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.     

Prostrepsiceros and Protragelaphus (Artiodactyla,Mammalia) from the Late Miocene locality of Chomateri (Attica,Greece)

The Late Miocene fossiliferous locality of Chomateri is located close to the classic locality of Pikermi, but unfortunately its faunal context is not sufficiently known. Some fossil remains of spiral-horned antelopes from Chomateri are assigned to Prostrepsiceros rotundicornis and Protragelaphus skouzesi. The co-existence of P. rotundicornis and P. skouzesi is relatively rare in the Late Miocene, but well-documented in Pikermi. Comparisons of the Chomateri material support a middle Turolian age for the Chomateri locality and suggest a close chronological relation to the Pikermi fauna.     

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.     

Dental evolution in Neophanomys (Rodentia, Octodontidae) from the late Miocene of central Argentina

The evolutionary pattern of the molar morphology of the small caviomorph (Octodontidae) Neophanomys from the late Miocene Cerro Azul Formation of central Argentina is analyzed. Two new species (chronomorphs) are recognized, which constitute an anagenetically evolving lineage with a gradual and directional pattern of increasing molar hypsodonty. Dental changes related to increasing hypsodonty are comparable to those of the octodontid lineage Chasichimys also recovered from the Cerro Azul Formation. However, Neophanomys shows comparatively less variation in gross morphology and there are no evidences that this lineage achieved euhypsodonty. In contrast, important changes in enamel microstructure (schmelzmuster) are observed among different populations of Neophanomys, supporting the hypothesis that these changes can occur at least partially independently from modifications in dental gross morphology. The patterns of dental evolution detected in the Neophanomys and Chasichimys-Xenodontomys lineages and the unequivocal polarity of the changes involved, related to increasing hypsodonty, reinforce the hypothesis that chronological differences exist among late Miocene outcroppings of Cerro Azul Formation in central Argentina.     

New material and revision of hipparions from the upper Miocene locality Ravin des Zouaves-5, RZO of Axios Valley (Macedonia,Greece)

The reconsideration of the hipparions from the locality Ravin des Zouaves-5 (RZO) of the lower Axios Valley provides a better taxonomy, based on the revision of the old known collection (up to 1987), the inclusion of new discoveries and the bibliographic data of the last 30 years related to the study of the hipparions in Greece and the neighboring countries. Based on the cranial material and metapodials, five different taxa have been distinguished. New cranial material confirms the presence of the large-sized Hipparion proboscideum and the small-sized Hipparion macedonicum in the RZO fauna. In the new middle-sized hipparion sample from RZO two species have been identified, Hipparion philippus, and another form similar to Hipparion mediterraneum, which is attributed to Hipparion cf. mediterraneum. Cranial remains of a second small-sized hipparion are also recognized in the RZO fauna, which is attributed to Hipparion sithonis. The relationships of H. mediterraneum and H. proboscideum are discussed; a cladistic analysis suggests that they are sister taxa, sharing a common ancestor with the Hipparion moldavicum group, and a form similar to H. mediterraneum is a possible ancestor of H. proboscideum.     

New data on late Miocene mammals of Kohfidisch, Austria

Cervids and bovids of the Early Turolian mammalian fauna of Kohfidisch (Austria) are reviewed. A new species, Micromeryx mirus sp. nov. (Moschidae) is described.     

Palaeoreas,Majoreas, and Stryfnotherium gen. nov. (Mammalia: Artiodactyla: Bovidae) from the Late Miocene of Greece

A new comparative study of the two crania, LGPUT RZO-03 and LGPUT RZO-68 attributed to Palaeoreas zouavei Bouvrain, 1980 from the Late Miocene of Axios Valley, Greece reveals great morphological and metric differences between them, which require the exclusion of LGPUT RZO-68 from this species and even from Palaeoreas Gaudry, 1861. This decision leaves unsupported many diagnostic features of P. zouavei applied by previous authors and along with recent data from Bulgaria put in doubt the validity of P. zouavei as a whole. As a result, the genus Palaeoreas is re-defined as monotypic and the cranial morphometric changes of P. lindermayeri (Wagner, 1848) are discussed on a temporal basis. An updated taxonomic analysis confirms the presence of P. lindermayeri in the Late Turolian mammal assemblage of Dytiko-1 (Axios Valley, Greece) but rejects its occurrence at Dytiko-2. Instead, a re-examination of old cranial specimens and a study of new ones suggest the presence in both sites of Majoreas Kostopoulos, 2004 a Late Miocene taxon previously mostly known from the eastern banks of the Aegean Sea. A reassessment of the LGPUT RZO-68 cranium, and its comparison with Palaeoreas and Criotherium Forsyth-Major, 1891, suggests that it deserves taxonomic distinction at genus and species levels: Stryfnotherium exophthalmon.     

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.     

Orbital signatures in a late Miocene dinoflagellate record from Crete (Greece)

A high-resolution palynological study of the cyclically bedded Faneromeni section (upper Tortonian-lower Messinian) on Crete (Greece) is presented. This study aims to recognize orbitally-driven variations in the palynological record and to validate the age model based on the astronomical calibration of the sedimentary cycles. Four palynology-based environmental proxies were utilised using interpretations of fossil dinoflagellate associations based on modern ecological characteristics. Cross-spectral analysis between the proxy records and astronomical target curve, the 65°N summer insolation, yielded in most cases significant spectral power and coherence in the precession and/or obliquity frequency bands. Precession-controlled variations in the proxy records are related to lithology and indicate that maxima in continental input and minima in sea surface salinity coincide with sapropel formation. The influence of obliquity is most clearly reflected in the index of continental versus marine palynomorphs (S-D). The absence of a distinct time lag relative to obliquity indicates that the 41-kyr component in continental input is controlled by oscillations in regional Mediterranean climate rather than by glacial cyclicity. Phase relations in the different astronomical frequency bands of the spectrum, as compared with the Mediterranean Pliocene, essentially confirm the validity of the Miocene astronomical time scale. Finally, a major non-cyclic change in the palynological assemblage at 6.68 Ma indicates enhanced salinity and decreased river discharge. This shift coincides with a significant drop in sedimentation rate informally termed the “Early Messinian Sediment starvation Event”.     

The deciduous lower dentition of Ouranopithecus macedoniensis (Primates, Hominoidea) from the late Miocene deposits of Macedonia, Greece

Two mandibular fragments with associated milk teeth assigned to the late Miocene hominoid primate Ouranopithecus macedoniensis are analyzed. The fossils, which belong to a single individual, were found in the Vallesian locality of "Ravin de la Pluie" of the Axios Valley (Macedonia, Greece). The material is described here and compared with extant and extinct hominoids, allowing assessment of the evolutionary trends in the deciduous lower dentition within the Hominoidea. Hylobatids represent the more primitive pattern. Gorilla is slightly more derived than hylobatids, but less derived than Pongo and Pan, the latter being the most derived. With relatively smaller deciduous canines and more molarized deciduous premolars, Ouranopithecus is more derived than both Pan and Gorilla. Among the fossil hominoids, Proconsul, representing the primitive condition, has a very simple dp(3)and a dp(4)that has a trigonid that is taller than the talonid and which lacks a hypoconulid. Griphopithecus is more derived than Proconsul in having a dp(4) with a lower trigonid, a hypoconulid, and a less oblique cristid obliqua. Australopithecus and Paranthropus possess a similar morphology to that of Homo, while Ardipithecus appears to be more primitive than the latter genera. Ouranopithecus has a more derived lower milk dentition than Proconsul and Griphopithecus, but less derived than Australopithecus and Paranthropus. The comparison of the lower milk dentition of Ouranopithecus confirms our previous conclusions suggesting that this fossil hominoid shares derived characters with Australopithecus and Homo.     

甘肃临夏盆地首次发现乌米兽(牛科,偶蹄类)头骨化石（英文）

报道并描述了一件来自甘肃临夏盆地的中间乌米兽(Urmiatherium intermedium)头骨化石新材料,该材料产自柳树组上部,属于晚中新世晚期杨家山动物群。中间乌米兽是一种大型的晚中新世牛科动物,角心特化,短且呈薄板状,并且在基部相互靠近。在角心前后方的额骨和顶骨上,发育大片赘生骨疣,这些骨疣与角心一起,合称角器。20世纪初,步林报道了中国北方晚中新世地层中的大量中间乌米兽化石,包括产自山西保德和甘肃庆阳的20多件头骨以及很多破碎的齿列和骨骼。本文报道的乌米兽头骨化石是乌米兽在甘肃临夏盆地的首次发现,将其在中国北方的分布向西扩展到了青藏高原东北缘地带。乌米兽被普遍认为与近旋角羊(Plesiaddax)、和政羊(Hezhengia)和柴达木兽(Tsaidamotherium)等晚中新世"麝牛类"牛科动物具有较近的亲缘关系,但其系统发育地位仍存有争议。相比其他晚中新世"麝牛类"牛科动物,乌米兽的分布较广,从伊朗至中国北方都有分布,但它鲜与其他晚中新世"麝牛类"牛科动物伴生。在临夏盆地,与其伴生的牛科动物目前仅发现中华羚(Sinotragus)一种。     

Miocene ostracods from the Tremiti Islands and Hyblean Plateau: biostratigraphy and description of new and poorly known species

The main taxonomic and stratigraphical results of a study of ostracods from the middle-late Miocene of the Tremiti Islands and the Hyblean Plateau are presented. The faunas are mostly typical of deep thermospheric or psychrospheric bathyal environments. Excluding a few, mostly juvenile shallow water contaminants, a total of 127 species were recorded. Twenty-four taxa are discussed herein; of these, 13 species (Cytherella dissimilis nov. sp., Cytherella parvula nov. sp., Cytherella subtilis nov. sp., Argilloecia triangularis nov. sp., Retibythere (Retibythere)? claudii nov. sp., Retibythere (Retibythere)? gibba nov. sp., Cytheropteron (Cytheropteron) joachinoi nov. sp., Cytheropteron (Aversovalva) consuetum nov. sp., Cytheropteron (Aversovalva) trimerumense nov. sp., Cytheropteron (Aversovalva) turgidum nov. sp., Loxoconcha julianii nov. sp., Buntonia whatleyi nov. sp., Pterygocythere nuda nov. sp.) and a subspecies (Costa tricostata dorsoarcuata nov. subsp.) are described as new, while six are left in open nomenclature. The stratigraphical distribution of ostracods is expressed in terms of planktonic foraminiferal zones. Information obtained from this study enriches biostratigraphical data on Mediterranean Miocene deep-sea ostracods, especially with respect to the Langhian.     

A new elasmothere (Perissodactyla, Rhinocerotidae) from the late Miocene of the Linxia Basin in Gansu, China

Ningxiatherium euryrhinus sp. nov. is a relatively large elasmothere rhino from the Linxia Basin in northwestern China found in the early Bahean, which corresponds to the early Vallesian (MN 9), ca. 11.1 Ma. It is much larger than the extant Ceratotherium simum and has a single horn. Ningxiatherium is similar to the late Miocene Parelasmotherium from Gansu and Shanxi, but differs in having partially ossified nasal septum, terminal nasal horn boss, shallower nasal notch above the P3/P4 boundary, and much shorter premolars. N. euryrhinus sp. nov. differs from N. longirhinus from the late Miocene of Zhongning, Ningxia in northwestern China by its larger size, relatively wider nasals, shallower nasal notch, and sub-quadrangular occlusal surface of M3. In the Linxia Basin, the other early late Miocene (Bahean or Vallesian) elasmotheres are known from complete skulls or isolated teeth. They include two genera and three species: Parelasmotherium linxiaense, P. simplum, and Iranotherium morgani. They lack a nasal septum, but Ningxiatherium has one. The age of N. euryrhinus is early late Miocene based on direct association with biochronologic indicator taxa, such as Dinocrocuta gigantea, Hipparion dongxiangense, and Chilotherium primigenius. N. euryrhinus is more primitive than N. longirhinus sp. nov. in having a sub-quadrangular M3, a shallower nasal notch, and the presence of DP1 in adults.     

Rodents and insectivores from the late Miocene of Romerales (Fortuna Basin,Southern Spain)

Abstract

The Neogene Fortuna Basin (Murcia Region, SE Spain) is rich in microvertebrate sites. Its continental sections include localities extended from the middle Turolian (MN12) to the early Ruscinian (MN14). However, there are few works dealing with the taxonomy of these fossil assemblages. In this paper we provide a complete taxonomic study of the rodents and insectivores from three levels (ROM-2B, ROM-2C and ROM-3A) within the Romerales section. We infer a late Turolian age (late Messinian) for these levels, among which the richest and most diverse level is ROM-C, including at least 11 different taxa. In addition, the paleoecological analysis of these fossil assemblages suggests the dominance of open herbaceous meadows under temperate climate during the formation of these sites, with a slight decrease in temperature and humidity from ROM-2B to ROM-2C.     

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.     

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.     

Stephanorhinus hundsheimensis (Mammalia,Rhinocerotidae) from the late early Pleistocene deposits of the Denizli Basin (Anatolia,Turkey)

Pleistocene rhinoceroses are poorly documented in Turkey where they have been reported only from the late early Pleistocene (1.3–1.1 Ma) travertine deposits of the Denizli Basin. In this work, new rhinoceros remains collected from this basin are assigned to a relatively large-sized Stephanorhinus hundsheimensis on the basis of their morphology and morphometry. The first Turkish record of this species is approximately coeval with the first appearance of S. hundsheimensis in Europe, chronologically referred to the late early Pleistocene, ca. 1.2 Ma. During that time, S. etruscus still survived in Iberian Peninsula, central Italy and Dacian Basin. The presence of two successive evolutionary morphs for S. hundsheimensis during the Pleistocene is not confirmed.     

Brachypotherium (Perissodactyla,Rhinocerotidae) from the late Miocene of Samburu Hills,Kenya

Several isolated cheek teeth and mandibular specimens of Rhinocerotidae (Mammalia, Perissodactyla) from the upper Miocene Namurungule Formation in Samburu Hills, Kenya, are redescribed. Previously, these specimens had been identified as Chilotheridium pattersoni, Chilotheridium sp., Paradiceros mukirii, and Paradiceros sp. They are reidentified here as documenting the genus Brachypotherium based on their bucco-lingually broad molariform upper premolars with short crochet and flattened buccal walls on both upper and lower molars, the latter having a shallow external groove. Comparisons with other Brachypotherium species suggest that the present specimens belong to Brachypotherium sp. cf. B. minor. The presence of Brachypotherium in the Samburu Hills, at ca. 9.5 Ma, is concordant with the paleoenvironment (presence of lacustrine and river environments) known for this locality during the early late Miocene.     

A new stage in the evolution of the mole rats (Rodentia,Spalacinae) from the Early Miocene of northern Greece

The cheek teeth of Debruijnia kostakii n. sp. from the Early Miocene of Karydia are only slightly smaller than those of the largest extant species of the Spalacinae (Spalax giganteus), but show characteristics that are reminiscent of its Paleogene cricetid ancestor. The peculiar combination of large size and primitive dental morphology of this species bridges the gap between Debruijnia and Pliospalax and thereby supports the hypothesis that these genera form the core of the complex evolutionary history leading to extant Spalax.     

Systematics and phylogeny of Vasseuromys (Mammalia,Rodentia, Gliridae) with a description of a new species from the late Miocene of eastern Europe

Vasseuromys is a species‐rich genus of small‐ to medium‐sized glirids spanning the latest Oligocene to late Miocene of Europe and western Asia. Despite extensive discoveries over the past 50 years, little phylogenetic work has been done on Vasseuromys. This study presents the first phylogenetic analysis of the genus that includes all the described species and a new taxon Vasseuromys tectus sp. nov. from the late Miocene of eastern Europe, providing the first insights into the evolutionary relationships within the clade. Results suggest that the genus is clearly paraphyletic. Two strongly supported genus‐level clades are recognized within ‘Vasseuromys’: a restricted Vasseuromys clade (containing the three species, V. pannonicus, V. rugosus and V. tectus) and the Glirulus clade that includes ‘Vasseuromys’ duplex. The remaining ‘Vasseuromys’ species are found to constitute a set of paraphyletic taxa, with the polyphyletic ‘Ramys’ nested within it. The genus Gliruloides is synonymized with Glirulus. Vasseuromys tectus sp. nov. is the most derived member of the genus in having a greater number of cheek teeth ridges including constantly present anterotrope, centrotrope, second prototrope on M1–2, third metatrope on M2, two to three posterotropids on p4 and strong ectolophids on lower molars. The results of the study confirm a European origin for Vasseuromys while suggesting that the late Miocene species of the genus dispersed from the east in the early Turolian.