Linda Van Elsacker

Relations between fruits and disperser assemblages in a Malagasy littoral forest : a community-level approach

Interactions among fleshy fruits and frugivore assemblages are presented from a 1-y study in the littoral forest of Sainte Luce, south-eastern Madagascar. This community-level approach allowed us to determine food selection by all consumer species and to evaluate the role different frugivores play in seed dispersal and predation. For this, interactions between 136 consumed fruit species and 13 frugivorous species were studied. Fruit and seed size were the most important physical factors determining food selection of all consumer species. Nutritionally birds favoured and mammals avoided lipid-rich fruits. For Cheirogaleus spp., that go into torpor, there was a trend to select sugar-rich fruit pulp. However, for numerous fruit traits the consumer species had no clear feeding preferences and they seemed to be quite flexible, eating whatever was available. This might be related to unpredictable fruit availability and low fruit productivity in the littoral forest, which may also partially explain the low number of frugivores present. Nevertheless frugivores have different impacts on seed dispersal. Eulemur fulvus collaris is particularly important for the dispersal of large-seeded species, while frugivorous birds and flying foxes ensure plant regeneration between and outside forest fragments. In terms of conservation, heterogeneous seed transport is particularly important for this severely degraded littoral forest.

Tree dispersal strategies in the littoral forest of Sainte Luce (SE-Madagascar)

Zoochory is the most common mode of seed dispersal for the majority of plant species in the tropics. Based on the assumption of tight plant-animal interactions several hypotheses have been developed to investigate the origin of life history traits of plant diaspores and their dispersers, such as species-specific co-evolution, the low/high investment model (low investment in single fruits but massive fruiting to attract many different frugivores versus high investment in single fruits and fruit production for extended periods to provide food for few frugivores), and the evolution of syndromes which represent plant adaptations to disperser groups (e.g. birds, mammals, mixed). To test these hypotheses the dispersal strategies of 34 tree species were determined in the littoral forest of Sainte Luce (SE-Madagascar) with the help of fruit traps and tree watches. The impact of fruit consumers on the seeds was determined based on detailed behavioral observations. Phenological, morphological and biochemical fruit traits from tree species were measured to look for co-variation with different types of dispersal. No indication for species-specific co-evolution could be found nor any support for the low/high investment model. However dispersal syndromes could be distinguished as diaspores dispersed by birds, mammals or both groups (mixed) differ in the size of their fruits and seeds, fruit shape, and seed number, but not in biochemical traits. Five large-seeded tree species seem to depend critically on the largest lemur, Eulemur fulvus collaris, for seed dispersal. However, this does not represent a case of tight species-specific co-evolution. Rather it seems to be the consequence of the extinction of the larger frugivorous birds and lemurs which might also have fed on these large fruits. Nevertheless these interactions are of crucial importance to conserve the integrity of the forest.

Feeding Ecology of Pteropus rufus (Pteropodidae) in the Littoral Forest of Sainte Luce, SE Madagascar

This paper examines bat-plant interactions by focusing on the fruit diet and food selection of flying foxes (Pteropus rufus) in the littoral forest fragments of Sainte Luce, SE Madagascar. Analyses of faecal samples and opportunistic observations revealed 40 endemic plant species in the diet. The bats mainly eat odorous ripe and juicy berries. No particular fruit colour was predominant in their diet. Both multi- and single-seeded fruits are eaten. Small seeds (1–3.5 mm in length) are usually swallowed whole. Passage through the digestive tract of the flying foxes does not reduce the germination rate of seeds nor the percentage of seeds germinated. This study indicates that the role of flying foxes in both short and long distance seed dispersal for a large number of endemic tree species of the littoral forest should not be underestimated when designing reforestation programs in particular or conservation action plans in general.

Function and Distribution of Coalitions in Captive Bonobos (Pan paniscus)

We examined the distribution of support behaviour within a captive group of bonobos. Most support was evoked by inter-sexual conflicts with the two highest ranking females. Within a dyad, the usual winner was most often supported. Individuals that challenged the rank order by aggressions and pestering were aggressed more often by their targets in the company of an ally. The two lowest ranking males served as scapegoats, receiving 80% of the contra-support. In coalitions, inviduals did not aggress victims they would not dare to attack without supporters. However, the victims of coalitions reacted more strongly with fear and rarely counteraggressed than when being attacked alone, indicative of the high impact of aggression in support. The alpha female showed some control behaviour when intervening in conflicts. The data fitted with several functional hypotheses: coalitions functioned to maintain existing ranks, to acquire ranks, to reduce tension, and to test or strengthen the bond. We suggest that support behaviour fulfilled a crucial role in the maintenance of the power of the two highest ranking females over the males. Among the females themselves the dominance relationships were not based on coalitions, but on individual attributes.

The influence of the steepness of dominance hierarchies on reciprocity and interchange in captive groups of bonobos (**Pan paniscus**)

Summary Biological market models explain variability in reciprocity and interchange between groups. In groups with a shallow dominance gradient, grooming will be mostly exchanged for itself (i.e. exchange will occur). In groups with steep dominance hierarchies, interchange is expected: individuals will groom higher ranking individuals to get access to limited resources or commodities such as support in conflicts, and grooming will be traded for these commodities. We examine patterns of reciprocity in grooming and support, and of interchange of grooming for support or for tolerance in six captive groups of bonobos. We test whether differences between groups in patterns of reciprocity and interchange can be attributed to differences in a measure of steepness of dominance hierarchies, which is based on dyadic agonistic interactions. We found that grooming was reciprocal in some, but not all groups. Support was highly reciprocal, but this was a side effect of dominance in most groups. Interchange between grooming and support was observed in some groups. Corroborating earlier findings, this was a side effect of individuals preferring high ranking individuals as grooming and support partners, possibly because these high-ranking individuals provide more efficient support in conflicts. There was no evidence for interchange of grooming for tolerance. Variability in grooming reciprocity was explained by differences in steepness of dominance hierarchies, as predicted by the biological market models. In groups with a shallow dominance hierarchy, grooming was more reciprocal. This was not true for reciprocity in

Sex Differences in the Steepness of Dominance Hierarchies in Captive Bonobo Groups

Bonobos have a reputation as a female-dominated and egalitarian species. We examined the 2 aspects of dominance in 6 captive bonobo groups. Females do not consistently evoke submission from all males in all contexts. Though females occupy the highest-ranking positions in the dominance hierarchy, there are in each group males that obtain rather high ranks and are able to dominate ≥1 female. Thus female dominance is not complete and hierarchies can be better described as nonexclusive female dominance. We studied egalitarianism by measuring linearity and steepness of dominance hierarchies. The hierarchies of all groups are highly linear. Hierarchies among males are steeper than among females. On average, male bonobos are more despotic than females, but females too can have despotic relations, both with other females and with males. Hence one can call bonobos in captivity semidespotic rather than egalitarian.

The pivotal role of rank in grooming and support behavior in a captive group of bonobos (Pan paniscus)

Summary We investigated dyadic grooming relationships in a captive group of bonobos (Pan paniscus) and questioned what social function grooming fule ls in the ‘market of services and favors’. Hereto we examined which of two theoretical models — grooming for support (Seyfarth, 1977, 1980) or grooming according to the similarity principle (de Waal & Luttrell, 1986) — best accounted for the observed grooming distribution. Similarity in traits did not correlate with increased grooming or close proximity among the individuals. Therefore, the similarity hypothesis was rejected. Seyfarth’s model of rank-related grooming was largely cone rmed. The animals distributed their grooming according to the rank of the receivers. We found an exchange between grooming and receipt of support. There was more grooming up than down the hierarchy. However, not all predictions about rank-related competition over grooming were cone rmed. We found that dyadic grooming reciprocity indeed increased with decreasing rank distance. Yet, there was no increase of grooming within the dyad with decreasing rank distance and high ranking individuals were not competed over at the highest rates. The observed correlation between grooming and support received represents an important e t with Seyfarth’s prediction, but does not allow for conclusions about underlying causal processes. Other causal explanations, besides the ‘groom to receive support’ hypothesis, that could explain a similar correlation are discussed.

New Evidence for Leaf Swallowing and Oesophagostomum Infection in Bonobos (Pan paniscus)

We collected data on parasitic prevalence and leaf-swallowing behavior of bonobos (Pan paniscus) between August 1998 and April 1999 at the Iyema research site, Lomako Forest, Democratic Republic of Congo. We report the first detailed observations of leaf-swallowing among bonobos and the first record of the behavior at Iyema-Lomako. Bonobo leaf-swallowing closely fits the description of the behavior among chimpanzees. Bonobos ingested leaves of Manniophyton fulvum, as occurs in two chimpanzee populations in Central and Western Africa and among bonobos at Wamba, about 200 km from Iyema-Lomako. All leaf-swallowing occured in the rainy season. In conformity with patterns among Mahale chimpanzees, the prevalence of Oesophagostomum sp. infection in bonobos increased after the onset of the rainy season.

Reactions of a group of pygmy chimpanzees (Pan paniscus) to their mirror-images: evidence of self-recognition

A group of seven pygmy chimpanzees (Pan paniscus) was tested for their mirror-image reactions during a ten-day experiment. The time spent viewing the mirror waned quickly. Little social responses directed towards the mirror were observed. Self-directed behaviors were shown from testday one on. It was concluded that four out of seven animals could correctly identify their mirror-image, one infant was not (yet) able to do so, and for two other individuals the results were inconclusive.

Social structures in Pan paniscus : testing the female bonding hypothesis

Based on previous research in captivity, bonobos, Pan paniscus, have been called a female-bonded species. However, genetic and behavioural data indicate that wild females migrate. Bonding between these unrelated females would then be in contradiction with socio-ecological models. It has been argued that female bonding has been overemphasized in captive bonobos. We examine patterns of proximity, grooming and support behaviour in six well established captive groups of bonobos. We find that female bonding was not a typical characteristic of all captive bonobo groups. In only two groups there was a trend for females to prefer proximity with other females over association with males. We found no evidence that following or grooming between females was more frequent than between males and unrelated females or between males. Only in coalitions, females supported each other more than male–female or male–male dyads. We also investigated five mother–son pairs. Grooming was more frequent among mothers and sons than in any other dyad, but sons did not groom their mothers more than males groomed unrelated females. Mothers groomed their sons, or provided more support to them than females groomed or supported unrelated males. Thus, while bonds between females were clearly present, intersexual relations between males and either unrelated females or their mothers are of more, or equal importance.