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In this guide


Macaques have a rich behavioural repertoire. Understanding the intricacies of macaque behaviour can better aid our understanding of their welfare and management in captivity.

Being able to recognise, interpret and respond appropriately to their communication signals (such as facial expressions, postures and vocalisations) provides a good basis for refining many aspects of their care and use. The advice of a primate behaviour specialist is recommended for helping staff and animals to interact positively within the laboratory environment. For example ethograms and advice on recording behaviour, see the Behavioural indicators of welfare section.


Assess your knowledge and understanding of the behaviour of macaques, take our quiz on macaque behaviour.


A healthy captive environment allows macaques to spend their time engaged in a wide range of ‘natural’ behaviours.

A healthy captive environment allows macaques to spend their time engaged in a wide range of ‘natural’ behaviours. The activity budgets of wild macaques vary [1-5], but in all studies the macaques spend the majority of their time foraging, resting, allogrooming and moving around their environment. In captivity, animals that cannot spend their time in this way are at risk of becoming bored, depressed, frustrated, aggressive and/or stressed [6]. While daily patterns vary, wild macaques forage and travel most intensively in the morning and rest and allogroom throughout the afternoon.

  • A pie chart showing rhesus macaque time budgets; they spend the majority of the time foraging and resting (often in social contact).
    A pie chart showing rhesus macaque time budgets; they spend the majority of the time foraging and resting (often in social contact). Data for rhesus macaques, averaged from Jaman & Huffman 2013 and Zhou et al. 2013


Macaques have a variety of facial expressions. Expressions can be an indication of a macaques emotional state. When working with macaques, having a good knowledge of their typical expressions can help in management and to prevent injuries.

Macaques have a variety of facial expressions which they use to communicate with conspecifics, composed of movements of the ears, brow, eyes and mouth from the neutral state [7-9]. Facial expressions can be used by laboratory staff to infer the attention, intention and internal state of animals. However, the purpose of all macaque facial expressions is not yet fully understood. Expressions may be performed at different intensities in different contexts; there may be individual differences in how expressions appear; and an animal will often perform several expressions and behaviours together, as seen from the videos in this section. About one third of facial expressions are accompanied by vocalisation [9].


Resting facial expression, shown in calm social contexts or when resting alone. All other expressions differ from this standard facial display.

Signs of a neutral expression:

  • Ears relaxed.
  • Eyes are open and relaxed.
  • Brow relaxed.
  • Mouth is closed and relaxed.
  • A macaque shows a neutral face, Ears relaxed. Eyes open and relaxed. Brow relaxed. Mouth is closed and relaxed.
    Neutral face


Affiliative facial expressions are often performed together with submissive and anxious or fearful expressions, depending on the context.

Signs of an affiliative expression:

  • Ears are back against the side of the head (may flap back and forth).
  • Eyes are open, attentive and looking towards the interactant.
  • Brow relaxed or may pull up and back with the ears.
  • Mouth makes rapid open-close movements (lip-smack) with or without tongue protrusion.
  • Signs of an affiliative expression:  Ears are back against the side of the head (may flap back and forth). Eyes are open, attentive and looking towards the interactant. Brow relaxed or may pull up and back with the ears. Mouth makes rapid open-close movements (lip-smack) with or without tongue protrusion.
    Macaque affiliative face
  • During affiliative contact, a cynomolgus macaque lip smacks to maintain peaceful contact; his companion lip smacks in return to confirm affiliation.
    Lip smack interaction (Michael Gumert)
  • A cynomolgus macaque showing an affiliative face, with ears and brow drawn back, eyes looking at the interactant and mouth puckered.
    A cynomolgus macaque showing an affiliative face (Michael Gumert)
In this video, you can clearly see components of the affiliative facial expression – eyes open and attentive, ears being moved back against the sides of the head, and tongue protrusion. (Video: MRC Centre for Macaques)

Lip smack

Lip smacking indicates peaceful intentions [7]. It may be performed between any individuals intent on maintaining a peaceful relationship. Lip smacking is often performed during approach towards another macaque, or even human caregivers. For additional videos of lip smacking, see the RHVIDEO site.

In this video, the macaque signals its affiliative intention towards a known human caregiver by lip smacking with ears pulled back and brow raised.
In this video, the macaque lip smacks to a conspecific whilst also being vigilant and chewing food from its cheek pouches. (Credit: MRC Centre for Macaques)

Play face

The play face stops rough and tumble play getting out of hand. It is seen mostly in younger animals during play. The ears and brow are pulled back, the mouth is open and the top lip is pulled over the teeth.

  • Two young cynomolgus macaques display play face.
    Infant play face (Michael Gumert)
  • Young rhesus macaque displaying play face
    Play face (Image: Claire Witham)

Submissive or fearful

Grimace/bared teeth

The grimace or bared teeth display signals submission and averts possible attack [10-12]. It is performed by low-ranked individuals towards high-ranked individuals; often performed by the subordinate when approached by a dominant animal. The brow is neutral or pulled back with the ears, and the teeth are exposed. The animal performing the grimace, will look towards the interactant or may turn and look away. For additional videos of the grimace, see the RHVIDEO site.

In this video, the macaque on the left grins towards a dominant animal approaching from the right.
In this video, a female macaque grins at a group member with a squeak vocalisation. (Credit: MRC Centre for Macaques)
In this video, a female macaque grins and remains vigilant to her group (Credit: Jade Brooks and David Massey)


Tense mouth

This expression signals fear and/or alarm. It may be seen in response to aggressive conspecifics, humans, startling stimuli or predators. For videos of tense mouth, see the RHVIDEO site.


Exaggerated yawning with full display of teeth is a threat behaviour or indicative of tension [7]. Yawning without full display of teeth may simply be an indication of tiredness. Often shown during periods of uncertainty and tension such as unstable hierarchy.

  • A cynomolgus macaque yawns, displaying its teeth
    Yawn (Image: Michael Gumert)
In this video, the male macaque yawns indicating tension (Credit: BPRC). 


In this video, the macaque yawns and stares at the camera operator, indicating tension. (Credit: MRC Centre for Macaques)
This yawn is probably a ‘true’ yawn associated with tiredness, since the animal is relaxed and it’s attention is not fixed on any conspecifics. (Credit: MRC Centre for Macaques)

Chin up

Chin up is an avoidant posture that allows covert monitoring of events in the environment while avoiding making eye contact with other animals. It is seen in situations of uncertainty or when an animal has reduced escape options. Males may also approach females with chin up when soliciting for sexual behaviour.

  • A macaque sits with chin up, possibly monitoring events from a sideways position
    A macaque sits with chin up, possibly monitoring events from a sideways position (Alexander Georgiev)

Agonistic or aggressive

During agonistic or aggressive expressions, typically the ears are forward or flapping (indicating intense interest), the brow is lowered or neutral, with eyes open and staring at the interactant, and the mouth is open (the teeth may or may not be exposed). The head may be lowered with both head and body pulled forward.

Open mouth stare

The open mouth stare is used to threaten other individuals. It is seen during tension between group members, and is often performed towards unwelcome humans.

A mother suckling her infant threatens the camera operator with an open mouth stare expression. (Credit: MRC Centre for Macaques)
Examples of two different responses to an intruder. In this video, the macaque on the left lowers its brow, flaps its ears, and moves away – signs of tension or annoyance. The younger, bolder animal on the right comes forward and shows an open mouth stare expression, before also moving away from the camera operator.


Macaques have a range of postures that can indicate about their welfare state.


Piloerection [13] is a reflex arousal response in which the macaque’s hair stands up, making it look bigger, triggered by negative situations of extreme fear or anger. Piloerection typically occurs in highly arousing situations, such as the appearance of predators, humans or aggressive conspecifics.


Head-cocking [14] is thought to help non-human primates observe objects from different angles, allowing them to encode the parameters during learning [15]. It is often observed in youngsters when viewing novel objects.


Crouch signals a readiness for action. It is seen in situations causing fear, such as the presence of predators or aversive stimuli. Crouching can also be seen before and after aggressive encounters with conspecifics [7]. Crouching may be accompanied by pant threats, screams or barks depending on the context (see Vocalisations). For videos of crouching, see the RHVIDEO site.

  • Two macaques crouching before an aggressive encounter. Both are maintaining eye contact.
    Two macaques crouching before an aggressive encounter. Both are maintaining eye contact. (Credit: Michael Gumert)
  • Cynomolgus macaque crouching, with bared teeth
    Cynomolgus macaque crouching, with bared teeth indicating submissiveness (Credit: Michael Gumert)


Freezing is a fear response to predators, aversive stimuli or unexpected noise. For videos of freezing, see the RHVIDEO site.

Tail up

An erect tail signals alertness and is often seen in animals exerting their dominance over conspecifics. It also accompanies sexual presentations.

  • A rhesus macaques standing with it's tail stiff in an up-right position to display dominance
    A rhesus macaques standing with it's tail stiff in an up-right position to display dominance (Credit: Prabal Sarkar)

Tail wave

Tail wave is seen in affiliative contexts, such as females playing with infants. The tail waves from side to side, with small or large movements.

Postures are also important in the recognition of agonistic/aggressive and affiliative behaviours (see Expressions and Social behaviour).

A diagram showing various postures of rhesus macaques
A diagram showing various postures of rhesus macaques. See below:

Key to letters in the figure above:

A to B – Movement from a high to a low position, indicating threat. An attack by a confident animal may begin with a bobbing of the head accompanied by an open mouth directed towards the victim, grade into a lunging of the shoulders toward the victim, and finally become a charge ending when the dominant individual bites the victim.
C and D – Examples of subordinate posturing. The victim of an attack may attempt to present its hind quarters and grimacing face towards the attacker simultaneously, so that the animal’s body is displayed laterally to the attacker.
E to F – Upward jerking of the head, seen during non-hostile, non-fearful encounters. The upward jerking display may frequently be accompanied by protrusion and smacking of the lips.
G – Neutral sitting posture.
H – One individual of a wrestling pair. Rotation of the head and torso is often seen during wrestling play behaviour.
I to L – Oblique bobbing movements of a querying animal, often directed towards other monkeys, humans or novel objects. The open mouth face of the monkey in figure K indicates an aggressive component.

(Adapted from Sade 1973).


Macaques have a wide repertoire of vocalisations used in a variety of social contexts. Understanding the meaning behind vocalisations can be crucial for effective colony management e.g. for identifying aggression within a group and then intervening.

Rhesus macaque

Listen to all vocalisations

You can find a playlist of all of the behavioural vocalisations for rhesus macaques shown below on Soundcloud.

Affiliative vocalisations

These vocalisations signal peaceful intentions and help maintain contact with group members. They are used when approaching higher-ranked animals and infants, approaching to groom another animal, and during group movements [16]. They often accompany lip smacking (see Expressions).


Around 400 ms in duration, coos are characterised by their harmonic structure. They are relatively quiet calls, with a variation on an OOOH sound.

Coo spectogram
Coo spectogram


Around 200 ms in duration, grunts are characterised by their noisy, time modulated structure.

Grunt spectogram
Grunt spectogram


Girneys are high-pitched, soft, sing-song vocalisations used by adult female rhesus macaques to establish friendly contact with infants which are not their own offspring [17].

Threatening vocalisations

These vocalisations are used to signify dominance to lower ranking animals, to enhance visual signals during aggressive, threat displays (see Expressions), and to scare away potential threats, such as predators [18].


Often the animal will have its head down and ears back, suggesting a degree of fear within the threatening situation [9].


Around 300 ms in duration, pant threats are characterised by their noisy and threatening sounding structure. They are often seen with head raised and ears forward, suggesting increased intention to attack. 

Bark pant threat spectogram
Bark pant threat spectogram

Submissive or alarm vocalisations

These vocalisations may be involuntarily produced as an immediate reaction to surprise, in response to pain or fear, or to appease aggressors. They are given to aggressive or dominant animals, potential threats, and/or alarming or painful stimuli.

Squeak / Alarm call

Often heard in conjunction with grimace (see Expressions), crouch (see Postures) and retreat [19,20].

Screech, scream and squeal

Loud, high pitched calls, often heard in conjunction with grimace (see Expressions), crouch (see Postures) and retreat.

Other vocalisations

Warble, harmonic arch and chirp

Around 500 ms in duration, warbles, harmonic arches and chirps are characterised by their harmonic and contoured frequencies that increase toward the middle or end of the call. They are possible food calls, given when high-quality, rare food items are found. 

Harmonic arch spectogram
Harmonic arch spectogram


Geckers area made by infants during weaning, when mothers refuse attempts to nurse [9,18].

Copulation call

In macaques, both males and females can produce vocalisations during or immediately following the completion of copulation. These are more common in cynomolgus macaques than in rhesus macaques [22].

Cynomolgus macaque


Description / acoustic structure


Kra-(c) [23]

Extremely common call, occurring in two forms. Kra-(c) comprises three or four more or less discrete sound pulses that appear on sonograms as vertical “columns”. Harsh, rasping quality [20].

Appears to act as mild form of threat, but occurs in a wide variety of contexts, including agonistic interactions.


Kra-(a) comprises two to four separate sound pulses, similar in duration to kra-(c) but higher in frequency (pitch).

Appears to function as an alert signal, by attracting the attention of nearby monkeys to modifications in the environment. Often accompany alarm calling. Can also occur during agonistic interactions.


Alarm 1 is a modification of the basic kra-(c) call, comprising three to five sound pulses. Alarm 2 sounds more tonal or “chirp-like” to humans than the alarm 1 call, probably because call energy is organized into five to eight narrow, horizontal “bands”.

Both types of alarm calls are performed repetitively and quite loudly by noticeably aroused and highly agitated macaques (e.g. in the presence of predators)


Moderately loud call composed of five to seven sound pulses, and longer than alarm 1 call. The pulse units of the harr are not delivered plosively, hence it sounds ‘growl-like’ to humans.

Emitted primarily by juvenile macaques. Usually performed when threatened by another monkey during agonistic interactions that may or may not involve physical contact (similar to the “geckering screech” of rhesus macaques) [24]. Juveniles playing with each other perform harr calls, especially as play becomes rambunctious and grades into aggression.


Kra calls grade into barks, which vary in shrillness depending on the amount of energy in the higher-frequency range.

Associated with agonistic interactions, particularly those involving adult males. Vocal threat via barking sometimes substitutes for aggressive chasing or physical contact. Bark-kras occur in a much wider variety of contexts than barks, including intra-group agonism.


Consists of an initial, broadband and characteristically “harsh” segment (the “kra”) followed by a somewhat shorter “trailing off” of call energy in the lower frequencies (the “hoo”). The loudest call in the cynomolgus macaque repertoire. Usually performed in series with pauses in between. Kra component often accompanied by rhythmic, abrupt jerking of the head and upper body.

Performed by adult males only. Possibly functions in inter-group spacing.


Comprises both tonal and harsh components, increasing in amplitude.

Performed by both sexes and all ages when agitated. Isolated, solitary macaques perform Wraggh especially frequently, so this call may function in re-establishing social/spatial relations.


Loud, usually plosive vocal discharges whose total energy is distributed over a wide range of frequencies between 1 and 16 kHz. The acoustic structure varies, as with rhesus macaques [25].

Performed by both sexes and all ages engaged in agonistic interactions.

Khreeet screech

The khreeet screech is a loud, partially tonal scream, increasing then decreasing in frequency, similar to the “arched scream” or “screech” of rhesus macaques [24,25]. Often occurs in long, repetitive series with whimper calls.

Given by distressed and agitated young macaques (juveniles and possibly older infants) in both agonistic and non-agonistic contexts.


Low-amplitude, high-pitched calls composed of multiple units of often dissimilar form and that characteristically experience marked changes in pitch.

Relative rare calls. Performed by young macaques only.

Copulation calls

Females give a staccato “ohoh” copulation call. A weak high-pitched bleep” is also performed infrequently by males during copulations [26].



Relatively quiet, tonal calls.

Given when calm and not agitated.


A specific, consistently heard patterning of coo call units analogous to the combining of barks to form the “pant threat” of rhesus macaques [7]. Contains a distinctive, pronounced upward modulation in frequency that achieves its highest peak in the last two thirds of the call.

Performed by distressed infant or adolescent macaques, such as young juveniles left behind by their mothers. Young monkeys engaged in tantrum behaviour also perform these calls. May be accompanied by screams or khreeet screeches, to which mothers usually respond with retrieval.


Macaques are social primates and interactions with conspecifics are an important part of their daily lives. Social behaviour varies with sex, age, reproductive status and dominance rank.



One animal picks through the hair of another with hands or teeth, removing skin, dirt or ticks [7]. Allo-grooming is used to develop and maintain bonds between individuals. Females form stable life-long attachments, maintained through grooming, touch and close proximity [28]. Males groom females more during the mating season [29,30]. Macaques may groom after conflicts to console each other or repair damaged relationships [31-34]. Both giving and receiving grooming releases B-endorphins; after conflict situations, these may reduce pain from injury and relieve stress more quickly [35-37].

In this video, a female grooms a males back.
Female rhesus macaque grooms its group mate whilst resting together at night. (Credit: Claire Witham)


Affiliative contact

Macaques often rest and sleep huddled in family groups; they will also huddle when under threat. In behavioural studies, affiliation is often measured as proximity (e.g. within an arm’s reach of each other), because macaques that are friendly with each other will remain close, to maintain bonds and provide assistance if needed. The neuromodulators oxytocin and serotonin are implicated in development and maintenance of social behaviours such as approach and close contact [38-41]. These contribute to a sense of security from being close to conspecifics and have health benefits, such as maintaining low heart rate.

In this video, a group of macaques are seen resting in contact with one dyad engaging in social grooming (Credit: Jade Brooks and David Massey).


Rhesus macaques sleeping in close contact
Rhesus macaques sleeping in close contact (Alexander Georgiev)

Present for grooming

An animal will approach or reposition itself in front of another, presenting an area of the body to be groomed. Typically seen during peaceful contexts, or by individuals seeking comfort after a fight.

In this video, a macaque presents its tail for grooming by another group member. (Credit: MRC Centre for Macaques)



Mostly seen in infants, juveniles and adolescents. Play develops social bonds and motor-coordination skills, allowing younger animals to develop and assess their relative strength to peers. There are sex differences in play: females engage in more play with infants and older females; males engage in more rough-and-tumble play [42-44]. Adult males may engage older juveniles in play [20].

Two infant rhesus macaques, playing on the forest floor.
In this video, a young cynomolgus macaque plays by himself.


Infant care and handling

Infant care is performed mainly by nursing mothers during the birthing season. However, other group members are very interested in new infants. Looking after the infants of higher ranking females, either by a low-ranking female or male, will help improve their standing with the mother. The mother-infant bond in macaques is strong, and mediated by oxytocin released during birth and lactation throughout the first year of life [41]. It is important that infants are not removed from the mother before natural weaning has occurred, as this will lead to social attachment problems later in life [45].

In this video, a female rhesus macaque carries her infant dorsally. Female macaques tend to carry their infants either dorsally or ventrally (Credit: Jade Brooks and David Massey)


In this video, a female rhesus macaque cradles her infant while foraging (Credit: Jade Brooks and David Massey) 


In this video, a large adult male cynomolgus macaque cradles an infant. (Credit: BFC)


Genital present

When in oestrous and proceptive, a female will present her rump to the male to solicit copulation [46]. Outside of mating, both males and females may present their rump as a signal of subordination [47].

Male cynomolgus macaque inspects female. He crouches and places his head near her genitals.
Male cynomolgus macaque inspects female (Image: Michael Gumert)

Mount and copulate

Reproductive behaviours are triggered by seasonal changes in sex hormones (e.g. oestrogen, testosterone). These may be accompanied by reddening of sexual skin on the face, genitals and rump. Mating is highly seasonal in the wild [48,49]. In captivity, mating seasons may be less pronounced.

Rhesus macaque copulation (Credit: Alexander Georgiev)

Agonistic and aggressive


Displacement most often occurs during competition for valued resources such as food, mates, shelter and good resting places. The dominant animal approaches a subordinate, who then moves away to avoid potential conflict. Dominant animals exert their dominance in subtle ways, which helps to conserve energy. In captivity, distributing resources so that they cannot be monopolised by dominant animals reduces stress for subordinates.

In this video, a rhesus macaque is displaced by the alpha female.


In this video, an older rhesus macaque is displaced from her position on the shelf


In this video, we see cynomolgus macaques being displaced for their foraging positions.


Aggressive approach

Approaching with staring face and an aggressive stance, including lunging with the tail up, are signs of dominance. Aggressive approach is typically performed by a dominant animal towards a subordinate.

In this video, a dominant animal approaches a subordinate with high posture and tail up; the subordinate then runs away.


Dominant animals will chase lower ranked animals away from resources where there is high competition, and use aggression to gain access.

Rhesus macaques on Cayo Santiago, showing low ranked animals being chased into the sea.


In this video, three rhesus macaques are in an agonistic encounter with another. Two of the three then proceed to chase the recipient into the wooded area (Credit: Jade Brooks and David Massey).

Slap ground

Threat action, often accompanied by open mouth stare. For videos of slap ground, see the RHVIDEO site.


Threat action

In this video, the alpha male lunges at members of his group in order to move them away from food he wishes to take for himself. (Credit: BFC)

Branch shake

Size and aggression display to drive away potential threats. Most frequently given by males in response to other males, humans or perceived threats. In captivity, macaques will sometimes shake the home cage structure [7].

An adult male rhesus macaque shakes a resting platform is a display of strength. (Credit: MRC Centre for Macaques)

Dominant mount

An animal may exert its dominance over another by mounting it. Dominant mounting often occurs during tense situations, as dominant animals seek to re-affirm the hierarchy.


Push, Grab, Fight/Wrestle, Bite, Hit

Physical contact behaviours which can result in injury only occur when aggressive interactions cannot settled by non-contact means.

In this video, a rhesus macaque female playing with water in a pool repeatedly pushes away a smaller member of the group.


The macaque on the left attempts to grab food from the feeding animal on the right. (Credit: MRC Centre for Macaques)


Flighting cynomolgus macaque males. One is lunging towards another while they display teeth
Flighting cynomolgus macaque males (Image: Michael Gumert)


Threat action

Rhesus macaques resting together on a raised platform begin to stare at the camera operator. (Credit: MRC Centre for Macaques)

Submissive and fearful


Subordinates will avoid or move away from dominant or aggressive animals, to avoid physical contact which may result in injury.

Rump present

Rump present indicates submission, performed by subordinates towards dominants [11]. Females may also present their rumps to males to solicit copulation.

Lean away

During potentially threatening social situations, macaques may lean away from a conspecific to avoid physical contact which may result in injury. For video of lean away, see the RHVIDEO site.

Look away

A subordinate will turn its back to a dominant individual to avoid eye contact and potential aggression.

A macaque turns its back to hide its face from another animal. Dominant mount is also seen in this video.


Flee is a fear response in which a macaque runs away from a perceived threat. Seen during aggressive encounters, in the presence of predators, and when surprised [7].


Freeze is a response in which a macaque stops all activity and remains still while assessing threat. Seen during aggressive encounters, in the presence of predators, and when surprised.

Non Social

Although social animals, macaques have a repertoire of non-social behaviours that is important they are able to express in a captive environment for optimum welfare.


Macaques are highly food motivated. In the wild they spend 20-40% of their day foraging for food, with variance between species and seasons [50]. In captivity, macaques will work harder to obtain preferred foods; they also value rewards that are hard to obtain.

These cynomolgus macaques foraging on the forest floor on Tinjil Island, Indonesia, call to each other to maintain peaceful contact.


These rhesus macaques foraging for small seeds and grains within the wood shavings continually monitor their surroundings. Note their position either side of the wooden visual barrier. (Video: MRC Centre for Macaques)


In the wild, over 20% of the day is spent moving between foraging and resting sites and approaching and avoiding other macaques [50]. Macaques are opportunists that have adapted to live in a greater range of habitats than any other primate except man. Daily path length varies from 1-3.5 km depending on the habitat [51].


Self-grooming (also called auto-grooming) aids an animal in maintaining its own health, without relying on a conspecific. Excessive self-grooming or scratching is a sign of underlying stress (see Behavioural signs of poor welfare). Grooming removes parasites and dirt or attends to wounds.

  • A young rhesus macaque perches on fire hose rope enrichment while it's hand is reaching around it's back to self scratch
    A young rhesus macaque perches on fire hose rope enrichment
In this video, a juvenile is seen self-scratching its back whilst sitting and foraging on the ground. Excessive self-scratching can be an indicator of negative welfare (Credit: Jade Brooks and David Massey).
In this video, a cynomolgus macaque is self-grooming


Wild macaques rest for around 17-26% of the day, with some seasonal variance [52,53]. It is important to allow macaques time to rest, and to not continually stimulate them in captivity. Macaques rest after periods of activity such as foraging and socialising.

  • A cynomolgus macaque lies ventrally on a wooden perch. It rests it's head on both of it's hands.
    A cynomolgus macaque lies ventrally on a wooden perch
These cynomolgus macaques are resting in the trees. The macaque on the right makes a coo call in response to a coo from another group member.


A female rhesus macaque resting on a wooden shelf in her enclosure (Credit: MRC Centre for Macaques)



Being alert in social situations prepares an animal to move away quickly or towards something. It also aids in making them look larger and more threatening, and can allow for a better view of conspecifics, humans, or the surrounding environment.

In this video, rhesus macaques are foraging and standing alert (Credit: MRC Centre for Macaques).


Vigilance helps macaques to avoid predators and keep an eye on the activity of conspecifics and other animals. It is often noticeable in foraging and resting animals.

This macaque remains vigilant, scanning its environment, while feeding.


Watch what happens to one of these macaques whose attention is solely focused on grooming the infant.

Swimming and diving

Swimming and diving have multiple purposes in macaques. They engage in these activities for locomotion, play and access to food across bodies of water, as well as for foraging underwater [54,55]. Macaques can swim from birth [56] and will enjoy playing and swimming in water if provided with the opportunity in captivity (see Sensory).

In this video, rhesus macaques are seen swimming and diving into a pool that was created after a period of rainfall (Credit: Jade Brooks and David Massey).


Dive bomb

Object handling, solitary play and tool use

Stone handling may enhance long-term neural and cognitive development in younger macaques, and maintain or repair neural pathways in older macaques [57,58]. Cynomolgus macaques have been observed using stone tools to open bivalves, nuts and sea snails [59], and to wash and rub clean certain foods, including potatoes and roots; they will also peel sweet potatoes [60]. In impoverished captive conditions, a desire to manipulate objects may be manifested in manipulation of the cage fixings.

In this video, a cynomolgus macaque is handling a stone (Credit: Michael Gumert).


In this video, a macaque is using a string like object to teeth floss, a rarely seen behaviour (Credit: Michael Gumert).


Abnormal behaviours in macaques can be qualitatively abnormal (those that occur in captivity but not in natural settings, such as stereotypies) or quantitatively abnormal (those that occur more/less often in captivity than in nature, such as excessive grooming, hair plucking or scratching). It is important to recognize that inactivity (doing nothing) may also be an abnormal behaviour.

Information on using abnormal behaviours to assess welfare, and preventing abnormal behaviour, is given in the Behavioural indicators section.

Hair plucking is one sign of abnormal behaviour:


In this video, a cynomolgus macaque can be seen plucking and ingesting its hair.


Macaques diets are highly varied consisting of a variety of foods that are readily available to provide them a naturalistic diet in a captive setting.

Macaques diets are highly varied:

  • Rhesus macaques eat fruits, young and mature leaves, stems, seeds, flowers, petioles, bark and roots of over 75 different plant species [62].
  • Cynomolgus macaques are predominantly frugivorous (fruit accounts for 67%-82% of their diet) [63,64], but also feed on other plant parts. Foraging is typically conducted below 20 m (65.6 ft); usually around 12 m (39.4 ft) in the lower canopy, understory, and on the ground [65]. At mangroves they have also been seen to consume bivalves, shrimp and octopus [66,67].
  • Both species feed opportunistically on grass, clay, mushrooms, invertebrates, eggs, crabs, and small vertebrates such as lizards, frogs, birds and fish [68].

 Non-exhaustive list of plant species and their component parts eaten by rhesus macaques

(Adapted from Zhou et al., 2014)

Plant: Latin name 

Plant: Common name 

Fruit eaten 

Stems eaten 

Young leaves eaten 

Mature leaves eaten 

Aristolochia longgangensis 

Dutchman’s pipe, Pipevine, Birthwort 





Capparis cahtohiesis 






Carvota ochlandra 

Chinese fishtail palm 





Clausena anisum 

Anis (Philippines) 





Clausena emarginata 

Powderpuff plant, Cat’s tail 





Cuscuta chinensis 

Cuscuta, Chinese dodder 





Dracontomelon duperreanum 






Embelia scandens 






Ficus gibbosa 






Ficus glaberrima 






Ficus macrocarpa 

Curtain fig, Chinese banyan, Indian laurel 





Ficus obscura 

Fig, Appolas 





Indocalamus calcicolus 






Iondes ovalis 






Pithecellobium clypearia 

Greater grasshopper tree 





Pothos repens 






Polygonum chinense 

Chinese knotweed 





Pueraria thunbergiana 

Kudzu, Japanese arrowroot 





Spondia lakonensis 






Urobotrya latisquama 






Diet composition of cynomolgus macaques

(Adapted from Brotcorne 2014)


Origin of food (%) 

Fruit (%) 

Leaves (%) 


Animal matter (%) 

Other (%) 








Natural (86) and anthropogenic (14) 




Natural (51) and anthropogenic (49) 







Natural (74) and anthropogenic (26) 






Kalimantan. Borneo 







Kalimantan. Borneo






Ubud, Bali

Natural (48) and anthropogenic (52) 






Ubud, Bali

Natural (30) and anthropogenic (70) 






TNBB, Bali

Natural (89) and anthropogenic (11) 





Uluwatu, Bali

Natural (52) and anthropogenic (48) 







Natural (75) and anthropogenic (25) 







Natural and anthropogenic 






  1. Jaman MF and Huffman MA (2013). The effect of urban and rural habitats and resource type on activity budgets of commensal rhesus macaques (macaca mulatta) in bangladesh. Primates 54(1): 49-59.
  2. Zhou Q et al. (2014). Niche separation of sympatric macaques, macaca assamensis and m. Mulatta, in limestone habitats of nonggang, china. Primates 55(1): 125-37.
  3. Li D et al. (2012). Sex-age differences in activity budget and position behavior of rhesus macaques (macaca mulatta). Acta Theriologica Sinica 32(1): 25-32.

  4. Hambali K et al. (2012). Daily activity budget of long-tailed macaques (macaca fascicularis) in kuala selangor nature park. International Journal of Basic & Applied Sciences 12(4): 47-52. 

  5. Majolo B et al. (2013). The effect of climatic factors on the activity budgets of barbary macaques (macaca sylvanus). International Journal of Primatology 34(3): 500-14.

  6. Mallapur A et al. (2005). Factors influencing the behaviour and welfare of captive lion-tailed macaques in indian zoos. Applied Animal Behaviour Science 91(3-4): 337-53.

  7. Hinde RA and Rowell TE (1962). Communication by postures and facial expressions in the rhesus monkey (macaca mulatta). Proceedings of the Zoological Society of London 138(1): 1-21.

  8. Parr LA et al. (2010). Brief communication: Maqfacs: A muscle-based facial movement coding system for the rhesus macaque. American Journal of Physical Anthropology 143(4): 625-30.

  9. Partan SR (2002). Single and multichannel signal composition: Facial expressions and vocalizations of rhesus macaques (macaca mulatta). Source: Behaviour 139(8): 993-1027.

  10. Rowe N (1996). The pictorial guide to the living primates. Pogonias Pr.

  11. Maestripieri D (1999). Primate social organization, gestural repertoire size, and communication dynamics: A comparative study of macaques. In: The evolution of language: Assessing the evidence from nonhuman primates (Ed., King BJ) (pp. 55-77). School of American Research.

  12. Maestripieri D (2005). Gestural communication in three species of macaques (macaca mulatta , m. Nemestrina , m. Arctoides). Gesture 5(1-2): 57-73.

  13. Moynihan M (1976). The new world primates: Adaptive rediation and the evolution of social behavior, languages, and intelligence. Princeton University Press.

  14. Menzel CR (1980). Head-cocking and visual perception in primates. Animal Behaviour 28(1): 151-IN10.

  15. Cantalupo C et al. (2001). Function of head-cocking in garnett’s greater bush baby (otolemur garnettii). International Journal of Primatology 11(1).

  16. Hauser MD (1998). Functional referents and acoustic similarity: Field playback experiments with rhesus monkeys. Anim Behav 55: 1647-58. 

  17. Whitham JC et al. (2007). Intended receivers and functional significance of grunt and girney vocalizations in free-ranging female rhesus macaques. Ethology 113(9): 862-74.

  18. Lindburg DG (1971). The rhesus monkey in north india : An ecological and behavioral study. In: Primate behavior: Developments in field and laboratory research (Ed., Rosenblum LA) (Vol. 2, pp. 1-106). Academic Pr.

  19. Palombit RA (1992). A preliminary study of vocal communication in wild long-tailed macaques (macaca fasc/adar/s). I. Vocal repertoire and call emission. International Journal of Primatology 13. 

  20. Wheatley BP (1999). The sacred monkeys of bali, 1st edition. Waveland Press.

  21. Lindburg DG (1971). The rhesus monkey in north india : An ecological and behavioral study. In: Primate behavior: Developments in field and laboratory research (Ed., Rosenblum LA) (Vol. 2, pp. 1-106). Academic Pr.

  22. Maestripieri D and Roney JR (2005). Primate copulation calls and postcopulatory female choice. Behavioral Ecology 16(1): 106-13.

  23. Raffles TS (1821). Xvii. Descriptive catalogue of a zoological collection, made on account of the honourable east india company, in the island of sumatra and its vicinity, under the direction of sir thomas stamford raffles, lieutenant-governor of fort marlborough; with addi. Transactions of the Linnean Society of London 13(1): 239-74.

  24. Rowell T (1962). Agonistic noises of the rhesus monkey (macaca mulatta). Sym Zool Soc London 8: 91-6.

  25. Gouzoules S et al. (1984). Rhesus monkey (macaca mulatta) screams: Representational signalling in the recruitment of agonistic aid. Animal Behaviour 32(1): 182-93.

  26. van Noordwijk MA (2010). Sexual behaviour of sumatran long-tailed macaques (macaca fascicularis). Zeitschrift für Tierpsychologie 70(4): 277-96.

  27. Augustsson H and Hau J (1999). A simple ethological monitoring system to assess social stress in group-housed laboratory rhesus macaques. Journal of Medical Primatology 28(2): 84-90.

  28. Massen JJM and Sterck EHM (2013). Stability and durability of intra- and intersex social bonds of captive rhesus macaques (macaca mulatta). International Journal of Primatology 34(4): 770-91.

  29. Rakhovskaya MV (2013). Correlates of male consortship rate in free-ranging rhesus macaques (macaca mulatta). International Journal of Primatology 34(4): 662-80.

  30. Gumert MD (2007). Grooming and infant handling interchange in macaca fascicularis: The relationship between infant supply and grooming payment. International Journal of Primatology 28(5): 1059-74.

  31. Cords M (1992). Post-conflict reunions and reconciliation in long-tailed macaques. Animal Behaviour 44(1): 57-61.

  32. Aureli F and Schaik CPv (2010). Post-conflict behaviour in long-tailed macaques (macaca fascicularis). Ethology 89(2): 89-100.

  33. Demaria C and Thierry B (2001). A comparative study of reconciliation in rhesus and tonkean macaques. Behaviour 138(3): 397-410.

  34. Call J et al. (1999). Reconciliation patterns among stumptailed macaques: A multivariate approach. Animal Behaviour 58(1): 165-72.

  35. Russell YI and Phelps S (2013). How do you measure pleasure? A discussion about intrinsic costs and benefits in primate allogrooming. In Biology and Philosophy (Vol. 28, pp. 1005-20).

  36. Dunbar RIM (2010). The social role of touch in humans and primates: Behavioural function and neurobiological mechanisms. Neuroscience & Biobehavioral Reviews 34(2): 260-8.

  37. Shutt K et al. (2007). Grooming in barbary macaques: Better to give than to receive? Biology Letters 3(3): 231-3.

  38. Simpson EA et al. (2014). Inhaled oxytocin increases positive social behaviors in newborn macaques. Proceedings of the National Academy of Sciences 111(19): 6922-7.

  39. Winslow JT et al. (2003). Rearing effects on cerebrospinal fluid oxytocin concentration and social buffering in rhesus monkeys. Neuropsychopharmacology 28(5): 910-8.

  40. Chang SWC et al. (2012). Inhaled oxytocin amplifies both vicarious reinforcement and self reinforcement in rhesus macaques (macaca mulatta). Proceedings of the National Academy of Sciences 109(3): 959-64.

  41. Higham JP et al. (2011). Mu-opioid receptor (oprm1) variation, oxytocin levels and maternal attachment in free-ranging rhesus macaques macaca mulatta. Behavioral Neuroscience 125(2): 131-6.

  42. Hinde RA and Spencer-Booth Y (1967). The behaviour of socially living rhesus monkeys in their first two and a half years. Animal Behaviour 15(1): 169-96.

  43. Brown GR and Dixson AE (2000). The development of behavioural sex differences in infant rhesus macaques (macaca mulatta). Primates 41(1): 63-.

  44. Hassett JM et al. (2009). Social segregation in male, but not female yearling rhesus macaques (macaca mulatta). American Journal of Primatology:.

  45. Prescott MJ et al. (2012). Laboratory macaques: When to wean? Applied Animal Behaviour Science 137(3-4): 194-207.

  46. Estes RD (1991). The behavior guide to African mammals. University of California Press.

  47. Wallen K et al. (1984). Periovulatory changes in female sexual behavior and patterns of ovarian steroid secretion in group-living rhesus monkeys. Hormones and Behavior 18(4): 431-50.

  48. Rowell TE (1963). Behaviour and female reproductive cycles of rhesus macaques. Reproduction 6(2): 193-203.

  49. Dunk RDP (2013). Seasonality of conceptions under varying conditions in a rhesus macaque breeding colony. Thesis. University of Wisconsin-Milwaukee

  50. Hambali K et al. (2012). Daily activity budget of long-tailed macaques (macaca fascicularis) in kuala selangor nature park. International Journal of Basic & Applied Sciences 12(4): 47-52.

  51. Fooden J (2000). Systematic review of the rhesus macaque, macaca mulatta (zimmermann, 1780) / jack fooden  1 ed. Field Museum of Natural History.

  52. Xu F et al. (2012). Construction and Validation of a Systematic Ethogram of Macaca fascicularis in a Free Enclosure. PLOS ONE 7(5): e37486. doi:10.1371/journal.pone.0037486

  53. Jaman MF and Huffman MA (2013). The effect of urban and rural habitats and resource type on activity budgets of commensal rhesus macaques (macaca mulatta) in bangladesh. Primates 54(1): 49-59.

  54. Dunbar DC (1989). Locomotor behavior of rhesus macaques (macaca mulatta) on cayo santiago. Puerto Rico health sciences journal 8(1): 79-85.

  55. Dunbar DC (1989). Locomotor behavior of rhesus macaques (macaca mulatta) on cayo santiago. Puerto Rico health sciences journal 8(1): 79-85.

  56. Riopelle AJ and Hubbard DG (1982). Vestibular function and development of swimming behavior in rhesus monkeys. Journal of Human Evolution 11(5): 377-82.

  57. Nahallage CAD and Huffman MA (2012). Macaque–human interactions in past and present-day sri lanka. In: The macaque connection. Developments in primatology: Progress and prospects (Eds. Radhakrishna S, Huffman MA, and Sinha A) (Vol. 43, pp. 135-48). Springer.

  58. Nahallage CAD and Huffman MA (2008). Comparison of stone handling behavior in two macaque species: Implications for the role of phylogeny and environment in primate cultural variation. American Journal of Primatology 70(12): 1124-32.

  59. Gumert MD et al. (2009). The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the andaman sea region of thailand. American Journal of Primatology 71(7): 594-608.

  60. Wheatley BP (1980). Feeding and ranging of east bornean macaca fascicularis. In: The macaques: Studies in ecology, behavior, and evolution (Ed., Lindburg DG) (pp. 215-46). Van Nostrand Reinhold.

  61. Erwin J and Deni R (1979). Strangers in a strange land: Abnormal behaviors or abnormal environments? In: Captivity and behavior: Primates in breeding colonies, laboratories and zoos (Eds. Maple TL, Erwin J, and Mitchell G) (pp. 1-28). Van Nostrand Reinhold.

  62. Zhou Q et al. (2014). Niche separation of sympatric macaques, macaca assamensis and m. Mulatta, in limestone habitats of nonggang, china. Primates 55(1): 125-37.

  63. Wich SA (2002). Measuring fruit patch size for three sympatric indonesian primate species. Primates 43: 19-27. 

  64. Ungar PS (1996). Feeding height and niche separation in sympatric sumatran monkeys and apes. Folia Primatologica 67(3): 163-8.

  65. Sussman RW and Tattersall I (1986). Distribution, abundance, and putative ecological strategy of macaca fascicularis on the island of mauritius, southwestern indian ocean. Folia Primatologica 46(1): 28-43.

  66. Son VD (2003). Diet of macaca fascicularis in a mangrove forest, vietnam. Laboratory Primate Newsletter 42(4): 1-5.

  67. Wheatley BP (1980). Feeding and ranging of east bornean macaca fascicularis. In: The macaques: Studies in ecology, behavior, and evolution (Ed., Lindburg DG) (pp. 215-46). Van Nostrand Reinhold.