The Role of Primate Sociality in Evolutionary

Theory
Some animals, and more specifically primates, have been noted for having brains of an
uncommonly large size. One explanation for this phenomenon is the ‘social brain’ or
‘social intelligence’ hypothesis, which states that brain sizes have increased in order to
handle the demands of sociality (Lindenfors 2005; Pérez-Barbería et al. 2007). The
increase in brain size can be attributed to the computational power needed to track others
in the group and ensure individual and species survival (Müller & Soligo 2005; Pérez-
Barbería et al. 2007). Sociality in primates – and thus also in humans – has then some
evolutionary basis.
This review will discuss the impact and possible reasons underlying the evolution of
sociality, in relation to primates as a close approximation to humans. In particular, it will
address the question of whether sociality in primates evolved as a means of survival.
Each article will be rated on a five star scale, where one star represents one mark. Five
stars would indicate that the article is highly relevant to the topic, while four stars would
indicate a very good association with the topic. Three stars would show a mild
association with the topic, while two would indicate that the article displays only a
passing reference. One star would mean that the article does not display much relevance
or only a part of the article is really relevant to the topic.

1
Social bonds of female baboons enhance infant survival
This article aimed to produce evidence to support the theory that social bonds have a
unique positive influence on primate adaptability, regardless of other factors. The
evidence was collected from a 16 year study of the Kenyan wild baboon population
Papio cynocephalus. Social contact was defined by three estimates: the amount of time a
baboon would be within five metres of another baboon, the amount of time a baboon
would be grooming others and the amount of time it would be groomed by others. These
factors encompass the social activities of the baboon population and are considered to be
an accurate measure of primate sociality. The three estimates were then combined to form
a combined sociality index score.
Female baboons engaging more frequently with other adult baboons, as well as those
occupying definite social positions in the group hierarchy, were found to be most likely
to succeed in raising their young. Females with higher index scores also produced
children with marginally higher survival rates than those females with lower scores.
Furthermore these observations were found to be independent of other factors (e.g.
female dominance) implying that primate sociality plays an important evolutionary role.

Silk, JB, Alberts, SC, Altmann, J 2003. Social bonds of female baboons enhance infant
survival. Science 302: 1231-1234.

Primate Sociality in Evolutionary Context

The aim of this article was to examine sociality from an evolutionary perspective and see
whether it affected certain factors that influence sociality. These factors were an
individual’s weight, eating habits, movement patterns and use of environment. Rodents
were used to create a model that could later be extrapolated to investigate primates,
where a rodent genus was considered social if male and female social interactions
occurred outside mating times and formed some kind of network with at least one adult
animal. Rodents were considered a suitable group for examination as they are the largest
group of mammals and display a wide range of characteristics over the given factors.

2
Data was collected from all rodent genera for which sociality could be measured using
these four factors. The study concluded that these four factors, all impacting on animal
sociality, were highly influenced by the evolution of the rodent. Large animals, those
living in trees, and animals feeding on fruits, vegetables or animal matter were found to
display the highest degree of sociality. This is believed to have evolved as a means for all
animals to obtain food and hence be able to survive. This research implies that animals
that mostly or exclusively consume fruit are instrumental in the development of social
networks.

Müller, AE & Soligo, C 2005. Primate sociality in evolutionary context. American
Journal of Physical Anthropology 128: 399-419.

Neocortex evolution in primates: the ‘social brain’ is for females

This study used the size of the neocortex, the component handling complex mental
processes in animal brains, to explore how these processes evolved. The research aimed
to show that the social intelligence is only applicable to females. This was because
females were known to cluster together as a result of environmental conditions, while for
males the only important factor was the position of the females.
As neocortex size and brain size were correlated, and the individual’s weight and the
group’s size were also factors affecting sociality, data was collected for all four variables.
The research concluded that those species containing large groups of social females
showed larger neocortex sizes. Relatively large measurements of neocortex size for males
was thought to show that it might be important for both sexes to know about female-
female bonds.The research also concluded that brain development occurs alongside the
social development of females. It also implies that in the more sociable groups, female
neocortex sizes were larger than the male equivalents.

Lindenfors, P 2005. Neocortex evolution in primates: the ‘social brain’ is for females.
Biology Letters 1: 407-410.

3
Evidence for Coevolution of Sociality and relative brain size in three orders of
mammals
The aim of this study was to investigate the pairing between sociality and brain size over
time. Though there is a correlation between these two variables, the literature does not
indicate how the two became linked over time. Current developments in statistical
analysis have now made it possible for this relationship to be understood.
Data was collected on the brain size and individual weight for 206 species comprising
four different mammalian orders, of which 42 were primates. Sociality was able to be
measured from all of these species, on the basis that a social group comprised at the least
two adult animals who regularly communicated with each other. As primates are
primarily social animals, the definition of sociality was further narrowed in their case – a
social group was observed if the geometric mean of the group size was larger than that
for the entire group.
While it seemed apparent that sociality and brain size evolved alongside each other in all
orders, the study concluded that the way in which this occurred varied for each order. The
increase in brain size could be attributed to the increase in mental processes needed to
track other group members for various reasons important for survival – e.g. to avoid
separation, or to generate ideas as to how to avoid competition, persecution etc. It is
implied that the evolution of sociality, and its close linkage with brain size, has definitely
lead to an increase in the chance of survival of a species.

Pérez-Barbería, FJ, Shultz, S, Dunbar, RIM 2007. Evidence for coevolution of sociality
and relative brain size in three orders of mammals. Evolution 61: 2811-2821.

4
Sociality in Callithrix penicillata: II Individual strategies during intergroup
encounters
This article investigated interactions between groups of the same animals to determine
their impact on various factors, e.g. territorial boundaries and access to various items.
The interactions of a group of Brazilian black tufted-ear marmosets with each other were
observed as part of the study. Both antagonistic and friendly behaviors were monitored,
with the latter being much more predominant. However antagonistic behavior can be
important when it comes to defense of territory, which is closely related to reproductive
success and hence survival.
Data was collected based on observations of the marmosets during morning and
afternoon periods. An intergroup meeting was recorded to have occurred if more than
one marmoset of another group came into contact with the test group.
The study concluded that most meetings between marmoset groups were harmonious, and
in many exchanges the marmosets did not seem to respond to the presence of the other
animal, instead feeding or resting as normal. Antagonistic behaviour without physical
violence was mostly displayed by males to males and when present in females, was either
directed towards males or against strange females. In the former, this was judged to occur
as a result of an unsuccessful copulatory attempt, but in the latter no clear reason was
evident. Where there was insufficient food available or intruders were present,
particularly sexually aggressive ones, antagonistic behavior was more prevalent. The
study implied that social interactions are used to protect and ensure the future survival of
the species.

Decanini, DP, Macedo, RM 2008. Sociality in Callithrix penicillata: II individual
strategies during intergroup encounters. International Journal of Primatology 29: 627-
639.

5
 Reference List

Decanini, DP, Macedo, RM 2008. Sociality in Callithrix penicillata: II individual

strategies during intergroup encounters. International Journal of Primatology 29: 627-
639.

Lindenfors, P 2005. Neocortex evolution in primates: the ‘social brain’ is for females.
Biology Letters 1: 407-410.

Müller, AE & Soligo, C 2005. Primate sociality in evolutionary context. American

Journal of Physical Anthropology 128: 399-419.

Pérez-Barbería, FJ, Shultz, S, Dunbar, RIM 2007. Evidence for coevolution of sociality
and relative brain size in three orders of mammals. Evolution 61: 2811-2821.

Silk, JB, Alberts, SC, Altmann, J 2003. Social bonds of female baboons enhance infant
survival. Science 302: 1231-1234.

6
 Abstracts
Primate Sociality in Evolutionary Context
Alexandra E. Müller1* and Christophe Soligo2
1Anthropological Institute and Museum, University of Zu¨ rich, 8057 Zu¨ rich, Switzerland
2Human Origins Group, Department of Palaeontology, Natural History Museum, London SW7
5BD, UK

KEY WORDS primate origins; rodents; social organization; social networks; frugivory

ABSTRACT Much work has been done to further our

understanding of the mechanisms that underlie the diversity
of primate social organizations, but none has addressed
the limits to that diversity or the question of what
causes species to either form or not form social networks.
The fact that all living primates typically live in social
networks makes it highly likely that the last common
ancestor of living primates already lived in social networks,
and that sociality formed an integral part of the
adaptive nature of primate origins. A characterization of
primate sociality within the wider mammalian context is
therefore essential to further our understanding of the
adaptive nature of primate origins. Here we determine
correlates of sociality and nonsociality in rodents as a
model to infer causes of sociality in primates. We found
sociality to be most strongly associated with large-bodied
arboreal species that include a significant portion of fruit
in their diet. Fruits and other plant products, such as
flowers, seeds, and young leaves, are patchily distributed
in time and space and are therefore difficult to find. These
food resources are, however, predictable and dependable
when their location is known. Hence, membership in a
social unit can maximize food exploitation if information
on feeding sites is shared. Whether sociality evolved in the
primate stem lineage or whether it was already present
earlier in the evolution of Euarchontoglires remains uncertain,
although tentative evidence points to the former
scenario. In either case, frugivory is likely to have played
an important role in maintaining the presence of a social
lifestyle throughout primate evolution. Am J Phys Anthropol
128:399–414, 2005. © 2005 Wiley-Liss, Inc.

Neocortex evolution in
primates: the ‘social brain’
is for females
Patrik Lindenfors*,†
Department of Biology, University of Virginia, Gilmer Hall,
Charlottesville, VA 22904-4328, USA
*( patrik.lindenfors@virginia.edu)
†Temporary address: P.O. Box 19423, 202 KNH-Nairobi, Kenya.

According to the social intelligence hypothesis,

relative neocortex size should be directly related

7
to the degree of social complexity. This hypothesis
has found support in a number of comparative
studies of group size. The relationship
between neocortex and sociality is thought to
exist either because relative neocortex size
limits group size or because a larger group size
selects for a larger neocortex. However,
research on primate social evolution has indicated
that male and female group sizes evolve
in relation to different demands. While females
mostly group according to conditions set by the
environment, males instead simply go where the
females are. Thus, any hypothesis relating to
primate social evolution has to analyse its
relationship with male and female group sizes
separately. Since sex-specific neocortex sizes in
primates are unavailable in sufficient quantity,
I here instead present results from phylogenetic
comparative analyses of unsexed relative neocortex
sizes and female and male group sizes.
These analyses show that while relative
neocortex size is positively correlated with
female group size, it is negatively, or not at all
correlated with male group size. This indicates
that the social intelligence hypothesis only
applies to female sociality.

Sociality in Callithrix penicillata: II. Individual

Strategies During Intergroup Encounters
Daniel P. Decanini & Regina H. Macedo
Received: 28 May 2007 / Accepted: 9 October 2007 /
Published online: 5 June 2008
# Springer Science + Business Media, LLC 2008
Abstract In social animals, intergroup interactions, whether through agonistic and
competitive behaviors or affiliative ones, can influence important parameters such as
home range, territory sizes, and access to resources, which may directly affect both
female and male fitness. We studied the intergroup interaction patterns of a wild
group of black-tufted-ear marmosets (Callithrix penicillata) in central Brazil.
Agonistic interactions occurred at low frequencies during intergroup encounters.
The marmosets directed agonistic interactions without physical aggression primarily
against same-sex individuals, suggesting that male and female aggression patterns
are shaped by their sexual interests. However, females of the focal group also
directed agonistic behavior toward extragroup males that attempted copulation. The
marmosets appeared to use intergroup encounters to gather information about
possible partners and extragroup reproductive opportunities. Intergroup sexual
interactions occurred mainly in the form of copulations or attempted copulations
by all adults, with the exception of the dominant female. Our results suggest that a
possible reproductive strategy used by males is to attempt fertilization of extragroup
females. Adult males copulated with the same extragroup female during several
opportunities, which suggests sperm competition or the establishment of social
bonds with neighboring females.

8
EVIDENCE FOR COEVOLUTION OF
SOCIALITY
AND RELATIVE BRAIN SIZE IN
THREE ORDERS
OF MAMMALS
F. Javier Pe´ rez-Barber´ıa,1,2 Susanne Shultz,3,4 and Robin I. M. Dunbar3,5
1The Macaulay Institute, Craigiebuckler, Aberdeen, AB15 8QH, Scotland, United Kingdom
2E-mail: j.pe´ rez-barberı´a@macaulay.ac.uk
3British Academy Centenary Research Project, School of Biological Sciences, University of
Liverpool, Crown St. Liverpool
L69 7ZB, United Kingdom
4E-mail: susanne.shultz@liv.ac.uk
5E-mail: rimd@liv.ac.uk
Received September 8, 2006
Accepted July 17, 2007
As the brain is responsible for managing an individual’s behavioral response to its
environment, we should expect that large relative
brain size is an evolutionary response to cognitively challenging behaviors. The “social brain
hypothesis” argues that maintaining
group cohesion is cognitively demanding as individuals living in groups need to be able to
resolve conflicts that impact on their
ability to meet resource requirements. If sociality does impose cognitive demands, we expect
changes in relative brain size and
sociality to be coupled over evolutionary time. In this study, we analyze data on sociality and
relative brain size for 206 species
of ungulates, carnivores, and primates and provide, for the first time, evidence that changes
in sociality and relative brain size
are closely correlated over evolutionary time for all three mammalian orders. This suggests a
process of coevolution and provides
support for the social brain theory. However, differences between taxonomic orders in the
stability of the transition between
small-brained/nonsocial and large-brained/social imply that, although sociality is cognitively
demanding, sociality and relative
brain size can become decoupled in some cases. Carnivores seem to have been especially
prone to this.

Social Bonds of Female Baboons

Enhance Infant Survival
Joan B. Silk,1* Susan C. Alberts,2,4 Jeanne Altmann3,4,5
Among nonhuman primates, females often form strong bonds with kin and
other group members. These relationships are thought to have adaptive value
for females, but direct effects of sociality on fitness have never been demonstrated.
We present 16 years of behavioral data from a well-studied population
of wild baboons, which demonstrate that sociality of adult females is positively
associated with infant survival, an important component of variation in female
lifetime fitness. The effects of sociality on infant survival are independent of
the effects of dominance rank, group membership, and environmental conditions.
Our results are consistent with the evidence that social support has
beneficial effects on human health and well-being across the life span. For
humans and other primates, sociality has adaptive value.

9
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Some animals, and more specifically primates, have been noted for having brains of an
uncommonly large size. One explanation for this phenomenon is the ‘social brain’
or ‘social intelligence’ hypothesis, which states that brain sizes have increased in
order to handle the demands of sociality (Lindenfors 2005, Dunbar et.al 2007). The
increase in brain size can be attributed to the computational power needed to track others
in the group and ensure individual and species survival (Müller & Soligo 2005; Dunbar
et al. 2007). Sociality in primates – and thus also in humans – has then some
evolutionary basis. This review will discuss the impact and possible reasons underlying
the evolution of sociality, in relation to primates as a close approximation to humans. In
particular, it will address the question of whether sociality in primates evolved as a means
of survival. Each article will be rated on a five star scale, where one star represents one
mark. Five stars would indicate that the article is highly relevant to the topic, while four
stars would indicate a very good association with the topic. Three stars would show a
mild association with the topic, while two would indicate that the article displays only a
passing reference. One star would mean that the article does not display much relevance
or only a part of the article is really relevant to the topic. This article aimed to produce
evidence to support the theory that social bonds have a unique positive influence on
primate adaptability, regardless of other factors. The evidence was collected from a 16
year study of the Kenyan wild baboon population Papio cynocephalus. Social contact was
defined by three estimates: the amount of time a baboon would be within five metres of
another baboon, the amount of time a baboon would be grooming others and the amount
of time it would be groomed by others. These factors encompass the social activities of
the baboon population and are considered to be an accurate measure of primate sociality.
The three estimates were then combined to form a combined sociality index score.
Female baboons engaging more frequently with other adult baboons, as well as those
occupying definite social positions in the group hierarchy, were found to be most likely
to succeed in raising their young. Females with higher index scores also produced
children with marginally higher survival rates than those females with lower scores.
Furthermore these observations were found to be independent of other factors (e.g.

10
female dominance) implying that primate sociality plays an important evolutionary role.
The aim of this article was to examine sociality from an evolutionary perspective and see
whether it affected certain factors that influence sociality. These factors were an
individual’s weight, eating habits, movement patterns and use of environment.
Rodents were used to create a model that could later be extrapolated to investigate
primates, where a rodent genus was considered social if male and female social
interactions occurred outside mating times and formed some kind of network with at least
one adult animal. Rodents were considered a suitable group for examination as they are
the largest group of mammals and display a wide range of characteristics over the given
factors. Data was collected from all rodent genera for which sociality could be measured
using these four factors. The study concluded that these four factors, all impacting on
animal sociality, were highly influenced by the evolution of the rodent. Large animals,
those living in trees, and animals feeding on fruits, vegetables or animal matter were
found to display the highest degree of sociality. This is believed to have evolved as a
means for all animals to obtain food and hence be able to survive. This research implies
that animals that mostly or exclusively consume fruit are instrumental in the development
of social networks. This study used the size of the neocortex, the component handling
complex mental processes in animal brains, to explore how these processes evolved. The
research aimed to show that the social intelligence is only applicable to females. This was
because females were known to cluster together as a result of environmental conditions,
while for males the only important factor was the position of the females. As neocortex
size and brain size were correlated, and the individual’s weight and the group’s
size were also factors affecting sociality, data was collected for all four variables. The
research concluded that those species containing large groups of social females showed
larger neocortex sizes. Relatively large measurements of neocortex size for males was
thought to show that it might be important for both sexes to know about female-female
bonds.The research also concluded that brain development occurs alongside the social
development of females. It also implies that in the more sociable groups, female
neocortex sizes were larger than the male equivalents. The aim of this study was to
investigate the pairing between sociality and brain size over time. Though there is a
correlation between these two variables, the literature does not indicate how the two
became linked over time. Current developments in statistical analysis have now made it
possible for this relationship to be understood. Data was collected on the brain size and
individual weight for 206 species comprising four different mammalian orders, of which
42 were primates. Sociality was able to be measured from all of these species, on the
basis that a social group comprised at the least two adult animals who regularly
communicated with each other. As primates are primarily social animals, the definition of
sociality was further narrowed in their case – a social group was observed if the
geometric mean of the group size was larger than that for the entire group. While it
seemed apparent that sociality and brain size evolved alongside each other in all orders,
the study concluded that the way in which this occurred varied for each order. The
increase in brain size could be attributed to the increase in mental processes needed to
track other group members for various reasons important for survival – e.g. to avoid
separation, or to generate ideas as to how to avoid competition, persecution etc. It is
implied that the evolution of sociality, and its close linkage with brain size, has definitely
lead to an increase in the chance of survival of a species. This article investigated

11
interactions between groups of the same animals to determine their impact on various
factors, e.g. territorial boundaries and access to various items. The interactions of a group
of Brazilian black tufted-ear marmosets with each other were observed as part of the
study. Both antagonistic and friendly behaviors were monitored, with the latter being
much more predominant. However antagonistic behavior can be important when it comes
to defense of territory, which is closely related to reproductive success and hence
survival. Data was collected based on observations of the marmosets during morning and
afternoon periods. An intergroup meeting was recorded to have occurred if more than one
marmoset of another group came into contact with the test group. The study concluded
that most meetings between marmoset groups were harmonious, and in many exchanges
the marmosets did not seem to respond to the presence of the other animal, instead
feeding or resting as normal. Antagonistic behaviour without physical violence was
mostly displayed by males to males and when present in females, was either directed
towards males or against strange females. In the former, this was judged to occur as a
result of an unsuccessful copulatory attempt, but in the latter no clear reason was evident.
Where there was insufficient food available or intruders were present, particularly
sexually aggressive ones, antagonistic behavior was more prevalent. The study implied
that social interactions are used to protect and ensure the future survival of the species.

12