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YEARBOOK OF PHYSICAL ANTHROPOLOGY 55:323 (2012)

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- chies, to insights from a fission that happened when the
els for human evolution. I utilize 40 years of data from foraging strategy of raiding and nonraiding baboons
my long-term study on baboons in Kenya to suggest that diverged, to evidence of adaptation after translocation,
baboons are once again relevant for understanding and finally to assessing two unusual fusions of baboon
human evolution, not as a referential model but to reset groups. Altogether, these natural histories yield baboon
the starting conditions of the human experiment. The principles of the social with insights about cognition,
baboon data also offer a critique of widely held ideas cooperation, and culture and suggest why baboons cant
about how natural selection might work by looking at become human. The data also support Weiss and
real lives in real time. This situates competition in a ma- Buchanans framework (The Mermaids Tale: Four Bil-
trix of collaboration and illustrates the critical role of lion Years of Cooperation in the Making of Living
chance, contingency, and history in baboon survival and Things. Cambridge, MA: Harvard University Press,
success. I make three methodological moves to reach 2009. 305 p) with its alternative view of natural selec-
these conclusions. The first is to focus on process not tion in which there is more slippage and tolerance, mul-
just outcome. The second is to look at time scales longer tiple solutions with larger acceptability spaces, and the
than usual studies but shorter than evolutionary time as possibility that an adaptive fit will be good enough
a way to open the black box that currently links behav- rather than seamless. However, capturing behavioral
ior to evolutionary value. The third is to use compara- complexity in the wild poses methodological challenges.
tive natural history, Darwins method, as a way to cap- Long-term field studies provide critical information but
ture and comprehend how complexity is generated and the current quantitative methods should be expanded
how baboons deal with it in their daily lives. These em- also include natural history observations of behaviors
pirical and methodological turns lead to conclusions that and events across time, space, groups, and landscapes.
run counter to widely held ideas about baboons, about Finally, the baboon natural histories illustrate how the
primates, and about the determinism of natural selec- evolutionary game has changed in the Anthropocene
tion. I follow my own research history to illustrate these yielding a cautionary tale about the future for many
points. The discussion ranges from alternative interpre- primate species. Yrbk Phys Anthropol 55:323, 2012.
tations of the male and the female dominance hierar- C 2012
V Wiley Periodicals, Inc.

Baboons have played a central role both in our under- males were the most reproductive. The male dominance
standing of primates and as models for human evolution hierarchy also was the political structure of the group; it
beginning with Zuckermans 1932 description of primate ordered male interactions reducing the need for frequent
society. Based on his observations of Hamadryas baboons aggressive encounters. Males defended and policed;
at the London Zoo, he argued for a society founded on females raised babies. Males were central to all major
sex, violence, and male domination of both females and group functions, whereas females were defined in terms
males (Zuckerman, 1932, 1933). Zuckerman can be for- of their relationships with males. Washburn and DeVore
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 Grant sponsors: National Science Foundation, Wenner-Gren
public) imagination three decades later through the Foundation for Anthropological Research, University of California,
early scientific work on savanna baboons done by San Diego, National Geographic Society, Fyseen Foundation, Wild-
Washburn, his student, DeVore, and the English psychol- life Conservation International, LSB Leakey Foundation, Art Orten-
ogist, Hall (DeVore, 1965). Once again baboons became berg and Liz Claiborne Foundation, African Conservation Centre,
African Conservation Fund.
emblematic of evolution; however, this time it was the fit
between behavior and anatomy which told a convincing
*Correspondence to: Shirley C. Strum, Department of Anthropol-
story about how natural selection worked (Washburn
ogy, University of California, 9500 Gillman Drive, San Diego, La
and DeVore, 1961, 1963; DeVore and Hall, 1965; Hall Jolla, CA 92093-0532, USA. E-mail: sstrum@africaonline.co.ke (or)
and DeVore, 1965; Strum and Fedigan, 2000b). Evolution sstrum@ucsd.edu
gave male baboons the anatomy of aggression. Baboon
males used those remarkable canines, large body size, Received 6 September 2012; accepted 7 September 2012
impressive mantle of hair, and white eyelids in displays
during sexual competition with other males and against DOI 10.1002/ajpa.22158
predators. Fighting ability determined male rank which Published online 17 October 2012 in Wiley Online Library
in turn governed reproductive success so that the best (wileyonlinelibrary.com).

C 2012
V WILEY PERIODICALS, INC.

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4 S.C. STRUM

(1961) argued that we could infer the challenges that the 1987; Strum, 2005). Translocation was an experiment
savanna presented to a puny primate living away from that provided persuasive evidence about baboon adapta-
the safety of the forest (read hominids) by looking bility. The new location, arid savanna occupied by no-
through the lens of the only other primate that managed madic pastoralists, has had both natural cycles of extreme
to do so successfully, baboons. events like droughts and in the last decade, human-
What follows is not a scholarly review of the baboon induced changes that also created extreme events. Else-
literature and its relevance to human evolution. where (Strum and Western, 1982), I have argued that the
Instead, I was invited to discuss my insights about baboons dont distinguish between the sources (natural
baboons based on 40 years of observing them in the vs. human) of environmental change in their responses to
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 processes I will explore. In this presentation, I touch on
(Goodall, 1988/1971) and then by bonobos (Zihlman, areas that are minefields of controversy, for example,
1978; Kano, 1992; Strum and Fedigan, 2000b). In con- what is culture or tradition and what is cooperation. For-
trast, I will suggest that baboons are once again rele- tunately, my argument does not depend on resolving
vant for those interested in human evolution but in these controversies, a task I leave to others.
new ways. What follows is an argument that is mainly In this article, I move through a series of steps that
bottom up, grounded in baboon observations. In the reflect my own research history rather than their order
end, I hope to illustrate that the baboons I have studied of importance. During this time, I have gone from mak-
live in the most complex nonhuman primate society yet ing baboons almost human by focusing on their social
described and that they reside in the most socioecologi- negotiation and politics (Strum, 2001/1987a) to under-
cal complexity documented so far for a nonhuman pri- standing better why baboons arent human. I have col-
mate. Furthermore, these baboon data suggest that fo- lected quantitative data for four decades; however, I find
cusing on real time and real lives provides a different these data inadequate to make sense of what the
view of how evolutionary processes work. Chance, con- baboons are doing. Instead, I turn to comparative natu-
tingency, and history play an important role in survival ral history for the missing insights. The title Darwins
and success. This offers a critique of widely held ideas Monkey refers both to the importance of Darwins meth-
about natural selection, particularly the centrality of odology as well as his prescient observations of baboons
competition. The data also argue for the existence of (Darwin, 1871/2004).
multiple options (including mistakes) made possible by My over-arching goal, in this article and in my
the intelligence and social skill that baboons use to research, is to recapture the complexity that exists in ba-
build their primate society. I am often asked why, if boon lives and then to consider the implications of this
baboons are so smart, they arent human. There are complexity for evolutionary interpretations and for our
many things wrong with the way this question is current scientific practice. I end with a discussion of why
phrased. However, the perspective I present below does understanding baboons is important for [physical/biologi-
add a new factor to the list of obvious differences: the cal] anthropologists today.
importance of managing complexity in daily life.
To reach these conclusions, I made three critical meth- PART 1: GENERATING COMPLEXITY
odological moves. The first was to focus on process not Remodeling the male dominance hierarchy
just outcome. The second was to look at time scales longer
makes baboons almost human (19721980)
than usual studies but not collapse everything into evolu-
tionary time. The third was to use comparative natural It is hard to believe in the 21st century how little we
history, Darwins method. Primate studies rejected natu- knew about primate behavior in the 1960s and how rudi-
ral history observations when growing knowledge about mentary were our methods. The pioneering work of the
primates grew and more sophisticated data collection classical ethologists like Tinbergen, Lorenz, and von
methods were available (Strum and Fedigan, 2000a). For Frisch starting in the 1930s (Tinbergen, 1942, 1963; Lor-
now, however, natural history may be the only way to enz, 1950) did two key things. It critiqued (previous)
capture and comprehend how complexity is generated anthropomorphic interpretations of behavior and created
and how baboons deal with it in their daily lives. a novel methodology that still relied on natural history
These empirical and methodological turns lead to con- [ethogram, fixed action patterns; critical periods; and
clusions that run counter to widely held ideas about interaction of field data with captive experiments (Tin-
baboons, about primates, and about the determinism of bergen, 1974/1951)].
natural selection. This is not simply an emphasis on When primate field studies resumed after World War
proximate rather than ultimate levels of causal explana- 2, scientists still used natural history descriptions but
tion (Tinbergen, 1963). Instead, it tries to open the black added the ethogram, a careful and detailed description
box linking behavior and evolutionary outcome and of the behavior of that species (or group, or popula-
raises the question of what can be legitimately assumed tion. . .these terms were interchangeable then). In the
about shorter time scales when we make evolutionary middle of last century, there were only dozens of studies
time arguments. I do not review the extensive literature of primates in the wild, not hundreds, and only a few
on all these topics. Instead, I include key references as a species had been observed in more than one location
starting point for readers interested in exploring further. (DeVore, 1965; Altmann, 1967; Jay, 1968).
My reference species is olive baboons, Papio anubis Variation in behavior seemed minimal, making broad
(Zinner et al., 2009). The reference material is the 40 generalizations about primate patterns seemed war-
years of my own research (see Strum and Fedigan, ranted (Washburn and DeVore, 1961; Strum and Fedi-
2000b) for a framework to situate most of these years). I gan, 2000b). However, by the early 1970s, variation in
track individuals, relationships, and social groups in two behavior and social organization began to accumulate.
populations. The baboons lived in two locations because I For example, patas monkeys solved the problems of sa-
translocated three groups in 1984 (Strum and Southwick, vanna living with an anatomy adapted for rapid escape

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DARWINS MONKEY 5
and behavior adapted for vigilant diversionary males 1967 study but hadnt yet appreciated their function.
and stealthy females and young (Hall, 1968). While Kekopey males created special relationships and later
Washburn and DeVores dominant males strode out to used them to advantage. For example, males used young
face a predator, Rowells male baboons were the first to infants as agonistic buffers against other males (Strum,
run to the trees, leaving the females and youngsters 1983c). Usually, it was the potential loser who got an
exposed (Rowell, 1966). Natural history methods were infant, returning to the protagonist with the baby on his
necessary whenever a new species was studied; however, belly. Most often, the aggressive male backed off. A male
disagreements of fact were hard to resolve. Were the could also use a female as a buffer (Strum, 1983b). Why
animals really different or were differences an artifact of did it work?because buffering allowed a male to mobi-
observers and ways of observing? lize troop support. Normally, other group members dont
When I began my baboon study on male and female get involved in malemale interactions. However, the
roles in olive baboon society (19721974, Kekopey near group will mob a male whose aggression causes distress
Gilgil, Rift Valley, Kenya), there was already evidence to infants and females. A pivotal element in buffering as
from rhesus macaques on Cayo Santiago Island in the Ca- a successful tactic is trust. Trust is a social construct for
ribbean that females, as well as males, had a dominance humans and for baboons. It makes social life predictable,
hierarchy and that matrilineal relationships were the creates a sense of community, and makes it easier for
core of the group (Sade, 1967, 1972). As well, there was a individuals to work together (Holmes and Rempel, 1989;
move to use more systematic ways of data collection in Misztal, 1996, 2001; Rousseau et al., 1998; Kappeler and
the field [well summarized by Jeanne Altmanns often van Schaik, 2006; Henrich et al., 2009; Sueur et al.,
cited paper (Altmann, 1974)]. All baboon projects from the 2011b). In economics, trust is a way to reduce transac-
early 1970s onward inherited a baboon ethogram from tion costs, creating new forms of collaboration and it is a
Hall, Washburn, DeVore, Rowell, and Ransom (Washburn form of social capital. For baboons in the context of ago-
and DeVore, 1961; DeVore and Hall, 1965; Hall and nistic buffering, trust implies a particular social history.
DeVore, 1965; Rowell, 1966; Ransom, 1981). This meant It means that the male has been predictably affiliative
that the data included the same basic elements of behav- with the infant. Both the infant and the male have
ior. When possible (meaning in contexts where the prima- expectations of each other based on their social history.
tes could actually be seen), investigators relied on focal If the buffer trusts the male friend, it will scream at the
animal samples (follows for a fixed duration of randomly aggressive opponent. Without trust, a male runs the risk
chosen individuals) as a way to minimize bias. of himself becoming the object of mobbing if the infant
I collected my data this way. The results confirmed screams at him rather than at the opponent. This
that matrilines (female based families) were the core of implies that the social relationship comes first; using the
the group and that a matrilineal dominance hierarchy relationships in a social strategy comes later.
was the troops main structure. Families, rather than Males got other benefits from friendships with
just adult males, protected and policed the group on females. Following ethological tradition, I refrained from
behalf of their own members. Females had political calling special relationships friendships until I was
functions besides motherhood (see also Hausfater, 1975; convinced that baboon and human friendships shared
Ransom, 1981). many characteristics (the term special relationship was
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 ning in the 1980s). There is no rape among baboons, and
reaching the asymptote of their growth (Strum, 1991) therefore, while males might fight to get a temporarily
and fully adult males ranging in age from young ([10 exclusive association with a sexually receptive female, or
years) to old (more than 20 years). Hausfater, on yellow consort, the consort male needs the females coopera-
baboons in Amboseli National Park in Kenya (Hausfater, tion to successfully copulate with her. Consorts may
1975; Hausfater et al., 1982), documented a relatively attract many male followers. Friendship between the
stable male dominance hierarchy that operated as pre- consort male and female also influenced whether a male
dicted. The dominant male(s) had priority of access to can keep the consort in the face of pressure from fol-
limited resources and mating opportunities. The lower males (Smuts, 1985; Strum, 2001/1987b).
Hausfater study was a careful and quantitative refine- Other social tactics relied on social knowledge and
ment of the previous baboon model. By contrast, I found assessment skills. For example, older males with less
male rank relationships exceedingly dynamic (Strum, aggressive potential still got consorts and copulations.
1982). Individuals spent time and energy working on One way was through a maneuver I called sidelines. In
their relationships with each other and tracking the sidelines, a male monitors the consort from a distance.
changes in all male relationships. Processes of male mat- He makes his move when aggression breaks out between
uration and male migration fueled the instability. the consort male and his close followers. Then, the more
Besides its dynamic nature, the existing male hierar- distant monitoring male dashes to the female. By the
chy did not correlate with access to resources such as time the male aggression is over, the older male has
receptive females. Often rank did not even predict the copulated with the female (becoming her new consort),
winner of an aggressive encounter (Strum, 1982). The has skillfully led her away from the other males, and is
reason was that baboons, including the males, had alter- deftly grooming her. Ironically, another avenue for males
natives to aggression. These social strategies of competi- with little aggressive potential is to manipulate aggres-
tion and defense (Strum, 1975a, 1976; Western and sion between consort male and followers. He can do this
Strum, 1983) could be as effective and were less risky as a follower or from the sidelines, escalating aggression
than aggressive approaches. However, they depended on by his involvement but usually leaving before another
a number of factors for success. First, some relied on male turns on him.
special social relationships. Ransom (1981) identified An individuals age and length of residency in the
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 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 a system with numerous elements and many forms of
life history from socially unskilled maturing male with relationships which are all entwined (Latin com-
growing physical powers (using aggressive strategies) to plexus); complex systems have more parts and more con-
the socially skillful mature male whose success depends nections than simple ones (Dictionary, 1971).
entirely on social expertise when he is old (using social
strategies of competition and defense).
Males werent the only beneficiaries of social strat- Remodeling socioecology: Group size does not
egies. Infants and females could rely on the support of determine group fission (19811984)
their male friend when aggression was directed at them.
Today, group size is thought to be a trade-off between
Even the close proximity of a male friend reduced social
the advantages (predator protection, resource discovery,
interference that a female or infant would receive based
on rank and age. and for primates, traditional knowledge of resources) and
disadvantages (feeding competition and inefficiency) of
Complexity 1: Social strategies in addition to living in a large group when compared with small group
aggressive strategies create multiple options (Wrangham, 1980; van Schaik, 1983; Isbell, 1994; Isbell
and Young, 2002). Group fission happens regularly
Science makes simplifying assumptions to be able to among baboons (although there is a dearth of published
begin studying complex phenomena such as social inter- information: for baboons, see Stoltz, 1972; Nash, 1976;
actions. The problem comes when the results of those Hamilton and Bulgar, 1993; Henzi et al., 1997; Wasser et
investigations slide into thinking that such systems are al., 2004; Van Horn et al., 2007; but for macaques, see
simple. Uncoupling the tight link between male domi- Furuya, 1969; Chepko-Sade, 1974; Chepko-Sade and
nance rank and male evolutionary payoffs was the first Sade, 1979; Dittus, 1988; Kuester and Paul, 1997; Oka-
evidence I had of social complexity among baboons. moto and Matsumura, 2001; Kawamoto, 2010). Theoreti-
Baboons created and managed special relationships. cally, a group should split when it gets too large to forage
They adjusted their behavior based on expectations from effectively (increased feeding competition and/or has to
past interactions (Strum, 2008). Social strategies were travel too far to feed that many individuals), or too large
also evidence of social collaboration (see section on col- to monitor and manage the many social interactions, or
laboration later) and exchange. Among baboons, even
for both reasons.
the physically impressive males needed the help of less
In 1981, the main study group, Pumphouse, fissioned.
imposing individuals to succeed. Baboons, it seemed, had
The split centered on a disagreement about foraging
no option but to play by the golden rule.
At this point in my research, negotiations between strategies not about group size as Pumphouse was well
baboons stood out in bold relief. Human negotiation is below the numbers recorded in other baboon fissions (see
usually a dialog between two or more individuals. The above). An agricultural cooperative bought Kekopey
dialog aims for a common understanding about a course Ranch, the study site, in 1976. Kekopey covered 45,000
of actions that satisfies the interests of the parties acres. Although it was a cattle ranch, there were few
involved (Homans, 1958; Festinger, 1985/1957; Misztal, cows, a large complement of wildlife, and minimal farm-
2001; Raiffa, 2003/1982; Pearce and Hans-Werner, 2008; ing potential. By 1981, small-hold farms (shambas)
West and Turner, 2009). I saw baboon negotiations as be- appeared in the baboons traditional home range. Then
havioral conversations about investments in, and payoffs humanbaboon conflict began (Strum, 2001/1987b, 2010).
of, special relationships. Clearly, such interactions select Young adult and large adolescent Pumphouse males
for very different skills than would aggressive competi- were the most motivated to raid. Troop movements away
tion in, for example, mating success (see also Sussman from the sleeping sites included extensive negotiations
and Garber, 2011). The conclusion: a primate with a big between these males and the rest of the group. Some
brain can create more alternatives than previous evolu- days, the raiders pulled the rest of the troop with them
tionary models suggested. I argued that these kinds of but on other days they failed. Then only the raiders
negotiations, social strategies, and sexual politics made went to the fields. There was a dramatic increase in
baboons almost human (Strum, 2001/1987b). anthropogenic injuries and deaths because of the conflict
A brief note about my use of the term complexity is (Strum, 2010). Pumphouse spent over 6 months in the
warranted. The study of social complexity has generated process of splitting. During this time, there were con-
a huge literature since these first baboon observations stant negotiations between raiders and nonraiders as
(e.g., see Byrne and Whiten, 1989; Whiten and Richard, well as among family members of raiders and nonraiders
1997; de Waal and Tyack, 2003; Cheney, 2007). Despite as they decided which group they would join. In the end,
this, there is no clear definition of social complexity Pumphouse, the larger descendent group remained pri-
(Strum et al., 1997). Whiten (2000) suggested that social marily natural foragers, whereas the smaller descendent
complexity must include the dimensions of variability of group (Wabaya) became dedicated raiders.
response, size of group, number of relationships, instabil- However, these werent the only responses. A third
ity, and predictability. Dunbar (1998) found a strong cor- study group, Eburru Cliffs (larger than the other two
relation between group size. Ironically, Robert Hindes and having the largest home range), tried human food,
1970s model of interacting levels that moves from indi- sustained the costs of conflict, and then completely aban-
viduals to interactions to relationships to group structure doned the settlement area. They remained natural forag-
is probably the best approximation to an operationalized ers using the part of their traditional home range that
definition of social complexity [Hinde, 1976, 1987; and was as far away from the settled area as they could get
see Forster and Rodriguez (2006) for an example of how (a distance of over 7 km as the crow flies).
this would work]. In another section, I offer a specific def- Why raid? Human food offers significant benefits to
inition of baboon social complexity. Until that point, I use baboons. Both males and females grew faster, reached
data to build the dimensions of baboon complexity and adulthood earlier, and had heavier final weights (Strum,
use an everyday definition of complexity: being intricate, 1991, 2010), a result found in other studies of food

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DARWINS MONKEY 7
enhanced baboons (Altmann and Muruthi, 1988; Muru- them made the switch. Not all females became raiders. I
thi et al., 1991; Altmann et al., 1993). Faster growth expect that the conservativeness of baboon females
meant earlier female reproduction. Although the sample played an important role and for a few females a special
size is small, raider females reached menarche sooner social relationship (friendship) with a raider male was
and produced their first baby earlier than nonraider the defining influence. However, even in this smaller
females. Interbirth intervals were significantly shorter subset, there were exceptions: the lowest ranking female
in raider females when compared with nonraiders. An and her daughter were not friends with any raider male
extreme example of this was the lowest ranking raider when they joined the raiding group. Equally important,
female. Her fecundity increased significantly when she not all troops with the opportunity chose to raid.
became a raider, which reversed her previous age-related Paying attention to the process, to the natural history
reproductive decline. Infants who grew up in food- of events and individuals, showed me baboon complexity
enhanced conditions reached menarche and first birth beyond social strategies and added to my appreciation of
earlier even though these landmarks occurred after they how evolution works at the level of real lives.
were translocated to a resource-poor environment. Infant
survivorship did not differ between the raiders and non- Translocation: Survival depends on others not on
raiders despite initially high rates of injury and death competition between individuals (19842000)
associated with raiding. With time, raiders adjusted to
the risks of being close to humans so that costs declined Darwin relied on the idea of survival of the fittest to
(Strum, 2010). explain evolution by natural selection (Darwin, 1859).
What were the baboons doing during those 6 months of Subsequent evolutionary interpretations of animal (pri-
the fission? The matriarch of the lowest ranking family mate) behavior focused on competition as the key to indi-
provides one illustration. One day she would be with the vidual survival. Sociobiology (Wilson, 1975) enlarged the
raiders; the next day she was with the nonraiders. This individual to include kin in the mid-1970s [inclusive fit-
went on for months. Her already independent infant, ness (Hamilton, 1964)], and recently, group selection has
who still closely associated with his mother, moved back been resuscitated as possible and important for humans
and forth with her unlike the females adolescent sons (Richerson and Boyd, 2005; Leigh, 2010; Nowak et al.,
who remained with the nonraiders. Eventually, after 2010; West et al., 2010). Usually, scientists infer what
many migrations back and forth, that female became a behavior is crucial to survival and reproductive success.
permanent member of the raiding troop but her infant By contrast, the translocation of the study baboons
did not. He returned to the nonraiders and stayed with became an actual test of survival where the process could
his brothers. Months later, the matriarchs subadult be documented and the outcome could be measured.
daughter also became a member of the raider group. Despite great success with crop-raiding research and
Females could have several reasons for joining the control techniques, in 1984, I had to translocate three
raiding group, Wabaya. Most often, the emigrating troops of wild baboons (Pumphouse, Wabayathe
female was following her closest male friend who hap- raiders, and a third troop called Cripple who were also
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 was the first scientific translocation of a primate group
the raider group were most often the oldest daughters (Strum and Southwick, 1987). The translocation pro-
and hence the lowest ranking females in their matriline. vided a unique opportunity to document adaptability
By joining the raider group, they traded kin support for (possibly adaptation) in action. I could compare the same
the benefits of male friendship and of raiding. They also individuals and the same troops in two radically differ-
avoided internal matrilineal disputes and reduced the ent environments. The translocation site, on the Eastern
number of females to compete with because Wabaya was Laikipia Plateau, is arid savanna with half the annual
smaller. Three pairs of sisters transferred to the raiding rainfall of Kekopey, the baboons previous home. Serious
troop. The social framework that underpinned their pre- droughts are common. The baboons were familiar with
vious rank relationship to each other disappeared and only about half of the foods available in the new location
their rank reversed in Wabaya. The older sister became and the unknown ones turned out to be critical to sur-
dominant to the younger one. By contrast, some things vival during seasonal bottlenecks. The release site was
didnt change. Despite having only a few matrilines rep- not ideal; however, it was the only site I could get.
resented in Wabaya, the original Pumphouse female hi- The descendants of the translocated baboons are alive
erarchy was perfectly reproduced albeit with some today. The process by which they survived provided
females missing. more evidence of the complexity of baboon life and illus-
Complexity 2: Context becomes history as baboons trated evolutionary raw material.
negotiate socioecology Primates are social; it is a key characteristic of the
Order. There are a number of theories about why most
The study troop did not split because it was too large. primates live in a social group (see above); however, it is
What had happened during the fission added evidence difficult to document the value of group living during the
about the socioecological complexity of baboon lives. course of most studies. The baboon translocation pro-
Most individuals selected not to raid despite the major vided one chance. The benefits of living in an intact
documented benefits of the raiding lifestyle. Individuals social group permeated almost every aspect of adjust-
with the same characteristics chose different paths. To ment during the translocation.
my surprise, other factors trumped evolutionary princi- To start, I manipulated existing social networks to
ples like optimal foraging and kinship as well as derived ensure that the troops stayed where they were released.
principles like hierarchy. I held adult males captive at the release site, whereas
Why didnt all males become raiders? Large subadult the females and youngsters were set free. As expected,
and young adult males had the most to gain in terms of the females did not go far without the males. By the
growth and weight (and reproduction?) yet not all of 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 the previous raiders (now called Malaika trading the bad
tendency to wander (Strum, 2002, 2005). guys for the the angels) who were translocated to the
Each of the three troops was released in a different same location stayed put from 1984 until 2007.
place and at different times. The troops maintained All but the first home-range shift tracked a resource
cohesion and coordination from the first days of freedom. gradient; however, information about and the attraction
The group represented a social learning resource in a of food resources werent enough to force a move. The
more basic sense than we usually assume. Individuals in shifts depended on social processes. Baboons are not ter-
the translocated groups demonstrated flexibility in learn- ritorial; adjacent groups have overlapping home ranges
ing in novel settings. Social cues figured prominently in with undefended boundaries, although intensively used
the process of acquiring information. For example, the core areas are sometimes defended. Normally, the con-
translocated troops had to construct a diet in this unfa- servativeness of females prevents a baboon group from
miliar place. Social facilitation played a role as was crossing home-range boundaries. Each subsequent
documented in another context before translocation Pumphouse range shift followed an influx of immigrant
(Strum, 1983a). The incorporation of new foods into the males [see Rowells suggestions about sex-biased knowl-
diet was strongly related to specific social events. For edge (Rowell, 1972)]. These immigrants had firsthand
example, adults in translocated troops watched and fol- experience of the resource gradient because they came
lowed indigenous baboon troops. The strangers built from there. However, even with that knowledge, one
their home range by tracking the path of local troops male was ineffective. It took a cohort of four or more
and foraging on some of same foods. In addition, when males, all indicating in the same direction during troop
local males migrated into the translocated troops, they movement, to succeed. There was a time lag because the
brought indigenous information. The translocated males had to first create friendships with females in
baboons behavior tracked the immigrant males behav- order to influence troop direction. Clearly, knowledge
ior. In this way, they learned to find water in dry sand was vital, but action relied on social processes.
rivers and began using several novel large succulents
(Sansevieria intermedia, Sansevieria abyssinica) for both Complexity 3: The social group is an organ of adap-
moisture and food.
tation; accidents of history create different socio-
Another value of the social group could only be called
ecological paths
emotional support (Hannah and McGrew, 1991). This
was evident during the accidental fission of one group at
The translocation was a real test not an imaginary
release and their subsequent reunion. Loud greetings,
one. If ever there was a situation where survival of the
extensive embracing, and other amplifications of typical
fittest should operate, where competition should have
baboon reunion signals punctuated this meeting (Strum,
had an upper hand, this was it. Instead other baboons,
2001/1987b). It was the exaggerations of these normal
either from within the translocated group or the indige-
behaviors that pointed to the stress of separation and
nous population facilitated both learning and survival.
the relief of reunion.
The translocation, even more than the crop raiding,
A comparison of the baboon translocation with subse-
pointed out how evolutionary principles get embedded in
quent primate reintroductions and translocations add
a specific time and place and why context matters. Each
force to my conclusions. Survival skills are lost in captiv-
groups natural history illustrated a variety of different
ity. Artificially created groups fall apart (Kawai, 1960;
paths. Chance (like an encounter with elephants), con-
de Vries, 1991; Vie and Richard-Hansen, 1997). Under
normal circumstances (and in extraordinary conditions), tingency (how many immigrant males from which area),
the social group provides the context for appropriate and evolutionary principles interact to generate baboon
learning and as a resource for that learning (King, 1994; options and baboon responses. The process was infinitely
Russon, 1999; Crast et al., 2010; Leca et al., 2010). Rein- more complex than explanations collapsed into evolution-
troduced captive primate groups fail to meet the learn- ary time might suggest.
ing challenges of their wild habitat even after extensive
pre-release training (Kleiman and Rylands, 2002). Fusion of two groups: Baboons negotiate the
The translocation gave other insights about adaptabil- nature of their social group (20002002)
ity, chance events, and the importance of the social. A
night-time encounter with elephants was an accident Fusion, when two groups merge to become one, is rare
that changed the future of one of the translocated troops in baboons, so rare that it has been observed only three
(Strum, 2001/1987b). Surrounded by elephants at their times, once in baboons in Amboseli National Park in
sleeping site, Pumphouse moved 5 km during the night, 1960s (Altmann and Altmann, 1970; Altmann et al.,
an unprecedented act for baboons and other diurnal pri- 1985), once among baboons in Mikumi National Park,
mates. I know this because they marched past the lone Tanzania (Wasser et al., 2004), and once at De Hoop in
human outpost in the area to reach another set of sleep- South Africa (Henzi et al., 2000). There are only a few
ing rocks at the edge of their home range. Subsequently, other cases reported for cercopithecine monkeys despite
Pumphouse abandoned more than half of their range the hundreds of thousands of hours that cercopithecines
including the best rainy season portion. Following that have been studied in the wild. These include two vervet
move, group size declined as males and even mothers fusions (Hauser et al., 1986; Isbell et al., 2002; Isbell,
with infants left to join other, indigenous, baboon troops. 2009), one toque macaque fusion (Dittus, 1987), and sev-
Just when it seemed that Pumphouse might simply be eral cases of fusion in Japanese macaques (Maruhashi,
absorbed into the local population, the exodus stopped. 1992; Takahata et al., 1994; Sugiura et al., 2002).
New males joined and the next stage in this troops his- Descriptions suggest that the fusion happened almost
tory began. instantaneously (or over a short time for toque maca-
There were subsequent home range shifts but none as ques). Two separate monkey groups come together in the
precipitous as the first. By 2000, Pumphouse ranged more evening at a shared sleeping site and leave the next day
than 20 km from their release site. By contrast, Wabaya, as an integrated merged group.

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DARWINS MONKEY 9
This is quite different from fission/fusion societies when Soit Oitashe noticed, they searched for them and
among primates where coming together and splitting aggressively mobbed Malaika when they found them.
into smaller, usually foraging groups, happens on a For two groups to merge, the two female hierarchies
daily or weekly basis (Aureli et al., 2008). Chimpanzees needed to be integrated. Initially, Malaikas high-ranking
(Lehmann and Boesch, 2004) and spider monkeys females kept pushing all of Soit Oitashes females. With
(Chapman et al., 1995) are well-known examples. the support of their male friends, Malaika females man-
I have documented four baboon fusions in just over a aged to be in the middle of the integrated dominance hier-
decade. The first fusion was 16 years after translocation archy and forcing their way upward. However, Malaika
and involved one of the translocated troops, Malaika (the males stopped supporting their females. Without male sup-
angels, previously the bad guys) and an indigenous port, Malaika females dropped to the bottom of the inte-
group, Soit Oitashe, that had been studied since 1985 grated female hierarchy while maintaining their original
(Barton, 1993). What made this (and subsequent fusions) rankings among themselves.
so distinct is that the process took more than 2 years. Specific individuals had disproportionate impact on
During that time, individuals in each group engaged in the degree of mixing between the two groups. This
extensive negotiations about merging both within the included two Malaika females and specific males from
group and across the groups. both groups. Low-ranking Soit Oitashe females also
The impetus for the first fusion involved several eco- became the most active and gave the most aggressive re-
logical events including the creation of a wildlife sanctu- sistance to any attempt by Malaika females to push up
ary on communally owned land that overlapped the the integrated female hierarchy. An illustration of the
baboons home range. The sanctuary led to an increase influence of specific baboons was the effort that one Soit
in prey and predator numbers. Later, the large mammal Oitashe male exerted to keep his troops females from
prey species left. This made local pastoralists dogs and interacting with Malaika. The day he emigrated, there
then the baboons the prey. Predation rates rose to un- was a noticeable increase in intertroop interactions and
precedented levels. in spatial proximity between members of the two groups.
Who did what and when during the fusion process Given the length and extent of the process, I had to
depended on specific events and on daily negotiations develop criteria to use to decide when the two troops had
become one group. The criteria reflected what I felt were
involving particular individuals and age-sex classes. The
aspects of normal group sociality. These included the ab-
translocated troop, Malaika, was the smaller of the two
sence of previous troop bias in proximity, grooming and
troops. Malaika shadowed the indigenous group, Soit other social interactions, and troop movement negotia-
Oitashe, initially, but roles switched as Soit Oitashe, the tions with indicators and followers responding across
indigenous group, lost members to heavy predation. previous troop lines. Foraging subgroups should be
The sum of the forces of attraction and dispersal deter- mixed, and the fused group should react as an inte-
mined how mixed the two groups might be at any one grated troop in encounters with other groups, during
time. For example, adult and subadult females were very predations by baboons, toward new male immigrants, to
attracted to new babies in the other troop. Sexually recep- separations and reunions, and to social disruptions. Of
tive females also approached the other troops males. Sev- course, the fused group had to have an integrated female
eral females from Malaika were drawn to a male friend dominance hierarchy and an integrated male hierarchy.
who had recently emigrated back to his natal troop, Soit It therefore should not be a surprise that integrating
Oitashe. Large subadult and adult males, in both troops, two social systems took such a long time.
herded their females away from the vicinity of the other Why fuse? Why fuse now? The answers to those ques-
troops males trying to keep the two groups apart. This tions offer a test of major socioecological theories of
was only a small part of the daily push and pull of indi- socialness (see above and Dittus, 1986). Some of the
viduals, relationships, and classes of animals. fusions reported in the literature took place because one
Leaving the sleeping rocks in the morning no longer or both groups were too small to be viable in terms of
followed a standard procedure because each group had social and sexual interactions and predator protection.
its own indicator animals and there was no certainty Protection against predation likely played a role in the
about who was going to follow whom. Foraging during first baboon fusion I documented, although the sizes of
the day was also problematic because Malaikas smaller both troops before merging were within the normal
home range was entirely contained within Soit Oitashes range for olive baboons and had all agesex classes rep-
larger home range. Initially, Malaika refused to cross its resented. Yet before the fusion, both lost many group
members to predators. By contrast, after the fusion, the
own boundaries when Soit Oitashe moved on. Instead,
merged group lost none despite the continued presence
Malaika waited, intercepting Soit Oitashe on its way
of predators in the area.
back to the sleeping site. As the fusion progressed, The details of negotiations suggest, however, that the
Malaika did cross this invisible line, albeit hesitantly, baboon reality was much more complex than simply
thereby benefiting from access to new resources. Eventu- defense against predation. Malaika traded off feeding
ally, the home range of the fused troop (now called Nabo, competition and protection from danger on a daily basis
Maasai for coming together) represented a compromise until the fusion was complete. Once fused, it bore the
between the two previous home ranges. cost of feeding competition but benefited from enhanced
Feeding competition also played a role during the predator protection. Soit Oitashe benefited from better
fusion process. For example, acacia flowers are a pre- protection while remaining dominant in feeding competi-
ferred and high-quality food (Barton, 1993). Flowers tion. The costs and benefits of fusion, therefore, were dif-
emerge during the dry season. Their distribution is lim- ferent for Malaika and Soit Oitashe. Costs and benefits
ited both at the start and at the end of blooming. During also diverged for different agesex classes and even for
these times, Soit Oitashe displaced Malaika from the few individual baboons. Moreover, the benefit to cost ratio
trees that had flowers. In response, Malaika would sneak changed based on season, location, and day. This push/
away to feed undisturbed and out of sight. However, pull of fusion was even more elaborate than the process
Malaika paid a high price for this maneuver because of fission I had documented during the crop-raiding era.

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10 S.C. STRUM

Complexity 4: Baboons negotiate almost everything, gaining limited insights about how behaviors fit together
even the nature of their social group into a complex whole or how that complex whole can
change during the lifetime of the individual. The baboon
Baboon negotiations during the fusion were about the data offer a dynamic and integrated view of real-life be-
nature of the social group to a degree not yet docu- havioral complexity. From this vantage point, the inevi-
mented for a nonhuman primate species. The fact that tability of an individuals choices and therefore the inevi-
the process took over 2 years and involved dyadic, tria- tability of the link between proximate and ultimate cau-
dic, and polyadic interactions in the context of every ba- sation seems less compelling.
sic aspect of baboon life suggests that the term nego-
tiation, in its human sense, is appropriate for baboons PART 2: MANAGING COMPLEXITY
as well.
The negotiations were complex. The socioecology of the
Managing complexity: Redefining the meanings
fusion was complex. The interaction of levels was com- of social
plex. However, there was a ladder of complexity In 1987, Bruno Latour and I proposed a different
[building on Hindes model of levels (Hinde, 1976, 1987)] approach to compare human and nonhuman primate so-
from dyadic interactions, cascading upward to relation- ciety (Strum and Latour, 1987). We were struck by the
ships and subgroupings, and clusters of subgroupings to unlikely similarities between his subjects, scientists in
troops each with their own histories and negotiating laboratories (see Latour and Woolgar, 1986/1979), and
interests relative to each other and to higher and lower mine, baboons in the wild.
levels. This was embedded in environmental complexity, Baboon negotiation, even at the minor level I had docu-
which operated at different spatial and temporal scales. mented back then, raised a profound question: does soci-
As ecological factors are often not correlated (e.g., the ety exist or is it created? At the time, we contrasted two
effects of rainfall, food availability, predation, disease, major views about society. What holds society together?
and human impacts are often independent of each Traditional approaches to this question assume that soci-
other), one simple principle cannot govern what the ety exists and actors enter society adhering to rules and
appropriate response will be. One translocated troop, a structure that are already determined (Strum and
Pumphouse, reacted as if the elephant risk was greater Latour, 1987, p. 785). This is the definition of an
than the loss of food resources. Another translocated ostensive society. For most descriptions of society, it cer-
troop, Malaika, accepted the cost of feeding competition tainly seems that way. By contrast, in a performative so-
in exchange for increased safety. This does not make ciety (Goffman, 1971/1959; Garfinkel, 2003/1967), individ-
selection random or arbitrary. Rather, it points out an uals actively create and define society for themselves and
important constraint and obstacle: selection must oper- for others. Therefore society, rather than being a preex-
ate on a range of point responses to challenges that are isting structure, is actually constructed through the
sometimes themselves complex and independent, sug- many efforts to define it; it is something achieved in
gesting that evolutionary explanations need to incorpo- practice by all actors (Strum and Latour, 1987, p. 785).
rate, rather than eliminate, complexity, chance, and con- We argued for a shift to examine how actors create (per-
tingency. At many times in a study troops history, it was form) their society. In hindsight, we were changing focus
possible to identify probable evolutionary causes of ba- from outcome to process motivated by our data on nego-
boon responses; however, these did not operate alone. tiation in baboon and human social groups. Many fields
For example, although it is tempting to conclude that have since embraced this shift (see below).
predation pressure explained the first fusion, another in- Although we argued that the process might be similar
digenous troop in the same area, Musul, lost members to for baboons and humans, it was clear that the outcomes
predators and yet didnt try to fuse. are different. Transforming baboons into active perform-
My argument so far can be summarized as follows. ers of their society doesnt put them on a par with
Looking at the scale of baboon daily life, focusing on pro- humans. We proposed that the difference is to be found
cess rather than only on outcome, unearths layers of com- in the practical means actors have to enforce their ver-
plexity and illustrates how complexity is generated. Out- sion of society or to organize others on a larger scale.
comes are neither random nor deterministic in a way of- Baboons have only themselves, their bodies (including
ten assumed when scaling up to make evolutionary brains and minds), social skills, and whatever social
arguments. Evolutionary time speculations of cause and strategies and relationships they can construct as
effect assume rather than clarify how evolutionary princi- resources. Some factors are embodied and change slowly;
ples are situated by chance, contingency, and history. This age, sex, perhaps kinship and dominance need not be
is because they either lack the information or ignore the continually reconsidered. However, social skills are soft
importance of context and black-box the relationships tools and with them baboons can build only soft soci-
between real behaviors and evolutionary outcomes. eties. Other factors and contingencies must be continu-
My approach is not just a focus on proximate rather ally tracked and performed. Baboons become the arche-
than ultimate explanations of behavior. The baboon data typical competent member of the ethnomethodologists
challenge some of the fundamental assumptions about (Garfinkel, 2003/1967) constantly subject to the interfer-
how behavior evolves. Tinbergen (1963) highlighted four ence of others who are trying to do the same thing. They
questions that must be addressed in any comprehensive live in a complex society.
explanation of a behavior. His framework identified both Humans, by contrast, bring extrasomatic resources to
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, simplify the task of creating society and of strengthening
had to assume the feedback between proximate and ulti- the social bond. Human society becomes a complicated
mate levels while studying static behaviors. Equally latticework, which looks ostensive because so much can
important, ethologists had to examine simplified contexts 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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12 S.C. STRUM

rhesus macaques on Cayo Santiago (Sade, 1967) pro- (Seyfarth, 1977). The female hierarchy thus easily fulfills
vided an alternative structure for a monkey group the need for a structure to reduce transaction costs. As
where males leave and females stay. Sociobiology later baboons practice female philopatry, this hierarchy also
injected kinship and the gene calculus of inclusive fit- organizes the largest cohort of animals in the group. Of
ness, sexual selection, and parenting strategies to create course, there are other evolutionarily generated struc-
socioecological determinants of group configurations tures in a baboon troop including the male dominance
(Williams, 1966; Trivers, 1972; Wilson, 1975). These hierarchy, malefemale friendships, and other affiliative
strengthened the idea that female matrilines and the bonds (like those between play group members), but all
female dominance hierarchy were evolutionary adapta- of these are more dynamic, more ephemeral, and incor-
tions and the primary structure of a baboon group. porate fewer individuals than does the female hierarchy.
Arguments about hierarchy, female or otherwise, have If the female hierarchy is co-opted as primary struc-
assumed that rank will be strongly correlated to repro- ture, then it is the conservative nature of females [based
ductive success (Hausfater, 1975; Altmann et al., 1988; on reproductive constraints of female mammals (Trivers,
Cheney et al., 2006). Yet, data since the 1970s show an 1972, 1974)] that helps to keep this hierarchy relatively
inconsistent and often poor fit between baboon female stable and predictable. A male baboon who loses an en-
rank and parameters of female evolutionary success. counter to another male might continue to contest the
Instead, other factors appear influential like female outcome throughout the day or even the week. A female
sociality (Silk et al., 2003, 2010b), female age-specific fe- who loses an encounter to another female rarely disputes
cundity (Strum and Western, 1982), environment the results. This contrast results from sex differences in
(Wasser et al., 2004), or a variety of factors that seem to risks and benefits.
cancel each other out (Brown and Silk, 2002; Wasser et However, the female hierarchy isnt invariant. Ive
al., 2004; Silk et al., 2005; Crockford et al., 2008; Silk seen what happens when the female hierarchy is unsta-
and Strum, 2010). ble. This evidence supports my argument. The female hi-
The imperfect fit between female rank and reproduc- erarchy (in the different study troops) has changed a few
tive success is not surprising if attention is on what times during the last 40 years. These have been mostly
actually happens. Then evolutionary principles operate adjustments in rank between mothers and daughters,
in a context also shaped by complexity, chance, contin- disruptions but not of major concern to anyone but the
gency, and history. The surprise is, instead, that there is members of the family. By contrast, the few major
a stable female hierarchy at all despite limited evolution- changes that have occurred in the overall female hierar-
ary benefits. chy are instructive. They elicited lots of aggression some
I suggest that the female dominance hierarchy has a resulting in injuries. The aggression also disrupted basic
different purpose. Its function is primarily to create pre- daily activities like foraging and travelling. Disorder
dictability (and stability) so that transaction costs for quickly spread from a few females through the group
females and others can be reduced. The complexity of ba- embroiling individuals not initially involved. Group life
boon social life needs to be simplified. If the transaction came to a standstill for a good part of several days while
costs of grooming limit the number of grooming partners the continued instability affected individuals and the
[and stress is lower when a female baboon has fewer troop for weeks and even months. It is not hard to imag-
rather than many grooming partners (Crockford et al., ine, from these periods of disruption, why a stable and
2008; Wittig et al., 2008)], imagine the social transaction predictable structure is crucial not just for females but
costs (and stress) involved in any part of baboon daily for the rest of the troop.
routine if an animal didnt know in advance its relation- The possibility exists that there may be a better fit
ship to others. Individuals, and by analogy the troop, between rank and individual reproductive success under
would be paralyzed by minute-to-minute negotiations specific conditions (Cheney et al., 2004, 2006). For exam-
involved in the process of where to feed, rest, or travel, ple, high rankers have preferred access when it is feasi-
whom to approach, and whom to avoid. There would be ble to monopolize key foods during a resource bottleneck
no time for the basics of survival or enough time and (Barton, 1993). However, during the decades of my study,
energy to meet any new challenges. such situations turned out to be few, short, and did not
If I am correct that the female hierarchy is primarily produce the expected fit between female rank and repro-
structure, then there need not be a strong correlation duction (Strum and Western, 1982; Silk and Strum,
between female rank and individual reproductive success 2010). This was true even during critical periods such as
and no conundrum about why low ranking females stay droughts probably because the foods left then are small,
in a group. All individuals benefit from predictability dispersed, and not defensible.
and stability no matter their rank. Given the complexity of baboon reality, individuals
However, why use the female hierarchy instead of (and groups) need a way to manage daily life. It makes
some other structure to generate predictability and sta- more sense, in this context, to view the female domi-
bility? Im not suggesting that the baboons think about nance hierarchy as a transactional principle than as a
the problem and consciously create the solution. Instead, genetic characteristic of individuals or as the genetic
it is obvious that the female hierarchy is easily gener- consequences of the behavior of individuals. It is the ba-
ated from evolutionary principles such as kin selection boon version of what humans do; . . .an economizing
and reproductive value (Vx) (Williams, 1966). In baboons behavior in the sense that it greatly reduces the transac-
and other cercopithecines, a mothers intervention on tion costs of social interactions and permits efficient col-
behalf of her younger offspring in altercations with an lective action (Fukuyama, 2011). In the same way that
older offspring creates an internal family hierarchy in manners in human societies make life livable, workable,
which the youngest sibling is the highest ranking. The and usually prevent descent into anarchy, the female
normal expansion of a family over time would fashion a hierarchy creates a livable, even, civil social life for
group hierarchy where only those closest in rank are baboons. Reproductive success would not be possible
likely to be related enough for kin selection to operate otherwise; however, this is not the same as saying that a

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DARWINS MONKEY 13
females rank in the hierarchy is somehow directly benefits of not fusing. Musul targeted Nabo (itself the
connected to her reproductive success. result of the previous fusion of Malaika and Soit
The importance of structure is intuitively obvious Oitashe discussed previously). By this time, Nabo had
when complexity and process are taken seriously. There operated as a well-integrated group for more than 7
are analogous arguments about structure from other years. Unlike the earlier fusion, Musul didnt shadow
fields. For example, the value of structure in biological Nabo. Instead, Musul tried to lead Nabo to the area
systems is well summarized by Weiss and Buchanan with the best cactus.
(2009). For humans, a variety of theories argue for the Initially, most of Nabo baboons resisted probably
benefit of structure to reduce uncertainty, minimize cog- because the area was outside of Nabos traditional
nitive dissonance, build social relationships, and facili- home range and already hotly contested (baboons can
tate social exchange (for cybernetics, see Bateson, 1967; hear intertroop interactions beyond where they nor-
Watzlawick and Bevin, 1967; Harries-Jones, 1995). In mally range). As the MusulNabo fusion progressed,
fact, Festinger (1985/1957; Harmon-Jones and Harmon- Musul females exerted more and more influence on
Jones, 2007) suggested that humans seek consistency Nabo through their growing network of friendships
and try to reduce dissonance in any new situation. with Nabo males. Occasionally, the combined group
Recent approaches to human cognition including distrib- made it to the cactus area only to be aggressively
uted cognition (Hutchins, 1995; Forster and Rodriguez, expelled by one resident baboon group in particular.
2006; Barnard, 2010) and situated action (Suchman, Aggressive mobbing is usually a very effective tactic to
1987; Rogoff, 1990, 2003; Lave and Wegner, 1991) rely displace another group from a location. In this case,
on a nontraditional idea of structure that is similar to while the intruders fled from the aggression, they kept
my argument. There may be indirect or downstream coming back. MusulNabo endured repeated evictions
impacts of structure on female baboon reproduction, but over more than a year but also developed evasive tac-
I suggest that the value of structure is not directly tics. For example, the invaders would leave the sleeping
reproductive success. site at first light to avoid being mobbed. This early de-
parture meant giving up resting and socializing time
Managing complexity: Extinction, tolerance, and even some feeding time clearly a disruption of their
slippage, and the evolution of the good enough normal daily routine. However, leaving early didnt
(20072011) always work. Often the resident troop would track
down the invaders and push them until they were
An underlying assumption of evolutionary arguments away from the disputed area.
is that there are winners and losers (Weiss and Bu- The MusulNabo fusion (the group was now called
chanan, 2009; Watts, 2010). Yet, in the vast span of life Namu) violated many rules of baboon life that I had
on earth, more than 99% of all species have gone extinct documented during the preceding decades. I considered
(Raup, 1986; Wilson, 2002; Barnosky et al., 2011). As ev- whether it might represent a baboon mistake. The
olutionary history is continuous and the end point in ev- quantitative data suggested that it was. No baboon was
olutionary time is most probably extinction, calculating actually killed because of the confrontations between
costs and benefits has to be sensitive to that history. groups, but during the first 18 months of the range shift,
Species who flourished at one point later disappeared. most of Nabos and Musuls babies did not survive the
This is relevant to my argument in two ways. First, a first year of life. Adult mortality also increased.
particular point in time may or may not be representa- Autopsies of adult females showed that previously nonfa-
tive of what has happened or what will happen. I recog- tal conditions had turned lethal. Nutritional analysis of
nized this about the baboons Im watching and it will the cactus fruits indicated that its value, and thus its
apply equally to other species. The power of evolutionary appeal, might be the energy bonus provided by such a
arguments will therefore depend on capturing as much large package of calories [see also crop raiding (Strum,
of the (natural) history as possible. Second, the adapta- 2010)]. Yet, the troop must have used up that bonus and
tionist paradigm (Gould and Lewontin, 1979; Laland and more evading attack. Unlike during crop raiding, body
Brown, 2011/2002) often assumes that animals in nature condition didnt improve.
are finely, even perfectly tuned to their environment Do baboons make mistakes? Was I observing the pro-
and, at least until recently, that what we observe are the cess of extinction or at least extirpation, one death at a
elements of this excellent fit. Yet, focusing on real lives time? My answer is different now, 4 years later, than it
in real time illustrates how difficult it is to know about was at the start of the fusion. Currently the troops have
the consequences of particular actions or to decide been fused (becoming Namu) for several years. Namu, as
whether, for example, a baboon individual or group invaders, eventually created a home range in the cactus
(since individuals dont exist outside of groups) is mak- area despite all the aggression. Although Namu is still
ing a mistake. subordinate to the main resident troop, they arent
I had the opportunity to consider this issue during a mobbed as often. Namu female reproductive parameters
second fusion event. This time the impetus appeared to are steadily improving along with female physical condi-
be food, not predators. The little group, Musul, that tion. Age at menarche is earlier, and interbirth intervals
hadnt fused 7 years earlier, initiated a fusion for a dif- are getting shorter. Namus overall mortality has
ferent reason, to gain access to an important new food declined while survivorship is improving. I had to con-
resource or so it seemed. This resource was dominated clude, against my previous assessment, that Namus per-
at the time by other, larger, baboon groups (who were sistence paid off.
not part of my study). The second fusion provides more evidence about the
The resource they sought was the fruit of Opuntia pitfalls of making evolutionary interpretations of behav-
stricta, an exotic invader which had only just appeared ior because these depend on history and time frame.
in Musuls home range. Ironically, before this fusion Flexibility and adaptability are more vital than I had
Musul had been my control as a test of the costs and 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 routes because ecological niches were a constantly mov-
evolutionary criteria like condition, reproduction, and ing target. They proposed that speciation might itself
survival are available as was evident in the Namu range be the outcome of unusual single events (emphasized by
shift. Too often, the link between the behavior and its ev- the author) and that to understand so-called adaptive
olutionary consequences passes through a black box radiations, it is necessary to study the multiple events in
where much of the work and most of the assumptions re- a species life that provide it with the opportunity to
side [see similar argument for the interpretation of the adapt, rather than studying wide and general processes
link between social complexity and the evolution of cog- (Venditti et al., 2011, p. 393).
nition (Strum et al., 1997)]. Lorenzen et al. (2011) offered another example. North
The MusulNabo fusion also clarifies that evolutionary American Pleistocene megafuana extinctions appear
processes may be more tolerant, have larger acceptabil- much more complex than previously assumed. Their
ity spaces (Weiss and Buchanan, 2009), and allow for analysis demonstrates that changes in the abundance of
more variance than most evolutionary time arguments megafauna were idiosyncratic because each species
assume. What these baboons did for more than 4 years (and even continental populations within species)
was not a perfect fit to the layers of complexity in their appeared to respond differently to the effects of climate
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 ment (Lorenzen et al., 2011, p. 364). Surprisingly, the
to answer that question and require measures of success species that went extinct did not have characteristics
situated in their natural history. that set them apart from those that survived. This
Since complexity generates a variety of options, it is implies a complex and contingent process (with behav-
not surprising that individuals in a baboon troop dis- ioral flexibility in some species) rather than a simple
agree about what to do. These disagreements must be and deterministic one.
resolved because the group is constrained to move and A recent study on the evolution of sociality in primates
act as a unit. Resolution requires negotiation. Therefore, provides another parallel. Shultz et al. (2011) modeled
managing the consequences of socioecological complexity the evolution of primate social grouping patterns using
is a serious daily challenge for baboons. This might be genetic and social information about 217 primate spe-
most apparent during the special circumstances I was cies. The conclusion contradicts previous speculations
able to document like the incursion of agriculture, peri- about the evolution of primate social groupings, which
ods of heavy predation, or the appearance of a new food. assumed that the original groups were small and simple
However, disagreements, behavioral discussions, and that larger more complex structures developed from
happen during negotiations about the more ordinary these groupings. The analysis indicates that the initial
parts of baboon life such as during troop movements, change in social grouping was probably caused by a shift
consorts, when a new infant is born, and the like. 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
Coming to grips with complexity has some phylogenetic inertia. In this view, the evolution
These are more than just interesting stories. They are of primate sociality was neither simple nor gradual and
evidence about how evolution actually works. The focus likely involved multifaceted interactions of group size,
on process and history reveals the importance of complex- social complexity, and cooperative sociality.
ity, chance, and contingency in the real lives of baboons. The argument for, and the value of, focusing on pro-
It also highlights the flexibility and adaptability (in cess and description is best illustrated by modern macro-
response to complexity) that pervade baboon daily life. ecology (Smith et al., 2011). Macroecologists look at com-
Other disciplines like paleontology, ecology, genetics, plex interactions embedded in space and time by exam-
and cellular biology have recently made similar argu- ining multiple factors and levels including morphology,
ments. The broad integrative approach of Weiss and Bu- physiology, behavior, ecology, phenology, and phylogeny.
chanan (2009; Weiss et al., 2011) documents the impor- Taking their cue from past natural historians and bio-
tance of chance, contingency, and history (and emer- geographers, macroecology uses natural history descrip-
gence) in the history of life. Their EcoDevoEvo tion as a critical scientific tool. Evolutionary change,
perspective advocates going from evolutionary to devel- they argue, is best understood in terms of complexity,
opmental and ecological time scales [see Fig. 2.2 in Weiss history, chance, and contingency (and emergence)
and Buchanan (2009, p. 12) for an instructive diagram], whether looking at the past or trying to predict the
and illustrates why chance is always built into concep- future.
tion, inheritance, birth, death, and much that happens Even studies of modern human behavior that usually
in between (Weiss and Buchanan, 2009, p. 197). Context begin with the assumption of complexity and history are
and history, for Weiss and Buchanan, are essential to moving further in that direction. For example, a recent
understanding how, for example, a gene network, cell, study of the effectiveness of co-management of global
organism, or ecosystem gets from point a to point b. fisheries identified 19 factors (summed into eight binary
Viewing evolution at these scales suggests that there are measures and their interactions) that were vital to suc-
multiple routes to success with more tolerance and slip- cess (Gutierrez et al., 2011). However, history and acci-
page and therefore more flexibility than modern Darwin- dent (not planned design) determined whether a fishery
ian interpretations often allow. had any of the set of these critical features. Similarly,
A recent example in paleontology is the work of Ven- Black et al. (2011) explained the inadequacy of previous
ditti et al. (2011) on body size in mammals. They con- models of human migration that rely only on simple
tend that in the evolution of mammalian body sizes, principles of causation. Instead, they argued that there
diversification did not speed up and then slow down are numerous drivers of migration that include social,
when ecological niches were invaded as previously economic, demographic, and political factors. Further-
assumed. Instead, body size diversification took multiple more, each driver interacts with the others as well as

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DARWINS MONKEY 15
each being influenced by environmental change. The my argument about the challenge of managing complex-
result is that they can make better predictions about ity and offered an alternative meaning for the female
who migrates, when, and why than previous attempts. dominance hierarchy.
Another recent example for humans is Fukuyamas As with all complex systems, the whole is greater
work on the origins of human political order (Fukuyama, than the sum of its parts (Johnson, 2011/2007; Mitchell,
2011). He convincingly demonstrated how similar starting 2009) even though we still lack the methodology to cap-
conditions result in radically different outcomes and that ture behavioral emergence. However, baboon natural
The factors driving the development of any given politi- histories help. They are a resource for understanding
cal institution are multiple, complex, and often dependent how complexity is generated. They show how solutions
on accidental or contingent events. Any causal factors one to challenges develop. They illustrate the enormity of
adduces for a given development are themselves caused the task that baboons face in managing complexity.
by prior conditions that extend backward in time in an This type of evidence also underscores the role that
endless regression (Fukuyama, 2011, p. 23). context, chance, and contingency play in producing spe-
Of course, Complexity Theory as it has developed over cific outcomes on the evolutionary stage. The baboon
the last three decades draws some of the same conclu- data also raise the possibility that there are multiple
sions (Waldrop, 1992; Thelen and Smith, 1994; Capra, ways for individuals to succeed, not just one optimal
1997; Oyama et al., 2001; Ward, 2002; Johnson, 2011/ evolutionary path.
2007). Complex systems models, including complex
adaptive systems and dynamical systems, have recently PART 3: WHY SHOULD ANTHROPOLOGISTS
been applied to primates using social network analysis TODAY BE INTERESTED IN BABOONS?
and analysis of collective behavior (King and Sueur,
2011a; Sueur and Deneubourg, 2011; Sueur et al., Washburn and DeVore (1961, 1963) used baboons spe-
2011a). Some of this work formally addresses the issues cifically to frame questions about human evolution.
of nonlinearity and of history, chance, and contingency Ironically, the intuitive power of DeVore and Washburns
in producing specific outcomes. Most work on complex model did a disservice to their goals as the simplified
systems still falls short of addressing the context of version was applied to other primates and even to
behavior in the real world. For primate social networks, humans (Strum et al., 1999; Strum and Fedigan, 2000b).
for example, this should include multiple interacting Nearly four decades later, I once again suggest that
contexts that set up the starting conditions and continue baboons may be relevant in highlighting the challenges
to influence the operation of the system. and solutions of early human evolution.
I began thinking about complexity in the early 1970s My first decade of baboon observations led me to
when I realized that male success rested on more than declare them almost human (Strum, 2001/1987b).
the male dominance hierarchy. Age, residency, context, Indeed, they exhibited remarkable social skills, social
and social relationships based on collaboration and be- strategies, and even sexual politics (for a current defini-
havioral exchange all played a role. This view of social tion of and reservations about using the word politics,
complexity was my first step in expanding out from the see Watts, 2010), we normally reserve for humans. How-
powerful simplifying assumptions of the previous ba- ever, the following decades of baboon quantitative data
boon models. Crop raiding (19761984) demonstrated and detailed natural histories suggested to me why, de-
the significance of ecology and how the interaction of spite having much in common in the process of building
social and ecological added to the complexity of baboon society, baboons and humans have diverged so much.
lives. The disagreement about whether to raid high- This renders the baboon vantage point particularly use-
lighted the way that social negotiations influenced ful both for understanding what makes us human and
choices about the basics of daily life. The translocation for refining concepts of how evolutionary processes
(1984) was both a provocative experiment and an un- actually work at the level of behavior.
precedented opportunity to see baboon adaptability in
action. The value of the social group was never as Summary of baboon principles of the social
obvious as after translocation; however, I also learned
that baboons had multiple ways to succeed and that 1. The social group is a major organ of adaptation. The
specific events and accidents of history set different group offers social resources useful in meeting new
groups on unique paths. challenges (raiding, translocation, etc.) and in solv-
The stone of baboon complexity that I had slowly ing daily problems (translocation, Opuntia, etc.).
pushed up the hill in the first decades picked up incredi- Baboons illustrate how this works not just why it
ble speed rolling down the other side in the last decade. should work this way.
The first fusion (19992001) showed the full extent of 2. Social negotiation is critical to baboon life but it
social negotiation about the nature of the social group, quickly reaches the limits of available time and
something scientists assume to be a given fact. The 2 comprehension. Baboons are also constrained by
years of negotiation embedded in specific social and eco- their inability to hold nonmaterial artifacts (rela-
logical events made the fusion understandable but not tionships, structure, etc.) over time and space.
predictable based on simple evolutionary rules. The sec- 3. Baboon society is restricted in scale because it has
ond fusion (20052008) certainly looked like a monumen- few means to simplify social negotiations. Therefore,
tal evolutionary mistake but turned out not to be. That baboons cannot escape social complexity to build a
fusion demonstrated the constantly changing balance of complicated social structure.
costs and benefits and the large list of factors that had 4. Context and history matter to baboons. This
to be considered. It also implied that evolution was more includes social actions at the individual and group
tolerant and allowed more slippage making simple evolu- levels.
tionary calculations problematic. Finally, the changes in 5. Collaboration (see below), not just competition, is a
the female dominance hierarchy (20082011) bolstered 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


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

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DARWINS MONKEY 17
(Dictionary, 1971) defines cooperation as to work to- other long-term baboon research sites also provide evi-
gether, a meaning that originated in 1616 (p. 963) dence of social collaboration even if the investigators do
which by 1830 also meant the practice of economic coop- not label the behaviors that way (e.g., Palombit et al.,
eration. For humans, it is used interchangeably with the 1997; Silk et al., 2006, 2010a; Swedell and Leigh, 2006).
term collaboration which first appeared in 1871. There This means that baboons regularly engage in collective
is no scientific consensus about the definition of the term action that goes beyond the coordination of individuals.
cooperation as it applies to animals and humans However, I propose that baboons have constraints. With-
(Richerson and Boyd, 2005; Kappeler and van Schaik, out additional resources, baboons cant move from collab-
2006; van Schaik et al., 2006; Tomasello, 2009; Weiss oration to cooperation just as they are limited in the
and Buchanan, 2009; West et al., 2010). Definitions have type of society they can build.
ranged from complex and restrictive economic criteria On the other hand, would baboons benefit from being
with analysis based on game theory (e.g., see Noe et al., able to cooperate? The origins of cooperation in humans
2001), to more real-life criteria like joint commitment to is often attributed to a change in lifestyle, particularly
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 comitant division of labor and necessity for sharing (Lee
possible meaning of cooperation: components working and DeVore, 1968; Bird, 1999; Wilson, 2012). Without
together successfully by some criterion, including that of anything in their diet that could be shared, except the
evolutionary viability (Weiss and Buchanan, 2009, p. infrequent prey carcass (Strum, 1975b), baboons may
38). One cross-cutting claim, however, is that human not have the necessary or sufficient conditions for the
cooperation is distinctly different from what other ani- evolution of cooperation. In the end, although baboons
mals or even other primates do. are great collaborators, they have neither the means nor
For the purpose of situating baboons, I find it useful to the reasons to cooperate.
think in terms of a continuum that starts with the ge-
ometry of the selfish herd. Here, individuals aggregate Culture as an organizing principle. The performative
and cohere by simple fact of trying to get away from view of society, augmented by the frameworks of situated
something, for instance, a predator (Hamilton, 1971). I action and distributed cognition, offers a novel perspec-
suggest that what comes next is called coordination, tive on tradition and culture among baboons. Baboons
which develops as the enhanced value of group living dont use or make material tools in the wild but they do
requires solutions to behaving as a collective in everyday skillfully craft friendships. These social tools are non-
life [see also the collective action problem (Nunn, 2000; material artifacts (Strum and Forster, 2001) because
Noe et al., 2001; Sussman and Garber, 2011)]. Collabo- making them requires the same cognitive achievement
ration is still further in the continuum from the selfish as making material artifacts: their shape and use isnt
herd. Although cooperation and collaboration are used intuitively obvious on the surface of the material. How-
interchangeably in both the common and scientific lexi- ever, here the similarity stops. Social tools, unlike mate-
cons, I make a distinction based on insights from rial tools, have difficulty expanding across space and
baboons. In 1973, one baboon male started to hunt time. They are soft tools and difficult to stabilize. Still,
rather than opportunistically collect gazelle prey. His since tool use is often a criterion for culture (see Fra-
actions were purposeful and his intent was evident in gaszy and Perry, 2003), baboons do have some of the ba-
the sequence of his behaviors. Then other males joined, sic skills (Strum and Latour, 1987).
initially alerted by his return to the troop covered in dry Another criterion used to identify a primate cultural
blood. Through a set of events, the males actually tradition is that the behavior should be invariant
learned that collaborating in the direction of the chase within a group but variant across groups (Tomasello,
paid dividends. Later, they practiced what they learned 1999; Rendell and Whitehead, 2001; Fragaszy and Perry,
(Strum, 1975b). I use the term collaboration in its orig- 2003; Richerson and Boyd, 2005). A number of baboon
inal meaning, to work in conjunction with others, and behaviors or behavioral systems may qualify. For exam-
not as a synonym for cooperation. Collaborating baboons ple, a hunting tradition developed in only one group
do not have a predetermined common goal. Rather they (Strum, 1975b, 1976). The raiding lifestyle, the selection
are a collection of individuals with similar goals; in the of sleeping sites after translocation, differences in diets
case of predation, each wanted meat and realized the ef- between adjacent groups in the same location, and even
ficacy of chasing prey one way rather than another. variations in aspects of the friendship configuration
Finally, in this scheme, cooperation is a special type of might be more controversial suggestions as baboon tradi-
collective action in which participants have a preexisting tions. I could go further and claim that every groups
common purpose (see also references above). Chimpan- female dominance hierarchy is a traditional system. Af-
zee predatory hunts (Boesch and Boesch-Achermann, ter all, each is unique to a group, carries forward across
2000) appear a rudimentary form of primate cooperation, generations, and is relatively stable over time.
particularly by comparison with collaborative baboon What sets baboons apart from humans, if we grant
hunting. The continuum ends with human cooperation, that these are social traditions, is the next step in the
which entails a variety of additional aspects not found in cultural process: the persistence and accumulation of
other species (but see Wilson, 2012), although the specif- innovations (Tomasello, 1999; Richerson and Boyd,
ics depend on whose definition is used. 2005). I have argued that for baboons, the means of ac-
Although competition is not as central to the baboons, quisition and transmission of a tradition depends on
I have studied, as previously assumed, neither can frequent exposure and visible actions. Thus, the hunt-
baboons cooperate. That they are expert social collabora- ing tradition disappeared when the innovator trans-
tors is visible in their social strategies of competition ferred out of the troop a year later. Why? Although the
and defense, in troop movements, intertroop encounters, rate of baboon predatory behavior in Pumphouse was
home-range shifts, changes in diet, as well as in intricate high when compared with other baboon and chimpan-
multi-individual social and sexual encounters. Data from 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 Finally, I conclude that the four decades of baboon
away from the troop so that most of the animals didnt research and the framework outlined above is helpful in
see or participate in the actual behavior. As a result, resetting the starting point for the human experiment.
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- phisticated or embedded in less complexity than these
sion, the entire raider troop, not just a few animals, baboons. Moreover, lowly baboons also serve as a re-
raided. This frequency and visibility, I propose, were minder that the group is a primary organ of adaptation
important contributors to the persistence of raiding for most primates, that collaboration/cooperation is likely
behavior. Raiding even accumulated some modifications as crucial as competition in daily life, and that social
as raiders learned to avoid the new anthropogenic risks intelligence, social skills, and social management were
and human-induced mortality declined over time at a premium long before humans or even the great
(Strum, 2010). apes.
Conversely, if culture is based, in part, on internalized
rules for behavior (DAndrade, 1981, 1990; Byrne and Summary of baboon principles of evolution
Russon, 1998), the same factors that may be important
facilitators of traditions for baboons (visibility and 1. Baboons demonstrate the value of examining process
involvement) could mitigate against the need for tradi- rather than just outcome and the pitfalls of uncriti-
tions. Baboons have constant access to each other as cally collapsing behavior into evolutionary time. This
social information resources (Strum, 1983a, 2010; King, is a critical methodological move. How evolutionary
1994; King and Cowlishaw, 2007; King and Sueur, principles work in real time and real lives can only be
2011b; King et al., in press). They can rely on distributed identified this way.
cognition and situated action rather than on individually 2. History, chance, and contingency are crucial for
internalized operating principles (Forster and Strum, understanding baboons embedded in social, ecological,
1994; Strum et al., 1997; Forster, 2002; Forster and and socioecological interactions. The extensive baboon
Rodriguez, 2006). Recently, Roediger and McDermott negotiations imply that there are multiple options not
(2011) made a similar suggestion for humans. They just one best evolutionary solution. In fact, the
argued that If one individual in a group forgets critical baboons add support to the assertion that behavioral
information (about food resources or dangers, for adaptability is a ubiquitous trait of life but that it is
instance), then it is wise to get an updated memory from often missed because of the existing scientific empha-
another group member (Roediger and McDermott, 2011, sis on evolutionary time and on evolutionary out-
p. 48). This perspective could help explain why the two comes (Weiss and Buchanan, 2009, p. 231).
most cultural primates so far are chimpanzees (Whiten 3. Thinking in terms of contingency and (natural) history
et al., 1999, 2007) and orangutans (van Schaik et al., helps to make sense of complexity. It is also easier to
2003; Jaeggi et al., 2010). If a baboon forgets how to do understand complexity by looking at its unfolding, how
something, it can just look around. Chimpanzees and it is generated, than by puzzling about existing com-
orangutans dont have that luxury. Chimpanzee fission plex structures (Weiss and Buchanan, 2009, p. 176).
fusion society (Chapman et al., 1995; Lehmann and
Boesch, 2004) and the nearly solitary existence of orang- Given these three principles, the baboon data that I
utans (van Schaik, 1999) deprive them of each other as have just presented dont fit the standard view of Dar-
social resources. This should increase the value of tradi- winian natural selection (Laland and Brown, 2011/2002).
tions and possibly of culture. By analogy, if early Instead, they correspond to Weiss and Buchanans alter-
humans lived in fissionfusion societies (Grove et al., in native evolutionary framework. Weiss and Buchanan
press), they would face challenges of remembering argued that chance (action without direction that has
more like chimpanzees than like baboons. major impacts; p. 36) is one of lifes important regular-
In summary, I am proposing that baboons have a soci- ities (p. 40), that life is a contingent phenomenon (p.
ety at the limits of social complexity and that they lack 35), and that adaptability in the face of changing cir-
the resources needed either to manage more complexity cumstances means that responses are flexible and not
or to simplify negotiations and build a complicated large- always prescribed in advance (p. 37). Predictability is
scale society. Baboons have sophisticated cognition that the basis of life processes (p. 21), and history and context
tracks social, ecological, and socioecological interactions. matter because these are the contingent conditions
It is likely that they appropriate solutions to new prob- upon which the future is built (p. 40). Weiss and Bu-
lems from the social context. If so, baboons may need chanan demonstrates how this applies at the cellular,
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 Buchanan, 2009; Weiss et al., 2011). I suggest that it
social traditions (culture?); however, baboons have diffi- also applies to baboon behavior.
culty with the persistence and accumulation of innova-
tions because, I suggest, for baboons this depends on fre- PART 4: CAPTURING AND EXPANDING
quent exposure and actions that are visible to everyone. COMPLEXITY IN THE ANTHROPOCENE
Although baboons are almost human in their negotia-
tion and politics, baboons cant become human because Anthropologists should be interested in baboons
they lack the ability to build a complicated society from because of what they tell us about the reality of daily
a complex one. To do this, they would need the behaviors lives, about the challenge of complexity, and about how
that are usually classified as hallmarks of humanness, evolutionary principles get embedded in contexts deter-
minimally the new resources that material culture, sym- mined by history, chance, and contingency. Baboons add
bols, language-related abilities provide, and a new sub- evidence to the suggestion that evolutionary processes
sistence base that both permits and requires division of contain more tolerance, multiple solutions, larger accept-
labor and sharing. ability spaces, and the possibility that an adaptive fit

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DARWINS MONKEY 19
will be good enough rather seamless. When the This article wouldnt be complete without a mention of
straight line of evolutionary scale opens up to capture what Darwins thought about baboons. His summary is
what actually happens, arguments become less reduc- particularly apt: For my own part I would as soon be
tionist and less deterministic [the pentagon (Weiss and descended from that heroic little monkey, who braved
Buchanan, 2009, p. 9)]. his dreaded enemy in order to save the life of his keeper,
Yet studying behavioral complexity in the wild poses or from that old baboon, who descending from the moun-
methodological challenges. How do we document com- tains, carried away in triumph his young comrade from
plexity and track the impact of chance, contingency, and a crowd of astonished dogsas from a savage. . . and I
context? I see no other way than by returning to compar- have given the evidence to the best of my ability
ative natural history as a scientific method (Dayton, (Darwin, 1871/2004, p. 791). I dont think Darwin would
2003; Smith et al., 2011). Although the plural of anec- be surprised by the picture of baboons that I have just
dote may not be data [but see recent shift in medicine presented.
(Aronson, 2003)], the plural of comparative natural his-
tory certainly is. I call baboons, Darwins monkey, not ACKNOWLEDGMENTS
just because he made frequent reference to them
(Darwin, 1871/2004) but because baboon lives once again I thank the editor for his generous invitation to com-
argue for the value of comparative natural history, the ment on the important lessons from my years of baboon
key method that Darwin used. Following Darwins lead, research. My insights also rest on the work done by
natural history observations should include well-docu- many other individuals associated first with the Gilgil
mented details tracked across time, space, groups, land- Baboon Project and later with the Uaso Ngiro Baboon
scapes, and species. They should be aided, when possi- Project. Since 1981, Kenyan paraethologists have been
ble, by quantitative data and experiment. Long-term the mainstay of the basic data collection. I thank them
studies become extremely vital because only they can all for their contributions. Finally, I thank David West-
generate natural histories and the kind of insights pre- ern for his distributed cognition; Katerina Semendeferi
sented here. for useful comments on the article; and Bruno Latour
Baboons are also useful to anthropologists for reasons who first helped me to see why complexity is so impor-
that Darwin could not have guessed. Today, it is common tant although it took many decades to collect the evi-
practice to end an article about primates with a com- dence.
ment about conservation. The baboons offer a singular
perspective on primate conservation because they illus-
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