Metailurus major Zdansky, 1924 (Carnivora,Mammalia) from the classical locality of Pikermi (Attica,Greece)

The species of Metailurus major has a large geographical extension and is known from localities spread out from West Europe to China. In Greece it is known from the Late Miocene locality of Halmyropotamos in Euboea, while some authors mention it also in Pikermi and Samos in faunal lists only. Two specimens from Pikermi prove its existence in this classical locality.     

An almost complete skeleton of Metailurus parvulus (Carnivora, Felidae) from the late Miocene of Kerassia (Northern Euboea, Greece)

We describe a partial skeleton of Metailurus parvulus from the Turolian site of Kerassia 1 (Northern Euboea, Greece). The material, which consists of a mandible, the anterior and posterior limb-bone elements, some sternal bones and some vertebrae, is the most complete known of this species. The dental material is compared to specimens from Pikermi and Chomateri (Greece), and China. The limb-bones available offer us the possibility to discuss the status of some previously described specimens from Pikermi. The limb proportions indicate that M. parvulus had elongated posterior limbs relative to the anterior ones, which reflects developed jumping skills. M. parvulus had moderately developed cursorial abilities, intermediate between open and closed habitat felids, and probably frequented primarily relatively open woodlands.     

Late Miocene (Turolian,MN13) squirrels from Moncucco Torinese,NW Italy

Squirrel remains are rare in the fossil record, which hampers a properly documented interpretation of their evolutionary history. The Late Miocene sciurids from Moncucco Torinese (NW Italy) are described based on a diverse tooth assemblage. The data presented herein confirm the presence of Sciurus warthae and Pliopetaurista pliocaenica, and also include the first unquestionable occurrence of Hylopetes hungaricus plus an indeterminate Sciurinae. The report of Sciurus warthae from Moncucco Torinese represents the earliest record of the genus Sciurus in Europe. Moreover, Moncucco Torinese is the first European locality recording the co-occurrence of Sciurini and Pteromyini in the Late Miocene. Comparative analyses support a close relationship between S. warthae and Sciurus vulgaris corroborating the hypothesis of an earliest divergence of the “Sciurus vulgaris clade” in the Late Miocene. Biochronological, paleobiogeographical and paleoecological implications are also discussed.     

Musteloids and feloids (Mammalia, Carnivora) from the late Miocene locality of Pikermi (Attica, Greece)

Some relatively rare carnivore remains from Pikermi are studied. The present specimens include Simocyon primigenius(ROTH and WAGNER, 1854), Promeles palaeatticus(WEITHOFER, 1888), Martes woodwardiPILGRIM, 1931, Felis atticaWAGNER, 1857 and Machairodus giganteus(WAGNER, 1848), as well as some postcranial remains that resemble Enhydriodon? latipesPILGRIM, 1931 and are assigned to Mustelidae gen. et spec. indet. Some of these species were not known until now but from their holotypes as M. woodwardi and E.? latipes. Other species such as F. attica and M. giganteus are better known but from relatively scant material. Since the material of the earlier mentioned species comes from Pikermi, their type locality, it is of especial importance and improves our knowledge for the Pikermi carnivores and their relationships to other species. In some cases where the material comes from ossiferous blocks, the accompanying faunal context is discussed.     

Sexual and individual morphometric variation in Libycosaurus (Mammalia, Anthracotheriidae) from the Maghreb and Libya

Anthracotheres of late Middle Miocene and Late Miocene age have been described from several localities in northern Africa, all of them currently assigned to the genus Libycosaurus Bonarelli, although in several previous works they were assigned to Merycopotamus Falconer and Cautley, a considerably younger and specialised form from the Indian subcontinent which has quite different dental and cranial morphology. Three species of Libycosaurus have been named, but there has been some doubt about the morphometric variation within the various species, with some authors such as Gaziry (1987) placing the fossils from Sahabi (Latest Miocene, Libya) and Beglia (end Middle Miocene to basal Late Miocene, Tunisia) into the same species despite marked size differences, and others (Ducrocq et al., 2001) creating a species for a restricted sample of small specimens from Nementcha (Late Middle Miocene, Algeria), but which overlaps with the range of size variation of Beglia fossils. The aim of this paper is to examine the available samples in greater depth in order to understand the morphometric variation in these anthracotheres. It is confirmed that the Beglia sample is quite variable (Black, 1972), both morphologically and metrically, but it is concluded that it nevertheless belongs to a single species, because specimens from some of the localities within the Beglia Formation (e.g. Loc. 17 in the lower levels at Beglia) span the entire range of variation (Pickford, 1994). The sample from Nementcha cannot be distinguished from the Beglia sample on any consistent metric or morphological basis, but in general the specimens fall at the low end of the range of variation of the Beglia sample. It is thus likely that L. algeriensis is a synonym of L. anisae. The Sahabi and Chad samples (L. petrocchii), in contrast, fall above the known range of variation of the Beglia material in almost all metric features, but are close to it morphologically, and they are considered to represent a species distinct from the Beglia sample.     

The earliest European badger (Meles meles), from the Late Villafranchian site of Fuente Nueva 3 (Orce, Granada, SE Iberian Peninsula)

Paleontological and molecular data suggest that the divergence of the European and Asian badgers (Meles meles and Meles leucurus, respectively) from their ancestor Meles thorali might have taken place in the Middle to Late Villafranchian boundary (ca. 1.8 Ma). However, the available record of Late Villafrachian European badgers is scanty and poorly known, and this hypothesis needs more paleontological data in order to be tested. The badger hemimandible from the Iberian locality of Fuente Nueva 3, a locality placed in the beginning of the Late Villafranchian, supports the idea of an early divergence between European and Asian badgers.     

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.     

Early Miocene cricetids (Rodentia) from the Junggar basin (Xinjiang, China) and their biochronological implications

Among the 14 small mammal species from the early Miocene locality of the northern area of the Junggar basin (northern Xinjiang, China), four species are cricetids (Rodentia): unnamed species of Cricetodon and Eumyarion, and two new species, Karydomys debruijni nov. sp. and Megacricetodon beijiangensis nov. sp. Some aspects of the morphology of Cricetodon sp. are shared by Eucricetodon from the late Oligocene, suggesting that these specimens could be of intermediate form between Eucricetodon and Cricetodon. One tooth of Eumyarion sp. was found, making its determination uncertain, but its morphology is clearly differentiated from the one of Cricetodon sp. The species K. debruijni nov. sp. is established based on its primitive features compared to the species known in Europe and Anatolia, and its specific association of characters compared to Karydomys dzerzhinskii. M. beijiangensis nov. sp. shows many plesiomorphic features compared to the species already described in the middle Miocene of China. Based on both the whole assemblage of rodents and the species of cricetids, the biochronologic position and the age of the locality are discussed. The locality appears to be biochronologically very close to the fauna from the Chul’adyr Formation in Aktau Mountains, but we propose an age slightly older than the one proposed for this fauna, probably equivalent to the MN3 biozone in Europe.     

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.     

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

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

Carnivores from the early/middle Miocene locality of Antonios (Chalkidiki, Macedonia, Greece)

The mammal locality of Antonios is one of the few known early-middle Miocene ones of Greece with large mammals. It is situated in Chalkidiki Peninsula (Macedonia, Greece) and includes both small and large mammals. The study of the carnivores found in Antonios indicates the presence of the following taxa: Proputorius cf. P. sansaniensis, Protictitherium gaillardi, Protictitherium cf. P. crassum, Percrocuta sp., Percrocutidae indet. and Pseusaelurus romieviensis. The material of each taxon is described, compared with other material from various Eurasian localities and determined. The species P. cf. sansaniensis and P. romieviensis are traced for first time in Eastern Mediterranean, while Percrocuta for first time in Greece. The presence of the other carnivoran taxa in the Antonios fauna is also interesting as they are recognized in older stratigraphic levels enriching our knowledge about their stratigraphic distribution. The carnivores of Antonios cannot offer significant data for the dating of the fauna but its age is discussed using the data coming from the carnivores as well as those from older studies of the micromammals and suoids.     

Ancylotherium pentelicum (Perissodactyla, Mammalia) from the Upper Miocene of central and western Turkey

We report here new fossils of A. pentelicum from two Turolian localities of central and western Turkey, Pinaryaka and Salihpasalar (Kemikalan). The material, which consists of well preserved distal limb-bones, is compared with the few previously described specimens, mainly from Samos and Pikermi, but the observed differences should probably be attributed to individual variation. There is no evidence of evolution of the species during the Middle Turolian of the eastern Mediterranean, where it filled a narrow ecological niche, before its extinction.     

A Late Miocene Paratethyan mollusc fauna from the Denizli Basin (southwestern Anatolia, Turkey) and its regional palaeobiogeographic implications

We report on a Late Miocene mollusc fauna from a single locality in the Denizli Basin in southwestern Turkey that is composed of ten presumably euryhaline species and three freshwater species. The fauna is remarkably distinct from faunas of the adjacent Late Miocene Aegean and Euxinian Basins, and has elements in common with the modern Caspian Sea fauna in the form of Didacna species. The suggested age of the fauna (Late Miocene) would extend the stratigraphic range of the lymnocardiid genus Didacna (hitherto Calabrian-Extant) considerably. The palaeobiogeographic significance of the Denizli fauna is discussed.     

The identity of Tetracyclus constrictus (Bacillariophyceae) and a description of Tetracyclusbradburyii, a new miocene species

The siliceous valves from species of Tetracyclus (Bacillariophyceae) have variable but characteristic shapes. In this paper, species characterised by having valves with a central constriction are discussed. This distinctive shape occurs during only a period of its lifecycle. Tetracyclus constrictus and Tetracyclus celatom have been applied to fossil specimens occurring in deposits from the west coast of the USA and the east coast of Russia. Do these names apply to two closely related species or are they one and the same taxon? Two further species, Tetracyclus javanicus and Tetracyclus peragalli, have been involved in the identity of T. constrictus–T. celatom. T. constrictus M. Peragallo is the correct name and author for specimens of the extinct Miocene, circum-Pacific (or northern) fossil species with valves panduriform with rounded poles; T. javanicus refers to a species found living on Java only; fossil specimens usually referred to T. javanicus (Java) belong to a new taxon, Tetracyclus bradburyii (USA); T. peragalli is an extinct Miocene fossil, with a northern hemisphere distribution; small (≤10 μm) circular/elliptical fossil specimens, found alongside other specimens, are difficult (if not impossible) to determine to species level.     

Earliest occurrence of Puma pardoides (Owen, 1846) (Carnivora,Felidae) at the Plio/Pleistocene transition in western Europe: New evidence from the Middle Villafranchian assemblage of Montopoli,Italy

The puma-like cat Puma pardoides (Owen, 1846) is reported from several Eurasian localities dated to the Late Pliocene–Early Pleistocene, but its Italian fossil record is really poor; in fact, the only possible occurrence comprises some fragmentary postcranial material from the Late Villafranchian locality of Pirro Nord (1.6–1.3 Ma). In the present paper, we describe an isolated left upper carnassial of a medium-sized felid belonging to the collection of fossil mammals from Montopoli (Tuscany, Italy), the type locality of the Middle Villafranchian Faunal Unit (∼2.6 Ma). All the morphological and morphometric characters of the tooth are consistent with an attribution to Puma pardoides (Owen, 1846), which comes out as the second felid species from Montopoli together with Acinonyx pardinensis (Croizet et Jobert, 1828) and is, possibly with the specimens from Perrier-Étouaries, the earliest occurrence in western Europe.     

Les Megacricetodon du gisement karstique miocène moyen de Blanquatère 1 (Pyrénées-Orientales, Sud de la France) : nouvelles espèces, implications biochronologique et phylogénique

Three Megacricetodon species are recognized in the Middle Miocene locality of Blanquatère 1 (Languedoc-Roussillon province, France). Two are new, the small-sized M. tautavelensis nov. sp. the medium to large-sized M. aunayi nov. sp., the third one being the already known M. “collongensis-gersii”, a medium-sized species found in other Miocene localities of the area. Size and morphology differentiate these species. Such a high number of contemporaneous species of the genus Megacricetodon is found for the first time in a Miocene locality in the region and indicate that the evolution of this genus in western Europe is likely more complicated. It may testify both of a biogeographical differentiation and of migration events. According to our calibration charts, the deposit is dated ca. 16 Ma ago. The position of Vieux-Collonges (MN 5) and Sansan (MN 6) on the biochronological scale is also discussed, as well as the estimated age of the biozonal boundaries MN 3/MN 4, MN 4/MN 5 and MN 5/MN 6.     

New material of Chalicotheriidae (Perissodactyla,Mammalia) from the Late Miocene of Axios Valley,Macedonia (Greece) with the description of a new species

Chalicotheriids are rare in the late Miocene mammal localities of Axios Valley, Macedonia (Greece). The new campaign of excavations, since 1972, has provided some specimens, which are studied in this article. They are coming from two different localities. The late early Vallesian locality of Pentalophos 1 (PNT) has provided a skull and a mandible of an Ancylotherium. The morphological characters of the PNT material as the small size, the long snout, the shallow mandibular corpus, the strong cingulum in the teeth, the short tooth rows and the short M3/m3 indicate that it differs from the known Turolian species A. pentelicum and allow the erection of a new species, named Ancylotherium hellenicum n. sp., which can be used as a biostratigraphic marker of the Vallesian. The middle Turolian locality Prochoma 1 (PXM) has provided only one M3, which is determined to the chalicotheriine Anisodon macedonicus. This species was earlier described from the middle Turolian locality Vathylakkos 3 (VAT) and the late Turolian one of Dytiko 3 (DKO) of Axios Valley. The biogeography and biostratigraphy of the late Miocene chalicotheres of the Greco-Iranian Palaeoprovince (GRIP), as well as their palaeoecology are also discussed. The common chalicothere of GRIP is A. pentelicum, expanded from the Balkans to Afganistan and ranging stratigraphically from the early to the late Turolian. Chalicotherium goldfussi is certainly present in GRIP and it also ranges from the early to the late Turolian; its possible Vallesian occurrence needs confirmation. The other two late Miocene chalicotheres of GRIP A. macedonicus and Kalimantsia bulgarica are restricted to the Turolian of the Balkan Peninsula.     

New cranial material of Shansirhinus (Rhinocerotidae, Perissodactyla) from the Lower Pliocene of the Linxia Basin in Gansu, China

A well-preserved skull and articulated mandible of Shansirhinus, a horned aceratherine rhinocerotid, is described from an Early Pliocene (Gaozhuangian; approximately 5.3-4.34 Ma) locality of the Linxia Basin in Gansu, China. Comparing the new material of Shansirhinus ringstromi Kretzoi, 1942, we confirm the synonymy proposed previously for the species Chilotherium yunnanensis Tang et al., 1974, Chilotherium cornutum Qiu and Yan, 1982, and Ch. tianzhuensis Zheng, 1982. New knowledge concerning its cranial and mandibular morphology allows insight into its phylogenetic position among aceratherine rhinocerotids. A sister group relationship between Shansirhinus and Chilotherium is proposed on the basis of cranial, mandibular and dental evidence. Shared derived characters that support this relationship include: an expanded mandibular symphysis with a concave ventral surface; retracted premaxillae lacking upper incisors; a robust and right-angled facial crest; a flat or slightly concave dorsal skull profile; a weak or absent parastyle fold; and a constricted protocone on the premolars. Shansirhinus was probably a grazer, which is evidenced by the high crown, strong wear, well-developed secondary folds, and enamel plications on its teeth. The age of S. ringstromi is Late Miocene to the Early Pliocene, corresponding to the Turolian to Ruscinian, MN12-MN15 of Europe. S. ringstromi is likely the ancestor of the more advanced S. brancoi (Schlosser, 1903).     

Pseudomeriones megistos nov. sp. (Gerbillinae, Mammalia) from the Latest Miocene of Northern Greece and its phylogenetic relationships

The study of the primitive gerbil remains of Pseudomeriones, recovered from the micromammalian sites of Paliambela-B and Monasteri (Latest Miocene, Northern Greece), gives evidence for a new species, which is characterized by its large size and simple molar morphology. A cladistic analysis based on dental characters affiliates this form with P. rhodius and P. pythagorasi and enables an intrageneric phylogenetic scenario.