Evolution And History

82 CHAPTER 4 | Primate Behavior

low-ranking females, and groups of females form alliances

in which they may cooperatively attack males, to the point

of inflicting blood-drawing injuries.^8 Thus instead of the

male dominance characteristic of chimps, one sees female

dominance.

Western primatologists’ focus on social rank and attack

behavior may be a legacy of the individualistic, competi-

tive nature of the societies in which evolutionary theory

originated. To a certain degree, natural selection relies

upon a struggle among living creatures rather than peace-

ful coexistence. By contrast, noted Japanese primatologist

Kinji Imanishi (see Anthropologists of Note) developed a

harmonious theory of evolution and initiated field stud-

ies of bonobos that have demonstrated the importance of

social cooperation rather than competition. As the work

of Dutch primatologist Frans de Waal illustrates in the fol-

lowing Original Study, reconciliation after an attack may

be even more important from an evolutionary perspective

than the actual attack.

like Goodall, the nuances of primate social behavior and the
importance of female primates have been documented. High-
ranking (alpha) females may dominate low-ranking males.
In groups such as bonobos, females dominate overall. While
strength and size contribute to an animal’s rank, other impor-
tant factors include the rank of its mother and effectiveness
at creating alliances with other individuals. For males, drive
or motivation to achieve high status also influences rank. For
example, in the community studied by Goodall, one male
chimp hit upon the idea of incorporating noisy kerosene cans
into his charging displays, thereby intimidating all the other
males.^7 As a result, he rose from relatively low status to the
number one (alpha) position.
Among bonobos, female–female bonds play an im-
portant role in determining rank. Further, the strength of
the bond between mother and son may interfere with the
ranking among males. Not only do bonobo males defer to
females in feeding, but alpha females have been observed
chasing high-ranking males. Alpha males even yield to

(^7) Goodall, J. (1986). The chimpanzees of Gombe: Patterns of behavior
(p. 424). Cambridge, MA: Belknap.
(^8) de Waal, F. B. M., Kano, T., & Parish, A. R. (1998). Comments. Current
Anthropology 39, 408, 410, 413.
Original Study

Reconciliation and Its Cultural Modification

in Primates by Frans B. M. de Waal

Despite the continuing popularity of the

struggle-for-life metaphor, it is increas-

ingly recognized that there are draw-

backs to open competition,

hence that there are sound

evolutionary reasons for curb-

ing it. The dependency of so-

cial animals on group life and

cooperation makes aggression

a socially costly strategy. The

basic dilemma facing many

animals, including humans,

is that they sometimes cannot

win a fight without losing a

friend.

This photo shows what may

happen after a conflict—in

this case between two female

bonobos. About 10 minutes

after their fight, the two fe-

males approach each other,

with one clinging to the other

and both rubbing their clitorises and

genital swellings together in a pattern

known as genito-genital rubbing, or GG-

rubbing. This sexual contact, typical

of bonobos, constitutes a so-called

reconciliation. Chimpanzees, which are

closely related to bonobos (and to us:

bonobos and chimpanzees are our clos-

est animal relatives), usually reconcile in

a less sexual fashion, with an embrace

and mouth-to-mouth kiss.

There is now evidence for reconcili-

ation in more than twenty-five different

primate species, not just in apes but

also in many monkeys. The

same sorts of studies have

been conducted on human

children in the schoolyard,

and of course children show

reconciliation as well. Re-

searchers have even found

reconciliation in dolphins,

spotted hyenas, and some

other non primates. Recon-

ciliation seems widespread:

a common mechanism found

whenever relationships need

to be maintained despite oc-

casional conflict.a,b

The definition of reconcili-

ation used in animal research

is a friendly reunion between

former opponents not long after

a conflict. This is somewhat different from

definitions in the dictionary, primarily be-

cause we look for an empirical definition

that is useful in observational studies—in

Two adult female bonobos engage in so-called GG-rubbing, a

sexual form of reconciliation typical of this species.

© Amy Parish/Anthro-Photo