Relationships between the hard and soft dimensions of the nose in Pan troglodytes and Homo sapiens reveal the positions of the nasal tips of Plio-Pleistocene hominids.

By identifying homogeneity in bone and soft tissue covariation patterns in living hominids, it is possible to produce facial approximation methods with interspecies compatibility. These methods may be useful for producing facial approximations of fossil hominids that are more realistic than currently possible. In this study, we conducted an interspecific comparison of the nasomaxillary region in chimpanzees and modern humans with the aim of producing a method for predicting the positions of the nasal tips of Plio-Pleistocene hominids. We addressed this aim by first collecting and performing regression analyses of linear and angular measurements of nasal cavity length and inclination in modern humans (Homo sapiens; n = 72) and chimpanzees (Pan troglodytes; n = 19), and then performing a set of out-of-group tests. The first test was performed on four subjects that belonged to the same genus as the training sample, i.e., Homo (n = 2) and Pan (n = 2), and the second test, which functioned as an interspecies compatibility test, was performed on Pan paniscus (n = 1), Gorilla gorilla (n = 3), Pongo pygmaeus (n = 1), Pongo abelli (n = 1), Symphalangus syndactylus (n = 3), and Papio hamadryas (n = 3). We identified statistically significant correlations in both humans and chimpanzees with slopes that displayed homogeneity of covariation. Prediction formulae combining these data were found to be compatible with humans and chimpanzees as well as all other African great apes, i.e., bonobos and gorillas. The main conclusion that can be drawn from this study is that our set of regression models for approximating the position of the nasal tip are homogenous among humans and African apes, and can thus be reasonably extended to ancestors leading to these clades.

The character of parietal and orbital alterations in the superfamily Hominoidea (families Hominidae [exclusive of Homo] and Hylobotidae).

Parietal external surface disruption routinely referred to as porotic hyperostosis, and orbital alterations (cribra orbitalia), have been attributed to anemia-related bone marrow hyperplasia in humans. A recent study in humans identified that they were actually vascular in nature. Skeletons were examined and epi-illumination surface microscopy was performed on the parietal region and orbit of 156 Hominidae and 123 Hylobotidae to assess if these phenomena were trans-phylogenetic. Trans-cortical channels were recognized on the basis of visualized ectocranial surface defects penetrating the parietal; cribra orbitalia, by alteration of the normally smooth orbital roof appearance. Trans-cortical parietal channels, ranging in size from 20 to 100 µm, are rare in Gorilla and Pan troglodytes and absent in Pan paniscus. They are universally present in adult Pongo abeli and in Hylobatidae, independent of species. Cribra orbitalia was common in Hylobotidae, Pongo pygmaeus and P. abelii, less prevalent in adult P. troglodytes, and not recognized in any Gorilla gorilla or P. paniscus examined. The proliferative form predominated, with the exception of Hylobates concolor and muelleri, in which uncalcified vascular grooves predominated. No correlation was observed between the presence of either trans-cortical channels or cribra orbitalia and fractures, osteoarthritis, or inflammatory arthritis. Parietal alterations observed in apes are trans-cortical channels, analogous to those observed in humans, and do not represent porosity. Similarly, cribra orbitalia in apes is confirmed as vascular in nature. The proliferative form apparently represents calcification of blood vessel walls, indistinguishable from observations in humans. Predominant presence in adults rather than in juveniles suggests that both forms are acquired rather than developmental in derivation. Sex and bone alteration/disease-independence suggests that mechanical, endocrine, and inflammatory phenomena do not contribute to the development of either. Further, independent occurrence of trans-cortical channels and cribra orbitalia suggests that they do not have a shared etiology.

Metameric variation of upper molars in hominoids and its implications for the diversification of molar morphogenesis.

Metameric variation of molar size is in part associated with the dietary adaptations of mammals and results from slight alterations of developmental processes. Humans and great apes exhibit conspicuous variation in tooth morphology both between taxa and across tooth types. However, the manner in which metameric variation in molars emerged among apes and humans via evolutionary alterations in developmental processes remains largely unknown. In this study, we compare the enamel-dentine junction of the upper molars of humans-which closely correlates with morphology of the outer enamel surface and is less affected by wear-with that of the other extant hominoids: chimpanzees, bonobos, gorillas, orangutans, and gibbons. We used the morphometric mapping method to quantify and visualize three-dimensional morphological variation, and applied multivariate statistical analyses. Results revealed the following: 1) extant hominoids other than humans share a common pattern of metameric variation characterized by a largely linear change in morphospace; this indicates a relatively simple graded change in metameric molar shape; 2) intertaxon morphological differences become less distinct from the mesial to distal molars; and 3) humans diverge from the extant ape pattern in exhibiting a distinct metameric shape change trajectory in the morphospace. The graded shape change and lower intertaxon resolution from the mesial to distal molars are consistent with the concept of a 'key' tooth. The common metameric pattern observed among the extant nonhuman hominoids indicates that developmental patterns underlying metameric variation were largely conserved during ape evolution. Furthermore, the human-specific metameric pattern suggests considerable developmental modifications in the human lineage.

Ecomorphological specialization leads to loss of evolvability in primate limbs.

Primate limb morphology is often described as either generalized, that is, suited to a range of locomotor and positional behaviors, or specialized for unique locomotor behaviors such as brachiation or bipedalism. The evolution of highly specialized limb morphology may result in loss of evolvability, that is, in a decreased capacity of the locomotor skeleton to evolve in response to selection towards alternative ecomorphological niches. Using evolutionary simulations, I show that the highly specialized limb anatomy of hominoids is associated with a significant loss of evolvability, defined as the number of generations to reach alternative adaptive peaks, and in parallel an increased risk of extinction, particularly in simulated evolution toward generalized quadrupedal limb proportions. Loss of evolvability in apes and humans correlates with three factors: (1) decreased correlation among limb bone lengths (i.e., integration), which slows the rate of change along lines of least evolutionary resistance; (2) limb specialization, which places apes and humans in relatively remote areas of morphospace; and (3) increased skeletal size as a proxy for body size. Thus, locomotor over-specialization can lead to evolutionary dead-ends that significantly increase the probability of hominoid populations going extinct before evolving new adaptive morphologies.

The gibbon's Achilles tendon revisited: consequences for the evolution of the great apes?

The well-developed Achilles tendon in humans is generally interpreted as an adaptation for mechanical energy storage and reuse during cyclic locomotion. All other extant great apes have a short tendon and long-fibred triceps surae, which is thought to be beneficial for locomotion in a complex arboreal habitat as this morphology enables a large range of motion. Surprisingly, highly arboreal gibbons show a more human-like triceps surae with a long Achilles tendon. Evidence for a spring-like function similar to humans is not conclusive. We revisit and integrate our anatomical and biomechanical data to calculate the energy that can be recovered from the recoiling Achilles tendon during ankle plantar flexion in bipedal gibbons. Only 7.5% of the required external positive work in a stride can come from tendon recoil, yet it is delivered at an instant when the whole-body energy level drops. Consequently, an additional similar amount of mechanical energy must simultaneously dissipate elsewhere in the system. Altogether, this challenges the concept of an energy-saving function in the gibbon's Achilles tendon. Cercopithecids, sister group of the apes, also have a human-like triceps surae. Therefore, a well-developed Achilles tendon, present in the last common 'Cercopithecoidea-Hominoidea' ancestor, seems plausible. If so, the gibbon's anatomy represents an evolutionary relict (no harm-no benefit), and the large Achilles tendon is not the premised key adaptation in humans (although the spring-like function may have further improved during evolution). Moreover, the triceps surae anatomy of extant non-human great apes must be a convergence, related to muscle control and range of motion. This perspective accords with the suggestions put forward in the literature that the last common hominoid ancestor was not necessarily great ape-like, but might have been more similar to the small-bodied catarrhines.

Hip extensor mechanics and the evolution of walking and climbing capabilities in humans, apes, and fossil hominins.

The evolutionary emergence of humans' remarkably economical walking gait remains a focus of research and debate, but experimentally validated approaches linking locomotor capability to postcranial anatomy are limited. In this study, we integrated 3D morphometrics of hominoid pelvic shape with experimental measurements of hip kinematics and kinetics during walking and climbing, hamstring activity, and passive range of hip extension in humans, apes, and other primates to assess arboreal-terrestrial trade-offs in ischium morphology among living taxa. We show that hamstring-powered hip extension during habitual walking and climbing in living apes and humans is strongly predicted, and likely constrained, by the relative length and orientation of the ischium. Ape pelves permit greater extensor moments at the hip, enhancing climbing capability, but limit their range of hip extension, resulting in a crouched gait. Human pelves reduce hip extensor moments but permit a greater degree of hip extension, which greatly improves walking economy (i.e., distance traveled/energy consumed). Applying these results to fossil pelves suggests that early hominins differed from both humans and extant apes in having an economical walking gait without sacrificing climbing capability. Ardipithecus was capable of nearly human-like hip extension during bipedal walking, but retained the capacity for powerful, ape-like hip extension during vertical climbing. Hip extension capability was essentially human-like in Australopithecus afarensis and Australopithecus africanus, suggesting an economical walking gait but reduced mechanical advantage for powered hip extension during climbing.

Body mass estimation in hominoids: Age and locomotor effects.

While there are a number of methods available for estimation of body mass in adult nonhuman primates, very few are available for juveniles, despite the potential utility of such estimations in both analyses of fossils and in museum collection based research. Furthermore, because of possible scaling differences, adult based body mass estimation equations may not be appropriate for non-adults. In this study, we present new body mass estimation equations for both adult and immature nonhuman hominoids based on joint and metaphyseal dimensions. Articular breadths of the proximal and distal femur, distal humerus and tibial plateau, and metaphyseal breadths of the distal femur and humerus were collected on a reference sample of 159 wild Pan, Gorilla, Pongo, Hylobates, and Symphalangus specimens of known body mass from museum and research collections. Scaling of dimensions with body weight was assessed in both the adult and the ontogenetic sample at several taxonomic levels using reduced major axis regression, followed by regression of each dimension against body mass to generate body mass estimation equations. Joint dimensions were found to be good predictors of body mass in both adult and immature hominoids, with percent prediction errors of 10-20%. However, subtle scaling differences between taxa impacted body mass estimation, suggesting that phylogeny and locomotor effects should be considered when selecting reference samples. Unlike patterns of joint growth in humans, there was little conclusive evidence for consistently larger joints relative to body mass in the non-adult sample. Metaphyseal breadths were strong predictors of body mass and, with some exceptions, gave more precise body mass estimates for non-adults than epiphyseal breadths.

Description of a new species of Hoolock gibbon (Primates: Hylobatidae) based on integrative taxonomy.

We describe a species of Hoolock gibbon (Primates: Hylobatidae) that is new to science from eastern Myanmar and southwestern China. The genus of hoolock gibbons comprises two previously described living species, the western (Hoolock hoolock) and eastern hoolock (H. leuconedys) gibbons, geographically isolated by the Chindwin River. We assessed the morphological and genetic characteristics of wild animals and museum specimens, and conducted multi-disciplinary analyses using mitochondrial genomic sequences, external morphology, and craniodental characters to evaluate the taxonomic status of the hoolock population in China. The results suggest that hoolocks distributed to the east of the Irrawaddy-Nmai Hka Rivers, which were previously assigned to H. leuconedys, are morphologically and genetically distinct from those to the west of the river, and should be recognized as a new species, the Gaoligong hoolock gibbon or skywalker hoolock gibbon (H. tianxing sp. nov.). We consider that the new species should be categorized as Endangered under IUCN criteria. The discovery of the new species focuses attention on the need for improved conservation of small apes, many of which are in danger of extinction in southern China and Southeast Asia.

Scapular shape of extant hominoids and the African ape/modern human last common ancestor.

Newly discovered early hominin fossil scapulae have bolstered investigations of scapular shape, which have long been used to interpret behavioral variation among primates. However, unexpected similarities between Pongo and Homo - particularly in scapular spine orientation - have raised questions about the functional utility of scapular morphology and its phylogenetic context in the hominin lineage. Not surprisingly, significant disagreement surrounds disparate morphological reconstructions of the modern human/African ape last common ancestor (LCA). Our study utilizes geometric morphometric (GM) approaches - two employing homologous, anatomical landmarks and a "spine-free" alternative using 98 sliding semilandmarks along the boundary of the subscapular fossa. The landmark-based "wireframe" GM analysis principally sorted groups by spine orientation: Homo and Pongo were similar to one another with more transversely-oriented spines as compared to Hylobates and the African apes. In contrast, Homo and Gorilla clustered together in our semilandmark analysis with superoinferiorly broad blades. Pan scapulae were similar, but had more mediolaterally compressed blades and laterally-positioned superior angles. Hylobates was superoinferiorly narrow, yet obliquely expanded relative to the vertebral border. Pongo scapulae were unique among hominoids in being nearly as broad as they were long. Previously documented 'convergence' of Homo and Pongo scapulae appears to be principally driven by similarities in spine orientation, rather than overall blade shape. Therefore, we contend that it is more parsimonious to reconstruct the African ape/Homo LCA scapula as being Gorilla-like, especially in light of similar characterizations of certain fossil hominin scapulae. Accordingly, the evolution of Pan (highly oblique spine and laterally-situated superior angle) and Homo (transversely-oriented spine) scapular morphology would have involved relatively minor shifts from this ancestral condition. These results support the prevailing molecular phylogeny and provide further insight into the behavioral implications of scapular shape in the LCA and fossil hominins.

Giant pandas (Carnivora: Ailuropoda melanoleuca) and living hominoids converge on lumbar vertebral adaptations to orthograde trunk posture.

Living hominoids share a common body plan characterized by a gradient of derived postcranial features that distinguish them from their closest living relatives, cercopithecoid monkeys. However, the evolutionary scenario(s) that led to the derived postcranial features of hominoids are uncertain. Explanations are complicated by the fact that living hominoids vary considerably in positional behaviors, and some Miocene hominoids are morphologically, and therefore probably behaviorally, distinct from modern hominoids. Comparative studies that aim to identify morphologies associated with specific components of positional behavioral repertoires are an important avenue of research that can improve our understanding of the evolution and adaptive significance of the hominoid postcranium. Here, we employ a comparative approach to offer additional insight into the evolution of the hominoid lumbar vertebral column. Specifically, we tested whether giant pandas (Carnivora: Ailuropoda melanoleuca) converge with living hominoids on lumbar vertebral adaptations to the single component of their respective positional behavioral repertoires that they share--orthograde (i.e., upright) trunk posture. We compare lumbar vertebral morphologies of Ailuropoda to those of other living ursids and caniform outgroups (northern raccoons and gray wolves). Mirroring known differences between living hominoids and cercopithecoids, Ailuropoda generally exhibits fewer, craniocaudally shorter lumbar vertebrae with more dorsally positioned transverse processes that are more dorsally oriented and laterally directed, and taller, more caudally directed spinous processes than other caniforms in the sample. Our comparative evidence lends support to a potential evolutionary scenario in which the acquisition of hominoid-like lumbar vertebral morphologies may have evolved for generalized orthograde behaviors and could have been exapted for suspensory behavior in crown hominoids and for other locomotor specializations (e.g., brachiation) in extant lineages.

Genetic comparisons yield insight into the evolution of enamel thickness during human evolution.

Enamel thickness varies substantially among extant hominoids and is a key trait with significance for interpreting dietary adaptation, life history trajectory, and phylogenetic relationships. There is a strong link in humans between enamel formation and mutations in the exons of the four genes that code for the enamel matrix proteins and the associated protease. The evolution of thick enamel in humans may have included changes in the regulation of these genes during tooth development. The cis-regulatory region in the 5' flank (upstream non-coding region) of MMP20, which codes for enamelysin, the predominant protease active during enamel secretion, has previously been shown to be under strong positive selection in the lineages leading to both humans and chimpanzees. Here we examine evidence for positive selection in the 5' flank and 3' flank of AMELX, AMBN, ENAM, and MMP20. We contrast the human sequence changes with other hominoids (chimpanzees, gorillas, orangutans, gibbons) and rhesus macaques (outgroup), a sample comprising a range of enamel thickness. We find no evidence for positive selection in the protein-coding regions of any of these genes. In contrast, we find strong evidence for positive selection in the 5' flank region of MMP20 and ENAM along the lineage leading to humans, and in both the 5' flank and 3' flank regions of MMP20 along the lineage leading to chimpanzees. We also identify putative transcription factor binding sites overlapping some of the species-specific nucleotide sites and we refine which sections of the up- and downstream putative regulatory regions are most likely to harbor important changes. These non-coding changes and their potential for differential regulation by transcription factors known to regulate tooth development may offer insight into the mechanisms that allow for rapid evolutionary changes in enamel thickness across closely-related species, and contribute to our understanding of the enamel phenotype in hominoids.

Micro-finite element (µFE) modeling of the siamang (Symphalangus syndactylus) third proximal phalanx: the functional role of curvature and the flexor sheath ridge.

Phalangeal curvature is a commonly used morphological feature for the interpretation of extant and fossil primate locomotor behaviour. Here, we build on a recent biomechanical study (Richmond, 2007) in two ways: first, we use a 3D micro-FE model, which models the real internal microstructure (i.e., cortical thickness and trabecular bone structure) and, second, we model four siamang third proximal phalanges. We test identical 2D homogenized FE models and two 3D micro-FE phalanx models that are mathematically straightened to isolate the biomechanical significance of curvature. We further investigate how varying the loading configuration (e.g., boundary constraints) and modeling (e.g., 2D versus 3D) affects the biomechanical behaviour of the phalanx. Finally, we examine how intraspecific variation in external and internal bony morphology affects the biomechanical behaviour of the phalanx. Simulation results demonstrate that the general pattern of strain and displacement is similar between the 3D micro-FE and 2D homogenized FE models but the absolute values differ substantially. The biomechanical behaviour of the 3D FE models more closely match the relative strain patterns from the validation experiment than the 2D homogenized FE models, indicating the 3D microstructure model is preferable. Varying the loading configuration can have dramatic effects on the biomechanical behaviour of the phalanx depending on individual morphology, but overall a cantilevered beam model is an equally valid, if not better, configuration for modeling the phalanx as other previously-proposed models. Variation in flexor ridge morphology has a substantial effect on phalanx strain; the taller the ridge, the less strain incurred by other regions of the palmar shaft. Finally, phalangeal curvature reduces overall strain experienced by the phalanx, but does not necessarily reduce bending or increase the compression-to-tension ratio. These results confirm the adaptive role of phalangeal curvature during flexed-finger grasping postures and demonstrate that modeling variation in cortical thickness and flexor ridge morphology improves the behaviour of the FE model, which has important implications for the functional interpretation of phalanx form.

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

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

A comparative study of crested gibbons (Nomascus).

Crested gibbons (Nomascus) are in the rarest genus of the family Hylobatidae, with the Hainan gibbon (Nomascus hainanus) being the rarest primate in the world. In the past, the number of species in this genus has been at the center of much controversy, in part, because their color changes during immaturity as well as other factors, such as physical similarities in genitalia, creating difficulties in accurately determining the sex of individuals. Furthermore, owing to their rarity, illusiveness, and the rough terrain that comprises their native habitat, Nomascus is one of the least studied Hylobatidae. This article represents the most comprehensive dissemination of visual characteristics of the genus Nomascus to assist in the accurate identification of captive and wild crested gibbons. Through differences in pelage color, skeletal anatomy, dentition, vocalizations, behavior, distribution, and genetic studies, we are able to determine more accurately whether or not a subspecies should be elevated to species level. From the current data, there are six species and one subspecies in the genus Nomascus. However, reports of a recently identified light-cheeked gibbon (Nomascus sp.) in northeast Cambodia, Central Vietnam, and South Lao PDR, will add additional taxa to this genus.

A comparative analysis of internal cranial anatomy in the hylobatidae.

Craniometric studies on the hylobatids using external metrics (Creel and Preuschoft, 1976, 1984) sorted hylobatid populations into primary species groupings which are in accordance with the four currently recognized generic-level groupings. The goal of the current study was to assess the relative orientations of the orbits, palate, and basioccipital clivus among the hylobatid genera in an effort to further clarify whether the lesser apes differ significantly in these internal cranial features and how that variation patterns across the groups. Nine angular variables quantifying orbital, palatal, and basioccipital clivus orientations were measured on lateral view radiographs of adults representing three of the four hylobatid genera: Hylobates; Nomascus; and, Symphalangus. The interspecific adult hylobatid means for the angular variables were analyzed using t-test contrasts. The total sample was further subjected to discriminant function analysis (DFA) to test for the ability of craniofacial angular variables to distinguish the hylobatid genera from one another. The three hylobatid genera displayed significant morphological differentiation in orbital, palatal, and posterior skull base orientations. Normal, jackknifed, and cross-validation DFA procedures correctly identified the hylobatids 50-100% of the time. The observed morphological patterns generally mapped onto the findings of earlier external craniometric hylobatid studies and suggest concordance between specific internal and external cranial features. This article is the first comprehensive study of variation in internal cranial anatomy of the Hylobatidae and includes the first published craniofacial angular data for Nomascus.

Technical note: Forearm pronation efficiency analysis in skeletal remains.

This work presents an original methodology for analyzing forearm-pronation efficiency from skeletal remains and its variation with regard to changes in the elbow position. The methodology is based on a biomechanical model that defines rotational efficiency as a mathematical function expressing a geometrical relationship between the origin and insertion of the pronator teres. The methodology uses humeral distal epiphysis photography, from which the geometrical parameters for the efficiency calculus can be obtained. Rotational efficiency is analyzed in a human specimen and in a living nonhuman hominoid (Symphalangus syndactylus) for a full elbow extension (180 degrees) and an intermediate elbow position (90 degrees). In both specimens, the results show that this rotational-efficiency parameter varies throughout the entire rotational range and show a dependency on the elbow joint position. The rotational efficiency of the siamang's pronator teres is less affected by flexion of the forearm than that of the human. The fact that forearm-pronation efficiency can be inferred, even quantified, allows us to interpret more precisely the functional and evolutionary significance of upper-limb skeletal design in extant and fossil primate taxa.

Systematic morphology and evolutionary anatomy of the autonomic cardiac nervous system in the lesser apes, gibbons (hylobatidae).

We examined the morphology of the autonomic cardiac nervous system (ACNS) on 20 sides of 10 gibbons (Hylobatidae) of three genera, and we have inferred the evolution of the anatomy of the primate ACNS. We report the following. (1) Several trivial intraspecific and interspecific variations are present in gibbons, but the general arrangement of the ACNS in gibbons is consistent. (2) Although the parasympathetic vagal cardiac nervous system is extremely consistent, the sympathetic cardiac nervous system, such as the composition of the sympathetic ganglia and the range of origin of the sympathetic cardiac nerves, exhibit topographical differences among primates. (3) The vertebral ganglion, seldom observed in the Old World monkeys (Cercopithecidae), was consistently present in gibbons as well as in humans. (4) There are fewer thoracic ganglia contributing to the cervicothoracic ganglion in humans than in gibbons and in gibbons than in Old World monkeys. (5) The superior cardiac nerve originating from the superior cervical ganglion, rarely observed in Old World monkeys but commonly observed in humans, was present in 13 of 20 sides (65%), mostly on the left. Accordingly, the ACNS morphology exhibits evolutionary changes within the primate lineage. These evolutionary differences between Old World monkeys, gibbons, and humans are most parsimoniously interpreted as resulting from regular changes in the lineages leading from their common ancestor to the extant species that we dissected. They include the reduction in the number of thoracic ganglia contributing to the cervicothoracic ganglion and the expansion of the range of the cardiac nervous origin.

The deciduous lower dentition of Ouranopithecus macedoniensis (Primates, Hominoidea) from the late Miocene deposits of Macedonia, Greece.

Two mandibular fragments with associated milk teeth assigned to the late Miocene hominoid primate Ouranopithecus macedoniensis are analyzed. The fossils, which belong to a single individual, were found in the Vallesian locality of "Ravin de la Pluie" of the Axios Valley (Macedonia, Greece). The material is described here and compared with extant and extinct hominoids, allowing assessment of the evolutionary trends in the deciduous lower dentition within the Hominoidea. Hylobatids represent the more primitive pattern. Gorilla is slightly more derived than hylobatids, but less derived than Pongo and Pan, the latter being the most derived. With relatively smaller deciduous canines and more molarized deciduous premolars, Ouranopithecus is more derived than both Pan and Gorilla. Among the fossil hominoids, Proconsul, representing the primitive condition, has a very simple dp(3)and a dp(4)that has a trigonid that is taller than the talonid and which lacks a hypoconulid. Griphopithecus is more derived than Proconsul in having a dp(4) with a lower trigonid, a hypoconulid, and a less oblique cristid obliqua. Australopithecus and Paranthropus possess a similar morphology to that of Homo, while Ardipithecus appears to be more primitive than the latter genera. Ouranopithecus has a more derived lower milk dentition than Proconsul and Griphopithecus, but less derived than Australopithecus and Paranthropus. The comparison of the lower milk dentition of Ouranopithecus confirms our previous conclusions suggesting that this fossil hominoid shares derived characters with Australopithecus and Homo.

Mosaic evolution in the origin of the Hominoidea.

The initial appearance of hominoids, or apes, and the selective pressures that led to their emergence are currently disputed. Central to the argument are the proconsulids, variously described as the earliest apes or as stem catarrhines, based on facial and postcranial data, respectively. The present paper reports on incongruence and parsimony analyses applied to a combined data set. The results demonstrate that proconsulids are cladistic hominoids, and that the apparent incongruence between the data sets is due to mosaic evolution; the earliest changes in Hominoidea occurred in the face. These results suggest that the initial divergence of hominoids involved selection for an ape-like face, and was not driven by an adaptive shift to below-branch locomotion.