Primate Sociality and Social Systems

Why be social? And, why not be? What are the costs and benefits of sociality, and what types of sociality
characterize nonhuman primates?

Why Be Social?

Many primates and other animals live in social groups. In social groups, individual members coordinate
their activities, communicate with one another, and interact in both affiliative (friendly) and agonistic
(aggressive or submissive) ways. In many social groups, individuals are gregarious; that is, they interact
with one another frequently, engage in a variety of types of social interactions, and typically form and
maintain social bonds (strong social relationships) with other individuals (Dunbar 1988, Silk 2007). These
bonds are often expressed in the form of grooming, a common social activity among primates in which
one individual carefully picks through the fur of another and removes any debris or ectoparasites (Figure
1). Grooming serves an important social function for most primates in addition to its hygienic function
(Henzi & Barrett 1999).

Two female geladas (<i>Theropithecus gelada</i>) grooming one another.

Figure 1: Two female geladas (Theropithecus gelada) grooming one another.

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Group life carries with it inevitable conflict and competition. Individuals must share food resources,
water resources, sleeping sites, and mates. While usually mediated by dominance hierarchies in which
higher-ranking individuals have priority of access to limited resources, aggressive competition over food
and mates is common in primates, and is not only energetically costly but can lead to injury and even
death (Mason & Mendoza 1993). In addition to energy costs and risk of injury, high levels of aggression-
given or received-can lead to chronic psychological stress. Chronic stress can adversely affect health and
reproduction in numerous ways (Sapolsky 2002). Other socially-induced events may also increase stress
levels: in chacma baboons, for example, infanticidal (infant killing) behavior by males-or even simply the
immigration of a potentially infanticidal male into a social group-increases stress hormone levels in
females with infants (Engh et al. 2006). In addition to these social costs of group living, close social
contact also increases the potential transmission of pathogens, which increases each individual's risk of
contracting infectious diseases. Finally, groups may be disadvantageous because they are more easily
detectable by predators compared to solitary individuals.
Given all of these potential costs of group living, why do so many primates-and other mammals-bother
living in groups at all? One might especially wonder about this for the lowest-ranking individuals in a
social group (i.e., those who are last in line for the available resources).

Despite the costs, sociality is important to these animals for several reasons. Probably most importantly,
living in a group likely decreases one's risk of falling victim to predation (van Schaik 1983). There are
three reasons for this. First, in social groups there are more individuals looking out for predators and
thus predators will be detected more quickly. Second, living in a group decreases each individual's
chance of being preyed upon due to an effect called "geometry for the selfish herd" (Hamilton 1971):
this states that the larger the group (e.g., 100 versus 10), the lower each individual's chance (1/100
versus 1/10) of becoming prey. Third, individuals in groups can collectively mob predators and
successfully drive them away, whereas lone individuals cannot.

Sociality also benefits animals via access to food and other resources. In groups, there are many
individuals looking for food simultaneously and thus detection of good food resources (e.g., ripe fruit;
Figure 2) will inevitably be communicated to others simply because group members are usually in close
proximity to one another. Group members will also benefit from cooperation over defense of food-or
other limited resources such as water holes and sleeping sites-as groups can out-compete individuals,
and larger groups can out-compete smaller groups (Wrangham 1980).

A moor macaque (<i>Macaca maurus</i>) feeding on ripe fruit in Sulawesi.

Figure 2: A moor macaque (Macaca maurus) feeding on ripe fruit in Sulawesi.

© 2012 Nature Education Courtesy of L. Swedell. All rights reserved. View Terms of Use

In addition, sociality is beneficial to group-living animals in that it makes it easier for them to find mates.
Animals that do not live in groups must either search for mates or opportunistically mate when they
encounter other individuals. Group-living animals simply choose mates within their social group.

Moreover, sociality allows cooperative socialization of offspring. In social groups, infants and juveniles
play with one another, which develops motor skills as well as the social skills necessary to survive and
reproduce in a social setting. For example, during social play juveniles receive reinforcement from adults
about how dominance hierarchies work, and what it means to have a given rank within a social group of
a given species. Thus when they reach adulthood they will have learned what they can and cannot do,
and thereby fit into the social fabric of their social group (Pereira & Fairbanks 1993).
Finally, sociality in and of itself appears to carry benefits for individuals. As noted above, one of the costs
of group living is the potentially high level of conflict and aggression that occurs among group members,
which involves greater energy expenditure, risk of injury, and chronic stress. In baboons, this socially-
induced stress appears to be alleviated by the receipt of affiliative vocalizations, as well as the
maintenance of grooming relationships with a small network of close associates (Crockford et al. 2008).
Ultimately, female baboons with strong social bonds (i.e., social relationships characterized by frequent
proximity and grooming; Figure 3) experience greater offspring survival and even longer lifespans than
females with weaker bonds (Silk et al. 2003, 2009, 2010). These studies demonstrate that strong social
relationships within groups, beyond group living alone, can carry important fitness benefits for
individuals.

Olive baboons (<i>Papio anubis</i>) grooming in Gombe Stream National Park, Tanzania.

Figure 3: Olive baboons (Papio anubis) grooming in Gombe Stream National Park, Tanzania.

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Primate Social Systems

The social system of a given species is an outcome of (1) its social structure, the size and composition of
a typical group of that species, and (2) its social organization, how those individuals are organized (i.e.,
the patterns of spacing, agonistic and affiliative social interactions, philopatry [whether one or both
sexes remain in their natal group], and dispersal [whether one or both sexes move to a new group to
reproduce]) that typify that species. Each species also tends to have a characteristic mating system (i.e.,
the pattern of mating among members of each sex). All of these aspects of primate societies vary widely
across the primate order.

The least gregarious primates have what is often referred to as a solitary dispersed social system. In
these primates, an adult male's territory overlaps the territory of one or more adult females, but each
individual forages alone and maintains social contact mainly through vocal and/or olfactory
communication. These primates are typically nocturnal, foraging at night and sleeping in trees during
the day. The mating system in these primates is usually polygynous (i.e., each male mates with multiple
females). This type of social system characterizes galagos, lorises, some lemurs, some tarsiers, and
orangutans. Notably, orangutans are the only anthropoid primates with a solitary social system.
Titi monkeys, owl monkeys, some callitrichids (marmosets and tamarins), and many hylobatids (gibbons
and siamangs) are characterized by a pair-bonded social system. Here, one adult male and one adult
female form a small social group and defend a territory from other pairs. The mating system in these
groups is usually monogamous (only one male mates with only one female), though extra-pair
copulations have been observed (Palombit 1994), and the male usually participates in offspring care,
which is unusual for male mammals (Fuentes 2002).

Many marmosets and tamarins live in one-female multi-male groups characterized by cooperative
breeding. In this type of system, usually only one female breeds, and that female suppresses the
reproduction of any subordinate females via aggression and/or pheromonal (olfactory) signals. Usually
there is more than one breeding male, thus the mating system is polyandrous, a rarity among mammals.
Some or all of the individuals in these groups participate in offspring care and this social system is thus
often called cooperative polyandry.

One of the most common primate social systems is the one-male group, which characterizes most
colobine monkeys, most guenons, patas monkeys, howler monkeys, and some gorillas. Here, a single
resident adult male defends a group of (usually) philopatric, related females from other males and,
while his tenure lasts, enjoys exclusive mating access to those females (i.e., polygyny). Sometimes called
harems, these groups are always at risk of takeover by non-resident males, who typically form all-male
groups while awaiting their chance to become a resident male. Often, takeovers are accompanied by
infanticide, in which the new resident male kills the young infants in the group. This behavior has the
effect of bringing the mothers back into estrus (sexual receptivity) sooner than they would have
otherwise.

Also common among primates are multi-male multi-female groups, in which multiple individuals of each
sex form large social groups in which the mating system is usually polygynandrous (i.e., both males and
females are polygamous in that they mate with multiple members of the opposite sex). These are the
largest groups of primates, and usually quite complex socially, with differentiated social and kin
relationships among group members. This type of social system characterizes many monkeys, including
macaques, most baboons, vervet monkeys, mangabeys, capuchins, squirrel monkeys, woolly monkeys,
and some colobine monkeys, as well as some lemurs-most notably the ringtailed lemur and sifaka. In
most of these species, females are philopatric and males disperse.

Similar to multi-male multi-female groups are the fission-fusion communities of chimpanzees, bonobos,
spider monkeys, and some other ateline monkeys. Fission-fusion communities are less cohesive than
typical multi-male multi-female groups. These groups occupy very large home ranges in which
temporary foraging parties cleave and coalesce over time with changes in resource availability and
female reproductive condition. These social systems are typically characterized by female dispersal and
male philopatry.

The most complex type of social system found in primates, and in mammals as whole, is the multi-level
society (also known as a hierarchical or modular society) characterizing hamadryas baboons (Figure 4),
geladas, snub-nosed monkeys, and a few other mammals such as elephants. In this type of system, there
are at least three levels of social structure: the one-male unit (OMU), the band, and the troop or herd.
The OMU is the reproductive unit and consists of one "leader" male, sometimes a follower male, and
several females; the band is the ecological unit that forages and sleeps together, and the troop or herd
is a temporary aggregation at a sleeping site or foraging area. In hamadryas baboons, there is a fourth
layer between the OMU and the band, the clan, which consists of OMUs and bachelor males linked by
social bonds and possibly kinship among males. In geladas, bachelor males join together to form all-male
groups. Reproduction in these societies is usually polygynous, and OMUs are always at risk of takeover
by bachelor males, who may commit infanticide after taking over females with young infants.

A hamadryas baboon (<i>Papio hamadryas</i>) one-male unit (OMU) at Filoha, Ethiopia.

Figure 3: A hamadryas baboon (Papio hamadryas) one-male unit (OMU) at Filoha, Ethiopia.

© 2012 Nature Education Courtesy M. Pines All rights reserved. View Terms of Use

It is important to note that the social organization of a species might not be immediately apparent from
its social structure. For example, both geladas (Figure 1) and hamadryas baboons (Figure 4) are
characterized by multiple layers of society, and their social structure is almost identical. However, the
social organization of these two species could not be more different (Kummer 1967, Dunbar 1983). In
geladas, cohesion of OMUs is maintained by philopatric females, as each OMU is a female kin group and
these kin groups are taken over as entire units by males. In hamadryas, by contrast, cohesion of OMUs is
maintained by philopatric males who take over females one by one, often exchanging them with other
males in their clans (Pines et al. 2011), and aggressively condition those females to remain in their OMU.
Thus, hamadryas social organization includes strong male-female bonds (the glue that holds together an
OMU) and strong male-male bonds (as males are philopatric and thus related to one another within
bands and clans), whereas gelada social organization is characterized by strong female-female bonds
(which hold together OMUs) but weak bonds between the sexes.
Moreover, one must be careful not to make assumptions about a species' mating system simply upon
observing its social system. For example, multi-male multi-female groups of gorillas may have an age-
graded dominance structure in which only the oldest, highest-ranking (alpha) silverback male is allowed
to copulate; in this case, the mating system is not polygynandrous (as one might expect in a multi-male
group) but instead polygynous (Robbins 2011). Another example can be found in guenons, which live in
one-male groups. During the mating season multi-male influxes occur in which outside males come into
the group, copulate with the females, and then leave again (Cords 1988). The mating system in this case
is not polygynous, which one would expect, but polygynandrous. Finally, gibbons and siamangs were
first thought to be monogamous, but then observations of extra-pair mating (i.e., copulations with
individuals outside of the pair-bond) confirmed that they are not always monogamous but sometimes
polygynous, polyandrous, or both (Palombit 1994).