Species Of Pan Research Articles

The endocast morphology of LES1, Homo naledi.

Homo naledi is near the extreme of small brain size within Homo but is easily recognized as Homo in other aspects of endocast morphology. This study adds new evidence of the endocast morphology of H. naledi by describing the Lesedi Hominin 1 (LES1) endocranium from the Lesedi Chamber and compares it to the previously known H. naledi individual Dinaledi Hominin 3 (DH3) as well as other hominin taxa. We examined interlandmark distances with both univariate and multivariate methods in multiple hominin taxa and both species of Pan. For each distance, we compared groups using adjusted Z-scores (Azs). Our multivariate analyses included both principal component analyses (PCA) and linear discriminant analyses (LDA). DH3 and LES1 each have absolute third frontal convolution measures that enter the upper half of the variation for Homo sapiens, Homo erectus, and Homo neanderthalensis. Examined relative to the cube root of endocranial volume, H. naledi ranks among the highest values in these samples of Homo. Both absolute and relative values for H. naledi specimens are far above Pan, Australopithecus, and Paranthropus, suggesting an expanded Broca's area. Both qualitative and quantitative analyses show consistency between LES1 and other H. naledi endocasts and confirm the shared morphology of H. naledi with H. sapiens, H. neanderthalensis, and some specimens of H. erectus.

The relativity of ontogenetic time on primate evolution and the emergence of language

Most reviews on primatology focus on the similarities between the species of Pan (chimpanzees and bonobos) and Homo (sapiens) genera. In this paper, however, we review the literature on heterochronic differences in ontogenetic development of species in order to raise a discussion about their behavioral differences, particularly in relation to the human communication system. A key concept discussed here is neoteny, which refers to the slowed rate of species development. Human biology exhibits high levels of neoteny, resulting in a prolonged period of development during the first decade of postnatal life. Human neoteny enables the brain to develop in conjunction with physical and social environments, emphasizing that biology and society should not be perceived as distinct perspectives of the same process, but rather as interdependent processes that collaborate for the normal development of human beings. In order to identify biological concepts that may have contributed to the emergence of language, throughout this paper we explore how neoteny can be a potential explanatory concept to some of the behavioral differences between species, and how this may relate to cognitive systems such as language acquisition in Homo sapiens.

Plasma Testosterone and Androstenedione Levels Follow the Same Sex-Specific Patterns in the Two Pan Species.

Simple SummaryGenerally male mammals are more aggressive than their female peers. In these males, aggressive behavior is linked to levels of androgens; higher levels of testosterone are predictive of higher aggression rates or more severe aggression. There are some species where the pattern of sex-specific aggression is reversed, and it was hypothesized that high levels of androgens may be responsible for social dominance and aggressiveness in these females. Studies so far found that females of species with sex-reversed aggression patterns (e.g., spotted hyenas and ring-tailed lemurs) had lower plasma testosterone levels than their male peers, but a precursor of testosterone, androstenedione, was comparable or even higher in females than in males. This supported the idea that selection for female aggressiveness may be facilitated through augmented androgen secretion. Here we show that in two sister species, bonobos and chimpanzees, that differ in terms of sex-specific aggression patterns, females have lower plasma testosterone levels and higher plasma androstenedione levels than their male peers. Thus, our data do not support a theory of a role of female androgen levels on the expression of sex-specific patterns of aggression.In most animals, males are considered more aggressive, in terms of frequency and intensity of aggressive behaviors, than their female peers. However, in several species this widespread male-biased aggression pattern is either extenuated, absent, or even sex-reversed. Studies investigating potential neuro-physiological mechanisms driving the selection for female aggression in these species have revealed an important, but not exclusive role of androgens in the expression of the observed sex-specific behavioral patterns. Two very closely related mammalian species that markedly differ in the expression and degree of sex-specific aggression are the two Pan species, where the chimpanzee societies are male-dominated while in bonobos sex-biased aggression patterns are alleviated. Using liquid chromatography–mass spectrometry (LC-MS) methods, we measured levels of plasma testosterone and androstenedione levels in male and female zoo-housed bonobos (N = 21; 12 females, 9 males) and chimpanzees (N = 41; 27 females, 14 males). Our results show comparable absolute and relative intersexual patterns of blood androgen levels in both species of Pan. Plasma testosterone levels were higher in males (bonobos: females: average 0.53 ± 0.30 ng/mL; males 6.70 ± 2.93 ng/mL; chimpanzees: females: average 0.40 ± 0.23 ng/mL; males 5.84 ± 3.63 ng/mL) and plasma androstenedione levels were higher in females of either species (bonobos: females: average 1.83 ± 0.87 ng/mL; males 1.13 ± 0.44 ng/mL; chimpanzees: females: average 1.84 ± 0.92 ng/mL; males 1.22 ± 0.55 ng/mL). The latter result speaks against a role of androstenedione in the mediation of heightened female aggression, as had been suggested based on studies in other mammal species where females are dominant and show high levels of female aggressiveness.

Morphological variation of the maxilla in modern humans and African apes

Differences in morphology among modern humans and African apes are frequently used when assessing whether hominin fossils should be attributed to a single species or represent evidence for taxic diversity. A good understanding of the degree and structure of the intergeneric, interspecific, and intraspecific variation, including aspects such as sexual dimorphism and age, are key in this context. Here we explore the variation and differences shown by the maxilla of extant hominines, as maxillary morphology is central in the diagnosis of several hominin taxa. Our sample includes adults of all currently recognized hominine species and subspecies, with a balanced species sex ratio. In addition, we compared the adults with a small sample of late juveniles. The morphology of the maxillae was captured using three-dimensional landmarks, and the size and shape were analyzed using geometric morphometric methods. Key observations are that 1) the maxillae of all extant hominine species and subspecies show statistically significant differences, but complete separation in shape is only seen at the genus level; 2) the degree of variation is not consistent between genera, with subspecies of Gorilla being more different from each other than are species of Pan; 3) the pattern of sexual shape dimorphism is different in Pan, Gorilla, and Homo, often showing opposite trends; and 4) differentiation between maxillary shapes is increased after adjustment for static intraspecific allometry. These results provide a taxonomically up-to-date comparative morphological framework to help interpret the hominin fossil record, and we discuss the practical implications in that context.

Geometric morphometric variability in the supraorbital and orbital region of Middle Pleistocene hominins: Implications for the taxonomy and evolution of later Homo

This study assessed variation in the supraorbital and orbital region of the Middle Pleistocene hominins (MPHs), sometimes called Homo heidelbergensis s.l., to test whether it matched the expectations of intraspecific variation. The morphological distinctiveness and relative variation of this region, which is relatively well represented in the hominin fossil record, was analyzed quantitatively in a comparative taxonomic framework. Coordinates of 230 3D landmarks (20) and sliding semilandmarks (210) were collected from 704 specimens from species of Homo, Australopithecus, Paranthropus, Gorilla, Pan, Papio, and Macaca. Results showed that the MPHs had expected levels of morphological distinctiveness and intragroup and intergroup variation in supraorbital and orbital morphology, relative to commonly recognized non-hominin catarrhine species. However, the Procrustes distances between this group and H. sapiens were significantly higher than expected for two closely related catarrhine species. Furthermore, this study showed that variation within the MPH could be similarly well contained within existing hypodigms of H. sapiens, H. neanderthalensis, and H. erectus s.l. Although quantitative assessment of supraorbital and orbital morphology did not allow differentiation between taxonomic hypotheses in later Homo, it could be used to test individual taxonomic affiliation and identify potentially anomalous individuals. This study confirmed a complicated pattern of supraorbital and orbital morphology in the MPH fossil record and raises further questions over our understanding of the speciation of H. sapiens and H. neanderthalensis and taxonomic diversity in later Homo.

Attractiveness of female sexual signaling predicts differences in female grouping patterns between bonobos and chimpanzees

Here we show that sexual signaling affects patterns of female spatial association differently in chimpanzees and bonobos, indicating its relevance in shaping the respective social systems. Generally, spatial association between females often mirrors patterns and strength of social relationships and cooperation within groups. While testing for proposed differences in female-female associations underlying female coalition formation in the species of the genus Pan, we find only limited evidence for a higher female-female gregariousness in bonobos. While bonobo females exhibited a slightly higher average number of females in their parties, there is neither a species difference in the time females spent alone, nor in the number of female party members in the absence of sexually attractive females. We find that the more frequent presence of maximally tumescent females in bonobos is associated with a significantly stronger increase in the number of female party members, independent of variation in a behavioural proxy for food abundance. This indicates the need to look beyond ecology when explaining species differences in female sociality as it refutes the idea that the higher gregariousness among bonobo females is driven by ecological factors alone and highlights that the temporal distribution of female sexual receptivity is an important factor to consider when studying mammalian sociality.

Frontal sinus variation in extant species of the genera Pan, Gorilla

Frontal pneumatisation is not present in all primates, and among extant species, ethmoidally-derived frontal sinuses are present only in the genera Pan, Gorilla and Homo. A simple and repeatable method is described here to quantify variation in the shape, size and bilateral variation of the frontal sinuses. This has allowed the first large study of these variables, including analyses of large samples of P. paniscus, P. troglodytes, G. gorilla and of several geographically diverse populations of H. sapiens. Frontal sinus shape and size are potential phylogenetic markers in primate systematics. We suggest that, in contrast to an allometric relationship between endocranial and frontal sinus form in Pan and Gorilla, H. sapiens is autapomorphic in having highly variable, supero-inferiorly large frontal sinuses, whose size is unrestricted by variation in cranial form. However, we also describe differences in frontal pneumatisation between those taxa. H. sapiens differs from the other taxa in having relatively smaller frontal sinuses that are more variable in shape and size. In P. troglodytes, P. paniscus and G. gorilla, the shape and size of frontal pneumatisation correlates positively with the overall size of the braincase. This suggests that the large space available in the bone structures of the non-human primates analysed allows the sinuses to develop under only slight constraint (if any) from surrounding structures. In H. sapiens, a significant correlation is observed between the asymmetry in the anterior extension of the frontal lobes of the brain and the shape and extension of the frontal sinuses. We suggest that the more vertical orientation of the H. sapiens frontal bone, its reduced thickness, the reduction of the supraorbital tori and the increased influence of the frontal lobes appear to have modified the integration between the skull, brain and sinuses in the frontal region in our species compared to our nearest extant relatives. Our results reduce the possibility of a strong functional origin for the variation in size and shape of the frontal sinuses in the primates analysed.

Sacrum morphology supports taxonomic heterogeneity of \u201cAustralopithecus africanus\u201d at Sterkfontein Member 4

The presence of multiple Australopithecus species at Sterkfontein Member 4, South Africa (2.07–2.61 Ma), is highly contentious, and quantitative assessments of craniodental and postcranial variability remain inconclusive. Using geometric morphometrics, we compared the sacrum of the small-bodied, presumed female subadult Australopithecus africanus skeleton Sts 14 to the large, alleged male adult StW 431 against a geographically diverse sample of modern humans, and two species of Pan, Gorilla, and Pongo. The probabilities of sampling morphologies as distinct as Sts 14 and StW 431 from a single species ranged from 1.3 to 2.5% for the human sample, and from 0.0 to 4.5% for the great apes, depending on the species and the analysis. Sexual dimorphism and developmental or geologic age could not adequately explain the differences between StW 431 and Sts 14, suggesting that they are unlikely to be conspecific. This supports earlier claims of taxonomic heterogeneity at Sterkfontein Member 4.

Epiphyseal fusion and dental development in Pan paniscus with comparisons with Pan troglodytes.

Compared with frequent studies of skeletal development in chimpanzees, relatively little is known about bonobo skeletal development. This study seeks to explore the relationship between skeletal and dental development in both species of Pan. New data are presented for fusion sites not previously observed in bonobos. In a sample of 34 Pan paniscus and 168 Pan troglodytes subadults, state of fusion was recorded for 30 epiphyseal fusion sites using a three-stage system of unfused, midfusion, and complete fusion based on Wintheiser, Clauser, and Tappen. Stage of dental development for permanent mandibular dentition was assessed using the Demrijian, Goldstein, and Tanner method. These data allowed for comparisons of both species of Pan and the two subspecies of P. troglodytes. The sequence of fusion events was generally consistent between the two species, but some exceptions may exist for the knee and ankle. The number of fusion events that occurred after complete dental mineralization was similar in both species. No statistically significant differences were found in the fusion timing for the subspecies of P. troglodytes. Bolter and Zihlman suggested that fusion at the acetabulum occurs earlier in Pan paniscus, while fusion of epiphyses at the knee are delayed, compared with P. troglodytes. Our data do not indicate earlier fusion of the acetabulum, but fusion events at the knee may complete later relative to dental mineralization in Pan pansicus. Compared with Homo sapiens, both P. troglodytes and Pan paniscus demonstrate later completion of epiphyseal fusion relative to dental mineralization.

Warfare, Ethics, Ethology

The aim of this article is to set a macro-historical narrative concerning the emergence of warfare and social ethics as symplesiomorphic features in the lineage of Homo sapiens. This means that these two behavioral aspects, representative of a very selected branch in the phylogenetic tree of the Primate order, are shared by the two lineages of great African apes that diverged from a common ancestor around six million years in the past, leading to extant humans and chimpanzees. Therefore, this article proposes an ethological understanding of warfare and social ethics, as both are innate to the social high-specialized modular mind present in the species of genera Pan and Homo. However behavioral restraints to intersocietal coalitionary violence seems to be an exclusive aspect of the transdominial modular cognition that characterizes modern humans. Thus, if in the evolutionary long duree, warfare and restrictions to intrasocial violence both appear to be ethologically common to humans and chimpanzees to a certain extent, an ethics of warfare - and, of course, the cognitive capability for intersocietal peace - seems to be distinctly human.

Social Attention in the Two Species of Pan: Bonobos Make More Eye Contact than Chimpanzees.

Humans’ two closest primate living relatives, bonobos and chimpanzees, differ behaviorally, cognitively, and emotionally in several ways despite their general similarities. While bonobos show more affiliative behaviors towards conspecifics, chimpanzees display more overt and severe aggression against conspecifics. From a cognitive standpoint, bonobos perform better in social coordination, gaze-following and food-related cooperation, while chimpanzees excel in tasks requiring extractive foraging skills. We hypothesized that attention and motivation play an important role in shaping the species differences in behavior, cognition, and emotion. Thus, we predicted that bonobos would pay more attention to the other individuals’ face and eyes, as those are related to social affiliation and social coordination, while chimpanzees would pay more attention to the action target objects, as they are related to foraging. Using eye-tracking we examined the bonobos’ and chimpanzees’ spontaneous scanning of pictures that included eyes, mouth, face, genitals, and action target objects of conspecifics. Although bonobos and chimpanzees viewed those elements overall similarly, bonobos viewed the face and eyes longer than chimpanzees, whereas chimpanzees viewed the other elements, the mouth, action target objects and genitals, longer than bonobos. In a discriminant analysis, the individual variation in viewing patterns robustly predicted the species of individuals, thus clearly demonstrating species-specific viewing patterns. We suggest that such attentional and motivational differences between bonobos and chimpanzees could have partly contributed to shaping the species-specific behaviors, cognition, and emotion of these species, even in a relatively short period of evolutionary time.

Morphometric and molecular analyses of two digenean species from the mullet: Haplosplanchnus pachysomus (Eysenhardt, 1892) from Vietnam and Provitellotrema crenimugilis Pan, 1984 from the Russian southern Far East.

Adults of Haplosplanchnus pachysomus (Eysenhardt, 1829) were found in the intestine of Liza engeli (Bleeker) from the coastal waters of Cat Ba Island, Ha Long Bay, northern Vietnam. Additionally, Provitellotrema crenimugilis Pan, 1984 was discovered in Liza haematocheila (Temminck & Schlegel) from Vostok Bay, Gulf of Peter the Great, southern Far-East Russia. Data concerning morphology, 18S rDNA and 28S rDNA of these worms were obtained.The molecular data confirmed the validity of these species and showed that specimens identified as H. pachysomus are closely related to specimens of H. pachysomus found in Spain, and that P. crenimugilis is closely related to Haplosplanchnus purii. Molecular differentiation of P. crenimugilis and H. purii was 0.92% by combined ribosomal gene sequences that confirmed species validity. Molecular differentiation between P. crenimugilis and H. purii, on the one hand, and H. pachysomus, on the other hand, was much higher, suggesting that the sequence for H. purii in GenBank is for a misidentified species of the genus Provitellotrema Pan, 1984.

A comparative assessment of handedness and its potential neuroanatomical correlates in chimpanzees (Pan troglodytes) and bonobos (Pan paniscus)

The evolutionary origins of human right-handedness remain poorly understood. Some have hypothesized that tool use served as an important preadaptation for the eventual evolution of population-level right-handedness. In contrast, others have suggested that complex gestural and vocal communication served as prerequisite for the evolution of human right-handedness. In this study, we tested these competing hypotheses by comparing the handedness of bonobos and chimpanzees, two closely related species of Pan, on three different measures of hand use including simple reaching, manual gestures and coordinated bimanual actions. Chimpanzees are well known for their tool using abilities whereas bonobos rarely use tools in the wild. In contrast, many have suggested that bonobos have a more flexible gestural and vocal communication system than chimpanzees. The overall results showed that chimpanzees were significantly more right-handed than bonobos for all three measures suggesting that adaptations for tool use rather than communication may have led to the emergence of human right-handedness. We further show that species differences in handedness may be linked to variation in the size and asymmetry of the motor-hand area of the precentral gyrus. The results are discussed within the context of evolutionary theories of handedness, as well as some limitations in the approach to handedness measurement in nonhuman primates.

The Denisova hominin need not be an out of Africa story

This study assessed variation in the supraorbital and orbital region of the Middle Pleistocene hominins (MPHs), sometimes called Homo heidelbergensis s.l., to test whether it matched the expectations of intraspecific variation. The morphological distinctiveness and relative variation of this region, which is relatively well represented in the hominin fossil record, was analyzed quantitatively in a comparative taxonomic framework. Coordinates of 230 3D landmarks (20) and sliding semilandmarks (210) were collected from 704 specimens from species of Homo, Australopithecus, Paranthropus, Gorilla, Pan, Papio, and Macaca. Results showed that the MPHs had expected levels of morphological distinctiveness and intragroup and intergroup variation in supraorbital and orbital morphology, relative to commonly recognized non-hominin catarrhine species. However, the Procrustes distances between this group and H. sapiens were significantly higher than expected for two closely related catarrhine species. Furthermore, this study showed that variation within the MPH could be similarly well contained within existing hypodigms of H. sapiens, H. neanderthalensis, and H. erectus s.l. Although quantitative assessment of supraorbital and orbital morphology did not allow differentiation between taxonomic hypotheses in later Homo, it could be used to test individual taxonomic affiliation and identify potentially anomalous individuals. This study confirmed a complicated pattern of supraorbital and orbital morphology in the MPH fossil record and raises further questions over our understanding of the speciation of H. sapiens and H. neanderthalensis and taxonomic diversity in later Homo.

Mitochondrial genomic divergence in coelacanths (Latimeria): slow rate of evolution or recent speciation?

Dating the divergence between the two known living species of coelacanths has remained a difficult issue because of the very ancient origin of this lineage of fish, which is more closely related to tetrapods than to other fishes. We sequenced the complete mitochondrial genome of a recently captured individual of the Indonesian coelacanth in order to solve this issue. Using an approach based on loglinear models, we studied the molecular divergence between the two species of coelacanths and three other pairs of species, one that has diverged recently (Pan) and two that have diverged more distantly in the past. The loglinear models showed that the divergence between the two species of coelacanths is not significantly different from the two species of Pan. A detailed gene by gene analysis of the patterns of nucleotide and amino acid substitutions between these two pairs of species further supports the similarity of these divergences. On the other hand, a molecular dating analysis suggested a much older origin of the two coelacanth species (between 20 and 30 million years ago). We discuss the potential reasons for this discrepancy. The analysis of new individuals of the Indonesian coelacanth will certainly help to solve this issue.

The vertebral formula of the last common ancestor of African apes and humans

The modal number of lumbar vertebrae in modern humans is five. It varies between three and four in extant African apes (mean=3.5). Because both chimpanzees (Pan troglodytes) and gorillas (Gorilla gorilla) possess the same distributions of thoracic, lumbar, and sacral vertebrae, it has been assumed from parsimony that the last common ancestor (LCA) of African apes and humans possessed a similarly short lower back. This "short-backed LCA" scenario has recently been viewed favorably in an analysis of the intra- and interspecific variation in axial formulas observed among African apes and humans (Pilbeam, 2004. J Exp Zool 302B:241-267). However, the number of bonobo (Pan paniscus) specimens in that study was small (N=17). Here we reconsider vertebral type and number in the LCA in light of an expanded P. paniscus sample as well as evidence provided by the human fossil record. The precaudal (pre-coccygeal) axial column of bonobos differs from those of chimpanzees and gorillas in displaying one additional vertebra as well as significantly different combinations of sacral, lumbar, and thoracic vertebrae. These findings, along with the six-segmented lumbar column of early Australopithecus and early Homo, suggest that the LCA possessed a long axial column and long lumbar spine and that reduction in the lumbar column occurred independently in humans and in each ape clade, and continued after separation of the two species of Pan as well. Such an explanation is strongly congruent with additional details of lumbar column reduction and lower back stabilization in African apes.

Volumetric and lateralized differences in selected brain regions of chimpanzees (Pan troglodytes) and bonobos (Pan paniscus)

The two species of Pan, bonobos and common chimpanzees, have been reported to have different social organization, cognitive and linguistic abilities and motor skill, despite their close biological relationship. Here, we examined whether bonobos and chimpanzee differ in selected brain regions that may map to these different social and cognitive abilities. Eight chimpanzees and eight bonobos matched on age, sex and rearing experiences were magnetic resonance images scanned and volumetric measures were obtained for the whole brain, cerebellum, striatum, motor-hand area, hippocampus, inferior frontal gyrus and planum temporale. Chimpanzees had significantly larger cerebellum and borderline significantly larger hippocampus and putamen, after adjusting for brain size, compared with bonobos. Bonobos showed greater leftward asymmetries in the striatum and motor-hand area compared with chimpanzees. No significant differences in either the volume or lateralization for the so-called language homologs were found between species. The results suggest that the two species of Pan are quite similar neurologically, though some volumetric and lateralized differences may reflect inherent differences in social organization, cognition and motor skills.

Population systematics of chimpanzees using molar morphometrics

When dental morphological variation within extant species is used as a guideline to partition variation within fossil samples into species, the underlying assumption is that fossil species are equivalent to extant species. This is the case despite the fact that dental morphology, which is commonly used to differentiate fossil species, is rarely used to differentiate extant species. Aspects of external morphology, ecology, behavior, breeding patterns, and molecular structure that are used to delineate living species are generally not available for fossils. In this paper, the utility of dental evidence for sorting fossil samples into species is evaluated by testing whether molar occlusal morphology is capable of sorting populations of Pan into the species and subspecies already well-established by nondental evidence. The dentitions of 341 chimpanzee individuals, sampled from regions throughout equatorial Africa, were sorted into 16 populations using rivers to demarcate the boundaries between populations. Digital-imaging software was used to measure 15 traits on the occlusal surface of each upper molar and 19 on each lower molar. After applying size adjustments, size-transformed and untransformed variables were subjected to discriminant analysis, with separate analyses carried out for each molar type. Results indicate that populations of Pan troglodytes and Pan paniscus are well differentiated at all molar positions. Populations of P. t. verus are distinct from other populations of P. troglodytes. Populations of P. t. troglodytes and P. t. schweinfurthii show close dental similarity. A distinct population is recognized at the Nigeria-Cameroon border, indicating the presence of P. t. vellerosus. The concordance between the patterns of diversity recognized by this study and other molecular and nonmolecular studies indicates that paleontological species that are similar to species of Pan in terms of size and patterns of diversification may be differentiated using molar morphology.

The development of representational play in chimpanzees and bonobos: Evolutionary implications, pretense, and the role of interspecies communication

This research investigates the development of symbolic or representational play in two species of the genus Pan, bonobos ( Pan paniscus) and chimpanzees ( Pan troglodytes) . The participants varied not only by species, but also as to whether they had become proficient in communicating with humans via a set of arbitrary visual symbols, called lexigrams. Using a developmental sequence of representational play based on McCune, we found every level that children manifest to be constructed by Pan. The most robust and regular ontogenetic sequence for both bonobos and chimpanzee was not McCune's five-level progression, but a three-step ontogenetic sequence: Level 1 (no representation, no pretense) precedes Levels 2–4 (representation but no pretense), which in turn precedes Level 5 (includes pretense as well as representation). A linguistic system for interspecies communication was necessary for Level 5 representational play and “true” pretense. Human scaffolding produced developmental progress within sequences for all the apes, except the bonobo who lacked a system of interspecies communication. This evidence suggests that the potential for representational play and its social stimulation were present in the common ancestor of bonobos, chimpanzees and humans five million years ago.

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.