Tooth Wear Difference Between the Yuanmou Hominoid and <Emphasis Type="Italic">Lufengpithecus</Emphasis>

The Late Miocene hominoids recovered from Lufeng (Lufengpithecus) and Yuanmou of Yunnan Province, China, are among the most numerous hominoid fossils in Eurasia. They have yielded critical evidence for the evolutionary history, biogeography and paleobiology of Miocene hominoids. We examined and compared the wear pattern and differences of 804 molars of the Yuanmou hominoid and Lufengpithecus. Our results indicate that both the upper and lower molars of the Yuanmou hominoids were more heavily worn than those of Lufengpithecus. The wear patterns of the individual molars between the Yuanmou hominoid and Lufengpithecus also are different. The heaviest wear of lower molars of the Yuanmou hominoid occur in M₂, followed by M₁ and M₃. In Lufengpithecus, M₁ and M₃ were more heavily worn than M₂. There are differences in wear between the upper and lower molars for the two hominoids. Among the various factors related to tooth wear, we suggest that the main reason for the tooth wear differences between the Yuanmou hominoid and Lufengpithecus may be that they had different diets. More soft dietary items like leaves and berries were probably consumed by Lufengpithecus, and the Yuanmou hominoid may mainly have feed on harder or frugivorous diets. This result complements findings from previous studies of tooth size proportion, and the development of lower molar shearing crests in the 2 samples. Enamel thickness, living environment, behavior patterns, and population structure also might account for dental wear differences between the Yuanmou hominoid and Lufengpithecus.     

Preliminary investigation of dental microstructure in the Yuanmou hominoid (Lufengpithecus hudienensis), Yunnan Province,China

Fieldwork in the Yuanmou Basin of southern China has uncovered a large assemblage of late Miocene hominoid fossils assigned to Lufengpithecus hudienensis. Two mandibular first molars from this species were made available for histological analysis as part of a larger ongoing study on the ontogeny of dental development in Miocene to Recent hominoids. Results are compared with published and unpublished data on tooth growth in a wide range of extant and extinct hominoids. The Yuanmou molars are smaller than those of Lufengpithecus lufengensis and have markedly shorter crown formation times, overlapping slightly with Pan, but most similar to Proconsul and Dryopithecus. In other aspects of molar development (including enamel extension rates and enamel thickness), L. hudienensis shows similarities with all extant hominoids, in particular, Pongo. Ultimately, charting the ontogeny of molar crown formation may help shed light on the relationship of Lufengpithecus hudienensis to orangutans, and other Miocene to Recent hominoids.     

New primate carpal bones from Rudabánya (late Miocene, Hungary): taxonomic and functional implications

We describe a scaphoid and two capitates from the late Miocene site of Rudabánya, Hungary using qualitative and quantitative comparisons to a large sample of hominoid, cercopithecoid, and platyrrhine primates. The scaphoid (RUD 202) is not fused to the os centrale and in this way is like most primates other than African apes and humans (hominines). Qualitatively, its morphology is most similar to Pongo, and univariate analyses generally confirm an ape-like morphology with an increased range of mobility. One capitate (RUD 167) is compatible in size to the scaphoid, and its morphology suggests a combination of monkey-like generalized arboreality and ape-like enhanced mobility. RUD 203 is a smaller, fragmentary capitate, about half the size of RUD 167, and preserves only the distal portion of the body with the third metacarpal articular surface. Its morphology is virtually identical to that of RUD 167, and an exact randomization test revealed that it is statistically likely to find two carpal bones of such disparate sizes within one taxon. However, due to morphological similarities with other Miocene hominoids as well as implications for size variation within one taxon and sex, we consider the taxonomic affiliation of RUD 203 to be unresolved. We attribute the scaphoid and RUD 167 capitate to the hominine Rudapithecus hungaricus (formerly Dryopithecus brancoi; see Begun et al., 2008) based on overall morphological similarity to extant apes, particularly Pongo, and not to the pliopithecoid Anapithecus hernyaki, the only other primate known from Rudabánya. The similarities in carpal morphology to suspensory taxa are consistent with previous interpretations of Rudapithecus positional behavior. The scaphoid and the RUD 167 capitate are consistent in size with a partial skeleton including associated postcranial and craniodental specimens from the same level at the locality and may be from the same individual. These are the first carpal bones described from Rudabánya and from this taxon, and they add to our understanding of the evolution of arboreal locomotion in late Miocene apes.     

Enamel thickness,microstructure and development in Afropithecus turkanensis

Afropithecus turkanensis, a 17-17.5 million year old large-bodied hominoid from Kenya, has previously been reported to be the oldest known thick-enamelled Miocene ape. Most investigations of enamel thickness in Miocene apes have been limited to opportunistic or destructive studies of small samples. Recently, more comprehensive studies of enamel thickness and microstructure in Proconsul, Lufengpithecus, and Dryopithecus, as well as extant apes and fossil humans, have provided information on rates and patterns of dental development, including crown formation time, and have begun to provide a comparative context for interpretation of the evolution of these characters throughout the past 20 million years of hominoid evolution. In this study, enamel thickness and aspects of the enamel microstructure in two A. turkanensis second molars were quantified and provide insight into rates of enamel apposition, numbers of cells actively secreting enamel, and the time required to form regions of the crown. The average value for relative enamel thickness in the two molars is 21.4, which is a lower value than a previous analysis of this species, but which is still relatively thick compared to extant apes. This value is similar to those of several Miocene hominoids, a fossil hominid, and modern humans. Certain aspects of the enamel microstructure are similar to Proconsul nyanzae, Dryopithecus laietanus, Lufengpithecus lufengensis, Graecopithecus freybergi and Pongo pygmaeus, while other features differ from extant and fossil hominoids. Crown formation times for the two teeth are 2.4-2.6 years and 2.9-3.1 years respectively. These times are similar to a number of extant and fossil hominoids, some of which appear to show additional developmental similarities, including thick enamel. Although thick enamel may be formed through several developmental pathways, most Miocene hominoids and fossil hominids with relatively thick enamel are characterized by a relatively long period of cuspal enamel formation and a rapid rate of enamel secretion throughout the whole cusp, but a shorter total crown formation time than thinner-enamelled extant apes.     

Cautious climbing and folivory: a model of hominoid differentation

Despite the large and growing number of Miocene fossil catarrhine taxa, suitable common ancestors of great apes and humans have yet to be agreed upon. Considering a) the conservative and primitive nature of the hominoid molar cusp pattern, and b) the variability of secondary dental features, it is difficult to discern whether a hominoid dentition is primitive, secondarily simplified to the primitive condition or too far derived to be ancestral to any of the living forms. Nonetheless, the inability to recognize a common ancestor is primarly due to the absence of a model of hominoid differentiation that provides a basis for its recognition. Vertical climbing as the limiting component of cautious climbing, explains all of the locomotor anatomy shared by living hominoids. Comparison of the shared derived characters of hominoids to those of forms which have converged on hominoidsi.e colobines, atelines, lorisines, paleopropithecines and sloths suggest that early hominoids were probably folivores. In arboreal forms there is a strong link between a large body size, folivory and cautious climbing. Comparison of craniodental characters of committed folivores to committed frugivores from among each of the compared groups with the exception of lorisines, indicates that many of the distinguishing craniodental characters of humans and great apes are adaptations to folivory. Many of these characters, however, are also present in Jolly's seed eating complex. As such folivory may be the heritage factor which Jolly hypothesized to account for differential reduction of canines in fossilTheropithecus and hominids.     

A morphometric mapping analysis of lower fourth deciduous premolar in hominoids: Implications for phylogenetic relationship between Nakalipithecus and Ouranopithecus

Clarifying morphological variation among African and Eurasian hominoids during the Miocene is of particular importance for inferring the evolutionary history of humans and great apes. Among Miocene hominoids, Nakalipithecus and Ouranopithecus play an important role because of their similar dates on different continents. Here, we quantify the lower fourth deciduous premolar (dp4) inner morphology of extant and extinct hominoids using a method of morphometric mapping and examine the phylogenetic relationships between these two fossil taxa. Our data indicate that early Late Miocene apes represent a primitive state in general, whereas modern great apes and humans represent derived states. While Nakalipithecus and Ouranopithecus show similarity in dp4 morphology to a certain degree, the dp4 of Nakalipithecus retains primitive features and that of Ouranopithecus exhibits derived features. Phenotypic continuity among African ape fossils from Miocene to Plio-Pleistocene would support the African origin of African apes and humans (AAH). The results also suggest that Nakalipithecus could have belonged to a lineage from which the lineage of Ouranopithecus and the common ancestor of AAH subsequently derived.     

Miocene hominoid palatofacial morphology

The palatofacial morphology of Proconsul africanus, P. nyanzae, P. major and Sivapithecus meteai is compared to extant catarrhines. The early Miocene hominoids (Proconsul) are unlike modern great apes, but retain a primitive catarrhine pattern more similar to some extant cercopthecoids. By middle Miocene times the typical hominoid palatofacial morphology can be recognized in at least one species (S. meteai) and this corresponds to the evolution of the postcranium in which the hominoid pattern is also only recognizable by the middle Miocene.     

Metric variation and sexual dimorphism in the dentition of Ouranopithecus macedoniensis

The fossil sample attributed to the late Miocene hominoid taxon Ouranopithecus macedoniensis is characterized by a high degree of dental metric variation. As a result, some researchers support a multiple-species taxonomy for this sample. Other researchers do not think that the sample variation is too great to be accommodated within one species. This study examines variation and sexual dimorphism in mandibular canine and postcanine dental metrics of an Ouranopithecus sample. Bootstrapping (resampling with replacement) of extant hominoid dental metric data is performed to test the hypothesis that the coefficients of variation (CV) and the indices of sexual dimorphism (ISD) of the fossil sample are not significantly different from those of modern great apes. Variation and sexual dimorphism in Ouranopithecus M(1) dimensions were statistically different from those of all extant ape samples; however, most of the dental metrics of Ouranopithecus were neither more variable nor more sexually dimorphic than those of Gorilla and Pongo. Similarly high levels of mandibular molar variation are known to characterize other fossil hominoid species. The Ouranopithecus specimens are morphologically homogeneous and it is probable that all but one specimen included in this study are from a single population. It is unlikely that the sample includes specimens of two sympatric large-bodied hominoid species. For these reasons, a single-species hypothesis is not rejected for the Ouranopithecus macedoniensis material. Correlations between mandibular first molar tooth size dimorphism and body size dimorphism indicate that O. macedoniensis and other extinct hominoids were more sexually size dimorphic than any living great apes, which suggests that social behaviors and life history profiles of these species may have been different from those of living species.     

The relationships among jaw‐muscle fiber architecture,jaw morphology,and feeding behavior in extant apes and modern humans

The jaw‐closing muscles are responsible for generating many of the forces and movements associated with feeding. Muscle physiologic cross‐sectional area (PCSA) and fiber length are two architectural parameters that heavily influence muscle function. While there have been numerous comparative studies of hominoid and hominin craniodental and mandibular morphology, little is known about hominoid jaw‐muscle fiber architecture. We present novel data on masseter and temporalis internal muscle architecture for small‐ and large‐bodied hominoids. Hominoid scaling patterns are evaluated and compared with representative New‐ (Cebus) and Old‐World (Macaca) monkeys. Variation in hominoid jaw‐muscle fiber architecture is related to both absolute size and allometry. PCSAs scale close to isometry relative to jaw length in anthropoids, but likely with positive allometry in hominoids. Thus, large‐bodied apes may be capable of generating both absolutely and relatively greater muscle forces compared with smaller‐bodied apes and monkeys. Compared with extant apes, modern humans exhibit a reduction in masseter PCSA relative to condyle‐M₁ length but retain relatively long fibers, suggesting humans may have sacrificed relative masseter muscle force during chewing without appreciably altering muscle excursion/contraction velocity. Lastly, craniometric estimates of PCSAs underestimate hominoid masseter and temporalis PCSAs by more than 50% in gorillas, and overestimate masseter PCSA by as much as 30% in humans. These findings underscore the difficulty of accurately estimating jaw‐muscle fiber architecture from craniometric measures and suggest models of fossil hominin and hominoid bite forces will be improved by incorporating architectural data in estimating jaw‐muscle forces. Am J Phys Anthropol 151:120–134, 2013. © 2013 Wiley Periodicals, Inc.     

Evolution of the second orangutan: phylogeny and biogeography of hominid origins

Aim To resolve the phylogeny of humans and their fossil relatives (collectively, hominids), orangutans (Pongo) and various Miocene great apes and to present a biogeographical model for their differentiation in space and time. Location Africa, northern Mediterranean, Asia. Methods Maximum parsimony analysis was used to assess phylogenetic relationships among living large‐bodied hominoids (= humans, chimpanzees, bonobos, gorillas, orangutans), and various related African, Asian and European ape fossils. Biogeographical characteristics were analysed for vicariant replacement, main massings and nodes. A geomorphological correlation was identified for a clade we refer to as the ‘dental hominoids’, and this correlation was used to reconstruct their historical geography. Results Our analyses support the following hypotheses: (1) the living large‐bodied hominoids represent a monophyletic group comprising two sister clades: humans + orangutans, and chimpanzees (including bonobos) + gorillas (collectively, the African apes); and (2) the human–orangutan clade (dental hominoids) includes fossil hominids (Homo, australopiths, Orrorin) and the Miocene‐age apes Hispanopithecus, Ouranopithecus, Ankarapithecus, Sivapithecus, Lufengpithecus, Khoratpithecus and Gigantopithecus (also Plio‐Pleistocene of eastern Asia). We also demonstrate that the distributions of living and fossil genera are largely vicariant, with nodes of geographical overlap or proximity between Gigantopithecus and Sivapithecus in Central Asia, and between Pongo, Gigantopithecus, Lufengpithecus and Khoratpithecus in East Asia. The main massing is represented by five genera and eight species in East Asia. The dental hominoid track is spatially correlated with the East African Rift System (EARS) and the Tethys Orogenic Collage (TOC). Main conclusions Humans and orangutans share a common ancestor that excludes the extant African apes. Molecular analyses are compromised by phenetic procedures such as alignment and are probably based on primitive retentions. We infer that the human–orangutan common ancestor had established a widespread distribution by at least 13 Ma. Vicariant differentiation resulted in the ancestors of hominids in East Africa and various primarily Miocene apes distributed between Spain and Southeast Asia (and possibly also parts of East Africa). The geographical disjunction between early hominids and Asian Pongo is attributed to local extinctions between Europe and Central Asia. The EARS and TOC correlations suggest that these geomorphological features mediated establishment of the ancestral range.     

On the systematic status of the late Neogene hominoids from Yunnan Province,China

Late Miocene and Pliocene hominoids from Yunnan Province in southern China have been recovered from four sites or site complexes: Xiaolongtan, Yangyi, Shihuiba and Yuanmou. Of these, Shihuiba and Yuanmou are among the most prolific fossil hominoid sites in Eurasia, and they have yielded important evidence that is critical for documenting the evolutionary history, biogeography and paleobiology of later Neogene hominids. The aim of this paper is to clarify their taxonomy and nomenclature, and to present a preliminary synthesis of their phylogenetic relationships and biogeography. The morphological pattern and degree of variation observed in the fossil samples is consistent with there being a single, sexually dimorphic species represented at each site. Provisionally, we consider the Shihuiba, Xiaolongtan and Yuanmou samples to belong to two separate species within a single genus. The valid names for these species are Lufengpithecus lufengensis (from Shihuiba) and L. keiyuanensis (from Xiaolongtan and Yuanmou). From a phylogenetic perspective, the currently available evidence suggests that Lufengpithecus is either a primitive hominid that represents the sister taxon of the Ponginae+Homininae or a primitive sister taxon to the Ponginae. We tend to favor the second alternative, but acknowledge that a more comprehensive comparative analysis is needed to substantiate the phylogenetic and taxonomic affinities of Lufengpithecus. Importantly, the Yunnan fossil apes provide a unique temporal perspective on the evolutionary history of hominoids. Their continued occurrence during the late Miocene and Pliocene (approximately 8-2Ma), when hominoids became extinct throughout the rest of Eurasia, suggests that southern China (and presumably southeast Asia in general) was an important refugium for hominoids, including the ancestors of the orang-utans and gibbons. The uplift of the Tibetan plateau and its impact on regional climatic conditions may have been an important contributing factor in isolating the hominoids geographically and ecologically. We speculate that changed climatic condition in the mid-Pliocene, and possibly the arrival of Homo soon after, may have precipitated the regional extinction of large hominoids in southern China and in mainland southeast Asia.     

Hominoid dietary evolution.

During the later Palaeocene and early Miocene, catarrhine primates and the evolving hominoids had adaptations for frugivorous diets, with the emphasis on soft foods. Early in the middle Miocene the hominoids underwent a major shift, both in morphology and in habitat, with the morphology characterized by thickened enamel on the molars, enlarged incisors and massive jaws. The diet indicated by this morphology is interpreted as still mainly frugivorous but with changed emphasis, possibly towards harder objects. The thick-enamelled hominoids are found associated with more open forest habitats, and the distribution of food resources in equivalent habitats today is discontinuous both in time and in space, leading to evolutionary pressures particularly affecting locomotion, brain size and social behaviour. The earliest known hominid fossils differed little in dental and mandibular morphology from the middle Miocene apes, and the implied dietary similarity, together with ape-like patterns of dental development and retained arboreal adaptations of the postcrania, suggests little change in the foraging strategies of the earliest hominids compared with their ape ancestors and further suggests similarity in evolutionary grade. This similarity may have extended to other aspects of behaviour, for example to patterns of tool making and use, which may have been similar in the common ancestor of apes and humans to the pattern shared by the earliest australopithecines and chimpanzees.     

Knee function through finite element analysis and the role of Miocene hominoids in our understanding of the origin of antipronograde behaviours: the Pierolapithecus catalaunicus patella as a case study

Although extensive research has been carried out in recent years on the origin and evolution of human bipedalism, a full understanding of this question is far from settled. Miocene hominoids are key to a better understanding of the locomotor types observed in living apes and humans. Pierolapithecus catalaunicus, an extinct stem great ape from the middle Miocene (c. 12.0 Ma) of the Vallès-Penedès Basin (north-eastern Iberian Peninsula), is the first undoubted hominoid with an orthograde (erect) body plan. Its locomotor repertoire included above-branch quadrupedalism and other antipronograde behaviours. Elucidating the adaptive features present in the Pierolapithecus skeleton and its associated biomechanics helps us to better understand the origin of hominoid orthogrady. This work represents a new biomechanical perspective on Pierolapithecus locomotion, by studying its patella and comparing it with those drawn from a large sample of extant anthropoids. This is the first time that the biomechanical patellar performance in living non-human anthropoids and a stem hominid has been studied using finite element analysis (FEA). Differences in stress distribution are found depending on body plan and the presence/absence of a distal apex, probably due to dissimilar biomechanical performances. Pierolapithecus’ biomechanical response mainly resembles that of great apes, suggesting a similar knee joint use in mechanical terms. These results underpin previous studies on Pierolapithecus, favouring the idea that a relevant degree of some antipronograde behaviour may have made up part of its locomotor repertoire. Moreover, our results corroborate the presence of modern great ape-like knee biomechanical performances back in the Miocene.     

Molar crown formation in the Late Miocene Asian hominoids, Sivapithecus parvada and Sivapithecus indicus

During the past decade, studies of enamel development have provided a broad temporal and geographic perspective on evolutionary developmental biology in Miocene hominoids. Here we report some of the first data for molar crown development in one hominoid genus, Sivapithecus. The data are compared to a range of extant and extinct hominoids. Crown formation times (CFTs), daily rates of enamel secretion (DSR), Retzius line number and periodicity, and relative enamel thickness (RET) were calculated in a mandibular first molar of Sivapithecus parvada and a maxillary first molar of Sivapithecus indicus from the Siwalik sequence of Pakistan. A CFT of 2.40 years for the protoconid of S. parvada and 2.25 years for the protocone of S. indicus lie within the range of first molar (M1) formation times for the majority of Miocene hominoids (1.96-2.40 years, excluding Proconsul heseloni), and are similar to an M(1) from Gorilla (2.31 years) and M(1)s from Pan (2.22-2.39 years). This is unlike the longer CFTs in modern humans, which appear to be linked with their extended growth period. In contrast to extant great apes and humans, daily rates of enamel secretion are rapid in the Sivapithecus M1s during the early stages of growth, which seems to be a common pattern for most Miocene apes. The rapid accumulation of cuspal enamel in the Sivapithecus molars produced thicker enamel than either Pan or Gorilla in a comparable period of time. Future studies on larger samples of living and fossil hominoids are needed to clarify trends in crown development, which may be better understood in the context of life history strategies coupled with good data on body mass and brain size.     

Variations of the mandibular shape in extant hominoids: Generic, specific, and subspecific quantification using elliptical fourier analysis in lateral view

While a number of studies have documented the mandibular variations in hominoids, few focused on evaluating the variation of the whole outline of this structure. Using an efficient morphometrical approach, i.e. elliptical Fourier analysis, mandibular outlines in lateral view from 578 adult hominoids representing the genera Hylobates, Pongo, Gorilla, Pan, and Homo were quantified and compared. This study confirms that elliptical Fourier analysis provides an accurate characterization of the shape of the mandibular profile. Differences in mandibular shape between hominoid genera, species, subspecies, and to a lesser extent between sexes were demonstrated. Mandibles in great apes and hylobatids subspecies were generally less distinct from each other than were species. However, the magnitudes of differences among subspecies of Gorilla and Pongo approached or exceeded those between Pan troglodytes and P. paniscus. The powerful discrimination between taxa from the genus down to subspecific level associated to the relatively low level of intrageneric mandibular polymorphism in great apes provides strong evidences in support of the taxonomic utility of the shape of the mandibular profile in hominoids. In addition, morphological affinities between Pongo and Pan and the clear distinction between Homo and Pan suggest that the mandibular outline is a poor estimate of phylogenetic relationships in great apes and humans. The sexual dimorphism in mandibular shape exhibits two patterns of expression: a high degree of dimorphism in Gorilla, Pongo, and H. s. syndactylus and a relatively low one in modern humans and Pan. Besides, degree of mandibular shape dimorphism can vary considerably among closely related subspecies as observed in gorillas, arguing against the use of mandibular shape dimorphism patterns as characters in phylogenetic analyses. However, the quantification of the mandibular shape and of the variations among hominoids provides an interesting comparative framework that is likely to supply further arguments for a better understanding of the patterns of differentiation between living hominoids.     

Hominoid dental variability and species number at the late Miocene site of Lufeng,China

The large hominoid sample from the late Miocene site of Lufeng, China, has been variously claimed to contain either one or two species, but very few metric data in support of either position have been published. We calculate coefficients of variation for the dental remains both for the two presumed species and for the pooled sample as a whole using the summary statistics published by Wu & Oxnard (Wu & Oxnard: American Journal of Primatology 5:303–344, 1983a, Nature 306:258–260, 1983b). These are compared to the same measures of single-sex and combined-sex samples of extant hominoids. We also present metric characterizations of male and female canines of extant great apes, with which we evaluate the gender composition of the Lufeng canine sample. In a two-species alternative, the two presumed species have measures of variability and canine representation that are more compatible with single-sex samples representing males and females, respectively. The pooled dental sample has measures of variability within the ranges of single species of extant great apes. We conclude there is a single large hominoid species represented at Lufeng that is highly sexually dimorphic. The phylogenetic relationships of this species are briefly considered. It is generally primitive in craniodental morhpology and is unlikely to be a member of the Sivapithecus-Pongo clade.     

Pierolapithecus and the functional morphology of Miocene ape hand phalanges: paleobiological and evolutionary implications

The partial skeleton of Pierolapithecus, which provides the oldest unequivocal evidence of orthogrady, together with the recently described phalanges from Pa?alar most likely attributable to Griphopithecus, provide a unique opportunity for understanding the changes in hand anatomy during the pronogrady/orthogrady transition in hominoid evolution. In this paper, we describe the Pierolapithecus hand phalanges and compare their morphology and proportions with those of other Miocene apes in order to make paleobiological inferences about locomotor evolution. In particular, we investigate the orthograde/pronograde evolutionary transition in order to test whether the acquisition of vertical climbing and suspension were decoupled during evolution. Our results indicate that the manual phalanges of Miocene apes are much more similar to one another than to living apes. In particular, Miocene apes retain primitive features related to powerful-grasping palmigrady on the basal portion, the shaft, and the trochlea of the proximal phalanges. These features suggest that above-branch quadrupedalism, inherited from stem hominoids, constituted a significant component of the locomotor repertories of different hominoid lineages at least until the late Miocene. Nonetheless, despite their striking morphological similarities, several Miocene apes do significantly differ in phalangeal curvature and/or elongation. Hispanopithecus most clearly departs by displaying markedly-curved and elongated phalanges, similar to those in the most suspensory of the extant apes (hylobatids and orangutans). This feature agrees with several others that indicate orang-like suspensory capabilities. The remaining Miocene apes, on the contrary, display low to moderate phalangeal curvature, and short to moderately-elongated phalanges, which are indicative of the lack of suspensory adaptations. As such, the transition from a pronograde towards an orthograde body plan, as far as this particular anatomical region is concerned, is reflected only in somewhat more elongated phalanges, which may be functionally related to enhanced vertical-climbing capabilities. Our results therefore agree with the view that hominoid locomotor evolution largely took place in a mosaic fashion: just as taillessness antedated the acquisition of an orthograde body plan, the emergence of the latter—being apparently related only to vertical climbing—also preceded the acquisition of suspensory adaptations, as well as the loss of primitively-retained, palmigrady-related features.     

Major features in the evolution of early hominoid locomotion

Evolution of hominoid locomotion is a traditional topic in primate evolution. Views have changed during the last decade because a number of crucial differences between early and advanced hominoid morphologies have been demonstrated. Increasing evidence on primate behaviour and ecology show that any direct analogies between living and fossil hominoids must be made extremely carefully. The necessity of synthesizing data on primate behaviour, locomotion, morphology and ecology and simultaneously defining the framework in which the data should be interpreted are explained. Results of our studies of ontogeny of locomotor and behavioural patterns (LBP) are presented that could help identify the main features of early hominoid locomotor patterns (LP) and the mechanisms of their changes. The early hominoid LP was different from those of pronograde monkeys and specialized antipronograde living apes. Some similar features could be expected between early hominoid LP and the LP of ceboid monkeys. Analogous mechanisms of change of LBP exist in all groups of living higher primates. Crucial early mechanisms of change are the ontogenetic shifts in LBP connected with ethoecological changes. Analysis of fossil evidence has shown that Miocene hominoids differ morphologically from any group of living primates. Certain features present in Miocene hominoids could be found in Atelinae and living Asian apes but they are limited to some functional regions of the postcrania only. Consequently the early hominoid general LP can not be strictly analogous either to that of any monkey group or to the LP of apes. We suppose that certain pronograde adaptations, such as climbing, bipedality, limited suspensory activity and sitting constituted the main part of their LP.