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.     

Sivapithecus simonsi,a new species of miocene hominoid,with comments on the phylogenetic status of the ramapithecinae

The Ramapithecinae are an extinct, mainly Miocene group of hominoids, whose relationship to modern taxa is disputed. Some regard them as hominids, while others view them as ancestral toPongo,or even as the group ancestral to both hominids and extant apes. In this paper a systematic revision of Ramapithecinae is undertaken. Sivapithecus sivalensis andRamapithecus punjabicus are considered the same species, with the former name having priority. A new Indian species,Sivapithecus simonsi,is recognized. Ramapithecine anatomy is reviewed and compared with that of gracileAustralopithecus, early and middle MioceneProconsul andDryopithecus, and living pongidsPan, Gorilla, andPongo.Ramapithecines are shown to be much more primitive or “ape-like” than some have argued. Anatomical data are evaluated cladistically with several results. Parallel evolution in the jaws, teeth, and facial structure of hominoids appears to be the rule rather than the exception. Bearing this in mind, nevertheless, from the available evidence of anatomy, ramapithecines are cladistically hominids.     

A new species of Anthracotheriidae, Merycopotamus medioximus nov. sp. from the Late Miocene of the Potwar Plateau, Pakistan

New anthracotheriid remains, discovered by the H-GSP in well-dated localities from the Potwar plateau in the North of Pakistan, between 10.4 and 8.6 Ma, are described and attributed to Merycopotamus medioximus nov. sp. This new species displays an intermediate morphology between the older M. pusillus and the more recent M. dissimilis. These results permit to emend the Merycopotamus diagnosis. To cite this article: F. Lihoreau et al., C. R. Palevol 3 (2004).

Résumé

Une nouvelle espèce d'Anthracotheriidae, Merycopotamus medioximus nov. sp. du Miocène récent du plateau du Potwar, Pakistan. Des restes d'Anthracotheriidae, découverts par le H-GSP dans des localités bien datées du plateau du Potwar, au Nord du Pakistan, entre 10,4 et 8,6 Ma, sont décrits et attribués à Merycopotamus medioximus nov. sp. Cette nouvelle espèce possède une morphologie intermédiaire entre M. pusillus, espèce plus ancienne, et M. dissimilis, plus récente. Ces résultats permettent en outre d'émender la diagnose du genre Merycopotamus. Pour citer cet article : F. Lihoreau et al., C. R. Palevol 3 (2004).     

New Sivapithecus postcranial specimens from the Siwaliks of Pakistan

Several new postcranial elements of Sivapithecus from the Siwaliks of Pakistan are described. These include a distal femur from the U-level of the Dhok Pathan Formation, a navicular from the Chinji Formation, and seven manual and pedal phalanges from the Nagri Formation. The functional morphology of these elements adds new detail to the reconstruction ofSivapithecus positional behavior. Femoral cross-sectional geometry indicates that the shaft was adapted to support mediolaterally directed loading. Femoral condylar asymmetry and a broad but shallow trochlea are distinctly ape-like, revealing capabilities for both rotation and withstanding eccentric loading in the knee. The navicular is characterized by features relating to a broad mid-tarsus and broad distal articulations for the cuneiforms. It also lacks a navicular tubercle as in Pongo. These features suggest that the foot was capable of a powerful grip on large supports, with an inversion/supination capability that would permit foot placement in a variety of positions. The morphology of the new phalanges, including evidence for a relatively large pollex, similarly suggests powerful grasping, consistent with prior evidence from the hallux and tarsus. The functional features of the new specimens permit refinement of previous interpretations of Sivapithecus positional capabilities. They suggest a locomotor repertoire dominated by pronograde activities and also such antipronograde activities as vertical climbing and clambering, but not by antipronograde suspensory activities as practiced by extant apes.     

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.     

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.     

Molar enamel thickness in European Miocene and extant hominoidea

Based on a roentgenographic analysis, the molar enamel of certain European Miocene dryopithecines is absolutely thick (r=1.03–1.30 mm in thickness); the molar enamel of certain European pliopithecines is thin (r=0.32–0.82 mm thick). The rank order for enamel thickness in extant hominoids (from thickest to thinnest) is confirmed to beHomo, Pongo, Gorilla, Pan, andHylobates. There is a great deal of enamel thickness variability within the great ape sample. Extant analogues suggest that dryopithecines were probably adapted to a frugivorous/gramnivorous dietary regimen, while pliopithecines were probably better suited to folivory.     

A juvenile skull of Acerorhinus yuanmouensis (Mammalia: Rhinocerotidae) from the Late Miocene hominoid fauna of the Yuanmou Basin (Yunnan,China)

A unique juvenile skull bearing both milk premolars and unerupted but fully developed permanent premolars and molars (observed using X-ray microcomputed tomography), and some isolated upper cheek teeth, all from the Late Miocene hominoid fauna of the Yuanmou Basin (Yunnan, China), closely resemble craniodental material of Acerorhinus yuanmouensis Zong, 1998 from the same locality, and are referred to this species. A phylogenetic analysis based on 214 craniodental morphological characters scored for 31 terminal taxa reveals that A. yuanmouensis should be assigned to the genus Acerorhinus indeed. The newly discovered specimens improve our understanding of this species, especially with respect to the morphology of the milk premolars and premolars. Two intraspecific variations in the upper premolars are noted: a lingual bridge may be present or absent, and the lingual cingulum continuous or reduced. The analysis also indicates that: the phylogenetic status of Acerorhinus lufengensis Deng and Qi, 2009 should be reconsidered; “Aceratherium” huadeensis Qiu, 1979 does neither belong to Aceratherium nor Acerorhinus, and its phylogenetic status remains debatable.     

Paleosols and paleoenvironments of the middle Miocene,Maboko Formation,Kenya

The middle Miocene (15 Ma) Maboko Formation of Maboko Island and Majiwa Bluffs, southwestern Kenya, has yielded abundant fossils of the earliest known cercopithecoid monkey (Victoriapithecus macinnesi), and of a kenyapithecine hominoid (Kenyapithecus africanus), as well as rare proconsuline (Simiolus leakeyorum, cf. Limnopithecus evansi) and oreopithecine apes (Mabokopithecus clarki, M. pickfordi), and galagids (Komba winamensis). Specific habitat preferences can be interpreted from large collections of primate fossils in different kinds of paleosols (pedotypes). Fossiliferous drab-colored paleosols with iron-manganese nodules (Yom pedotype) are like modern soils of seasonally waterlogged depressions (dambo). Their crumb structure and abundant fine root-traces, as well as scattered large calcareous rhizoconcretions indicate former vegetation of seasonally wet, wooded grassland. Other fossiliferous paleosols are evidence of nyika bushland (Ratong), and early-successional riparian woodland (Dhero). No fossils were found in Mogo paleosols interpreted as saline scrub soils. Very shallow calcic horizons (in Yom, Ratong, and Mogo paleosols) and Na-montmorillonite (in Mogo) are evidence of dry paleoclimate (300-500 mm MAP=mean annual precipitation). This is the driest paleoclimate and most open vegetation yet inferred as a habitat for any Kenyan Miocene apes or monkeys. Victoriapithecus was abundant in dambo wooded grassland (Yom) and riparian woodland (Dhero), a distribution like that of modern vervet monkeys. Kenyapithecus ranged through all these paleosols, but was the most common primate in nyika bushland paleosols (Ratong), comparable to baboons and macaques today. Mabokopithecus was virtually restricted to riparian woodland paleosols (Dhero), and Simiolus had a similar, but marginally wider, distribution. Habitat preferences of Mabokopithecus and Simiolus were like those of modern colobus monkeys and mangabeys. A single specimen of Komba was found in dambo wooded grassland paleosol (Yom), a habitat more like that of the living Senegal bushbaby than of rainforest galagids. A shift to non-forest habitats may explain the terrestrial adaptations of Victoriapithecus, basal to the cercopithecid radiation, and of Kenyapithecus, basal to the hominoid radiation. Both taxa are distinct from earlier Miocene arboreal proconsulines, oreopithecines and galagids.     

A new species of Megacerini (Cervidae, Artiodactyla) from the Late Miocene of Taralyk-Cher, Tuva (Russia), and remarks on the relationships of the group

A new species of a megacerine deer, Praesinomegaceros venustus, from the Taralyk-Cher locality in the Tuva region on the south of Eastern Siberia, Russia, is described. P. venustus nov. sp., the oldest known megacerine, occurred in Asia in the Late Turolian, about 7 Ma. The early history of megacerines is discussed. The existence of the phyletic lineage Cervavitus-Praesinomegaceros-Sinomegaceros in Asia is confirmed.     

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.     

Late Miocene insular mice from the Tusco-Sardinian palaeobioprovince provide new insights on the palaeoecology of the Oreopithecus faunas

Oreopithecus bambolii is one of the few hominoids that evolved under insular conditions, resulting in the development of unique adaptations that have fueled an intensive debate. The palaeoenvironment associated with this great ape has been the subject of great controversy as well. On the one hand, palaeobotanical data indicate that Oreopithecus likely inhabited mixed mesophytic forests interrupted by swamps; on the other hand, an abundance of hypsodont bovids points towards the existence of dry and open environments. Here, we provide a new approach based on the ecomorphology of the extinct endemic Muridae (rats and mice) of the so-called Oreopithecus faunas. Our results show that the successive species of endemic insular murids (Huerzelerimys and Anthracomys) evolved a number of adaptations observed only in extant family members that include significant proportions of grass in their diet. While this fits the pattern exhibited by large mammals, it contrasts with the available palaeobotanical information, which indicates that grasses were minor components of the vegetation. This contradiction may be explained because these endemic murids may have been adapted to the consumption of particular food items such as hard parts of aquatic plants (as shown by some extant murid species). However, because it is unlikely that the remaining herbivore mammals were adapted to this diet as well, we favour an alternative hypothesis that takes into account the peculiar ecological conditions of insular ecosystems leading to a density-dependent selective regime with strong competition. Such a regime would promote the selection of dental adaptations to increase feeding efficiency and durability of the dentition (such as hypsodonty) as seen in some fossil insular ruminants. This hypothesis requires further testing, but may partly account for parallel evolution of dental traits in phylogenetically unrelated insular mammals.     

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.     

Manual proximal phalanx of Laccopithecus robustus from the Latest Miocene site of Lufeng

An isolated proximal phalanx recovered from the Latest Miocene site of Lufeng, China, is assigned to the species Laccopithecus robustus. It shows a number of features associated with grasping and manual suspension, such as dorsoventral curvature of the shaft, pronounced flexor sheath ridges, dorsoventrally deep trochlear condyles, and a prominent epicondylar ridge. In addition, the phalanx displays a combination of features that is similar to extant hylobatids, including asymmetry of the flexor ridges and transverse concavity of the ventral surface of the distal shaft, and thick cortices. Despite the many dental similarities between Laccopithecus and Pliopithecus, the phalangeal morphology of the former is quite distinct and shows many similarities to the extant hylobatids, particularly the siamang.     

Late Miocene Microstonyx remains (Suidae, Mammalia) from Northern China

Several large suid cranial remains attributed to Microstonyx major are part of a new Hipparion Fauna collection from the Hezheng area, Northern China. The new material confirms the presence of Microstonyx in the late Miocene of the area. The Chinese form belongs to a small-sized eastern population with reduced premolar row and clear sexual bimodality. Statistical comparison shows that Microstonyx major was a polymorphic species and reinforces recognition of Hippopotamodon as a separate genus, defined by relatively stout premolars resulting from a different underlying pattern of allometric growth. The presence of Microstonyx in North China and the distinct suid assemblage that lived there suggest biogeographic connections between Northern China and Western Eurasia in contrast to isolation from Southern China and the Indian subcontinent. The suid fauna of the late Miocene of Northern China seems to have been restricted to the later, more humid phase represented by the Red Clay faunas.     

A fossil hominoid proximal femur from Kikorongo Crater, southwestern Uganda

The external morphology of a fragmentary right proximal femur from southwestern Uganda is described here. Discovered in the Kikorongo Crater of Queen Elizabeth National Park in 1961, this specimen was informally assigned to Homo sapiens (although never described) and tentatively dated to the late Pleistocene. However, because aspects of the external morphology of the femur align the fossil with the African great apes, we suggest that the Kikorongo femur may be the first postcranial fossil of the genus Pan. Like the African apes, the Kikorongo specimen lacks both an obturator externus groove and an intertrochanteric line. It has a short femoral neck with a circular cross section, and a narrow and deep superior notch. Using resampling statistics and discriminant function analysis, the Kikorongo femur clustered with the genus Pan, as opposed to Gorilla or Homo. However, if the specimen is from Pan, it would be large for this taxon. Furthermore, features that clearly distinguish the external morphology of Plio-Pleistocene hominin proximal femora from African ape femora, such as the shape of the femoral neck in cross section and femoral neck length, have converged in Holocene humans and African apes. Unfortunately, the internal morphology of the femoral neck of the Kikorongo fossil was not discernable. Although we hypothesize that the Kikorongo femur is from the genus Pan, there is such variability in the proximal femora of modern humans that, although it would be an unusual human, it remains possible that this fossil represents H. sapiens.