Masticatory form and function in the African apes

This study examines variability in masticatory morphology as a function of dietary preference among the African apes. The African apes differ in the degree to which they consume leaves and other fibrous vegetation. Gorilla gorilla beringei, the eastern mountain gorilla, consumes the most restricted diet comprised of mechanically resistant foods such as leaves, pith, bark, and bamboo. Gorilla gorilla gorilla, the western lowland gorilla subspecies, consumes leaves and other terrestrial herbaceous vegetation (THV) but also consumes a fair amount of ripe, fleshy fruit. In contrast to gorillas, chimpanzees are frugivores and rely on vegetation primarily as fallback foods. However, there has been a long-standing debate regarding whether Pan paniscus, the pygmy chimpanzee (or bonobo), consumes greater quantities of THV as compared to Pan troglodytes, the common chimpanzee. Because consumption of resistant foods involves more daily chewing cycles and may require larger average bite force, the mechanical demands placed on the masticatory system are expected to be greater in folivores as compared to primates that consume large quantities of fleshy fruit. Therefore, more folivorous taxa are predicted to exhibit features that improve load-resistance capabilities and increase force production. To test this hypothesis, jaw and skull dimensions were compared in ontogenetic series of G. g. beringei, G. g. gorilla, P. t. troglodytes, and P. paniscus. Controlling for the influence of allometry, results show that compared to both chimpanzees and bonobos, gorillas exhibit some features of the jaw complex that are suggestive of improved masticatory efficiency. For example, compared to all other taxa, G. g. beringei has a significantly wider mandibular corpus and symphysis, larger area for the masseter muscle, higher mandibular ramus, and higher mandibular condyle relative to the occlusal plane of the mandible. However, the significantly wider mandibular symphysis may be an architectural response to increasing symphyseal curvature with interspecific increase in size. Moreover, Gorilla and Pan do not vary consistently in all features, and some differences run counter to predictions based on dietary variation. Thus, the morphological responses are not entirely consonant with predictions based on hypothesized loading regimes. Finally, despite morphological differences between bonobos and chimpanzees, there is no systematic pattern of differentiation that can be clearly linked to differences in diet. Results indicate that while some features may be linked to differences in diet among the African apes, diet alone cannot account for the patterns of morphological variation demonstrated in this study. Allometric constraints and dental development also appear to play a role in morphological differentiation among the African apes.     

Systematic relationships of gorillas from Kahuzi Tshiaberimu and Kayonza

Previous classifications of gorillas from Mt. Kahuzi, Mt. Tshiaberimu and the Kayonza Forest, placing them in Gorilla gorilla graueri or G.g. beringei, somewhat over-simplify a complex situation. Both Kahuzi and Tshiaberimu gorillas are close to graueri and should be placed in that subspecies, but each (in different ways) shows some approach to beringei, either through independent adaptation to extreme montane conditions, or because they may be points along a (now disrupted) cline from one race to the other. A hypothesis for the dispersal of beringei is presented, making use of geophysical data on the movement of the African plate over the Virunga 'hot-spot'.     

Diet and Mandibular Morphology in African Apes

Investigations seeking to understand the relationship between mandibular form, function, and dietary behavior have focused on the mandibular corpus and symphysis. African apes vary along a gradient of folivory/frugivory, yet few studies have evaluated the morphology of the mandibular corpus and symphysis in these taxa, and the investigations have yielded mixed results. Specifically, studies using external metrics have identified differences in mandibular proportions that analysis of cortical bone distribution has not substantiated. I contribute to the ongoing debate on the relationship between jaw form and dietary behavior by comparing mandibular corporal and symphyseal shapes in African apes. Importantly, and in contrast to previous studies of African ape internal geometry, I include the Virunga mountain gorillas (Gorilla beringei beringei), the ape most specialized toward a folivorous diet. I test the hypotheses that 1) Gorilla beringei beringei always has significantly more robust mandibular corpora and symphyses, relative to mandibular length, than all other African apes and 2) all gorillas have significantly more robust mandibular corpora and symphyses, relative to mandibular length, than Pan. Results demonstrate that the folivorous mountain gorillas consistently exhibit a relatively wider mandibular symphysis and corpus than all other African apes. Furthermore, all gorillas consistently exhibit relatively more robust mandibular corporal and symphyseal dimensions than Pan. The results indicate that among African apes, mountain gorillas are better able to counter lateral transverse bending (wishboning) loads at the symphysis and torsional loads at the corpus. All gorillas are likewise better able to resist wishboning and vertical bending at the symphysis, and sagittal bending and torsion at the corpus, than Pan, findings that are consistent with masticating relatively tougher foods, repetitive loading of the jaws, or both. I offer possible explanations for the lack of concordance in results between studies that have analyzed the biomechanical properties of African ape mandibles and others that have relied on external metrics. More comprehensive study of the internal geometry of the mandible is needed to resolve whether African apes differ morphologically in ways predicted by dietary variation.

Tooth cusp sharpness as a dietary correlate in great apes

Mammalian molars have undergone heavy scrutiny to determine correlates between morphology and diet. Here, the relationship between one aspect of occlusal morphology, tooth cusp radius of curvature (RoC), and two broad dietary categories, folivory and frugivory, is analyzed in apes. The author hypothesizes that there is a relationship between tooth cusp RoC and diet, and that folivores have sharper teeth than frugivores, and further test the correlation between tooth cusp RoC and tooth cusp size. Eight measures of tooth cusp RoC (two RoCs per cusp) were taken from 53 M²s from four species and subspecies of frugivorous apes (Pongo pygmaeus, Pan troglodytes troglodytes, Pan troglodytes schweinfurthii, and Gorilla gorilla gorilla) and two subspecies of folivorous apes (Gorilla beringei beringei, and Gorilla beringei graueri). Phylogenetically corrected ANOVAs were run on the full dataset and several subsets of the full dataset, revealing that, when buccolingual RoCs are taken into account, tooth cusp RoCs can successfully differentiate folivores and frugivores. PCAs revealed that folivores consistently had duller teeth than frugivores. In addition, a weak, statistically significant positive correlation exists between tooth cusp size and tooth cusp RoC. The author hypothesizes differences in tooth cusp RoC are correlated with wear rates, where, per vertical unit of wear, duller cusps will have a longer length of exposed enamel ridge than sharper cusps. More data need to be gathered to determine if the correlation between tooth cusp RoC and tooth cusp size holds true when small primates are considered. Am J Phys Anthropol 153:226–235, 2014. © 2013 Wiley Periodicals, Inc.     

Patterns of mandibular variation in Pan and Gorilla and implications for African ape taxonomy

Pan and Gorilla taxonomy is currently in a state of flux, with the number of existing species and subspecies of common chimpanzee and gorilla having been recently challenged. While Pan and Gorilla systematics have been evaluated on the basis of craniometric and odontometric data, only a handful of studies have evaluated multivariate craniometric variation within P. troglodytes, and none have evaluated in detail mandibular variation in either P. troglodytes or Gorilla gorilla. In this paper, we examine ontogenetic and adult mandibular variation in Pan and Gorilla. We test the hypothesis that patterns and degrees of mandibular variation in Pan and Gorilla closely correspond to those derived from previous analyses of craniometric variation. We then use these data to address some current issues surrounding Pan and Gorilla taxonomy. Specifically, we evaluate the purported distinctiveness of P.t. verus from the other two subspecies of Pan troglodytes, and the recent proposals to recognize Nigerian gorillas as a distinct subspecies, Gorilla gorilla diehli, and to acknowledge mountain and lowland gorillas as two separate species. Overall, patterns and degrees of multivariate mandibular differentiation parallel those obtained previously for the cranium and dentition. Thus, differences among the three conventionally recognized gorilla subspecies are somewhat greater than among subspecies of common chimpanzees, but differences between P. paniscus and P. troglodytes are greater than those observed between any gorilla subspecies. In this regard, the mandible does not appear to be more variable, or of less taxonomic value, than the face and other parts of the cranium. There are, however, some finer differences in the pattern and degree of morphological differentiation in Pan and Gorilla, both with respect to cranial and dental morphology, and in terms of the application and manner of size adjustment. Mandibular differentiation supports the conventional separation of bonobos from chimpanzees regardless of size adjustment, but size correction alters the relative alignment of taxa. Following size correction, intergroup distances are greatest between P.t. verus and all other groups, but there is considerable overlap amongst chimpanzee subspecies. Amongst gorillas, the greatest separation is between eastern and western gorillas, but adjustment relative to palatal vs. basicranial length results in a greater accuracy of group classification for G.g. gorilla and G.g. graueri, and more equivalent intergroup distances amongst all gorilla groups. We find no multivariate differentiation of the Nigerian gorillas based on mandibular morphology, suggesting that the primary difference between Nigerian and other western lowland gorillas lies in the nuchal region. Though intergroup distances are greatest between P.t. verus and other chimpanzee subspecies, the degree of overlap amongst all three groups does not indicate a markedly greater degree of distinction in mandibular, as opposed to other morphologies. Finally, mandibular differentiation corroborates previous craniodental studies indicating the greatest distinction amongst gorillas is between eastern and western groups. Thus, patterns and degrees of mandibular variation are in agreement with other kinds of data that have been used to diagnose eastern and western gorillas as separate species.     

Mitochondrial DNA phylogeography of western lowland gorillas (Gorilla gorilla gorilla)

The geographical distribution of genetic variation within western lowland gorillas (Gorilla gorilla gorilla) was examined to clarify the population genetic structure and recent evolutionary history of this group. DNA was amplified from shed hair collected from sites across the range of the three traditionally recognized gorilla subspecies: western lowland (G. g. gorilla), eastern lowland (G. g. graueri) and mountain (G. g. beringei) gorillas. Nucleotide sequence variation was examined in the first hypervariable domain of the mitochondrial control region and was much higher in western lowland gorillas than in either of the other two subspecies. In addition to recapitulating the major evolutionary split between eastern and western lowland gorillas, phylogenetic analysis indicates a phylogeographical division within western lowland gorillas, one haplogroup comprising gorilla populations from eastern Nigeria through to southeast Cameroon and a second comprising all other western lowland gorillas. Within this second haplogroup, haplotypes appear to be partitioned geographically into three subgroups: (i) Equatorial Guinea, (ii) Central African Republic, and (iii) Gabon and adjacent Congo. There is also evidence of limited haplotype admixture in northeastern Gabon and southeast Cameroon. The phylogeographical patterns are broadly consistent with those predicted by current Pleistocene refuge hypotheses for the region and suggest that historical events have played an important role in shaping the population structure of this subspecies.     

Dental and phylogeographic patterns of variation in gorillas

Gorilla patterns of variation have great relevance for studies of human evolution. In this study, molar morphometrics were used to evaluate patterns of geographic variation in gorillas. Dental specimens of 323 adult individuals, drawn from the current distribution of gorillas in equatorial Africa were divided into 14 populations. Discriminant analyses and Mahalanobis distances were used to study population structure.Results reveal that: 1) the West and East African gorillas form distinct clusters, 2) the Cross River gorillas are well separated from the rest of the western populations, 3) gorillas from the Virunga mountains and the Bwindi Forest can be differentiated from the lowland gorillas of Utu and Mwenga-Fizi, 4) the Tshiaberimu gorillas are distinct from other eastern gorillas, and the Kahuzi-Biega gorillas are affiliated with them. These findings provide support for a species distinction between Gorilla gorilla and Gorilla beringei, with subspecies G. g. diehli, G. g. gorilla, G. b. graueri, G. b. beringei, and possibly, G. b. rex-pygmaeorum. Clear correspondence between dental and other patterns of taxonomic diversity demonstrates that dental data reveal underlying genetic patterns of differentiation.Dental distances increased predictably with altitude but not with geographic distances, indicating that altitudinal segregation explains gorilla patterns of population divergence better than isolation-by-distance. The phylogeographic pattern of gorilla dental metric variation supports the idea that Plio-Pleistocene climatic fluctuations and local mountain building activity in Africa affected gorilla phylogeography. I propose that West Africa comprised the historic center of gorilla distribution and experienced drift-gene flow equilibrium, whereas Nigeria and East Africa were at the periphery, where climatic instability and altitudinal variation promoted drift and genetic differentiation. This understanding of gorilla population structure has implications for gorilla conservation, and for understanding the distribution of sympatric chimpanzees and Plio-Pleistocene hominins.     

An allometric study of the frontal sinus in Gorilla, Pan and Pongo

There is considerable speculation about the role and significance of the paranasal sinuses in the Hominoidea, and this study aims to present new data about an old problem from cephalograms of dried crania. Measurements of frontal sinus volumes were determined for Gorilla gorilla gorilla; G. gorilla beringei and Pan troglodytes. By adopting an allometric approach it was determined that the frontal sinus volume of Gorilla is relatively smaller than that of Pan, and that the frontal sinus of G. g. gorilla is relatively smaller than that of G. g. beringei. Frontal sinus volume scales in a positive allometric fashion relative to skull length. Since the slope is steeper for Pan, frontal sinus volume is increasing at a faster rate than in Gorilla. Sexual dimorphism in frontal sinus volume is present. Thirty crania of Pongo were investigated for evidence of pneumatization of the frontal bone. In no case was secondary invasion of the frontal bone by the maxillary antrum observed. In Gorilla, the nasal cavity volume scales isometrically with skull length. The scaling relationships discussed do not support any 'functional' role of the frontal sinus associated with nasal function but suggest a 'structural' role associated with craniofacial architecture.     

Degenerative joint disease in African great apes: an evolutionary perspective

Degenerative joint disease is investigated in the spine and major peripheral joints (shoulder, elbow, hip and knee) in samples of chimpanzees (Pan troglodytes schweinfurthii; P. troglodytes troglodytes), lowland gorillas (Gorilla gorilla gorilla), and bonobos (P. paniscus). The P. troglodytes schweinfurthii sample comes from Gombe National Park, Tanzania, while the other samples are derived from museum materials originally collected in west/central Africa. Total data for African ape samples include 5807 surfaces for ascertainment of vertebral osteophytosis, 12,479 surfaces for determination of spinal osteoarthritis, and 1211 joints for evaluation of peripheral joint osteoarthritis. All apes display significantly less spinal disease than in a comparable human sample, and these differences are most likely a consequence of human biomechanical adaptations for bipedal locomotion. Apes are also generally less involved in the major peripheral joints than are humans, but human groups are themselves highly variable in prevalence of peripheral osteoarthritis. These data agree with other findings of low prevalence of degenerative joint prevalence in free-ranging apes, but contrast markedly with evidence derived from colony-reared Old World monkeys.     

Anthrax in Western and Central African great apes

During the period of December 2004 to January 2005, Bacillus anthracis killed three wild chimpanzees (Pan troglodytes troglodytes) and one gorilla (Gorilla gorilla gorilla) in a tropical forest in Cameroon. While this is the second anthrax outbreak in wild chimpanzees, this is the first case of anthrax in gorillas ever reported. The number of great apes in Central Africa is dramatically declining and the populations are seriously threatened by diseases, mainly Ebola. Nevertheless, a considerable number of deaths cannot be attributed to Ebola virus and remained unexplained. Our results show that diseases other than Ebola may also threaten wild great apes, and indicate that the role of anthrax in great ape mortality may have been underestimated. These results suggest that risk identification, assessment, and management for the survival of the last great apes should be performed with an open mind, since various pathogens with distinct characteristics in epidemiology and pathogenicity may impact the populations. An animal mortality monitoring network covering the entire African tropical forest, with the dual aims of preventing both great ape extinction and human disease outbreaks, will create necessary baseline data for such risk assessments and management plans.     

Identifying metameric variation in extant hominoid and fossil hominid mandibular molars

Landmark data were collected from cross sections and occlusal images of mandibular molar crowns, and Euclidean distance matrix analysis (EDMA) was used to identify metameric morphological variation between the first and second mandibular molars of living taxa: Gorilla gorilla (n = 30), Pan troglodytes (n = 34), and Homo sapiens (n = 26). Two patterns of metameric variation were identified, one unique to humans and the other shared by chimpanzees and gorillas. In order to assess the utility of this type of analysis for the interpretation of the hominid fossil record, 19 mandibular molars from Sterkfontein Member 4, South Africa, were examined. The pattern of metameric variation of the Sterkfontein molars resembled that of the African great apes, and differed from the modern human pattern. These results demonstrate that data on metameric variation may provide information regarding function or developmental processes previously indiscernible from fossil material.     

Selective logging, habitat quality and home range use by sympatric gorillas and chimpanzees: a case study from an active logging concession in Southeast Cameroon

We examined range use by great apes during logging activities and investigated associations between local variations in ape abundance and changes in the structure of the habitat or in the availability of fruits after disturbances. We carried out two annual censuses of western lowland gorilla (G. g. gorilla) and chimpanzee populations (Pan t. troglodytes) in an active logging concession in Southeast Cameroon. The results suggest that gorillas may adapt their range use to avoid most recently logged compartments, while chimpanzees appear to be more spatially resilient to logging. In our study site, selective logging affected 10% of the forest. After logging, gorillas nested in all types of vegetation, while chimpanzees nested exclusively in mixed mature forest. Fruit availability was not affected by logging and did not explain the distribution of ape nests in the study area.     

Species association of hepatitis B virus (HBV) in non-human apes; evidence for recombination between gorilla and chimpanzee variants

Hepatitis B virus (HBV) infections are widely distributed in humans, infecting approximately one third of the world's population. HBV variants have also been detected and genetically characterised from Old World apes; Gorilla gorilla (gorilla), Pan troglodytes (chimpanzee), Pongo pygmaeus (orang-utan), Nomascus nastusus and Hylobates pileatus (gibbons) and from the New World monkey, Lagothrix lagotricha (woolly monkey). To investigate species-specificity and potential for cross species transmission of HBV between sympatric species of apes (such as gorillas and chimpanzees in Central Africa) or between humans and chimpanzees or gorillas, variants of HBV infecting captive wild-born non-human primates were genetically characterised. 9 of 62 chimpanzees (11.3%) and two from 11 gorillas (18%) were HBV-infected (15% combined frequency), while other Old world monkey species were negative. Complete genome sequences were obtained from six of the infected chimpanzee and both gorillas; those from P. t .ellioti grouped with previously characterised variants from this subspecies. However, variants recovered from P. t. troglodytes HBV variants also grouped within this clade, indicative of transmission between sub-species, forming a paraphyletic clade. The two gorilla viruses were phylogenetically distinct from chimpanzee and human variants although one showed evidence for a recombination event with a P.t.e.-derived HBV variant in the partial X and core gene region. Both of these observations provide evidence for circulation of HBV between different species and sub-species of non-human primates, a conclusion that differs from the hypothesis if of strict host specificity of HBV genotypes.     

Comparison of cranial ontogenetic trajectories among great apes and humans

Molecular data suggest that humans are more closely related to chimpanzees than either is to the gorillas, yet one finds the closest similarity in craniofacial morphology to be among the great apes to the exclusion of humans. To clarify how and when these differences arise in ontogeny, we studied ontogenetic trajectories for Homo sapiens, Pan paniscus, Pan troglodytes, Gorilla gorilla and Pongo pygmaeus. A total of 96 traditional three-dimensional landmarks and semilandmarks on the face and cranial base were collected on 268 adult and sub-adult crania for a geometric morphometric analysis. The ontogenetic trajectories are compared by various techniques, including a new method, relative warps in size-shape space. We find that adult Homo sapiens specimens are clearly separated from the great apes in shape space and size-shape space. Around birth, Homo sapiens infants are already markedly different from the great apes, which overlap at this age but diverge among themselves postnatally. The results suggest that the small genetic differences between Homo and Pan affect early human ontogeny to induce the distinct adult human craniofacial morphology. Pure heterochrony does not sufficiently explain the human craniofacial morphology nor the differences among the African apes.     

Behavioral Ecology of Sympatric Chimpanzees and Gorillas in Bwindi Impenetrable National Park,Uganda: Diet

Via a field study of chimpanzees (Pan troglodytes schweinfurthii) and gorillas (Gorilla gorilla beringei) in Bwindi Impenetrable National Park, Uganda, we found that their diets are seasonally similar, but diverge during lean seasons. Bwindi chimpanzees fed heavily on fruits of Ficus sp., which were largely ignored by the gorillas. Bwindi gorilla diet was overall more folivorous than chimpanzee diet, but was markedly more frugivorous than that of gorillas in the nearby Virunga Volcanoes. During 4 mo of the year Bwindi gorilla diet included more food species than that of the chimpanzees. Three factors in particular—seasonal consumption of fibrous foods by gorillas, interspecific differences in preferred fruit species, and meat consumption by chimpanzees—contributed to dietary divergence between the two species. When feeding on fruits, gorillas ate Myrianthus holstii more frequently than chimpanzees did, while chimpanzees included more figs in their annual diet. Chimpanzee diet included meat of duikers and monkeys; gorilla frequently consumed decaying wood.     

Food material properties and mandibular load resistance abilities in large-bodied hominoids

Numerous comparative studies have sought to demonstrate a functional link between feeding behavior, diet, and mandibular form in primates. In lieu of data on the material properties of foods ingested and masticated, many investigators have relied on qualitative dietary classifications such as "folivore" or "frugivore." Here we provide the first analysis of the relationship between jaw form, dietary profiles, and food material properties in large-bodied hominoids. We employed ratios of area moments of inertia and condylar area to estimate moments imposed on the mandible in order to evaluate and compare the relative ability to counter mandibular loads among central Bornean orangutans (Pongo pygmaeus wurmbii), Virunga mountain gorillas (Gorilla beringei beringei), and east African chimpanzees (Pan troglodytes schweinfurthii). We used data on elastic modulus (E) of fruit, fracture toughness (R) of fruit, leaves, and non-fruit, non-leaf vegetation, and derived fragmentation indices ( radicalR/E and radicalER), as proxies for bite force. We generated bending and twisting moments (forcexmoment arm) for various mandibular loading behaviors using food material properties to estimate minimally required bite forces. Based on E and R of foods ingested and masticated, we hypothesized improved resistance to mandibular loads in Pongo p. wurmbii compared to the African apes, and in G. b. beringei compared to Pan t. schweinfurthii. Results reveal that our predictions are borne out only when bite forces are estimated from maximum R of non-fruit, non-leaf vegetation. For all other tissues and material properties results were contrary to our predictions. Importantly, as food material properties change, the moments imposed on the mandible change; this, in turn, alters the entire ratio of relative load resistance to moment. The net effect is that species appear over- or under-designed for the moments imposed on the mandible. Our hypothesis, therefore, is supported only if we accept that maximum R of these vegetative tissues represents the relevant mechanical property influencing the magnitude of neuromuscular activity, food fragmentation, and mandibular morphology. A general implication is that reliable estimates of average and maximum bite forces from food material properties require that the full range of tissues masticated be tested. Synthesizing data on ingestive and masticatory behaviors, the number of chewing cycles associated with a given food, and food mechanical properties, should inform the broader question of which foods and feeding behaviors are most influential on the mandibular loading environment.     

Skeletal evidence of trauma in African apes, with special reference to the gombe chimpanzees

Incidence of cranial and postcranial skeletal trauma was investigated in samples of chimpanzees (Pan troglodytes troglodytes, P. troglodytes schweinfurthii), lowland gorillas (Gorilla gorilla gorilla), and bonobos (P. paniscus). The larger (adult) samples of chimpanzees (N=127 crania, 92 postcrania) and gorillas (N=136 crania, 62 postcrania) are curated at the Powell-Cotton Museum, Birchington, U.K. The bonobo collection (N=71 crania, 15 postcrania) is housed the Musée Royal de l'Afrique Centrale in Tervuren, Belgium. In addition, data were collected on the small but extremely well-documented skeletal sample from Gombe National Park (N=14 crania, 13 postcrania — including adults and adolescents). Cranial injuries, including healed fractures and bite wounds, were fairly frequent in the museum collection of chimpanzees (5.5% of individuals), but were twice as frequent in gorillas (11.0%). In the Gombe sample an even higher incidence was observed (28.6% of individuals). Bonobos, however, showed the lowest incidence of cranial trauma found among any of the African ape samples (1.4% of individuals). Postcranial trauma, documented most clearly by healed fractures, was seen in 21.7% of the Powell-Cotton chimpanzees, 30.8% of Gombe chimpanzees, 17.7% of gorillas, and in 13.3% of bonobos. Most of these lesions were found in the upper appendage. Nevertheless, highly debilitating healed fractures of the femur were also noted, most frequently and severe in female gorillas. The pattern of injuries suggests serious risks of falling in all free-ranging African apes, but also (in chimpanzees and gorillas) considerable risk from interindividual aggression, especially for males.     

Electromyography of pronators and supinators in great apes.

We obtained electromyographic recordings from the supinator, biceps brachii, pronator quadratus, and pronator teres muscles of a chimpanzee and a gorilla and from the supinator, pronator quadratus, and biceps brachii muscles of an orangutan as they stood and walked quadrupedally on horizontal and inclined surfaces, engaged in suspensory behavior, reached overhead, and manipulated a variety of foods and artifacts. In Pan troglodytes and Pan gorilla, as in Homo sapiens, the supinator muscle is the prime supinator, with the biceps brachii muscle serving to augment speed or force of supination. Primary of the pronator quadratus muscle over the pronator teres muscle during pronation is less clear in the African apes than in humans. Possibly, pongid radial curvature or forelimb elongation or both factors are related to the somewhat different patterns of activity that we observed in the pronator muscles of Pan versus those reported for Homo sapiens. In Pongo pygmaeus, as in P. troglodytes and P. gorilla, the pronator quadratus muscle acts as a pronator and the supinator muscle acts to supinate the hand at the radioulnar joints. The biceps brachii muscle is active at low levels as the orangutan supinates its hand with the elbow flexed.     

Occlusal relief changes with molar wear in Pan troglodytes troglodytes and Gorilla gorilla gorilla

Most research on primate tooth form-function relationships has focused on unworn teeth. This study presents a morphological comparison of variably worn lower second molars (M(2)s) of lowland gorillas (Gorilla gorilla gorilla; n=47) and common chimpanzees (Pan troglodytes troglodytes; n=54) using dental topographic analysis. High-resolution replicas of occlusal surfaces were prepared and scanned in 3D by laser scanning. The resulting elevation data were used to create a geographic information system (GIS) for each tooth. Occlusal relief, defined as the ratio of 3D surface area to 2D planometric area of the occlusal table, was calculated and compared between wear stages, taxa, and sexes. The results failed to show a difference in occlusal relief between males and females of a given taxon, but did evince differences between wear stages and between taxa. A lack of significant interaction between wear stage and taxon factors suggests that differences in occlusal relief between chimpanzees and gorillas are maintained throughout the wear sequence. These results add to a growing body of information on how molar teeth change with wear, and how differences between primate species are maintained at comparable points throughout the wear sequence. Such studies provide new insights into form-function relationships, which will allow us to infer certain aspects of diet in fossils with worn teeth.     

Dispersed male networks in western gorillas

Although kin-selection theory has been widely used to explain the tendency of individuals to bias beneficial behaviors towards relatives living within the same social group, less attention has focused on kin-biased interactions between groups. For animal societies in which females emigrate, as is the case for mountain gorillas (Gorilla beringei beringei), encounters between males in different groups often involve aggressive displays that can escalate to physical violence and fatal injuries. However, recent findings on the little-studied western gorilla (Gorilla gorilla) indicate that interactions between social groups occur more frequently than they do in mountain gorillas and are often, although not always, surprisingly nonaggressive. We investigated the pattern of genetic relationships between individuals of different groups and found evidence suggesting a previously unrecognized "dispersed male network" social structure in western gorillas in which the single males leading social groups were usually related to one or more nearby males. We propose that this provides a basis for extra-group, kin-biased behaviors and may explain the reported peaceful intergroup interactions. Furthermore, these results suggest that a patrilocal social structure, in which males remain in their natal region and potentially benefit from kin associations, is a feature unifying African apes and humans.