A new hipparion (Mammalia,Perissodactyla) from the Vallesian (Late Miocene) of Greece

A small hipparion from the Vallesian (early Late Miocene) of Northern Greece is studied. It is coming from the locality “Ravin de la Pluie” of the lower Axios valley, near Thessaloniki. Its characteristics as the small size, the short symphysis, the long snout, the high enamel plication, the well developed protostylid, the large m₃ and the elongated m₁-m₃ series distinguish it from the other known eurasiatic small hipparions, allowed us to derive a new species,Hipparion macedonicum. It lived during Vallesian and can be used as a stratigraphic species for this period.     

Correlation ofHipparion faunal components in the East mediterranean and paratethys areas

Several species ofHipparion from Turolian/Middle Sarmatian-Meotian faunas in the East Mediterranean and Paratethys areas are compared. The large hipparions common to the faunas are referred toH. primigenium (v. Meyer), the smaller hipparions discussed belong to the speciesH. verae Gabunia,H. mediterraneum Roth & Wagner,H. cf.moldavicum Gromova, andH.schlosseri Antonius. A correlation of the faunas in the two areas is attempted on the basis of evolutionary events (e. g. radiation) and the presence of taxonomically similar species.     

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.     

A new protictithere from the late Miocene hominoid locality Ravin de la Pluie of Axios Valley (Macedonia, Greece)

This article concerns the study of a protictithere from the late Miocene of Axios Valley (Macedonia, Greece). The material was found in the hominoid-bearing mammal locality Ravin de la Pluie (RPl), which is corellated with the late Vallesian, MN 10. The available material, including maxillary and mandibular elements, is described and compared morphologically and metrically with the known Eurasian protictitheres. The small size, the low cusps(-ids) of the teeth, the protocone of the P4, which is in line with the mesial border of the parastyle, the large molars (especially the M2), the small buccal projection of the M1 paracone, the strongly molarized p4, the strong metaconid and large talonid with high entoconid of the m1 separate the RPl protictithere from the known Eurasian protictitheres and allow the erection of a new species, named P. thessalonikensis n. sp.     

New material of Carnivora (Mammalia) from the Late Miocene of Axios Valley,Macedonia, Greece

A new collection of carnivores from the Late Miocene deposits of Axios Valley, Macedonia, Greece was collected in 2004–2009 at the localities named Xirochori 1 (XIR) and Ravin de la Pluie (RPl), dated to the Late Vallesian (MN 10), as well as at Ravin des Zouaves 5 (RZO), dated to the Early Turolian (MN 11). The studied material is described and compared morphologically and biometrically with other materials from Greece and its neighboring area. The following species are determined: XIR: Dinocrocuta sp.; RPl: Eomellivora wimani, Adcrocuta eximia leptoryncha, ?Hyaenictis sp., Metailurus parvulus; RZO: Plioviverrops orbignyi, Machairodus giganteus. This collection is interesting because it includes the upper deciduous dentition of A. eximia and Dinocrocuta, which are only known from limited material until now, especially the latter taxon. E. wimani is recognized for the first time in Greece and the Balkans, while M. parvulus is identified for the first time in the Vallesian of Greece. The possible presence of Hyaenictis in RPl is the first indication of its presence in the Vallesian. The guild structure diagrams of the carnivore fauna from the three studied localities provide some indications of their palaeoenvironment, which fits quite well with the results of previous studies on the palaeoenvironments of these localities and of the wider Eastern Mediterranean region.     

Study of turolian hipparions of the lower axios valley (Macedonia,Greece) 2. locality »prochoma-1« (PXM)

The hipparions of the locality Prochoma-1 of the lower Axios valley (Macedonia, Greece) are studied. Two species can be distinguished, the medium-sizedH. dietrichi (Wehrli) and the smallsizedH. macedonicum Koufos. The presence of the large-sizedH. proboscideum Studer, is possible. The locality dates back to Latest Vallesian-Early Turolian as it is proved by some morphological characteristics of hipparions, by the faunal composition and by its similarity with the faunas of the other localities of the lower Axios valley.     

Late Miocene Proboscidea (Mammalia) from Macedonia and Samos Island, Greece: preliminary results

The abundant Late Miocene proboscidean remains of Greece have never been studied in detail and compared with those of Eurasia in order to determine their taxonomy and their biostratigraphical and palaeoecological significance. The first results of such study are given in this article. During the past decades, several new proboscidean specimens have been added to the old collections, significantly enriching the available material. The Axios Valley (Macedonia, Greece) proboscidean fossils belong mainly to two species of Choerolophodon: C. anatolicus of early Vallesian age and C. pentelici of late Vallesian–Turolian age. Deinotherium giganteum is rare and recognized only in the late Vallesian locality Ravin de la Pluie of Axios Valley. A zygodont form has also been identified in the Turolian of Axios Valley, attributed to “Mammut” sp. The Late Miocene localities of Nikiti (Macedonia, Greece) revealed several remains of C. pentelici, which are similar to the Turolian ones of Axios Valley. The Samos proboscidean collection includes C. pentelici, “Tetralophodon” atticus, “Mammut” sp. and Deinotherium gigantissimum. The taxonomy of the Late Miocene peri-Mediterranean Choerolophodon is given, and the biostratigraphy and palaeoecology of the Greek Late Miocene proboscideans are discussed.     

A hominid tooth from Bulgaria: the last pre-human hominid of continental Europe

A hominid upper premolar was discovered in the Azmaka quarry, near Chirpan (Bulgaria). The associated fauna, especially the co-occurrence of Choerolophodon and Anancus among the proboscideans, and Cremohipparion matthewi and Hippotherium brachypus among the hipparions, constrains the age of the locality to the second half of the middle Turolian (ca. 7 Ma), making it the latest pre-human hominid of continental Europe and Asia Minor. The available morphological and metric data are more similar to those of Ouranopithecus from the Vallesian of Greece than to those of the early to middle Turolian hominids of Turkey and Georgia, but the time gap speaks against a direct phyletic link, and Turolian migration from the east cannot be rejected.     

The hipparions (Mammalia,Perissodactyla) from the Turolian locality “Nikiti-2” (NIK), Macedonia,N. Greece

The hipparions from the late Miocene locality Nikiti-2 (NIK), Macedonia, Greece are described and compared with those from the other Greek and Eurasian localities. Two species have been determined, the medium-sized Hipparion dietrichi and the small-sized Hipparion macedonicum, while a third large-sized Hipparion is also recognized. The scanty material of the latter species indicates similarities with Hipparion proboscideum, as well as with Hipparion mediterraneum and it is referred to as Hipparion sp. The locality is dated to early Turolian as this is proved by the resemblance of the hipparions from “Nikiti 2” faunal assemblage with those from the neighbouring localities of “Ravin des Zouaves 5” and “Prochoma 1”, of Axios Valley, Greece. Interesting differences, which are inferred by the comparison of the studied material with those of Axios Valley, Samos and Turkey, are also discussed.     

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.     

Early Late Miocene insectivores (Eulipotyphla,Mammalia) from the Cañada section (Province of Zaragoza,east Central Spain)

Vallesian (early Late Miocene) strata from the recently introduced Cañada section (province of Zaragoza, east Central Spain) have yielded fairly large insectivore assemblages. These show that, after the generally dry Aragonian, the Vallesian gave rise to more humid conditions that were favourable to insectivores, both in number of taxa, and in overall number of specimens. The assemblage of Cañada 8 (Biozone H) is dominated by shrews, whereas the assemblage of Cañada 10 (uppermost Biozone H) contains the oldest record of Desmanella in the area. This seems to signify a bioevent in which after millions of years of absence, talpids return to the area. In addition to the Vallesian assemblages, a small Turolian insectivore fauna has been recovered. On the basis of the rodents, Cañada 12 was assigned to Biozone L, and the insectivore assemblage is very similar to the assemblages from the Teruel basin of that zone. This implies that the discovery of Postpalerinaceus in Cañada 12 is the youngest published record of this large spiny hedgehog.     

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.     

The Hipparion of the Rhone valley

A revision is given of the Rhone valley Hipparion. One new species is described from Montredon. It is noted that in the lower Vallesian localities one big heavily built Hipparion is present. In the late Vallesian a medium sized Hipparion with short limbs is found; in the Turolian a slender form of medium size and perhaps also a heavy form is present; finally in the Ruscinian two forms are found of which one has a very big size and has cabaloid characteristics.Some remarks are made about the paleoecology of Hipparion in general.     

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.     

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.     

New materials of Dinocrocuta (Percrocutidae, Carnivora) from Lantian, Shaanxi Province, China, and remarks on Chinese Late Miocene biochronology

New materials from the middle part of the Bahe formation are described as Dinocrocuta gigantea. Review of the species reveals that it is derived in the evolutionary lineage of Dinocrocuta, and biochronologically later than Vallesian records from Turkey. The only possibly related Vallesian species from China is Crocuta gigantea xizangensis from Biru, Tibet, which may prove to be conspecific with D. senyureki. Based on the mammalian faunal sequence from Lantian, and with reference to Red Clay paleomagnetic data, the duration of D. gigantea in China should be later late Miocene, rather than the previously postulated early late Miocene (Vallesian equivalent) age.     

Supraspecific grouping of Old World hipparions (Mammalia,Equidae)

Morphological characters useful for supraspecific grouping of hipparionid horses are discussed. On the basis of such characters supraspecific groups are distinguished among Old World hipparions. These are the generaHipparion, Proboscidipparion, andNeohipparion, the latter with the subgeneraNeohipparion andStylohipparion. The relationship of the New World hipparions to the Old World groups is briefly discussed.     

New material of Muntiacinae (Artiodactyla,Mammalia) from the Late Miocene of the northeastern Qinghai-Tibetan Plateau,China

Recent expeditions in the northeastern Qinghai-Tibetan Plateau have produced additional mammalian fossils from the Lower Baodean (equivalent to Vallesian, Late Miocene), and, significantly, a new large and the earliest muntjac, Muntiacus noringenensis sp. nov. It is phylogenetically closer to the Late Pliocene M. fenghoensis and extant M. vuqangensis than the other fossil and extant muntjacs are. The existence of this muntjac and other folivores in the Late Miocene Qaidam Basin suggests a forested period in the basin and the evolution from a forested and humid environment to a desert one today. Such a dramatic evolution in environments is an evidence of the effect of the uplift of the Qinghai-Tibetan Plateau and of its influence on continental climates.     

The paleoenvironment of Hispanopithecus laietanus as revealed by paleobotanical evidence from the Late Miocene of Can Llobateres 1 (Catalonia, Spain)

The early Vallesian site of Can Llobateres 1 (Vallès-Penedès Basin, Catalonia, Spain) is one of the richest localities of the European Late Miocene, having yielded the most complete remains of the fossil great ape Hispanopithecus laietanus (Primates: Hominidae). Fossil plant remains had been previously reported from this site but mostly remained unpublished. Here we describe an assemblage of plant megaremains recovered in 2010, which provides valuable paleoenvironmental data. This assemblage consists of a mixture of parautochthonous and allochthonous detached organs (leaves, stems, reproductive structures) deposited in marshy areas. The source vegetation mainly consisted of abundant reeds, palms, evergreen laurels and figs that probably grew in or near the marsh boundaries or nearby riparian forests. This environmental picture is consistent with the mammalian fauna, which shows the prevalence of humid forested environments, although somewhat more open woodlands might have been present away from the wet areas. The occurrence of mega-mesothermal taxa, together with the absence of deciduous elements, is consistent with a subtropical to warm-temperate climate. Within this mosaic environment, H. laietanus would have preferred the more humid and forested habitats, which probably were still quite common in the Vallès-Penedès during the early Vallesian. Such habitats would have provided a continuous ripe fruit supply throughout the year to these frugivorous great apes. Paleobotanical data from older sites of the same area and nearby basins show that the zonal vegetation was a warm-temperate mixed forest defined by evergreen laurels, together with leguminous trees and shrubs as well as a significant proportion of deciduous elements. Tropical and subtropical taxa would have been restricted to humid areas in the lowlands. From the late Vallesian onwards, many of these taxa disappeared from the Vallès-Penedès, whereas deciduous trees became dominant in the forested areas and wetlands, thus likely having driven Hispanopithecus to extinction in the study area.