Caroline E. G. Tutin

Cultures in chimpanzees

As an increasing number of field studies of chimpanzees (Pan troglodytes) have achieved long-term status across Africa, differences in the behavioural repertoires described have become apparent that suggest there is significant cultural variation. Here we present a systematic synthesis of this information from the seven most long-term studies, which together have accumulated 151 years of chimpanzee observation. This comprehensive analysis reveals patterns of variation that are far more extensive than have previously been documented for any animal species except humans. We find that 39 different behaviour patterns, including tool usage, grooming and courtship behaviours, are customary or habitual in some communities but are absent in others where ecological explanations have been discounted. Among mammalian and avian species, cultural variation has previously been identified only for single behaviour patterns, such as the local dialects of song-birds,. The extensive, multiple variations now documented for chimpanzees are thus without parallel. Moreover, the combined repertoire of these behaviour patterns in each chimpanzee community is itself highly distinctive, a phenomenon characteristic of human cultures but previously unrecognised in non-human species.

Mating patterns and reproductive strategies in a community of wild chimpanzees (Pan troglodytes schweinfurthii)

Summary1.The sexual behaviour of a chimpanzee community in the Gombe National Park, Tanzania, was studied intensively for 16 months. Additional information came from 15 years of demographic and behavioural data accumulated by Jane Goodall and members of the Gombe Stream Research Centre.2.The mating system of the Gombe chimpanzees is flexible and comprises three distinct mating patterns: (a) opportunistic, non-competitive mating, when an oestrous female may be mated by all the community males; (b) possessiveness, when a male forms a special short-term relationship with an oestrous female and may prevent lower-ranking males from copulating with her; and (c) consortships, when a male and a female leave the group and remain alone, actively avoiding other chimpanzees. While males took the initiative in possessive behaviour and consortships, females had to cooperate for a successful relationship to develop.3.Data from 14 conceptions indicated that the majority of females (9) became pregnant while participating in the restrictive mating patterns, possessiveness and consorting. It could be established definitely that seven of these females were consorting during the cycle in which they conceived. As 73% of the 1137 observed copulations occurred during opportunistic mating, 25% during possessiveness, and only 2% during consortships, there was no correlation between copulation frequency and reproductive success.4.Adult males showed differential frequencies of participation in the restrictive mating patterns. Male age, dominance rank, and the amount of agonistic behaviour directed to females showed no correlation with participation in the restrictive mating patterns. The following male characteristics did show significant, positive correlations with involvement in the restrictive mating patterns: (a) the amount of time spent in the same group as oestrous females, (b) the proportion of that time spent grooming oestrous females in groups, and (c) the frequency with which males shared food with females. While dominance ranks of the adult males showed no consistent correlation with involvement in the restrictive mating patterns, it was clear that the most dominant male did gain an advantage. He was the only male able to monopolise oestrous females by showing possessive behaviour.5.Consortships appeared to be the optimal reproductive strategy for males (with the exception of the most dominant) and females, as they gave males the highest probability of reproductive success, and allowed females to exercise choice. However, there appeared to be disadvantages associated with consort formation; the greatest of these was the increased risk of intercommunity encounters. While all individuals have the potential to practice each mating pattern, the strategy actually used at any moment will be determined by variables both within the individual, e.g. age, physical condition, dominance position; and by social factors in the group, e.g. general stability of male dominance relationships, presence of a strong alpha male, existence of special male-female relationships.

The primate community of the Lopé reserve, Gabon: Diets, responses to fruit scarcity, and effects on biomass

The diets of all diurnal primates (Gorilla g. gorilla, Pan t. troglodytes, Mandrillus sphinx, Colobus satanas, Cercocebus albigena, Cercopithecus nictitans, C. pogonias, C. cephus) in the Lopé Reserve, central Gabon, are described from qualitative and quantitative data collected over 10 years. A total of 397 foods were recorded, of which 91% were from plants. The diets of seven of the eight species were numerically dominated by fruit, the exception being Colobus satanas with a diet dominated by seeds. When proportion of feeding time was examined, fruit remained the dominant food class for six species, while Cercocebus albigena, like the black colobus, spent most time feeding on seeds. The number of foods recorded per species ranged from 46–220, but dietary breadth of the four species that have not been subjects of detailed study was certainly underestimated. Almost half of the foods (46%) were eaten by only one species, and only four (1%) were recorded for all eight species. At Lopé, fruit is scarce for 2–3 months during the annual dry season, and fruit crop failures can prolong this period of scarcity to as many as 8 months in some years. When fruit is scarce, the diets of all species changed to include more nonfruit foods, but chimpanzees and the three species of guenons maintained a fruit dominated diet. The total biomass of the Lopé primate community is 374 kg/km2, which is low compared to other sites in tropical Africa. Within the Lopé community, biomass correlates negatively with degree of frugivory, suggesting that fruit availability may be critical, but this does not explain the low total biomass compared to other sites. Neither floristic diversity nor the composition of the primate community at Lopé can account for the low biomass. While historical events such as climate‐induced changes in forest cover, disease, or impact of human activities cannot be ruled out, we conclude that the most likely explanation of low primate biomass is the occurrence of an ecological “catastrophe” in the fairly recent past from which populations have not yet recovered. We believe that the most likely scenario was climate‐mediated disturbance of flowering of important fruit food species. Prolonged fruit scarcity would have caused mortalities and affected the most frugivorous species more severely. Am. J. Primatol. 42:1–24, 1997.

Gorilla diet in the Lope Reserve, Gabon: A nutritional analysis

SummaryThe results of an analysis of gorilla diet in the Lopé Reserve, Gabon are presented. Samples were assayed for nutrients and plant secondary compounds (total phenols, condensed tannins and alkaloids) in an attempt to explain gorilla food choice. The diet is the most diverse so far analysed for gorillas; it seems to be a balance between sugary fruit, proteinaceous leaves, and relatively fibrous stems. Most fruits and herbaceous stems are succulent, but some drier, fibrous fruit and bark is also consumed. Seeds are another component of the diet, including unripe ones. Fruit, seeds, leaves and bark may all contain very high levels of total phenols and condensed tannins; but all herbaceous stems assayed contain low levels of these compounds. Alkaloids are not apparently a significant component of gorilla foods, and may be avoided. Gorillas at Lopé tend to avoid fatty fruit, and select leaves which are high in protein and low in fibre compared to the general vegetation. When fruit and preferred young leaves are scarce, proteinaceous barks and mature leaves, and sugary pith, are important sources of nutrients. We conclude that gorillas exploit the broad frugivore niche in West African lowland forests, and are part of the frugivore community there. What distinguishes them is their ability to eat large fibrous fruit, mature leaves and stems, and to overcome high levels of phenolics (we use “phenolics” as an umbrella term for both total phenols and condensed tannins). Gorilla diet at Lopé overlaps greatly with that of sympatric, frugivorous, primates, and resembles more closely that of chimpanzees than it does gorilla diet studied elsewhere in Africa.

The role of Pleistocene refugia and rivers in shaping gorilla genetic diversity in central Africa

The role of Pleistocene forest refugia and rivers in the evolutionary diversification of tropical biota has been the subject of considerable debate. A range-wide analysis of gorilla mitochondrial and nuclear variation was used to test the potential role of both refugia and rivers in shaping genetic diversity in current populations. Results reveal strong patterns of regional differentiation that are consistent with refugial hypotheses for central Africa. Four major mitochondrial haplogroups are evident with the greatest divergence between eastern (A, B) and western (C, D) gorillas. Coalescent simulations reject a model of recent east–west separation during the last glacial maximum but are consistent with a divergence time within the Pleistocene. Microsatellite data also support a similar regional pattern of population genetic structure. Signatures of demographic expansion were detected in eastern lowland (B) and Gabon/Congo (D3) mitochondrial haplogroups and are consistent with a history of postglacial expansion from formerly isolated refugia. Although most mitochondrial haplogroups are regionally defined, limited admixture is evident between neighboring haplogroups. Mantel tests reveal a significant isolation-by-distance effect among western lowland gorilla populations. However, mitochondrial genetic distances also correlate with the distance required to circumnavigate intervening rivers, indicating a possible role for rivers in partitioning gorilla genetic diversity. Comparative data are needed to evaluate the importance of both mechanisms of vicariance in other African rainforest taxa.

Chimpanzees in a hot, dry and open habitat: Mt. Assirik, Senegal, West Africa

The habitat of the chimpanzees of Mt. Assirik, in the Parc National du Niokolo-Koba, Senegal, is described in terms of rainfall, temperature and vegetation. The results are compared with those collected at five other sites of study elsewhere in Africa. Mt. Assirik is the driest site at which chimpanzees have been studied, in terms of annual rainfall, proportion of dry months, and number of rainy days. Mt. Assirik is also the hottest such site: the coolest mean maximum temperature at Mt. Assirik exceeds the hottest such temperature at any other site. Mt. Assirik is the only site where chimpanzees have been studied in which the majority of vegetation is grassland. Forest consitutes less than 3% of the surface area. In summary, Mt. Assirik presents a truly open savanna habitat and is thus unique amongst sites where chimpanzees have been studied. These results are compared with data from a tropical foraging human society, the !Kung San of southern Africa. The !Kung Sans habitat is drier on most (but not all) criteria, but Mt. Assirik is hotter. The climate and vegetation of Mt. Assirik strikingly resemble those reconstructed for the Plio-Pleistocene in eastern Africa. This suggests that the chimpanzees of Mt. Assirik provide a useful model for inferring the processes of adaptation in early hominids.

Nest building by lowland gorillas in the Lope Reserve, Gabon : Environmental influences and implications for censusing

We analyzed data from 373 fresh nest-sites (containing 2435 nests) of lowland gorillas (Gorilla g. gorilla)during a 4-year period in the Lopé Reserve, Gabon, to determine whether the observed variability in nest building was due to environmental influences. We recognized and defined seven types of nest in terms of the degree of construction and the raw materials used. Overall, nests built on the ground from herbaceous plants are the most common type (40%), followed by tree nests (35%). Frequencies of the different nest-types vary significantly between eight habitat-types. In habitat-types with high densities of understory herbs, ground nests predominated, but when herbs were rare, the majority of nests were in trees. A general preference for sleeping in herbaceous ground nests is indicated since trees are abundant in all habitat-types, except savanna. The frequency of nesting in trees shows a significant positive correlation with rainfall, but effects of climate are confounded by seasonal variation in use of different habitat-types. When elephants were attracted to the same localized food sources as gorillas, many tree nests were built even when herbs were available. We conclude that different nest-types reflect a variety of solutions to maximize comfort, depending on available raw materials and the probability of rainfall or disturbance by elephants or both factors. Nests are a powerful tool for population censuses and demographic studies of great apes, but problems exist in interpreting data on lowland gorilla nests. Results from this analysis show that only a third of nest-sites accurately reflects group size (of weaned individuals) and that 26% of all gorilla nest-sites could be mistaken for those of chimpanzees, as all nests, or all those visible from a transect, were in trees. Gorilla nests at Lopé were nonrandomly distributed with respect to habitat-types, and nest construction varied seasonally, thereby introducing sources of bias to transect nest counts. We discuss these problems and ones related to assessing the decay rate of nest-sites and make recommendations relevant to census work.

FRAGMENTED LIVING : BEHAVIOURAL ECOLOGY OF PRIMATES IN A FOREST FRAGMENT IN THE LOPE RESERVE, GABON

A 17-month study was made of the primates using a 9-ha “island” of forest, surrounded by savanna, in the northern part of the Lopé Reserve, Gabon. One group ofCercopithecus cephus (plus a young maleCercopithecus nictitans who was in permanent association with them) were resident in the fragment and groups of five other species of primates made visits during 127 days of observation:Pan troglodytes, 15 visits;Cercocebus albigena, 10;Colobus satanas, 3;Cercopithecus nictitans, 2;C. pogonias, 1. Visits were also made by lone males of three species,C. nictitans, Cercocebus albigena, andMandrillus sphinx. The eighth species of diurnal primate present at Lopé,Gorilla g. gorilla, did not visit the fragment during the study. Compared to conspecific groups in neighbouring continuous forest, primates in the fragment ate less fruit, seeds and flowers and more insects and leaves. The local population density of primates resident in the fragment was equivalent to that of the neighbouring continuous forest where all eight species occur, despite the diversity and abundance of fruit being less in the fragment. The costs imposed on the resident group by the reduced diversity and availability of preferred fruit foods appeared to be offset by a number of benefits that increased individual feeding efficiency for monkeys residing within a single fragment. These included lower travel costs, reduced feeding competition between individuals through group fission, and excellent knowledge of the location and quality of food resources in the small home range. It is also possible that the overall negative impact of inter-specific feeding competition was lower in fragments than in continuous forest and that micro-habitat differences resulted in an increased availability of palatable insect and leaf fallback foods in the fragment.

Wide-Ranging Chimpanzees at Mt. Assirik, Senegal

A 4-year study of the ecology and ethology of wild chimpanzees (Pan troglodytes verus)was carried out in far western Africa. Contacts with chimpanzees and the locations of their nests were noted to determine which types of habitat were most used and to estimate the density of the population and the size of its home range. The results show that this community has one of the lowest densities and largest home ranges of all populations of chimpanzees studied so far. As such, it may provide a useful model for the reconstruction of hominid evolution in the Plio-Pleistocene.

Social organization of savanna-dwelling chimpanzees,Pan troglodytes verus, at Mt. Assirik, Senegal

A community of chimpanzees at Mt. Assirik in south-eastern Senegal subsists in a hot, dry and open environment. This wide-ranging, savanna-living group provides an opportunity for comparisons of social organization with other population elsewhere in Africa living in forest and woodland. The group numbered about 28 over the four-year study, and its composition by age and sex was typical. The average size of parties (i.e., temporary sub-groups) did not differ from other populations. However, a high proportion of the group tended to remain together in such parties. The composition of parties resembled that found elsewhere, but some differences emerged between the wet and dry seasons. Larger, mixed parties containing adult males were much more common in open, non-forested habitats than were solitary individuals or parties without adult males. Large parties tended to form for travelling, especially for rapid movement over long distances. Such aspects of social organization seem unlikely to be related to the availability of food in any simple way. Instead, they seem to be adaptations to threat from predators and patchily distributed food, water and nesting sites. This results in occasional bivouacs and mass-migrations from one part of the home-range to another, especially in the dry season.