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Clasping behaviour in Macaca tonkeana

Article in Behaviour · September 1984

DOI: 10.1163/156853984X00010

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 CLASPING BEHAVIOUR IN MACACA TONKEANA

by
BERNARD THIERRY1)
(Laboratoire de Psychophysiologie, Université Louis Pasteur, Strasbourg, France)
(With 7 Figures)
(Acc.1-IX-1983)

Introduction

Judging by the frequency and diversity of physical contact between in-

dividuals, tactile communication is very important in non-human
primates. However, few authors have studied this type of communication
specifically (ANTHONEY, 1968; WEBER, 1973; DE WAAL & VAN
ROOSMALEN, 1979; PELAEZ, 1982). With the exception of behaviours such
as mounting or social grooming, our knowledge of the forms and contexts
of tactile signals has progressed little, in comparison to the knowledge
which has accumulated in the fields of vocalizations and facial expres-
sions. This may be accounted for by two different facts. On the one hand,
it is not possible to record directly a tactile signal: by definition, only the
individuals involved in contact receive the signal, while the observer only
perceives the visual component of the contact. On the other hand, the
tactile signal is frequently accompanied by other types of signal; among
these, olfactory signals sometimes have a role which is difficult to
evaluate (MARLER, 1965; SMITH, 1977).
Clasping is a form of tactile communication encountered in many
primate species. It consists of a more or less close contact between two in-
dividuals, of grasping or embracing for example. Various forms have
been described in Macaca nigra (DixsoN, 1977; NICKLESON & LOCKARD,
1978; BRAMBLETT, 1973) but no studies have been undertaken in the
other six Sulawesi stumptails (cf. FOODEN, 1969, for taxonomy). The aim
of this study was to examine the forms, distribution and contexts of oc-
currence of clasping behaviour in Macaca tonkeana, in order to reveal its
functions in the species' social system.

1) The author is grateful to Pr. Ph. ROPARTZ and to Drs J. J. ROEDERand N. HER-
RENSCHMIDT for their helpful comments on the manuscript. Special thanks are due to Dr
J. R. ANDERSON for his fruitful criticisms and English translation.
2

TABLE 1. Identity, date of birth and genealogies of individuals

in groups A and B

Parentheses indicate estimated birthdates for subjects born in the wild. Other subjects
were born in captivity.

Methods

Subjects.
Two groups of Macacatonkeanawere studied. They originate from the division of one troop
whose origin has previously been described (HERRENSCHMIDT, 1977). The first group (A:
ten individuals) is semi-free ranging in a wooded park of approximately half a hectare.
The second group (B: twelve individuals) is kept in a cage of approximately 50 m2. The
composition of each group and kinship relations are shown in Table 1...

Sampling of data.
Sequences comprising a clasp, a mount or an agonistic interaction were recorded scoring
the identity of the individuals involved. In an initial phase, group A was observed during
April and May 1982, for 66 hours distributed equally between 0800 and 1900 h; the
sampling method used focal sampling (ALTMANN, 1974). In a subsequent phase, the two
grous were observed from July to October 1982, totalling 48 hours for group A and 30
hours for group B. For each group the observations were equally distributed between 0900
and 1300 h, and 1400 and 1600 h. The sampling method used behaviour dependent
sampling (ALTMANN, 1974).
 3

Observational bias: in some cases, the faces of individuals involved in a clasping

episode could not be seen by the observer and thus facial expression was not recorded.
Behavioural units.

Only a part of the behavioural repertoire of Macaca tonkeanais briefly described here. The
species' repertoire is the subject of a study in progress (HERRENSCHMIDT, in prep.).
a. Behaviours associated with threat or attack.
- Stare (ST): visual fixation of the opponent.
- Half open mouth (HOM): mouth slightly open, corners of the mouth retracted, teeth
partly visible; this facial expression is occasionally silent but is most often accompanied
by a sharp vocalization (SHV).
- Openmouth,baredteeth,screech(OMBTS): mouth open wide, lips retracted vertically, cor-
ners of the mouth retracted, teeth visible; this facial expression is accompanied by
strong, shrill screams.
- Jaw movements UM): rapid, rhythmic movements of the lower jaw, the tongue is moved
backwards and forwards but is never protuded, the lips are not protuded; this facial ex-
pression is silent.
- Sharp vocalization(SHV): a short, high-pitched sound, of varying intensity, usually
repeated several times ("ee-ee-ee").).
- Bark (B): a short and loud sound, mouth slightly open, usually repeated several times
("wa-wa-wa").).
- Snarl (SN): a long and throaty sound, not loud, emitted without facial expressions
other than stare; it can be repeated several times.
- Slap (SL): a forceful movement of the open hand towards an opponent, who may or
may not be reached.
- Grab (GR): the fur of the opponent is gripped and pulled roughly.

b. Behaviours not associated with threat or attack.

- Lipsmacking(L): mouth opens and shuts rapidly and rhythmically, lips protruding; ac-
companied by movements of the tongue which is sometimes partly protuded.
- Protudedlips (PL): lips tightened and extended forwards; the head is raised.
- Closemouth, baredteeth(CMBT): mouth closed, lips retracted vertically, corners of the
mouth retracted, teeth visible; this facial expression is silent.
- Openmouth, bared teeth(OMBT): mouth open, lips retracted vertically, corners of the
mouth retracted, teeth visible; this expression can be silent or accompanied by GU or
STV (see below).
- Grunt (Go: a soft, brief sound emitted in a series; the mouth is either half opened
(teeth not visible) or open with teeth bared (OMBT).
- Staccatovocalization(STV): a strong, guttural sound, a rapid succession of rhythmical
exhalations ("hau-hau-hau"). ).
- Turnface away (TFA): in the course of an interaction, an individual briskly turns the
head to one side or backwards, the head is generally brought back in the direction of
the partner; this behaviour may be repeated several times, it may be accompanied by
L, PL, CMBT, OMBT, or GU.
- Expressiverun (ER): after a non-agonistic interaction, one of the animals involved
quickly runs for several metres, and often comes back towards the individual from
which it moved away; this behaviour may be accompanied by CMBT, OMBT, GU,
or STV.
- Mouth approach (MA): an individual brings its mouth towards that of another,
sometimes contact is made.
- Mouthing(MO): an individual mouths the fur of another.
4

Quantitative analysis.
Unless otherwise stated, numbers in the text represent means (:t± SD) for the two groups.
Statistical tests are chi-square test (Yates correction), Wilcoxon matched pairs test, Fried-
man two-way analysis of variance and Spearman rank correlation coefficient.

Results

Description of clasping behaviour.

By the term clasping behaviour we mean a form of interaction which lasts

only a few seconds, and which differs from the various types of contact
which may arise in play, mounts, social grooming, or resting (huddling).
The behaviour of infants (individuals below one year of age) has not been
taken into account in order to avoid confusion with clinging to the mother
as well as other types of parental or alloparental behaviour. Our defini-
tion of clasping will therefore be both formal and contextual.

'
1. Forms of clasping behaviour.

Five types of clasping have been distinguished: grasping the hind-

quarters, grasping, reaching around, embracing, hugging.
a) Graspingthe hindquarters:an individual grasps another at the level of the hindquarters
(rump and hind limbs) with one or both hands (Fig. la, b and c); grasping by individuals
in a bipedal position have not been included, in order to distinguish so-called grasping the
hindquarters from grasping which normally takes place during mounting behaviour.
b) Grasping: an individual grasps another with one or both hands; grasping may be
directed towards any part of the body except the hindquarters.
c) Reachingaround: an individual passes one or both hands, or one or both arms, around
the body of another: around the head, neck, back (Fig. 2a) or below the chest or abdomen;
there may be a simultaneous grasping of the fur.
d) Embracing:two individuals clasp each other face to face, there is contact between the
heads or chests (Fig. ld and 3b); embracing may be unilateral or mutual; there may be a
simultaneous grasping of the fur.
e) Hugging: an individual clasps another one while keeping the latter below him or her
(Fig. 2b); there may be a simultaneous grasping of the fur.

2. Contexts of clasping behaviour.

Three contexts have been distinguished : greeting, sexual harassment and
aggression.
a) Greeting:greeting generally refers to the coming together of two or more individuals,
leading to non-agonistic interactions; here, we shall mean by clasping in greeting all clasp-
ing not preceded by an agonistic interaction or occurring in the context of sexual harass-
ment (cf. Fig. 1 and 2); 138 instances were observed in group A, 125 in group B.
b) Sexualharassment:the term sexual harassment refers to a third party interfering in a
mount between two animals; among the responses of the third party clasping sometimes
 5

Fig. 1. Sequenceof four clasps betweentwofemales. Between each clasp, Bou makes an ex-
pressive run then returns and presents to Mam who clasps her, with associated behaviours
(open mouth bared teeth, staccato vocalization, mouthing); a, b and c: grasping the hind-
quarters ; d: embracing (note behaviour of the infant, Mam's daughter).

'
occurs, most often directed towards the animal being mounted; 4 instances were observed
in group A, 15 in group B: these instances of clasping during harassment represent
25,9 ± 6, 2 % of the total number of instances of sexual harassment observed.
c) Aggression:clasping in the context of aggression is defined as all clasping occurring
either during aggression or in the 10 seconds following aggression; aggression itself is
defined as any interaction containing at least one behaviour associated with attack or
threat (cf. Methods section); 35 instances were observed in group A, 40 in group B.
- Opponent-directed clasping: This type of clasping occurs in the course of an agonistic
interaction between two individuals, one clasping the other; clasping may begin during
the aggression or within 10 seconds after the end of the aggression:
- Clasping during aggression (A: 8 instances observed; B:
23): in all observed instances,
clasping was initiated by the receiver of the aggression;in 26 cases out of the 31, aggres-
sion stopped during clasping: the probability of an association between the end of aggres-
6

Fig. 2. Forms of clasps. a: Reaching around: Bou passes an arm around the back of Mam
who is carrying Mar; b: Hugging: Mou hugs Bou (Cho harasses with open mouth bared
teeth and slap).
 7

Fig. 3. Protectingembrace.a: Mam threatens (half open mouth, sharp vocalization) Cho
who withdraws; b:. Ver, Cho's mother, embraces Mam with lipsmacking: Mam lipsmacks
also while Cho approaches the group (the two photographs were taken at about three
seconds interval).
8

sion and the beginning occurring by chance is so small that the observation of several in-
stances is sufficient to show there was a true association; this type of clasping may be inter-
preted as functioning as an "appeasement" of the aggressor.
- Clasping after aggression (A : 7 B:7):
; in all of the observed instances, clasping was in-
itiated by the aggressor; this type of clasping may be interpreted as leading to "reconcilia-
tion" between the opponents.
- Side-directedclasping:during an agonistic interaction, an animal clasps an individual
who is not an opponent; the animal who grasps may be the aggressor or the agressee:
- Clasping by the agressor: three types were observed: either two individuals
aggressingg
against a third animal might clasp each other mutually (A : 0 B:1);; or one individual ag-
gresses against another while simultaneously clasping a third individual (A : 0 ; an
individual while aggressing against another might go and clasp a third animal not irivolv-
ed in the aggression (A:1; B : 2) in
: the three cases observed, a juvenile received aggression
and its mother was immediately clasped by the aggressor.
- Clasping by the aggressee (A:O;
B:2): the aggressee flees from the aggressor and
clasps a third individual.
- Claspingas non-agonisticinterference
(A:19; B:4): during an agonistic interaction between
two animals, a third individual intervenes by clasping the aggressor (Fig. 3); in 17 cases
the end of the aggression coincided with the onset of clasping; as with clasping between
opponents, this association cannot be attributed to chance. The intervention of the
third individual has the effect of calmly stopping the aggression and may therefore be
described as "non-agonistic protection". Furthermore, in 18 instances out of the 23, the
protector was related to the aggressee: comparison between this distribution and the
distribution of agonistic interactions between related and non-related individuals shows a
highly significant difference (P < 0.001, X2= 51.7, 1 d.f.); although the non-agonistic pro-
tection might be in favour of non-related individuals, it most often results in the ending of
aggression directed towards a related individual (cf. Fig. 3).

3. Intervention of a third individual during clasping.

Individuals may direct a behaviour towards two individuals engaged in
clasping (cf. Fig. 1, 2b and 3b). Two particular types of interaction were
distinguished: clasping and harassment.
a) Clasping:a third individual clasps an animal already involved in a clasp, either during
this first clasp or immediately afterwards. Clasping may be directed towards the initiator
of the previous clasp (A:3; B:6) or towards the receiver (A:2; B:4).
b) Harassment:harassment was only observed on three occasions in group A (Fig. 2b).
In all respects this behaviour resembles sexual harassment: the third individual shows
ambivalent approach-withdrawal behaviours accompanied by slaps without hitting and by
vocalizations and facial expressions such as GU, L, CMBT and OMBT.

Distribution of clasping behaviour

1. Rate.
The rate of initiating clasping was established for each age- and sex-class
from the data obtained from the behaviour-dependent sampling: for each
class represented in the two groups, the rate was always higher in group
 9

A, living in semi-liberty (ad. 0.69; sub. (7: 0.15; juv. 0": 0.17; ad. 9:
0.45; sub. 9: 0.17; juv. 9: 0.92) than in group B, kept in captivity (ad.
0": 0.97; sub. 0.20; juv. o': 0.27; ad. 9: 0.72; sub. 9: 1.60).

2. Distribution.
The distribution of clasping among the different age- and sex-classes has
been calculated from the entire set of data (Table 2). With the aim of
evaluating the contribution of each class in the initiation and receipt of
clasps, the expected distribution was established in order to permit com-
parison with the observed frequencies (ALTMANN, 1968; DITTUS, 1977).
Under the null hypothesis, clasps are randomly distributed among in-
dividuals, that is, each individual clasps and is clasped equally frequent-
ly. The probability of an individual belonging to class i clasping an in-
dividual belonging to class j is:

P;j = total number of individuals in the group.

N(N-1) i : number of individuals in class i.
j : number of individuals in class j.

(within a class, the probability is Pii = ).

N(Nl l)
If F is the total number of clasps observed in the group, the expected fre-
quency of clasps in each class is f = Pij. F.
The expected frequencies shown in Table 2 represent the summed fre-
quencies over the two groups (f = fA + fB).
The observed and expected frequencies are compared for each row and
column of the matrix using the X2 test. The analysis leads to the following
conclusions: adult males, and adult and subadult females initiate
significantly more clasps than expected under the null hypothesis,
juvenile males initiate significantly fewer clasps than predicted, subadult
males show no significant trend; adult and subadult females receive more
clasps than predicted, juvenile males receive fewer clasps than expected
but the difference is not significant (the low scores concerning clasps
received by adult and subadult males do not permit statistical analysis).
The comparison of the number of initiations with the number of
receipts completes this analysis. For the five age- and sex-classes
represented in the two groups, comparison of frequencies of clasps in-
itiated (column totals) with those of clasps received (row totals) shows a
highly significant difference (P < 0.001, XZ = 65.8, 4 d.f.). There is no
evidence of a significant difference for adult and subadult females
10

TABLE 2. Distribution of clasping behaviour

In each cell of the matrix: numbers in small type represent the frequencies respectively
observed in groups A and B; numbers in large type represent the sum of the frequencies
observed over both groups; numbers in middle type represent the expected frequencies
over both groups.

(X2 = 0.03, 1 d.f.), while for males (adults, subadults and juveniles) the
difference is highly significant (P< 0.001, X2 = 22,4, 2 d.f.). Females,
therefore, give as many clasps as they receive, while males initiate more
clasps than they receive.
Furthermore, females frequently initiate clasps during aggression (ag-
gression : 21.9 + 0.8 % of all clasps; greeting: 31.6 + 4.4 % ), whereas
males initiate very few (aggression: 2.0 + 0.9 % ; greeting: 31.5 + 3.8 % )
'
(P< 0.001, X2 = 50.8, 1 d.f.).
 11

3. Correlation with the distribution of agonistic interactions.

The frequency of clasps (initiation and receipt) was compared with the
frequency of agonistic interactions for each individual: there is a signifi-
cant correlation between the two variables in group A (rs = 0. 74,
P < 0.05) as well in group B (rs = 0.67, P < 0.05) (Spearman rank correla-
tion coefficient, one-tailed). Thus individuals most often involved in
clasps are also those most often involved in conflicts.

Fig. 4. Form of claspingbehaviouras a function of context.Ordinate: relative importance of

clasping forms expressed as a percentage. Abscissa: contexts in which clasping occurs.

Variations in form of clasping behaviour as a function of

context and age- and sex-class.

1. Context.

Fig. 4 shows that the different forms of clasping were utilized with equal
frequency in the context of greeting and aggression, only hugging was
restricted to greeting. This is confirmed by a partial statistical analysis
concerning adult and subadult females (these individuals form a class
which is both homogeneous and frequently involved in the two contexts,
cf. above): comparison of the most frequent forms (grasping the hind-
quarters, grasping, reaching around, embracing) reveals no significant
difference between the two contexts (Xz = 3.5, 3 d.f.).
With regard to sexual harassment, Fig. 4 shows that only grasping and
reaching around were observed in this situation. This finding, however,
is of limited importance: the posture of the animals involved in the
12

mount, likewise frontal approach of the harassing individual, prohibits

forms such as grasping the hindquarters or hugging.
No direct relation between forms of clasping and the previously de-
fined contexts has, therefore, been demonstrated.

2. Age- and sex-classes.

The distribution of forms of clasping was established for each age- and
sex-class (Fig. 5). Comparison of the six classes shows that use of the
repertoire varies as a function of age- and sex-class, in initiation
(P < 0.001, X2 r =29.9) as well as receipt (P < 0.001, X2 r =36.9) (Friedman
two-way analysis of variance).

Fig. 5. Form of claspingbehaviouras a functionof age- and sex-class. The percentage ordinate
shows the relative importance of clasping forms for each age- and sex-class.

a) Initiation: the most frequent form in adult and subadult males is

grasping the hindquarters: they use this form more than the other age-
and sex-classes (P<0.001, X2=22.3, 1 d.f.). In contrast, reaching
around is the most frequent form in these latter classes, which they use
more frequently than adult and subadult males (P < 0.001, X2 = 17.5, 1
d.f.). Grasping and embracing show no noticeable trends. Hugging was
observed only in group A: it was initiated by the adult male or the adult
females and was always directed towards a female.
b) Receipt: the small number of observed instances in males does not
permit a definitive conclusion. In females, reaching around remains the
most frequent form.
 13

In the context of greeting and aggression grasping the hindquarters oc-

curred more frequently between sexes (males and adult and subadult
females) than between females (adult and subadult females) (P < 0.001,
x2 = 15.7, 1 d.f.).

Behaviours associated with clasping behaviour.

1. Behaviours accompanying clasping.

Non-tactile communicatory signals frequently accompany clasping
behaviour. They may immediately precede or follow the clasp, or occur
simultaneously with it. No attempt has been made to distinguish between
these three categories, since this type of signal very often began before the
clasp and continued during it, or else it began during the clasp and con-
tinued for some seconds afterwards.

Fig. 6. Behavioursaccompanyingclaspingbehaviour.Values represent the percentage of clasps

in which each of the behaviours occurred.

Nine types of signal may accompany clasping behaviour (Fig. 6):

visual (L, PL, CMBT, OMBT, TFA), auditory (GU, STV) or tactile
(MA, MO); the possibility is not excluded that an olfactory exchange oc-
curs during mouthing or mouth approach. The initiator shows associated
behaviours more often than the receiver (P<0.01, Wilcoxon-matched
pairs-test) (Fig. 6). Furthermore, there exist several variations as a func-
tion of context; a comparison was carried out between greeting and ag-
gression, based on all the adult and subadult females and dealing with
the most frequently encountered behaviours (L, CMBT, OMBT, GU):
no significant difference emerges for L d.f.) or OMBT
(X2 = 2.2, 1 d.f.) while CMBT and GU more
appeared frequently during
greeting than aggression (P < 0.05, X2 = 5.4 and 4.4 respectively, 1 d.f.).
The behaviours accompanying clasping also vary as a function of age-
and sex-class. L, for example, is more characteristic of females
(24. 2 ± 9. 5 % of clasps received or initiated) than males (2.0 ± 2.0 % )
14

(P < 0.001, x2 = 27. I , 1 d. f.), while MO is more characteristic of males

probable that such differences are not specific to the context of clasping.

2. Behaviours follow ing clas,bing.

Nine other behaviours which may follow clasping can be added to those
described above.
a) Expressive run: expressive runs occurred more frequently in the
receiver than the initiator, however, the difference is not significant
(X2 = 1.9, 1 d.f.).

followingclaspingbehaviour.Values represent the percentage of clasps in

Fig. 7. Behaviours
which each of the behaviours occurred.

b) Branch-shaking: branch-shaking was observed on one single occa-

sion, in an adult female who had first performed a protecting clasp.
c) Aggression (between the clasping partners): three instances of ag-
gression after clasping were observed (A:2, B:1); each time, the clasp
was directed towards an animal carrying an infant, this individual then
responding with a threat (OMBTS).
d) Presenting: presenting was observed exclusively in the context of
greeting; it occurs more frequently in the receiver than the initiator
(P< 0.01, X2 = 10.1, 1 d.f.); in addition there exists a relationship be-
tween the form of the clasp and the occurrence of presenting:
17.8 + 1.0% of instances of grasping of the hindquarters (receipt) were-
followed by presenting (P < 0.02, X2 = 6.2, 1 d. f. ). * )

*) Sometimes presenting precedes clasping. As previously, this behaviour is unique to

adult and subadult females and it is more specific to grasping the hindquarters
(24.4 ± 4.5% of the grasps of the hindquarters and 3.7 ± 0.6%of other clasps received;
P<0.001, X2= 12.7, 1 d.f.), it was always performed by the receiver of the clasp.
 15

e) Genital exploration: genital exploration (sniffing or touching the

genital region) was observed exclusively in the initiator, and it may occur
in both males and females.
f) Mounting: any mount is always performed by the initiator of a
clasp; it was observed exclusively in males, and it more often followed
grasping of the hindquarters (78.6 + 2.4% of the latter) than the other
forms of clasping (3.9 ± 3.9 % ) (P < 0.001, X2 = 20.2, 1 d.f.).
g) Clasping: a clasp could be followed by another clasp; the initiator of
the first clasp seems to be the more likely to initiate the second clasp,
however the difference is not significant (X2 = 3.1, 1 d.f.). Two cases may
occur: one of the two animals involved in the first clasp initiates a clasp
with a third individual ( 1. 7 ± 0.9 % ), or a second clasp occurs between
two individuals already involved in the first (7.3 + 0.7 % ). Over the 26
clasping sequences observed between two individuals, the transition
always led towards contact between a greater surface area of the animals:
grasping the hindquarters or grasping - reaching around - embracing
or hugging (an embrace or a hug may directly follow grasping) (Fig. 1).
h) Social play: wrestling may follow clasping, but no attempt has been
made to identify the individual initiating play.
i) Social grooming: the groomer is most often the individual who in-
itiated clasping (P<0.02, X2 = 6. 1, 1 d.f.).

Discussion .

Bibliographical data on clasping behaviour are presented in Table 3. The

existing literature does not always permit precise determination of which
forms were used, thus only the context of clasping and the principal ac-
companying behaviours are reported in the table.

Frequency, form and distribution of clasping behaviour.

There exists great variation in the frequency of clasping behaviour. It

seems to be rare in the genus Cercopithecus while it is frequent in the
genera Cercocebus and Papio (STRUHSAKER, 1967; ROWELL, 1971;
CHALMERS & ROWELL, 1971; BOLWIG, 1978; WALLIS, 1981; PELAEZ,
1982) as well as in the chimpanzee (Pan troglodytes) (NISHIDA, 1970; VAN
LAwICK-GOODALL, 1972). It seems to be particularly frequent in Macaca
tonkeana but, without quantified data on other species of macaque, it is
impossible to make comparisons.
Five categories of clasping were distinguished in this study. However,
clasping behaviour is very variable within each category; in addition,
 16

TABLE3. Bibliographical data on clasping behaviour in non-human primate species

 17

TABLE3. Continued

Behaviourswhich accompanyclaspingmost frequently:lipsmacking,bared teeth (closedor open mouth),

mouthing.Contexts:greeting;sexualharassment;"tension", impreciseor poorlydefinedby the authors;ag-
: opponent-directedclasping;2, side-directedclasping:a, towardsa non-opponentduring the ag-
gression 1,
; betweenthe aggressorsprecedingor followingthe aggression;c, towardsa non-opponentafterthe
gression b,
aggression;3, claspingby a non-opponent:a, during the aggression;b, after the aggression.
18

numerous transitions were observed trom one category to the other,

leading towards a greater contact between individuals. Embracing, for
example, could be a more intense form than grasping: this is accordance
with ANDERSON & CHAMOVE's data (1979) according to which ventro-
ventral contact is one of the most intimate contacts between individuals.
In Macaca tonkeana, it is conceivable that clasping behaviour is a graded
signal in a manner similar to vocalizations or facial expressions
throughout the macaques (ROWELL, 1962; SHIREK-ELLEFSON, 1972;
GAUTIER & GAUTIER, 1977).
The form of clasping varies with the species. It is very varied in Macaca
tonkeana but in other macaque species too (for example: Macaca nigra,
DIXSON, 1977; NICKELSON & LOCKARD, 1978; BRAMBLETT, 1973; Macaca
arctoides: BERTRAND, 1969; CHEVALIER-SKOLNIKOFF, 1974), in certain ba-
boons (patio cynocephalus superspecies: HALL & DEVORE, 1965; ANTHONEY,
1968; BoLwIG, 1978; PELAEZ, 1982), mangabeys (Cercocebus albigena:
GAUTIER & GAUTIER, 1977; WALLIS, 1981), langurs (Presbytis entellus:
WEBER, 1973; DoLHINOw, 1978; BOGGESS, 1982), and in the chimpanzee
(Pan troglodytes: VAN LAWICK-GOODALL, 1968; SUGIYAMA, 1969; NISHIDA,
1970; VAN HOOFF, 1974). In some other species, only some forms of
clasping are used; in the long-tailed macaque (Macaca fascicularis), for
example, grasping is used almost exclusively (ANGST, 1974; CHANCE et al. ,
1977; KOYAMA et al., 1981); sometimes a particular form of clasp is added,
namely grasping the penis or scrotum (Macaca fascicularis: KOYAMA et al.,
1981; Macaca fuscata: KAWAI, 1960; Macaca radiata: SUGIYAMA, 1971;
Macaca arctoides: BERTRAND, 1969; CHEVALIER-SKOLNIKOFF, 1974; Thero-
pithecus gelada: MORI, 1979; Macaca tonkeana: pers. obs.*); Patio anubis:
HALL & DEVORE, 1965; Papio cynocephalus: PELAEZ, 1982; Pan troglodytes:
SUGIYAMA, 1969; DE WAAL, 1982); in contrast, neither reaching around
nor embracing have been reported for the gorilla (Gorilla gorilla) (BYGOTT,
1979). In general, grasping the hindquarters and embracing are the two
most commonly reported forms in the majority of species.
No direct relation was found between the form of clasping behaviour
and contexts such as greeting or aggression in Macaca tonkeana. However,
hugging was observed exclusively during greeting. Furthermore, grasp-
ing the hindquarters frequently preceded mounting, however it did not
necessarily occur in a sexual context, being observed in most age- and
sex-classes, and it was not always followed by a mount (Fig. 1). Such a
sequential association between this type of clasping and mounting has

*) Such behaviour was not observed in the course of the present study.
 19

been observed in several species, and it may lead to solicitation, orienting

or even to raising the hindquarters in the receiver (KAUFMAN &
ROSENBLUM, 1966; BERTRAND, 1969; HANBY & BROWN, 1974).
In Macaca tonkeana, each age- and sex-class used most forms of clasping,
the differences being essentially quantitative. Grasping the hindquarters
constitutes the dominant form of clasping in adult and subadult males,
this may be related to its function as a preliminary to mounting. In
females, reaching around and embracing are the most frequent forms.
Such tendencies have already been reported in several species of primate
(Colobus guereza: OATES, 1977; HORWICH & WURMAN, 1978; Cercopithecus
aethiops: BRAMBLETT, 1980; Patio anubis; ROWELL, 1966; Thero,bithecus
gelada, Macaca nigra: BRAMBLETT, 1973; Macaca radiata, M. nemestrina:
KAUFMAN & ROSENBLUM, 1966).

Behaviours associated with clasping behaviour.

Several behaviours may accompany clasping. In Macaca tonkeana the most

frequent are also those which are more widespread among other species
(Table 3): lipsmacking, bared-teeth (closed or open mouth), mouthing.
Other, more specific behaviours may also occur: PL, TFA, GU and STV
in Macaca tonkeana, tongue-clicking in Macaca radiata (SUGIYAMA, 1971)
and Miopithecus talapoin (GAUTIER & GAUTIER, 1977), or teeth-chattering
in several species of macaque (Macaca sylvanus: LAHIRI & SOUTHWICK,
1966; Macaca arctoides: CHEVALIER- SKOLNIKOFF, 1974; Macacajascicularis:
pers. obs.). In Macaca tonkeana, associated behaviours occurred most fre-
quently in the initiator of the clasp than the receiver, this could reflect a
difference in the motivational state of the clasping partners. Further-
more, the presence or absence of these behaviours seemed to modulate
the meaning of the clasping behaviour: hence the greater frequency of
OMBT and GU in the context of greeting compared to the context of ag-
gression.

Origin of clasping behaviour.

The origin of clasping may be studied ontogenetically. A popular

hypothesis is that clasping originates from the infant's behaviour of cling-
ing to the mother (KUMMER, 1957; ANTHONEY, 1968; POIRIER, 1970a): it
is commonly observed that the infant takes refuge on the mother in situa-
tions of alarm, the same behaviour similarly occurring towards a cloth
surrogate-mother (HARLOW & HARLOW, 1965). In other contexts too, it is
generally thought that physical contact between conspecifics plays a ma-
20

jor role in the reduction of social "tension" (NISHIDA, 1970; DE WAAL &
VAN ROOSMALEN, 1979; cf. Table 3). It would be desirable to verify these
interesting hypotheses through systematic, ontogenetic studies. The use
of clasping may vary with age: in Cercopithecus aethiops, for example,
BRAMBLETT (1980) has shown that grasping the hindquarters increases in
frequency in males on reaching adult age. In Macaca tonkeana, clasping
towards individuals other than the mother appears quite precociously: we
have observed it during sexual harassment in a female of three and a half
months, and during aggression (grasping the aggressor) in a female of
eight months.

Similarities between mounting behaviour and clasping

behaviour.

Taken together, the characteristics of clasping behaviour are very similar

to those of mounting behaviour, as ANTHONEY (1968) has already noted
in the baboon. In Macaca tonkeana, the same behaviours (CMBT,
OMBT, L, PL, MO) may accompany clasping and mounting; and
presenting, play and grooming may be sequentially associated with each
of these two behaviours. Harassment, identical to sexual harassment,
may occur during clasping (Fig. 2b): such behaviour has likewise
been described during embracing in the langur (Presbytis entellus: WEBER,
1973).
Additionally, non-copulatory mounting like clasping may occur in
greeting or aggression; in the latter case the mount appears after aggres-
sion or, in contrast, during aggression in several species of primate (BER-
TRAND, 1969; HANBY & BROWN, 1974; HINDE, 1974; DIXSON, 1977). In
Macaca tonkeana, mounting during aggression occurs principally in males
(THIERRY, unpublished data) while in the same context females use clasp-
ing. Such a sex-difference in the use of clasping also occurs with respect
to the forms used: grasping the hindquarters, like mounting, is a
behaviour which occurs more frequently between sexes (male and adult
or subadult females) than between females (adult or subadult females); in
contrast, the other forms of clasping are more frequent between females
than between sexes; there was no evidence of differences in the frequen-
cies between males and between sexes: the small number of adult and
subadult males present in the two groups does not permit strong conclu-
sions to be drawn.
The principal feature distinguishing clasping from close forms of non-
copulatory mounting may be the presence or absence of pelvic thrusts.
 21

This criterion seems satisfactory in Macaca tonkeana: of 357 clasps ob-

served in the course of this study, a single one contained pelvic thrusts
(ventro-dorsal reaching around between two juvenile males). However,
such a distinction is less clear in other species such as Macaca fuscata
(HANBY & BROWN, 1974) or Papio spp. (ANTHONEY, 1968), for example, in
which pelvic thrusts are not rare during embracing; then it becomes dif-
ficult to decide whether a mount or a clasp is taking place. Incidentically,
it may be noted that in apes, copulation itself may be ventroventral
(MCKINNON, 1974; NADLER, 1975; SAVAGE & BAKEMAN, 1978; CHIVERS,
1978).
It is known in primates and canids that certain patterns of infants'
responses to the mother or adult sexual behaviour may develop into
social signals (WICKLER, 1967; ANTHONEY, 1968; Fox, 1971). It is in-
teresting to note that the behaviours of clasping and mounting, although
of different origin, have acquired similar social functions.

The function of clasping in the control of aggression.

Clasping behaviour plays an important role in the regulation of aggres-

sion in many species of primates (Table 3). It's function varies according
to the context and species: in the genus Cebus, for example, clasping
seems to be used very often in "enlisting", one animal clasping another
while threatening a third individual (BERNSTEIN, 1965; DEFLER, 1979;
ISAWA, 1980; LORENZ, cited by OPPENHEIMER, 1977); in the hamadryas
baboon (Papio hamadryas), the most frequent function is "seeking refuge
and reassurance": the aggressee clasps an individual not involved in the
conflict (KUMMER, 1957); but it is in the chimpanzee that the greatest
diversity is reached: according to DE WAAL et at. (DE WAAL & VAN Roos-
MALEN, 1979; DE WAAL & VAN HOOFF, 1981 ; DE WAAL, 1982), clasping
occurs most often as "enlisting", "seeking refuge and reassurance",
"appeasement" and "reconciliation". In Macaca tonkeana, the role of
clasping is also quite varied; three principal functions have been found in
this study: "appeasement", in which the aggressee ends the conflict by
clasping the aggressor, "reconciliation", in which the aggressor clasps the
aggressee after the aggression; and finally "non-agonistic protection", in
which a third individual intervenes by clasping the aggressor. The occur-
rence of these different behaviours is, however, dependent on the inter-
individual relations: in group B, one subadult female often tried to touch
the infant of another female, the latter then threatened the subadult female
thus provoking appeasement on its part; in group A, one female often
22

threatened a juvenile male: this frequently provoked intervention by the

mother of the juvenile male, and thus a high frequency of non-agonistic
protection in the group.
Non-agonistic protection appears to be an exceptional behaviour, since
in non-human primates when an individual intervenes in a conflict be-
tween two animals it is frequently an agonistic form of interaction (KUM-
MER, 1967; DE WAAL, 1977). In contrast, intervention through a non-
agonistic behaviour has until now only been reported in the chimpanzee,
for which DE WAAL et al. (op. cit. ) have described several cases. However
the effects of non-agonistic protection are not as easily interpretable as in
Macaca tonkeana. In fact, while in the latter species non-agonistic protec-
tion results directly in the end of aggression, in chimpanzees this
behaviour seems to have more subtle consequences, probably on account
of their more complex social mechanisms.
Clasping behaviour constitutes a good model for the study of the con-
trol of aggression: contrary to other types of signal such as auditory or
visual signals, it is always possible to determine unequivocally who is the
receiver of the signal in a clasp. It would be desirable to study this
behaviour thoroughly in order to permit phylogenetic comparisons and to
reveal the evolution of its function. The existence of non-agonistic protec-
tion, for example, may be viewed as a further feed-back in the
homeostasis of a system of social relationships.

Summary
Clasping behaviour was studied in two groups of Macacatonkeana,one being confined in a
cage, the other living in a half-hectare park.
Five patterns have been distinguished: grasping the hindquarters, grasping, reaching
around, embracing and hugging. Clasping may occur in three contexts: greeting, aggres-
sion and sexual harassment. There seems to be no direct relationship between context and
pattern of clasping. However, there is substantial variation in form and context according
to age- and sex-class. Females are the individuals mostly involved in clasps, being in-
itiators as well as receivers; adult males also initiate numerous clasps but receive few.
Adult and subadult males especially use grasping the hindquarters while other age- and
sex-classes more often use reaching around. Clasping in aggression is more characteristic
of females than males.
Several behaviours may be associated with clasping. The accompanying behaviours are
more frequent in initiator than in receiver. Clasp is often followed by social grooming,
mount, wrestle or another clasp.
The results of this study in Macaca tonkeanamay be compared with information from
other studies of non-human primate species. This allows one to draw several conclusions:
1. Frequency, form and distribution of clasping vary from one species to another.
There is great variation in certain species, for instance, Macaca tonkeana.
2. Several behaviours usually accompany clasping and may modulate its meaning.
3. Clasping behaviour shows many similarities to mounting behaviour. Although these
two behaviours have different origins, they have acquired similar social functions.
 23

4. Clasping plays a prominent role in control of aggression. In Macaca tonkeana,this

behaviour is especially used in appeasement (an individual receiving aggression clasps its
aggressor during the aggression), reconciliation (an aggressor clasps its antagonist after
the aggression) and non-agonistic protection (a third individual intervenes in an agonistic
interaction by clasping the aggressor). Non-agonistic protection is an interaction which
was until now described only in chimpanzees.

References
ALTMANN, J. (1974). Observational study of behaviour: sampling methods. - Behaviour
49, p. 227-267.
ALTMANN, S. A. (1968). Sociobiology of rhesus monkeys. III: The basic communication
network. - Behaviour 32, p. 17-32.
ANDERSON, R. J. &CHAMOVE, A. S. (1979). Contact and separation in adult monkeys.-
S. Afr. J. Psychol. 9, p. 49-53.
ANDREW,R. J. (1944). The displays of the primates. - In: Evolutionary and genetic
biology of primates 2 (BUETTNER-JANUSCH, J., ed.). Academic Press, New York,
p. 227-309.
ANGST,W. (1974). Das Ausdrucksverhalten des Javaneraffen MacacafascicularisRaffles
1821. - Paul Parey, Berlin.
ANTHONEY, T. R. (1968). The ontogeny of greeting, grooming, and sexual motor patterns
in captive baboons (superspecies Papio cynocephalus).- Behaviour 31, p. 358-372.
BALDWIN,L. A. & TELEKI,G. (1976). Patterns of gibbon behavior on Hall's island,
Bermuda. A preliminary ethogram for Hylobateslar. - In: Gibbon and Siamang 4
(RUMBAUGH, D. M. ed.). Karger, Basel, p. 21-105.
BENEDICTUS, T. DE (1973). The behavior of young primates during adult copulation:
observations of a Macaca irus colony. - Am. Anthrop. 75, p. 1469-1484.
BERKSON, G., Ross, B. A. &JATINANDANA, S. (1971). The social behavior of gibbons in
relation to a conservation program. - In: Primate behavior: developments in field
and laboratory research 2 (ROSENBLUM, L. A. ed.). Academic Press, New York,
p. 225-255.
BERNSTEIN, I. S. (1965). Activity patterns in a cebus monkey group. - Folia primatol. 3,
p. 211-224.
-- (1968). The lutong of Kuala Selangor. - Behaviour 32, p. 1-16.
-- (1969). Introductory techniques in the formation of pigtail monkey troops. - Folia
primatol. 10, p. 1-19.
-- (1975). Activity patterns in a gelada monkey group. - Folia primatol. 23, p. 50-71.
-- (1976). Activity patterns in a sooty mangabey group. - Folia primatol. 26,
p. 185-206.
- - & SCHUSTERMAN, R. J. (1964). The activity of gibbons in a social group. - Folia
primatol. 2, p. 161-170.
BERTRAND, M. (1969). The behavioral repertoire of the stumptail macaque. - Bibl.
primatol. 11, Karger, Basel.
BOGGESS, J. (1982). Immature male and adult male interactions in bisexual langur
(Presbytisentellus)troops. - Folia primatol. 38, p. 19-38.
BOLWIC,N. (1978). Communication signals and social behaviour of some African
monkeys: a comparative study. - Primates 19, p. 61-99.
BRAMBLETT, C. A. (1973). Social organization as an expression of role behavior among
old world monkeys. - Primates 14, p. 101-112.
-- (1980). A model for development of social behavior in vervet monkeys. -
Developm. Psychobiol. 13, p. 205-223.
BYGOTT,J. D. (1979). Agonistic behavior, dominance, and social structure in wild
chimpanzees of the Gombe National Park. - In: The great apes: perspectives on
24

human evolution 5 (HAMBURG,D. & MCGOWN,E., eds). Benjamin/Cummings,

Menlo Park, p. 405-427.
CARPENTER, C. R. (1940). A field study in Siam of the behavior and social relations of the
gibbon (Hylobateslar). - Comp. Psychol. Monogr. 16, p. 1-212.
CHALMERS, N. R. &ROWELL,T. E. (1971). Behaviour and female reproductive cycles in a
captive group of mangabeys. - Folia primatol. 14, p. 1-14.
CHANCE,M. R. A., EMORY,G. R. &PAYNE,R. G. (1977). Status referents in long-tailed
macaques (Macacafascicularis): precursors and effects of a female rebellion. -
Primates 18, p. 611-632.
CHEVALIER-SKOLNIKOFF, S. (1974). The ontogeny of communication in the stumptail
macaque (Macacaarctoides).- Contrib. Primatol. 9, Karger, Basel.
CHIVERS,D. J. (1978). Sexual behaviour of wild siamang. - In: Recent advances in
primatology 1 (CHIVERS,D. J. & HERBERT,J., eds). Academic Press, London,
p. 609-610.
DEAG,J. M. & CROOK,J. H. (1971). Social behaviour and "agonistic buffering" in the
wild barbary macaque Macaca sylvanaL. - Folia primatol. 15, p. 183-200.
DEFLER,T. R. (1979). On the ecology and behavior of Cebusalbifronsin Eastern Co-
lombia : II. Behavior. - Primates 20, p. 491-502.
DEPUTTE,B. L. (1982). Duetting in male and female songs of the white-cheeked gibbon
(Hylobatesconcolorleucogenys).- In: Primate communication (SNOWDON, C. T.,
BROWN,C. H. & PETERSEN,M. R., eds). Cambridge Univ. Press, Cambridge,
p. 67-73.
DITTUS,W. P. J. (1977). The social regulation of population density and age-sex distribu-
tion in the toque monkey. - Behaviour 63, p. 281-322.
DIXSON,A. F. (1977). Observations on the displays, menstrual cycles and sexual behav-
iour of the "black ape" of Celebes (Macacanigra). - J. Zool. Lond. 182, p. 63-84.
--, SCRUTON, D. M. & HERBERT, J. (1975). Behaviour of the talapoin monkey (Miopi-
thecustalapoin)studied in groups, in the laboratory. -J. Zool. Lond. 176, p. 177-210.
DOLHINOW, P. (1978). A behavior repertoire for the Indian langur monkey (Presbytisentel-
lus). - Primates 19, p. 449-472.
EHRLICH,A. & MUSICANT, A. (1977). Social and individual behaviors in captive slow
lorises. - Behaviour 60, p. 195-220.
EISENBERG, J. F. (1973). Mammalian social systems: are primate social systems unique?
- Symp. IVth Int. Congr. Primat. 1: precultural primate behavior (MENZEL,E.
W., ed.). Karger, Basel, p. 232-249.
-- (1976). Communication mechanisms and social integration in the black spider
monkey, Ateles fusciceps robustus,and related species. - Smithson. Contr. Zool, 213.
ELLEFSON, J. O. (1974). A natural history of white-handed gibbons in the Malayan
peninsula. - Gibbon and siamang 3 (RUMBAUGH, D. M., ed.). Karger, Basel,
p. 1-136.
EMORY,G. R. (1975). The patterns of interaction between the young males and group
members in captive groups of Mandrillus sphinxand Theropithecus gelada. - Primates
16, p. 317-334.
FOODEN, J. (1969). Taxonomy and evolution of the monkeys of Celebes. - Bibl. prima-
tol. 10, Karger, Basel.
Fox, M. W. (1971). Socio-infantile and socio-sexual signals in canids: a comparative
and ontogenetic study. - Z. Tierspychol. 28, p. 185-210.
FURUYA,Y. (1965). Social organization of the crab-eating monkey. - Primates 6,
p. 285-336.
GAUTIER, J. P. &GAUTIER,A. (1977). Communication in old world monkeys. - In: How
animals communicate (SEBEOK,T. A., ed.). Indiana Univ. Press, U.S.A.,
p. 890-964.
GAUTIER-HION, A. (1971). Répertoire comportemental du talapoin (Miopithecustalapoin).
- Biol. gabon. 7, p. 295-391.
 25

HALL,K. R. L. (1962). The sexual, agonistic and derived social beaviour patterns in the
wild chacma baboon, Papio ursinus. - Proc. Zool. Soc. Lond. 139, p. 283-327.
-- &DEVORE,I. (1965). Baboon social behavior. - In: Primate behavior: field studies
of monkeys and apes (DEVORE,I., ed.). Holt, Rinehart & Winston, New York,
p. 53-110.
HANBY,J. P. & C. E. BROWN,(1974). The development of sociosexual behaviours in
Japanese macaques Macaca fuscata.- Behaviour 49, p. 152-196.
HARLOW,H. F. & HARLOW,M. K. (1965). The affectional systems. - In: Behavior of
non-human primates 2 (SCHRIER,A. M., HARLOW,H. F. & STOLLNITZ, F., eds).
Academic Press, New York, p. 287-334.
HAUSFATER, G. (1975). Dominance and reproduction in baboons (Papio cynocephalus). A
quantitative analysis. - Contr. Primatol. 7, Karger, Basel.
HERRENSCHMIDT, N. (1977). Semi-free breeding of tropical celebes macaques (Macaca
tonkeana)in a continental European climate. - J. med. Primatol. 6, p. 58-65.
HINDE, R. A. (1974). Biological bases of human social behaviour. - McGraw-Hill,
U.S.A.
HOOFF,J. A. R. A. M. VAN(1974). A structural analysis of the social behaviour of a
semi-captive group of chimpanzees. - In: Social communication and movement
(CRANACH, M. VON& VINE, I., eds). Academic Press, London, p. 75-162.
HOPF, S., HARTMANN-WIESNER, E., KUHLMORGEN, B. & MAYER, S. (1974). The
behavioral repertoire of the squirrel monkey (Saimiri). - Folia primatol. 21,
p. 225-249.
HORWICH,R. H. & WURMAN,C. (1978). Socio-maternal behaviors in response to an
infant birth in Colobusguereza.- Primates 19, p. 693-713.
ISAWA,K. (1980). Social behavior of the wild black-capped capuchin (Cebusapella). -
Primates 21, p. 443-467.
JAY, P. (1965). The common langur of North India. - In: Primate behavior: field
studies of monkeys and apes (DEVORE,I., ed.). Holt, Rinehart & Winston, New
York, p. 53-110.
KAUFMAN,I. C. & ROSENBLUM, L. A. (1966). A behavioral taxonomy for Macaca
nemestrinaand Macaca radiata: based on longitudinal observation of family groups in
the laboratory. - Primates 7, p. 205-258.
KAVANAGH, M. & DRESDALE, L. (1975). Observations on the wooly monkey (Lagothrix
lagothricha)in Northern Colombia. - Primates 16, p. 285-294.
KAwAI,M. (1960). A field experiment on the process of group formation in the Japanese
monkey (Macaca fuscata)and the releasing of the group at Ohirayama. - Primates 2,
p. 181-253.
KLEIN,L. L. (1971). Observations on copulation and seasonal reproduction of two species
of spider monkeys, Atelesbelzebuthand A. geoffroyi.- Folia primatol. 15, p. 233-248.
KOYAMA, N., ASNAN,A. & NATSIR,N. (1981). Socio-ecological study of the crab-eating
monkeys in Indonesia. - Kyoto university overseas research report of studies on
Indonesian macaques 1, p. 1-10.
KUMMER,H. (1957). Soziales Verhalten einer Mantelpavian-Gruppe. - Verlag Hans
Huber, Bern.
-- (1967). Tripartite relations in hamadryas baboons. - In: Social communication
among primates (ALTMANN, S. A., ed.). Univ. Chicago Press, Chicago, p. 63-71.
--, GOTZ, W. & ANGST,W. (1974). Triadic differentiation: an inhibitory process
protecting pair bonds in baboons. - Behaviour 49, p. 62-87.
LAHIRI,R. K. & SOUTHWICK, C. H. (1966). Parental care in Macaca sylvana. - Folia
primatol. 4, p. 257-264.
LAWICK-GOODALL,VAN J. (1968). The behaviour of free-living chimpanzees of the Gombe
Stream Reserve. - Anim. Behav. Monogr. 1, p. 161-311.
-- (1972). A preliminary report on expressive movements and communication in the
26

Gombe Stream chimpanzees. - In: Primate patterns (DOLHINOW, P., ed.). Holt,
Rinehart & Winston, New York, p. 25-84.
LESKES,A. &ACHESON, N. H. (1971). Social organization of a free-ranging troop of black
and white colobus monkeys (Colobusabyssinicus).- Proc. 3rd Int. Congr. Primat. 3
(KUMMER,H., ed.). Karger, Basel, p. 22-31.
LOY,J. &LOY, K. (1977). Sexual harassment among captive patas monkeys (Erythrocebus
patas) in the Wasa Reserve, Cameroon. - J. Zool. 161, p. 49-63.
MACKINNON, J. (1974). The behaviour and ecology of wild orang-utans (Pongopygmaeus).
- Anim. Behav. 22, p. 3-74.
MCGUIRE,M. T. (1974). The St. Kitts vervet. - Contr. Primatol. 1. Karger, Basel.
MCKENNA,J. J. (1978). Biosocial functions of grooming behavior among the common
Indian langur monkey (Presbytisentellus).- Am. J. Phys. Anthrop. 48, p. 503-510.
MARLER,P. (1965). Communication in monkeys and apes. - In: Primate behavior: field
studies of monkeys and apes (DEVORE,I., ed.). Holt, Rinehart & Winston, New
York, p. 544-584.
MORI,A. (1982). An ethological study on chimpanzees at the artificial feeding place in the
Mahale Mountains, Tanzania - with special reference to the booming situation. -
Primates 23, p. 45-65.
MORI, U. (1979). Development of sociability and social status. - In: Ecological and
sociological studies of gelada baboons (KAWAI,M., ed.), Contr. Primatol. 16.
Karger, Basel, p. 125-154.
NADLER,R. D. (1975). Sexual cyclicity in captive lowland gorillas. - Science 189,
p. 813-814.
NICKELSON, S. A. & LOCKARD, J.S. (1978). Ethogram of celebes monkeys (Macacanigra)
in two captive habitats. - Primates 19, p. 437-447.
NISHIDA,T. (1970). Social behavior and relationship among wild chimpanzees of the
Mahali Mountains. - Primates 11, p. 47-87.
OATES, J.F. (1977). The social life of black-and-white colobus monkey, Colobus guereza. -
Z. Tierpsychol. 45, p. 1-60.
OPPENHEIMER, J. R. (1977). Communication in new world monkeys. - In: How animals
communicate (SEBEOK,T. A., ed.). Indiana Univ. Press, Bloomington, London,
p. 851-889.
PELAEZ,F. (1982). Greeting movements among adult males in a colony of baboons: Papio
hamadryas,P. cynocephalus and their hybrids. - Primates 23, p. 233-244.
POIRIER,F. E. (1970a). Dominance structure of the nilgiri langur (Presbytis johnii) of South
India. - Folia primatol. 12, p. 161-186.
-- (1970b). The communication matrix of the nilgiri langur (Presbytis johnii)of South
India. - Folia primatol. 13, p. 92-136.
POOK,A. G. &POOK,G. (1981). A field study of the socio-ecology of theGoeldi's monkey
(Callimicogoeldii)in Northern Bolivia. - Folia primatol. 35, p. 288-312.
RANSOM,T. M. & RANSOM,B. S. (1971). Adult male-infant relations among baboons
(Papio anubis). - Folia primatol. 16, p. 179-195.
ROSE,M. D. (1977). Positional behaviour of olive baboons (Papio anubis)and its relation-
ship to maintenance and social activities. - Primates 18, p. 59-116.
ROWELL,T. E. (1962). Agonistic noises of the rhesus monkey (Macacamulatta). - Symp.
Zool. Soc. Lond. 8, p. 91-96.
-- (1966). Hierarchy in the organization of a captive baboon group. - Anim. Behav.
14, p. 430-443.
-- (1971). Organization of caged groups of cercopithecus monkeys. - Anim. Behav.
19, p. 625-645.
SAAYMAN, G. S. (1971). Behaviour of the adult males in a troop of free-ranging chacma
baboons (Papio ursinus). - Folia primatol. 15, p. 36-57.
SAVAGE, S. & BAKEMAN, R. (1978). Sexual morphology and behavior in Pan paniscus. -
 27

In: Recent advances in primatology 1 (CHIVERS,D. J. & HERBERT,J., eds).

Academic Press, London, p. 609-610.
SEYFARTH, R. M. (1976). Social relationships among adult female baboons. - Anim.
Behav. 24, p. 917-938.
SHIREK-ELLEFSON, J. (1972). Social communication in some old world monkeys and
gibbons. - In: Primate patterns (DOLHINOW, P., ed.). Holt, Rinehart & Winston,
New York, p. 297-311.
SIMONDS, P. E. (1965). The bonnet macaque in South India. - In: Primate behavior:
field studies of monkeys and apes (DEVORE,I., ed.). Holt, Rinehart &Winston, New
York, p. 175-196.
SKINNER,S. W. & LOCKARD, J. J. (1979). An ethogram of the liontail macaque (Macaca
silenus)in captivity. - Appl. Anim. Ethol. 5, p. 241-253.
SMITH,W. J. (1977). The behavior of communicating: an ethological approach. -
Harvard Univ. Press, Cambridge.
SPIVAK,H. (1971). Ausdrucksformen und soziale Beziehungen in einer Dschelada-
Gruppe (Theropithecus gelada) im Zoo. - Tierpsychol. 28, p. 279-296.
STEVENSON, M. F. & POOLE,T. B. (1976). An ethogram of the common marmoset
(Callithrix jacchus jacchus): general behavioural repertoire. - Anim. Behav. 24,
p. 428-445.
STRAYER, F. F. & HARRIS,P. J. (1979). Social cohesion among captive squirrel monkeys
(Saimiri sciureus).- Behav. Ecol. Sociobiol. 5, p. 93-110.
STRUHSAKER, T. T. (1967). Behavior of vervet monkeys and other Cercopithecines. -
Science 156, p. 1197-1203.
SUGAWARA, K. (1979). Sociological study of a wild group of hybrid baboons between
Papio anubis and P. hamadryas in the Awash Valley, Ethiopia. - Primates 20,
p. 21-56.
SUGIYAMA, Y. (1969). Social behavior of chimpanzee in the Budongo forest, Uganda. -
Primates 10, p. 197-225.
-- (1970). Characteristics of the social life of bonnet macaques (Macaca radiata). -
Primates 12, p. 247-266.
TARTABINI, A. (1978). An analysis of dyadic interactions of male Japanese monkeys
(Macaca fuscata fuscata) in a cage-room observation. - Primates 19, p. 423-436.
TILSON,R. L. (1981). Family formation strategies of Kloss' gibbons. - Folia primatol.
35, p. 259-287.
TUTIN,C. E. G. (1979). Responses of chimpanzees to copulation, with special reference
to interference by immature individuals. - Anim. Behav. 27, p. 845-854.
-- &McGREW, W. C. (1973). Chimpanzee copulatory behavior. - Folia primatol. 19,
p. 237-256.
VICK, L. G. & CONLEY,J. M. (1976). An ethogramm for Lemur fulvus. - Primates 17,
p. 125-144.
WAAL,F. B. M. DE (1977). The organization of agonistic relations within two captive
groups of Java-monkeys (Macaca fascicularis).- Z. Tierpsychol. 44, p. 225-282.
-- (1982). Chimpanzee politics. - Jonathan Cape, London.
- - & HOOFF,J. A. R. A. M. VAN(1981). Side-directed communication and agonistic
interactions in chimpanzees. - Behaviour 77, p. 164-198.
- - & ROOSMALEN, A. VAN(1979). Reconciliation and consolation among chimpanzees.
- Behav. Ecol. Sociobiol. 5, p. 55-66.
WALLIS,S. J. (1981). The behavioural repertoire of the grey-cheeked mangabey Cercocebus
albigena johnstoni.- Primates 22, p. 523-532.
WEBER,I. (1973). Tactile communication among free-ranging langurs. - Am. J. Phys.
Anthrop. 38, p. 481-486.
WICKLER,W. (1967). Socio-sexual signals and their intraspecific imitation among
primates. - In: Primate ethology (MORRIS,D., ed.). Aldine, Chicago, p. 89-189.
 28

WILLIAMS,L. (1967). Breeding Humbold's wooly monkey (Lagothrixlagothricha)at Mur-

rayton wooly monkey sanctuary. - Int. Zoo. Yb. 7, p. 86-89.
WOLFHEIM,J. H. & ROWELL,T. E. (1972). Communication among captive talapoin
monkeys (Miopithecustalapoin). - Folia primatol. 18, p. 224-255.

Résumé
Le comportement d'étreinte est étudié dans deux groupes de Macaca tonkeana,l'un entre-
tenu en cage, l'autre élevé dans un parc d'un demi hectare.
Cinq formes d'étreintes ont été distinguées: l'agrippement de l'arrière-train, l'agrippe-
ment, l'enlacement, l'embrassement et l'enserrement. L'étreinte peut survenir dans trois
contextes: l'accueil, l'agression et le harcèlement sexuel. Il ne semble pas y avoir de rela-
tion directe entre ces formes et contextes. En revanche, il existe des variations importantes
de la forme et du contexte d'utilisation en fonction de la classe d'âge et de sexe. Les femel-
les sont les individus les plus souvent impliqués dans l'étreinte aussi bien en tant qu'ini-
tiateurs qu'en tant que récepteurs; les mâles adultes initient également de nombreuses
étreintes mais en reçoivent peu. Les mâles adultes et subadultes emploient essentiellement
l'agrippement de l'arrière-train alors que les autres classes d'âge et de sexe utilisent sur-
tout l'enlacement. L'étreinte lors de l'aggression est un comportement plus spécifique des
femelles que des mâles.
Plusieurs comportements peuvent être associés à l'étreinte. Les comportements qui
l'accompagnent sont plus fréquents chez l'initiateur que chez l'animal récepteur de
l'étreinte. L'étreinte est souvent suivi d'un toilettage social, d'une monte, d'une lutte ou
d'une autre étreinte.
Les résultats de ce travail chez Macaca tonkeanasont comparés avec les connaissances
établies chez d'autres espèces de primates. Cela permet de tirer les conclusions suivantes:
1. La fréquence, la forme et la distribution de l'étreinte varient d'une espèce à l'autre.
Il existe des variations importantes au sein de certaines espèces telles que Macaca tonkeana,
par exemple.
2. Plusieurs types de comportements accompagnent généralement l'étreinte et certains
peuvent en moduler la signification.
3. Le comportement d'étreinte présente de nombreuses similitudes avec le comporte-
ment de monte. Bien que ces deux comportements aient des origines différentes, ils ont
acquis une fonction sociale semblable.
4. L'étreinte joue un rôle important dans le contrôle de l'agression. Chez Macaca ton-
keana, ce comportement est utilisé principalement dans l'apaisement (un individu agressé
étreint son agresseur pendant l'agression), la réconciliation (un individu agresseur étreint
son adversaire après l'agression) et la protection non agonistique (un individu tiers inter-
vient dans une interaction agonistique en étreignant l'agresseur). La protection non ago-
nistique est un type d'interaction qui jusqu'ici n'a été décrit que chez le chimpanzé.

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