Further evidence on relative dental maturation and somatic developmental rate in hominoids.

New data on hominoid dental development are presented. Individual bivariate pairings of all mandibular teeth were made for African apes and humans. Data were analyzed with a full linear regression model. No statistically significant differences were found among apes, although a consistent pattern of earlier incisal development was observed in Pan relative to Gorilla. This is concordant with an earlier fusion of the premaxillary:maxillary suture in Pan. Only one tooth pair differed significantly by sex among apes. Two biologically distinct human samples (Libben and Hamann-Todd), although assessed differently (extraction and radiography) yielded virtually identical results. Humans differ from apes only by earlier relative calcification of their anterior teeth. This can be viewed as a consequence of reduced facial prognathism and a shift in hominid canine function.     

Ontogeny of body size variation in African apes

Size variation in African apes (Gorilla gorilla [gorilla], Pan paniscus [pygmy chimpanzee], and Pan troglodytes [“common” chimpanzee]) is substantial, both within and between species. We investigate the possible evolutionary significance of this variation through an analysis of the ontogeny of size variation in this group. In addition, we highlight possible areas of future endocrinological research, and evaluate recently proposed alternative models that attempt to account for ontogenetic variation in apes. The present study shows that intergeneric variation in size is largely a consequence of differences among species in the rate of body weight growth. Interspecific size variation in Pan is a product of both rate and duration differences in growth. The ontogenetic bases of sexual dimorphism vary in this group. Dimorphism is largely a result of sex differences in the duration of body weight growth in gorillas and pygmy chimpanzees, but results from differences in the rate of growth in common chimpanzees. Ontogenetic analyses largely confirm earlier interpretations, but with better data and methods. The great degree of ontogenetic variation within and among these species, especially in the timing and magnitude of “pubertal” growth spurts, implies that studies of endocrine growth control in African apes could be a productive line of future research. We also suggest that ontogenetic variation can be understood with respect to ecological risks. Growth rates seem to be negatively correlated with ecological risk in African apes, suggesting links between ontogenetic patterns and social and ecological variables. High growth rates in gorillas compared to Pan are most consistent with this model. Variation between chimpanzees and pygmy chimpanzees (especially females) also seem to fit predictions of this model. © 1996 Wiley-Liss, Inc.     

Ontogeny and the evolution of adult body size dimorphism in apes

This analysis investigates the ontogeny of body size dimorphism in apes. The processes that lead to adult body size dimorphism are illustrated and described. Potential covariation between ontogenetic processes and socioecological variables is evaluated. Mixed-longitudinal growth data from 395 captive individuals (representing Hylobates lar [gibbon], Hylobates syndactylus [siamang], Pongo pygmaeus [orangutan], Gorilla gorilla [gorilla], Pan paniscus [pygmy chimpanzee], and Pan troglodytes [“common” chimpanzee]) form the basis of this study. Results illustrate heterogeneity in the growth processes that produce ape dimorphism. Hylobatids show no sexual differentiation in body weight growth. Adult body size dimorphism in Pongo can be largely attributed to indeterminate male growth. Dimorphism in African apes is produced by two different ontogenetic processes. Both pygmy chimpanzees (Pan paniscus) and gorillas (Gorilla gorilla) become dimorphic primarily through bimaturism (sex differences in duration of growth). In contrast, sex differences in rate of growth account for the majority of dimorphism in common chimpanzees (Pan troglodytes). Diversity in the ontogenetic pathways that produce adult body size dimorphism may be related to multiple evolutionary causes of dimorphism. The lack of sex differences in hylobatid growth is consistent with a monogamous social organization. Adult dimorphism in Pongo can be attributed to sexual selection for indeterminate male growth. Interpretation of dimorphism in African apes is complicated because factors that influence female ontogeny have a substantial effect on the resultant adult dimorphism. Sexual selection for prolonged male growth in gorillas may also increase bimaturism relative to common chimpanzees. Variation in female growth is hypothesized to covary with foraging adaptations and with differences in female competition that result from these foraging adaptations. Variation in male growth probably corresponds to variation in level of sexual selection. © 1995 Wiley-Liss, Inc.     

Enamel hypoplasia in deciduous teeth of great apes: do differences in defect prevalence imply differential levels of physiological stress?

This paper presents new data on enamel hypoplasia in the deciduous canine teeth of great apes. The enamel defect under consideration is known as localized hypoplasia of primary canines (LHPC), and is characterized by an area of thin or missing enamel on the labial surface of deciduous canine teeth (Skinner [1986a] Am. J. Phys. Anthropol. 69:59-69). Goals of this study are: 1) to determine if significant differences in the frequency of LHPC occur among three genera of great apes, and 2) to evaluate variation in LHPC prevalence among great apes as evidence of differential physiological stress. Infant and juvenile apes with deciduous teeth were examined at the Cleveland Museum of Natural History (n = 100) and at the Smithsonian Institution, National Museum of Natural History (n = 36). Deciduous teeth were observed under oblique incandescent light, with the naked eye and with a 10x hand lens. Enamel hypoplasia was scored using Federation Dentaire International (FDI)-Defects of Dental Enamel (DDE) standards. Hypoplasias were recorded by drawing defect location and size on a dental chart, and by measuring defect size and location with Helios needlepoint dial calipers. The prevalence of LHPC is reported by genus and sex, using two approaches: 1) the frequency of affected individuals-those having one or more deciduous canine teeth scored positive for LHPC; and 2) the number of canine teeth scored positive for LHPC as a percentage of all canine teeth observed. Variation in defect size and location will be described elsewhere. Localized hypoplasia of primary canine teeth was found in 62.5% of 128 individual apes, and in 45.5% of 398 great ape deciduous canines. As in humans, LHPC is the most common form of enamel hypoplasia in deciduous teeth of great apes, while LEH is rare or absent. The distribution and pattern of expression of LHPC in great apes is similar to that described in humans: side differences are not significant, but mandibular canines exhibit the defect two to five times more often than maxillary canine teeth. Differences in LHPC prevalence by sex are small and not significant. Intergeneric differences are large and non-random: chimpanzees (Pan) exhibit a significantly lower frequency of LHPC (22%, n = 50) by individual count, than either the orangutan (Pongo, 88.0%, n = 25) or the gorilla (Gorilla, 88.7%, n = 53). Tooth count prevalences exhibit a similar pattern of variation and are also statistically significant. These findings suggest that large bodied great apes (gorilla and orangutan) may be under greater physiological stress during perinatal and early postnatal development than the chimpanzee. The size, position, and timing of LHPC lesions are currently under analysis and may yield more insight into the etiological origin of this enamel defect.     

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.     

Sexual dimorphisms in dental dimensions of higher primates

Dental dimensions and distributions of dental dimensions of males and females were compared for great apes (Pan, Gorilla, and Pongo, and humans (Homo). The results were examined and discussed with reference to fossil primates Sivapithecus and Ramapithecus. The analyses focused on patterns of sexual dimorphism, both with regard to mean dimensions and the distribution of those dimensions. Sex differences in mean canine dimensions were large and significant for Gorilla and Pongo, significant but smaller for Pan, and small but occasionally significant for Homo. The dispersions of measures were greater for males than for females in Gorilla and Pan but did not differ significantly for Pongo or Homo. Examination of the noncanine teeth revealed complex sex differences. In the anterior teeth, sex differences in mean dimensions were generally apparent for Gorilla and Pongo, less so for Pan, and least of all in Homo. The patterns of dispersion of measures of anterior teeth differed markedly from those of the canines. Pan exhibited the same pattern for anterior and canine teeth. Gorilla showed the opposite pattern. Pongo and Homo showed similar dispersions for males and females in many cases. Sex differences in posterior teeth followed the pattern of the canines for Gorilla and were absent for Pan. Pongo exhibited mean differences in dimensions across sex, but dispersions were similar. The pattern for Homo was most like that of Pongo, but with fewer significant differences. The genera differed with regard to the number of significant differences in means or dispersions along the tooth row. It is clear that the patterns of dimorphism differ qualitatively across all extant genera of great apes and humans. It appears that the pattern for Homo most closely resembles that of Ramapithecus, whereas Pongo most closely resembles Sivapithecus. The patterns for Gorilla and Pan appear to be unlike either of the fossil forms. It is suggested that the qualitatively distinct patterns of dental sexual dimorphism indicate substantial flexibility during recent primate evolution and that the degree of structural flexibility demonstrated provides a basis for appreciating potential for plasticity of gender differences in behavioral, social, and cultural systems.     

Molecular evolution of IgG subclass among nonhuman primates: implication of differences in antigenic determinants among Apes

The cross-reactivity of five different rabbit polyclonal antibodies to human IgG and IgG subclass (IgG1, IgG2, IgG3, and IgG4) was determined by competitive ELISA with nine nonhuman primate species including five apes, three Old World monkeys, and one New World monkey. As similar to those previously reported, the reactivity of anti-human IgG antibody with plasma from different primate species was closely related with phylogenic distance from human. Every anti-human IgG subclass antibody showed low cross-reactivity with plasma from Old World and New World monkeys. The plasma from all apes except for gibbons (Hylobates spp.) showed 60 to 100% of cross-reactivity with anti-human IgG2 and IgG3 antibodies. On the other hand, chimpanzee (Pan troglodytes andPan paniscus) and orangutan (Pongo pygmaeus) plasma showed 100% cross-reactivity with anti-human IgG1 antibody, but gorilla (Gorilla gorilla) and gibbon plasma showed no cross-reactivity. The chimpanzee and gorilla plasma cross-reacted with anti-human IgG4 antibody at different reactivity, 100% in chimpanzee and 50% in gorilla, but no cross-reactivity was observed in orangutan and gibbon plasma. These results suggest the possibilities that the divergence of “human-type” IgG subclasses might occur at the time of divergence ofHomo sapience fromHylobatidae, and that the molecular evolution of IgG1 as well as IgG4 is different from that of IgG2 and IgG3 in great apes, this is probably caused by different in development of immune function in apes during the course of evolution.     

Sexual dimorphism in canine shape among extant great apes

There have been numerous attempts to sex fossil specimens using the canine dentition. Whether focused on canine size or canine shape, most of these efforts share two deficiencies: lack of quantification of male-female differences in the adopted criteria and a failure to adequately explore among extant species the discriminatory power of these criteria. Here, canine shape indices relating to relative canine height, upper canine root/crown proportionality, and relative length of the lower canine mesial ridge were calculated for males and females of all species and subspecies of extant great apes and two species of gibbons. The accuracy of these indices for identifying the sex of the extant ape specimens was investigated through discriminant analysis and the use of bivariate plots of the two upper and two lower canine indices. The indices were found to be highly accurate in identifying the sex of great ape individuals, not only in single-species and subspecies samples but in mixed-species samples as well; assignment error rates were mostly between 0 and 4%. Accuracy was lowest in Pan (error rates as high as 15%) and highest in Pongo (one error). In most cases, error rates were lower in the upper canines. The effectiveness of these shape indices for sexing might be related to the degree of absolute canine size dimorphism; the indices did not effectively segregate males and females among minimally canine-dimorphic gibbons. The mixed-species results reveal that same-sex index values are remarkably concordant across great ape species, as are the patterns of spatial segregation of males and females in the bivariate plots. Results suggest that, while the indices can be used with some confidence to sex individual fossil specimens, their greatest utility will be for identifying the sex of groups of canines united by size and morphology. © 1995 Wiley-Liss, Inc.     

Hominoid seminal protein evolution and ancestral mating behavior

Hominoid mating systems show extensive variation among species. The degree of sexual dimorphism in body size and canine size varies among primates in accordance with their mating system, as does the testes size and the consistency of ejaculated semen, in response to differing levels of sperm competition. To investigate patterns of evolution at hominoid seminal proteins and to make inferences regarding the mating systems of extinct taxa, we sequenced the entire coding region of the prostate-specific transglutaminase (TGM4) gene in human, chimpanzee, bonobo, western lowland gorilla, eastern lowland gorilla, orangutan, and siamang, including multiple humans, chimps, and gorillas. Partial DNA sequence of the coding regions was also obtained for one eastern lowland gorilla at the semenogelin genes (SEMG1 and SEMG2), which code for the predominant proteins in semen. Patterns of nucleotide variation and inferred protein sequence change were evaluated within and between species. Combining the present data with previous studies demonstrates a high rate of amino acid substitutions, and low intraspecific variation, at seminal proteins in Pan, presumably driven by strong sperm competition. Both gorilla species apparently possess nonfunctional TGM4, SEMG1, and SEMG2 genes, suggesting that gorillas have had low sperm competition, and therefore their current polygynous mating system, for a long time before their divergence. Similarly, orangutans show longstanding stasis at TGM4, which may be interpreted as evidence for an unchanging mating system for most of their evolution after their divergence from African apes. In contrast to the great apes, the data from humans could be interpreted as evidence of fluctuations between different mating systems or alternatively as a relaxed functional constraint in these proteins. It is our hope that this study is a first step toward developing a model to predict ancestral mating systems from extant molecular data to complement interpretations from the fossil record.     

The AluI-induced bands in great apes and man: implication for heterochromatin characterization and satellite DNA distribution

Restriction endonucleases have recently been proved to be active on fixed chromatin, producing differences in staining of metaphase chromosomes. In this paper we show the results obtained by treating the metaphase chromosomes of Pan troglodytes, Pan paniscus, and Gorilla gorilla with the restriction enzyme AluI. These results demonstrate qualitative differences in the telomeric heterochromatin between Pan and Gorilla despite the fact that these areas appear homogeneous in the two genera by the C-banding method. The results found with individual chromosomes in the different species also appear relevant, in the light of the evolutionary relationships between these nonhuman primates and man. Lastly, the results suggest the presence, in great apes, of some highly repetitive DNA sequences different from the human satellites I-IV.     

Geographic Variation in Nonmetric Dental Traits of the Deciduous Molars of Pan and Gorilla

Physical anthropologists often use nonmetric dental traits to trace the movement of human populations, but similar analysis of the teeth of nonhuman primates or the deciduous teeth is rare. Because nonmetric dental characteristics are manifestations of genetic differences among groups, they vary among geographically distant members of the same species and subspecies. We use 28 nonmetric dental traits in the deciduous molars to compare genetically and geographically distinct groups of extant African apes (Gorilla and Pan). Previous researchers have studied these traits in the adult or juvenile teeth of great apes and humans, and we score our observations according to established standards for hominins. We observe marked differences in trait frequencies between Gorilla and Pan, Pan troglodytes and P. paniscus, and two P. troglodytes subspecies but we find no significant differences between geographically isolated groups within the subspecies. Trait frequencies differ from those found in previous studies that contained fewer individuals. We find that the deciduous molars show similar variation to adult premolars and molars within Pan and Gorilla. This suggests that the deciduous dentition of these and other apes may contain diagnostic traits that are not currently in use.     

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.     

New approaches in hominoid taxonomy: morphometrics

We report here on new cranial data relevant to hominoid taxonomic analyses, based on a study of 438 skulls belonging to 13 nonhuman living hominoid taxa. Nineteen landmarks were selected to describe the overall shape of the maxillofacial complex, in order to investigate its discriminative power in taxonomic analyses. We used a geometric morphometrics approach to depict morphological variation from the genus down to the subspecific level, and we evaluated whether our morphologic criteria are relevant to discriminating species and subspecies among living hominoids. Considering previous genetic studies, we discuss whether our results can be extrapolated to the hominin fossil record, providing a reference for species and subspecies morphologic differentiation. Our results indicate that the relative warp method, as applied to facial landmarks, provides a powerful tool to discriminate taxa down to a subspecific level. Results show a noticeable divergence of P. t. verus compared to P. t. troglodytes and P. t. schweinfurthii. According to our data, the distance between eastern and western gorilla populations as well as between Bornean and Sumatran orangutan subspecies is as great as between the two species of Pan. In the same manner, differences between Hylobates and Symphalangus are similar to those between Pan and Gorilla genera. Congruence between the morphological distances computed in this study and previous morphological and genetical studies strongly supports their relevance for morphological species recognition in paleoanthropology. Our data provide an objective standard for assessing taxonomic differences among hominoids, and will enable us to define more precisely the significance of morphological differences in the fossil record.     

Patterns of tooth crown size and shape variation in great apes and humans and species recognition in the hominid fossil record

It has been suggested that patterns of craniodental variation in living hominids (Gorilla, Homo, Pan, and Pongo) may be useful for evaluating variation in fossil hominid assemblages. Using this approach, a fossil sample exhibiting a pattern of variation that deviates from one shared among living taxa would be regarded as taxonomically heterogeneous. Here we examine patterns of tooth crown size and shape variation in great apes and humans to determine 1) if these taxa share a pattern of dental variation, and 2) if such a pattern can reliably discriminate between samples that contain single species and those that contain multiple species. We use parametric and nonparametric correlation methods to establish the degree of pattern similarity among taxa, and randomization tests to assess their statistical significance. The results of this study show that extant hominids do not share a pattern of dental size variation, and thus these taxa cannot be used to generate expectations for patterns of size variation in fossil hominid species. The hominines (Gorilla, Homo, and Pan) do share a pattern of shape variation in the mandibular dentition; however, Pongo is distinct, and thus it is unclear which, if either, pattern should be expected in fossil hominids. Moreover, in this case, most combined-species samples exhibit patterns of shape variation that are similar to those for single hominine species samples. Thus, although a common pattern of shape variation is present in the mandibular dentition, it is not useful for recognizing taxonomically mixed paleontological samples.     

Meningeal arterial patterns in great apes: Implications for hominid vascular evolution

Arterial meningeal patterns were observed for 100 hemispheres from great ape endocasts (Pan paniscus, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus). Eight patterns emerged based on the relative contributions to the walls and dura mater of the middle part of the braincase of meningeal arteries that stem from two sources. These arteries enter the braincase through either the orbit (delivering blood from the internal carotid artery) or through the base of the middle cranial fossa (via the middle meningeal artery whose blood comes from the external carotid artery). The three genera of apes manifest different frequencies of the eight, patterns, with orangutans highly dependent on orbital meningeal arteries at one extreme, and chimpanzees showing the greatest reliance on the middle meningeal artery at the other. As was the case in an earlier study of rhesus monkeys, there is a trend across the two genera of African apes for increased mean cranial capacity to be associated with increased reliance on the internal carotid artery for supplying the middle portion of the braincase. However, unlike the case for macaques, this trend does not reach statistical significance in African apes. Because it is rare for humans to manifest significant arterial contributions from the orbit to the middle cranial fossa, the comparative data on monkeys, apes, and humans suggest that, during the course of vascular evolution in Homo, the middle meningeal artery eventually took over supply of the entire middle cranial fossa. This hypothesis should be tested in the hominid fossil record. Earlier work on meningeal arterial patterns in apes has traditionally relied on Adachi's system that was determined from humans and focuses on the origin of the middle branch of the middle meningeal artery. As a result, the extensive orbital contributions to the middle portion of the braincase that characterize apes were not recognized and the eight patterns described in this paper were often erroneously assigned to the three patterns that adequately describe only humans. Adachi's system should therefore be abandoned for nonhuman primates and early hominids. A correct understanding of meningeal arterial evolution cannot be achieved until the orbital contributions to the meningeal arteries are recognized and incorporated into an evolutionary study that spans from apes to fossil hominids to living people. © 1993 Wiley-Liss, Inc.     

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.     

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.     

Longitudinal study of dental development in chimpanzees of known chronological age: Implications for understanding the age at death of Plio-Pleistocene hominids

Reconstruction of life history variables of fossil hominids on the basis of dental development requires understanding of and comparison with the pattern and timing of dental development among both living humans and pongids. Whether dental development among living apes or humans provides a better model for comparison with that of Plio-Pleistocene hominids of the genus Australopithecus remains a contentious point. This paper presents new data on chimpanzees documenting developmental differences in the dentitions of modern humans and apes and discusses their significance in light of recent controversies over the human or pongid nature of australopithecine dental development. Longitudinal analysis of 299 lateral head radiographs from 33 lab-reared chimpanzees (Pan troglodytes) of known chronological age allows estimation of means and standard deviations for the age at first appearance of 8 developmental stages in the mandibular molar dentition. Results are compared with published studies of dental development among apes and with published standards for humans. Chimpanzees are distinctly different from humans in two important aspects of dental development. Relative to humans, chimpanzees show advanced molar development vis a vis anterior tooth development, and chimpanzees are characterized by temporal overlap in the calcification of adjacent molar crowns, while humans show moderate to long temporal gaps between the calcification of adjacent molar crowns. In combination with recent work on enamel incremental markers and CAT scans of developing dentitions of Plio-Pleistocene hominids, this evidence supports an interpretation of a rapid, essentially “apelike” ontogeny among australopithecines. © 1996 Wiley-Liss, Inc.     

Sex determination in Miocene catarrhine primates

Canines of fossil hominoids and primitive catarrhines from several early, middle, and late Miocene sites were analyzed according to the shape indices described in Kelley (1995) and compared to those of males and females of extant great apes. In bivariate plots of the fossil canines utilizing the indices, 90% of the upper canines and 85% of the lower canines fell within or just outside the exclusively male or exclusively female territories delimited by the extant great apes. The remainder fell in the male-female overlap zones. Sex assignments based on these distributions were nearly 100% concordant with classifications according to canine height, suggesting a high degree of accuracy. There were various taxon-specific shifts in bivariate space among fossil genera, reflecting subtle differences in canine shape between taxa within the overall pattern of similarity to extant great apes as a whole. In many cases these shifts are matched by particular extant-ape species and subspecies, while other fossil taxa have no exact analogue for canine shape among the extant great apes. However, the pattern of spatial segregation of canines identified as either male or female at each of the sites largely mirrors that of males and females within the extant-ape sample, indicating that Miocene catarrhines shared with extant great apes a common pattern of shape differences between male and female canines, regardless of taxonspecific morphologies. These observations demonstrate that the canines of fossil catarrhines can be sexed with a high degree of confidence based solely on intrinsic features of shape. This will permit more reliable characterizations of morphological sexual dimorphism among fossil species. It is also argued that canine shape is a more reliable indicator of sex in fossil taxa than are canine/molar size ratios. © 1995 Wiley-Liss, Inc.