The ecology of primate material culture

Tool use in extant primates may inform our understanding of the conditions that favoured the expansion of hominin technology and material culture. The ‘method of exclusion’ has, arguably, confirmed the presence of culture in wild animal populations by excluding ecological and genetic explanations for geographical variation in behaviour. However, this method neglects ecological influences on culture, which, ironically, may be critical for understanding technology and thus material culture. We review all the current evidence for the role of ecology in shaping material culture in three habitual tool-using non-human primates: chimpanzees, orangutans and capuchin monkeys. We show that environmental opportunity, rather than necessity, is the main driver. We argue that a better understanding of primate technology requires explicit investigation of the role of ecological conditions. We propose a model in which three sets of factors, namely environment, sociality and cognition, influence invention, transmission and retention of material culture.     

Hominization in the rainforest: The chimpanzee's piece of the puzzle

Many models of human evolution propose that key behavioral innovations were involved in the divergence of the human line from the ape line. An ecological reason, the shift from dense forests to a more open habitat, is suggested as the basis for these innovations. Primate models can be useful to our understanding of how environmental factors can affect such key behaviors. New results on forest chimpanzees demonstrate that for most patterns of behavior considered to favor a savanna model, the environmental influences are in opposition to the expectations. A new evaluation of the environmental influences on human evolution is required. We propose that either patterns of key human behaviors were inaccurately distinguished or that the hominization process started while the common ancestor of the chimpanzee and man was alive.     

Are behavioral differences among wild chimpanzee communities genetic or cultural? An assessment using tool‐use data and phylogenetic methods

Over the last 30 years it has become increasingly apparent that there are many behavioral differences among wild communities of Pan troglodytes. Some researchers argue these differences are a consequence of the behaviors being socially learned, and thus may be considered cultural. Others contend that the available evidence is too weak to discount the alternative possibility that the behaviors are genetically determined. Previous phylogenetic analyses of chimpanzee behavior have not supported the predictions of the genetic hypothesis. However, the results of these studies are potentially problematic because the behavioral sample employed did not include communities from central Africa. Here, we present the results of a study designed to address this shortcoming. We carried out cladistic analyses of presence/absence data pertaining to 19 tool‐use behaviors in 10 different P. troglodytes communities plus an outgroup (P. paniscus). Genetic data indicate that chimpanzee communities in West Africa are well differentiated from those in eastern and central Africa, while the latter are not reciprocally monophyletic. Thus, we predicted that if the genetic hypothesis is correct, the tool‐use data should mirror the genetic data in terms of structure. The three measures of phylogenetic structure we employed (the Retention Index, the bootstrap, and the Permutation Tail Probability Test) did not support the genetic hypothesis. They were all lower when all 10 communities were included than when the three western African communities are excluded. Hence, our study refutes the genetic hypothesis and provides further evidence that patterns of behavior in chimpanzees are the product of social learning and therefore meet the main condition for culture. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Why Don't Chimpanzees in Gabon Crack Nuts?

Some populations of wild chimpanzees (Pan troglodytes) use hammers and anvils of stone or wood to crack open nuts for food. Others do not. The aim of this study was to ask why one non-nut-cracking population, in the Lopé Reserve, Gabon, lacks this useful form of tool use. We tested 10 hypotheses: (1) nuts are absent; (2) nuts are few; (3) nuts are unsuitable; (4) hammers are absent; (5) hammers are unsuitable; (6) anvils are absent; (7) anvils are unsuitable; (8) nuts are displaced by better food items; (9) intelligence is insufficient; and (10) knowledge is insufficient. All but the last are clearly falsified, leaving by exclusion the likelihood that Lopé's chimpanzees lack the technology—knowledge of appropriate technique—to exploit this resource. Thus, the behavioral differences across populations of these apes are cultural and not environmentally dictated. This explanation is congruent with the distribution of chimpanzee nut-cracking across Africa.     

Digital dermatoglyphic patterns in a sample of the Nigerian population

Eight hundred Nigerians (400 males and 400 females) were screened for the analysis of digital dermatoglyphic patterns. The frequency distribution of the patterns are 52.09%, 27.67%, 19.20% and 1.13% for ulnar loops, whorls, arches and radial loops respectively. Except for arches, the males have more of the patterns than females. The pattern intensity indices for males and females are 11.2 and 9.9 respectively. Bilateral symmetry is similar in both sexes and the percent distribution is as high as 81.5% and 84% on digit V in males and females respectively.     

Flexible and Persistent Tool-using Strategies in Honey-gathering by Wild Chimpanzees

Several populations of wild chimpanzees use tools to raid bee nests, but preliminary observations of chimpanzees in the Congo Basin indicate that they may have developed sophisticated technical solutions to gather honey that differ from those of apes in other regions. Despite the lack of habituated groups within the range of the central subspecies, there have been several reports of different types of tools used by chimpanzees to open beehives and gather honey. Researchers have observed some of these behaviors (honey dipping) in populations of western and eastern chimpanzees, whereas others (hive pounding) may be limited to this region. Toward evaluating hypotheses of regional tool using patterns, we provide the first repeated direct observations and systematic documentation of tool use in honey-gathering by a population of Pan troglodytes troglodytes. Between 2002 and 2006, we observed 40 episodes of tool use in honey-gathering by chimpanzees in the Goualougo Triangle, Republic of Congo. Pounding was the most common and successful strategy to open beehives. Chimpanzees at this site used several tools in a single tool-using episode and could also attribute multiple functions to a single tool. They exhibited flexibility in responses toward progress in opening a hive and hierarchical structuring of tool sequences. Our results support suggestions of regional tool using traditions in honey-gathering, which could be shaped by variation in bee ecology across the chimpanzee range. Further, we suggest that these chimpanzees may have an enhanced propensity to use tool sets that could be related to other aspects of their tool repertoire. Clearly, there is still much to be learned about the behavioral diversity of chimpanzees residing within the Congo Basin.     

Systematics value of stomata in some Nigerian hardwood species of Fabaceae

ABSTRACT

Epidermal morphology, and the structure and development of stomata in 10 species of Fabaceae are described. The epidermal cells varied from irregular to straight-walled and in some taxa sinuous patterns were observed. The leaves are hypostomatic. Anomocytic stomata follow an agenous ontogenetic pathway, while paracytic stomata are eumesogenously derived. These stomatal features are considered to be of systematic value in delimiting hardwood species in this family.     

Socioecological Influences on Tool Use in Captive Chimpanzees

Some tool use in wild chimpanzees is seasonal because they follow the patterns of availability of resources. The type and abundance of vegetation affect the selection of materials used for tools. Besides triggering, does seasonality also shape tool use? We tested the tool performance of 2 chimpanzee groups in a large outdoor compound, enriched with hundreds of plants. We conducted the experiments in 4 seasons to test their behavioural adaptation to the environmental changes. Initially the groups showed preference for different tool materials, one group using grass, abundant in summer, the other using twigs. While twigs were constant through the year, the availability of grass fluctuated greatly, affecting the number of insertions and success of the individuals that used it. Therefore, seasonality did not affect the performance of the group that preferred twigs (less abundant but constant) as it affected the performance of the group that preferred grass (plentiful in certain seasons and scarce in others). We recorded several cases of observation, which may have biased the choices of material. Analysis of the episodes and first insertions of some subjects suggests social transmission, which might explain the initial intergroup difference in tool preference.     

Measurement of tree diversity in the Nigerian rainforest

The diversity of trees 5 cm in diameter and above at breast height was studied at five sites within the Nigerian rainforest. The sites were Gambari, Oban, Omo, Owan and Sapoba forest reserves. Diversity was measured in terms of similarity and heterogeneity and explored at beta and gamma levels. Over 315 species were encountered during the study. In terms of species richness, there are 80, 226, 163, 137 and 159 species in Gambari, Oban, Omo, Owan and Sapoba, respectively. The occurrence of the species showed that, while 26 species are common to all the sites, 62, 17, 15, 8 and 6 species are restricted to Oban, Omo, Sapoba, Gambari and Owan, respectively. Ordination by detrended correspondence analysis indicated gradients along the first two axes. The first axis reflecting rainfall gradient which decreases from Oban through to Sapoba, Owan and Gambari. The second axis showed a separation of Omo, and Owan and Sapoba. This suggests a gradient of soil parent materials, the former site is on the basement complex while the latter is on the cretaceous rock. It is suggested that each forest should be managed based on its particular characteristics, such as species composition and abundance. Thus, forests with high species diversity but fewer numbers of individuals, such as in Oban, should be earmarked for conservation while species-rich forests with relatively more economic species, as in Sapoba and Omo, should be managed for sustained timber production. Floristically poor forest, such as those in Gambari and Owan, should be converted to plantations of fast growing species but with care taken to avoid exposing such sites to soil erosion.     

New Cases of Intergroup Violence Among Chimpanzees in Gombe National Park,Tanzania

Despite considerable attention to chimpanzee intergroup violence, the number of observed cases remains small. We report 4 cases of intergroup violence that occurred in Gombe National Park, Tanzania, between 1993 and 2002. We observed (3 cases) or inferred (1 case) males from the Kasekela community to attack members of their 2 neighboring communities: Mitumba and Kalande. In 1993, Kasekela males killed and ate a female infant from Mitumba. In 1998, Kasekela males captured 2 infants (sex unknown) from Kalande, one of which escaped and the other was killed and eaten. Also in 1998, Kasekela males attacked an adolescent male from Kalande. The victim was alive but severely injured by the end of the attack. The intensity and duration of the attack are comparable to other attacks that resulted in fatal injuries. In 2002, observers found the body of an adolescent male from Mitumba following an incursion by Kasekela males into the area. The injuries inflicted on the Mitumba male together with circumstantial evidence suggest that Kasekela males killed him. The attacks support the view that intergroup violence is a persistent feature of chimpanzee societies and that the primary benefit attackers gain from them is reduced competition for resources.     

Soils Consumed by Chimpanzees of the Kanyawara Community in the Kibale Forest,Uganda

We previously reported on a study of 4 soils that chimpanzees of the Kanyawara community in the Kibale National Park, Uganda consumed on a near-daily basis. We suggested that iron was a possible chemical stimulus in association with high quantities of Si:Al = 1:1-dominated clay minerals in the consumed material. To test our initial findings, we analyzed 18 samples from the same general area including 7 samples that the chimpanzees did not eat. Among the chemical elements, As, Au, Br, Ca, Cl, Dy, Mg, Ni, Sb, Sr, and I are below detection limits. Only Fe stands out as a potentially important nutritional element present in sufficient quantity to provide a physiological stimulus for chimpanzees living at high elevations near the flanks of the Ruwenzori Mountains. Along with Fe, metahalloysite is present in high amounts in these soils. In its pure crystalline form as a pharmaceutical grade clay mineral metahalloysite may well counteract the debilitating effects of diarrhea, with an effect similar to what is achieved with kaolinite (cf. Kaopectate^TM). An unexpected result, the relatively high nitrogen and carbon in the eaten samples relative to the uneaten group, indicates the chimpanzees may have a higher threshold for organic-rich material than previously believed. Contrarily, the color of the ingested material, depicts a material with less humus than in the uneaten group, a finding that is compatible with previous work reported at other geophagy sites in Africa. Of all the choices of soil available to them, the chimpanzees appear to be selecting highly homogeneous chemical natural earths with well-leached and uniform mineralogical material similar to the uneaten group, but with higher relative amounts of clay size material.