Primate cognition

Primate cognition is the study of the intellectual and behavioral skills of non-human primates, particularly in the fields of psychology, behavioral biology, primatology, and anthropology.

Primates are capable of high levels of cognition; some make tools and use them to acquire foods and for social displays; some have sophisticated hunting strategies requiring cooperation, influence and rank; they are status conscious, manipulative and capable of deception; they can recognise kin and conspecifics; they can learn to use symbols and understand aspects of human language including some relational syntax, concepts of number and numerical sequence.

Studies in primate cognition

Theory of mind

Theory of mind (also known as mental state attribution, mentalizing, or mindreading) can be defined as the "ability to track the unobservable mental states, like desires and beliefs, that guide others' actions". Premack and Woodruff's 1978 article "Does the chimpanzee have a theory of mind?" sparked a contentious issue because of the problem of inferring from animal behavior the existence of thinking, of the existence of a concept of self or self-awareness, or of particular thoughts.

Non-human research still has a major place in this field, however, and is especially useful in illuminating which nonverbal behaviors signify components of theory of mind, and in pointing to possible stepping points in the evolution of what many claim to be a uniquely human aspect of social cognition. While it is difficult to study human-like theory of mind and mental states in species which we do not yet describe as "minded" at all, and about whose potential mental states we have an incomplete understanding, researchers can focus on simpler components of more complex capabilities.

For example, many researchers focus on animals' understanding of intention, gaze, perspective, or knowledge (or rather, what another being has seen). Part of the difficulty in this line of research is that observed phenomena can often be explained as simple stimulus-response learning, since mental states can often be inferred based on observed behavioural cues. Recently, most non-human theory of mind research has focused on monkeys and great apes, who are of most interest in the study of the evolution of human social cognition. Research can be categorized in to three subsections of theory of mind: attribution of intentions, attribution of knowledge (and perception), and attribution of belief.

1. Attribution of intentions. Research on chimpanzees, capuchin monkeys, and Tonkean macaques (Macaca tokeana) has provided evidence that they are sensitive to the goals and intentions of others and are able to differentiate between when an experimenter is unable to give them food versus when the experimenter is just unwilling to.
2. Attribution of knowledge (and perception). Hare et al. (2001) demonstrates that chimpanzees are aware of what other individuals know. They can also understand what another perceives, and they selectively choose food that is not visible to their competitor.
3. Attribution of belief. A false-belief test is a comprehensive test used to test for an individual's theory of mind. Understanding language is a key component to being able to understand the directions for the false-belief test, and researchers have had to get creative to utilize this test in the research of non-human primates' theory of mind. Recent technology has enabled researchers to closely resemble the false-belief task without needing to use language. In Krupenye et al. (2016), an advanced eye-tracking technology was used to test for false-belief understanding in apes. The findings of this experiment showed that apes understood and accurately anticipated the behavior of an individual who held a false belief.

There has been some controversy over the interpretation of evidence purporting to show theory of mind ability—or inability—in animals. Part of this debate has involved whether animals are really able to associate cognitive abilities with another individual, or if they are just able to read and understand behavior. Povinelli et al. (1990) points out that most evidence in support of great ape theory of mind involves naturalistic settings to which the apes have already adapted through past learning. Their "reinterpretation hypothesis" explains away evidence supporting attribution of mental states to others in chimpanzees as merely evidence of risk-based learning; that is, the chimpanzees learn through experience that certain behaviors in other chimpanzees have a probability of leading to certain responses, without necessarily attributing knowledge or other intentional states to those other chimpanzees. They have proposed testing theory of mind abilities in great apes in novel, and not naturalistic settings. Experimenters since then, such as demonstrated in Krupenye et al. (2016), have gone to extensive lengths to control for behavioral cues by placing the apes in novel settings as suggested by Povinelli and colleagues. Research has shown that there is substantial evidence for some non-human primates to track the mental state, like desires and beliefs, of other individuals that cannot be deduced to a response of learned behavioural cues.

Communication in the wild

For most of the 20th century, scientists who studied primates thought of vocalizations as physical responses to emotions and external stimuli. The first observations of primate vocalizations representing and referring to events in the exterior world were observed in vervet monkeys in 1967. Calls with specific intent, such as alarm calls or mating calls has been observed in many orders of animals, including primates. Researchers began to study vervet monkey vocalizations in more depth as a result of this finding. In the seminal study on vervet monkeys, researchers played recordings of three different types of vocalizations they use as alarm calls for leopards, eagles, and pythons. Vervet monkeys in this study responded to each call accordingly: going up trees for leopard calls, searching for predators in the sky for eagle calls, and looking down for snake calls. This indicated a clear communication that there is a predator nearby and what kind of predator it is, eliciting a specific response. The use of recorded sounds, as opposed to observations in the wild, gave researchers insight into the fact that these calls contain meaning about the external world. This study also produced evidence that suggests vervet monkeys improve in their ability to classify different predators and produce alarm calls for each predator as they get older. Further research into this phenomenon has discovered that infant vervet monkeys produce alarm calls for a wider variety of species than adults. Adults only use alarm calls for leopards, eagles, and pythons while infants produce alarm calls for land mammals, birds, and snakes respectively. Data suggests that infants learn how to use and respond to alarm calls by watching their parents.

A different species of monkeys, the wild Campbell's monkeys have also been known to produce a sequence of vocalization that require a specific order to elicit a specific behaviour in other monkeys. Changing the order of the sounds changes the resulting behaviour, or meaning, of the call. Diana monkeys were studied in a habituation-dishabituation experiment that demonstrated the ability to attend to the semantic content of calls rather than simply to acoustic nature. Primates have also been observed responding to alarm calls of other species. Crested Guinea fowl, a ground-dwelling fowl, produce a single type of alarm call for all predators it detects. Diana monkeys have been observed to respond to the most likely reason for the call, typically a human or leopard, based on the situation and respond according to that. If they deem a leopard is the more likely predator in the vicinity, they will produce their own leopard-specific alarm call, but if they think it is a human, they will remain silent and hidden.

The ability for non-human primates to understand call systems that belong to a different species of monkey happens but to a limited extent. In this case Diana monkeys and Campbell's monkeys often form mixed species groups but they seem only to respond to each other's danger related calls.

Tool use

There are many reports of primates making or using tools, both in the wild or when captive. Chimpanzees, gorillas, orangutans, capuchin monkeys, baboons, and mandrills have all been reported as using tools. The use of tools by primates is varied and includes hunting (mammals, invertebrates, fish), collecting honey, processing food (nuts, fruits, vegetables and seeds), collecting water, weapons and shelter.

Tool making is much rarer, but has been documented in orangutans, bonobos and bearded capuchin monkeys. Research in 2007 shows that chimpanzees in the Fongoli savannah sharpen sticks to use as spears when hunting, considered the first evidence of systematic use of weapons in a species other than humans. Captive gorillas have made a variety of tools. In the wild, mandrills have been observed to clean their ears with modified tools. Scientists filmed a large male mandrill at Chester Zoo (UK) stripping down a twig, apparently to make it narrower, and then using the modified stick to scrape dirt from underneath its toenails.