Vernon Reynolds

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.

CHARTING CULTURAL VARIATION IN CHIMPANZEES

Cultural variation among chimpanzee communities or unit-groups at nine long-term study sites was charted through a systematic, collaborative procedure in which the directors of the sites first agreed a candidate list of 65 behaviour patterns (here fully defined), then classified each pattern in relation to its local frequency of occurrence. Thirty-nine of the candidate behaviour patterns were discriminated as cultural variants, sufficiently frequent at one or more sites to be consistent with social transmission, yet absent at one or more others where environmental explanations were rejected. Each community exhibited a unique and substantial profile of such variants, far exceeding cultural variation reported before for any other non-human species. Evaluation of these pan-African distributions against three models for the diffusion of traditions identified multiple cases consistent with cultural evolution involving differentiation in form, function and targets of behaviour patterns.

Aging and fertility patterns in wild chimpanzees provide insights into the evolution of menopause

Human menopause is remarkable in that reproductive senescence is markedly accelerated relative to somatic aging, leaving an extended postreproductive period for a large proportion of women. Functional explanations for this are debated, in part because comparative data from closely related species are inadequate. Existing studies of chimpanzees are based on very small samples and have not provided clear conclusions about the reproductive function of aging females. These studies have not examined whether reproductive senescence in chimpanzees exceeds the pace of general aging, as in humans, or occurs in parallel with declines in overall health, as in many other animals. In order to remedy these problems, we examined fertility and mortality patterns in six free-living chimpanzee populations. Chimpanzee and human birth rates show similar patterns of decline beginning in the fourth decade, suggesting that the physiology of reproductive senescence was relatively conserved in human evolution. However, in contrast to humans, chimpanzee fertility declines are consistent with declines in survivorship, and healthy females maintain high birth rates late into life. Thus, in contrast to recent claims, we find no evidence that menopause is a typical characteristic of chimpanzee life histories.

To what extent does living in a group mean living with kin

Chimpanzees live in large groups featuring remarkable levels of gregariousness and cooperation among the males. Because males stay in their natal communities their entire lives and are hence expected to be living with male relatives, cooperation is therefore assumed to occur within one large ‘family’ group. However, we found that the average relatedness among males within several chimpanzee groups as determined by microsatellite analysis is in fact rather low, and only rarely significantly higher than average relatedness of females in the groups or of males compared across groups. To explain these findings, mathematical predictions for average relatedness according to group size, reproductive skew and sex bias in dispersal were derived. The results show that high average relatedness among the philopatric sex is only expected in very small groups, which is confirmed by a comparison with published data. Our study therefore suggests that interactions among larger number of individuals may not be primarily driven by kin relationships.

Censusing Chimpanzees in the Budongo Forest, Uganda

It is difficult to make valid comparisons of chimpanzee densities among sites because observers calculate them using different methods. We argue that nest count estimates of density are preferable to densities from home range estimates because of the problems of defining home range. There are many problems associated with nest count methods, some of which have not been addressed in previous studies. In 1992, we censused chimpanzees in the Budongo Forest using three methods;the standing crop nest count (SCNC), the marked nest count (MNC), and visual sightings of the animals (VS). Each method is based on standard line transect techniques. Of 96 nests monitored for decay rate,those constructed in the dry seasons decayed faster than those in the wet seasons. All- day follows of individual chimpanzees and observations of nesting chimpanzees at dusk showed that about 15.8% of night nests were reused,17.5% of the population did not build nests, and 18.8% of nests were first constructed as day nests. Given the variability in nest decay rates, we argue that MNC is a better method than SCNC because it avoids having to calculate decay rates.

Nesting Behavior of Chimpanzees: Implications for Censuses

Chimpanzees have been censused using nest counting techniques since the mid 1970s. The use of nest counts makes several assumptions which have never been tested: (a) that the visibility of nests does not change with the height of the nest in the canopy; (b) that weaned chimpanzees construct on average one nest per day; (c) that establishing census lines has no effect on the nesting behavior of the chimpanzees; and (d) that the presence of snare injuries, common in forests in eastern Africa, does not affect nest construction. Tests of these assumptions in the Budongo Forest in Uganda showed that visibility of nests at different heights in the canopy is not different from the true distribution of nests but that the other assumptions are false. Minimizing human use could limit the effects of line transects on censuses, but the other two assumptions need to be corrected for.

Genetic and ‘cultural’ similarity in wild chimpanzees

The question of whether animals possess ‘cultures’ or ‘traditions’ continues to generate widespread theoretical and empirical interest. Studies of wild chimpanzees have featured prominently in this discussion, as the dominant approach used to identify culture in wild animals was first applied to them. This procedure, the ‘method of exclusion,’ begins by documenting behavioural differences between groups and then infers the existence of culture by eliminating ecological explanations for their occurrence. The validity of this approach has been questioned because genetic differences between groups have not explicitly been ruled out as a factor contributing to between-group differences in behaviour. Here we investigate this issue directly by analysing genetic and behavioural data from nine groups of wild chimpanzees. We find that the overall levels of genetic and behavioural dissimilarity between groups are highly and statistically significantly correlated. Additional analyses show that only a very small number of behaviours vary between genetically similar groups, and that there is no obvious pattern as to which classes of behaviours (e.g. tool-use versus communicative) have a distribution that matches patterns of between-group genetic dissimilarity. These results indicate that genetic dissimilarity cannot be eliminated as playing a major role in generating group differences in chimpanzee behaviour.

Food Supply and Chimpanzee (Pan troglodytes schweinfurthii) Party Size in the Budongo Forest Reserve, Uganda

A central issue in socioecology is the nature of the relationship between an organisms environment and its social structure. In chimpanzees, the fission-fusion social system is thought to minimize feeding competition for primary dietary components: ephemeral, dispersed patches of ripe fruit. Intragroup feeding competition is thought to force individuals into small parties. Informal observations in the Sonso region of the Budongo forest had suggested that in this habitat, food supply was such that feeding competition was less important in determining grouping patterns than elsewhere. We used data collected on food supply and party sizes over a 4-year period to investigate this suggestion. In accord with theoretical expectation, sizes of foraging parties fluctuated with the size of food patches. However, party sizes showed either negative or no relationship with habitat-wide measures of food abundance. Likewise party sizes showed little relationship to overall measures of food dispersion. For important dietary items, both fruit and leaves had patchy distributions, though the degree of clumping was not strong, and fruit was not more clumped than leaves. Generally, abundant food appeared to be less patchy, and chimpanzees appeared to use more patches as food became more abundant rather than forming larger parties. We suggest that both dispersal and abundance need to be considered when investigating the impact of food supply on grouping patterns, and that the importance of food as a factor in determining chimpanzee grouping patterns declines with increasing levels of abundance.

The Genetic Signature of Sex-Biased Migration in Patrilocal Chimpanzees and Humans

A large body of theoretical work suggests that analyses of variation at the maternally inherited mitochondrial (mt)DNA and the paternally inherited non-recombining portion of the Y chromosome (NRY) are a potentially powerful way to reveal the differing migratory histories of men and women across human societies. However, the few empirical studies comparing mtDNA and NRY variation and known patterns of sex-biased migration have produced conflicting results. Here we review some methodological reasons for these inconsistencies, and take them into account to provide an unbiased characterization of mtDNA and NRY variation in chimpanzees, one of the few mammalian taxa where males routinely remain in and females typically disperse from their natal groups. We show that patterns of mtDNA and NRY variation are more strongly contrasting in patrilocal chimpanzees compared with patrilocal human societies. The chimpanzee data we present here thus provide a valuable comparative benchmark of the patterns of mtDNA and NRY variation to be expected in a society with extremely female-biased dispersal.

Paternity and social rank in wild chimpanzees (Pan troglodytes) from the Budongo Forest, Uganda

We analyzed patterns of paternity and male dominance rank in the Sonso community of wild East African chimpanzees (Pan troglodytes schweinfurthii) in the Budongo Forest, Uganda. Our major objective was to determine whether and how social rank influenced paternity success. We successfully genotyped 52 individuals at up to nine microsatellite loci, using DNA extracted from fecal samples. Of 24 offspring analyzed, we identified sires for 21. Paternity success was significantly correlated with social rank, with alpha males siring a disproportionate number of offspring. However, both middle- and low-ranking males also fathered offspring, and the priority-of-access model provided a relatively poor prediction of which males would be successful and under what circumstances. The concentration of paternities among only seven males and the tendency for high-ranking males to sire offspring of multiparous females suggest that both individual variation in male quality and the resource value of particular females may be mediating factors. In comparison with other chimpanzee studies, our results support the hypothesis that larger male cohort size reduces the ability of the alpha male to monopolize females, though within our study, male number did not affect the success of the alpha. Successful sires were not necessarily those who achieved the highest mating success with the females whose offspring they sired, but were those who demonstrated higher investment by spending significantly more time in association with these females. Finally, we estimate extra-group paternity at 0-5%, supporting other evidence that the community serves as the primary reproductive unit in chimpanzees.