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Extant colobine monkeys are specialized leaf eaters. But during the late Miocene, western Eurasia was home to colobines that were less efficient at chewing leaves than they were at breaking seed shells. To understand the link between folivory and granivory in this lineage, the dietary niche of Mesopithecus delsoni and Mesopithecus pentelicus was investigated in southeastern Europe, where a major environmental change occurred during the late Miocene. We combined dental topographic estimates of chewing efficiency with dental microwear texture analysis of enamel wear facets. Mesopithecus delsoni was more efficient at chewing leaves than M. pentelicus, the dental topography of which matches an opportunistic seed eater. Concurrently, microwear complexity increases in M. pentelicus, especially in the northernmost localities corresponding to present-day Bulgaria. This is interpreted as a dietary shift toward hard foods such as seeds or tubers, which is consistent with the savanna and open mixed forest biomes that covered Bulgaria during the Tortonian. The fact that M. delsoni was better adapted to folivory and consumed a lower amount of hard foods than M. pentelicus suggests that colobines either adapted to folivory before their dispersal to Europe or evolved adaptations to leaf consumption in multiple occurrences.

Apart from a juvenile hominoid, the locality of Shuitangba (southwestern China, 6.5–6.0 Ma) has yielded a mandible and proximal femur attributed to the colobine genus Mesopithecus. A complete colobine calcaneus also accompanies this material, but its association with the other Mesopithecus material remains to be confirmed. These fossil elements are very important as they represent the oldest known colobines from East Asia, extend the dispersal of Mesopithecus to southwestern China, and underscore its close affinities and potential ancestry to the odd-nosed colobines. The present article focuses on the functional morphology of this complete calcaneus to reconstruct the positional habits, infer the paleocology, and understand the dispersal patterns of this fossil colobine. The studied characters corroborate the attribution of this element to colobines and support potential affinities with the Mesopithecus remains of the same locality. Functionally, characters such as the long and narrow tuber calcanei, the short proximal calcaneal region, and the relatively extended and long and narrow proximal calcaneoastragalar facet appear to enable habitual pedal flexion with conjunct inversion that accommodate the foot on diversely oriented and differently sized arboreal substrates. On the other hand, the relatively short distal calcaneal region is functionally related to (mainly terrestrial) quadrupedal activities, wherein thrust and rapid flexion are required. This combination of characters suggests that the Shuitangba colobine could move at ease on arboreal substrates and was also able to occasionally use terrestrial substrates. The potential affinities of this calcaneus to Mesopithecus and its positional profile most likely imply an eastward migration via forested corridors. In Shuitangba, this fossil colobine could trophically and positionally exploit a wide range of habitats successfully coexisting with resident hominoids.

Macaques dispersed out of Africa into Eurasia in the framework of a broader intercontinental faunal exchange that coincided in time with the sea level drop associated with the Messinian Salinity Crisis. They are first recorded in Europe (Italy and Spain) by the latest Miocene, being subsequently recorded all over Europe, albeit sparsely, throughout the Pliocene and Pleistocene. These fossil European macaques are attributed to several (sub)species of the extant Barbary macaque (Macaca sylvanus). In Iberia, fossil macaques are best documented by Macaca sylvanus florentina from various Early Pleistocene sites, whereas their published Pliocene record is very scarce. Here we report the oldest post-Messinian occurrence of macaques in the Iberian Peninsula, based on the description and metrical comparisons of two upper teeth (a male canine and a third molar of two different individuals) from the early Pliocene (MN14, 5.0–4.9 Ma) site of Puerto de la Cadena (Murcia, SE Spain). The male C1 is fully comparable in morphology with those of extant and fossil M. sylvanus, and larger than those of Mesopithecus. The M3, in turn, displays the typical papionin morphology that characterizes the dentally-conservative genus Macaca—thereby discounting an alternate assignment to either the extinct colobine monkey Mesopithecus or the more dentally-derived papionin Theropithecus. Dental size and proportions of the M3 further support an attribution to an extinct subspecies of M. sylvanus instead of the larger papionin Paradolichopithecus. Mostly on biochronologic grounds, the two macaque teeth from Puerto de la Cadena are here assigned to Macaca sylvanus cf. prisca, albeit tentatively, given the lack of clear-cut criteria to distinguish this subspecies from the younger Macaca sylvanus florentina. The described material represents the oldest well-dated Pliocene record of macaques in Iberia, predating the record of Paradolichopithecus by almost 1.5 million years., This work has been funded by the Generalitat de Catalunya (CERCA Program), and the Agencia Estatal de Investigación (Ministerio de Economía, Industria y Competitividad)–European Regional Development Fund of the European Union (CGL2015-68333-P, MINECO/FEDER-UE; and CGL2016-76431-P and CGL2017-82654-P, AEI/FEDER-UE). Fieldwork at Puerto de la Cadena was carried out with funding from the Fundación Séneca–Agencia de Ciencia y Tecnología de la Región de Murcia (11891/PHCS/09), and it was possible thanks to the collaboration of the Demarcación de Carreteras del Estado in Murcia (Ministerio de Fomento) and the Dirección General de Bienes Culturales of the Comunidad Autónoma de la Región de Murcia, and the Asociación Cultural Paleontológica Murciana.

New material of the Mio-Pliocene colobine Mesopithecus from the Turolian locality of Kryopigi (Greece) is described here. It includes a complete skull with the atlas attached and other dental and postcranial elements representing at least five individuals (four males and one female). The material is compared with Mesopithecus delsoni, Mesopithecus pentelicus, Mesopithecus monspessulanus and intermediate forms from more than a dozen Turolian localities of the Greco-Iranian province. These comparisons support the attribution of the Kryopigi material to M. pentelicus. The chronostratigraphic distribution of Mesopithecus species and intermediate forms suggests that the Kryopigi fauna could be dated as younger than the Perivolaki locality with M. delsoni/pentelicus (7.1–7.3 Ma, MN12) and older than the Dytiko localities with M. aff. pentelicus, M. cf. pentelicus and M. cf. monspessulanus (?middle MN13). The dimensions of the atlas are within the distribution of extant colobines. The skull shows bite-marks, probably caused by the hyaena Adcrocuta eximia.

A dentate mandible and proximal femur of Mesopithecus pentelicus Wagner, 1839 are described from the Shuitangba lignite mine in Zhaotong Prefecture, northeastern Yunnan Province, China. The remains were retrieved from sediments just below those that yielded a juvenile Lufengpithecus cranium and are dated at about ∼6.4 Ma. The mandible and proximal femur were found in close proximity and are probably of the same individual. The lower teeth are metrically and morphologically closely comparable with those of confirmed M. pentelicus from Europe, and on this basis, the specimen is assigned to this species. The anatomy of the proximal femur indicates that the Shuitangba Mesopithecus was a semiterrestrial quadruped that engaged in a range of mostly arboreal activities, including walking, climbing, and occasional leaping, with an abducted hip joint. The Shuitangba Mesopithecus is dentally typical for the genus but may have been more arboreal than previously described for M. pentelicus. M. pentelicus is well known from late Miocene (MN 11–12) sites in Europe and southwest Asia. Its estimated average rate of dispersal eastward was relatively slow, although it could have been episodically more rapid. The presence of a colobine, only slightly lower in the same section at Shuitangba that produced Lufengpithecus, is one of the only two well-documented instances of the near or actual co-occurrence of a monkey and ape in the Miocene of Eurasia. At Shuitangba, M. pentelicus occupied a freshwater-margin habitat with beavers, giant otters, swamp rabbits, and many aquatic birds. The presence of M. pentelicus in southwest China near the end of the Miocene further attests to the ecological versatility of a species long recognized as widespread and adaptable. The modern colobines of Asia, some or all of which are probable descendants of Mesopithecus, have gone on to inhabit some of the most highly seasonal and extreme habitats occupied by nonhuman primates.

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 δ18Op 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.

Extant colobine monkeys have been historically described as specialized folivores. However, reports on both their behavior and dental metrics tend to ascribe a more varied diet to them. In particular, several species, such as Pygathrix nemaeus and Rhinopithecus roxellana, are dedicated seasonal seed eaters. They use the lophs on their postcanine teeth to crack open the hard endocarp that protects some seeds. This raises the question of whether the bilophodont occlusal pattern of colobine monkeys first evolved as an adaptation to folivory or sclerocarpic foraging. Here, we assess the sclerocarpic foraging ability of the oldest European fossil colobine monkey, Mesopithecus. We use computed microtomograpy to investigate the three-dimensional (3D) dental topography and enamel thickness of upper second molars ascribed to the late Miocene species Mesopithecus pentelicus from Pikermi, Greece. We compare M. pentelicus to a sample of extant Old World monkeys encompassing a wide range of diets. Furthermore, we combine classic dietary categories such as folivory with alternative categories that score the ability to crack, grind and shear mechanically challenging food. The 3D dental topography of M. pentelicus predicts an ability to crack and grind hard foods such as seeds. This is consistent with previous results obtained from dental microwear analysis. However, its relatively thin enamel groups M. pentelicus with other folivorous cercopithecids. We interpret this as a morphological trade-off between the necessity to avoid tooth failure resulting from hard food consumption and the need to process a high amount of leafy material. Our study demonstrates that categories evaluating the cracking, grinding or shearing ability, traditional dietary categories, and dental topography combine well to make a powerful tool for the investigation of diet in extant and extinct primates.

A large sample of the Pikermi and Samos ungulates was examined by microwear analysis using a light stereomicroscope (561 extinct and 809 extant comparative specimens). The results were used to infer the dietary adaptations of individual species and to evaluate the Pikermian Biome ungulate fauna. Many of the bovids have wear consistent with mixed feeding, although a few mesodont taxa apparently enjoyed an exclusive browsing and or grazing diet. The giraffids spanned the entire dietary spectrum of browsing, mixed feeding, and grazing, but most of the three-toed horses (Hippotherium) were hypsodont grazers. The colobine monkeyMesopithecus pentelicidisplays microwear consistent with a mixed fruit and leaf diet most likely including some hard objects. Similar results were obtained from prior scanning electron microscopy microwear studies at 500 times magnification and from the light microscope method at 35 times magnification for the same species. Results show that diet can differ between species that have very similar gross tooth morphology. Our results also suggest that the Pikermian Biome was most likely a woodland mosaic that provided a diversity of opportunities for species that depended on browsing as well as species that ate grass. The grasses were most likely C3grasses that would grow in shaded areas of the woodland, glades, and margins of water. The ungulate component of the Pikermi and Samos fauna was more species-rich and more diverse in diet than the ungulates observed in modern African forests, woodlands, or savannas, yet dietarily most similar to the ungulates found in woodland elements of India and to some extent of Africa. It is unlikely that the Pikermi and Samos ungulates inhabited dense forests because we find no evidence for heavy fruit browsing. Conversely, a pure savanna is unlikely because many mixed feeders are present as well as browsers. Extant woodland African species are morphologically and trophically very similar to the African savanna species. Therefore the evolution of grazing and of hypsodont morphology for Africa may have evolved within the Plio-Pleistocene woodlands of Africa. Our results show that major dietary and morphologic ungulate evolution may take place within woodlands rather than as a consequence of species moving into savannas both during the late Miocene of Pikermi and Samos and during the Pleistocene–Recent of Central Africa.

Th e presence of the cercopithecids in the Neogene of Greece is known since the beginning of the 19th century. Th e excavations of the last 20 years increase their number in Greece. Th e main taxon of the cercopithecids is Mesopithecus Wagner, 1839 found originally in the middle Turolian (MN 12) locality of Pikermi, near Athens. Th e Pikermi Mesopithecus sample is rich and belongs to a medium-sized form, M. pentelicus Wagner, 1839. Besides the well-known M. pentelicus, two other species were recognized. Th e new species M. delsoni Bonis, Bouvrain, Geraads & Koufos, 1990, a large-sized form found in the early Turolian (MN 11) locality Ravin des Zouaves-5 of Axios Valley (Macedonia, Greece) has several diff erences from the type species. In the middle Turolian (MN 12) localities of Vathylakkos (Axios Valley) and Perivolaki (Th essaly) a large to medium-sized Mesopithecus form with “delsoni” and “pentelicus” characters was found and is referred to as M. delsoni/pentelicus. A small-sized form named M. cf. monspessulanus was recognized by a mandibular fragment in the late Turolian (MN 13) locality Dytiko-2 of Axios Valley and indicates the early appearance of the taxon at the end of Miocene. Th e rest of the material found in the late Turolian localities of Dytiko has some diff erences from the typical M. pentelicus. Moreover, Mesopithecus was traced in several late Miocene localities, indicating its wide distribution in Greece. Two other cercopithecids were also found in the Pliocene of Greece. Dolichopithecus ruscinensis Deperet, 1889 was recognized in the locality of Megalon Emvolon and in Ptolemais Basin (Macedonia, Greece); both are dated to late Ruscinian (MN 15). Th e second Pliocene cercopithecid is Paradolichopithecus arvernensis (Deperet, 1929), found in the locality of Vatera (Lesvos Island) and dated to late Pliocene. Th e stratigraphic distribution and the palaeoenvironment of these cercopithecids are also discussed.

Unlike most primates, extant cercopithecoids lack maxillary sinuses, which are pneumatic spaces in the facial skeleton lateral of the nasal cavity proper. Character state analysis of living cercopithecoids across well-supported topologies suggests that the sinus was lost at the origin of the superfamily, only to have evolved again convergently in extant macaques. Recent work has shown that a) the 'early loss' hypothesis is supported by the lack of any pneumatization in Victoriapithecus, a stem cercopithecoid, b) like extant macaques, the fossil cercopithecine Paradolichopithecus shows evidence of presence of the maxillary sinus (MS), and c) unlike extant colobines, the fossil colobine Libypithecus also possesses a maxillary sinus. To more fully assess the pattern of cercopithecoid sinus evolution, fossil taxa from both subfamilies (Colobinae, Cercopithecinae) were examined both visually and by computed tomography (CT). The observations were evaluated according to standard anatomical criteria for defining sinus spaces, and compared with data from all extant Old World monkey genera. Most taxa examined conformed to the pattern already discerned from extant cercopithecoids. Maxillary sinus absence in Theropithecus oswaldi, Mesopithecus, and Rhinocolobus is typical for all extant cercopithecids except Macaca. The fossil macaque Macaca majori possesses a well-developed maxillary sinus, as do all living species of the genus. Cercopithecoides, on the other hand, differs from all extant colobines in possessing a maxillary sinus. Thus, paranasal pneumatization has reemerged a minimum of two and possibly three times in cercopithecoids. The results suggest that maxillary sinus absence in cercopithecoids is due to suppression, rather than complete loss.

(2007). Paranasal pneumatization of two late Miocene colobines: Mesopithecus and Libypithecus (Cercopithecidae: Primates) Journal of Vertebrate Paleontology: Vol. 27, No. 3, pp. 768-771.

Remains of the cercopithecid Mesopithecus monspessulanus are relatively rare. Two previously unpublished mandibles of M. monspessulanus (housed in the Basel Naturhistorisches Museum), from the Italian locality of Villafranca d’Asti are described. These remains belong to the assemblage of the Triversa Faunal Unit, dated to the Early Villafranchian, that is to the unit MN16a (Middle Pliocene) of the European mammal biochronology. According to this recently revised biochronological attribution, Villafranca d’Asti records the last Mesopithecus occurrence in Europe. The NOW (Neogene Old World) database has been used as a basis to evaluate both the mammalian faunal and the palaeoenvironmental context at the time surrounding this last occurrence of Mesopithecus. The comparison (taxonomical composition and ungulate hypsodonty), between the Villafranca d’Asti assemblage and other Plio-Pleistocene mammal communities of Europe, shows that the extinction of Mesopithecus is related to a faunal turnover and a change toward more open landscapes during the Early–Middle Villafranchian transition. This is consistent with the “Elephant-Equus event”, that occurred in Europe at 2.5 Ma. Furthermore, the co-occurring but more terrestrial cercopithecid Macaca crossed this faunal turnover. This strengthens the assumption that the latest Mesopithecus species, M. monspessulanus, had woodland-oriented adaptations.

We report dental remains of the extinct colobine monkey Mesopithecus from the Turolian (MN13, Late Miocene, ca. 6.23 Ma) locality of Venta del Moro (Valencia, Spain). They include most of the deciduous dentition and the unerupted germs of the first molars of a single infantile individual, as well as two lower left lateral incisors from two additional individuals. On the basis of morphometric comparisons, mainly based on the Ms, these remains are attributed to the Late Miocene species Mesopithecus pentelicus. They represent a significant addition to the knowledge of the deciduous dentition of this taxon, much less well-known than the permanent dentition. Although this genus was widely distributed from the Late Miocene through the Pliocene across Europe, southwestern Asia, Pakistan, and China, until now its occurence in the Late Miocene of the Iberian Peninsula had not been documented conclusively. Hence, the reported remains considerably enlarge southwestwards the known geographic distribution of Mesopithecus. The presence of this genus at Venta del Moro must be understood within the framework of the significant faunal turnover that took place in European faunas during the latest Turolian (the second Messinian mammalian dispersal), which is further documented at this locality by the occurrence of other eastern immigrants. At the same time, the presence of M. pentelicus at this site agrees well with previous paleoenvironmental and sedimentological evidence, indicating a lacustrine depositional environment with strong hydrologic seasonality., This work has been funded by the Spanish Ministerio de Economía y Competitividad (CGL0211-28681, CGL2011-25754, CGL2011-27343, CGL2014-54373-P, and RYC-2009-04533 to D.M.A.), the Spanish Ministerio de Educación (AP2010-4579 to M.P.), and the Generalitat de Catalunya (2014 SGR 416 GRC).

We present here a study of European Neogene primate occurrences in the context of changing humidity. We studied the differences of primate localities versus non-primate localities by using the mammal communities and the ecomorphological data of the taxa present in the communities. The distribution of primates is influenced by humidity changes during the whole Neogene, and the results suggest that the primates track the changes in humidity through time. The exception to this is the Superfamily Cercopithecoidea which shows a wider range of choices in habitats. All primate localities seem to differ from non-primate localities in that the mammal community structure is more closed habitat oriented, while in non-primate localities the community structure changes towards open-habitat oriented in the late Neogene. The differences in primate and non-primate localities are stronger during the times of deep environmental change, when primates are found in their preferred habitats and non-primate localities have faunas better able to adapt to changing conditions.

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.

The genus Macaca (Primates: Cercopithecidae) originated in Africa, dispersed into Europe in the Late Miocene and resided there until the Late Pleistocene. In this contribution, we provide an overview of the evolutionary history of Macaca in Europe, putting it into context with the wider late Miocene, Pliocene and Pleistocene European monkey fossil record (also comprising Mesopithecus, Paradolichopithecus, Dolichopithecus and Theropithecus). The Pliocene and Pleistocene European Macaca fossil material is largely regarded as Macaca sylvanus, the same species as the extant Barbary macaque in North Africa. The M. sylvanus specimens found at West Runton in Norfolk (53°N) during the Middle Pleistocene are among the most northerly euprimates ever discovered. Our simple time-budget model indicates that short winter day lengths would have imposed a significant constraint on activity at such relatively high latitudes, so macaque populations in Britain may have been at the limit of their ecological tolerance. Two basic models using climatic and topographic data for the Last Interglacial and the Last Glacial Maximum alongside Middle and Late Pleistocene fossil distributions indicate that much of Europe may have been suitable habitat for macaques. The models also indicate that areas of southern Europe in the present day have a climate that could support macaque populations. However, M. sylvanus became locally extinct in the Late Pleistocene, possibly at a similar time as the straight-tusked elephant, Palaeoloxodon antiquus, and narrow-nosed rhinoceros, Stephanorhinus hemitoechus. Its extinction may be related to vegetation change or increased predation from Homo, although other factors (such as stochastic factors occurring as a result of small population sizes) cannot be ruled out. Notwithstanding the cause of extinction, the European macaque may thus be a previously overlooked member of the Late Pleistocene faunal turnover.

A newly discovered fossil monkey (AUH 1321) from the Baynunah Formation, Emirate of Abu Dhabi, United Arab Emirates, is important in a number of distinct ways. At ∼ 6.5-8.0 Ma, it represents the earliest known member of the primate subfamily Cercopithecinae found outside of Africa, and it may also be the earliest cercopithecine in the fossil record. In addition, the fossil appears to represent the earliest member of the cercopithecine tribe Cercopithecini (guenons) to be found anywhere, adding between 2 and 3.5 million y (∼ 50-70%) to the previous first-appearance datum of the crown guenon clade. It is the only guenon--fossil or extant--known outside the continent of Africa, and it is only the second fossil monkey specimen so far found in the whole of Arabia. This discovery suggests that identifiable crown guenons extend back into the Miocene epoch, thereby refuting hypotheses that they are a recent radiation first appearing in the Pliocene or Pleistocene. Finally, the new monkey is a member of a unique fauna that had dispersed from Africa and southern Asia into Arabia by this time, suggesting that the Arabian Peninsula was a potential filter for cross-continental faunal exchange. Thus, the presence of early cercopithecines on the Arabian Peninsula during the late Miocene reinforces the probability of a cercopithecoid dispersal route out of Africa through southwest Asia before Messinian dispersal routes over the Mediterranean Basin or Straits of Gibraltar.

Cercopithecid fossil remains from the post-evaporitic Messinian (5.40-5.33 Ma, MN13, latest Turolian, latest Miocene) locality of Moncucco Torinese (Tertiary Piedmont Basin, NW Italy) are described. A talus is assigned to the fossil colobine Mesopithecus pentelicus, while a proximal fragment of ulna and a male lower canine are attributed to cf. Me. pentelicus. An isolated I(2) and M3 are assigned to the papionin cf. Macaca sp., and two cercopithecid phalanges are left unassigned even to the subfamily level. The record of Mesopithecus at Moncucco Torinese agrees well with the previously-known range of this species in Italy and elsewhere in Europe, whereas that of cf. Macaca constitutes only the second occurrence of macaques in the Miocene of Eurasia. Although the co-occurrence of these two genera in a single locality had been previously reported in the Pliocene, this is the first instance in which macaques are associated with the Late Miocene M. pentelicus instead of Mesopithecus monspessulanus. The record of cf. Macaca and Mesopithecus-and especially the latter's talar morphology, similar to that of extant arboreal colobines-fits well with paleoenvironmental reconstructions of Moncucco Torinese based on the associated fauna, which indicate a humid and densely-forested environment, probably with more open and drier habitats nearby. From a paleobiogeographic viewpoint, the record of Macaca at Moncucco Torinese, together with the previously reported occurrence at Almenara-Casablanca M (Spain), supports the contention that macaques dispersed from Africa into Europe during the latest Miocene (ca. 5.9-5.3 Ma) at the same time as the sea level drop associated with the Messinian Salinity Crisis.

12 pages; International audience; 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.

The aim of this study is to describe the environments where the cercopithecid Mesopithecus was found during latest Miocene in Europe. For this purpose, we investigate the paleoecology of the herbivorous ungulate mesofauna of three very rich late Miocene fossil localities from southwestern Bulgaria: Hadjidimovo, Kalimantsi and Strumyani. While Mesopithecus has been found in the two first localities, no primate remains have yet been identified in Strumyani. Comparison between localities with and without primates using the herbivore mesofauna allows the cross-corroboration of paleoenvironmental conditions where this primate did and did not live. A multi-parameter statistical approach involving 117 equid and 345 bovid fossil dental and postcranial (phalanges, metapodia, astragali) remains from these three localities provides species to generic-level diet and locomotor habit information in order to characterize the environment in which Mesopithecus evolved. The analysis of dental mesowear indicates that the bovids were mainly mixed feeders, while coeval equids were more engaged in grazing. Meanwhile, postcranial remains show that the ungulate species from Hadjidimovo and Kalimantsi evolved in dry environments with a continuum of habitats ranging from slightly wooded areas to relatively open landscapes, whereas the Mesopithecus-free Strumyani locality was in comparison reflecting a rather contrasted mosaic of environments with predominant open and some more closed and wet areas. Environments in which Mesopithecus is known during the late Miocene were not contrasted landscapes combining open grassy areas and dense forested patches, but instead rather restricted to slightly wooded and homogeneous landscapes including a developed grassy herbaceous layer.

Mesopithecus pentelicus Wagner, 1839 HOLOTYPE. — Maxillary fragment with M1-M3 described by Wagner (1839) and figured by Wagner (1840). The specimen is housed in BSPM numbered as BSPM ASII. 11. LOCALITIES. — Classical Pikermi ravine (PIK), Attica, Greece (near Athens); Chomateres or Kisdari (CHO), Attica, Greece (near Athens). AGE. — Middle Turolian, MN 12 (late Miocene); more precisely it is referred to the uppermost MN 12 with an age of ~7.0 Ma. Th e fauna of Chomateres is considered slightly older than the Pikermi one. DIAGNOSIS. — Medium-sized colobine monkey; short, upright face; sexual dimorphism in the skull, canines and postcranials; absent or very small sagittal crest in the males; enlarged mandibular angle; shallow mandibular corpus with constant height between p4 and m3; convex anterior symphysis without symphyseal constriction; small and deeply inclined planum alveolare; absent or weak fossa genioglossa; small lingual cusp in the P3, 4; small honing facet in the p3; small hypoconulid in the m3., Published as part of Koufos, George D., 2009, The Neogene cercopithecids (Mammalia, Primates) of Greece, pp. 817-850 in Geodiversitas 31 (4) on page 839, DOI: 10.5252/g2009n4a817, http://zenodo.org/record/5381502, {"references":["WAGNER A. 1839. - Fossile Uberreste von einem Affenschadel und andern Saugethierreste aus Griechenland. Gelehrte Anzeigen Bayerische Akademie der Wissenschaften 38: 301 - 312.","WAGNER A. 1840. - Fossile Uberreste von einem Affenschadel und andern Saugethierreste aus Griechenland. Abhandlungen der Mathematisch-Physikalischen classe der Bayerischen Academie der Wissenschaften 3: 153 - 170."]}

Mesopithecus delsoni Bonis, Bouvrain, Geraads & Koufos, 1990 HOLOTYPE. — Mandible of male adult individual with both tooth rows, RZO-159. It is housed in the Laboratory of Geology and Palaeontology, University of Thessaloniki. LOCALITY. — Ravin des Zouaves-5 (RZO), Axios Valley, Macedonia, Greece. AGE. — Early Turolian, MN 11 (late Miocene); estimated magnetostratigraphic age ~8.2 Ma. DIAGNOSIS. — Large size; deep mandibular corpus; flattened anterior symphysis; strong symphyseal constriction; slightly inclined alveolar plane; large fossa genioglossa; thick inferior transverse torus; large honing facet in the p3; well-developed and bicuspid talonid in the m3., Published as part of Koufos, George D., 2009, The Neogene cercopithecids (Mammalia, Primates) of Greece, pp. 817-850 in Geodiversitas 31 (4) on page 839, DOI: 10.5252/g2009n4a817, http://zenodo.org/record/5381502, {"references":["BONIS L. DE, BOUVRAIN G., GERAADS D. & KOUFOS G. D. 1990. - New remains of Mesopithecus (Primates, Cercopithecidae) from the late Miocene of Macedonia with the description of a new species. Journal of Vertebrate Paleontology 10: 473 - 483."]}

International audience; 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.

This study aims to characterize the feeding habits of the first European colobines, Mesopithecus (late Miocene), through the analysis of its molar microwear pattern. Fifty-seven adult individuals of Mesopithecus (from Greece and Bulgaria) are compared to 162 wild-shot specimens representing nine modern species of African and Asian cercopithecids. Through the combination of a principal component analysis and analyses of variances, Mesopithecus displays significant differences with its extant colobine relatives. Actually, there is no significant difference between the Mesopithecus samples and Chlorocebus aethiops and Papio hamadryas. An intra-specific test dedicated to Mesopithecus pentelicus reveals no difference between the Bulgarian and the Greek samples. To sum up, the species of Mesopithecus were not leaf eaters like their present-day colobine relatives. Considering its wide paleogeographic range, its post-cranial anatomy, the paleoenvironmental data, and the present dietary reconstruction as ...

A cercopithecid mandible is described from the locality Megalo Emvolon in Macedonia (Greece). The main morphological characters of the mandible and teeth as well as the dimensions allow us to attribute this specimen to Dolichopithecus ruscinensis . The mandible is compared with known material from Europe. The similarity of the Megalo Emvolon specimen to the material from Perpignan indicates a late Ruscinian age (MN 15) for Megalo Emvolon. This age is also confirmed by the other faunal and stratigraphical data. Three different primate biozones in Macedonia are discussed: the Ouranopithecus biozone of the Vallesian, the Mesopithecus biozone of the Turolian and the Dolichopithecus biozone of the Ruscinian.

We use new data on the timing and extent of the early Pleistocene dispersal of Homo erectus to estimate diffusion coefficients of early Homo from Africa. These diffusion coefficients indicate more rapid and efficient dispersals than those calculated for fossil Macaca sp., Theropithecus darti, and Mesopithecus pentelicus. Increases in home range size associated with changes in ecology, hominid body size, and possibly foraging strategy may underlay these differences in dispersal efficiency. Ecological data for extant primates and human foragers indicate a close relationship between body size, home range size, and diet quality. These data predict that evolutionary changes in body size and foraging behavior would have produced a 10-fold increase in the home range size of H. erectus compared with that of the australopithecines. These two independent datasets provide a means of quantifying aspects of the dispersal of early Homo and suggest that rapid rates of dispersal appear to have been promoted by changes in foraging strategy and body size in H. erectus facilitated by changes in ecosystem structure during the Plio-Pleistocene.

This paper reports on new finds and re-discovered old collections of the latest Miocene colobine Mesopithecus from Italian localities. New finds are reported from the Baccinello V3 faunal assemblage and from the Monticino gypsum quarry (Brisighella) and newly rediscovered teeth from the Casino basin are herein described. The latter two samples are attributed to the species Mesopithecus pentelicus, while the allocation of the Baccinello V3 sample is unclear and for the moment is attributed to Mesopithecus sp. indet. A fourth Italian locality that yielded Mesopithecus remains is Gravitelli. Unfortunately the latter specimens are lost, and from the literature it is impossible to furnish an accurate specific attribution. The taxonomic allocation of latest Miocene Mesopithecus species is briefly discussed.

The phyletic relationships of cercopithecid higher taxa are fairly well understood. By reference to the outgroup Hominoidea, character state distributions for a variety of dental, cranial and posteranial features are reviewed and morphotypes are presented for supra-generic taxa. Cercopithecids show 13 derived conditions relative to the eucatarrhine morphotype. Cercopithecines are further derived by eight features while the colobine morphotype shows an additional nine synapomorphies, including three newly identified pedal features. Within Cercopithecinae, the tribes Papionini and Cercopithecini are each characterized by several derived states. In turn, each tribe includes a relatively conservative and a more derived clade, recognized at the subtribal level: Macacina and Papionina (including the previously recognized Theropithecina) for Papionini: and the newly defined Allenopithecina and Cercopithecina among Cercopithecini. The distinctions within Colobinae appear comparable to those defining cercopithecine subtribes, and two taxa are accepted at that rank: the African Colobina, defined by three derived states: and the possibly paraphyletic Asian Presbytina. Piliocolobus shares four synapomorphies with Procolobus and therefore is ranked as a subgenus of that taxon. The hypothesis that Dolichopithecus is the direct descendant of Mesopithecus is weakened on the basis of contrary pedal evidence.

UNDER this poetic title Prof. Gaudry has added one more to his valuable works on the fossil mammalia of the Tertiary Period. Some sixteen years ago he carried on the explorations on the classic ground of Pikermi, which proved that the plains of Marathon were haunted by antelopes, gazelles, giraffes, and hipparions, in the upper miocene age, while the forests then overshadowing rocky Attica sheltered the mastodon and rhinoceros, and yielded food to innumerable monkeys (Mesopithecus) allied to the Barbary ape. During the last five years he has been painting a like picture of the upper miocene mammalia of Mont Leberon, in which, as in Pikermi, the remains of the animals are preserved in an abundance and a perfection somewhat like those discovered in the Western States, by Prof. Marsh. The two elaborate quartos on these discoveries are now followed by an elegant octavo full of woodcuts, treating of the links that bind together some of the more important groups of tertiary and living mammalia. In it the author deals with the European mammalia without treating exhaustively the accumulation of facts brought to light by Marsh, Hayden, Leidy, and others, in the American eocenes and miocenes. Norare these as yet published in sufficient detail to allow of their being worked into the book before us. The impression made upon my mind by the vast stores of remains in the museum at Yale is one of profound melancholy; for so numerous and varied are the mammalia that many years must elapse before they can be brought into complete relation with those of Europe. Up to the present time the fragmentary notices which have appeared bear the same relation to the systematic treatment to be expected from Prof. Marsh, that isolated tesserae bear to the whole design of a mosaic. Prof. Gaudry, therefore, has acted well in confining his work mainly to the Tertiary mammalia of Europe. Les Enchainements du Monde Animal dans les Temps Geologiques, Mammijeres, Tertiares. Par Albert Gaudry. 8vo. (Paris: F. Savy, 1878.)