Une étude des relations de dominance a été réalisée dans un groupe de Lemur fulvus mayottensis élevé au Centre de Primatologie de Strasbourg. L'analyse a porté sur différents critères relatifs a l'agression, ainsi qu'à la

        préséance alimentaire lors de l'accès à une source de nourriture ou de boisson. Les résultats révèlent essentiellement une relation de dominance marquée entre les deux mâles adultes du groupe, ainsi qu'une absence de dominance des femelles sur les mâles contrairement au schéma classiquement

        décrit chez les Lémuridés; ils sont discutés dans le cadre de données récentes concernant l'organisation sociale de ces espèce

FORNASIERI, Isabelle

ROEDER, Jean-Jacques

I-Revues

DOMINANCE RELATIONSHIPS IN A LEMUR FUL VUS GROUP 
Isabelle FORNASIERI & Jean-Jacques ROEDER* 
Female dominance is considered to be a characteristic of social organization 
in many prosimian species (Richard, 1 987). Explanatory models proposed for 
female dominance are based mainly on energetic costs of breeding during a short 
annual season (see Richard & Nicoll, 1 987). 
In the Lemuridae, a family of gregarious malagasy prosimians, female 
dominance was first described by Jolly ( 1 966) in ringtailed lemurs, Lemur calta. L. 
calta females were reported to win most agonistic encounters with males, and to 
have priority of access to feeding and drinking sites. However, conventional 
criteria based on the direction of asymmetric dyadic agonistic interactions 
(Bernstein, 1 98 1 )  proved to be difficult to use for other Lem ur species, such as the 
more common L. fulvus, due to the low frequency and intensity of spontaneous 
aggression (Harrington, 1 975 ; Sussman, 1 975 ; Tattersall, 1 977). Consequently, 
dominance in these species has remained little documented in field studies. 
More recent data from semi free-ranging groups (Pereira et al. , 1 990) indicate 
considerable interspecific variability in dominance relationships within the Lemu­
ridae. Compared to unambiguous « female-dominant » species like L. calta, other 
species such as L. coronatus and L. fu/vus appear to show different patterns of 
dominance, without any systematic effect of gender. 
Here, social dominance was studied in L. fulvus mayottensis by examining 
factors related to aggression and to priority of access to food in two competitive 
contexts. Like most other Lemuridae, L.fulvus mayottensis lives in multimale 
groups, but is characterized by a grea ter lability of social units (Tattersall, 1 983).  
The aims of the study were to test the hypothesis of female dominance, using 
experimental situations designed to increase interindividual competition and 
aggression. These situations were adapted from those traditionally used to 
measure dominance in other primates (e .g. macaques, Belzung & Anderson, 1 986 ; 
Boelkins, 1 967). 
MATERIAL 
The study was conducted on a group of L.fulvus mayottensis, at the Centre 
de Primatologie de l'Université Louis Pasteur de Strasbourg, in a spacious 
* Laboratoire de Psychophysiologie, Université Louis Pasteur, 7 ,  rue de l'Université, F-67000
Strasbourg. 
Rev. Eco/. (Terre Vie) , vol. 48, 1 993 .  
1 55 -
indoor-outdoor cage. The group contained two adult males (B, M), one subadult 
male (f), two adult females (H, V), and the subadult daughter (h) of one of the
adult females. This group bas evolved from an original group formed three years 
before from presumedly unrelated wild or captive group-reared adults. The 
animais received commercial primate pellets and water ad libitum. 
METHODS 
The group was studied in two experimental situations involving limited 
access to 1 )  a box containing banana slices, and 2) a bottle of orange juice. Both 
apparatus were attached to the wire-mesh of the cage. In order to reach the 
banana slices, the animais bad to extend an arm through the wire-mesh. Drinking 
from the bottle was via a metal spout. 
For each situation, a series of 1 0  experimental sessions was conducted, during 
the non-breeding season. Each session within a series was separated by 1 2  to 
48 hours. The following behavioural data were collected during each session, 
using checksheets marked off into 30 sec-intervals : 
- the identity of the animais present at the food-box or at the bottle, that 
is, subjects standing within one meter from the apparatus and attempting to gain 
access to the resource (« one-zero sampling » ,  Altmann, 1 974), 
- the acquisition of banana slices or of orange juice, 
- ali occurrences of aggressive interactions in the immediate vicinity of the 
resource (noting initiator, receiver, and outcome of aggression : fleeing, counter­
aggression, or no response) . 
Data collection lasted until ali banana slices bad been eaten or until ali 
individuals bad drank for at !east ten 30 sec intervals, which usually took about 
thirty minutes. 
Aggressive interactions were also recorded using the same observation 
procedure for 1 0  hours during non-experimental sessions. 
RESULTS 
No clear dominance of one individual over ali the others appeared in terms 
of access to banana slices or orange juice. 
During feeding tests, four individuals showed similar degrees of access to the 
bananas, whereas two others, namely the two males M and f, virtually never 
obtained any (Fig. 1 ). The male M was prevented from approaching by frequent 
aggression from B, whereas f was frequently present at the food-box but was 
rarely successful at grasping fruit, despite frequent attempts . 
During drinking tests ali individuals, excepted the male M,  bad access to the 
bottle for severa! periods throughout each session (Fig. 2). On sorne occasions 
severa! subjects were present simultaneously at the bottle and drank together or 
in alternation . Such mutual tolerance was especially characteristic of the females. 
However, ali individuals showed one main period of drinking sooner or later 
during the test. The females h and V bad priority of access at the start of each test. 
The male B and the male f drank next, while the male M was always last to drink, 
once ali others were satiated. Rank order based on either total drinking duration 
1 56 -
M 0.4% 
f D.4% 
Figure l .  - Distribution of banana slices among individuals in the group. Males are underlined.
or drinking dura ti on during the first five minutes of the test was not significantly 
correlated with the feeding rank arder (Spearman rank-correlation test, N = 6, 
rs = 0 .0 1  / rs = 0. 52, n .s . ) .
30 
en QI 
1 -I'IS 8 QI IJ. 
<( h 
<( v 
� f
<( H 
-4-- ll 
5 10 15 20 25 30 Min
Figure 2 . - Mean individual drinking scores during the 10  experimental sessions (number of 30-sec 
intervals) . Males are underlined. 
Eighty percent of ali aggressive interactions occurring at the drinking or 
feeding site were unidirectional, with no marked behavioural reaction by the 
receiver. Submissive behaviours were never observed, and displacement of one 
individual by another was very rarely observed, and only in males . Aggressive acts 
1 57 -
were most frequently initiated by the male B and were directed towards the male 
M, resulting in M fleeing without counter-aggression (Table 1). The male M
virtually never showed aggression against females, whereas the majority of this 
type of aggression was initiated by B. Although only a low proportion of 
aggressive episodes between B and a female resulted in a withdrawal, the 
« winner » was more frequently the male . The subadult male f received aggression 
from females as often as he threatened them, with the exception of H .  Intrafemale 
aggression was more limited both in frequency and intensity, consisting usually of 
vocal threats or mild cuffs . Thus, in the absence of decisive aggression, no 
dominance hierarchy could be inferred from the analysis of intrafemale aggressive 
interactions. In order to correlate the feeding and drinking rank orders with a 
rank order based on aggression, we used Zumpe and Michael's  dominance index 
( 1 986). This index, based on the proportions of aggressive and submissive 
behaviors (fleeing in the present study) between ali possible pairs of animais was 
found to be concordant with other dominance measures, and it is suitable for 
situations where aggression is infrequent. No significant correlations emerged 
from this analysis, although ali were positive (Spearman, N = 6, feeding test :
rs = 0.58 ,  drinking test rs = 0.7 1 for total drinking duration and rs = 0 .3 1 for
drinking during the first five minutes) . 
During non-testing periods, aggression was even more infrequent, and the 
only dominance relation that appeared clearly was between the two males B and 
M, again showing that B was dominant. 
DISCUSSION 
In the group of L. fu/vus mayottensis studied here, we found no agonistic 
dominance or feeding priority of one sex relative to the other. Priority of access 
to an incentive in a restricted-access situation appeared in sorne L. fulvus 
individuals, but the feeding rank order varied between situations and was not 
significantly correlated with rank order based on aggressive interactions. 
The most striking result was the marked dominance relationship observed 
between the two adult males, which was evident from the asymmetry of aggression 
and displacement as weil as in priority of access to a limited food resource . In the 
context of another food acquisition task - opening a baited food-box - a similar 
dominance-subordination relationship occurred between adult male L. fu/vus 
mayottensis, whereas female dominance emerged in L. macaco and L. catta 
(Fornasieri et al. , 1 990) . Moreover, increased evidence of intermale dominance 
appeared from long-term observations of L. fulvus social behaviour, with escala­
ting aggression occurring during the breeding season (Colquhoun, 1 987 ; Forna­
sieri , 1 99 1  ) . 
Apart from adult males, social conflict in the group was characterized by 
infrequent decisive interactions. However, when considering only interactions 
intense enough to result in withdrawal, these were performed most frequently by 
a male, in agreement with the results of Pereira et al. ( 1 990) in L. fulvus rufus. In 
L. fulvus females, social relationships seem to be especially influenced by prefe­
rential associations with individual males (see Fornasieri, 1 99 1 )  and by occasional 
episodes of social rejection. The phenomenon of « targeting aggression » (Vick & 
Pereira, 1 989), initiated by most group-members against one individual and 
1 58 -
TABLE 1 
Aggression in limited-access situation to afeeding resource (pooled data for feeding 
and drinking tests) . 
Top li ne : number of aggressive acts initiatedjreceived ; bottom li ne : percentage of decisive aggressions 
initiatedjreceived (aggression resulting in withdrawal) . Males are underlined. 
M f H v h 
134 1 1 27 1 0 54 1 0 128 1 9 25 1 6 
B 
63 0 7 - 20 0 7 44 8 17 
9 1 6 2 1 0 0 1 0 0 1 0 
M 
0 50 0 - - - - -
54 1 7 55 1 41 53 1 52 
f 
20 0 4 0 2 2 
3 1 4 0 1 0 
H 
0 0 - -
2 1 16 
v 
0 0 
resulting in the latter's exclusion from the group, has been described in ali captive 
rearing conditions and in the field, and appears to be an important social 
regulatory mechanism in Lemuridae. 
In conclusion, the absence of any significant correlation between the 
measures used to evaluate dominance in the two situations studied here raises the 
problem of the nature of social dominance in the Lemuridae, in which the forms 
of aggression limit the applicability of traditional methods of deciding of 
dominance used in other primates. Compared to most other primates which 
respond to restricted food-access situations by increased competition, L. fulvus 
1 59 -
mayottensis appear to show only mild aggression and greater tolerance in similar 
contexts. In contrast to L. calta, which exhibit a clear dominance of females over 
males, as weil as a marked dominance hierarchy among females and to a lesser 
extent in males (Kappeler, 1 990), in L. fulvus mayottensis, only males could be 
ranked reliably in terms of dominance status. Other dominance relationships 
varied great! y as a function of the criteria used and the situation, suggesting low 
interindividual rank order differences in terms of agonistic or feeding precedence 
in this species. 
RÉSUMÉ 
Une étude des relations de dominance a été réalisée dans un groupe de Lemur 
fu/vus mayottensis élevé au Centre de Primatologie de Strasbourg. L'analyse a 
porté sur différents critères relatifs à l'agression, ainsi qu'à la préséance alimen­
taire lors de l'accès à une source de nourriture ou de boisson. Les résultats révèlent 
essentiellement une relation de dominance marquée entre les deux mâles adultes 
du groupe, ainsi qu'une absence de dominance de� femelles sur les mâles 
contrairement au schéma classiquement décrit chez les Lémuridés ; ils sont 
discutés dans le cadre de données récentes concernant l'organisation sociale de ces 
espèces.  
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BERNSTEIN, I .S .  ( 198 1 ) . - Dominance : The baby and the bathwater. Behav. Brain Sei. , 4 :  4 1 9-457.  
BOELKINS, R.C. ( 1 967). - Determination of dominance hierarchies in monkeys. Psychonom. Sei. , 7 :
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1 6 1  



Dominance relationships in a Lemur fulvus group

Une étude des relations de dominance a été réalisée dans un groupe de Lemur fulvus mayottensis élevé au Centre de Primatologie de Strasbourg. L’analyse a porté sur différents critères relatifs à l’agression, ainsi qu’à la préséance alimentaire lors de l’accès à une source de nourriture ou de boisson. Les résultats révèlent essentiellement une relation de dominance marquée entre les deux mâles adultes du groupe, ainsi qu’une absence de dominance des femelles sur les mâles contrairement au schéma classiquement décrit chez les Lémuridés ; ils sont discutés dans le cadre de données récentes concernant l’organisation sociale de ces espècesFornasieri Isabelle, Roeder Jean-Jacques. Dominance relationships in a Lemur fulvus group. In: Revue d'Écologie (La Terre et La Vie), tome 48, n°2, 1993. pp. 155-161

Fornasieri, Isabelle

Roeder, Jean-Jacques

Périodiques Scientifiques en Édition Électronique

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Dominance relationships in a Lemur fulvus group

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