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Darwins Monkey: Why Baboons Cant Become Human
Shirley C. Strum1,2,3*
1
Department of Anthropology, University of California, San Diego, La Jolla, CA 92093-0532
2
Uaso Ngiro Baboon Project, Box 62844, Nairobi, 00200, Kenya
3
Institute of Primate Research, National Museums of Kenya, Box 24481-00502, Karen, Nairobi, Kenya
KEY WORDS baboons; Papio anubis; natural history; complexity; negotiation; dominance;
collaboration; natural selection; human evolution; Anthropocene
ABSTRACT Baboons were used in the past as mod-
els for human evolution. I utilize 40 years of data from
my long-term study on baboons in Kenya to suggest that
baboons are once again relevant for understanding
human evolution, not as a referential model but to reset
the starting conditions of the human experiment. The
baboon data also offer a critique of widely held ideas
about how natural selection might work by looking at
real lives in real time. This situates competition in a ma-
trix of collaboration and illustrates the critical role of
chance, contingency, and history in baboon survival and
success. I make three methodological moves to reach
these conclusions. The first is to focus on process not
just outcome. The second is to look at time scales longer
than usual studies but shorter than evolutionary time as
a way to open the black box that currently links behav-
ior to evolutionary value. The third is to use compara-
tive natural history, Darwins method, as a way to cap-
ture and comprehend how complexity is generated and
how baboons deal with it in their daily lives. These em-
pirical and methodological turns lead to conclusions that
run counter to widely held ideas about baboons, about
primates, and about the determinism of natural selec-
tion. I follow my own research history to illustrate these
points. The discussion ranges from alternative interpre-
tations of the male and the female dominance hierar-
Baboons have played a central role both in our under-
standing of primates and as models for human evolution
beginning with Zuckermans 1932 description of primate
society. Based on his observations of Hamadryas baboons
at the London Zoo, he argued for a society founded on
sex, violence, and male domination of both females and
males (Zuckerman, 1932, 1933). Zuckerman can be for-
given for his mistakes because we knew very little about
primates in nature at that time (Strum and Fedigan,
2000b). Baboons continued to capture the scientific (and
public) imagination three decades later through the
early scientific work on savanna baboons done by
Washburn, his student, DeVore, and the English psychol-
ogist, Hall (DeVore, 1965). Once again baboons became
emblematic of evolution; however, this time it was the fit
between behavior and anatomy which told a convincing
story about how natural selection worked (Washburn
and DeVore, 1961, 1963; DeVore and Hall, 1965; Hall
and DeVore, 1965; Strum and Fedigan, 2000b). Evolution
gave male baboons the anatomy of aggression. Baboon
males used those remarkable canines, large body size,
impressive mantle of hair, and white eyelids in displays
during sexual competition with other males and against
predators. Fighting ability determined male rank which
in turn governed reproductive success so that the best
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YEARBOOK OF PHYSICAL ANTHROPOLOGY 55:323 (2012)
chies, to insights from a fission that happened when the
foraging strategy of raiding and nonraiding baboons
diverged, to evidence of adaptation after translocation,
and finally to assessing two unusual fusions of baboon
groups. Altogether, these natural histories yield baboon
principles of the social with insights about cognition,
cooperation, and culture and suggest why baboons cant
become human. The data also support Weiss and
Buchanans framework (The Mermaids Tale: Four Bil-
lion Years of Cooperation in the Making of Living
Things. Cambridge, MA: Harvard University Press,
2009. 305 p) with its alternative view of natural selec-
tion in which there is more slippage and tolerance, mul-
tiple solutions with larger acceptability spaces, and the
possibility that an adaptive fit will be good enough
rather than seamless. However, capturing behavioral
complexity in the wild poses methodological challenges.
Long-term field studies provide critical information but
the current quantitative methods should be expanded
also include natural history observations of behaviors
and events across time, space, groups, and landscapes.
Finally, the baboon natural histories illustrate how the
evolutionary game has changed in the Anthropocene
yielding a cautionary tale about the future for many
primate species. Yrbk Phys Anthropol 55:323, 2012.
C 2012
V Wiley Periodicals, Inc.
males were the most reproductive. The male dominance
hierarchy also was the political structure of the group; it
ordered male interactions reducing the need for frequent
aggressive encounters. Males defended and policed;
females raised babies. Males were central to all major
group functions, whereas females were defined in terms
of their relationships with males. Washburn and DeVore
Grant sponsors: National Science Foundation, Wenner-Gren
Foundation for Anthropological Research, University of California,
San Diego, National Geographic Society, Fyseen Foundation, Wild-
life Conservation International, LSB Leakey Foundation, Art Orten-
berg and Liz Claiborne Foundation, African Conservation Centre,
African Conservation Fund.
*Correspondence to: Shirley C. Strum, Department of Anthropol-
ogy, University of California, 9500 Gillman Drive, San Diego, La
Jolla, CA 92093-0532, USA. E-mail: sstrum@africaonline.co.ke (or)
sstrum@ucsd.edu
Received 6 September 2012; accepted 7 September 2012
DOI 10.1002/ajpa.22158
Published online 17 October 2012 in Wiley Online Library
(wileyonlinelibrary.com).
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4 S.C. STRUM
(1961) argued that we could infer the challenges that the
savanna presented to a puny primate living away from
the safety of the forest (read hominids) by looking
through the lens of the only other primate that managed
to do so successfully, baboons.
What follows is not a scholarly review of the baboon
literature and its relevance to human evolution.
Instead, I was invited to discuss my insights about
baboons based on 40 years of observing them in the
wild. Baboons have long since been dethroned as the change. This is particularly relevant to the scale and the
preferred species, first by common chimpanzees
(Goodall, 1988/1971) and then by bonobos (Zihlman,
1978; Kano, 1992; Strum and Fedigan, 2000b). In con-
trast, I will suggest that baboons are once again rele-
vant for those interested in human evolution but in
new ways. What follows is an argument that is mainly
bottom up, grounded in baboon observations. In the
end, I hope to illustrate that the baboons I have studied
live in the most complex nonhuman primate society yet
described and that they reside in the most socioecologi-
cal complexity documented so far for a nonhuman pri-
mate. Furthermore, these baboon data suggest that fo-
cusing on real time and real lives provides a different
view of how evolutionary processes work. Chance, con-
tingency, and history play an important role in survival
and success. This offers a critique of widely held ideas
about natural selection, particularly the centrality of
competition. The data also argue for the existence of
multiple options (including mistakes) made possible by
the intelligence and social skill that baboons use to
build their primate society. I am often asked why, if
baboons are so smart, they arent human. There are
many things wrong with the way this question is
phrased. However, the perspective I present below does
add a new factor to the list of obvious differences: the
importance of managing complexity in daily life.
To reach these conclusions, I made three critical meth-
odological moves. The first was to focus on process not
just outcome. The second was to look at time scales longer
than usual studies but not collapse everything into evolu-
tionary time. The third was to use comparative natural
history, Darwins method. Primate studies rejected natu-
ral history observations when growing knowledge about
primates grew and more sophisticated data collection
methods were available (Strum and Fedigan, 2000a). For
now, however, natural history may be the only way to
capture and comprehend how complexity is generated
and how baboons deal with it in their daily lives.
These empirical and methodological turns lead to con-
clusions that run counter to widely held ideas about
baboons, about primates, and about the determinism of
natural selection. This is not simply an emphasis on
proximate rather than ultimate levels of causal explana-
tion (Tinbergen, 1963). Instead, it tries to open the black
box linking behavior and evolutionary outcome and
raises the question of what can be legitimately assumed
about shorter time scales when we make evolutionary
time arguments. I do not review the extensive literature
on all these topics. Instead, I include key references as a
starting point for readers interested in exploring further.
My reference species is olive baboons, Papio anubis
(Zinner et al., 2009). The reference material is the 40
years of my own research (see Strum and Fedigan,
2000b) for a framework to situate most of these years). I
track individuals, relationships, and social groups in two
populations. The baboons lived in two locations because I
translocated three groups in 1984 (Strum and Southwick,
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1987; Strum, 2005). Translocation was an experiment
that provided persuasive evidence about baboon adapta-
bility. The new location, arid savanna occupied by no-
madic pastoralists, has had both natural cycles of extreme
events like droughts and in the last decade, human-
induced changes that also created extreme events. Else-
where (Strum and Western, 1982), I have argued that the
baboons dont distinguish between the sources (natural
vs. human) of environmental change in their responses to
processes I will explore. In this presentation, I touch on
areas that are minefields of controversy, for example,
what is culture or tradition and what is cooperation. For-
tunately, my argument does not depend on resolving
these controversies, a task I leave to others.
In this article, I move through a series of steps that
reflect my own research history rather than their order
of importance. During this time, I have gone from mak-
ing baboons almost human by focusing on their social
negotiation and politics (Strum, 2001/1987a) to under-
standing better why baboons arent human. I have col-
lected quantitative data for four decades; however, I find
these data inadequate to make sense of what the
baboons are doing. Instead, I turn to comparative natu-
ral history for the missing insights. The title Darwins
Monkey refers both to the importance of Darwins meth-
odology as well as his prescient observations of baboons
(Darwin, 1871/2004).
My over-arching goal, in this article and in my
research, is to recapture the complexity that exists in ba-
boon lives and then to consider the implications of this
complexity for evolutionary interpretations and for our
current scientific practice. I end with a discussion of why
understanding baboons is important for [physical/biologi-
cal] anthropologists today.
PART 1: GENERATING COMPLEXITY
Remodeling the male dominance hierarchy
makes baboons almost human (19721980)
It is hard to believe in the 21st century how little we
knew about primate behavior in the 1960s and how rudi-
mentary were our methods. The pioneering work of the
classical ethologists like Tinbergen, Lorenz, and von
Frisch starting in the 1930s (Tinbergen, 1942, 1963; Lor-
enz, 1950) did two key things. It critiqued (previous)
anthropomorphic interpretations of behavior and created
a novel methodology that still relied on natural history
[ethogram, fixed action patterns; critical periods; and
interaction of field data with captive experiments (Tin-
bergen, 1974/1951)].
When primate field studies resumed after World War
2, scientists still used natural history descriptions but
added the ethogram, a careful and detailed description
of the behavior of that species (or group, or popula-
tion. . .these terms were interchangeable then). In the
middle of last century, there were only dozens of studies
of primates in the wild, not hundreds, and only a few
species had been observed in more than one location
(DeVore, 1965; Altmann, 1967; Jay, 1968).
Variation in behavior seemed minimal, making broad
generalizations about primate patterns seemed war-
ranted (Washburn and DeVore, 1961; Strum and Fedi-
gan, 2000b). However, by the early 1970s, variation in
behavior and social organization began to accumulate.
For example, patas monkeys solved the problems of sa-
vanna living with an anatomy adapted for rapid escape
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DARWINS MONKEY
and behavior adapted for vigilant diversionary males
and stealthy females and young (Hall, 1968). While
Washburn and DeVores dominant males strode out to
face a predator, Rowells male baboons were the first to
run to the trees, leaving the females and youngsters
exposed (Rowell, 1966). Natural history methods were
necessary whenever a new species was studied; however,
disagreements of fact were hard to resolve. Were the
animals really different or were differences an artifact of
observers and ways of observing?
When I began my baboon study on male and female
roles in olive baboon society (19721974, Kekopey near
Gilgil, Rift Valley, Kenya), there was already evidence
from rhesus macaques on Cayo Santiago Island in the Ca-
ribbean that females, as well as males, had a dominance
hierarchy and that matrilineal relationships were the
core of the group (Sade, 1967, 1972). As well, there was a
move to use more systematic ways of data collection in
the field [well summarized by Jeanne Altmanns often
cited paper (Altmann, 1974)]. All baboon projects from the
early 1970s onward inherited a baboon ethogram from
Hall, Washburn, DeVore, Rowell, and Ransom (Washburn
and DeVore, 1961; DeVore and Hall, 1965; Hall and
DeVore, 1965; Rowell, 1966; Ransom, 1981). This meant
that the data included the same basic elements of behav-
ior. When possible (meaning in contexts where the prima-
tes could actually be seen), investigators relied on focal
animal samples (follows for a fixed duration of randomly
chosen individuals) as a way to minimize bias.
I collected my data this way. The results confirmed
that matrilines (female based families) were the core of
the group and that a matrilineal dominance hierarchy
was the troops main structure. Families, rather than
just adult males, protected and policed the group on
behalf of their own members. Females had political
functions besides motherhood (see also Hausfater, 1975;
Ransom, 1981).
However, the males I studied didnt behave as pre- replaced by the term friendship in the literature begin-
dicted. The category males includes large adolescents
reaching the asymptote of their growth (Strum, 1991)
and fully adult males ranging in age from young ([10
years) to old (more than 20 years). Hausfater, on yellow
baboons in Amboseli National Park in Kenya (Hausfater,
1975; Hausfater et al., 1982), documented a relatively
stable male dominance hierarchy that operated as pre-
dicted. The dominant male(s) had priority of access to
limited resources and mating opportunities. The
Hausfater study was a careful and quantitative refine-
ment of the previous baboon model. By contrast, I found
male rank relationships exceedingly dynamic (Strum,
1982). Individuals spent time and energy working on
their relationships with each other and tracking the
changes in all male relationships. Processes of male mat-
uration and male migration fueled the instability.
Besides its dynamic nature, the existing male hierar-
chy did not correlate with access to resources such as
receptive females. Often rank did not even predict the
winner of an aggressive encounter (Strum, 1982). The
reason was that baboons, including the males, had alter-
natives to aggression. These social strategies of competi-
tion and defense (Strum, 1975a, 1976; Western and
Strum, 1983) could be as effective and were less risky
than aggressive approaches. However, they depended on
a number of factors for success. First, some relied on
special social relationships. Ransom (1981) identified
special relationships between males and females and troop predicted whether he used aggression or social
males and infants in the Gombe Stream baboons in his
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5
1967 study but hadnt yet appreciated their function.
Kekopey males created special relationships and later
used them to advantage. For example, males used young
infants as agonistic buffers against other males (Strum,
1983c). Usually, it was the potential loser who got an
infant, returning to the protagonist with the baby on his
belly. Most often, the aggressive male backed off. A male
could also use a female as a buffer (Strum, 1983b). Why
did it work?because buffering allowed a male to mobi-
lize troop support. Normally, other group members dont
get involved in malemale interactions. However, the
group will mob a male whose aggression causes distress
to infants and females. A pivotal element in buffering as
a successful tactic is trust. Trust is a social construct for
humans and for baboons. It makes social life predictable,
creates a sense of community, and makes it easier for
individuals to work together (Holmes and Rempel, 1989;
Misztal, 1996, 2001; Rousseau et al., 1998; Kappeler and
van Schaik, 2006; Henrich et al., 2009; Sueur et al.,
2011b). In economics, trust is a way to reduce transac-
tion costs, creating new forms of collaboration and it is a
form of social capital. For baboons in the context of ago-
nistic buffering, trust implies a particular social history.
It means that the male has been predictably affiliative
with the infant. Both the infant and the male have
expectations of each other based on their social history.
If the buffer trusts the male friend, it will scream at the
aggressive opponent. Without trust, a male runs the risk
of himself becoming the object of mobbing if the infant
screams at him rather than at the opponent. This
implies that the social relationship comes first; using the
relationships in a social strategy comes later.
Males got other benefits from friendships with
females. Following ethological tradition, I refrained from
calling special relationships friendships until I was
convinced that baboon and human friendships shared
many characteristics (the term special relationship was
ning in the 1980s). There is no rape among baboons, and
therefore, while males might fight to get a temporarily
exclusive association with a sexually receptive female, or
consort, the consort male needs the females coopera-
tion to successfully copulate with her. Consorts may
attract many male followers. Friendship between the
consort male and female also influenced whether a male
can keep the consort in the face of pressure from fol-
lower males (Smuts, 1985; Strum, 2001/1987b).
Other social tactics relied on social knowledge and
assessment skills. For example, older males with less
aggressive potential still got consorts and copulations.
One way was through a maneuver I called sidelines. In
sidelines, a male monitors the consort from a distance.
He makes his move when aggression breaks out between
the consort male and his close followers. Then, the more
distant monitoring male dashes to the female. By the
time the male aggression is over, the older male has
copulated with the female (becoming her new consort),
has skillfully led her away from the other males, and is
deftly grooming her. Ironically, another avenue for males
with little aggressive potential is to manipulate aggres-
sion between consort male and followers. He can do this
as a follower or from the sidelines, escalating aggression
by his involvement but usually leaving before another
male turns on him.
An individuals age and length of residency in the
strategies as well as his success (Strum, 1978, 1982). All
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6 S.C. STRUM
males followed the same social trajectory through their
life history from socially unskilled maturing male with
growing physical powers (using aggressive strategies) to
the socially skillful mature male whose success depends
entirely on social expertise when he is old (using social
strategies of competition and defense).
Males werent the only beneficiaries of social strat-
egies. Infants and females could rely on the support of
their male friend when aggression was directed at them.
Even the close proximity of a male friend reduced social
interference that a female or infant would receive based
on rank and age.
Complexity 1: Social strategies in addition to
aggressive strategies create multiple options
Science makes simplifying assumptions to be able to
begin studying complex phenomena such as social inter-
actions. The problem comes when the results of those
investigations slide into thinking that such systems are
simple. Uncoupling the tight link between male domi-
nance rank and male evolutionary payoffs was the first
evidence I had of social complexity among baboons.
Baboons created and managed special relationships.
They adjusted their behavior based on expectations from
past interactions (Strum, 2008). Social strategies were
also evidence of social collaboration (see section on col-
laboration later) and exchange. Among baboons, even
the physically impressive males needed the help of less
imposing individuals to succeed. Baboons, it seemed, had
no option but to play by the golden rule.
At this point in my research, negotiations between
baboons stood out in bold relief. Human negotiation is
usually a dialog between two or more individuals. The
dialog aims for a common understanding about a course
of actions that satisfies the interests of the parties
involved (Homans, 1958; Festinger, 1985/1957; Misztal,
2001; Raiffa, 2003/1982; Pearce and Hans-Werner, 2008;
West and Turner, 2009). I saw baboon negotiations as be-
havioral conversations about investments in, and payoffs
of, special relationships. Clearly, such interactions select
for very different skills than would aggressive competi-
tion in, for example, mating success (see also Sussman
and Garber, 2011). The conclusion: a primate with a big
brain can create more alternatives than previous evolu-
tionary models suggested. I argued that these kinds of
negotiations, social strategies, and sexual politics made
baboons almost human (Strum, 2001/1987b).
A brief note about my use of the term complexity is
warranted. The study of social complexity has generated
a huge literature since these first baboon observations
(e.g., see Byrne and Whiten, 1989; Whiten and Richard,
1997; de Waal and Tyack, 2003; Cheney, 2007). Despite
this, there is no clear definition of social complexity
(Strum et al., 1997). Whiten (2000) suggested that social
complexity must include the dimensions of variability of
response, size of group, number of relationships, instabil-
ity, and predictability. Dunbar (1998) found a strong cor-
relation between group size. Ironically, Robert Hindes
1970s model of interacting levels that moves from indi-
viduals to interactions to relationships to group structure
is probably the best approximation to an operationalized
definition of social complexity [Hinde, 1976, 1987; and
see Forster and Rodriguez (2006) for an example of how
this would work]. In another section, I offer a specific def-
inition of baboon social complexity. Until that point, I use
data to build the dimensions of baboon complexity and
use an everyday definition of complexity: being intricate,
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a system with numerous elements and many forms of
relationships which are all entwined (Latin com-
plexus); complex systems have more parts and more con-
nections than simple ones (Dictionary, 1971).
Remodeling socioecology: Group size does not
determine group fission (19811984)
Today, group size is thought to be a trade-off between
the advantages (predator protection, resource discovery,
and for primates, traditional knowledge of resources) and
disadvantages (feeding competition and inefficiency) of
living in a large group when compared with small group
(Wrangham, 1980; van Schaik, 1983; Isbell, 1994; Isbell
and Young, 2002). Group fission happens regularly
among baboons (although there is a dearth of published
information: for baboons, see Stoltz, 1972; Nash, 1976;
Hamilton and Bulgar, 1993; Henzi et al., 1997; Wasser et
al., 2004; Van Horn et al., 2007; but for macaques, see
Furuya, 1969; Chepko-Sade, 1974; Chepko-Sade and
Sade, 1979; Dittus, 1988; Kuester and Paul, 1997; Oka-
moto and Matsumura, 2001; Kawamoto, 2010). Theoreti-
cally, a group should split when it gets too large to forage
effectively (increased feeding competition and/or has to
travel too far to feed that many individuals), or too large
to monitor and manage the many social interactions, or
for both reasons.
In 1981, the main study group, Pumphouse, fissioned.
The split centered on a disagreement about foraging
strategies not about group size as Pumphouse was well
below the numbers recorded in other baboon fissions (see
above). An agricultural cooperative bought Kekopey
Ranch, the study site, in 1976. Kekopey covered 45,000
acres. Although it was a cattle ranch, there were few
cows, a large complement of wildlife, and minimal farm-
ing potential. By 1981, small-hold farms (shambas)
appeared in the baboons traditional home range. Then
humanbaboon conflict began (Strum, 2001/1987b, 2010).
Young adult and large adolescent Pumphouse males
were the most motivated to raid. Troop movements away
from the sleeping sites included extensive negotiations
between these males and the rest of the group. Some
days, the raiders pulled the rest of the troop with them
but on other days they failed. Then only the raiders
went to the fields. There was a dramatic increase in
anthropogenic injuries and deaths because of the conflict
(Strum, 2010). Pumphouse spent over 6 months in the
process of splitting. During this time, there were con-
stant negotiations between raiders and nonraiders as
well as among family members of raiders and nonraiders
as they decided which group they would join. In the end,
Pumphouse, the larger descendent group remained pri-
marily natural foragers, whereas the smaller descendent
group (Wabaya) became dedicated raiders.
However, these werent the only responses. A third
study group, Eburru Cliffs (larger than the other two
and having the largest home range), tried human food,
sustained the costs of conflict, and then completely aban-
doned the settlement area. They remained natural forag-
ers using the part of their traditional home range that
was as far away from the settled area as they could get
(a distance of over 7 km as the crow flies).
Why raid? Human food offers significant benefits to
baboons. Both males and females grew faster, reached
adulthood earlier, and had heavier final weights (Strum,
1991, 2010), a result found in other studies of food
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DARWINS MONKEY
enhanced baboons (Altmann and Muruthi, 1988; Muru-
thi et al., 1991; Altmann et al., 1993). Faster growth
meant earlier female reproduction. Although the sample
size is small, raider females reached menarche sooner
and produced their first baby earlier than nonraider
females. Interbirth intervals were significantly shorter
in raider females when compared with nonraiders. An
extreme example of this was the lowest ranking raider
female. Her fecundity increased significantly when she
became a raider, which reversed her previous age-related
reproductive decline. Infants who grew up in food-
enhanced conditions reached menarche and first birth
earlier even though these landmarks occurred after they
were translocated to a resource-poor environment. Infant
survivorship did not differ between the raiders and non-
raiders despite initially high rates of injury and death
associated with raiding. With time, raiders adjusted to
the risks of being close to humans so that costs declined
(Strum, 2010).
What were the baboons doing during those 6 months of
the fission? The matriarch of the lowest ranking family
provides one illustration. One day she would be with the
raiders; the next day she was with the nonraiders. This
went on for months. Her already independent infant,
who still closely associated with his mother, moved back
and forth with her unlike the females adolescent sons
who remained with the nonraiders. Eventually, after
many migrations back and forth, that female became a
permanent member of the raiding troop but her infant
did not. He returned to the nonraiders and stayed with
his brothers. Months later, the matriarchs subadult
daughter also became a member of the raider group.
Females could have several reasons for joining the
raiding group, Wabaya. Most often, the emigrating
female was following her closest male friend who hap-
pened to be a raider. A females rank in her matriline in conflict with people), a total of 132 individuals. This
also played a role. The females who left Pumphouse for
the raider group were most often the oldest daughters
and hence the lowest ranking females in their matriline.
By joining the raider group, they traded kin support for
the benefits of male friendship and of raiding. They also
avoided internal matrilineal disputes and reduced the
number of females to compete with because Wabaya was
smaller. Three pairs of sisters transferred to the raiding
troop. The social framework that underpinned their pre-
vious rank relationship to each other disappeared and
their rank reversed in Wabaya. The older sister became
dominant to the younger one. By contrast, some things
didnt change. Despite having only a few matrilines rep-
resented in Wabaya, the original Pumphouse female hi-
erarchy was perfectly reproduced albeit with some
females missing.
Complexity 2: Context becomes history as baboons
negotiate socioecology
The study troop did not split because it was too large.
What had happened during the fission added evidence
about the socioecological complexity of baboon lives.
Most individuals selected not to raid despite the major
documented benefits of the raiding lifestyle. Individuals
with the same characteristics chose different paths. To
my surprise, other factors trumped evolutionary princi-
ples like optimal foraging and kinship as well as derived
principles like hierarchy.
Why didnt all males become raiders? Large subadult
and young adult males had the most to gain in terms of
growth and weight (and reproduction?) yet not all of
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them made the switch. Not all females became raiders. I
expect that the conservativeness of baboon females
played an important role and for a few females a special
social relationship (friendship) with a raider male was
the defining influence. However, even in this smaller
subset, there were exceptions: the lowest ranking female
and her daughter were not friends with any raider male
when they joined the raiding group. Equally important,
not all troops with the opportunity chose to raid.
Paying attention to the process, to the natural history
of events and individuals, showed me baboon complexity
beyond social strategies and added to my appreciation of
how evolution works at the level of real lives.
Translocation: Survival depends on others not on
competition between individuals (19842000)
Darwin relied on the idea of survival of the fittest to
explain evolution by natural selection (Darwin, 1859).
Subsequent evolutionary interpretations of animal (pri-
mate) behavior focused on competition as the key to indi-
vidual survival. Sociobiology (Wilson, 1975) enlarged the
individual to include kin in the mid-1970s [inclusive fit-
ness (Hamilton, 1964)], and recently, group selection has
been resuscitated as possible and important for humans
(Richerson and Boyd, 2005; Leigh, 2010; Nowak et al.,
2010; West et al., 2010). Usually, scientists infer what
behavior is crucial to survival and reproductive success.
By contrast, the translocation of the study baboons
became an actual test of survival where the process could
be documented and the outcome could be measured.
Despite great success with crop-raiding research and
control techniques, in 1984, I had to translocate three
troops of wild baboons (Pumphouse, Wabayathe
raiders, and a third troop called Cripple who were also
was the first scientific translocation of a primate group
(Strum and Southwick, 1987). The translocation pro-
vided a unique opportunity to document adaptability
(possibly adaptation) in action. I could compare the same
individuals and the same troops in two radically differ-
ent environments. The translocation site, on the Eastern
Laikipia Plateau, is arid savanna with half the annual
rainfall of Kekopey, the baboons previous home. Serious
droughts are common. The baboons were familiar with
only about half of the foods available in the new location
and the unknown ones turned out to be critical to sur-
vival during seasonal bottlenecks. The release site was
not ideal; however, it was the only site I could get.
The descendants of the translocated baboons are alive
today. The process by which they survived provided
more evidence of the complexity of baboon life and illus-
trated evolutionary raw material.
Primates are social; it is a key characteristic of the
Order. There are a number of theories about why most
primates live in a social group (see above); however, it is
difficult to document the value of group living during the
course of most studies. The baboon translocation pro-
vided one chance. The benefits of living in an intact
social group permeated almost every aspect of adjust-
ment during the translocation.
To start, I manipulated existing social networks to
ensure that the troops stayed where they were released.
I held adult males captive at the release site, whereas
the females and youngsters were set free. As expected,
the females did not go far without the males. By the
time the males were released, the females had already
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8 S.C. STRUM
settled in and female conservatism checked the male
tendency to wander (Strum, 2002, 2005).
Each of the three troops was released in a different
place and at different times. The troops maintained
cohesion and coordination from the first days of freedom.
The group represented a social learning resource in a
more basic sense than we usually assume. Individuals in
the translocated groups demonstrated flexibility in learn-
ing in novel settings. Social cues figured prominently in
the process of acquiring information. For example, the
translocated troops had to construct a diet in this unfa-
miliar place. Social facilitation played a role as was
documented in another context before translocation
(Strum, 1983a). The incorporation of new foods into the
diet was strongly related to specific social events. For
example, adults in translocated troops watched and fol-
lowed indigenous baboon troops. The strangers built
their home range by tracking the path of local troops
and foraging on some of same foods. In addition, when
local males migrated into the translocated troops, they
brought indigenous information. The translocated
baboons behavior tracked the immigrant males behav-
ior. In this way, they learned to find water in dry sand
rivers and began using several novel large succulents
(Sansevieria intermedia, Sansevieria abyssinica) for both
moisture and food.
Another value of the social group could only be called
emotional support (Hannah and McGrew, 1991). This
was evident during the accidental fission of one group at
release and their subsequent reunion. Loud greetings,
extensive embracing, and other amplifications of typical
baboon reunion signals punctuated this meeting (Strum,
2001/1987b). It was the exaggerations of these normal
behaviors that pointed to the stress of separation and
the relief of reunion.
A comparison of the baboon translocation with subse-
quent primate reintroductions and translocations add
force to my conclusions. Survival skills are lost in captiv-
ity. Artificially created groups fall apart (Kawai, 1960;
de Vries, 1991; Vie and Richard-Hansen, 1997). Under
normal circumstances (and in extraordinary conditions),
the social group provides the context for appropriate
learning and as a resource for that learning (King, 1994;
Russon, 1999; Crast et al., 2010; Leca et al., 2010). Rein-
troduced captive primate groups fail to meet the learn-
ing challenges of their wild habitat even after extensive
pre-release training (Kleiman and Rylands, 2002).
The translocation gave other insights about adaptabil-
ity, chance events, and the importance of the social. A
night-time encounter with elephants was an accident
that changed the future of one of the translocated troops
(Strum, 2001/1987b). Surrounded by elephants at their
sleeping site, Pumphouse moved 5 km during the night,
an unprecedented act for baboons and other diurnal pri-
mates. I know this because they marched past the lone
human outpost in the area to reach another set of sleep-
ing rocks at the edge of their home range. Subsequently,
Pumphouse abandoned more than half of their range
including the best rainy season portion. Following that
move, group size declined as males and even mothers
with infants left to join other, indigenous, baboon troops.
Just when it seemed that Pumphouse might simply be
absorbed into the local population, the exodus stopped.
New males joined and the next stage in this troops his-
tory began.
There were subsequent home range shifts but none as
precipitous as the first. By 2000, Pumphouse ranged more
than 20 km from their release site. By contrast, Wabaya,
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the previous raiders (now called Malaika trading the bad
guys for the the angels) who were translocated to the
same location stayed put from 1984 until 2007.
All but the first home-range shift tracked a resource
gradient; however, information about and the attraction
of food resources werent enough to force a move. The
shifts depended on social processes. Baboons are not ter-
ritorial; adjacent groups have overlapping home ranges
with undefended boundaries, although intensively used
core areas are sometimes defended. Normally, the con-
servativeness of females prevents a baboon group from
crossing home-range boundaries. Each subsequent
Pumphouse range shift followed an influx of immigrant
males [see Rowells suggestions about sex-biased knowl-
edge (Rowell, 1972)]. These immigrants had firsthand
experience of the resource gradient because they came
from there. However, even with that knowledge, one
male was ineffective. It took a cohort of four or more
males, all indicating in the same direction during troop
movement, to succeed. There was a time lag because the
males had to first create friendships with females in
order to influence troop direction. Clearly, knowledge
was vital, but action relied on social processes.
Complexity 3: The social group is an organ of adap-
tation; accidents of history create different socio-
ecological paths
The translocation was a real test not an imaginary
one. If ever there was a situation where survival of the
fittest should operate, where competition should have
had an upper hand, this was it. Instead other baboons,
either from within the translocated group or the indige-
nous population facilitated both learning and survival.
The translocation, even more than the crop raiding,
pointed out how evolutionary principles get embedded in
a specific time and place and why context matters. Each
groups natural history illustrated a variety of different
paths. Chance (like an encounter with elephants), con-
tingency (how many immigrant males from which area),
and evolutionary principles interact to generate baboon
options and baboon responses. The process was infinitely
more complex than explanations collapsed into evolution-
ary time might suggest.
Fusion of two groups: Baboons negotiate the
nature of their social group (20002002)
Fusion, when two groups merge to become one, is rare
in baboons, so rare that it has been observed only three
times, once in baboons in Amboseli National Park in
1960s (Altmann and Altmann, 1970; Altmann et al.,
1985), once among baboons in Mikumi National Park,
Tanzania (Wasser et al., 2004), and once at De Hoop in
South Africa (Henzi et al., 2000). There are only a few
other cases reported for cercopithecine monkeys despite
the hundreds of thousands of hours that cercopithecines
have been studied in the wild. These include two vervet
fusions (Hauser et al., 1986; Isbell et al., 2002; Isbell,
2009), one toque macaque fusion (Dittus, 1987), and sev-
eral cases of fusion in Japanese macaques (Maruhashi,
1992; Takahata et al., 1994; Sugiura et al., 2002).
Descriptions suggest that the fusion happened almost
instantaneously (or over a short time for toque maca-
ques). Two separate monkey groups come together in the
evening at a shared sleeping site and leave the next day
as an integrated merged group.
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DARWINS MONKEY
This is quite different from fission/fusion societies
among primates where coming together and splitting
into smaller, usually foraging groups, happens on a
daily or weekly basis (Aureli et al., 2008). Chimpanzees
(Lehmann and Boesch, 2004) and spider monkeys
(Chapman et al., 1995) are well-known examples.
I have documented four baboon fusions in just over a
decade. The first fusion was 16 years after translocation
and involved one of the translocated troops, Malaika (the
angels, previously the bad guys) and an indigenous
group, Soit Oitashe, that had been studied since 1985
(Barton, 1993). What made this (and subsequent fusions)
so distinct is that the process took more than 2 years.
During that time, individuals in each group engaged in
extensive negotiations about merging both within the
group and across the groups.
The impetus for the first fusion involved several eco-
logical events including the creation of a wildlife sanctu-
ary on communally owned land that overlapped the
baboons home range. The sanctuary led to an increase
in prey and predator numbers. Later, the large mammal
prey species left. This made local pastoralists dogs and
then the baboons the prey. Predation rates rose to un-
precedented levels.
Who did what and when during the fusion process
depended on specific events and on daily negotiations
involving particular individuals and age-sex classes. The
translocated troop, Malaika, was the smaller of the two
troops. Malaika shadowed the indigenous group, Soit
Oitashe, initially, but roles switched as Soit Oitashe, the
indigenous group, lost members to heavy predation.
The sum of the forces of attraction and dispersal deter-
mined how mixed the two groups might be at any one
time. For example, adult and subadult females were very
attracted to new babies in the other troop. Sexually recep-
tive females also approached the other troops males. Sev-
eral females from Malaika were drawn to a male friend
who had recently emigrated back to his natal troop, Soit
Oitashe. Large subadult and adult males, in both troops,
herded their females away from the vicinity of the other
troops males trying to keep the two groups apart. This
was only a small part of the daily push and pull of indi-
viduals, relationships, and classes of animals.
Leaving the sleeping rocks in the morning no longer
followed a standard procedure because each group had
its own indicator animals and there was no certainty
about who was going to follow whom. Foraging during
the day was also problematic because Malaikas smaller
home range was entirely contained within Soit Oitashes
larger home range. Initially, Malaika refused to cross its
own boundaries when Soit Oitashe moved on. Instead,
Malaika waited, intercepting Soit Oitashe on its way
back to the sleeping site. As the fusion progressed,
Malaika did cross this invisible line, albeit hesitantly,
thereby benefiting from access to new resources. Eventu-
ally, the home range of the fused troop (now called Nabo,
Maasai for coming together) represented a compromise
between the two previous home ranges.
Feeding competition also played a role during the
fusion process. For example, acacia flowers are a pre-
ferred and high-quality food (Barton, 1993). Flowers
emerge during the dry season. Their distribution is lim-
ited both at the start and at the end of blooming. During
these times, Soit Oitashe displaced Malaika from the few
trees that had flowers. In response, Malaika would sneak
away to feed undisturbed and out of sight. However,
Malaika paid a high price for this maneuver because
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9
when Soit Oitashe noticed, they searched for them and
aggressively mobbed Malaika when they found them.
For two groups to merge, the two female hierarchies
needed to be integrated. Initially, Malaikas high-ranking
females kept pushing all of Soit Oitashes females. With
the support of their male friends, Malaika females man-
aged to be in the middle of the integrated dominance hier-
archy and forcing their way upward. However, Malaika
males stopped supporting their females. Without male sup-
port, Malaika females dropped to the bottom of the inte-
grated female hierarchy while maintaining their original
rankings among themselves.
Specific individuals had disproportionate impact on
the degree of mixing between the two groups. This
included two Malaika females and specific males from
both groups. Low-ranking Soit Oitashe females also
became the most active and gave the most aggressive re-
sistance to any attempt by Malaika females to push up
the integrated female hierarchy. An illustration of the
influence of specific baboons was the effort that one Soit
Oitashe male exerted to keep his troops females from
interacting with Malaika. The day he emigrated, there
was a noticeable increase in intertroop interactions and
in spatial proximity between members of the two groups.
Given the length and extent of the process, I had to
develop criteria to use to decide when the two troops had
become one group. The criteria reflected what I felt were
aspects of normal group sociality. These included the ab-
sence of previous troop bias in proximity, grooming and
other social interactions, and troop movement negotia-
tions with indicators and followers responding across
previous troop lines. Foraging subgroups should be
mixed, and the fused group should react as an inte-
grated troop in encounters with other groups, during
predations by baboons, toward new male immigrants, to
separations and reunions, and to social disruptions. Of
course, the fused group had to have an integrated female
dominance hierarchy and an integrated male hierarchy.
It therefore should not be a surprise that integrating
two social systems took such a long time.
Why fuse? Why fuse now? The answers to those ques-
tions offer a test of major socioecological theories of
socialness (see above and Dittus, 1986). Some of the
fusions reported in the literature took place because one
or both groups were too small to be viable in terms of
social and sexual interactions and predator protection.
Protection against predation likely played a role in the
first baboon fusion I documented, although the sizes of
both troops before merging were within the normal
range for olive baboons and had all agesex classes rep-
resented. Yet before the fusion, both lost many group
members to predators. By contrast, after the fusion, the
merged group lost none despite the continued presence
of predators in the area.
The details of negotiations suggest, however, that the
baboon reality was much more complex than simply
defense against predation. Malaika traded off feeding
competition and protection from danger on a daily basis
until the fusion was complete. Once fused, it bore the
cost of feeding competition but benefited from enhanced
predator protection. Soit Oitashe benefited from better
protection while remaining dominant in feeding competi-
tion. The costs and benefits of fusion, therefore, were dif-
ferent for Malaika and Soit Oitashe. Costs and benefits
also diverged for different agesex classes and even for
individual baboons. Moreover, the benefit to cost ratio
changed based on season, location, and day. This push/
pull of fusion was even more elaborate than the process
of fission I had documented during the crop-raiding era.
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Complexity 4: Baboons negotiate almost everything,
even the nature of their social group
Baboon negotiations during the fusion were about the
nature of the social group to a degree not yet docu-
mented for a nonhuman primate species. The fact that
the process took over 2 years and involved dyadic, tria-
dic, and polyadic interactions in the context of every ba-
sic aspect of baboon life suggests that the term nego-
tiation, in its human sense, is appropriate for baboons
as well.
The negotiations were complex. The socioecology of the
fusion was complex. The interaction of levels was com-
plex. However, there was a ladder of complexity
[building on Hindes model of levels (Hinde, 1976, 1987)]
from dyadic interactions, cascading upward to relation-
ships and subgroupings, and clusters of subgroupings to
troops each with their own histories and negotiating
interests relative to each other and to higher and lower
levels. This was embedded in environmental complexity,
which operated at different spatial and temporal scales.
As ecological factors are often not correlated (e.g., the
effects of rainfall, food availability, predation, disease,
and human impacts are often independent of each
other), one simple principle cannot govern what the
appropriate response will be. One translocated troop,
Pumphouse, reacted as if the elephant risk was greater
than the loss of food resources. Another translocated
troop, Malaika, accepted the cost of feeding competition
in exchange for increased safety. This does not make
selection random or arbitrary. Rather, it points out an
important constraint and obstacle: selection must oper-
ate on a range of point responses to challenges that are
sometimes themselves complex and independent, sug-
gesting that evolutionary explanations need to incorpo-
rate, rather than eliminate, complexity, chance, and con-
tingency. At many times in a study troops history, it was
possible to identify probable evolutionary causes of ba-
boon responses; however, these did not operate alone.
For example, although it is tempting to conclude that
predation pressure explained the first fusion, another in-
digenous troop in the same area, Musul, lost members to
predators and yet didnt try to fuse.
My argument so far can be summarized as follows.
Looking at the scale of baboon daily life, focusing on pro-
cess rather than only on outcome, unearths layers of com-
plexity and illustrates how complexity is generated. Out-
comes are neither random nor deterministic in a way of-
ten assumed when scaling up to make evolutionary
arguments. Evolutionary time speculations of cause and
effect assume rather than clarify how evolutionary princi-
ples are situated by chance, contingency, and history. This
is because they either lack the information or ignore the
importance of context and black-box the relationships tools and with them baboons can build only soft soci-
between real behaviors and evolutionary outcomes.
My approach is not just a focus on proximate rather
than ultimate explanations of behavior. The baboon data
challenge some of the fundamental assumptions about
how behavior evolves. Tinbergen (1963) highlighted four
questions that must be addressed in any comprehensive
explanation of a behavior. His framework identified both
the how proximate level (mechanism and ontogeny) bear in performing society. Ironically, according to this
and the why ultimate level (function and phylogenetic framework, material resources, language, and symbols
history). Yet, the ethologists, like Darwin before them,
had to assume the feedback between proximate and ulti-
mate levels while studying static behaviors. Equally latticework, which looks ostensive because so much can
important, ethologists had to examine simplified contexts
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gaining limited insights about how behaviors fit together
into a complex whole or how that complex whole can
change during the lifetime of the individual. The baboon
data offer a dynamic and integrated view of real-life be-
havioral complexity. From this vantage point, the inevi-
tability of an individuals choices and therefore the inevi-
tability of the link between proximate and ultimate cau-
sation seems less compelling.
PART 2: MANAGING COMPLEXITY
Managing complexity: Redefining the meanings
of social
In 1987, Bruno Latour and I proposed a different
approach to compare human and nonhuman primate so-
ciety (Strum and Latour, 1987). We were struck by the
unlikely similarities between his subjects, scientists in
laboratories (see Latour and Woolgar, 1986/1979), and
mine, baboons in the wild.
Baboon negotiation, even at the minor level I had docu-
mented back then, raised a profound question: does soci-
ety exist or is it created? At the time, we contrasted two
major views about society. What holds society together?
Traditional approaches to this question assume that soci-
ety exists and actors enter society adhering to rules and
a structure that are already determined (Strum and
Latour, 1987, p. 785). This is the definition of an
ostensive society. For most descriptions of society, it cer-
tainly seems that way. By contrast, in a performative so-
ciety (Goffman, 1971/1959; Garfinkel, 2003/1967), individ-
uals actively create and define society for themselves and
for others. Therefore society, rather than being a preex-
isting structure, is actually constructed through the
many efforts to define it; it is something achieved in
practice by all actors (Strum and Latour, 1987, p. 785).
We argued for a shift to examine how actors create (per-
form) their society. In hindsight, we were changing focus
from outcome to process motivated by our data on nego-
tiation in baboon and human social groups. Many fields
have since embraced this shift (see below).
Although we argued that the process might be similar
for baboons and humans, it was clear that the outcomes
are different. Transforming baboons into active perform-
ers of their society doesnt put them on a par with
humans. We proposed that the difference is to be found
in the practical means actors have to enforce their ver-
sion of society or to organize others on a larger scale.
Baboons have only themselves, their bodies (including
brains and minds), social skills, and whatever social
strategies and relationships they can construct as
resources. Some factors are embodied and change slowly;
age, sex, perhaps kinship and dominance need not be
continually reconsidered. However, social skills are soft
eties. Other factors and contingencies must be continu-
ally tracked and performed. Baboons become the arche-
typical competent member of the ethnomethodologists
(Garfinkel, 2003/1967) constantly subject to the interfer-
ence of others who are trying to do the same thing. They
live in a complex society.
Humans, by contrast, bring extrasomatic resources to
simplify the task of creating society and of strengthening
the social bond. Human society becomes a complicated
be held constant over space and time. These material and
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DARWINS MONKEY 11
symbolic assets create a material and symbolic structure
that human actors now enter (the traditional view of soci-
ety). As a result, humans accomplish two new aspects of
the social link. They can simplify social negotiations by
using extrasomatic resources. This makes it possible to
build a larger more complicated social structure from the
complex baboon society. (Our terminology may be confus-
ing in light of the developments of complexity science in
the last three decades and the interchangeable use of the
terms complicated and complex in everyday language.
However, in the 1980s, we felt the contrast embedded in
our use of the two terms was appropriate and I continue
to draw on that distinction. Baboons live in a complex so-
ciety where many things impinge simultaneously, needing
attention constantly. Humans live in a complicated
society which like a snowflake is built of simple units
expanded out to make a large complicated but not com-
plex entity.) Baboons are limited in the ways they can
manage complexity because they cant simplify much of
their negotiation. This in turn limits the size and scale of
their society when compared with humans.
An amusing illustration of this contrast between
baboons and humans can be found in hats (Fig. 1). If
baboons wore hats, they could simplify their social nego-
tiations saving a lot of time and effort. These material
symbols contain information about roles and expecta-
tions of behavior. If only baboons had hats, they could
begin to shift from complex society where almost every-
thing impinges simultaneously and needs to be continu-
ally negotiated toward a complicated society where nego-
tiations are simplified, more focused, and produce more
stable outcomes that can travel over space and time.
In summary, redefining the social link suggests that
baboons and humans engage in the same process of cre-
ating society but use different resources. This produces
divergent societies. Baboon society is complex because
the ways to simplify, hold constant, black box are lim-
ited. Baboon complexity often verges on the edge of
chaos as so many factors impinge simultaneously on Fig. 1. What if baboons wore hats? (Credit: Dr. Shirley C.
negotiations. The baboon evidence for complexity and Strum C ) [Color figure can be viewed in the online issue, which
negotiation that Ive gathered since we wrote the article is available at wileyonlinelibrary.com.]
bolsters the argument. The volume and scale of negotia-
tions that exist each day create the need for structure
deal with complexity and how much complexity they can
that goes beyond kinship. Structure creates relative sta-
handle? Dunbar (1996) suggested that the transaction
bility and predictability, reducing what otherwise would
be intolerable transaction costs of living in a large and costs of a typical primate grooming encounter limit the
heterogeneous social group. number of grooming partners an individual can have
Recently, Shultz et al. (2011) modeled the evolution of hence humans switch to gossip in the service of a much
stable sociality in primates. Their conclusion that there larger social network. Economics and political science
was a step-wise evolution from solitary to multi-male/ also see human interactions in terms of transaction costs
multi-female groups, which only later reverted to one- (North, 1987; Coase, 1998; Beccevra and Gupta, 1999;
male and pair-living family groups, contracts previous Putnam, 2000; Rao, 2002). I suggest that baboons, too,
speculations but fits our framework very well. One solu- have transaction costs similar to those identified for
tion to the complexity problem is to revert to smaller humans: finding the information needed for a particular
and less heterogeneous groupings. In fact, recent events interaction, the bargaining costs of reaching an agree-
in one baboon troop illustrate just this point. This troop ment with another individual, and the policing and
appears to be simplifying social complexity through per- enforcement costs involved in monitoring and reassess-
manent one-male subgroupings that are not ecological ing the relationship and the interaction. These seem ru-
foraging units found in fissionfusion societies or in mod- dimentary to any complex primate social relationship
ular multilevel societies like hamadrayas baboons (Swe- and exchange.
dell and Plummer, in press). Historically, those studying primates generally
assumed that social structure and social grouping exist.
Managing complexity: Remodeling the female For example, DeVore/Washburn/Hall proposed the male
hierarchy so that rank simplifies complexity but dominance hierarchy as the organizing principle of the
is not linked to reproductive success (20012010) baboon group (Washburn and DeVore, 1961; Hall and
DeVore, 1965; Strum and Fedigan, 2000b). At the same
The evidence just presented (and more that I dont time, Chance (1967) suggested attention as the mecha-
have space to review) raises the question of how baboons nism. The discovery of a female matrilineal hierarchy in

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rhesus macaques on Cayo Santiago (Sade, 1967) pro-
vided an alternative structure for a monkey group
where males leave and females stay. Sociobiology later
injected kinship and the gene calculus of inclusive fit-
ness, sexual selection, and parenting strategies to create
socioecological determinants of group configurations
(Williams, 1966; Trivers, 1972; Wilson, 1975). These
strengthened the idea that female matrilines and the
female dominance hierarchy were evolutionary adapta-
tions and the primary structure of a baboon group.
Arguments about hierarchy, female or otherwise, have
assumed that rank will be strongly correlated to repro-
ductive success (Hausfater, 1975; Altmann et al., 1988;
Cheney et al., 2006). Yet, data since the 1970s show an
inconsistent and often poor fit between baboon female
rank and parameters of female evolutionary success.
Instead, other factors appear influential like female
sociality (Silk et al., 2003, 2010b), female age-specific fe-
cundity (Strum and Western, 1982), environment
(Wasser et al., 2004), or a variety of factors that seem to
cancel each other out (Brown and Silk, 2002; Wasser et
al., 2004; Silk et al., 2005; Crockford et al., 2008; Silk
and Strum, 2010).
The imperfect fit between female rank and reproduc-
tive success is not surprising if attention is on what
actually happens. Then evolutionary principles operate
in a context also shaped by complexity, chance, contin-
gency, and history. The surprise is, instead, that there is
a stable female hierarchy at all despite limited evolution-
ary benefits.
I suggest that the female dominance hierarchy has a
different purpose. Its function is primarily to create pre-
dictability (and stability) so that transaction costs for
females and others can be reduced. The complexity of ba-
boon social life needs to be simplified. If the transaction
costs of grooming limit the number of grooming partners
[and stress is lower when a female baboon has fewer
rather than many grooming partners (Crockford et al.,
2008; Wittig et al., 2008)], imagine the social transaction
costs (and stress) involved in any part of baboon daily
routine if an animal didnt know in advance its relation-
ship to others. Individuals, and by analogy the troop,
would be paralyzed by minute-to-minute negotiations
involved in the process of where to feed, rest, or travel,
whom to approach, and whom to avoid. There would be
no time for the basics of survival or enough time and
energy to meet any new challenges.
If I am correct that the female hierarchy is primarily
structure, then there need not be a strong correlation
between female rank and individual reproductive success
and no conundrum about why low ranking females stay
in a group. All individuals benefit from predictability
and stability no matter their rank.
However, why use the female hierarchy instead of
some other structure to generate predictability and sta-
bility? Im not suggesting that the baboons think about
the problem and consciously create the solution. Instead,
it is obvious that the female hierarchy is easily gener-
ated from evolutionary principles such as kin selection
and reproductive value (Vx) (Williams, 1966). In baboons
and other cercopithecines, a mothers intervention on
behalf of her younger offspring in altercations with an
older offspring creates an internal family hierarchy in
which the youngest sibling is the highest ranking. The
normal expansion of a family over time would fashion a
group hierarchy where only those closest in rank are
likely to be related enough for kin selection to operate
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(Seyfarth, 1977). The female hierarchy thus easily fulfills
the need for a structure to reduce transaction costs. As
baboons practice female philopatry, this hierarchy also
organizes the largest cohort of animals in the group. Of
course, there are other evolutionarily generated struc-
tures in a baboon troop including the male dominance
hierarchy, malefemale friendships, and other affiliative
bonds (like those between play group members), but all
of these are more dynamic, more ephemeral, and incor-
porate fewer individuals than does the female hierarchy.
If the female hierarchy is co-opted as primary struc-
ture, then it is the conservative nature of females [based
on reproductive constraints of female mammals (Trivers,
1972, 1974)] that helps to keep this hierarchy relatively
stable and predictable. A male baboon who loses an en-
counter to another male might continue to contest the
outcome throughout the day or even the week. A female
who loses an encounter to another female rarely disputes
the results. This contrast results from sex differences in
risks and benefits.
However, the female hierarchy isnt invariant. Ive
seen what happens when the female hierarchy is unsta-
ble. This evidence supports my argument. The female hi-
erarchy (in the different study troops) has changed a few
times during the last 40 years. These have been mostly
adjustments in rank between mothers and daughters,
disruptions but not of major concern to anyone but the
members of the family. By contrast, the few major
changes that have occurred in the overall female hierar-
chy are instructive. They elicited lots of aggression some
resulting in injuries. The aggression also disrupted basic
daily activities like foraging and travelling. Disorder
quickly spread from a few females through the group
embroiling individuals not initially involved. Group life
came to a standstill for a good part of several days while
the continued instability affected individuals and the
troop for weeks and even months. It is not hard to imag-
ine, from these periods of disruption, why a stable and
predictable structure is crucial not just for females but
for the rest of the troop.
The possibility exists that there may be a better fit
between rank and individual reproductive success under
specific conditions (Cheney et al., 2004, 2006). For exam-
ple, high rankers have preferred access when it is feasi-
ble to monopolize key foods during a resource bottleneck
(Barton, 1993). However, during the decades of my study,
such situations turned out to be few, short, and did not
produce the expected fit between female rank and repro-
duction (Strum and Western, 1982; Silk and Strum,
2010). This was true even during critical periods such as
droughts probably because the foods left then are small,
dispersed, and not defensible.
Given the complexity of baboon reality, individuals
(and groups) need a way to manage daily life. It makes
more sense, in this context, to view the female domi-
nance hierarchy as a transactional principle than as a
genetic characteristic of individuals or as the genetic
consequences of the behavior of individuals. It is the ba-
boon version of what humans do; . . .an economizing
behavior in the sense that it greatly reduces the transac-
tion costs of social interactions and permits efficient col-
lective action (Fukuyama, 2011). In the same way that
manners in human societies make life livable, workable,
and usually prevent descent into anarchy, the female
hierarchy creates a livable, even, civil social life for
baboons. Reproductive success would not be possible
otherwise; however, this is not the same as saying that a
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females rank in the hierarchy is somehow directly
connected to her reproductive success.
The importance of structure is intuitively obvious
when complexity and process are taken seriously. There
are analogous arguments about structure from other
fields. For example, the value of structure in biological
systems is well summarized by Weiss and Buchanan
(2009). For humans, a variety of theories argue for the
benefit of structure to reduce uncertainty, minimize cog-
nitive dissonance, build social relationships, and facili-
tate social exchange (for cybernetics, see Bateson, 1967;
Watzlawick and Bevin, 1967; Harries-Jones, 1995). In
fact, Festinger (1985/1957; Harmon-Jones and Harmon-
Jones, 2007) suggested that humans seek consistency
and try to reduce dissonance in any new situation.
Recent approaches to human cognition including distrib-
uted cognition (Hutchins, 1995; Forster and Rodriguez,
2006; Barnard, 2010) and situated action (Suchman,
1987; Rogoff, 1990, 2003; Lave and Wegner, 1991) rely
on a nontraditional idea of structure that is similar to
my argument. There may be indirect or downstream
impacts of structure on female baboon reproduction, but
I suggest that the value of structure is not directly
reproductive success.
Managing complexity: Extinction, tolerance,
slippage, and the evolution of the good enough
(20072011)
An underlying assumption of evolutionary arguments
is that there are winners and losers (Weiss and Bu-
chanan, 2009; Watts, 2010). Yet, in the vast span of life
on earth, more than 99% of all species have gone extinct
(Raup, 1986; Wilson, 2002; Barnosky et al., 2011). As ev-
olutionary history is continuous and the end point in ev-
olutionary time is most probably extinction, calculating
costs and benefits has to be sensitive to that history.
Species who flourished at one point later disappeared.
This is relevant to my argument in two ways. First, a
particular point in time may or may not be representa-
tive of what has happened or what will happen. I recog-
nized this about the baboons Im watching and it will
apply equally to other species. The power of evolutionary
arguments will therefore depend on capturing as much
of the (natural) history as possible. Second, the adapta-
tionist paradigm (Gould and Lewontin, 1979; Laland and
Brown, 2011/2002) often assumes that animals in nature
are finely, even perfectly tuned to their environment
and, at least until recently, that what we observe are the
elements of this excellent fit. Yet, focusing on real lives
in real time illustrates how difficult it is to know about
the consequences of particular actions or to decide
whether, for example, a baboon individual or group
(since individuals dont exist outside of groups) is mak-
ing a mistake.
I had the opportunity to consider this issue during a
second fusion event. This time the impetus appeared to
be food, not predators. The little group, Musul, that
hadnt fused 7 years earlier, initiated a fusion for a dif-
ferent reason, to gain access to an important new food
resource or so it seemed. This resource was dominated
at the time by other, larger, baboon groups (who were
not part of my study).
The resource they sought was the fruit of Opuntia
stricta, an exotic invader which had only just appeared
in Musuls home range. Ironically, before this fusion
Musul had been my control as a test of the costs and
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13
benefits of not fusing. Musul targeted Nabo (itself the
result of the previous fusion of Malaika and Soit
Oitashe discussed previously). By this time, Nabo had
operated as a well-integrated group for more than 7
years. Unlike the earlier fusion, Musul didnt shadow
Nabo. Instead, Musul tried to lead Nabo to the area
with the best cactus.
Initially, most of Nabo baboons resisted probably
because the area was outside of Nabos traditional
home range and already hotly contested (baboons can
hear intertroop interactions beyond where they nor-
mally range). As the MusulNabo fusion progressed,
Musul females exerted more and more influence on
Nabo through their growing network of friendships
with Nabo males. Occasionally, the combined group
made it to the cactus area only to be aggressively
expelled by one resident baboon group in particular.
Aggressive mobbing is usually a very effective tactic to
displace another group from a location. In this case,
while the intruders fled from the aggression, they kept
coming back. MusulNabo endured repeated evictions
over more than a year but also developed evasive tac-
tics. For example, the invaders would leave the sleeping
site at first light to avoid being mobbed. This early de-
parture meant giving up resting and socializing time
and even some feeding time clearly a disruption of their
normal daily routine. However, leaving early didnt
always work. Often the resident troop would track
down the invaders and push them until they were
away from the disputed area.
The MusulNabo fusion (the group was now called
Namu) violated many rules of baboon life that I had
documented during the preceding decades. I considered
whether it might represent a baboon mistake. The
quantitative data suggested that it was. No baboon was
actually killed because of the confrontations between
groups, but during the first 18 months of the range shift,
most of Nabos and Musuls babies did not survive the
first year of life. Adult mortality also increased.
Autopsies of adult females showed that previously nonfa-
tal conditions had turned lethal. Nutritional analysis of
the cactus fruits indicated that its value, and thus its
appeal, might be the energy bonus provided by such a
large package of calories [see also crop raiding (Strum,
2010)]. Yet, the troop must have used up that bonus and
more evading attack. Unlike during crop raiding, body
condition didnt improve.
Do baboons make mistakes? Was I observing the pro-
cess of extinction or at least extirpation, one death at a
time? My answer is different now, 4 years later, than it
was at the start of the fusion. Currently the troops have
been fused (becoming Namu) for several years. Namu, as
invaders, eventually created a home range in the cactus
area despite all the aggression. Although Namu is still
subordinate to the main resident troop, they arent
mobbed as often. Namu female reproductive parameters
are steadily improving along with female physical condi-
tion. Age at menarche is earlier, and interbirth intervals
are getting shorter. Namus overall mortality has
declined while survivorship is improving. I had to con-
clude, against my previous assessment, that Namus per-
sistence paid off.
The second fusion provides more evidence about the
pitfalls of making evolutionary interpretations of behav-
ior because these depend on history and time frame.
Flexibility and adaptability are more vital than I had
imagined. It is hard to do the evolutionary calculus
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14 S.C. STRUM
without knowing the specific history even when popular
evolutionary criteria like condition, reproduction, and
survival are available as was evident in the Namu range
shift. Too often, the link between the behavior and its ev-
olutionary consequences passes through a black box
where much of the work and most of the assumptions re-
side [see similar argument for the interpretation of the
link between social complexity and the evolution of cog-
nition (Strum et al., 1997)].
The MusulNabo fusion also clarifies that evolutionary
processes may be more tolerant, have larger acceptabil-
ity spaces (Weiss and Buchanan, 2009), and allow for
more variance than most evolutionary time arguments
assume. What these baboons did for more than 4 years
was not a perfect fit to the layers of complexity in their
life but it seems to have been good enough. Are they change, habitat redistribution, and human encroach-
on the road to success or to extirpation? It will take time
to answer that question and require measures of success
situated in their natural history.
Since complexity generates a variety of options, it is
not surprising that individuals in a baboon troop dis-
agree about what to do. These disagreements must be
resolved because the group is constrained to move and
act as a unit. Resolution requires negotiation. Therefore,
managing the consequences of socioecological complexity
is a serious daily challenge for baboons. This might be
most apparent during the special circumstances I was
able to document like the incursion of agriculture, peri-
ods of heavy predation, or the appearance of a new food.
However, disagreements, behavioral discussions, and that larger more complex structures developed from
happen during negotiations about the more ordinary
parts of baboon life such as during troop movements,
consorts, when a new infant is born, and the like.
Coming to grips with complexity
These are more than just interesting stories. They are
evidence about how evolution actually works. The focus
on process and history reveals the importance of complex-
ity, chance, and contingency in the real lives of baboons.
It also highlights the flexibility and adaptability (in
response to complexity) that pervade baboon daily life.
Other disciplines like paleontology, ecology, genetics,
and cellular biology have recently made similar argu-
ments. The broad integrative approach of Weiss and Bu-
chanan (2009; Weiss et al., 2011) documents the impor-
tance of chance, contingency, and history (and emer-
gence) in the history of life. Their EcoDevoEvo tion as a critical scientific tool. Evolutionary change,
perspective advocates going from evolutionary to devel-
opmental and ecological time scales [see Fig. 2.2 in Weiss
and Buchanan (2009, p. 12) for an instructive diagram],
and illustrates why chance is always built into concep-
tion, inheritance, birth, death, and much that happens
in between (Weiss and Buchanan, 2009, p. 197). Context
and history, for Weiss and Buchanan, are essential to
understanding how, for example, a gene network, cell,
organism, or ecosystem gets from point a to point b.
Viewing evolution at these scales suggests that there are
multiple routes to success with more tolerance and slip-
page and therefore more flexibility than modern Darwin-
ian interpretations often allow.
A recent example in paleontology is the work of Ven-
ditti et al. (2011) on body size in mammals. They con-
tend that in the evolution of mammalian body sizes,
diversification did not speed up and then slow down
when ecological niches were invaded as previously
assumed. Instead, body size diversification took multiple
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routes because ecological niches were a constantly mov-
ing target. They proposed that speciation might itself
be the outcome of unusual single events (emphasized by
the author) and that to understand so-called adaptive
radiations, it is necessary to study the multiple events in
a species life that provide it with the opportunity to
adapt, rather than studying wide and general processes
(Venditti et al., 2011, p. 393).
Lorenzen et al. (2011) offered another example. North
American Pleistocene megafuana extinctions appear
much more complex than previously assumed. Their
analysis demonstrates that changes in the abundance of
megafauna were idiosyncratic because each species
(and even continental populations within species)
appeared to respond differently to the effects of climate
ment (Lorenzen et al., 2011, p. 364). Surprisingly, the
species that went extinct did not have characteristics
that set them apart from those that survived. This
implies a complex and contingent process (with behav-
ioral flexibility in some species) rather than a simple
and deterministic one.
A recent study on the evolution of sociality in primates
provides another parallel. Shultz et al. (2011) modeled
the evolution of primate social grouping patterns using
genetic and social information about 217 primate spe-
cies. The conclusion contradicts previous speculations
about the evolution of primate social groupings, which
assumed that the original groups were small and simple
these groupings. The analysis indicates that the initial
change in social grouping was probably caused by a shift
from nocturnal to diurnal activity that generated large
multi-male and multi-female groups. Smaller social
structures are derived later in a stepwise fashion that
has some phylogenetic inertia. In this view, the evolution
of primate sociality was neither simple nor gradual and
likely involved multifaceted interactions of group size,
social complexity, and cooperative sociality.
The argument for, and the value of, focusing on pro-
cess and description is best illustrated by modern macro-
ecology (Smith et al., 2011). Macroecologists look at com-
plex interactions embedded in space and time by exam-
ining multiple factors and levels including morphology,
physiology, behavior, ecology, phenology, and phylogeny.
Taking their cue from past natural historians and bio-
geographers, macroecology uses natural history descrip-
they argue, is best understood in terms of complexity,
history, chance, and contingency (and emergence)
whether looking at the past or trying to predict the
future.
Even studies of modern human behavior that usually
begin with the assumption of complexity and history are
moving further in that direction. For example, a recent
study of the effectiveness of co-management of global
fisheries identified 19 factors (summed into eight binary
measures and their interactions) that were vital to suc-
cess (Gutierrez et al., 2011). However, history and acci-
dent (not planned design) determined whether a fishery
had any of the set of these critical features. Similarly,
Black et al. (2011) explained the inadequacy of previous
models of human migration that rely only on simple
principles of causation. Instead, they argued that there
are numerous drivers of migration that include social,
economic, demographic, and political factors. Further-
more, each driver interacts with the others as well as
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DARWINS MONKEY
each being influenced by environmental change. The
result is that they can make better predictions about
who migrates, when, and why than previous attempts.
Another recent example for humans is Fukuyamas
work on the origins of human political order (Fukuyama,
2011). He convincingly demonstrated how similar starting
conditions result in radically different outcomes and that
The factors driving the development of any given politi-
cal institution are multiple, complex, and often dependent
on accidental or contingent events. Any causal factors one
adduces for a given development are themselves caused
by prior conditions that extend backward in time in an
endless regression (Fukuyama, 2011, p. 23).
Of course, Complexity Theory as it has developed over
the last three decades draws some of the same conclu-
sions (Waldrop, 1992; Thelen and Smith, 1994; Capra,
1997; Oyama et al., 2001; Ward, 2002; Johnson, 2011/
2007). Complex systems models, including complex
adaptive systems and dynamical systems, have recently
been applied to primates using social network analysis
and analysis of collective behavior (King and Sueur,
2011a; Sueur and Deneubourg, 2011; Sueur et al.,
2011a). Some of this work formally addresses the issues
of nonlinearity and of history, chance, and contingency
in producing specific outcomes. Most work on complex
systems still falls short of addressing the context of
behavior in the real world. For primate social networks,
for example, this should include multiple interacting
contexts that set up the starting conditions and continue
to influence the operation of the system.
I began thinking about complexity in the early 1970s
when I realized that male success rested on more than
the male dominance hierarchy. Age, residency, context,
and social relationships based on collaboration and be-
havioral exchange all played a role. This view of social
complexity was my first step in expanding out from the
powerful simplifying assumptions of the previous ba-
boon models. Crop raiding (19761984) demonstrated
the significance of ecology and how the interaction of
social and ecological added to the complexity of baboon
lives. The disagreement about whether to raid high-
lighted the way that social negotiations influenced
choices about the basics of daily life. The translocation
(1984) was both a provocative experiment and an un-
precedented opportunity to see baboon adaptability in
action. The value of the social group was never as
obvious as after translocation; however, I also learned
that baboons had multiple ways to succeed and that
specific events and accidents of history set different
groups on unique paths.
The stone of baboon complexity that I had slowly
pushed up the hill in the first decades picked up incredi-
ble speed rolling down the other side in the last decade.
The first fusion (19992001) showed the full extent of
social negotiation about the nature of the social group,
something scientists assume to be a given fact. The 2
years of negotiation embedded in specific social and eco-
logical events made the fusion understandable but not
predictable based on simple evolutionary rules. The sec-
ond fusion (20052008) certainly looked like a monumen-
tal evolutionary mistake but turned out not to be. That
fusion demonstrated the constantly changing balance of
costs and benefits and the large list of factors that had
to be considered. It also implied that evolution was more
tolerant and allowed more slippage making simple evolu-
tionary calculations problematic. Finally, the changes in
the female dominance hierarchy (20082011) bolstered
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15
my argument about the challenge of managing complex-
ity and offered an alternative meaning for the female
dominance hierarchy.
As with all complex systems, the whole is greater
than the sum of its parts (Johnson, 2011/2007; Mitchell,
2009) even though we still lack the methodology to cap-
ture behavioral emergence. However, baboon natural
histories help. They are a resource for understanding
how complexity is generated. They show how solutions
to challenges develop. They illustrate the enormity of
the task that baboons face in managing complexity.
This type of evidence also underscores the role that
context, chance, and contingency play in producing spe-
cific outcomes on the evolutionary stage. The baboon
data also raise the possibility that there are multiple
ways for individuals to succeed, not just one optimal
evolutionary path.
PART 3: WHY SHOULD ANTHROPOLOGISTS
TODAY BE INTERESTED IN BABOONS?
Washburn and DeVore (1961, 1963) used baboons spe-
cifically to frame questions about human evolution.
Ironically, the intuitive power of DeVore and Washburns
model did a disservice to their goals as the simplified
version was applied to other primates and even to
humans (Strum et al., 1999; Strum and Fedigan, 2000b).
Nearly four decades later, I once again suggest that
baboons may be relevant in highlighting the challenges
and solutions of early human evolution.
My first decade of baboon observations led me to
declare them almost human (Strum, 2001/1987b).
Indeed, they exhibited remarkable social skills, social
strategies, and even sexual politics (for a current defini-
tion of and reservations about using the word politics,
see Watts, 2010), we normally reserve for humans. How-
ever, the following decades of baboon quantitative data
and detailed natural histories suggested to me why, de-
spite having much in common in the process of building
society, baboons and humans have diverged so much.
This renders the baboon vantage point particularly use-
ful both for understanding what makes us human and
for refining concepts of how evolutionary processes
actually work at the level of behavior.
Summary of baboon principles of the social
1. The social group is a major organ of adaptation. The
group offers social resources useful in meeting new
challenges (raiding, translocation, etc.) and in solv-
ing daily problems (translocation, Opuntia, etc.).
Baboons illustrate how this works not just why it
should work this way.
2. Social negotiation is critical to baboon life but it
quickly reaches the limits of available time and
comprehension. Baboons are also constrained by
their inability to hold nonmaterial artifacts (rela-
tionships, structure, etc.) over time and space.
3. Baboon society is restricted in scale because it has
few means to simplify social negotiations. Therefore,
baboons cannot escape social complexity to build a
complicated social structure.
4. Context and history matter to baboons. This
includes social actions at the individual and group
levels.
5. Collaboration (see below), not just competition, is a
building block of baboon society.
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16 S.C. STRUM

Fig. 2. The iconic male baboon: friendship and trust rather than aggressive might. A: Male grooming his yearling friend (credit:
Dr. Timothy Ransom C ). B: Male resting with yearling friend (credit: Dr. Timothy Ransom C ). C: Male with his infant and juvenile
friends (credit: Dr. Shirley C. Strum C ). D: Male resting with one of his female friends and her juvenile daughter (credit Dr. Shir-
ley C. Strum C ). [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.]

Examples never do alone. If Vygotsky (1978) is right, that there is

Cognition in the world and in the head. Baboons illus-
trate why the social and the ecological cannot be sepa-
rated in real life. The greatest cognitive challenge for
baboons is probably how to navigate both simultaneously
[see navigational intelligence (Strum et al., 1997)].
Because the two domains cannot be separated, baboons
may reach their cognitive limits faster than predicted by
either social or ecological hypotheses about the evolution
of primate cognition (Byrne and Whiten, 1988). By con-
trast, the human ability to simplify social negotiations
and build a complicated larger scale society may reduce
the cognitive demands of moment-to-moment life and
effectively segregate the social from the ecological for
humans compared to baboons.
Yet, the social is also a great baboon cognitive
resource. Several recent frameworks for human cogni-
tion also seem relevant to baboons (Strum et al., 1997;
Forster, 2002). Situated action and distributed cognition
extend the unit of cognition and cognitive analysis
beyond the individual. They suggested that knowledge
and cognitive processes can reside in the world (situated
in time and space and distributed between social actors
and their environment) rather than just inside the head
of an individual. Hutchins (1995) argued that distributed
cognition makes it possible for social actors to solve
problems (and think thoughts) that an individual could
a mind in society, then the social realm of baboons
becomes a resource for problem solving. Individual
baboons, like individual humans, can later appropriate
socially created solutions. This offers a mechanism for
how plans and schemas might get into the mind of an
individual.
Although the social resources available to baboons may
ameliorate some of the cognitive challenges of an inte-
grated socioecology, baboons eventually reach the limits of
their skills and abilities to manage complexity. Going
beyond this requires additions to brain/mind and a social
world that offers more resources for solving problems.
Competition versus collaboration versus cooperation. Re-
cently, Weiss and Buchanan (2009) and Sussman and
Garber (2011) have illustrated a growing shift in evolu-
tionary arguments from an emphasis on competition to
the importance of cooperation in daily life. Even in
baboons, a species where males have acquired the evolu-
tionary anatomy for aggression, competition is just one
factor and often not the most critical. The iconic picture
of baboon society should not be two males fighting;
rather it is a big male baboon grooming his totally
relaxed infant friend (Fig. 2). As Sussman and Garber
suggested, competition is embedded in a larger social
matrix (Sussman and Garber, 2011).
Yet, cooperation is a controversial term when applied
to nonhuman primates. The Oxford English Dictionary

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DARWINS MONKEY
(Dictionary, 1971) defines cooperation as to work to-
gether, a meaning that originated in 1616 (p. 963)
which by 1830 also meant the practice of economic coop-
eration. For humans, it is used interchangeably with the
term collaboration which first appeared in 1871. There
is no scientific consensus about the definition of the term
cooperation as it applies to animals and humans
(Richerson and Boyd, 2005; Kappeler and van Schaik,
2006; van Schaik et al., 2006; Tomasello, 2009; Weiss
and Buchanan, 2009; West et al., 2010). Definitions have
ranged from complex and restrictive economic criteria
with analysis based on game theory (e.g., see Noe et al.,
2001), to more real-life criteria like joint commitment to
a goal with mutual support and the ability to reverse the development of gathering and hunting and its con-
roles (Tomasello and Moll, 2010), and to the broadest
possible meaning of cooperation: components working
together successfully by some criterion, including that of
evolutionary viability (Weiss and Buchanan, 2009, p.
38). One cross-cutting claim, however, is that human
cooperation is distinctly different from what other ani-
mals or even other primates do.
For the purpose of situating baboons, I find it useful to
think in terms of a continuum that starts with the ge-
ometry of the selfish herd. Here, individuals aggregate
and cohere by simple fact of trying to get away from
something, for instance, a predator (Hamilton, 1971). I
suggest that what comes next is called coordination,
which develops as the enhanced value of group living
requires solutions to behaving as a collective in everyday
life [see also the collective action problem (Nunn, 2000;
Noe et al., 2001; Sussman and Garber, 2011)]. Collabo-
ration is still further in the continuum from the selfish
herd. Although cooperation and collaboration are used
interchangeably in both the common and scientific lexi-
cons, I make a distinction based on insights from
baboons. In 1973, one baboon male started to hunt
rather than opportunistically collect gazelle prey. His
actions were purposeful and his intent was evident in
the sequence of his behaviors. Then other males joined,
initially alerted by his return to the troop covered in dry
blood. Through a set of events, the males actually
learned that collaborating in the direction of the chase
paid dividends. Later, they practiced what they learned
(Strum, 1975b). I use the term collaboration in its orig-
inal meaning, to work in conjunction with others, and
not as a synonym for cooperation. Collaborating baboons
do not have a predetermined common goal. Rather they
are a collection of individuals with similar goals; in the
case of predation, each wanted meat and realized the ef-
ficacy of chasing prey one way rather than another.
Finally, in this scheme, cooperation is a special type of
collective action in which participants have a preexisting
common purpose (see also references above). Chimpan-
zee predatory hunts (Boesch and Boesch-Achermann,
2000) appear a rudimentary form of primate cooperation,
particularly by comparison with collaborative baboon
hunting. The continuum ends with human cooperation,
which entails a variety of additional aspects not found in
other species (but see Wilson, 2012), although the specif-
ics depend on whose definition is used.
Although competition is not as central to the baboons,
I have studied, as previously assumed, neither can
baboons cooperate. That they are expert social collabora-
tors is visible in their social strategies of competition
and defense, in troop movements, intertroop encounters,
home-range shifts, changes in diet, as well as in intricate
multi-individual social and sexual encounters. Data from
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17
other long-term baboon research sites also provide evi-
dence of social collaboration even if the investigators do
not label the behaviors that way (e.g., Palombit et al.,
1997; Silk et al., 2006, 2010a; Swedell and Leigh, 2006).
This means that baboons regularly engage in collective
action that goes beyond the coordination of individuals.
However, I propose that baboons have constraints. With-
out additional resources, baboons cant move from collab-
oration to cooperation just as they are limited in the
type of society they can build.
On the other hand, would baboons benefit from being
able to cooperate? The origins of cooperation in humans
is often attributed to a change in lifestyle, particularly
comitant division of labor and necessity for sharing (Lee
and DeVore, 1968; Bird, 1999; Wilson, 2012). Without
anything in their diet that could be shared, except the
infrequent prey carcass (Strum, 1975b), baboons may
not have the necessary or sufficient conditions for the
evolution of cooperation. In the end, although baboons
are great collaborators, they have neither the means nor
the reasons to cooperate.
Culture as an organizing principle. The performative
view of society, augmented by the frameworks of situated
action and distributed cognition, offers a novel perspec-
tive on tradition and culture among baboons. Baboons
dont use or make material tools in the wild but they do
skillfully craft friendships. These social tools are non-
material artifacts (Strum and Forster, 2001) because
making them requires the same cognitive achievement
as making material artifacts: their shape and use isnt
intuitively obvious on the surface of the material. How-
ever, here the similarity stops. Social tools, unlike mate-
rial tools, have difficulty expanding across space and
time. They are soft tools and difficult to stabilize. Still,
since tool use is often a criterion for culture (see Fra-
gaszy and Perry, 2003), baboons do have some of the ba-
sic skills (Strum and Latour, 1987).
Another criterion used to identify a primate cultural
tradition is that the behavior should be invariant
within a group but variant across groups (Tomasello,
1999; Rendell and Whitehead, 2001; Fragaszy and Perry,
2003; Richerson and Boyd, 2005). A number of baboon
behaviors or behavioral systems may qualify. For exam-
ple, a hunting tradition developed in only one group
(Strum, 1975b, 1976). The raiding lifestyle, the selection
of sleeping sites after translocation, differences in diets
between adjacent groups in the same location, and even
variations in aspects of the friendship configuration
might be more controversial suggestions as baboon tradi-
tions. I could go further and claim that every groups
female dominance hierarchy is a traditional system. Af-
ter all, each is unique to a group, carries forward across
generations, and is relatively stable over time.
What sets baboons apart from humans, if we grant
that these are social traditions, is the next step in the
cultural process: the persistence and accumulation of
innovations (Tomasello, 1999; Richerson and Boyd,
2005). I have argued that for baboons, the means of ac-
quisition and transmission of a tradition depends on
frequent exposure and visible actions. Thus, the hunt-
ing tradition disappeared when the innovator trans-
ferred out of the troop a year later. Why? Although the
rate of baboon predatory behavior in Pumphouse was
high when compared with other baboon and chimpan-
zee groups observed at that time, predation was still a
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18 S.C. STRUM

rare event in daily life. Furthermore, hunting happened
away from the troop so that most of the animals didnt
see or participate in the actual behavior. As a result,
hunting just didnt stick. Now contrast raiding. The It is improbable that the earliest humans were less so-
opportunities to raid were frequent and after the fis-
sion, the entire raider troop, not just a few animals,
raided. This frequency and visibility, I propose, were
important contributors to the persistence of raiding
behavior. Raiding even accumulated some modifications
as raiders learned to avoid the new anthropogenic risks
and human-induced mortality declined over time
(Strum, 2010).
Conversely, if culture is based, in part, on internalized
rules for behavior (DAndrade, 1981, 1990; Byrne and
Russon, 1998), the same factors that may be important
facilitators of traditions for baboons (visibility and
involvement) could mitigate against the need for tradi-
tions. Baboons have constant access to each other as
social information resources (Strum, 1983a, 2010; King,
1994; King and Cowlishaw, 2007; King and Sueur,
2011b; King et al., in press). They can rely on distributed
cognition and situated action rather than on individually
internalized operating principles (Forster and Strum,
1994; Strum et al., 1997; Forster, 2002; Forster and
Rodriguez, 2006). Recently, Roediger and McDermott
(2011) made a similar suggestion for humans. They
argued that If one individual in a group forgets critical
information (about food resources or dangers, for
instance), then it is wise to get an updated memory from
another group member (Roediger and McDermott, 2011,
p. 48). This perspective could help explain why the two
most cultural primates so far are chimpanzees (Whiten
et al., 1999, 2007) and orangutans (van Schaik et al.,
2003; Jaeggi et al., 2010). If a baboon forgets how to do
something, it can just look around. Chimpanzees and
orangutans dont have that luxury. Chimpanzee fission
fusion society (Chapman et al., 1995; Lehmann and
Boesch, 2004) and the nearly solitary existence of orang-
utans (van Schaik, 1999) deprive them of each other as
social resources. This should increase the value of tradi-
tions and possibly of culture. By analogy, if early
humans lived in fissionfusion societies (Grove et al., in
press), they would face challenges of remembering
more like chimpanzees than like baboons.
In summary, I am proposing that baboons have a soci-
ety at the limits of social complexity and that they lack
the resources needed either to manage more complexity
or to simplify negotiations and build a complicated large-
scale society. Baboons have sophisticated cognition that
tracks social, ecological, and socioecological interactions.
It is likely that they appropriate solutions to new prob-
lems from the social context. If so, baboons may need
less in the head because they have so much in the genetic, developmental, and ecological levels (Weiss and
world. The performative process already creates some
social traditions (culture?); however, baboons have diffi-
culty with the persistence and accumulation of innova-
tions because, I suggest, for baboons this depends on fre-
quent exposure and actions that are visible to everyone.
Although baboons are almost human in their negotia-
tion and politics, baboons cant become human because
they lack the ability to build a complicated society from
a complex one. To do this, they would need the behaviors
that are usually classified as hallmarks of humanness,
minimally the new resources that material culture, sym-
bols, language-related abilities provide, and a new sub-
sistence base that both permits and requires division of
labor and sharing.
Finally, I conclude that the four decades of baboon
research and the framework outlined above is helpful in
resetting the starting point for the human experiment.
phisticated or embedded in less complexity than these
baboons. Moreover, lowly baboons also serve as a re-
minder that the group is a primary organ of adaptation
for most primates, that collaboration/cooperation is likely
as crucial as competition in daily life, and that social
intelligence, social skills, and social management were
at a premium long before humans or even the great
apes.
Summary of baboon principles of evolution
1. Baboons demonstrate the value of examining process
rather than just outcome and the pitfalls of uncriti-
cally collapsing behavior into evolutionary time. This
is a critical methodological move. How evolutionary
principles work in real time and real lives can only be
identified this way.
2. History, chance, and contingency are crucial for
understanding baboons embedded in social, ecological,
and socioecological interactions. The extensive baboon
negotiations imply that there are multiple options not
just one best evolutionary solution. In fact, the
baboons add support to the assertion that behavioral
adaptability is a ubiquitous trait of life but that it is
often missed because of the existing scientific empha-
sis on evolutionary time and on evolutionary out-
comes (Weiss and Buchanan, 2009, p. 231).
3. Thinking in terms of contingency and (natural) history
helps to make sense of complexity. It is also easier to
understand complexity by looking at its unfolding, how
it is generated, than by puzzling about existing com-
plex structures (Weiss and Buchanan, 2009, p. 176).
Given these three principles, the baboon data that I
have just presented dont fit the standard view of Dar-
winian natural selection (Laland and Brown, 2011/2002).
Instead, they correspond to Weiss and Buchanans alter-
native evolutionary framework. Weiss and Buchanan
argued that chance (action without direction that has
major impacts; p. 36) is one of lifes important regular-
ities (p. 40), that life is a contingent phenomenon (p.
35), and that adaptability in the face of changing cir-
cumstances means that responses are flexible and not
always prescribed in advance (p. 37). Predictability is
the basis of life processes (p. 21), and history and context
matter because these are the contingent conditions
upon which the future is built (p. 40). Weiss and Bu-
chanan demonstrates how this applies at the cellular,
Buchanan, 2009; Weiss et al., 2011). I suggest that it
also applies to baboon behavior.
PART 4: CAPTURING AND EXPANDING
COMPLEXITY IN THE ANTHROPOCENE
Anthropologists should be interested in baboons
because of what they tell us about the reality of daily
lives, about the challenge of complexity, and about how
evolutionary principles get embedded in contexts deter-
mined by history, chance, and contingency. Baboons add
evidence to the suggestion that evolutionary processes
contain more tolerance, multiple solutions, larger accept-
ability spaces, and the possibility that an adaptive fit

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DARWINS MONKEY
will be good enough rather seamless. When the
straight line of evolutionary scale opens up to capture
what actually happens, arguments become less reduc-
tionist and less deterministic [the pentagon (Weiss and
Buchanan, 2009, p. 9)].
Yet studying behavioral complexity in the wild poses
methodological challenges. How do we document com-
plexity and track the impact of chance, contingency, and
context? I see no other way than by returning to compar-
ative natural history as a scientific method (Dayton,
2003; Smith et al., 2011). Although the plural of anec-
dote may not be data [but see recent shift in medicine
(Aronson, 2003)], the plural of comparative natural his-
tory certainly is. I call baboons, Darwins monkey, not
just because he made frequent reference to them
(Darwin, 1871/2004) but because baboon lives once again
argue for the value of comparative natural history, the
key method that Darwin used. Following Darwins lead,
natural history observations should include well-docu-
mented details tracked across time, space, groups, land-
scapes, and species. They should be aided, when possi-
ble, by quantitative data and experiment. Long-term
studies become extremely vital because only they can
generate natural histories and the kind of insights pre-
sented here.
Baboons are also useful to anthropologists for reasons
that Darwin could not have guessed. Today, it is common
practice to end an article about primates with a com-
ment about conservation. The baboons offer a singular
perspective on primate conservation because they illus-
trate how the evolutionary game has changed in the
Anthropocene (Steffen et al., 2007; Zalasiewicz et al.,
2008), the current era of human-dominated ecosystems
and landscapes. The heart of the current biodiversity cri-
sis is the presence of a super-dominant species (Western,
2001), humans. People either directly or indirectly create
the current threats (Wilson, 2002; Barnosky et al., 2011).
The baboon data suggest that the speed of human-
induced changes and the diversity of their impacts are
likely to create more complexity and introduce an even
larger element of chance and contingency into evolution-
ary processes (as illustrated by some of the baboon natu-
ral histories). Baboons are not typical primates. Their
flexibility and adaptability and their range of ecological
and social strategies seem preadapted to new human-
modified landscapes which have an acceleration of pace
and fragmentation of space. This may create selective
pressures for weedy species (Richard et al., 1989); how-
ever, the baboon evidence suggests that it also puts a
premium on smart species [a benefit that may extend to
other taxa: see trends in urban passerine birds
(Maklakov et al., 2011)]. Human activity fragments ba-
boon populations when it fragments space. These diver-
sified but isolated conditions would select for new behav-
ioral responses in baboons, perhaps even at the level of
new traditions. Yet the baboon scenario is not all rosy.
The history of the Cape baboons in South Africa
(Hoffman and ORiain, 2010, 2012; Kaplan et al., 2011)
illustrates the likely future trajectory for many baboons.
The current reality for Cape baboons is both genetic iso-
lation and unsustainable humanwildlife conflict. If the
future of baboons is uncertain despite their special abil-
ities, the situation may be even direr for the majority of
species who are not as adaptable as baboons. Therefore,
those interested in primate conservation will need to pay
special attention to the accelerating humanwildlife con-
flict over old and new resources in the Anthropocene.
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19
This article wouldnt be complete without a mention of
what Darwins thought about baboons. His summary is
particularly apt: For my own part I would as soon be
descended from that heroic little monkey, who braved
his dreaded enemy in order to save the life of his keeper,
or from that old baboon, who descending from the moun-
tains, carried away in triumph his young comrade from
a crowd of astonished dogsas from a savage. . . and I
have given the evidence to the best of my ability
(Darwin, 1871/2004, p. 791). I dont think Darwin would
be surprised by the picture of baboons that I have just
presented.
ACKNOWLEDGMENTS
I thank the editor for his generous invitation to com-
ment on the important lessons from my years of baboon
research. My insights also rest on the work done by
many other individuals associated first with the Gilgil
Baboon Project and later with the Uaso Ngiro Baboon
Project. Since 1981, Kenyan paraethologists have been
the mainstay of the basic data collection. I thank them
all for their contributions. Finally, I thank David West-
ern for his distributed cognition; Katerina Semendeferi
for useful comments on the article; and Bruno Latour
who first helped me to see why complexity is so impor-
tant although it took many decades to collect the evi-
dence.
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