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Species: Liontail Macaque (Macaca Silenus)

Use: General behavior

Authors: Samuel W. Skinner and Joan S. Lockard, Department of Psychology and Neurological Surgery, University of Washington

Methods: N/A

Publications/Presentations:Ê S.W. Skinner, J.S. Lockard (1979). Title: An Ethogram of the Liontail Macaque (Macaca Silenus) in captivity, Applied Animal Ethology, 5, 241:253.Affiliative Behavior

Huddle: Most often a huddle was composed of mature females and their infants, though huddles of up to 8 animals at a time were observed. The alpha male was rarely seen to join a huddle; rather he would sit by himself or would be joined by another animal- usually a mature female.

Embrace: Ventral- ventral hugging, were observed only twice. Once, two females embraced while the cage was being sprayed for insects. Another time, one female hit another of lower rank and then immediately embraced her. The behavior appears to serve as a reassurance gesture as it does in the stump-tail macaque (Bertrand, 1969).

Nuzzle: Nuzzling in the liontail is an affiliative behavior directed primarily towards mothers and their infants. In the stumptail it functions as an appeasement behavior (Blurton-Jones and Trollope, 1968). Its role in the bonnet macaque (Macaca radiata) is rather different, for Simonda (1965) found that nuzzling was a certain indication of dominance on the part of the performing animal.

Crouch: Crouching was a common accoutrement of nuzzling when the latter was directed at an infant. It may have served merely to place the directing animal at the eye level of the infant, but when concurrent with head bobbing, it appeared to have a more definite function.

Head bob, with crouch: This occurred only with crouching behavior but was not an inevitable part of crouching. Sometimes an animal lip smacked with the pink of the eyelids exposed while head bobbing. Allomothers (ÒauntsÓ) were the animals most frequently seen to crouch and head-bob.

Head bobbing and crouching are affiliative in the liontail but in the stumptail and the vervet monkey (Cercopithecus aethiops), head bobbing is used in threat contexts (Bertrand, 1969; Struhsaker, 1967) and the crouch shows even greater variability. When it accompanies head bobbing in a vervet, it is a high intensity threat. In the rhesus, bonnet, and pigtail (M.nemestrina), it has been associated with fear (Hinde and Rowell, 1962; Kaufmen and Rosenblum, 1966). Hinde and Rowell describe crouching present where the rear is oriented towards the fear-provoking animal. Apparently both arms and legs are flexed as occurs in the affiliative crouch in the liontail.Ê A behavior similar to Hinde and RowellÕs crouch is present in the liontail. Here the animal presents in response to aggression while flexing the arms only. Often the animal will turn its head back towards the aggressing individual and display the silent bared-teeth face.

Self-grooming: Was similar in topography to that seen in other macaques (Chevalier-Skolnikoff, 1974).

Self-Inspection: Of the vulva occurred only just prior to parturition in the pregnant females, as has been described for pigtail and bonnet macaques (Rosenbaum, 1971).

Allogrooming: Was elicited by proximity, by sprawling, and, in the alpha male by touching or cuffing a female with his hand. It was sometimes accompanied by lip smacking or, more rarely, by the pout face. Grooming in macaques is generally thought to sub serve two functions, i.e. the removal of ectoparasites and the maintenance of social contacts (Hutchins and Barash, 1977). Its frequency of occurrence may vary from species to species, as can the principal participants involved. Thus, in the bonnet macaque it is quite common and individuals of all sex/ age classes (except the very young) participate in extended grooming bouts. In other macaque species, it is less common, e.g. the rhesus and Japanese macaques (Simonds, 1965). BertrandÊ (1969) presents quantitative data, which suggests that allogrooming occurs infrequently in the liontail She also notes that it was harder to elicit in this species than in the rhesus or stumptail.

The topography of grooming and invitations to groom appear to be the same in the liontail as in other macaques (Chevalier- Skilnikoff, 1974), as well as the situational contexts and sex/ age classes involved (adult females groom more than adult males and mostly between themselves) (Bertrand, 1969).

Mainly preliminary behaviors to sexual displays were seen during the study period as all three mature females were pregnant and gave birth during this time.

Inspection: consisted of a male putting or brushing aside the tail of a female and then visually inspecting her vulva and anus while fingering and sniffing them, This was seen once in the sub-adult male and several times in the alpha male.

Masturbation: was observed in the alpha male and once in the juvenile male. This consisted largely of tugging on the flaccid penis and rubbing it between the hands. Alternately, it might be held in one hand while being picked at with the other. If the penis became erect, the tip would be touched with the fingers, which were then sniffed and licked. Ejaculation was never observed.

Mounting: took a typical macaque form (Bertrand, 1969). With the exception of infants, it was seen to occur between all sex/age classes in all permutations. In juveniles it was occasionally incomplete, and, too, was sometimes misdirected by them - being oriented towards the face or hand of the mounted animal.

Facial displays accompanying agonistic behavior are described below. Agonistic behaviors include supplanting, hitting, grabbing, shaking, attacking (running at), biting and wrestling with an opponent. With the exception of supplanting, their incidence was low.

Threat leap: A threat leap was seen in the dominant male and had two orientations and perhaps two functions. On occasion when the immature had been playing vigorously for some time or when antagonism broke out between them, he would leap into the middle of the group in a forceful manner causing all to scatter. In its second form, he would leap forcefully against the cagewire following aggressive interactions with another adult male liontail in a neighboring cage. This leap against the cagewire produced a loud noise and was followed by a yawn. Struhsaker (1967) discussed a branch shaking display in the vervet, and one in several macaque species, which may be the feral equivalent of this display. The branch shaking display is used as a dominance display, according to Struhsaker, as is the threat leap discussed here.

Threat present: The threat present was observed a few times in the alpha male. It served to terminate aggressive encounters between him and the adult male in the neighboring cage. In performing this display he would ÒpresentÓ to the other male with his tail arched over his back towards his head. He would then lift his head high and wave it from side to side while making the forward bared-teeth face. He was oriented away from the recipient at all times so it is doubtful if the recipient could see this facial expression.

The threat present is similar in some respects to the Òred, white, and blueÓ display of the vervet monkey (Struhmaker, 1967) and the Òbacking threatÓ described by Hinde and Rowell (1962) for the rhesus. In the vervet, the red, white, and blue display is typically given by a dominant male who holds his tail erect and paces back and forth before a seated monkey, displaying his red perianus, his blue scrotum, and the white medial stripe of fur extending between the perianus and the scrotum. Similarly, in aggressive bouts between two rhesus, the dominant animal may show his hindquarters as a threat.

The sequence of play initiation, chasing, wrestling and play termination is common for many macaque species although different forms of play initiation are described and the extent to which different sex/ age classes play may vary from species to species.

Bouncing gait: The bouncing gait as a form of play initiation may be common to all macaques (Bertrand, 1969), while tail pulling, another form of play initiation, may be restricted to the longer tailed species such as the bonnet (Kaufman and Rosenblum, 1966) and the liontail. The relaxed opened-mouth face, often termed the play face, is discussed below.

Play biting: Liontail play consisted mostly of chases, wrestling and play biting. In play biting the roles of pursuer and pursued were often reversed. Simonds (1965) reports that in the bonnet macaque all sex-age classes were observed to play extensively, even the adult males. In the Celebes, this is also true, though the juveniles and adult females played more than did the adult males (Nickelson and Lockard, 1978). Play was frequent between the juveniles and sub-adults of both sexes in the liontail. Additionally, the alpha males regularly engaged in play although the bouts were shorter, less frequent, and were predominantly between him and the next oldest male. A female monkey was observed to play several times about three weeks after the theft of her infant. Bertrand (1969) presents data, which indicate that the liontail may play more frequently than either the stumptail or rhesus macaque.

Play was first observed in the one-month-old infant as a series if hopping motions that ay have been a rudimentary form of bouncing gait. Such is also the case on the bonnet (Simonds, 1965). The next stage consisted of mutual mouthings between two of the infants.

Observations were too intermittent to allow an accurate ontogeny of the infantsÕ behaviors to be described. However a number of infant specific and infant-related behaviors were observed.

Root: This occurred when an infant of up to three weeks lost the nipple. Then it moved its head from side to side until contact with the nipple was regained and nursing commenced. The infant spent many hours on the nipple without actual sucking motions being observed.

Kneading grasp: A kneading grasp was observed in the one and two two-day-old infants. The infant kneaded its motherÕs fur with its hands and/ or feet, until a firm grip was established.

Ventral carry: The above behaviors were generally seen when the infant was in a ventral carry position, the most common carrying position.

Clutch: Sometimes a mother or allomother would clutch an infant to her and walk bipedally. Often the infant was disoriented or even upside down when clutched.

Push start: If the infant was not in a ventral carry position when the mother started to move, or if its grip was lax, she would push start it by pushing on the infantÕs back, thus pressing it to her chest. The infant responded by clasping her fur. This was performed in the same manner by all three females and contrasts with the stumptail in which Chevalier-Skolnikoff (1974) reports several different forms of Òpush startÓ.

Ventral contact: Ventral contact with the infant sitting in loose ventral contact with its mother, appeared after the infantÕs first week. It was followed shortly by hand contact where the infant moved around the mother while keeping a grip on her. About the same time the mother began to restrain the infant or, when the infant did not break contact, to retrieve it.

Steal: Stealing occurred when an alloparent took, or attempted to take an infant from its mother. Usually the infant was retrieved at once, but if it were not it was passed from one alloparent to another or held by one and groomed.

In Macaca species with long tails, the tail presumably could play a greater communicative role than it plays in short-tailed species. It is more conspicuous and could perhaps take a greater number of discrete postures. However, this does not seem to be the case. In the liontail, the tail is used to tug or to swing on in play invitation, similar to the observation by Kaufman and Rosenblum (1966) for the bonnet monkey- another long-tailed species. Female alloparents in the liontail also used the tail to drag infants to them, or behind them, as they walk about the enclosure.

In its more communicative roles, the tail positions and their apparent meanings in the liontail are the same as, or similar to, those described in bonnet, pigtail (Kaufman and Rosenblum, 1966) and stumptail macaques (Bertrand, 1969). In the alpha male, the tail was held rigid arching away from the head. In all other group members it hung loosely from the rear. Occasionally, when aroused, the alpha maleÕs tail was arched towards his head. This was the position it took in the threat present. In a submissive present, the presenting animalÕs tail was usually arched to one side and away from its head. A straight tail, held pointing rigidly away from the head, was seen in the mature females in fear-provoking situations. Once it was seen in the alpha male and was accompanied by defecation. This occurred when intense fighting broke out between all three mature females ten days after the third infant was born. This tail display has not been described by other investigators.

The facial displays of macaques are numerous and show variability in their component elements between species, although interspecific uniformities for each display can be identified. The terminology of van Hooff (1967) is used here, when applicable, as his descriptions are among the most complete for each display. The relaxed and the alert faces were the most commonly seen; they represented a neutral backgroung from which other displays were distinguished.

Four displays with a strong aggressive tendency were observed in the liontail.

Tense mouth: The tense mouth face, as described by van Hooff, accompanied attack or merely threatened it. Its usage was similar to that described for the bonnet, the pigtail, and the stumptail macaques (Kaufman and Rosenblum, 1966; Bertrand, 1969).

Forward-bared teeth: The more common threatening facial display is termed the forward-bared teeth face, which has not been described in other species. The display consisted of continued or repeated baring of the teeth with the mouth open partly to fully. The head and lower jaw were frequently thrust toward the recipient. It was used by the alpha male in aggressive interactions, in the threat present, and to defend prized food items. The response by other animals was to present and lip smack, or the silent bared-teeth face.

Yawns: ÊYawns are another facial display not described by van Hooff; however, several other observers have discussed their functions. Of those referenced here, only Rosenblum and Kaufman (1966) suggest a non-aggressive function: as an expression of fatigue in bonnet and pigtailed macaques. In the liontail, the females were seen to yawn repeatedly following parturition and then, as Kaufman and Rosenblumm note, the canines were seldom fully exposed.

Yawning with an aggressive association may be divided into two categories, depending upon the orientation of the display. When a yawn is not directed at the other individuals it is usually termed a tension yawn (Bertrand, 1969). In the Celebes monkey, the liontail, and the stumptail, the ÒtensionÓ yawn is associated with dominance assertion- it was given once following a dominance display (Nickelson and Lockard,1978 ) in the first two species and was repeated a number of times in the stumptail(Bertrand, 1966). Use of the term ÒtensionÓ may be slightly inappropriate here as, especially in the liontail group, it was given in a situation when the alpha male was not challenged. However, in other contexts described in these species and in the bonnet, the pigtail and rhesus macaques (Kaufman and Rosenblum, 1966; Hinde and Rowell,ÉÉÉÉÉÉÉÉÉÉÉÉ)

Teeth-chatter: The teeth-chattering face appears in the liontail as an amalgam of the lip smacking face interspersed with the silent bared-teeth face. However, the mouth is widely opened in the liontail teeth chatter, whereas it is usually closed in the silent bared-teeth face. Additionally, the pink of the eyelids may be exposed in the teeth chattering face. It generally occurred following mild physical aggression directed at the performer (e.g. a light slap or cuffing) and thus appears to represent a mixture of appeasement and fearful threat behaviors. This context is similar to its employment in other macaques species (van Hooff, 1967).

Pout face: In the pout face, the mouth corners are brought forward and the lips pursed to form a small ÒoÓ shape. Van Hooff (1967) restricts its use to infants; however, Bertrand (1969) found that in the stumptail it could be used in greeting and in reassurance when a special relationship existed between the animals involved. Liontail infants were observed to give the pout face on a number of occasions, but it was also seen in the sub-adult male, as he allogroomed his mother. This suggests that it may- albeit rarely Ð occur in the liontail in the special contexts described by Bertrand.

Relaxed open mouth: The relaxed open-mouthed face or play face resembles the staring opened mouth face-and aggressive display-except that the eyes and muscles of the face in general are relaxed. There is variability in the extent to which the teeth are covered and in the position of the eyebrows and lids between species. Nickelson and Lockard (1978), for example, found that in the Celebes the lips were retracted, exposing the teeth. The pink of the eyelids was not seen. In the stumptail, both Bertrand (1969) and Blurton-Jones and Trollope (1968) report that the teeth remain covered as well as the pink of the eyelids. The expression shows greater variability in the liontail in that the lips may or may not be retracted and the pink of the eyelid may or may not be exposed. Retraction of the lips and exposure of the eyelids appeared to occur when the animal was highly excited. This display served to initiate or to continue a play bout.