Late Turolian Mesopithecus (Mammalia: Cercopithecidae) from Axios Valley (Macedonia,Greece): earliest presence of M. monspessulanus in Europe

The fossil mammal localities of the Axios Valley (Macedonia, Greece) yielded a rich collection of Mesopithecus remains, which were first described at the beginning of the 1990s. The late Turolian Dytiko localities include several specimens of Mesopithecus, which were originally separated in two size-related forms: the relatively larger-sized M. cf. or aff. pentelicus, and the relatively smaller-sized M. cf. monspessulanus. However, some Dytiko specimens were only partially described, or remained even undescribed because the cranium either was still in connection with the mandible or was markedly deformed. Later, careful cleaning of the materials and their revision based on larger comparative samples indicate that they represent two distinct species: the relatively larger-sized specimens belong to M. pentelicus, while the relatively smaller-sized ones to M. monspessulanus. The Dytiko M. pentelicus has slightly smaller dental dimensions compared to M. pentelicus from Pikermi, indicating a possible trend for size decrease. The Dytiko M. monspessulanus, although close to the typical form of this species, has somewhat larger dimensions. The semi-terrestrial M. pentelicus disappeared at the end of the Miocene, while some of its populations adapted to the wetter and more woody Pliocene habitats, finally giving origin to M. monspessulanus, whose elbow-joint indicates an arboreal lifestyle. M. monspessulanus was widely distributed in Europe, but its remains are very scarce and are all from the Pliocene. Some questionable indications for the presence of M. monspessulanus in the latest Miocene come from some Italian Messinian sites (Gravitelli, Baccinello V3), but this scenario still needs verification. However, even if its presence in some Italian localities was confirmed, the Dytiko M. monspessulanus nonetheless represents the earliest known occurrence of this species in Europe, as the Dytiko fauna is considered as latest Turolian (pre-Messinian, 7.0–6.0 Ma).     

Stable isotope ecology of Miocene bovids from northern Greece and the ape/monkey turnover in the Balkans

Eurasia was home to a great radiation of hominoid primates during the Miocene. All were extinct by the end of the Miocene in Western Eurasia. Here, we investigate the hypothesis of climate and vegetation changes at a local scale when the cercopithecoid Mesopithecus replaced the hominoid Ouranopithecus along the Axios River, Greece. Because they are herbivorous and were much more abundant than primates, bovids are preferred to primates to study climate change in the Balkans as a cause of hominoid extinction. By measuring carbon stable isotope ratios of bovid enamel, we conclude that Ouranopithecus and Mesopithecus both evolved in pure C₃ environments. However, the large range of δ¹³C values of apatite carbonate from bovids combined with their molar microwear and mesowear patterns preclude the presence of dense forested landscapes in northern Greece. Instead, these bovids evolved in rather open landscapes with abundant grasses in the herbaceous layer. Coldest monthly estimated temperatures were below 10 °C and warmest monthly temperatures rose close to or above 20 °C for the two time intervals. Oxygen isotope compositions of phosphate from bulk samples did not show significant differences between sites but did show between-species variation within each site. Different factors influence oxygen isotope composition in this context, including water provenience, feeding ecology, body mass, and rate of amelogenesis. We discuss this latter factor in regard to the high intra-tooth variations in δ¹⁸O_p reflecting important amplitudes of seasonal variations in temperature. These estimations fit with paleobotanical data and differ slightly from estimations based on climate models. This study found no significant change in climate before and after the extinction of Ouranopithecus along the Axios River. However, strong seasonal variations with relatively cold winters were indicated, conditions quite usual for extant monkeys but unusual for great apes distributed today in inter-tropical regions.     

Additional specimens of Hyracoidea (Mammalia) from the Early and Middle Miocene of Kenya

We describe unpublished material of the poorly known hyracoid, Brachyhyrax aequatorialis from the Early Miocene of Songhor and Koru, Kenya, on the basis of specimens stored in the Community Museums of Kenya, the National Museums of Kenya and the Natural History Museum, London. As a result, we added 17 specimens to the hypodigm of this species which was previously known from only seven specimens. In addition, we describe further material of another hyracoid, Afrohyrax championi, from the Early Miocene of Mfwangano and the Middle Miocene of Kipsaraman. The chronological distributions of the two hyracoids do not overlap. Brachyhyrax appears to have lived in forest environments, a suggestion supported by its brachyodont dentition and the associated molluscan fauna, whereas Afrohyrax seems to have lived in more open wooded habitats which agree with its slightly more hypsodont dentition and the cursorial postcranial skeleton as well as with the land snails that occur with it.     

The Hand of Cercopithecoides williamsi (Mammalia,Primates): Earliest Evidence for Thumb Reduction among Colobine Monkeys

Thumb reduction is among the most important features distinguishing the African and Asian colobines from each other and from other Old World monkeys. In this study we demonstrate that the partial skeleton KNM-ER 4420 from Koobi Fora, Kenya, dated to 1.9 Ma and assigned to the Plio-Pleistocene colobine species Cercopithecoides williamsi, shows marked reduction of its first metacarpal relative to the medial metacarpals. Thus, KNM-ER 4420 is the first documented occurrence of cercopithecid pollical reduction in the fossil record. In the size of its first metacarpal relative to the medial metacarpals, C. williamsi is similar to extant African colobines, but different from cercopithecines, extant Asian colobines and the Late Miocene colobines Microcolobus and Mesopithecus. This feature clearly links the genus Cercopithecoides with the extant African colobine clade and makes it the first definitive African colobine in the fossil record. The postcranial adaptations to terrestriality in Cercopithecoides are most likely secondary, while ancestral colobinans (and colobines) were arboreal. Finally, the absence of any evidence for pollical reduction in Mesopithecus implies either independent thumb reduction in African and Asian colobines or multiple colobine dispersal events out of Africa. Based on the available evidence, we consider the first scenario more likely.     

Folivory or fruit/seed predation for Mesopithecus, an earliest colobine from the late Miocene of Eurasia?

Here we compare dental microwear textures from specimens of the fossil genus Mesopithecus (Cercopithecidae, Colobinae) from the late Miocene of Eastern Europe with dental microwear textures from four extant primate species with known dietary differences. Results indicate that the dental microwear textures of Mesopithecus differ from those of extant leaf eaters Alouatta palliata and Trachypithecus cristatus and instead resemble more closely those of the occasional hard-object feeders Cebus apella and Lophocebus albigena. Microwear texture data presented here in combination with results from previous analyses suggest that Mesopithecus was a widespread, opportunistic feeder that often consumed hard seeds. These data are consistent with the hypothesis that early colobines may have preferred hard seeds to leaves.     

A cercopithecid primate from the late miocene of Molayan,Afghanistan, with remarks onMesopithecus

The first fossil primate discovered in Afghanistan comes from the Late Miocene of Molayan, Khurdkabul Basin. The materials consist of an almost-complete juvenile mandible and an isolated P₃. These two specimens do not significantly differ from thePikermi Mesopithecus pentelicus and are assigned to this species. The primate mandible from the Late Miocene of Maragheh, Iran, which has always been referred by all authors toM. pentelicus, differs from the Pikermi and Molayan materials. It must be assigned to another taxon, probably a new one. The geographic range ofM. pentelicus turns out to be considerably wider in view of the discovery of the species in Molayan, eastern Afghanistan.     

A succession of Miocene rodent assemblages from fissure fillings in southern France: palaeoenvironmental interpretation and comparison with Spain

An Early to Late Miocene sequence of rodent assemblages from southern France has been quantitatively studied. The resulting pattern seems very similar to a contemporary sequence from central Spain (Calatayud–Teruel Basin). The fossil mammal-bearing localities are of different types: mainly karst infills in France and localities situated in sedimentary basins in Spain. In order to interpret the fossil record, a comparison has been made between southern France faunas of similar age but collected in karst infills and in basin deposits. There seems to be no difference between the two kinds of faunas and thus there is no indication that karst infills systematically give a picture of drier and more open environments. Both types of localities may give a similar relative abundance of taxa and when differences exist they can be attributed to local conditions. The comparison between southern France and the Calatayud–Teruel Basin (central Spain) shows that: (1) similar trends occurred in the two areas; (2) differences between spectra were more important during the late Early Miocene than during the Middle Miocene; (3) the shift between the late Early Miocene and the Middle Miocene environments in southern France does not seem to be correlated with a general drop in temperatures as inferred from the analysis of central Spain faunas.     

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.     

Cercopithecoid primate postcranial fossils from Cooper's D,South Africa

Among several highly fossiliferous localities in the Bloubank Valley (Gauteng, South Africa), the Cooper's Cave System has been known since 1938 and has produced a rich fossil assemblage, including some remains of the early hominin Paranthropus robustus. In 2001, excavations began at a new locality, Cooper's D, which dates to ～1.4-1.5 Ma. Although hominins are relatively rare in the assemblage, remains of cercopithecoid primates are much more common. Craniodental fossils currently indicate the possible presence of at least three large-bodied cercopithecoid primate genera at Cooper's D: Gorgopithecus, Papio, and Theropithecus. In this study, we identify and describe > 100 cercopithecoid primate postcranial fossils representing all regions of the appendicular skeleton. The specimens come from several age classes and size morphs; more than one third of the fossils described are from sub-adult and juvenile individuals. The adult postcranial fossils vary substantially in size, with body masses estimated between 30 and 60 kg (from 16 of the better preserved specimens). The functional morphology of the postcranial remains indicate that these elements come from animals that likely utilized terrestrial substrates, but they remain difficult to definitively attribute to Gorgopithecus, Theropithecus, or Papio given the absence of associated skeletons. The smaller specimens likely belong to Papio while the larger ones can be attributed to the other two genera. Because Cooper's D has also yielded fossils of the early hominin Paranthropus robustus, this raises the question of how these four large-bodied, mostly terrestrial primates sympatrically utilized the landscape.     

Brief communication: New primate remains from the Miocene of Namibia,Southern Africa

Miocene primates from southern Africa are extremely rare. For this reason we wish to place on record several interesting new fossil primate specimens recently recovered from the Miocene sites of Berg Aukas and Harasib in the Otavi Mountain region of northern Namibia. The new finds consist of a virtually complete atlas vertebra from Berg Aukas attributable to the hominoid Otavipithecus namibiensis and two teeth and four postcranial fragments from Harasib referrable to Cercopithecoidea. The atlas vertebra exhibits anatomical characteristics intermediate between those of modern cercopithecoids and hominoids which may be indicative of a transition from pronograde to orthograde postures. The cercopithecoid remains show that the earliest Old World monkeys known from southern Africa were small, approximately the size of vervet monkeys. These new specimens are important because they provide the first evidence relating to possible positional behaviors of Otavipithecus and the earliest fossil record of cercopithecoids from southern Africa. © 1996 Wiley-Liss, Inc.     

Brief communication: Plio‐Pleistocene eagle predation on fossil cercopithecids from the Humpata Plateau,southern Angola

Recent studies suggest a large raptor such as the crowned eagle (Stephanaoetus coronatus) was responsible for collecting at least a portion of the primate fauna from the South African fossil site of Taung, including its lone hominin specimen. This taphonomic signature at Taung is currently regarded as a unique and, most likely, isolated case in primate and human evolution. However, the activities of large, carnivorous birds should also be detectable at other primate fossil localities in Africa if raptors have been a strong selective force throughout primate evolution. Over the last 60 years, a collection of extinct cercopithecids has been assembled from several cave breccias on the Humpata Plateau in southern Angola. The material, dated near the Plio‐Pleistocene boundary, includes an assortment of craniodental and postcranial remains variably assigned to Papio (Dinopithecus) cf. quadratirostris, Parapapio, Cercopithecoides, and Theropithecus. We compare the Angolan and Taung material to remains of extant primates killed by crowned eagles in the Ivory Coast's Tai National Park. Our analysis indicates that the size distribution and composition of fauna from the localities is quite similar and that there are striking consistencies in damage to the crania from each site. The absence of large bodied (>20 kg) primates and other mammalian taxa at the Taung hominin locality and Tai, and their rarity in Angola, combined with the strong likelihood that raptor nests were positioned near fissure openings at both fossil localities, provides additional support for eagle involvement. On the basis of this evidence, we conclude that at least some of the Angolan cercopithecids were most likely raptor prey and hypothesize that raptor predation has been a strong and perhaps underappreciated selective force during the course of primate evolution. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Paleoecology of the Miocene fossil mammal fauna from Bełchatów (Poland)

Miocene fauna of land mammals is known from numerous fossil localities in western Europe but its remains are rare in the north-eastern part of this continent. The open brown coal mine at Bełchatów in central Poland revealed numerous remains of small and large mammals representing three Miocene biozones from MN4 to MN8/9. They indicate the presence of relatively stable subtropical conditions during all the time of the Miocene deposition in the Bełchatów basin with only slight cooling before the end of the sedimentation.

     

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.     

Die erste Wirbeltierfauna aus der miozänen Braunkohle der Niederrheinischen Bucht (Ville-Schichten, Tagebau Hambach)

This paper describes the first Miocene vertebrate fauna from the Ville Series of the Lower Rhine Embayment in western Germany. The fossil material was discovered in the vast Hambach open cast lignite mine ofRheinbraun. The numerous remains, mostly isolated teeth and disarticulated bones, were accumulated in a channel-fill within the main lignite seam. The fauna consists of chondrichthyans, teleosteans, amphibians, reptiles, and mammals which inhabited brackish-estuarine, lacustrine-fluviatile, and terrestrial environments. This is the first fauna to provide an insight into the diversity of vertebrates which lived in the forested peat swamps of the Rhenish Tertiary. Based on the rich association of more than 70 mammalian taxa, including about 30 rodent species, this late Orleanian fauna can be correlated with the upper part of the Mammalian Neogene biozone MN 5. Therefore the middle part of the main lignite seam (horizon 6C of the local lithostratigraphy) can be placed in the early middle Miocene. This means an age of 15.2-16.0 Ma (Langhian or Reinbekian of the stratigraphy of the NW German Tertiary). For some mammalian taxa (Lanthanotherium, Plesiosorex,Miopetaurista, Myoglis, Fahlbuschia, ‘Lartetomys’, Anchitheriomys, Dorcatherium), Hambach 6C is one of the richest European localities. In addition this assemblage contains some very rare mammals (‘Lartetomys’, Anchitheriomys, Orygotherium, Pliopithecus). The Hambach vertebrates are of great palaeobiogeographic importance as they represent the north westernmost outpost of terrestrial Miocene faunas in Europe.     

Palaeozoological insights into management options for a threatened mammal: southern Africa’s Cape mountain zebra (Equus zebra zebra)

Aim Promoting population growth of genetically distinct subpopulations of Cape mountain zebra (Equus zebra zebra) is crucial to the survival of the subspecies. Several important Cape mountain zebra reserves are dominated by fynbos vegetation, and population growth is limited by a lack of grassland habitat. A fossil ungulate sequence spanning the last c. 18,000 years is examined to understand the long‐term history of this conservation challenge. Location Boomplaas Cave (BPA), South Africa. Methods The fossil sequence from BPA is examined to reconstruct ungulate community dynamics in relation to climate and vegetation change over the last 18,000 years. Results Ungulates from 18,000 to 12,000 years ago suggest an expansion of open grasslands that supported a grazing ecosystem dominated by an extinct caprine antelope and equid remains attributed to E. zebra and E. quagga. At the onset of the Holocene, the grazing ungulate community disappears and small browsers and mixed feeders dominate the assemblage, indicating the loss of open grassland vegetation. Several open‐habitat grazers go extinct at this time, and Equus persists at much lower abundances. This shift can be explained by global climate change across the Pleistocene–Holocene transition. Main conclusions The fossil sequence supports contemporary observations indicating that access to open grassland is crucial to maintaining large Cape mountain zebra subpopulations. Although fynbos is abundant throughout the historic range of the Cape mountain zebra, fossil evidence suggests that such vegetation is unlikely to support dense populations. It has been suggested that the acquisition of agricultural lands that were historically converted to open grasslands for livestock could promote Cape mountain zebra population growth. Results presented here support this management option, as the open grasslands in these converted landscapes likely approximate the vegetation structure during latest Pleistocene, when grasslands were widespread and grazing ungulates abundant.     

A functional multivariate analysis of Mesopithecus (Primates: Colobinae) humeri from the Turolian of Greece

The genus Mesopithecus is well represented in the late Miocene of Greece by several recognized species. The present paper investigates functional aspects of the humeri of Mesopithecus delsoni/pentelicus, M. pentelicus and M. aff. pentelicus of several Turolian sites from central and northern Greece, using multivariate approaches. For these purposes, we selected significant humeral functional features, which were represented by 23 linear dimensions and three angles on 14 fossil humeri and 104 humeri from 10 genera and 22 species of extant African and Asian Colobines. All size-adjusted measurements were examined through a principal components analysis, followed by a discriminant function analysis, and a canonical variates analysis. All analyses revealed that the selected characters were able to discriminate between extant colobine genera. Functional groups, such as arboreal walking/climbing, arboreal walking/suspensory and semi-terrestrial walking, were set apart from a central cluster formed by the arboreal walking and arboreal walking/terrestrial groups. This cluster also grouped the three studied Mesopithecus species, which were mainly classified as arboreal walkers with significant terrestrial activities. These observations match with paleoenvironmental reconstructions and the suggested opportunistic feeding habits. Moreover, the overall arboreal/terrestrial locomotor tendencies of these fossil forms are discussed in relation to their earlier migration from Africa and later dispersal to eastern and southern Asia.     

Knuckle-walking in Sivapithecus? The combined effects of homology and homoplasy with possible implications for pongine dispersals

Sivapithecus is a Miocene great ape from South Asia that is orangutan-like cranially but is distinctive postcranially. Work by others shows that the humerus resembles large terrestrial cercopithecoids proximally and suspensory hominoids distally, but most functional interpretations nevertheless situate Sivapithecus in an arboreal setting. We present a new quantitative analysis of the Sivapithecus capitate and hamate. Though the functional morphology of both bones suggests some degree of arboreality, the overall morphology is most similar to knuckle-walking African apes. Other features of the Sivapithecus humerus and hind limb are also functionally consistent with knuckle-walking, and we suggest that this locomotor behavior is a valid alternative functional interpretation of the postcranial morphology. We speculate that knuckle-walking in Sivapithecus would have evolved independently from African apes, perhaps for similar ecological reasons. The discovery of a possible pongine knuckle-walker challenges the hypotheses that (1) knuckle-walking evolved only once in hominoids and (2) knuckle-walking is too highly specialized to be the positional behavior from which human bipedalism evolved. The possibility of knuckle-walking in Sivapithecus may help to explain not only the curious combination of characters that typify the postcranium but also the unique postcranial morphology of extant Pongo. Furthermore, it may clarify the distribution of fossil pongines across many ecological zones in Eurasia in the Miocene and Pleistocene, as well as, independently, the spread of African apes across a diversity of environments in equatorial Africa.     

A fossil record of the Eastern clade of Blanus (Amphisbaenia: Blanidae) from the late Miocene of Ukraine

The fossil history of worm lizards in Eastern Europe is very patchy because of the small number of localities and the fragmentary vertebral material available. A new record of the genus Blanus from the late Miocene of the Egorovka 1 and 2 localities in Ukraine is reported here. Based on its dental morphology, Blanus from Egorovka differs from all extinct species of the genus and from the extant B. cinereus complex. It is most closely related to the Eastern clade of Blanus (B. strauchi complex) in having enlarged second and small third dentary teeth. Blanus from Egorovka is the second fossil record of the B. strauchi complex and the first record of the Eastern clade outside of its current geographical range. It indicates a wider past range of the Eastern European Blanus, including the northern Black Sea coastal area.