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THE LIVING PRIMATES | Lead Editor: Jessica Rothman

Primate Cognition
By: Lydia M. Hopper, Ph.D. (Lester E. Fisher Center for the Study and Conservation of Apes, Lincoln Park
Zoo) & Sarah F. Brosnan, Ph.D. (Departments. of Psychology and Philosophy, Neuroscience Institute,
Georgia State University) 2012 Nature Education
Citation: Hopper, L. M. & Brosnan, S. F. (2012) Primate Cognition. Nature Education
Knowledge 5(8):3

What has driven the need for humans? Intelligence? To gain insight into our own cognitive evolution, we can look to our
closest living ancestors; our nonhuman primate cousins.
Aa

Aa

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Cognition is the ability to interpret and connect information that is perceived in the w orld and to then apply this personal know ledge adaptively to future situations and problems (Cheney & Seyfarth,
1990). Importantly, this allow s the animal to improve their fitness. All animals have to negotiate their environment to allow them to forage for food, avoid predation and find mates. Primates stand out
among other taxa for their flexibility in how they deal w ith the w orld around them. They inhabit both complex physical and social w orlds, w hich have each been posited as the major selective
pressures driving the advancement of primate brain size and, by extension, their cognitive abilities (Byrne, 2000; Reader

et al., 2011); accordingly w e first describe their understanding of their

physical environment and then go on to discuss social cognition.

Primate Cognition about the Physical World


Cognitive mapping
Across the w orld, primates have to master an array of environmental topographies, negotiate w eather extremes, and move about their territory safely and efficiently. Within their hom e range,
primates need to understand and recall the features that lie w ithin it, w hich requires both cognitive m apping skills and a flexible memory. Importantly, cognitive mapping allow s primates to
remember not just w here physical landmarks are (e.g., trees) but also salient and changing features (w hen that tree w ill bear fruit). When navigating their environment, primates do not travel
aimlessly, but move about it in a directed manner. Monkeys (e.g. Hylobates
routes, maximizing the available food that they pass (Asensio, et

lar and Cebus apella) and apes (e.g. Pan troglodytes and Gorilla gorilla)w ill take the most efficient
al., 2011; Bates & Byrne, 2009; Janson, 1998; Vedder, 1984), to reach preferred fruiting trees (Ban et al., 2014). Cognitive

mapping is essential for increasing fitness, because in dense environments vision is often extremely limited,and fruiting trees (or other valuable resources) may not be easily visible; planning travel
paths thus reduces energy expenditure and search costs. Of course, the foci of individual interest may vary, even w ithin species, revealing flexibility in primate decision-making. For instance,
chimpanzee females are more likely to plan routes follow ing paths that w ill lead them to food sources w hereas males appear more concerned w ith monitoring the border of their territory (Bates &
Byrne, 2009).

Memory and future planning


The memory of primates has been studied fairly extensively, revealing, for example, that apes are able to recall locations of items after delays of up to 16 hours (Menzel, 1999) and replicate novel
actions demonstrated to them over 24 hours previously (Hopper, 2010). Furthermore, like humans, monkeys ( Macaca

mulatta) and apes (Pan troglodytes ) are able to recall serial lists of

images (Inoue & Matsuzaw a, 2007; Sands & Wright, 1980). Indeed, the w orking m em ory capacities of captive chimpanzees, in some cases, may outdo the performance of humans (Inoue &
Matsuzaw a, 2007; but see Cook & Wilson, 2010). Primates' memory also allow s for future planning. Chimpanzees, for example, are able to select specific tools and save them to solve particular
tasks at a later time (Mulcahy & Call, 2006a) and, w hen foraging for fruits in their environment, monkeys ( Lophocebus
temperatures to determine on w hich trees ripe fruit w ill now be likely found (Janmaat et
monkey ( Macaca

fascicularis , Papio anubis

and Saimiri

albigena)have been reported to apply know ledge of the past days'


al., 2006). This flexibility may require mental representation and both ape (Pan troglodytes ) and

sciureus ) species are able to apply Roman numerals to specific quantities in w ays comparable w ith young children, and
al., 1997; Schmitt & Fischer, 2011).

use such numerals for simple arithmetic (Boysen & Berntson, 1989; Olthof, et

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Tool-use behavior and causal understanding


The sensory-motor cognition of primates is highlighted by their dexterous use of tools. Both ape and monkey species use tools to eat otherw ise inaccessible foods (Ottoni & Mannu, 2001; Whiten,
2011). The ability of primates to manufacture tools to specific requirements demonstrates an understanding of causal relations and physical properties (causal understanding). White bearded
capuchin monkeys ( Cebus

libidinosus ), for example, select substrates of specific w eight and material to use as hammers w hen cracking nuts (Visalberghi et al., 2008). This skill appears to
apella) show ing indiscriminate and ineffectual preferences for tool material, and their skill increasing w ith

be learnt during the monkey's development, w ith young brow n capuchin monkeys ( C.
age (Pouydebat et

al., 2006). This learnt skill highlights the interplay of the material and social w orld that primates negotiate; individuals learn about the causal relations of their physical environment

from both personal interaction and social observation. More formal tests of causal understanding w ith apes in captivity report that all four apes species show a clear understanding of causal
relations (Mulcahy & Call, 2006b, but see Horner & Whiten, 2007), using tools flexibly to negotiate food around obstacles.

Social Cognition
The social w orld that primates inhabit adds another dimension to their cognitive demands. Tests of self-recognition have led to the hypothesis that some primate species have an understanding of

et
al., 2000), retain know ledge about social relations (Byrne & Whiten, 1988), and that chimpanzees (Pan troglodytes , Hare et al., 2001), but not capuchin monkeys (Cebus apella, Hare et
al., 2003), recognize w hat others know . This know ledge of fellow members of their social group enables primates to determine w ho to associate w ith, cooperate w ith, and potentially learn from,
theory of m ind (Gallup, 1970), w hich further enables them to monitor their social w orld and their relation to it. Research reveals that primates can recognize and distinguish individuals (Parr

and how to manipulate individuals or situations (Brosnan & Hopper, 2013).

Cooperation and prosocial behaviour


Primates cooperate on tasks as diverse as group hunting, territory defence, and rank acquisition through alliances, and, in some cases, show ing quite complex behaviours (e.g., Boesch 1994). In
some tasks, at least, non-human primates reach outcomes equivalent to those of humans (e.g. Brosnan et

al., 2011). Primates are sensitive to task demands, for instance cooperating at a higher

rate w hen rew ards cannot easily be dominated by another individual (de Waal & Davis, 2006). Intriguingly, species w hich routinely cooperate w ith non-kin are also more likely to respond negatively
to inequity, indicating that cooperation and an abilityto recognise inequity may have co-evolved in some species (Brosnan, 2011). Primates also appear to engage in only limited reciprocity;

et al., 2008; Gomes & Boesch, 2009), short-term laboratory experiments have found little evidence for such
et al., 2008; but see Melis et al., 2011). This may indicate that most reciprocity is based on longer-term relationships rather than immediate contingency.
While cooperation is common, prosocial behaviour is more rare. There is little evidence in apes that individuals change their behaviour to bring a partner food (e.g. Silk et al., 2005; but see Horner
et al., 2011), although several monkey species do so (e.g. de Waal et al., 2008; Burkart et al., 2007; Massen et al., 2010), even in cases in w hich being prosocial actually results in inequity
tow ards the provider (Brosnan et al., 2010). On the other hand, even among chimpanzees, behaviours w hich help a partner are seen outside of the food context (e.g., helping; Warneken &
although reciprocity is seen in long-term analyses of w ild populations (Gomes
behaviour (Melis

Tomasello, 2006). One hypothesis is that prosocial behaviours are more common among cooperative breeders, due to the unique costs and benefits related to their interactions w ith each other
(Blaffer-Hrdy, 2009), leading them to more actively share food w ith partners (Jaeggi et

al., 2010).

Machiavellian and social Intelligence


Primates are able to use their skills in social cognition to their advantage by manipulating the behaviour of others. In fact, one hypothesis for primates' unusual intelligence is that cognitive skills w ere
strongly selected because of the necessity of outsmarting rivals (Byrne & Whiten, 1988). This may be most evident in the case of deception. Deception is rarely observed, partially because it is
expected to be uncommon to avoid habituation by potential targets (Cheney & Seyfarth 1990). How ever, a field experiment w ith capuchin monkeys ( Cebus

apella) has demonstrated tactical

deception (Wheeler, 2009). In cases of artificial provisioning, low er-ranking monkeys are more likely to give alarm calls in contested situations in w hich the departure of the dominant could result in
more food for the caller. Thus the absence of an extensive literature on deception may equally be due to an absence of deception, or to the challenges of demonstrating w hat is by necessity a rare
behaviour.

Social learning
The ability of primates to learn socially from one another, taking advantage of the know ledge of others, blurs the distinction betw een how they learn about their physical and social w orlds (van
Schaik & Burkart, 2011). The major importance of social learning is that it facilitates the transfer of information betw een individuals w ithout the need for genetic inheritance and can bypass
potentially costly and time-consuming trial-and-error learning. In humans, this has allow ed for the emergence of our diverse cultural w orld. Social learning has been reported for many primate
species but do they have cultures like us? Primates do indeed show evidence for socially-learned, behavioural traditions. Although undoubtedly not as rich as human culture, certain w ild primate
communities sustain multiple site-specific behaviours analogous to our culture w ith such traditions identified in ape ( Pan

fuscata, Cebus capucinus

and Ateles

troglodytes , Pongo pygmaeus ) and monkey (Macaca

geoffroyi) species in the w ild (Santorelli et al., 2011).

There is clearly a marked difference, how ever, betw een human culture and the behavioural traditions of primates, and w hat underlies this contrast is currently a topic of much debate. Cum ulative
culture has been proposed to describe the underlying mechanism that has allow ed for the development of our complex technologies. Without the ability for im itation or teaching, researchers

et al., 2014; Tomasello,


et al., 2012). Given this, along

argue, primates may not be capable of cumulative learning because the complex information required for such intricate behaviours cannot be transmitted faithfully (Dean
1999). Experimental studies w ith humans, how ever, reveal that teaching and imitation are not prerequisite for cumulative culture (Caldw ell & Millen, 2009; Caldw ell
w ith the mounting evidence that primates are able to copy even complex tool-use behaviours from observing others (Price et

al., 2009), w hat can explain the relative paucity of the cultural w orlds

of our closest-living relatives, especially for the chimpanzee? It may w ell be that the very ability of chimpanzees to copy others leads to their limited ability to build on the know ledge of previous
generations. After learning a specific method from observations of others, chimpanzees appear to become entrenched and unable to transition to a new behaviour, even if the introduced method is
more efficient (Hopper

et al., 2011). Although seemingly a limitation of their learning capacities, such as conformity and conservatism may enable chimpanzees to maintain strong social bonds
al., 2011).

and potentially avoid risks in the w ild. Thus, the key ability here is not just in copying others, but determining w hen to do so (Rendell et

Conclusions
We are still learning much about the cognitive capacities of primates, informing us not just about our ow n evolutionary past and the capabilities of our common ancestor, but also about the specific
and unique adaptations each species has for its ow n physical and social w orld. The cognitive capacities of primates should not just be used as a baseline for comparison to human capabilities, but
also studied in their ow n right in order to better understand the cognitive repertoire of each species and the effects of differing environments on the evolution of cognition.

Glossary
causal understanding: The ability to "understand not just that tw o events are associated w ith one another in space and time, but also that there is some mediating force' that binds the tw o
events to one another w hich may be used to predict or control those events" (Visalberghi & Tomasello, 1998, p.189).
cognitive m apping: "The representation embodied in a cognitive map is typically assumed to encode distances and directions and to enable mental operations of them" (Shettlew orth, 1998, p.296)
allow ing animals to calculate the optimum routes from point A to point B.
conservatism : Mastery of a skill inhibits further exploration to new alternative options or behavioural strategies w hich may be more efficient
cooperation: Tw o or more individuals increase their direct fitness more by w orking together than individually (Brosnan, Salw iczek & Bshary, 2010).

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cooperative breeders: Social system in w hich young are cared for by both parents and, often, their mature offspring.
cum ulative culture: The process by w hich generations build upon the know ledge of previous ones creating increasingly complex artefacts and technologies through the accumulation of design
the so-called "ratchet effect" (Tomasello, 1999).
deception: The provisioning of false information w hich affects another individual's behaviour in favour of the deceiver. While this may occur w ithout intention on the part of the deceiver, resulting
in functional, or tactical, deception, individuals may also do so intentionally.
fission-fusion: In w hich a group of individuals breaks ups and reforms over the course of a day or w eek meaning that each party size represents a subset of total group size.
fitness: The ability to survive and successfully reproduce (e.g., evolutionary fitness)
hom e range: The territory w hich a primate, or group of primates, occupies. Note, primates move about their territory but may only travel w ithin small subsections each day. Furthermore, the
degree to w hich a primates' home range overlaps w ith conspecifics and different primate species is affected not just by the specific primate but also by environmental and space pressures (e.g.,
food availability and deforestation).
im itation: Copying the exact behaviours demonstrated by another. Compare w ith emulation' in w hich an individual merely replicates the goal or end-state of a novel behaviour performed by others.
inequity: A situation in w hich unequal rew ards are received. The behavioural response studied is inequity aversion' vis--vis a sense of unfairness (see Brosnan, 2006, for further discussion).
prosocial behaviour: Behaviour w hich results in a benefit to another individual.
teaching: "An individual actor A can be said to teach if it modifies its behaviour only in the presence of a naive observer, B, at some cost or at least w ithout obtaining an immediate benefit for itself.
A's behaviour thereby encourages or punishes B's behaviour, or provides B w ith experience, or sets an example for B. As a result, B acquires know ledge or learns a skills earlier in life or more
rapidly or efficiently than it might otherw ise do, or that it w ould not learn at all" (Caro & Hauser, 1992, p. 153).
theory of m ind: The attribution of mental states such as beliefs, thoughts, desires and intentions to oneself and others. See Heyes (1998) for a detailed discussion about primate understanding of
theory of mind.
traditions: "enduring behaviour patterns shared among members of a group that depend to a measurable degree on social contributions to individual learning, resulting in shared practices among
members of a group" (Fragaszy & Perry, 2003, p.3)
w orking m em ory: The short-term retention of information w hich may sometimes incorporate elements of information processing.

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Outline | Keywords

Explore This Subject


BASIC

INTERMEDIATE

Primate Conservation: Is the Cup Half


Empty or Half Full?

Male Reproductive Strategies

Primate Sociality and Social Systems

Primate Cognition

ADVANCED

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Primates in Communities: The Ecology of
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