Micropithecus clarki, a small ape from the Miocene of Uganda.

Micropithecus clarki, from Miocene sediments of Napak, Uganda, is the smallest known hominoid primate, living or fossil. In facial morphology it is very similar to extant gibbons. Dentally, it is most similar to the small apes from the Miocene of Kenya, Dendropithecus and Limnopithecus. All of the apes from the early Miocene of East Africa seem to represent a single phyletic group that could be easily derived from the Oligocene apes known from the Fayum of Egypt. Pliopithecus from the Miocene of Europe is more closely allied with the Oligocene radiation than with the later East African radiation.     

Similarities in Aegyptopithecus and Afropithecus facial morphology

Recently discovered cranial fossils from the Oligocene deposits of the Fayum depression in Egypt provide many details of the facial morphology of Aegyptopithecus zeuxis. Similar features are found in the Miocene hominoid Afropithecus turkanensis. Their presence is the first good evidence of a strong phenetic link between the Oligocene and Miocene hominoids of Africa. A comparison of trait lists emphasizes the similarities of the two fossil species, and leads us to conclude that the two fossil genera share many primitive facial features. In addition, we studied facial morphology using finite-element scaling analysis and found that the two genera show similarities in morphological integration, or the way in which biological landmarks relate to one another in three dimensions to define the form of the organism. Size differences between the two genera are much greater than the relatively minor shape differences. Analysis of variability in landmark location among the four Aegyptopithecus specimens indicates that variability within the sample is not different from that found within two samples of modern macaques. We propose that the shape differences found among the four Aegyptopithecus specimens simply reflect individual variation in facial characteristics, and that the similarities in facial morphology between Aegyptopithecus and Afropithecus probably represent a complex of primitive facial features retained over millions of years.     

On aspects of skull form in African apes and orangutans, with implications for hominoid evolution

The study of hominoid phylogeny is currently in a state of controversy and debate due to the discovery of new fossil material and reanalysis of the morphology of extant apes. An important key to the resolution of these debates lies in attaining a fuller understanding of the morphological differences in skull form between the African and Asian great apes. In this paper I have analyzed aspects of facial morphology and internal cranial anatomy in the great apes. Results from this study and previous ones suggest that Pongo is characterized by a marked dorsal deflection of the face relative to the basicranium. Many aspects of circumorbital, midfacial, palatal, and mandibular morphology in Pongo may be related to this airorynchous condition. This hypothesis is supported by Enlow's work on form and pattern in the primate and mammalian skull. The position of the face in known Sivapithecus appears to be similar to that seen in Pongo. Although Pongo may be specialized in its marked degree of airorynchy, it seems likely that an important derived feature linking African apes and hominids is a ventral rotation of the splanchnocranium on the neurocranium. The appearance of marked supraorbital tori and ethmofrontal sinuses are probably correlated developments. Additional implications of this work for debates about hominoid phylogeny are discussed.     

元谋小河-竹棚与雷老地点古猿牙齿特征的对比分析

针对学术界持有不同看法的在元谋盆地发现的古猿化石是否代表着两个种的争议 ,本文把迄今在元谋发现的全部古猿牙齿化石按地点分为小河 -竹棚和雷老两组 ,从牙齿形态特征、测量数据分析及犬齿形态指数三方面对出自元谋不同地点的古猿在牙齿特征上的相似及差异程度进行了对比。结果显示在本文所对比的牙齿特征中 ,不同地点的古猿牙齿在形态特征、牙齿大小及犬齿形态比例的表现非常相似 ,未发现它们彼此之间在这些特征上存在任何明显的差别。作者认为本研究不支持在元谋不同地点发现的古猿之间存在不同种类的观点。     

Nacholapithecus and its importance for understanding hominoid evolution

Nacholapithecus kerioi is a large‐sized hominoid from the Aka Aiteputh Formation (15 Ma) in Nachola, northern Kenya. 1 While eight large‐sized hominoid species dating to the late Early to early Middle Miocene (17‐14 Ma) are known in Afro‐Arabia and western Eurasia, 2 - 6 the facial and postcranial anatomy of these apes is poorly known. However, much has been learned of the craniodental and postcranial anatomy of N. kerioi over the last ten years (A list of published specimens is available online, accompanying this article), and it plays a key role in our understanding of hominoid evolution in the Early to Middle Miocene of Africa and Eurasia. Importantly, it bears on the interpretation of the hominoid Morotopithecus bishopi from 20.6 my‐old Uganda. 7 - 10 In the article, we provide information on the anatomy and adaptations of N. kerioi as well as on the site of Nachola, and discuss how our current knowledge of N. kerioi can be incorporated into scenarios of hominoid evolution.     

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.     

Anatomy of the hominoid wrist joint: Its evolutionary and functional implications

Observations on the behavior of living hominoids show generic differences in the use and posture of the wrist joint. Both orang-utans and hylobatids usually use the wrist in suspensory behaviors. However, orang-utans emphasize markedly adducted and flexed wrist postures, while hylobatids emphasize violent forearm and wrist rotation. African apes, especially the gorilla, use the wrist more frequently than other hominoids for terrestrial quadrupedal weight-bearing. Humans use the wrist less frequently for supportive purposes than do other hominoids. These behavioral differences correspond to structural specializations in the proximal carpal joint of each of the hominoid genera. Although each of the hominoid genera has apparently modified its proximal carpal joint best to serve its characteristic behaviors, all hominoids share a unique proximal carpal joint that permits approximately 160ℴ of forearm rotation. The hylobatid proximal carpal joint is specialized in exhibiting a marked development of those structures limiting forearm rotation, but it is in most respects the least derived— that is, closest to the nonhominoid anthropoids. Chimpanzees show a proximal carpal joint that is more generalized than those of the other great apes but more derived than that of hylobatids. The human and gorilla proximal wrist joints, on the other hand, show marked modifications for weight-bearing in terrestrial behaviors. Orang-utans have the most derived proximal carpal joint, which in many respects parallels that of the slow-climbing nonhominoid primates. The comparative anatomy and structural specializations of the wrist joint support (a) an early divergence of hylobatids from the common hominoid stock, (b) a common ancestry for gorillas and humans separate from the other hominoids, and (c) a long independent evolutionary period for orang-utans since their divergence from the common hominoid stock, or one that was marked by strong selection pressures for wrist specializations. Unfortunately, the generalized condition of the chimpanzee’s wrist joint and the very derived condition of the orang-utan wrist provide uncertain evidence as to which of the two was first to diverge from the common hominoid stock. Identification of hominoid wrist specializations as reflecting real phylogenetic relationships or parallelisms depends on how well the phytogeny inferred from wrist morphology accords with those arrived at from the study of other systems.     

Hominoid phalanges from the middle Miocene site of Paşalar, Turkey

Eleven proximal and ten intermediate partial or complete hominoid phalanges have been recovered from the middle Miocene site of Pa?alar in Turkey. Based on species representation at Pa?alar, it is likely that most or all of the phalanges belong to Griphopithecus alpani rather than Kenyapithecus kizili, but both species may be represented. All of the complete or nearly complete phalanges appear to be manual, so comparisons to extant and other fossil primate species were limited to manual phalanges. Comparisons were made to extant hominoid and cercopithecoid primate genera expressing a variety of positional repertoires and varying degrees of arboreality and terrestriality. The comparisons consisted of a series of bivariate indices derived from previous publications on Miocene catarrhine phalangeal morphology. The proximal phalanges have dorsally expanded proximal articular surfaces, which is characteristic of cercopithecoids and most other Miocene hominoids, and indicates that the predominant positional behaviors involved pronograde quadrupedalism. Among the extant primates, many of the proximal and intermediate phalangeal indices clearly distinguish more habitually terrestrial taxa from those that are predominantly arboreal, and especially from taxa that commonly engage in suspensory activities. For nearly every index, the values of the Pa?alar phalanges occupy an intermediate position-most similar to values for Pan and, to a lesser extent, Macaca-indicating a generalized morphology and probably the use of both arboreal and terrestrial substrates. At least some terrestrial activity is also compatible with reconstructions of the Pa?alar habitat. Most proximal and intermediate phalanges of other middle and late Miocene hominoids have similar index values to those of the Pa?alar specimens, revealing broadly similar manual phalangeal morphology among many Miocene hominoids.     

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.     

Dental remains of Equatorius africanus from Kipsaramon, Tugen Hills, Baringo District, Kenya.

Forty-one isolated large hominoid teeth, as well as most of the mandibular and three maxillary teeth associated with a partial skeleton, were recovered from middle Miocene Muruyur sediments near Kipsaramon in the Tugen Hills, Baringo District, Kenya. The isolated teeth were collected as surface finds and the skeleton was excavated in situ at locality BPRP#122 dated between 15.58 Ma and 15.36 Ma. The majority of the teeth recovered at BPRP#122 are referable to a minimum of five individuals of the hominoid Equatorius africanus. Three of the teeth, however, are provisionally assigned to Nyanzapithecus sp. The new hominoids from Kipsaramon add to an increasing inventory of specimens that suggest greater large hominoid taxonomic diversity from the middle Miocene of Kenya than was previously recognized. It is suggested that there are two large-bodied hominoid species present at Mabako, only one of which is assignable to Equatorius.     

DA/DAPI fluorescent bands in the chromosomes of Pan paniscus

The fluorochrome pattern produced by DA/DAPI double staining in Pan paniscus chromosomes is reported. The location of DA/DAPI prominent bands differs from that reported for all other hominoid species. However, the pattern in the pygmy chimpanzee is most similar to that seen in Pan troglodytes. Comparison of the DA/DAPI pattern of the other hominoid species allows the construction of a proposed hominoid ancestral karyotype and a preliminary phylogenetic reconstruction of DA/DAPI bands for the great apes and man.     

Three-dimensional analysis of mandibular dental root morphology in hominoids

Although often preserved in the fossil record, mandibular dental roots are rarely used for evolutionary studies. This study qualitatively and quantitatively characterizes the three-dimensional morphology of hominoid dental roots. The sample comprises extant apes as well as two fossil species, Khoratpithecus piriyai and Ouranopithecus macedoniensis. The morphological differences between extant genera are observed, quantified and tested for their potential in systematics. Dental roots are imaged using X-ray computerized tomography, conventional microtomography and synchrotron microtomography. Resulting data attest to the high association between taxonomy and tooth root morphology, both qualitatively and quantitatively. A cladistic analysis based on the dental root characters resulted in a tree topology congruent with the consensus phylogeny of hominoids, suggesting that tooth roots might provide useful information in reconstructing hominoid phylogeny. Finally, the evolution of the dental root morphology in apes is discussed.     

Cranio-dental evidence of a hominin-like hyper-masticatory apparatus in Oreopithecus bambolii. Was the swamp ape a human ancestor?

The phylogeny of Oreopithecus bambolii has been controversial since Johannes Hurzeler first argued that the Late Miocene (Tortonian) primate was a fossil hominin. While most paleontologist currently exclude Oreopithecus from human ancestry, recent postcranial evidence of hominin-like bipedalism and power precision grips in Oreopithecus has rekindled interest in the fossil Italian hominoid.In this study, a comparative review of hominoid cranio-dental morphology indicates that Oreopithecus possessed a suite of hominin-like characteristics that were apparently functionally related to powerful folivorous mastication. Since the oreopithecine dentition exhibited exceptional adaptations for folivory relative to most other extant and extinct hominoids, the accessory development of a hominin-like hyper-masticatory cranio-mandibular apparatus to further enhance plant comminution and digestibility is not surprising. However, the combination of hominin-like locomotive, manual, and masticatory functional attributes appears to substantiate Hurzeler's designation of O. bambolii as a Late Miocene hominin. Additionally, an extensive compilation of hominoid cranio-dental and postcranial characteristics strongly supports a close phylogenetic relationship between Oreopithecus and the earliest African hominins Sahelanthropus and Australopithecus.The wetland paleoecology of the Tortonian island of Tuscany–Sardinia suggests that Oreopithecus was a specialized semiaquatic folivore who apparently waded bipedally into freshwater swamps to feed on aquatic plants. However, the extensive wear on the oreopithecine canines and incisors along with their manual precision grips may indicate that freshwater invertebrates were also included in their diets. Such specialized aquatic feeding behavior by these ancient Italian swamp apes seems to support Alister Hardy's hypothesis that human bipedalism and power precision grips were inherited from Late Miocene hominin ancestors who originally utilized such functional attributes for aquatic foraging in shallow water environments. Additionally, the remarkably hominin-like cranio-dental morphology of O. bambolii suggests that modern omnivorous humans probably inherited a significant number of their cranio-dental characteristics from these highly specialized aquatic plant eating hominins.     

Morphometric analysis of hominoid lower molars from Yuanmou of Yunnan Province, China

Shape analyses of cross-sectional mandibular molar morphology, using Euclidean Distance Matrix Analysis, were performed on 79 late Miocene hominoid lower molars from Yuanmou of Yunnan Province, China. These molars were compared to samples of chimpanzee, gorilla, orangutan,Lufengpithecus lufengensis, Sivapithecus, Australopithecus afarensis, and human mandibular molars. Our results indicate that the cross-sectional shape of Yuanmou hominoid lower molars is more similar to the great apes that to humans. There are few differences between the Yuanmou,L. lufengensis, andSivapithecus molars in cross-sectional morphology, demonstrating strong affinities between these three late Miocene hominoids. All three of the fossil samples show strong similarities to orangutans. From this, we conclude that these late Miocene hominoids are more closely related to orangutants than to either the African great apes or humans.     

Development and evolution of the helicoidal plane of dental occlusion

The helicoidal plane of dental occlusion is a composite feature involving axial inclination of teeth and effects of dental attrition. Recent studies disagree on its distribution and significance in hominoid primates. The distribution, development, and functional basis of the helicoidal plane are investigated here, based on quantitative analysis of dental morphology and attrition in 667 human and 60 chimpanzee dentitions. Helicoidal planes are nearly universal in the human and chimpanzee dentitions studied. Increasing axial inclination of molars from M1 to M3 is primarily responsible for the helicoidal plane, although attrition acts to increase its expression. In hominoids, increased molar axial tilt appears to be associated with facial shortening and dental reduction. Population and species comparisons suggest a functional relationship with cranial structure. Progressive axial tilt of molars producing a helicoidal plane is found consistently in mammals with cheek teeth positioned partly under the cranium, as in hominids, pongids, some cebids, macropodids, ursids, and sciurids. Facial shortening is an important trend in hominid evolution and axial inclination of molars might be expected to show progressive change from Australopithecus afarensis to recent Homo sapiens.     

A new hominoid species from the middle Miocene site of Paşalar, Turkey

A new species of fossil hominoid is described from the middle Miocene deposits at Pa?alar, Turkey. It is the less common of the two Pa?alar species discussed by Martin and Andrews (1993), making up approximately 10% of the individuals in the Pa?alar hominoid sample according to analyses of the minimum number of individuals. To the diagnostic features of I(1) described by Alpagut et al. (1990) and Martin and Andrews (1993) can now be added further diagnostic features of all the anterior teeth, as well as both upper premolars and P(3). These include discrete, nonmetric features and metric differences at all the noted tooth positions. Attempts to distinguish the upper and lower molars of the two species have so far been unsuccessful, with the possible exception of M(3). The morphology of the new species is similar in most respects to that of Kenyapithecus wickeri from Fort Ternan, especially concerning maxillary morphology. They share robust and moderately deep maxillary alveolar processes, a restricted maxillary sinus with an elevated and uncomplicated floor, lacking the compartmentalization evident to varying degrees in many other taxa, and a zygomatic process that originates and turns laterally fairly high above the alveolar margin. There are also a number of distinctive similarities in the dentition, particularly for I(1), C(1), P(4) and P(3). The I(1) morphology in particular, with greatly hypertrophied lingual marginal ridges bounding a uniformly thickened basal crown area, is distinctive among Miocene hominoids. All of these similarities serve to reinforce the differences noted by others between the derived morphology of K. wickeri and the more primitive morphology of Equatorius africanus from Maboko and Kipsaramon. The new species differs from K. wickeri in morphological details of most of the anterior and premolar teeth that are known for both species, despite the general morphological similarity, and in the size of I(1) versus I(2). One striking feature of the new species is a relatively large incisive fossa, although it cannot be determined if this is associated with an open palatine fenestra, as in many early Miocene hominoids, or a minimally overlapping palate and nasoalveolar clivus, as in some middle and late Miocene hominoids.     

A Genome-Wide Association Study Identifies Five Loci Influencing Facial Morphology in Europeans

Inter-individual variation in facial shape is one of the most noticeable phenotypes in humans, and it is clearly under genetic regulation; however, almost nothing is known about the genetic basis of normal human facial morphology. We therefore conducted a genome-wide association study for facial shape phenotypes in multiple discovery and replication cohorts, considering almost ten thousand individuals of European descent from several countries. Phenotyping of facial shape features was based on landmark data obtained from three-dimensional head magnetic resonance images (MRIs) and two-dimensional portrait images. We identified five independent genetic loci associated with different facial phenotypes, suggesting the involvement of five candidate genes—PRDM16, PAX3, TP63, C5orf50, and COL17A1—in the determination of the human face. Three of them have been implicated previously in vertebrate craniofacial development and disease, and the remaining two genes potentially represent novel players in the molecular networks governing facial development. Our finding at PAX3 influencing the position of the nasion replicates a recent GWAS of facial features. In addition to the reported GWA findings, we established links between common DNA variants previously associated with NSCL/P at 2p21, 8q24, 13q31, and 17q22 and normal facial-shape variations based on a candidate gene approach. Overall our study implies that DNA variants in genes essential for craniofacial development contribute with relatively small effect size to the spectrum of normal variation in human facial morphology. This observation has important consequences for future studies aiming to identify more genes involved in the human facial morphology, as well as for potential applications of DNA prediction of facial shape such as in future forensic applications.     

The phylogenetic affinities of Otavipithecus namibiensis

The middle Miocene hominoid Otavipithecus namibiensis is the first and most complete fossil ape from sub-equatorial Africa and represents a significant addition to the taxonomically sparse African middle Miocene hominoid fossil record. The Otavipithecus hypodigm comprises the holotype mandible, which presents a unique mosaic of dental and gnathic characters, and several attributed cranial and postcranial elements which resemble the stem hominoid Proconsul. Contrary to initial hopes that this discovery would provide new insights into hominoid morphological diversity and phylogenetic relationships, a variety of conflicting phylogenetic hypotheses have been advanced suggesting ties to virtually every major large-bodied hominoid group (Conroy et al., 1992; Andrews, 1992 a; Conroy, 1994; Pickford et al., 1994; Begun, 1994 a). Cladistic analysis of a matrix of 22 qualitative and ten quantitative characters of the mandible and mandibular dentition found no support for a close phylogenetic relationship between Otavipithecus and either the African ape or great ape clades, or with any of the Eurasian fossil hominoids with which it has previously been compared. A close relationship between Otavipithecus and Kenyapithecus cannot be ruled out, but is deemed unlikely on the basis both of morphological comparisons and the absence of support within a cladistic framework. The present analysis indicates that Otavipithecus is most closely related to Afropithecus, as previously suggested by Andrews (1992 a) among others. Due to lack of statistical support for this result, a conservative interpretation, that these taxa represented related but divergent lineages of a late early Miocene hominoid radiation, is currently favored. Findings are consistent with the allocation of Otavipithecus to Andrews' (1992 a) tribe Afropithecini which represents the sister group to Kenyapithecus and the extant ape clade.     

The relative congruence of cranial and genetic estimates of hominoid taxon relationships: implications for the reconstruction of hominin phylogeny

Previous analyses of extant catarrhine craniodental morphology have often failed to recover their molecular relationships, casting doubt on the accuracy of hominin phylogenies based on anatomical data. However, on the basis of genetic, morphometric and environmental affinity patterns, a growing body of literature has demonstrated that particular aspects of cranial morphology are remarkably reliable proxies for neutral modern human population history. Hence, it is important to test whether these intra-specific patterns can be extrapolated to a broader primate taxon level such that inference rules for understanding the morphological evolution of the extinct hominins may be devised. Here, we use a matrix of molecular distances between 15 hominoid taxa to test the genetic congruence of 14 craniomandibular regions, defined and morphometrically delineated on the basis of previous modern human analyses. This methodology allowed us to test directly whether the cranial regions found to be reliable indicators of population history were also more reliable proxies for hominoid genetic relationships. Cranial regions were defined on the basis of three criteria: developmental-functional units, individual bones, and regions differentially affected by masticatory stress. The results found that all regions tested were significantly and strongly correlated with the molecular matrix. However, the modern human predictions regarding the relative congruence of particular regions did not hold true, as the face was statistically the most reliable indicator of hominoid genetic distances, as opposed to the vault or basicranium. Moreover, when modern humans were removed from the analysis, all cranial regions improved in their genetic congruence, suggesting that it is the inclusion of morphologically-derived humans that has the largest effect on incongruence between morphological and molecular estimates of hominoid relationships. Therefore, it may be necessary to focus on smaller intra-generic taxonomic levels to more fully understand the effects of neutral and selective evolutionary processes in generating morphological diversity patterns.