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Communal roosting and formation of sleeping clusters in barbary macaques (Macaca sylvanus).

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American Journal of Primatology 28271-280 (1992)
Communal Roosting and Formation of Sleeping Clusters
in Barbary Macaques (Macaca sylvanus)
VIVEKA ANSORGE, KURT HAMMERSCHMIDT, AND DIETMAR TODT
Institut fur Verhaltensbiologie,Freie Uniuersitat Berlin, Berlin 41, Germany
Sleeping cluster composition and distribution were studied in a semi-freeranging population of 174 Barbary macaques (Macaca sylvanus) in Rocamadour/France. Over a period of 2 months 341 sleeping clusters comprising 754 animals were recorded as animals left the sleeping trees. To
control for nocturnal activities five observation periods were conducted,
each of which covered a complete night. Sleeping partners were selected
from a particular set of individuals. From night to night there was considerable fluctuation among the individuals, which finally formed a sleeping cluster. Preferred size of sleeping clusters was two and three individuals per cluster. The majority of females spent the nights in body contact
to infants or female juveniles; whereas males either slept alone or with
older male juveniles. Adult males and females never shared a sleeping
cluster. During the night the animals showed hardly any locomotive and
vocal activity. Sleeping clusters remained stable until dissociation the
next morning. o 1992 Wiley-Liss, Inc.
Key words: social behavior, nocturnal activity, roosting sites
INTRODUCTION
Sleeping clusters, in which individuals spend extended time in body contact,
are characteristic of most primate species [citations in Anderson, 19841 and represent an important part of social behavior. Spacing behavior, in the context of
communal roosting as well as individual sleeping partner choice, is closely correlated to the species-specific social network, as documented for Perodicticus potto
[Clauss et al., 19811,Miopithecus talapoin [Gautier-Hion, 19701,Papio papio [Dunbar & Nathan, 1972; Anderson & McGrew, 19841, Papio hamadryas [Kummer,
19681,Papio cynecephalus [Rasmussen, 19791,Cercopithecus petaurista [Todt et al.,
19823,Macaca mulatta [Vessey, 19731,Macaca sylvanus [Todt et al., 1984; Ansorge
& Hammerschmidt, 1987; Ansorge, 19891, Symphalangus syndactylus [Chivers,
19711, and Pan troglodytes [Riss & Goodall, 19761. There is no doubt about the
ecoethological functions of communal roosting [review in Anderson, 19841, but the
social correlates and the intra-group consequences are unclear.
Whereas there are several studies on the daytime social network of Barbary
Received for publication April 9, 1991; revision accepted April 17, 1992.
Address reprint requests to Viveka Ansorge, Institut fur Verhaltensbiologie, Freie Universitat Berlin,
Haderslebenerstr. 9, 1000 Berlin 41, Germany.
0 1992 Wiley-Liss, Inc.
272 I Ansorge et al.
ENTRANCE
C
-A
200 m
Fig. 1. “La For&t des Singes” i n RocamadouriFrance lmodified from De Turckheim & Merz, 19841. The park
consists of 10 fenced ha ofjuniper scrub, open meadow, and oak forest (part A,B,C). The preferred sleeping site
was part A. Data recording took place in two areas ( a l , a2). For a detailed description of “La for& des singes”
see De Turckheim and Merz [ 19841.
macaques [Deag, 1974; Deag & Crook, 1971; Taub, 1978,1984; Wolff & Todt, 1985;
Kuster & Paul, 1986; Timme, 1989; Small, 1990a,bl little has been published about
their nighttime relationships [Ansorge & Hammerschmidt, 1987; Ansorge, 19891.
In this paper we present data on Barbary macaque’s nocturnal behavior: 1)nocturnal activity, 2) age and sex composition of sleeping clusters, and 3) constancy in
partner choice.
MATERIALS AND METHODS
Study Animals and Study Site
The study was conducted on semi-free-ranging Barbary macaques (Macaca
sylvanus) a t “La For& des Singes,” Rocamadour, France (Fig. 1). The population of
Rocamadour was established in 1974, and has been the subject of various research
projects [Todt et al., 1984; Todt & Pohl, 1985; Wolff, 1985; Lehmann, 1989; van den
Bergh, 1989; Ansorge, 1989; Hammerschmidt & Ansorge, 1989; Small, 1990a,b;
Hammerschmidt, 19901. “La Foret des Singes” is a visitor park were the monkeys
range freely while the visitors are restricted to a path (see dashed line in Fig. 1).
Thus the animals can easily avoid contact with humans but are completely habituated to their presence.
The size of the park allows the monkeys to choose different areas for sleeping
and for daytime activities. The oak trees (part A, B, C in Fig. 1)have a height of
about 10-15 m and are easy to survey. The animals feed on natural vegetation, on
monkey chow which is placed in feeder huts, on grain seeds, and on apples which
are spread throughout the park. For further details of park management see Small
[199Obl.
Data were recorded during the birth season in spring 1987 (April to July). At
Roosting Behavior in Barbary Macaques / 273
TABLE I. M m a c a syluanus Population of “La For& des Singes” During the Birth
Season 1987 (Newborns excluded)*
Males
Females
Sum
12
12
24
9
17
26
7
10
17
9
6
15
13
17
30
11
25
36
*The discrete age classes are due to the birth season.
N = 148.
the end of the season the population consisted of 174 individuals, including 26
newborns. For the age profile used in this paper we distinguished the following
classes: infants (0-3 months), juveniles (age 1-3 years for females and 1-4 years
for males) and adults (Table I).
As Barbary macaques are seasonal breeders [Kiister & Paul, 19841siblings are
at least 1or 2 years apart. Therefore observed differences between young group
members (juveniles and infants) permit the marking of clear age differences.
Data Sampling
A sleeping cluster was defined as animals roosting in body contact. A sleeping
group was defined as an aggregation of several clusters which were spatially separated. According to the different properties of the areas a l and a2 (Fig. 1) the
distances between clusters within a group as well as the distances between sleeping groups varied. In attempting to note the sleeping places and constellations it
was impossible to survey all 174 animals a t once. Hence we had to find a way to
take a random sample each day. To do this we selected five adult individuals (all
were males) as focals and followed one of them each evening (focal animal observation) until it formed a stable sleeping cluster. Then we did a statistical survey
the next morning of the animals that surrounded the observed male. We selected
male focals as earlier observations in 1986 had shown that, although they frequently slept alone, they are just as spatially well integrated in the sleeping group
as females.
Each morning, before sleeping clusters disbanded we noted sex, age-class,
spatial position, and, where possible, the identity (30% of a total of 754 registered
animals) of all animals roosting up to 20 m around the focal animal. Because of the
darkness only adult animals could be identified. Data were recorded on 47 days;
754 animals were recorded distributed in 341 sleeping clusters.
We also conducted 5 all-night observations lasting from dusk until dawn (9
hours). A selected number of subjects (selection see above) was observed (event
sampling) after the termination of the focal observation. All activities including
vocalizations (mean: 21.8 per night) were recorded. As a measure of vocal activity
the minutes where any vocal pattern was uttered were counted throughout the
night. Each animal that vocalized in the same minute was counted as if it were a
seperate event. During the five nights we totaled 109 observations of animals.
RESULTS
Nocturnal Activity
In all night observations the monkeys were generally quiet and rarely showed
activity. They did not change their sleeping places. If a short dissociation occurred
they always returned to their original places. The majority of nocturnal disturbances were caused by young group members. Usually one member of a cluster
274 I Ansorge et al.
changed position. As a result other members of that cluster, in most cases juveniles
or infants, vocalized or even left the group for a short moment (<1min). This kind
of movement was observed 184 times (1.5 movements per animal in five nights);
62.5% of these movements were due to juvenile or infant activities. Accordingly
94.9% of the registered vocalizations (total: 361) were uttered by juveniles or infants. Vocalizations usually ended when a cluster had reassembled after a short
dissociation or in, other words, when the adult of a group had reaccepted the
protesting youngster in his huddle.
Roosting Sites
In spring 1987 the entire population roosted in part A of the park (Fig. 1). Our
recorded data refer to the areas a1 and a2. These areas were separated by a trail
and a number of trees which were not used for roosting. Within these areas some
trees were used more frequently than others, and there was considerable fluctuation in the occupation of marginal trees. Within the same area, spatial distances
between sleeping clusters varied between 0.5 and 30 m (al: mean distance of the
nearest neighbour of each group: 4.84, SD: 4.16; a2: mean: 5.84, SD: 4.48). Both
areas had a batch of young saplings which were not suitable for sleeping purposes;
thus there were always unoccupied trees, on average one-third. The 341 sleeping
clusters were distributed among 286 trees; 225 trees were occupied by a single
cluster. More than three clusters per tree were never observed.
Most of the individually known animals preferred one area. This distribution
of sleeping places correlated with the membership in different subgroups which
developed during the following years. In spring 1989 a stable subgroup roosted
separately from the other group in part B of the park. Most animals that preferred
area a1 in spring 1987 were now members of this subgroup.
Composition of Sleeping Clusters
In the choice of cluster sizes the animals showed clear preferences. In the
following the observed and expected number of clusters for each size is given: 80
single clusters (expected: 188.51, 175 dyads (exp.: 94.25), 80 triads (exp.: 62.71, 21
tetrades (exp.: 47). This result is significant (chi2 = 149.6, P < .001). Whereas
dyads and triads were preferred the macaques avoided to sleep alone or in a tetrade.
Most of those individuals that spent the night alone were adult males. These
singly sleeping males constituted 40% of all males, whereas only 10% of the adult
females or older juveniles were found roosting alone (Fig. 2).
Both adult males and females preferred t o form a cluster with one or two
non-adults. Here, males showed a preference for older juveniles (2 to 4 years of
age), whereas females preferred yearlings or infants. In triads and tetrades the two
younger sleeping partners of an adult female predominantly differed in age (83%
of 107 juveniles). In contrast only about half of the juveniles roosting together or
in a sleeping cluster with an adult male differed in age (male-juvenile-clusters:
49%, n = 31; all-juvenile-clusters: 53%, n = 79).
With respect to the sex of sleeping group members we found a clear cut distribution: Adult females did not roost together with adult males and the majority
of adult-juvenile clusters were comprised of members belonging to the same sex
(86%of 126juveniles). However, due to the difficulty in identifying a juvenile’s sex
at dawn, the latter result refers to a limited amount of material (n = 126 juveniles).
Exceptions from clusters with members of the same sex were observed in
Roosting Behavior in Barbary Macaques I 275
Sleepingpartner choice
Distributionof
s l e e p i n g c l u s t e r size
'A
1'"
QP
50
dc7
ns
2s
155
J1
n = 133
--SO
J2
n = 122
25
.-
-
lnf J1
11.1
JZ
J3&
'$ C?
I
-so
J3/4
n = 93
25
Fig. 2. Sleeping partner choice in dyads. Bars indicate the percentage of sleeping partners belonging to the
respective age and sex class. The circles give the distribution of sleeping cluster size for each age and sex class.
Size = 1,W, 2, M; 3, m; 4, 0.
mother-offspring clusters (male infants), or as individual idiosyncrasies: For example, the oldest adult male in Rocamadour had developed the unusual habit of
spending the night with one or two juvenile females.
276 I Ansorge et al.
2
C
W
2
W
E
60
.c
5)
W
>
L
W
50
0
40
30
20
10
10
20
30
40
50
expected frequency
Fig. 3. Correlation of the observed and expected frequency of cluster composition types. Cluster types with
observed or expected frequency below 5 are as follows: F + F + ;F + FJ; MMM; F + J J ; F J J J ; MJJJ; MMMJ; JJJJ.
Expected frequency: e = p ( n ai)*q. p ( n a,) = probability that a member of a certain age and sex class (i) is in
the cluster (based on the age and sex profile of the entire population, Table I); nJ = observed frequency (n) of
different cluster sizes 0’).
Figure 3 illustrates the preference for certain cluster types and reveals a
further aspect: Females with newborns (F+ 1 avoided additional sleeping partners, whereas those without newborns favoured clusters with more than one partner.
Constancy in Sleeping Partner Choice
We analyzed data of all individually known animals. As mentioned above
these were only adults and covered 30%of the whole population. Sleeping partners
were not selected at random. Figure 4 shows for 31 individually known animals the
following correlation: The greater the number of observation days, the greater is
the number of differing sleeping partners of an animal. The adult males that sleep
alone constitute an exception (diamonds on the zero-axis). If one excludes these
adult males from the sample in Figure 4 this correlation is highly significant (r =
5 7 , P < .01; Pearson’s correlation coefficient). Of course, the number of different
sleeping partners is limited, because the animals most probably choose from a
confined circle of animals with whom they are well familiar. But only a few days
of observation suffice to register a considerable fluctuation in sleeping partner
choice.
A similar dynamic was apparent in the choice of roosting places: Normally, a
monkey did not spend two or more successive nights in the same tree although
Roosting Behavior in Barbary Macaques / 277
q
5
* *
*
*
. .
0
0
3
6
9
12
15
days of observation
Fig. 4. Fluctuation in sleeping partner choice. Correlation of the number of different sleeping partners with
days of observation. The data sample refers to all individually known animals which were seen more than once
in the morning sample.
there seemed to be individual preferences for certain trees. Changes in roosting
places were presumably related to changes in partner choice.
DISCUSSION
It is clear that communal roosting in primates contributes to a number of
ecological and social functions [Anderson, 19841. Communal roosting occurs in
connection with common access to food resources or restricted sleeping lodges such
as trees or cliffs. Obvious consequences of communal roosting in terms of social
functions are an increased number of potential contact partners and therefore
increased information transfer about various kinds of social states (e.g., rank, role,
kinship, sexual state).
Establishment of Subgroups
Among primates several roosting sites in the home range of a group are common and different groups of the same population with overlapping home ranges
may compete over roosting sites [see Anderson, 19841. The occurrence of changes
in the use of certain roosting sites may not only depend on basic properties of the
respective home ranges or habitats. They may point to other factors which might
have caused them. During our study on the Barbary macques in Rocamadour we
observed the establishment of a novel subgroup. This consisted of 40 individuals,
which 1 year later roosted some 100 meters apart from the original roosting site
still used by the rest of the population (see part A and B, Fig. 1). The spatial
distribution of sleeping clusters reflects the beginning of the splitting process as
described also for other species [Bouliere et al., 1970; Eisenberg et al., 1972;
McGuire et al., 1974; Ransom, 19811. This indicates that changes of social relations
preceeded the group splitting.
Adult-Juvenile Male Bonds
Males showed a strong inclination to associate with older juvenile males in a
sleeping cluster and vice versa. A comparable result could not be found in other
species [Gautier-Hion, 1970; Dunbar & Nathan, 1972; Vessey, 1973; Clauss et al.,
1981; Anderson and McGrew, 19841.
Barbary macaque males build close relationships with infants and the bonds
between adult males and juvenile males continue with growing age of the juveniles
278 I Ansorge et al.
[Paul, 1984; Bartecki, 19861. However, still 40%of the adult males slept alone. A
similar result was found for rhesus macaques [Macaca mutlatta: Vessey, 19731,
Talapoins [Miopithecus talapoin: Gautier-Hion, 19701, and Pottos [Perodicticus
potto M: Clauss et al., 19811.
Fluctation in Partner Choice
Our data show that the animals alternate between sleeping partners. Still we
cannot present enough observations about individually known subjects (Fig. 4) to
draw a conclusion about the total number of potential sleeping partners for one
individual. It would be an interesting enrichment of the knowledge of social behavior to compare the social circle a t night with that during daytime.
However, we can state that the observed fluctuation in partner choice arose
partly from juveniles looking for alternative sleeping partners after rejection by
their mothers. The likelihood that a mother will reject her older infant may increase after she gives birth to her next offspring since mothers with newborns
avoid additional sleeping partners. Our data suggest that juvenile males are therefore forced to find new sleeping partners whereas juvenile females can return to
their mother’s sleeping cluster. The separation of infants from their mothers as
their nightly sleeping partner takes place later than weaning, which has its peak
around the age of 6 months when Barbary macaque mothers become sexually
active [Paul, 1984; Timme, 19891.
Matrilineal Kinship
As in other species of the genus Macaca, there is a strong matriline in Barbary
macaque populations and a linear rank order dominates the social life of females
[Deag, 1974; Kiister & Paul, 1986; Small, 1990bl. Daytime behavior shows that
familarity between individuals corresponds to kinship and is reflected by the
amount of affiliative behavior among group members [Wrangham, 1983; Bartecki,
1986; Steuckardt, 1989; Timme, 1989; Small, 1990a,bl. We cannot present data
about kin since it was not possible to identify young group members in the dark.
However, we found out that with the exception of infants, youngsters in a sleeping
cluster were predominantly of the same sex. In triads and tetrades the two younger
sleeping partners of an adult female predominantly differed in age (83% of 107
juveniles). In contrast only about half of the juveniles roosting together or in a
sleeping cluster with an adult male differed in age (male-juvenile-clusters:49%, n
= 31; all-juvenile-clusters: 53%, n = 79).
The age and sex ratio of female-juvenile triads and tetrades allow the assumption that most of them were sisters with their mother.
CONCLUSIONS
1. Compared to other species there was a high frequency of male-juvenile
clusters. This might be a consequence of the close male-infant relationship which
are characteristic for Barbary macaques.
2. Juveniles that roosted with adult females or males were predominantly of
the same sex as the adults. The sex-specific division in sleeping partner choice as
well as the age structure of female-juvenile clusters may correlate to the matrilineal structure of Barbary macaque populations.
3. The fluctation in sleeping partner choice may enhance the variety of social
relationships beyond the mother-infant dyad. This increases the social knowledge
about other group members and strengthens group bonds.
Roosting Behavior in Barbary Macaques / 279
4. A spatial aggregation of sleeping clusters might indicate the establishment
of a new social subgroup.
ACKNOWLEDGMENTS
We thank Ellen Merz and Gilbert de Turckheim for permission to study a t “La
Foret des Singes,” and Lucien Ambiehl for cooperation a t the park and hospitality
at his campground; Henrike Hultsch and Gottfried Wiedenmann for comments on
the manuscript; and Kurt Kaemmerer for assistance with the English version.
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