Late Miocene hominoid from Niger

African Late Miocene hominoids are rare, having been reported from six localities in Kenya, Ethiopia and Chad ranging in age from 10.5 to 5.5 Ma. We here report the occurrence of a hominoid in Niger associated with a vertebrate fauna which indicates an age of between 11–5 Ma. The Niger fossil locality is 940 km north of the nearest known extant hominoids, 1000 km west of the nearest recorded fossil hominoid from Chad.     

Mechanisms of hominoid dispersal in Miocene East Africa

The thesis of this essay is that the ecological changes which occurred during the Miocene in East Africa, created both the need and the conditions for the development of dispersal behaviour and other adaptations. The major ecological change in East Africa during the Miocene was the break-up of the relatively homogenous, lowland, tropical rainforest. The forest break-up was due to the combined effects of climatic change and rifting. The result of the fragmentation for species adapted to the forest, was increasing resource limitation. Species responded either by retreating with the forests, becoming extinct, or by evolving or developing new behaviours to cope with the new environment. The focus of this paper is upon the development of dispersal as a behaviour pattern for locating patchy resources. An ecological model is advanced to explain the development of dispersal and to place it in the context of other competing evolutionary responses to the ecological changes of the Miocene. The anatomical evolution of the Old World monkeys and apes during the Miocene is looked at in terms of the model. Bovid evolution over the same period is also looked at as evidence of a broad, episodic turnover in response to the ecological changes. It is suggested that hominid bipedalism may also fit the model and may have evolved as an adaptation for dispersion.     

Functional morphology and anatomy of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene hominoid from Kenya

This paper describes the morphology of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene primate species excavated from Nachola, Kenya in 1999-2002. The cervical vertebrae in Nacholapithecus are larger than those of Papio cynocephalus. They are more robust relative to more caudal vertebral bones. Since Nacholapithecus had large forelimbs, it is assumed that strong cervical vertebrae would have been required to resist muscle reaction forces during locomotion. On the other hand, the vertebral foramen of the lower cervical vertebrae in Nacholapithecus is almost the same size as or smaller than that of P. cynocephalus. Atlas specimens of Nacholapithecus resemble those of extant great apes with regard to the superior articular facet, and they have an anterior tubercle trait intermediate between that of extant apes and other primate species. Nacholapithecus has a relatively short and thick dens on the axis, similar to those of extant great apes and the axis body shape is intermediate between that of extant apes and other primates. Moreover, an intermediate trait between extant great apes and other primate species has been indicated with regard to the angle between the prezygapophyseal articular facets of the axis in Nacholapithecus. Although the atlas of Nacholapithecus is inferred as having a primitive morphology (i.e., possessing a lateral bridge), the shape of the atlas and axis leads to speculation that locomotion or posture in Nacholapithecus involved more orthograde behavior similar to that of extant apes, and, in so far as cervical vertebral morphology is concerned, it is thought that Nacholapithecus was incipiently specialized toward the characteristics of extant hominoids.     

A hominoid distal tibia from the Miocene of Pakistan

A distal tibia, YGSP 1656, from the early Late Miocene portion of the Chinji Formation in Pakistan is described. The fossil is 11.4 million years old and is one of only six postcranial elements now assigned to Sivapithecus indicus. Aspects of the articular surface are cercopithecoid-like, suggesting some pronograde locomotor activities. However, YGSP 1656 possesses an anteroposteriorly compressed metaphysis and a mediolaterally thick medial malleolus, ape-like features functionally related to orthograde body postures and vertical climbing. YGSP 1656 lacks specializations found in the ankle of terrestrial cercopithecoids and thus Sivapithecus may have been primarily arboreal. Nevertheless, the morphology of this tibia is unique, consistent with other interpretations of Sivapithecus postcranial functional morphology that suggest the locomotion of this ape lacks a modern analog. Based on the limited postcranial remains from S. indicus, we hypothesize that this taxon exhibited substantial body size dimorphism.     

Miocene hominoid biogeography: pulses of dispersal and differentiation

Aim  To test the hypothesis that the ancestor of the hominines (African apes and humans) had an African origin by comparing the historical biogeographical patterns of hominoids with those of two other large land mammal clades, namely the hyaenids and proboscideans.
Location  Global, primarily the Old World over the last 25 Myr (Miocene to present).
Methods  Creation of a general area cladogram using pact , a new method for generating area cladograms, and interpretation of general and clade-specific speciation events involving hominoids, proboscideans and hyaenids.
Results  The analysis of the areas using pact reveals both general patterns and clade-specific exceptions to these patterns. All three groups share a general episode of species formation in Africa in the early Miocene, followed by 'out of Africa' expansion into Europe, Asia and North America, and a second general episode of species formation in Asia in the mid-Miocene, followed by 'out of Asia' expansion into Africa, Europe and North America. Finally, there were two additional 'out of Africa' events during the late Miocene and into the Pliocene, the last one setting the stage for the emergence and spread of Homo . In addition to these shared episodes of vicariance and dispersal, each group exhibits clade-specific within-area and peripatric speciation events.
Main conclusions  The complex history of dispersal and speciation over large areas exhibited by hominoids is part of a more general history of biotic diversification by taxon pulses. Refining this scenario will require the integration of additional clades from the same areas and times, as well as more detailed palaeoclimatological, palaeoenvironmental and geological evidence.     

Khoratpithecus piriyai, a Late Miocene hominoid of Thailand

A Khoratpithecus piriyai lower jaw corresponds to a well-preserved Late Miocene hominoid fossil from northeastern Thailand. Its morphology and internal structure, using a microcomputed tomography scan, are described and compared to those of other known Miocene hominoids. It originated from fluviatile sand and gravel deposits of a large river, and was associated with many fossil tree trunks, wood fragments, and large vertebrate remains. A biochronological analysis by using associated mammal fauna gives an estimated geological age between 9-6 Ma. The flora indicates the occurrence of a riverine tropical forest and wide areas of grassland. K. piriyai displays many original characters, such as the great breadth of its anterior dentition, suggesting large incisors, large lower M3, a canine with a flat lingual wall, and symphysis structure. Several of its morphological derived characters are shared with the orangutan, indicating sister-group relationship with that extant ape. This relationship is additionally strongly supported by the absence of anterior digastric muscle scars. These shared derived characters are not present in Sivapithecus, Ankarapithecus, and Lufengpithecus, which are therefore considered more distant relatives to the orangutan than Khoratpithecus. The Middle Miocene K. chiangmuanensis is older, displays more primitive dental characters, and shares several dental characters with the Late Miocene form. It is therefore interpreted as its probable ancestor. But its less enlarged M3 and more wrinkled enamel may suggest an even closer phylogenetic position to orangutan ancestors, which cannot yet be supported because of the incomplete fossil record. Thus Khoratpithecus represents a new lineage of Southeast Asian hominoids, closely related to extant great ape ancestors.     

Comparative and functional morphology of hominoid fingers.

Comparisons of hominoid metacarpals and phalanges reveal differences, many of which are closely linked to locomotor hand postures. The African apes display features of the metacarpals and phalanges which distinguish them from the other Hominoidea. These features are most evident in digits III and IV. The orangutan hand is demonstrably less well adapted to knuckle-walking and is distinctive in its adaptation to power and hook grasping of vertical and horizontal supports, respectively. Orangutan fingers possess a "double-locking" mechanism (Napier, '60), and a slight ulnad shift in the axis of the hand which results in lengthened phalanges of ray IV. Hylobatid apes are more like orangutans in their finger morphology than any of the other Hominoidea, but exhibit unique features of their own. These include elongate phalanges of fingers II-V. Human metacarpals II-V form two sets composed of II-III, and IV-V. The heads of both metacarpals II and III are characterized by axial torsion. This reflects the enhanced manipulatory role of the third finger in humans. Human distal phalanges are unique in the development of pronounced apical tufts. Multivariate analysis of metacarpal III and proximal III yields variables that array the extant apes along an arboreal-terrestrial axis, from hylobatid apes to male gorillas. The positions of taxa on this discriminant concur with observations on the locomotion of free-ranging apes.     

A methodological investigation of hominoid craniodental morphology and phylogenetics

The evolutionary relationships of extant great apes and humans have been largely resolved by molecular studies, yet morphology-based phylogenetic analyses continue to provide conflicting results. In order to further investigate this discrepancy we present bootstrap clade support of morphological data based on two quantitative datasets, one dataset consisting of linear measurements of the whole skull from 5 hominoid genera and the second dataset consisting of 3D landmark data from the temporal bone of 5 hominoid genera, including 11 sub-species. Using similar protocols for both datasets, we were able to 1) compare distance-based phylogenetic methods to cladistic parsimony of quantitative data converted into discrete character states, 2) vary outgroup choice to observe its effect on phylogenetic inference, and 3) analyse male and female data separately to observe the effect of sexual dimorphism on phylogenies. Phylogenetic analysis was sensitive to methodological decisions, particularly outgroup selection, where designation of Pongo as an outgroup and removal of Hylobates resulted in greater congruence with the proposed molecular phylogeny. The performance of distance-based methods also justifies their use in phylogenetic analysis of morphological data. It is clear from our analyses that hominoid phylogenetics ought not to be used as an example of conflict between the morphological and molecular, but as an example of how outgroup and methodological choices can affect the outcome of phylogenetic analysis.     

First hominoid from the Late Miocene of the Irrawaddy Formation (Myanmar)

For over a century, a Neogene fossil mammal fauna has been known in the Irrawaddy Formation in central Myanmar. Unfortunately, the lack of accurately located fossiliferous sites and the absence of hominoid fossils have impeded paleontological studies. Here we describe the first hominoid found in Myanmar together with a Hipparion (s.l.) associated mammal fauna from Irrawaddy Formation deposits dated between 10.4 and 8.8 Ma by biochronology and magnetostratigraphy. This hominoid documents a new species of Khoratpithecus, increasing thereby the Miocene diversity of southern Asian hominoids. The composition of the associated fauna as well as stable isotope data on Hipparion (s.l.) indicate that it inhabited an evergreen forest in a C3-plant environment. Our results enlighten that late Miocene hominoids were more regionally diversified than other large mammals, pointing towards regionally-bounded evolution of the representatives of this group in Southeast Asia. The Irrawaddy Formation, with its extensive outcrops and long temporal range, has a great potential for improving our knowledge of hominoid evolution in Asia.     

An additional specimen of a large-bodied Miocene hominoid from Chiang Muan, northern Thailand

Chiang Muan is the first Miocene fossil site in Southeast Asia, from which large-bodied Miocene hominoids have been discovered (Kunimatsu et al., Primate Res 16:299, 2000a). In this article, we describe a hominoid lower molar (CMu15-501) recovered from the Upper Lignite Member of Chiang Muan. The age of Chiang Muan is estimated to be latest Middle or earliest Late Miocene (around 11 Ma), based on the mammalian fauna.This revised version was published online in April 2005 with corrections to the cover date of the issue.     

First hominoid from the Miocene of Ethiopia and the evolution of the catarrhine elbow

The first known fossil ape from the early-middle Miocene of Fejej, Ethiopia, is described here. The specimen, FJ-18SB-68, is a partial ulna from a locality dated by ⁴⁰Ar/³⁹Ar and paleomagnetic methods to a minimum age of 16.18 MYA. Compared to a variety of extant and fossil ulnae, FJ-18SB-68 is most similar to Turkanapithecus, Proconsul, and Pliopithecus, and appears to have been an arboreal quadruped with substantial forearm rotational mobility. Among the extant ulnae, canonical variates analysis successfully discriminates platyrrhines from catarrhines and within the latter, cercopithecoids from hominoids. Basal catarrhines (e.g., Aegyptopithecus) are platyrrhine-like in their morphology. Two basic trends appear to evolve from this generalized template: one with less mobile and more habitually pronated forearms, as seen in living and fossil cercopithecoids (including Victoriapithecus and Paracolobus), and another with greater forearm rotational mobility in fossil and modern hominoids. Primitive Miocene apes, including Proconsul, Turkanapithecus, and FJ-18SB-68, share with extant hominoids a more laterally positioned and laterally facing radial notch and an incipient trochlear keel. This morphology, along with a large insertion area for m. brachialis, suggests a departure from the more habitually pronated hand posture of monkeys and may indicate greater climbing abilities in these arboreally quadrupedal apes. Later Miocene apes, such as Oreopithecus and Dryopithecus share additional morphological features with hominoids, indicating considerable suspensory and climbing capabilities. Am J Phys Anthropol 105:257–277, 1998. © 1998 Wiley-Liss, Inc.     

A new hominoid hamate and first metacarpal from the Late Miocene Nagri Formation of Pakistan

A hamate and the proximal part of a first metacarpal from the type locality of the Nagri Formation in Pakistan, and attributed to Sivapithecus parvada, are described. In overall proportions, the hamate is rather robust, showing most similarity to that of Gorilla. Unlike extant hominoids it lacks a well-developed hamulus, and its triquetral facet is morphologically dissimilar to that in extant anthropoids. The morphology of the hamate indicates effective weight transmission through the ulnar side of the wrist, limited ulnar deviation and restricted extension in the triquetrohamate joint, and stability of the hamatometacarpal joints. The morphology of the partial first metacarpal is most similar to that of Pan. Previously described postcranial bones of S. parvada indicate that its locomotor behaviour included both quadrupedalism and climbing. This is consistent with the limited evidence of the first metacarpal, whereas the hamate strongly emphasizes the quadrupedal aspect of the locomotor repertoire.     

Geology and fauna of the middle Miocene hominoid site at Muruyur,Baringo district,Kenya

The middle Miocene sediments assigned to the Muruyur Beds have yielded abundant faunal remains which indicate an age somewhere near the early part of the middle Miocene, perhaps being earlier in time than Fort Ternan but probably coeval or slightly later than Maboko. Available radioisotopic age determinations suggest that the beds are between 13.5 and 14 m.y. old, which seems to be too young when compared with the biostratigraphic estimate. The importance of Muruyur Beds lies in their rich fossil content which includes hominoids of an age which is in general poorly represented in East Africa’s fossil record. This article places the fossil discoveries on record, and discusses their geological context.     

Ontogeny of the hominoid scapula: The influence of locomotion on morphology

Primate shoulder morphology has been linked with locomotor habits, oftentimes irrespective of phylogenetic heritage. Among hominoids, juvenile African apes are known to climb more frequently than adults, while orangutans and gibbons maintain an arboreal lifestyle throughout ontogeny. This study examined if these ontogenetic locomotor differences carry a morphological signal, which should be evident in the scapulae of chimpanzees and gorillas but absent in taxa that do not display ontogenetic behavioral shifts. The scapular morphology of five hominoid primates and one catarrhine outgroup was examined throughout ontogeny to evaluate if scapular traits linked with arboreal activities are modified in response to ontogenetic behavioral shifts away from climbing. Specifically, the following questions were addressed: 1) which scapular characteristics distinguish taxa with different locomotor habits; and 2) do these traits show associated changes during development in taxa known to modify their behavioral patterns? Several traits characterized suspensory taxa from nonsuspensory forms, such as cranially oriented glenohumeral joints, obliquely oriented scapular spines, relatively narrow infraspinous fossae, and inferolaterally expanded subscapularis fossae. The relative shape of the dorsal scapular fossae changed in Pan, Gorilla, and also Macaca in line with predictions based on reported ontogenetic changes in locomotor behavior. These morphological changes were mostly distinct from those seen in Pongo, Hylobates, and Homo and imply a unique developmental pattern, possibly related to ontogenetic locomotor shifts. Accordingly, features that sorted taxa by locomotor habits and changed in concert with ontogenetic behavioral patterns should be particularly useful for reconstructing the locomotor habits of fossil forms. Am J Phys Anthropol 152:239–260, 2013. © 2013 Wiley Periodicals, Inc.     

Bio-magnetostratigraphy and environment of the oldest Eurasian hominoid from the Early Miocene of Engelswies (Germany)

The paleobiogeography of hominoids exhibits a puzzling pattern of migrations between and within Africa and Eurasia. A precise dating of hominoid-bearing localities is therefore essential to reveal the timing, direction and possible causes of dispersals. Here, we present a bio-magnetostratigraphic analysis of the section of Engelswies (Southern Germany, Upper Freshwater Molasse, North Alpine Foreland Basin) where the oldest Eurasian hominoid was found. Our paleomagnetic results reveal a very short normal and a reverse magnetic polarity for the entire section. The polarity record is correlated to the Astronomical Tuned Neogene Time Scale using an integrated stratigraphic approach. This approach follows the chronostratigraphic framework for the Upper Freshwater Molasse, which combines magnetostratigraphy with biostratigraphic, lithostratigraphic and ⁴⁰Ar/³⁹Ar dating results. According to this outcome, the reverse polarity of the Engelswies section most likely correlates to magnetochron C5Cr. The origin of the short normal polarity remains enigmatic. The magnetostratigraphic calibration and the evolutionary level of the Engelswies small mammal fauna suggest an age of 17.1-17.0 Ma (Early Karpatian, Early Miocene) for the oldest Eurasian hominoid, and roughly confirm the estimates of Heizmann and Begun (2001). The estimated age suggests that the first hominoids in Eurasia are contemporaneous with Afro-Arabian afropithecins, and dispersal may have been facilitated by intra-Burdigalian (∼18-17 Ma) sea-level low stands and the beginning of the Miocene Climate Optimum. The paleoclimatic and environmental reconstruction of the Engelswies locality indicates a lakeshore environment near dense subtropical rain forest vegetation, where paratropical temperatures (mean annual temperature around 20 °C) and humid conditions (mean annual precipitation > 1.100 mm) prevailed.     

云南昭通水塘坝中新世末期古猿化石点的云南貘化石(英文)

此前,中国晚中新世到早上新世的貘化石发现较为稀少。最近几年,在云南省昭通市水塘坝禄丰古猿化石点发现了数量可观的中新世晚期貘化石,其时代为6~6.5 Ma。新发现材料包括左上颌骨带P2-M2、4件下颌骨带基本完整颊齿列、若干零散牙齿。新材料可归人先前报道的小型貘类——云南貘Tapirus yunnanensis。云南貘比甘肃的和政貘Tapirus hezhengensis时代较晚,体形也稍大;两者都比上新世一更新世的貘类明显较小。新发现的化石材料对更全面认识云南貘的特征提供了新信息。在中一晚中新世期间,云南是我国貘类动物的演化中心,先后发现的化石点有开远(小龙潭)、禄丰(石灰坝)、元谋(小河、竹棚、雷老)、昭通(水塘坝)及其他地点。在晚新生代期间,我国貘类动物演化的主要变化趋势是体形逐渐增大,这一特点或许具有时代意义。在中国发现的中新世貘类毫无例外都是体形较小的类型,这与欧洲及北美的情况迥异。