand gestures, many of which are also present in the
nonverbal communication of humans. For example, bonobos will beg by
stretching out an open hand (or, sometimes, a foot) to a possessor
of food and will pout their lips and make whimpering sounds if the
effort is unsuccessful. But bonobos make different sounds than
chimpanzees do. The renowned low-pitched, extended "huuu- huuu"
pant-hooting of the latter contrasts with the rather sharp,
high-pitched barking sounds of the bonobo.
My own interest in bonobos came not from an inherent fascination
with their charms but from research on aggressive behavior in
primates. I was particularly intrigued with the aftermath of
conflict. After two chimpanzees have fought, for instance, they may
come together for a hug and mouth-to-mouth kiss. Assuming that such
reunions serve to restore peace and harmony, I labeled them
Peace it is good for the economy
Any species that combines close
bonds with a
potential for conflict needs such conciliatory mechanisms.
Thinking how much faster marriages would break up if people had no
way of compensating for hurting each other, I set out to investigate
such mechanisms in several primates, including bonobos. Although I
expected to see peacemaking in these apes, too, I was little
prepared for the form it would take.
For my study, which began in 1983, I chose the
San Diego Zoo. At the time, it housed the world's largest
captive bonobo colony – 10 members divided into three groups. I
spent entire days in front of the enclosure with a video camera,
which was switched on at feeding time. As soon as a caretaker
approached the enclosure with food, the males would develop
erections. Even before the food was thrown into the area, the
bonobos would be inviting each other for sex: males would invite
females, and females would invite males and other females.
it turned out, is the key to the social life of the bonobo. The
first suggestion that the sexual behavior of bonobos is different
had come from observations at European zoos. Wrapping their findings
in Latin, primatologists Eduard Tratz and Heinz Heck reported in
1954 that the chimpanzees at Hellabrun mated more canum (like dogs)
and bonobos more hominum (like people). In those days, face-to- face
copulation was considered uniquely human, a cultural innovation that
needed to be taught to preliterate people (hence the term
"missionary position"). These early studies, written in German, were
ignored by the international scientific establishment. The bonobo's
humanlike sexuality needed to be rediscovered in the 1970s before it
became accepted as characteristic of the species.
become sexually aroused remarkably easily, and they express this
excitement in a variety of mounting positions and genital contacts.
Although chimpanzees virtually never adopt face-to-face positions,
bonobos do so in one out of three copulations in the wild.
Furthermore, the frontal orientation of the bonobo vulva and
clitoris strongly suggest that the female genitalia are adapted for
Another similarity with humans is increased female sexual
receptivity. The tumescent phase of the female's genitals, resulting
in a pink swelling that signals willingness to mate, covers a much
longer part of estrus in bonobos than in chimpanzees. Instead of a
few days out of her cycle, the female bonobo is almost continuously
sexually attractive and active.
Perhaps the bonobo's most typical sexual pattern, undocumented in
any other primate, is genito-genital rubbing (or GG rubbing) between
adult females. One female facing another clings with arms and legs
to a partner that, standing on both hands and feet, lifts her off
the ground. The two females then rub their genital swellings
laterally together, emitting grins and squeals that probably reflect
orgasmic experiences. (Laboratory experiments on stump- tailed
macaques have demonstrated that women are not the only female
primates capable of physiological orgasm.)
Male bonobos, too, may engage in pseudocopulation but generally
perform a variation. Standing back to back, one male briefly rubs
his scrotum against the buttocks of another. They also practice
so-called penis-fencing, in which two males hang face to face from a
branch while rubbing their erect penises together.
The diversity of erotic contacts in bonobos includes sporadic
oral sex, massage of another individual's genitals and intense
tongue-kissing. Lest this leave the impression of a pathologically
oversexed species, I must add, based on hundreds of hours of
watching bonobos, that their sexual activity is rather casual and
relaxed. It appears to be a completely natural part of their group
life. Like people, bonobos engage in sex only occasionally, not
continuously. Furthermore, with the average copulation lasting 13
seconds, sexual contact in bonobos is rather quick by human
That sex is connected to feeding, and even appears to make food
sharing possible, has been observed not only in zoos but also in the
wild. Nancy Thompson-Handler, then at the State University of New
York at Stony Brook, saw bonobos in Zaire's Lomako Forest engage in
sex after they had entered trees loaded with ripe figs or when one
among them had captured a prey animal, such as a small forest
duiker. The flurry of sexual contacts would last for five to 10
minutes, after which the apes would settle down to consume the food.
One explanation for the sexual activity at feeding time could be
that excitement over food translates into sexual arousal. This idea
may be partly true. Yet another motivation is probably the real
cause: competition. There are two reasons to believe sexual activity
is the bonobo's answer to avoiding conflict.
First, anything, not just food, that arouses the interest of more
than one bonobo at a time tends to result in sexual contact. If two
bonobos approach a cardboard box thrown into their enclosure, they
will briefly mount each other before playing with the box. Such
situations lead to squabbles in most other species. But bonobos are
quite tolerant, perhaps because they use sex to divert attention and
to diffuse tension.
Second, bonobo sex often occurs in aggressive contexts totally
unrelated to food. A jealous male might chase another away from a
female, after which the two males reunite and engage in scrotal
rubbing. Or after a female hits a juvenile, the latter's mother may
lunge at the aggressor, an action that is immediately followed by
genital rubbing between the two adults.
I once observed a young male, Kako, inadvertently blocking an
older, female juvenile, Leslie, from moving along a branch. First,
Leslie pushed him; Kako, who was not very confident in trees,
tightened his grip, grinning nervously. Next Leslie gnawed on one of
his hands, presumably to loosen his grasp. Kako uttered a sharp peep
and stayed put. Then Leslie rubbed her vulva against his shoulder.
This gesture calmed Kako, and he moved along the branch. It seemed
that Leslie had been very close to using force but instead had
reassured both herself and Kako with sexual contact.
During reconciliations, bonobos use the same sexual repertoire as
they do during feeding time. Based on an analysis of many such
incidents, my study yielded the first solid evidence for sexual
behavior as a mechanism to overcome aggression. Not that this
function is absent in other animals – or in humans, for that matter
– but the art of sexual reconciliation may well have reached its
evolutionary peak in the bonobo. For these animals, sexual behavior
is indistinguishable from social behavior. Given its
appeasement functions, it is not surprising that sex among bonobos
occurs in so many different partner combinations, including between
juveniles and adults. The need for peaceful coexistence is obviously
not restricted to adult heterosexual pairs.
Apart from maintaining harmony, sex is also involved in creating the
singular social structure of the bonobo. This use of sex becomes
clear when studying bonobos in the wild. Field research on bonobos
started only in the mid-1970s, more than a decade after the most
important studies on wild chimpanzees had been initiated. In terms
of continuity and invested (wo)manpower, the chimpanzee projects of
Jane Goodall and Toshisada Nishida, both in Tanzania, are
unparalleled. But bonobo research by Takayoshi Kano and others of
Kyoto University is now two decades under way at Wamba in Zaire and
is beginning to show the same payoffs.
Both bonobos and chimpanzees live in so-called fission – fusion
societies. The apes move alone or in small parties of a few
individuals at a time, the composition of which changes constantly.
Several bonobos traveling together in the morning might meet another
group in the forest, whereupon one individual from the first group
wanders off with others from the second group, while those left
behind forage together. All associations, except the one between
mother and dependent offspring, are of a temporary character.
Initially this flexibility baffled investigators, making them
wonder if these apes formed any social groups with stable
membership. After years of documenting the travels of chimpanzees in
the Mahale Mountains, Nishida first reported that they form large
communities: all members of one community mix freely in ever
changing parties, but members of different communities never gather.
Later, Goodall added territoriality to this picture. That is, not
only do communities not mix, but males of different chimpanzee
communities engage in lethal battles.
In both bonobos and chimpanzees, males stay in their natal group,
whereas females tend to migrate during adolescence. As a result, the
senior males of a chimpanzee or bonobo group have known all junior
males since birth, and all junior males have grown up together.
Females, on the other hand, transfer to an unfamiliar and often
hostile group where they may know no one. A chief difference between
chimpanzee and bonobo societies is the way in which young females
integrate into their new community.
On arrival in another community, young bonobo females at Wamba
single out one or two senior resident females for special attention,
using frequent GG rubbing and grooming to establish a relation. If
the residents reciprocate, close associations are set up, and the
younger female gradually becomes accepted into the group. After
producing her first offspring, the young female's position becomes
more stable and central. Eventually the cycle repeats with younger
immigrants, in turn, seeking a good relation with the now
established female. Sex thus smooths the migrant's entrance into the
community of females, which is much more close-knit in the bonobo
than in the chimpanzee.
Bonobo males remain attached to their mothers all their lives,
following them through the forest and being dependent on them for
protection in aggressive encounters with other males. As a result,
the highest-ranking males of a bonobo community tend to be sons of
What a contrast with chimpanzees! Male chimpanzees fight their
own battles, often relying on the support of other males.
Furthermore, adult male chimpanzees travel together in same-sex
parties, grooming each other frequently. Males form a distinct
social hierarchy with high levels of both competition and
association. Given the need to stick together against males of
neighboring communities, their bonding is not surprising: failure to
form a united front might result in the loss of lives and territory.
The danger of being male is reflected in the adult sex ratio of
chimpanzee populations, with considerably fewer males than females.
Serious conflict between bonobo groups has been witnessed in the
field, but it seems quite rare. On the contrary, reports exist of
peaceable mingling, including mutual sex and grooming, between what
appear to be different communities. If intergroup combat is indeed
unusual, it may explain the lower rate of all-male associations.
Rather than being male- bonded, bonobo society gives the impression
of being female- bonded, with even adult males relying on their
mothers instead of on other males. No wonder Kano calls mothers the
"core" of bonobo society.
The bonding among female bonobos violates a fairly general rule,
outlined by Harvard University anthropologist Richard W. Wrangham,
that the sex that stays in the natal group develops the strongest
mutual bonds. Bonding among male chimpanzees follows naturally
because they remain in the community of their birth. The same is
true for female kinship bonding in Old World monkeys, such as
macaques and baboons, where males are the migratory sex.
Bonobos are unique in that the migratory sex, females, strongly
bond with same-sex strangers later in life. In setting up an
artificial sisterhood, bonobos can be said to be secondarily bonded.
(Kinship bonds are said to be primary.) Although we now know HOW
this happens – through the use of sexual contact and grooming – we
do not yet know WHY bonobos and chimpanzees differ in this respect.
The answer may lie in the different ecological environments of
bonobos and chimpanzees – such as the abundance and quality of food
in the forest. But it is uncertain if such explanations will
society is, however, not only female-centered but also appears to be
female-dominated. Bonobo specialists, while long suspecting such a
reality, have been reluctant to make the controversial claim. But in
1992, at the 14th Congress of the International Primatological
Society in Strasbourg, investigators of both captive and wild
bonobos presented data that left little doubt about the issue.
Amy R. Parish of the University of California at Davis reported
on food competition in identical groups (one adult male and two
adult females) of chimpanzees and bonobos at the Stuttgart Zoo.
Honey was provided in a "termite hill" from which it could be
extracted by dipping sticks into a small hole. As soon as honey was
made available, the male chimpanzee would make a charging display
through the enclosure and claim everything for himself. Only when
his appetite was satisfied would he let the females fish for honey.
In the bonobo group, it was the females that approached the honey
first. After having engaged in some GG rubbing, they would feed
together, taking turns with virtually no competition between them.
The male might make as many charging displays as he wanted; the
females were not intimidated and ignored the commotion.
Observers at the Belgian animal park of Planckendael, which
currently has the most naturalistic bonobo colony, reported similar
findings. If a male bonobo tried to harass a female, all females
would band together to chase him off. Because females appeared more
successful in dominating males when they were together than on their
own, their close association and frequent genital rubbing may
represent an alliance. Females may bond so as to outcompete members
of the individually stronger sex.
The fact that they manage to do so not only in captivity is
evident from zoologist Takeshi Furuichi's summary of the relation
between the sexes at Wamba, where bonobos are enticed out of the
forest with sugarcane. "Males usually appeared at the feeding site
first, but they surrendered preferred positions when the females
appeared. It seemed that males appeared first not because they were
dominant, but because they had to feed before the arrival of
females," Furuichi reported at Strasbourg.
Occasionally, the role of sex in relation to food is taken one
step further, bringing bonobos very close to humans in their
behavior. It has been speculated by anthropologists – including C.
Owen Lovejoy of Kent State University and Helen Fisher of Rutgers
University – that sex is partially separated from reproduction in
our species because it serves to cement mutually profitable
relationships between men and women. The human female's capacity to
mate throughout her cycle and her strong sex drive allow her to
exchange sex for male commitment and paternal care, thus giving rise
to the nuclear family.
This arrangement is thought to be favored by natural selection
because it allows women to raise more offspring than they could if
they were on their own. Although bonobos clearly do not establish
the exclusive heterosexual bonds characteristic of our species,
their behavior does fit important elements of this model. A female
bonobo shows extended receptivity and uses sex to obtain a male's
favors when – usually because of youth – she is too low in social
status to dominate him.
the San Diego Zoo, I observed that if Loretta was in a sexually
attractive state, she would not hesitate to approach the adult male,
Vernon, if he had food. Presenting herself to Vernon, she would mate
with him and make high- pitched food calls while taking over his
entire bundle of branches and leaves. When Loretta had no genital
swelling, she would wait until Vernon was ready to share.
Primatologist Suehisa Kuroda reports similar exchanges at Wamba: "A
young female approached a male, who was eating sugarcane. They
copulated in short order, whereupon she took one of the two canes
held by him and left."
Despite such quid pro quo between the sexes, there are no
indications that bonobos form humanlike nuclear families. The burden
of raising offspring appears to rest entirely on the female's
shoulders. In fact, nuclear families are probably incompatible with
the diverse use of sex found in bonobos. If our ancestors started
out with a sex life similar to that of bonobos, the evolution of the
family would have required dramatic change.
Human family life implies paternal investment, which is unlikely
to develop unless males can be reasonably certain that they are
caring for their own, not someone else's, offspring. Bonobo society
lacks any such guarantee, but humans protect the integrity of their
family units through all kinds of moral restrictions and taboos.
Thus, although our species is characterized by an extraordinary
interest in sex, there are no societies in which people engage in it
at the drop of a hat (or a cardboard box, as the case may be). A
sense of shame and a desire for domestic privacy are typical human
concepts related to the evolution and cultural bolstering of the
Yet no degree of moralizing can make sex disappear from every
realm of human life that does not relate to the nuclear family. The
bonobo's behavioral peculiarities may help us understand the role of
sex and may have serious implications for models of human society.
Just imagine that we had never heard of chimpanzees or baboons
and had known bonobos first. We would at present most likely believe
that early hominids lived in female- centered societies, in which
sex served important social functions and in which warfare was rare
or absent. In the end, perhaps the most successful reconstruction of
our past will be based not on chimpanzees or even on bonobos but on
a three-way comparison of chimpanzees, bonobos and humans.
Social Organization among Various Primates
- Bonobo communities are peace-loving and generally
egalitarian. The strongest social bonds are those among females,
although females also bond with males. The status of a male
depends on the position of his mother, to whom he remains
closely bonded for her entire life.
- In chimpanzee groups the strongest bonds are established
between the males in order to hunt and to protect their shared
territory. The females live in overlapping home ranges within
this territory but are not strongly bonded to other females or
to any one male.
- Gibbons establish monogamous, egalitarian relations, and one
couple will maintain a territory to the exclusion of other
- Human society is the most diverse among the primates. Males
unite for cooperative ventures, whereas females also bond with
those of their own sex. Monogamy, polygamy and polyandry are all
- The social organization of gorillas provides a clear example
of polygamy. Usually a single male maintains a range for his
family unit, which contains several females. The strongest bonds
are those between the male and his females.
- Orangutans live solitary lives with little bonding in
evidence. Male orangutans are intolerant of one another. In his
prime, a single male establishes a large territory, within which
live several females. Each female has her own, separate home
To find out more about bonobos, check out
The Pygmy Chimpanzee : Evolutionary Biology and Behavior
Edited by Randall L. Susman. Plenum Press, 1984.
- THE COMMUNICATIVE REPERTOIRE OF CAPTIVE BONOBOS (PAN
PANISCUS) COMPARED TO THAT OF CHIMPANZEES. F.B.M. de Waal in "Behaviour,"
Vol. 106, Nos. 3-4, pages 183-251; September 1988.
Peacemaking Among Primates F.B.M. de Waal. Harvard
University Press, 1989.
Understanding Chimpanzees (Special Publication) Edited by
Paul Heltne and Linda A. Marquardt. Harvard University Press,
The Last Ape : Pygmy Chimpanzee Behavior and Ecology by
Takayoshi Kano. Stanford University Press, 1992.
Chimpanzee Cultures by R. Wrangham, W. C. McGrew, F.B.M. de
Waal and P. Heltne. Harvard University Press, 1994.
Bonobo The Forgotten Ape by Frans de Waal, 1997.
- Brass Monkey
Mystery Linked To The Bonobos Tribe
- Are the
Bonobos the Brass Monkey
- I found the Brass
- Bonobo Sex and Society
The Brass Monkey
- Is Sex and Touch the
answer to World Conflict ? Is this the Brass Monkey?
What is the Brass Monkey