A comparative analysis of temporomandibular joint morphology in the African apes

A number of researchers have suggested a functional relationship between dietary variation and temporomandibular joint (TMJ) morphology, yet few studies have evaluated TMJ form in the African apes. In this study, I compare TMJ morphology in adults and during ontogeny in Gorilla (G.g. beringei, G.g. graueri, and G.g. gorilla) and Pan (P. paniscus, P. troglodytes troglodytes, P.t. schweinfurthii, and P.t. verus). I test two hypotheses: first, compared to all other African apes, G.g. beringei exhibits TMJ morphologies that would be predicted for a primate that consumes a diet comprised primarily of moderately to very tough, leafy vegetation; and second, all gorillas exhibit the same predicted morphologies compared to Pan. Compared to all adult African apes, G.g. beringei has higher rami and condyles positioned further above the occlusal plane of the mandible, relative to jaw length. Thus, mountain gorillas have the potential to generate relatively more muscle force, more evenly distribute occlusal forces along the postcanine teeth, and generate relatively greater jaw adductor moment. G.g. beringei also exhibits relatively wider mandibular condyles, suggesting these folivorous apes are able to resist relatively greater compressive loads along the lateral and/or medial aspect of the condyle. All gorillas likewise exhibit these same shape differences compared to Pan. These morphological responses are the predicted consequences of intensification of folivory and, as such, provide support for functional hypotheses linking these TMJ morphologies to degree of folivory. The African apes to not, however, demonstrate a systematic pattern of divergence in relative condylar area as a function of intensification of folivory. The ontogenetic trajectories for gorillas are significantly elevated above those of Pan, and to a lesser but still significant degree, mountain gorillas similarly deviate from lowland gorillas (G.g. gorilla and G.g. graueri). Thus, adult shape differences in ramal and condylar heights do not result from the simple extrapolation of common growth allometries relative to jaw length. As such, they are suggestive of an adaptive shift towards a tougher, more folivorous diet. However, the allometric patterning for condylar area and condylar width does not systematically conform to predictions based on dietary specialization. Thus, while differences in condylar shapes may confer functional advantages both during growth and as adults, there is no evidence to suggest selection for altered condylar proportions, independent of the effects of changes in jaw size.     

Growth changes in measurements of upper facial positioning

Growth changes in the position of the midline upper face are examined for samples of Pan troglodytes, Gorilla gorilla, and modern humans. Horizontal and vertical distances between nasion and the anterior end of the cribriform plate are plotted against stage of dental development. Kendall's nonparametric correlations between facial positioning and stage of dental development are tested for significance. In African apes, the upper face becomes more projecting and positioned higher relative to the anterior cranial base. The extent of this horizontal and vertical separation reflects primarily facial size. In modern humans, the upper face becomes more projecting but is relatively stable in its vertical position. Comparison of Pan and modern human crania in the youngest dental age category indicates that the upper face of modern humans is positioned lower early in postnatal life. The position of the upper face (glabella) relative to the anterior and posterior cranial base is presented for several fossil hominid crania. The fossil crania are similar to Pan and modern humans in facial projection relative to the anterior cranial base. However, glabella is positioned low in the fossil crania. Total facial projection (relative to hormion) for Sts 5 is similar to the mean for Gorilla. Fossil Homo and robust australopithecine crania display very projecting upper faces. We suggest that the upper face of Homo is projecting due to the length of the anterior cranial fossa, while robust australopithecines possess a thick frontal bone.     

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.     

Comparison of arthritis characteristics in lowland Gorilla gorilla and mountain Gorilla beringei

Gorilla gorilla and the less-studied G. beringei occupy very different, geographically separate habitats. We studied the occurrence of various forms of arthritis to examine possible nature/nurture causality. The macerated skeletons of 38 G. beringei and 99 G. gorilla individuals were examined macroscopically for the presence of articular and osseous pathologies. Contrasting with only isolated osteoarthritis and infectious arthritis was the frequent occurrence of a form of erosive arthritis associated with joint fusion. Twenty-one percent of the G. beringei and 20% of G. gorilla specimens were afflicted, which are statistically indistinguishable frequencies. While both had prominent axial disease, they differed in patterns of peripheral arthritis. Whereas G. beringei showed a pauciarticular pattern, the pattern in G. gorilla was more often polyarticular. Susceptibility to spondyloarthropathy was apparently genetically imprinted before Gorilla separated into G. gorilla and G. beringei. However, the different patterns of peripheral joint involvement suggest a causality resulting from lifestyle (e.g., the presence/absence or extent of knuckle walking) or a habitat-related infectious agent.     

Growth changes in internal and craniofacial flexion measurements.

Growth changes in both internal and craniofacial flexion angles are presented for Pan troglodytes, Gorilla gorilla, and modern humans. The internal flexion angle (IFA) was measured from lateral radiographs, and the craniofacial flexion angle (CFA) was calculated from coordinate data. Stage of dental development is used as a baseline for examination of growth changes and nonparametric correlations between flexion angles and dental development stage are tested for significance. In Gorilla, the IFA increases during growth. The IFA is relatively stable in Pan and modern humans. Pan and Gorilla display an increase in the CFA. However, this angle decreases during growth in modern humans. Flexion angles were derived from coordinate data collected for several early hominid crania. Measurements for two robust australopithecine crania indicate strong internal flexion. It has been suggested that cerebellar expansion in this group may relate to derived features of the posterior cranial base. In general, australopithecine crania exhibit craniofacial flexion intermediate between great apes and modern humans. The "archaic" Homo sapiens specimen from Kabwe is most similar to modern humans.     

Are the gorillas in Bwindi Impenetrable National Park “true” mountain gorillas?

The gorillas that inhabit Bwindi Impenetrable National Park in Uganda are the least known of the eastern gorillas. Because they are an allopatric population living a minimum of 25 km from the well‐studied population of mountain gorillas (Gorilla beringei beringei) in Rwanda and have certain morphological and ecological differences from these gorillas, their taxonomic status has been in question in recent years. This study presents new craniodental metrics from Bwindi individuals and compares them to Virunga individuals as well as to eastern lowland gorillas, G. gorilla graueri. Multivariate statistics, including MANCOVA, least‐squares, regression, and principal components analyses, were used to evaluate how closely the Bwindi crania resemble the Virunga crania and how both relate to G. g. graueri. Results indicate that the Bwindi gorillas have generally smaller crania than the Virunga gorillas, but when metrics are log‐transformed, the only variable that distinguishes the Bwindi individuals is a longer face. When both populations are compared to G. g. graueri, they cluster together separately from the eastern lowland gorillas, sharing such features as higher rami, wider bigonia, longer mandibles, and wider and shorter mandibular symphyses in relation to G. g. graueri. Functional morphological explanations for these differences are discussed, but lacking measurements of the physical properties of G. g. graueri, they cannot fully be explained. Results clearly indicate that at least pertaining to the cranium, upon which most gorilla taxonomy is based, the Bwindi gorillas are proper mountain gorillas (G. b. beringei). Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Brain organization of gorillas reflects species differences in ecology

Gorillas include separate eastern (Gorilla beringei) and western (Gorilla gorilla) African species that diverged from each other approximately 2 million years ago. Although anatomical, genetic, behavioral, and socioecological differences have been noted among gorilla populations, little is known about variation in their brain structure. This study examines neuroanatomical variation between gorilla species using structural neuroimaging. Postmortem magnetic resonance images were obtained of brains from 18 captive western lowland gorillas (Gorilla gorilla gorilla), 15 wild mountain gorillas (Gorilla beringei beringei), and 3 Grauer's gorillas (Gorilla beringei graueri) (both wild and captive). Stereologic methods were used to measure volumes of brain structures, including left and right frontal lobe gray and white matter, temporal lobe gray and white matter, parietal and occipital lobes gray and white matter, insular gray matter, hippocampus, striatum, thalamus, each hemisphere and the vermis of the cerebellum, and the external and extreme capsules together with the claustrum. Among the species differences, the volumes of the hippocampus and cerebellum were significantly larger in G. gorilla than G. beringei. These anatomical differences may relate to divergent ecological adaptations of the two species. Specifically, G. gorilla engages in more arboreal locomotion and thus may rely more on cerebellar circuits. In addition, they tend to eat more fruit and have larger home ranges and consequently might depend more on spatial mapping functions of the hippocampus. Am J Phys Anthropol 156:252–262, 2015. © 2014 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.     

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'.     

Phylogeny, neoteny and growth of the cranial base in hominoids

This study tests the hypothesis that there is a general pattern in the growth of the cranial base of Homo sapiens that is 'essentially neotenous' [Gould, 1977]. Juvenile and adult crania of Homo sapiens, Gorilla gorilla, Pan troglodytes and Pongo pygmaeus were studied and the cross-sectional growth curves for 10 measurements made on the cranial base (as viewed in norma basilaris) were compared. The results of this study suggest that relatively simple modifications to the timing or pattern of growth are insufficient to explain the observed morphological differences between the cranial base of modern Homo sapiens and the great apes.     

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.     

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.     

Effects of habitat fragmentation, population size and demographic history on genetic diversity: the Cross River gorilla in a comparative context

In small and fragmented populations, genetic diversity may be reduced owing to increased levels of drift and inbreeding. This reduced diversity is often associated with decreased fitness and a higher threat of extinction. However, it is difficult to determine when a population has low diversity except in a comparative context. We assessed genetic variability in the critically endangered Cross River gorilla (Gorilla gorilla diehli), a small and fragmented population, using 11 autosomal microsatellite loci. We show that levels of diversity in the Cross River population are not evenly distributed across the three genetically identified subpopulations, and that one centrally located subpopulation has higher levels of variability than the others. All measures of genetic variability in the Cross River population were comparable to those of the similarly small mountain gorilla (G. beringei beringei) populations (Bwindi and Virunga). However, for some measures both the Cross River and mountain gorilla populations show lower levels of diversity than a sample from a large, continuous western gorilla population (Mondika, G. gorilla gorilla). Finally, we tested for the genetic signature of a bottleneck in each of the four populations. Only Cross River showed strong evidence of a reduction in population size, suggesting that the reduction in size of this population was more recent or abrupt than in the two mountain gorilla populations. These results emphasize the need for maintaining connectivity in fragmented populations and highlight the importance of allowing small populations to expand.     

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.     

A complete sequence of the mitochondrial genome of the western lowland gorilla

The complete mitochondrial DNA (mtDNA) molecule of the gorilla was sequenced. The entire sequence, 16,412 nucleotides, was determined by analysis of natural (not polymerase chain reaction) restriction fragments covering the whole molecule. The sequence was established from one individual and thus nonchimeric. After comparison with the COII gene of gorilla specimens with known geographical origin, the sequence was identified as characteristic of the Western lowland gorilla, Gorilla gorilla gorilla. With the exception of the NADH2 gene, all genes have a methionine start codon. The inferred start codon of NADH2 is ATT (isoleucine). The COIII, NASDH4, and cytochrome b genes are not terminated by a stop codon triplet, and the COI gene is probably terminated by an AAA triplet rather than by a regular stop codon. The great majority of genic sequences (rRNAs, peptide-coding genes, tRNAs) of the complete mtDNAs of Gorilla, Pan, and Homo show a greater similarity between Pan and Homo than between either of these genera to Gorilla. The analysis of the peptide-coding genes suggest that relative to comparison between Homo and Pan a certain degree of transition saturation has taken place in codon position 3 in comparisons between Gorilla to either Homo or Pan.      

腊玛古猿和西瓦古猿的形态特征及其系统关系——颅骨的形态与比较

本文通过具体的形态比较指出,禄丰腊玛古猿和西瓦古猿应为同一类型的雌雄个体,它们与现代猩猩比较相似,而与大猩猩和黑猩猩差别较大,因此,它们可看作是猩猩的祖先。猩猩这一支大约是在一千二百万年前开始从人猿超科的进化主干上分化出来的。     

Repeated loss of frontal sinuses in arctoid carnivorans

Many mammal skulls contain air spaces inside the bones surrounding the nasal chamber including the frontal, maxilla, ethmoid, and sphenoid, all of which are called paranasal sinuses. Within the Carnivora, frontal sinuses are usually present, but vary widely in size and shape. The causes of this variation are unclear, although there are some functional associations, such as a correlation between expanded frontal sinuses and a durophagous diet in some species (e.g., hyenas) or between absent sinuses and semiaquatic lifestyle (e.g., pinnipeds). To better understand disparity in frontal sinus morphology within Carnivora, we quantified frontal sinus size in relationship to skull size and shape in 23 species within Arctoidea, a clade that is ecologically diverse including three independent invasions of aquatic habitats, by bears, otters, and pinnipeds, respectively. Our sampled species range in behavior from terrestrial (rarely or never forage in water), to semiterrestrial (forage in water and on land), to semiaquatic (forage only in water). Results show that sinuses are either lost or reduced in both semiterrestrial and semiaquatic species, and that sinus size is related to skull size and shape. Among terrestrial species, frontal sinus size was positively allometric overall, but several terrestrial species completely lacked sinuses, including two fossorial badgers, the kinkajou (a nocturnal, arboreal frugivore), and several species with small body size, indicating that factors other than aquatic habits, such as space limitations due to constraints on skull size and shape, can limit sinus size and presence. J. Morphol. 276:22–32, 2015. © 2014 Wiley Periodicals, Inc.     

What's Your Angle? Size Correction and Bar–Glenoid Orientation in “Lucy” (A.L. 288-1)

There has been much debate as to the locomotor repertoire of Lucy (A.L. 288-1) and other specimens of Australopithecus afarensis, ranging from fully committed bipeds to species that spent a significant time in the trees as well as on the ground. We examined the bar–glenoid angle, a character purported to indicate arboreal propensities, and its implications for this specific debate and the more general challenge of extracting behavioral information from fossils. We examined the bar–glenoid angle in ontogenetic samples of Pan paniscus, Pan troglodytes, Gorilla gorilla gorilla, Gorilla gorilla beringei, Pongo pygmaeus, Homo sapiens, and A.L. 288-1 (Lucy). We found that there is no allometry in the bar–glenoid angle for the great apes, but a weak correlation for humans. Moreover, the data scatters for the African apes and humans converge at the smaller size ranges, and Lucy's value for bar–glenoid angle falls precisely in this area of overlap. Therefore, we conclude that the bar–glenoid angle is not tightly correlated with function and, as such, cannot be used as a morphological signal of arboreal behavior, especially in the smaller size ranges, at which arboreal and nonarboreal species overlap. Our work does not resolve issues concerning Lucy's precise locomotor repertoire but adds new information to consider. The total morphological pattern, plus an appreciation of the underlying variance in morphological and behavioral characters in extant species, is key for making functional inferences from the morphology of fossils.     

Metrical analysis of the basicranium of extant hominoids and Australopithecus

The size and shape of the basicranium (seen in norma basilaris) in Homo, Gorilla, Pan, Pongo, and Australopithecus have been studied by recording the relative disposition of midline and bilateral bony landmarks. Fifteen linear measurements and two angles were used to relate the landmarks. The relatively longer and narrower cranial base of Gorilla, Pan, and Pongo is clearly contrasted with the wider, shorter cranial base in Homo sapiens. When the same observations were made on two “robust” and two “gracile” australopithecine crania, marked differences were found between the taxa. In the two “robust” specimens, the foramen magnum is located relatively further forward, and the axis of the petrous temporal bone is aligned more nearly with the coronal plane than in the two “gracile” crania. The implications of this apparent parallelism in basicranial morphology between Homo sapiens and the “robust” australopithecines are discussed.     

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.