Behaviour

In this guide

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. 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). Data for rhesus macaques, averaged from Jaman & Huffman 2013 and Zhou et al. 2013

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].

Neutral

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.

Neutral face

Affiliative

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.

Macaque affiliative face

Lip smack interaction (Michael Gumert)
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.

Infant play face (Michael Gumert)
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.

Female rhesus grimaces as she turns from a dominant male (Image: Alexander Georgiev)
A subordinate gives a bared teeth display to an approaching dominant (Image: Gumert)
Bared teeth display with tongue protrusion in cynomolgus macaque (Image: Michael Gumert)
A young cynomolgus macaque performs a bared teeth display (Image: Michael Gumert)

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)

Anxious

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.

Yawn

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.

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 (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.

Open mouth stare (Image: Michael Gumert)
Open mouth stare
Intense open mouth stare
Cynomolgus macaque stare (Image: Michael Gumert)

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.

Piloerection

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.

Piloerection in a male rhesus macaque (Image: MRC Centre for Macaques)
Piloerection during aggressive interaction (Image: Michael Gumert)
Piloerection in a cynomolgus macaque (Image: Michael Gumert)

Head-cock

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

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. (Credit: Michael Gumert)
Cynomolgus macaque crouching, with bared teeth indicating submissiveness (Credit: Michael Gumert)

Freeze

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 (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. 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).

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).

Coo

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

NC3RsComms · CBC Coo 10 (1)

Grunt

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

NC3RsComms · CBC Grunt 10

Girney

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 [1 7].

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 [1 8].

Bark

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

Pant-threat

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.

NC3RsComms · CBC PantThreat 4

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.

NC3RsComms · Scream

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.

NC3RsComms · CBC HarmonicArch 1

Gecker

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 [2 2].

Cynomolgus macaque

Call	Description / acoustic structure	Context
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)	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	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)
Harr	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.
Bark	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.
Krahoo	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.
Wraggh	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.
Scream	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.
Squeal	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].	Copulation.
Coo	Relatively quiet, tonal calls.	Given when calm and not agitated.
Whimper	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.

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:

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	Yes	Yes	No	No
Capparis cahtohiesis	–	Yes	No	Yes	Yes
Carvota ochlandra	Chinese fishtail palm	Yes	No	Yes	No
Clausena anisum	Anis (Philippines)	Yes	No	No	No
Clausena emarginata	Powderpuff plant, Cat’s tail	Yes	No	Yes	Yes
Cuscuta chinensis	Cuscuta, Chinese dodder	No	Yes	No	No
Dracontomelon duperreanum	Yanmin	Yes	No	Yes	No
Embelia scandens	Mez	No	No	Yes	Yes
Ficus gibbosa	Fig	Yes	No	Yes	Yes
Ficus glaberrima	Fig	Yes	No	No	Yes
Ficus macrocarpa	Curtain fig, Chinese banyan, Indian laurel	Yes	No	Yes	Yes
Ficus obscura	Fig, Appolas	Yes	No	No	No
Indocalamus calcicolus	Bamboo	No	No	Yes	No
Iondes ovalis	–	Yes	No	Yes	Yes
Pithecellobium clypearia	Greater grasshopper tree	No	No	Yes	Yes
Pothos repens	–	Yes	No	Yes	No
Polygonum chinense	Chinese knotweed	Yes	No	No	No
Pueraria thunbergiana	Kudzu, Japanese arrowroot	No	Yes	Yes	Yes
Spondia lakonensis	–	Yes	No	Yes	No
Urobotrya latisquama	Hiepko	Yes	No	Yes	Yes

Diet composition of cynomolgus macaques

(Adapted from Brotcorne 2014)

Location	Origin of food (%)	Fruit (%)	Leaves (%)	Flowers (%)	Animal matter (%)	Other (%)
Malaysia	Natural	52.4	19	5.4	23.2	0
Singapore	Natural (86) and anthropogenic (14)	44	8	7	0	41
Singapore	Natural (51) and anthropogenic (49)	20.9	21	6.3	0	51.8
Singapore	Natural (74) and anthropogenic (26)	44.8	19.6	9.7	0	28.3
Kalimantan. Borneo	Natural	66.7	17.2	8.9	4.1	3.1
Kalimantan. Borneo	Natural	87	4	3	4	2
Ubud, Bali	Natural (48) and anthropogenic (52)	32	34	5	29	0
Ubud, Bali	Natural (30) and anthropogenic (70)	25.7	14.7	0	7.5	52.1
TNBB, Bali	Natural (89) and anthropogenic (11)	60	21	0	3	16
Uluwatu, Bali	Natural (52) and anthropogenic (48)	32	19	0	14	35
Vietnam	Natural (75) and anthropogenic (25)	15	20	1	35	29
Mauritius	Natural and anthropogenic	42.2	30.8	7	2.4	17.6

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Overview

Activity

Expressions

In this section

Neutral

Affiliative

Lip smack

Play face

Submissive or fearful

Grimace/bared teeth

Anxious

Tense mouth

Yawn

Chin up

Agonistic or aggressive

Open mouth stare

Postures

In this section:

Piloerection

Head-cock

Crouch

Freeze

Tail up

Tail wave

Vocalisations

In this section

Rhesus macaque

Listen to all vocalisations

Affiliative vocalisations

Coo

Grunt

Girney

Threatening vocalisations

Bark

Pant-threat

Submissive or alarm vocalisations

Squeak / Alarm call

Screech, scream and squeal

Other vocalisations

Warble, harmonic arch and chirp

Gecker

Copulation call

Cynomolgus macaque

Social

In this section

Affiliative

Allo-grooming

Affiliative contact

Present for grooming

Play

Infant care and handling

Sexual

Genital present

Mount and copulate

Agonistic and aggressive

Displacement

Aggressive approach

Chase

Slap ground

Lunge

Branch shake

Dominant mount

Push, Grab, Fight/Wrestle, Bite, Hit

Stare

Submissive and fearful

Avoidance

Rump present

Lean away

Look away

Flee

Freeze

Non Social

In this section

Foraging/eating

Locomotion

Self-grooming/scratching

Inactive/sleeping

Stand

Vigilance