Could woodworking have driven lithic tool selection?

Understanding early stone tools, particularly relationships between form and function, is fundamental to understanding the behavioral evolution of early hominins. The oldest-claimed flake tools date to ca. 3.3 million years ago, and their development may represent a key step in hominin evolution. Flake form, and its relationship to function, has long been a focus of Paleolithic studies, almost exclusively with respect to meat acquisition. However, evidence for woodworking is now known from sites dating to 1.5 Ma. Additionally, Pan troglodytes are known to manufacture wooden tools for hunting and foraging, thus creating a phylogenetic (parsimony) argument for more ancient woodworking. However, few studies examining woodworking and Paleolithic tools have been completed to date. Indeed, it remains an open question whether woodworking may have instigated specific selective demands on the form of early stone tools. Here, we conducted an experiment testing the comparative woodworking efficiency (measured by time) of small and large flakes. Two groups of participants used either a relatively small or large unretouched flake to remove a predefined area from standardized samples of wood. Those using larger flakes were significantly more efficient (i.e., required less time) during this woodworking task. Our results demonstrate that larger flakes could have been preferentially chosen by hominins for woodworking, consistent with previous data generated experimentally in other (non-woodworking) tasks. Moreover, the production of relatively large flakes, such as those at Lomekwi, could have been motivated by woodworking, rather than, or in addition to, butchery. Such issues may also have encouraged the use of Levallois production strategies in later times.

Underestimating Kanzi? Exploring Kanzi-Oldowan comparisons in light of recent human stone tool replication.

The knapping experiments with Kanzi, a bonobo, are among the most insightful experiments into Oldowan technology ever undertaken. Comparison of his artifacts against archeological material, however, indicated he did not produce Oldowan lithic attributes precisely, prompting suggestions that this indicated cognitive or biomechanical impediments. The literature describing the learning environment provided to Kanzi, we suggest, indicates alternative factors. Based on consideration of wild chimpanzee learning environments, and experiments with modern knappers that have looked at learning environment, we contend that Kanzi's performance was impeded by an impoverished learning environment compared to those experienced by novice Oldowan knappers. Such issues are precisely those that might be tested via a repeat study, but in this case, practical and ethical constraints likely impede this possibility. We propose experiments that may be relevant to drawing conclusions from Kanzi's experiments that may not need to use non-human primates, thus bypassing some of these issues.

The relative efficacy of the cranium and os coxa for taxonomic assessment in macaques.

OBJECTIVES: The cranium is generally considered more reliable than the postcranium for assessing primate taxonomy, although recent research suggests that pelvic shape may be equally reliable. However, little research has focused on intrageneric taxonomic discrimination. Here, we test the relative taxonomic efficacy of the cranium and os coxa for differentiating two macaque species, with and without considering sexual dimorphism. MATERIALS AND METHODS: Geometric morphometric analyses were performed on cranial and os coxa landmarks for 119 adult Macaca fascicularis, M. mulatta, and Chlorocebus pygerythrus. Among-group shape variation was examined using canonical variates analyses. Cross-validated discriminant function analysis provided rates of correct group classification. Additionally, average morphological distances were compared with neutral genetic distances. RESULTS: Macaque species were clearly differentiated, both cranially and pelvically, when sex was not considered. Males were more often correctly classified based on the os coxa, while female classification rates were high for both morphologies. Female crania and male os coxa were differentiated approximately the same as genetic distance, while male crania were more similar (convergent), and female os coxa were more divergent than expected based on genetic distance. DISCUSSION: The hypothesis that cranial and os coxal shape can be used to discriminate among macaque species was supported. The cranium was better at differentiating females, while the os coxa was better at differentiating male macaques. Hence, there is no a priori reason for preferring the cranium when assessing intragenetic taxonomic relationships, but the effects of high levels of sexual dimorphism must be corrected for to accurately assess taxonomic signatures.

Assessing the relative impact of historical divergence and inter-group transmission on cultural patterns: a method from evolutionary ecology.

In the study of cultural evolution, observed among-group affinity patterns reflect the effects of processes such as mutation (e.g. innovation and copying error), between-group interaction (culture flow), drift and selection. As in biology, cultural affinity patterns are often spatially correlated, making it difficult to distinguish between the opposing geographically mediated forces of divergence and interaction, which cause groups to become more distinct or similar over time, respectively. Analogous difficulties are faced by evolutionary biologists examining the relationship between biological affinity and geography, particularly at lower taxonomic levels where the potential for gene flow between lineages is greatest. Tree models are generally used to assess the fit between affinity patterns and models of historical divergence. However, factors driving lineage divergence are often spatially mediated, resulting in tree models that are themselves geographically structured. Here, we showcase a simple method drawn from evolutionary ecology for assessing the relative impact of both geographically mediated processes simultaneously. We illustrate the method using global human craniometric diversity and material culture from the northern coast of New Guinea as example case studies. This method can be employed to quantify the relative importance of history (divergence) and geographically mediated between-group interaction (culture flow) in explaining observed cultural affinity patterns.This article is part of the theme issue 'Bridging cultural gaps: interdisciplinary studies in human cultural evolution'.

Transmission of biology and culture among post-contact Native Americans on the western Great Plains.

The transmission of genes and culture between human populations has major implications for understanding potential correlations between history, biological, and cultural variation. Understanding such dynamics in 19th century, post-contact Native Americans on the western Great Plains is especially challenging given passage of time, complexity of known dynamics, and difficulties of determining genetic patterns in historical populations for whom, even today, genetic data for their descendants are rare. Here, biometric data collected under the direction of Franz Boas from communities penecontemporaneous with the classic bison-hunting societies, were used as a proxy for genetic variation and analyzed together with cultural data. We show that both gene flow and "culture flow" among populations on the High Plains were mediated by geography, fitting a model of isolation-by-distance. Moreover, demographic and cultural exchange among these communities largely overrode the visible signal of the prior millennia of cultural and genetic histories of these populations.

Are human hands and feet affected by climate? A test of Allen's rule.

OBJECTIVES: In recent years, several studies have shown that populations from cold, high-latitude regions tend to have relatively shorter limbs than populations from tropical regions, with most of the difference due to the relative length of the zeugopods (i.e., radius, ulna, tibia, fibula). This pattern has been explained either as the consequence of long-term climatic selection or of phenotypic plasticity, with temperature having a direct effect on bone growth during development. The aims of this study were to test whether this pattern of intra-limb proportions extended to the bones of the hands and feet, and to determine whether the pattern remained significant after taking into account the effects of neutral evolutionary processes related to population history. MATERIALS AND METHODS: Measurements of the limb bones, including the first metatarsal and metacarpal, were collected for 393 individuals from 10 globally distributed human populations. The relationship between intra-limb indices and minimum temperature was tested using generalized least squares regression, correcting for spatial autocorrelation. RESULTS: The results confirmed previous observations of a temperature-related gradient in intra-limb proportions, even accounting for population history. This pattern extends to the hands, with populations from cold regions displaying a relatively shorter and stockier first metacarpal; however, the first metatarsal appears to be wider but not shorter in cold-adapted populations. DISCUSSION: The results suggest that climatic adaptation played a role in shaping variation in limb proportions between human populations. The different patterns shown by the hands and feet might be due to the presence of evolutionary constraints on the foot to maintain efficient bipedal locomotion.

A comparison of catarrhine genetic distances with pelvic and cranial morphology: implications for determining hominin phylogeny.

Inferring the evolutionary history of the hominins is necessarily reliant on comparative analyses of fossilized skeletal anatomy. However, the reliability of different primate skeletal regions for recovering phylogenetic relationships is currently poorly understood. Historically, postcranial variation has largely been conceived of as reflecting locomotory and postural adaptation. The shape of the os coxae is central to such discussions given the divergent morphology displayed by the bipedal hominin pelvis relative to other primate taxa. While previous cladistic studies have suggested that postcranial and cranial datasets do not differ in terms of their propensity for homoplasy, methodological issues such as the numbers of characters and their quantification make it difficult to evaluate these findings. Here, we circumvent these problems by constructing morphological distance matrices based on cranial, mandibular and os coxae three-dimensional shape. Statistical comparisons of these morphological distance matrices against a single genetic distance matrix for 11 catarrhine taxa show that cranial and os coxae shape reflect genetic relationships better than the mandible when humans are included, and that the cranium and os coxae do not differ statistically in terms of their genetic correlations. When humans were excluded from the analyses, all three anatomical regions were equally strongly correlated with genetic distance. Moreover, a second analysis focusing solely on os coxae variation of 16 taxa demonstrated that os coxae shape correctly recovers catarrhine taxonomic relationships at the sub-family level, even when humans are included. Taken together, our results suggest that there is no a priori reason to favor cranial shape data over os coxae morphology when inferring the genetic relationships of extant or extinct primate taxa. Morphological similarities between humans and other primates differ depending on the skeletal element, suggesting that combining skeletal elements into a single analysis may provide more accurate reconstructions of genetic relationships.

Dynamics of cultural transmission in Native Americans of the high Great Plains.

Culture is a phenomenon shared by all humans. Attempts to understand how dynamic factors affect the origin and distribution of cultural elements are, therefore, of interest to all humanity. As case studies go, understanding the distribution of cultural elements in Native American communities during the historical period of the Great Plains would seem a most challenging one. Famously, there is a mixture of powerful internal and external factors, creating-for a relatively brief period in time-a seemingly distinctive set of shared elements from a linguistically diverse set of peoples. This is known across the world as the "Great Plains culture." Here, quantitative analyses show how different processes operated on two sets of cultural traits among nine High Plains groups. Moccasin decorations exhibit a pattern consistent with geographically-mediated between-group interaction. However, group variations in the religious ceremony of the Sun Dance also reveal evidence of purifying cultural selection associated with historical biases, dividing down ancient linguistic lines. The latter shows that while the conglomeration of "Plains culture" may have been a product of merging new ideas with old, combined with cultural interchange between groups, the details of what was accepted, rejected or elaborated in each case reflected preexisting ideological biases. Although culture may sometimes be a "melting pot," the analyses show that even in highly fluid situations, cultural mosaics may be indirectly shaped by historical factors that are not always obvious.

The interaction of neutral evolutionary processes with climatically-driven adaptive changes in the 3D shape of the human os coxae.

Differences in the breadth of the pelvis among modern human populations and among extinct hominin species have often been interpreted in the light of thermoregulatory adaptation, whereby a larger pelvic girdle would help preserve body temperature in cold environments while a narrower pelvis would help dissipate heat in tropical climates. There is, however, a theoretical problem in interpreting a pattern of variation as evidence of selection without first accounting for the effects of neutral evolutionary processes (i.e., mutation, genetic drift and migration). Here, we analyse 3D configurations of 27 landmarks on the os coxae of 1494 modern human individuals representing 30 male and 23 female populations from five continents and a range of climatic conditions. We test for the effects of climate on the size and shape of the pelvic bone, while explicitly accounting for population history (i.e., geographically-mediated gene flow and genetic drift). We find that neutral processes account for a substantial proportion of shape variance in the human os coxae in both sexes. Beyond the neutral pattern due to population history, temperature is a significant predictor of shape and size variation in the os coxae, at least in males. The effect of climate on the shape of the pelvic bone, however, is comparatively limited, explaining only a small percentage of shape variation in males and females. In accordance with previous hypotheses, the size of the os coxae tends to increase with decreasing temperature, although the significance of the association is reduced when population history is taken into account. In conclusion, the shape and size of the human os coxae reflect both neutral evolutionary processes and climatically-driven adaptive changes. Neutral processes have a substantial effect on pelvic variation, suggesting such factors will need to be taken into account in future studies of human and fossil hominin coxal variation.

From cultural traditions to cumulative culture: parameterizing the differences between human and nonhuman culture.

Diverse species exhibit cultural traditions, i.e. population-specific profiles of socially learned traits, from songbird dialects to primate tool-use behaviours. However, only humans appear to possess cumulative culture, in which cultural traits increase in complexity over successive generations. Theoretically, it is currently unclear what factors give rise to these phenomena, and consequently why cultural traditions are found in several species but cumulative culture in only one. Here, we address this by constructing and analysing cultural evolutionary models of both phenomena that replicate empirically attestable levels of cultural variation and complexity in chimpanzees and humans. In our model of cultural traditions (Model 1), we find that realistic cultural variation between populations can be maintained even when individuals in different populations invent the same traits and migration between populations is frequent, and under a range of levels of social learning accuracy. This lends support to claims that putative cultural traditions are indeed cultural (rather than genetic) in origin, and suggests that cultural traditions should be widespread in species capable of social learning. Our model of cumulative culture (Model 2) indicates that both the accuracy of social learning and the number of cultural demonstrators interact to determine the complexity of a trait that can be maintained in a population. Combining these models (Model 3) creates two qualitatively distinct regimes in which there are either a few, simple traits, or many, complex traits. We suggest that these regimes correspond to nonhuman and human cultures, respectively. The rarity of cumulative culture in nature may result from this interaction between social learning accuracy and number of demonstrators.

Considering the role of time budgets on copy-error rates in material culture traditions: an experimental assessment.

Ethnographic research highlights that there are constraints placed on the time available to produce cultural artefacts in differing circumstances. Given that copying error, or cultural 'mutation', can have important implications for the evolutionary processes involved in material culture change, it is essential to explore empirically how such 'time constraints' affect patterns of artefactual variation. Here, we report an experiment that systematically tests whether, and how, varying time constraints affect shape copying error rates. A total of 90 participants copied the shape of a 3D 'target handaxe form' using a standardized foam block and a plastic knife. Three distinct 'time conditions' were examined, whereupon participants had either 20, 15, or 10 minutes to complete the task. One aim of this study was to determine whether reducing production time produced a proportional increase in copy error rates across all conditions, or whether the concept of a task specific 'threshold' might be a more appropriate manner to model the effect of time budgets on copy-error rates. We found that mean levels of shape copying error increased when production time was reduced. However, there were no statistically significant differences between the 20 minute and 15 minute conditions. Significant differences were only obtained between conditions when production time was reduced to 10 minutes. Hence, our results more strongly support the hypothesis that the effects of time constraints on copying error are best modelled according to a 'threshold' effect, below which mutation rates increase more markedly. Our results also suggest that 'time budgets' available in the past will have generated varying patterns of shape variation, potentially affecting spatial and temporal trends seen in the archaeological record. Hence, 'time-budgeting' factors need to be given greater consideration in evolutionary models of material culture change.

Understanding the comparative catarrhine context of human pelvic form: a 3D geometric morphometric analysis.

Comparative studies of catarrhine pelvic morphology in an evolutionary framework play an important role in paleoanthropology, especially since this is the context from which human bipedalism eventually arose. Given the abundance of potentially confounding evolutionary and mechanical factors influencing pelvic form, it is important to tease apart the effects of shape and size in the major component of the primate pelvis, the os coxae. However, os coxae form is difficult to assess via traditional morphometric methods. Here, we adopt a 3D geometric morphometric approach to landmark data. Our analyses included data from 30 extant catarrhine taxa. Data were transformed and registered using Procrustes analysis and analyzed via examination of principal components. Two analyses were performed: one excluding Homo sapiens, and a second including them. Results of the first analysis demonstrate that the total diversity of os coxae morphology is significantly greater in hominoids than it is in cercopithecoids. This appears to be driven by the greater effects of size diversity (i.e., allometric effects) in the case of the hominoids. This analysis also revealed a clear taxonomic/phylogenetic distinction between hominoids and cercopithecoids in terms of os coxae shape. The second analysis showed that Procrustes distances in shape space are significantly greater between extant Pan and Homo than they are between any two non-human catarrhine taxa. This analysis thus quantifies, on a comparative basis, the dramatic effect that the course of hominin evolution had upon the morphology of the human pelvis, within what is - even by catarrhine standards--a relatively short span of evolutionary time.

Global geometric morphometric analyses of the human pelvis reveal substantial neutral population history effects, even across sexes.

Recent applications of population genetic models to human craniodental traits have revealed a strong neutral component to patterns of global variation. However, little work has been undertaken to determine whether neutral processes might also be influencing the postcranium, perhaps due to substantial evidence for selection and plastic environmental responses in these regions. Recent work has provided evidence for neutral effects in the pelvis, but has been limited in regard to shape data (small numbers of linear measurements) and restricted only to males. Here, we use geometric morphometric methods to examine population variation in the human os coxae (pelvic bone) in both males and females. Neutrality is examined via apportionment of variance patterns and fit to an Out-of-Africa serial founder effect model, which is known to structure neutral genetic patterns. Moreover, we compare males and females directly, and the true versus false pelvis, in order to examine potential obstetrical effects. Our results indicate evidence for substantial neutral population history effects on pelvic shape variation. They also reveal evidence for the effect of obstetrical constraints, but these affect males and females to equivalent extents. Our results do not deny an important role for selection in regard to specific aspects of human pelvic variation, especially in terms of features associated with body size and proportions. However, our analyses demonstrate that at a global level, the shape of the os coxae reveals substantial evidence for neutral variation. Our analyses thus indicate that population variation in the human pelvis might be used to address important questions concerning population history, just as the human cranium has done.

Late Pleistocene climate change and the global expansion of anatomically modern humans.

The extent to which past climate change has dictated the pattern and timing of the out-of-Africa expansion by anatomically modern humans is currently unclear [Stewart JR, Stringer CB (2012) Science 335:1317-1321]. In particular, the incompleteness of the fossil record makes it difficult to quantify the effect of climate. Here, we take a different approach to this problem; rather than relying on the appearance of fossils or archaeological evidence to determine arrival times in different parts of the world, we use patterns of genetic variation in modern human populations to determine the plausibility of past demographic parameters. We develop a spatially explicit model of the expansion of anatomically modern humans and use climate reconstructions over the past 120 ky based on the Hadley Centre global climate model HadCM3 to quantify the possible effects of climate on human demography. The combinations of demographic parameters compatible with the current genetic makeup of worldwide populations indicate a clear effect of climate on past population densities. Our estimates of this effect, based on population genetics, capture the observed relationship between current climate and population density in modern hunter-gatherers worldwide, providing supporting evidence for the realism of our approach. Furthermore, although we did not use any archaeological and anthropological data to inform the model, the arrival times in different continents predicted by our model are also broadly consistent with the fossil and archaeological records. Our framework provides the most accurate spatiotemporal reconstruction of human demographic history available at present and will allow for a greater integration of genetic and archaeological evidence.

Human pelvis and long bones reveal differential preservation of ancient population history and migration out of Africa.

One of the main events in the history of our species has been our expansion out of Africa. A clear signature of this expansion has been found on global patterns of neutral genetic variation, whereby a serial founder effect accompanied the colonization of new regions, in turn creating a wilhin-pupulation decrease in neutral genetic diversity with increasing distance from Africa. This same distinctive pattern has also been described for cranial and dental morphological variation in human populations distributed across the globe. Here, we used a data set of postcranial linear measurements for 30 globally distributed human populations, and a climatic data set of minimum annual temperature, maximum annual temperature, and precipitation in order to separate for the first time the relative effect of neutral demographic processes and climatic selection on four long (limb) bones (femur, tibia, radius, and humerus) versus the pelvic bones of the human appendicular skeleton. We implemented a stepwise regression procedure in which phenotypic variance is assumed to be affected by the iterative founder events that accompanied human expansion from Africa, as well as by climate. This model included, as independent factors, geographic distance from central Africa, the three climatic variables, and all possible interactions between the three climatic variables. We excluded all nonsignificant factors by backward stepwise elimination with the aim of identifying the minimal model significantly explaining variation in the phenotypic data. Our results indicate a sharp difference in the way the pelvis and the limb bones reflect the neutral signature of the out-of-Africa expansion. Consistent with previous analyses of the cranium and dentition, pelvic shape variation shows a significant within-population decrease with increasing distance from Africa. However, no such pattern could be found in the long bones. Rather, in the case of both the tibia and the femur, a significant relationship between population-level variance and minimum temperature was demonstrated. Hence, in the case of these limb bones, it is probable that the effects of climatic selection have obliterated the demographic signature of human dispersal from Africa. Our finding mat pelvic variation exhibits the neutral effects of demographic history suggests that consideration of this skeletal element might be used to shed light on factors of human population history, just as the cranium has done.

Comparison of handaxes from Bose Basin (China) and the western Acheulean indicates convergence of form, not cognitive differences.

Alleged differences between Palaeolithic assemblages from eastern Asia and the west have been the focus of controversial discussion for over half a century, most famously in terms of the so-called 'Movius Line'. Recent discussion has centered on issues of comparability between handaxes from eastern Asian and 'Acheulean' examples from western portions of the Old World. Here, we present a multivariate morphometric analysis in order to more fully document how Mid-Pleistocene (i.e. ∼803 Kyr) handaxes from Bose Basin, China compare to examples from the west, as well as with additional (Mode 1) cores from across the Old World. Results show that handaxes from both the western Old World and Bose are significantly different from the Mode 1 cores, suggesting a gross comparability with regard to functionally-related form. Results also demonstrate overlap between the ranges of shape variation in Acheulean handaxes and those from Bose, demonstrating that neither raw material nor cognitive factors were an absolute impediment to Bose hominins in making comparable handaxe forms to their hominin kin west of the Movius Line. However, the shapes of western handaxes are different from the Bose examples to a statistically significant degree. Moreover, the handaxe assemblages from the western Old World are all more similar to each other than any individual assemblage is to the Bose handaxes. Variation in handaxe form is also comparatively high for the Bose material, consistent with suggestions that they represent an emergent, convergent instance of handaxe technology authored by Pleistocene hominins with cognitive capacities directly comparable to those of 'Acheulean' hominins.

Why Levallois? A morphometric comparison of experimental 'preferential' Levallois flakes versus debitage flakes.

BACKGROUND: Middle Palaeolithic stone artefacts referred to as 'Levallois' have caused considerable debate regarding issues of technological predetermination, cognition and linguistic capacities in extinct hominins. Their association with both Neanderthals and early modern humans has, in particular, fuelled such debate. Yet, controversy exists regarding the extent of 'predetermination' and 'standardization' in so-called 'preferential Levallois flakes' (PLFs). METHODOLOGY/PRINCIPAL FINDINGS: Using an experimental and morphometric approach, we assess the degree of standardization in PLFs compared to the flakes produced during their manufacture. PLFs possess specific properties that unite them robustly as a group or 'category' of flake. The properties that do so, relate most strongly to relative flake thicknesses across their surface area. PLFs also exhibit significantly less variability than the flakes generated during their production. Again, this is most evident in flake thickness variables. A further aim of our study was to assess whether the particular PLF attributes identified during our analyses can be related to current knowledge regarding flake functionality and utility. CONCLUSIONS/SIGNIFICANCE: PLFs are standardized in such a manner that they may be considered 'predetermined' with regard to a specific set of properties that distinguishes them statistically from a majority of other flakes. Moreover, their attributes can be linked to factors that, based on current knowledge, are desirable features in flake tools (e.g. durability, capacity for retouch, and reduction of torque). As such, our results support the hypothesis that the lengthy, multi-phase, and hierarchically organized process of Levallois reduction was a deliberate, engineered strategy orientated toward specific goals. In turn, our results support suggestions that Levallois knapping relied on a cognitive capacity for long-term working memory. This is consistent with recent evidence suggesting that cognitive distinctions between later Pleistocene hominins such as the Neanderthals and anatomically modern humans were not as sharp as some scholars have previously suggested.

The importance of history in definitions of culture: Implications from phylogenetic approaches to the study of social learning in chimpanzees.

Some have claimed that wild chimpanzees possess multiple socially learned traditions that might constitute cultural patterns. Others, however, have suggested that even fundamental alternative explanations, such as proximate genetic mechanisms, have not been addressed satisfactorily. Multiple analyses using phylogenetic (cladistic) methods, however, have been shown not to support the genetic proposition. Rather, such analyses are more consistent with the growing body of evidence from studies of both wild and captive animals suggesting that behavioral patterns in wild chimpanzees are socially learned. The question remains, however, as to whether, from a scientific viewpoint, it is useful to term such patterns cultural. It is argued here that cultural mosaics of multiple behaviors that differ intercommunally, both in humans and chimpanzees, are an emergent property of a phylogenetic (i.e., historical) process of descent with modification, mediated by mechanisms of social transmission, variation, and sorting through time. This historical perspective is productive when attempting to consider the phenomenon of culture across species.

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.