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).     

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.     

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.     

Calcaneal features of the Greek Miocene primate Mesopithecus pentelicus (Cercopithecoidea: Colobinae)

This paper investigates substrate preferences of the Greek Colobine Mesopithecus pentelicus WAGNER, from the Miocene of Pikermi, by examining selected functional features of the calcaneus that distinguish between arboreal and terrestrial Cercopithecidae. Mesopithecus possesses a relatively long proximal calcaneal region associated with a slightly low and wide surface for the insertion of m. triceps surae. These features approximate that of semi-terrestrial Cercopithecidae and would suggest terrestrial cursorial activities. On the other hand, the relatively long and narrow proximal calcaneo-astragalar facet, similar to that of most arboreal species, would provide ampler subtalar movements. The mosaic of these features implies a semi-terrestrial way of life and conforms to the savanna-woodland paleoenvironment of Pikermi, Greece. These features appear to be well associated with foot function and change of habitus and are used to examine scenarios of the evolutionary history of the Colobinae.     

<Emphasis Type="Italic">Mesopithecus</Emphasis> (Primates,Cercopithecidae) from the turolian locality of Vathylakkos 2 (Macedonia,Greece)

Two skulls, two mandibles and some postcranials ofMesopithecus from the locality “Vathylakkos 2” of Axios Valley (Macedonia, Greece) enlarge our knowledge about the genus and its differentiation. The new material together with an old skull from the same locality is compared with the Pikermi, “Ravin des Zouaves 5” (Axios Valley), and Maramena (Serres basin) samples. It is concluded that the VathylakkosMesopithecus resemblesM. pentelicus from Pikermi, as wellM. delsoni from “Ravin des Zouaves 5” and it is referred asMesopithecus sp. aff.M. pentelicus. The biochronological age of “Vathylakkos 2” has been considered as MN 12, while recent magnetostratigraphic data indicate an age of about 7.5 Ma. This age confirms the position of the VathylakkosMesopithecus between Pikermi and “Ravin des Zouaves 5”. Some dental indices compared with those of the RecentCercopithecus suggest that the Vathylakkos sample is monospecific.      

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.     

Plio-Pleistocene cercopithecids from Kanam East, western Kenya

This paper describes cercopithecid craniodental and postcranial fossils recovered by L. S. B. Leakey at Kanam East, Kenya during the early 1930s. These fossil monkeys have been generally assumed to have been derived from early Pliocene horizons, but their exact geographical and stratigraphical provenience is unknown. Although the question of the evolutionary significance of these specimens must await the recovery of more securely dated material from Kanam East, some general conclusions can be drawn concerning their taxonomic affinities and paleobiology. Based on comparative studies of the craniodental material, at least three extant genera are represented—Colobus,Lophocebus, andCercopithecus. The postcranial fossils include a number of hindlimb specimens, as well as the manubrium of a sternum and a caudal vertebra. Identification of the postcranial remains to particular genera is not possible, but they are similar in morphology to modern arboreal and semiterrestrial cercopithecid monkeys of small to medium size. It is evident that Kanam East had a diverse cercopithecid community, similar to those found today in forested and woodland habitats, and this may be of some significance in reconstructing the paleoecology of the site. Because the fossil record of most extant cercopithecid genera is rather sparse at Plio-Pleistocene sites in Africa, Kanam East represents one of only a few sites that has yielded material that can be assigned toColobus,Lophocebus, orCercopithecus. The fossil monkeys from the site, therefore, provide additional evidence to help reconstruct the paleobiology, as well as the patterns of species diversity and community structure that characterized the cercopithecid radiation during the Plio-Pleistocene.     

Victoriapithecus: The key to Old World monkey and catarrhine origins

The past ten years have witnessed major changes in reconstructions of the history of Old World monkeys, most of them driven by new material of the Miocene monkey Victoriapithecus from Maboko Island, Kenya. Before the mid-1980s, predictions about the morphological and ecological adaptations of the earliest cercopithecoids relied heavily on evidence from extant colobine and cercopithecine monkeys. It was argued that the earliest cercopithecoids were largely or at least partly folivorous, had short colobine-like faces, and were arboreal. The only studies suggesting that some of these arguments were not true were based on limited knowledge of the anatomy of Victoriapithecus. The presence of semi-terrestrial adaptations in middle Miocene monkeys hinted to some that early monkeys may not have been arboreal. Others attempted to cope with the discrepancy between neontological predictions and the fossil evidence by proposing that limb bones with stronger terrestrial adaptations within the Maboko sample were derived cercopithecine remains, while those with more arboreal features belonged in the subfamily Colobinae and should be regarded as primitive.     

Proboscideans from the upper Miocene localities of Thermopigi,Neokaisareia and Platania (Northern Greece)

In the present article, we study the proboscidean remains from three upper Miocene localities of Northern Greece: Thermopigi (Serres), Neokaisareia (Pieria) and Platania (Drama). The material from the Turolian locality of Thermopigi includes only postcranial specimens. The morphological features of the scapula indicate the presence of the deinotheriid Deinotherium sp., whereas the rest of the specimens are morphologically distinct from Deinotherium and can be referred to Elephantimorpha indet. The material from Neokaisareia consists of a partial skeleton of a single individual and is attributed to the mammutid Mammut sp. (M. obliquelophus?). This taxon is known in Greece from the early–middle Turolian. The Platania proboscidean belongs to the tetralophodont amebelodontid Konobelodon cf. atticus. The genus Konobelodon was already present during the Vallesian of the wider area, but the lower tusk of the Platania shovel-tusker presents some morphological and metrical differences from the Vallesian representative, yet it has also smaller dimensions in its deciduous dentition than the morphologically similar Turolian specimens. The type locality of K. atticus is Pikermi (Attica, Greece), correlated to the middle Turolian, but the known biostratigraphic range of this species covers the entire Turolian. Platania is possibly correlated close to the Vallesian/Turolian boundary and the possible record of this species could document one of its earliest occurrences.     

Positional behavior of free-ranging Japanese macaques (<Emphasis Type="Italic">Macaca fuscata</Emphasis>)

Positional behavior was quantitatively studied in identified free-ranging Japanese macaques (Macaca fuscata). Five male and 11 female adults were observed in a forested mountain habitat. Data were analyzed for proportion of bout distance, number and time of each locomotion and postural type. Japanese macaques are semiterrestrial, and mainly walk and run quadrupedally. This supports the notion that Macaca are generally quadrupeds. Sex differences in positional behavior were found in the preference of substrate and types of positional behavior. Males and females tend to be terrestrial and arboreal, respectively. Males leap more frequently and longer in distance than do females when they are feeding in trees. These sex differences are considered to be related to differences in morphology, food choice, social activity, and the nursing of infants. Frequencies of leaping and the distance covered by leaping in Japanese macaques are more than those of long-tailed macaques which are arboreal quadrupeds. However, Japanese macaques leap shorter distances at a time than do long-tailed macaques, which indicates that body size may be related to leaping distance more than the frequency of leaping and the distance covered by leaping. Japanese macaques are not as specialized for terrestrial locomotion as pig-tailed macaques. They use both terrestrial and arboreal supports, and are considered to be semi-terrestrial quadrupeds, somewhere between the arboreal long-tailed macaque and the terrestrial pig-tailed macaque. Electronic Publication     

The late Miocene bovids from Platania (Drama Basin,Greece), with description of a new species of Palaeoryx

This study presents the new fossil material of bovids from the recently discovered upper Miocene locality of Platania, Drama (Greece). The material was excavated from 2012 to 2016 and yielded approximately 760 specimens attributed to hipparions, rhinos, cervids, giraffids, suids, proboscideans, hyaenids, and turtles. The bovid material described here includes six taxa of Antilopinae and one of Bovinae. Antilopines are represented by Gazella sp., Gazella cf. ancyrensis, Prostrepsiceros aff. syridisi, cf. Palaeoreas, Palaeoryx minor nov. sp., and Tragoreas? aff. oryxoides. Bovines are recorded by a single boselaphine attributed to Miotragocerus sp. Departing from other Palaeoryx species, the new species P. minor has rather straight and weakly divergent horn-cores tilted backwards, obtuse facial and occipito-parietal angles, and smaller cranial and horn-core size, though associated with a proportionally large toothrow. Miotragocerus sp. from Platania seems to be conspecific to the Miotragocerus sp. from the end-Vallesian Nikiti-1 fauna (Greece). The bovid assemblage of Platania shows a mix of both Vallesian and Turolian taxa indicating a likely late Vallesian-early Turolian age.     

A new terrestrial amphipod from tertiary amber deposits of Chiapas province,Southern Mexico

Transitroides morsei new genus, new species, is described from late Oligocene Early Miocene amber deposits of Chiapas Province from southern Mexico. This new taxon is the first known fossil member of the amphipod family Talitridae and superfamily Talitroidea. It appears intermediate between regional extant palustral (salt‐marsh) genera and fully terrestrial landhoppers of the genus Caribitroides. These beetle‐like crustaceans form a significant part of the forest‐floor leaf‐litter communities in some tropical parts of the globe. Extant terrestrial amphipods occur in south‐central Mexico and adjacent rain forest habitats in Central America and the Caribbean. Their occasional use of arboreal habitats in search for decaying organic matter, which serves as food, explains their occurrence in fossilized resin.     

Dental Eruption Sequences in Fossil Colobines and the Evolution of Primate Life Histories

Unlike other catarrhines, colobines show early molar eruption relative to that of the anterior dentition. The pattern is variable, with Asian genera (Presbytina) showing a greater variability than the African genera (Colobina). The polarity of early relative molar eruption, as well as the degree to which it is related to phylogeny, are unclear. Schultz (1935) suggested that the trend reflects phylogeny and is primitive for catarrhines. More recently, however, researchers have proposed that life history and dietary hypotheses account for early relative molar eruption. If the colobine eruption pattern is primitive for catarrhines, it implies that cercopithecines and hominoids converged on delayed relative molar eruption. Alternatively, if the colobine condition is derived, factors such as diet and mortality patterns probably shaped colobine eruption patterns. Here we update our knowledge on eruption sequences of living colobines, and explore the evolutionary history of the colobine dental eruption pattern by examining fossil colobine taxa from Eurasia (Mesopithecus) and Africa (Kuseracolobus aramisi and Colobus sp.) and the basal cercopithecoid Victoriapithecus macinnesi. We scored specimens per Harvati (2000). The Late Miocene-Early Pliocene Mesopithecus erupts the second molar early relative to the incisors, while the Early Pliocene Kuseracolobus aramisi does not. These results demonstrate that the common colobine tendency for early molar eruption relative to the anterior dentition had appeared by the Late Miocene, and that some of the diversity observed among living colobines was already established in the Late Miocene/Early Pliocene. We discuss the implications of these results for phylogenetic, life history, and dietary hypotheses of dental development.     

Miocene cercopithecoidea from the Tugen Hills, Kenya

Miocene to Pleistocene fossiliferous sediments in the Tugen Hills span the time period from at least 15.5 Ma to 0.25 Ma, including time periods unknown or little known elsewhere in Africa. Consequently, the Tugen Hills deposits hold the potential to inform us about crucial phylogenetic events in African faunal evolution and about long-term environmental change. Among the specimens collected from this region are a number of discoveries already important to the understanding of primate evolution. Here, we describe additional cercopithecoid material from the Miocene deposits in the Tugen Hills sequence, including those from securely dated sites in the Muruyur Beds (16-13.4 Ma), the Mpesida Beds (7-6.2 Ma) and the Lukeino Formation (∼6.2-5.7 Ma). We also evaluate previously described material from the Ngorora Formation (13-8.8 Ma). Identified taxa include Victoriapithecidae gen. et sp. indet., cf. Parapapio lothagamensis, and at least two colobines. Specimens attributed to cf. Pp. lothagamensis would extend the species’ geographic range beyond its type locality. In addition, we describe specimens sharing derived characters with modern African colobines (Tribe: Colobina), a finding that is congruent with previous molecular estimates of colobine divergence dates. These colobine specimens represent some of the earliest known members of the modern African colobine radiation and, in contrast to previous hypotheses, suggest that early African colobines were mainly arboreal and that semi-terrestrial Late Miocene and Plio-Pleistocene colobine taxa were secondarily derived in their locomotor adaptations.     

福建漳浦中中新世几种藓类植物研究*

亚洲新生代藓类植物化石非常稀少,漳浦琥珀是目前已知的我国低纬度地区唯一含苔藓植物化石的新生代琥珀矿床。漳浦琥珀源自中中新世气候适宜期(~14-17 Ma)的龙脑香树脂,温暖潮湿的热带季雨林气候和广泛分布的龙脑香科植物,为藓类植物的生长、多样性演化和三维立体保存提供了条件。本研究共报道了漳浦县中中新世藓类植物化石3个属种,包括:白发藓属Leucobryum Hampe、牛舌藓属Anomodon Hook.et Taylor和扁枝藓属Homalia(Brid.)B.S.G.。首次利用三维X射线断层扫描技术(Micro-CT)研究了白发藓属化石,发现其与现生种短枝白发藓Leucobryum humillimum Cardot非常相似,因此将其命名为短枝白发藓(相似种)Leucobryum cf.humillimum。此外,当前牛舌藓属Anomodon和扁枝藓属Homalia化石是东亚地区的首例记录。漳浦琥珀中的藓类植物化石揭示了中中新世时期苔藓植物的多样性,丰富了东亚低纬度地区新生代苔藓植物的化石记录。     

Diversification of South American spiny rats (Echimyidae): a multigene phylogenetic approach

Fabre, P.‐H., Galewski, T., Tilak, M.‐k. & Douzery, E.J.P. (2012) Diversification of South American spiny rats (Echimyidae): a multigene phylogenetic approach. —Zoologica Scripta, 00, 000–000. We investigated the phylogenetic relationships of 14 Echimyidae (spiny rats), one Myocastoridae (nutrias) and one Capromyidae (hutias) genera based on three newly sequenced nuclear genes (APOB, GHR and RBP3) and five previously published markers (the nuclear RAG1 and vWF, and the mitochondrial cytochrome b, 12S rRNA and 16S rRNA). We recovered a well‐supported phylogeny within the Echimyidae, although the evolutionary relationships among arboreal echimyid taxa remain unresolved. Molecular divergence times estimated using a Bayesian relaxed molecular clock suggest a Middle Miocene origin for most of the extant echimyid genera. Echimyidae seems to constitute an example of evolutionary radiation with high species diversity, yet they exhibit only narrow skull morphological changes, and the arboreal and terrestrial taxa are shown to retain numerous plesiomorphic features. The most recent common ancestor of spiny rats is inferred to be a ground‐dwelling taxon that has subsequently diverged into fossorial, semiaquatic and arboreal habitats. The arboreal clade polytomy and ancestral character estimations suggest that the colonization of the arboreal niche constituted the keystone event of the echimyid radiation. However, biogeographical patterns suggest a strong influence of allopatric speciation in addition to ecology‐driven diversification among South American spiny rats.