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.     

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.     

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.     

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.     

A new Late Miocene ovibovine-like bovid (Bovidae,Mammalia) from the Kassandra Peninsula (Chalkidiki,Northern Greece) and implications to the phylogeography of the group

A new Late Miocene bovid, Urmiatherium kassandriensis sp. nov., from Northern Greece is described. The material comes from the Fourka locality in the Kassandra Peninsula (Chalkidiki), and the included fauna is estimated to be of Vallesian age. The two preserved crania represent a medium-sized taxon with short, conical horn cores, a flat cranial roof (consisting of the posterior part of the frontals, parietal and occipital), thick and porous frontals and pneumatized short parietals, an extremely thick basioccipital with voluminous posterior tuberosities and accessory articular facets for the atlas. The specialized atlanto-occipital joint recalls Pleistocene and extant ovibovines, but the braincase structure as a whole and the horn core features closely match Late Miocene ovibovine-like taxa, especially Plesiaddax and even more Urmiatherium. Nevertheless, the Kassandra bovid differs from representatives of both genera in the simpler horn core morphology and external brain anatomy. Urmiatherium is known to appear first in China and Iran at about 7.8 Ma, whereas its westernmost appearance on Samos Island (Greece) is dated much later. The presence of Urmiatherium kassandriensis sp. nov. in N. Greece suggests a farther west and earlier (Vallesian at least) first appearance of the genus. This would justify a basic geographic and phylogenetic split of Urmiatherium into two main Turolian lineages: a central-eastern Asian one leading to the sister species U. polaki and U. intermedium and a western one leading to U. rugosifrons.     

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.     

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.     

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.     

Mandible of Ouranopithecus macedoniensis (Hominidae,primates) from a new late Miocene locality of Macedonia (Greece)

In 1990, a new late Miocene locality named “Nikiti 1” or NKT, was discovered near the village of Nikiti (Chalkidiki, Macedonia, Greece) about 100 km east of Thessaloniki City. The locality is situated in the Nikiti Formation, which consists of yellowish sands, gravels and pebbles and has been dated to late Vallesian–early Turolian. Among the initially collected fossils there is a mandible of a hominoid primate, which is described, compared, and attributed to Ouranopithecus macedoniensis. A short review of the species in Macedonia and its phylogenetic relationships are also given. © 1993 Wiley-Liss, Inc.     

The Bovidae of Dorn-Dürkheim 1, Germany (Turolian age)

Three bovid species are present at Dorn-Dürkheim 1. The overwhelmingly abundant species is a boselaphine, Miotragocerus sp., with smaller and less advanced teeth than Tragoportax amalthea from Pikermi, Greece. Miotragocerus was present in the latest middle Miocene and Vallesian of western and central Europe and survived into the Turolian. The other two bovid species remain enigmatic and of uncertain tribal affiliation. Each is represented by very few teeth, none of them associated. One species is larger and the other smaller than the Miotragocerus sp. Mesowear analysis of M²s and M³s was used to investigate the dietary regime of Miotragocerus sp. from Dorn-Dürkheim 1. Miotragocerus was found to be linked to extant browsing ungulates close to the transition to mixed feeders. The percentage of abrasive food components like grass in the diet of this species was probably close to 10%. This ratio suggests Miotragocerus is intermediate between the two hipparionine horse species of this early Turolian (MN11) palaeoenvironment.     

Qurliqnoria (Bovidae,Mammalia) from the Upper Miocene of Çorakyerler (Central Anatolia,Turkey) and its biogeographic implications

New bovid material from the Upper Miocene site of Çorakyerler (Çankırı basin, Anatolia, Turkey) is described and compared here. The described taxon is identified as a representative of the stem caprine genus Qurliqnoria, previously known from the peri-Tibetan area exclusively. The stronger horn-core divergence, weaker anterior keel, smoother horn-core surface, stronger lateral horn-core curvature, stronger and thicker interfrontal suture, less flexed and less pneumatized frontals, and smaller supraorbital foramina differentiate the Çorakyerler Qurliqnoria from the type and only known species of the genus, Q. cheni from China, and demand the erection of a new species, Qurliqnoria chorakensis n. sp. A review of other late Miocene bovid records allows the recognition of Qurliqnoria in Sinap Tepe (Turkey) and Platania (Greece), suggesting a westward propagation of the genus during the Vallesian.     

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.     

The hipparions (Mammalia,Equidae) from Gülpinar (Canakkale,Turkey)

The mammalian fauna from Gülpinar (Canakkale, Turkey), dated Turolian, comprises threeHipparion taxa:H. cf.matthewi, H. sp. medium-sized, andH. sp. large. Two metatarsals may represent a fourth taxon or may derive from a different locality. The hipparion material is described here.     

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.     

A contribution to the chronology of some Old World miocene faunas based on hipparionine horses

The Maragheh Formation, northwestern Iran,provides a local biostratigraphic base upon which to build a biochronology of hipparionine horses that has potential regional importance in ordering faunas of Vallesian and Turolian age in other areas of the Old World. This, as well as faunal and radiometric analysis of the Maragheh sequence, is compared with those aspects of other districts. The radiometric age of the so-called «Hipparion Datum» is 12 Ma, if not slightly older, and more than one species of «Hipparion» may be associated with it. Based on cranial morphology, hipparionine horses of Vallesian age consist of a single group; faunas of Turolian age contain four hipparionine groups by these criteria. A group composed of Hipparion prostylum and later members appears to be the most useful in compiling a possible biochronology. This group may have had an endemic European rather than allochthonous (American) origin. Based on the various assessments discussed here the following temporal sequence of certain faunal localities is proposed (oldest to youngest): Vallesian-Höwenegg (Germany), Bou Hanifia I (N. Africa) and Hostalets de Pierola (Spain). Early Turolian-Kopran, Lower Maragheh (Iran). Medial Turolian-Mont Luberon (France), Kerjabad or Ketschawa, Middle Maragheh (Iran), Saloniki (Macedonia), Pikermi (Greece), Shol'avand, Upper Maragheh (Iran), Samos (Greece). Our assessment of the age of the Mont Luberon and Samos faunas differs from conventional assignments.     

New data on Mio-Pliocene Sciuridae (Rodentia,Mammalia) from southern Spain

Sciurids are very scarce in the fossil record, especially in the basins of southern Spain. The aim of this paper is to review the Sciuridae record in these basins. The Granada and Guadix basins have yielded specimens of Xerini and Pteromyinae, which represent the largest collection of fossil Sciuridae in southern Spain from the Middle Turolian to the Upper Ruscinian. The new discoveries change the currently known geographical and temporal range of some taxons, since we find the oldest evidence of Pliopetaurista pliocaenica in localities from the Late Turolian and of Heteroxerus mariatheresae in a locality of the Middle Turolian. Furthermore, we record the first evidence of Atlantoxerus margaritae in southern Spain.     

The presence ofsus strozzii in the villafranchian (villanyian) of macedonia (greece)

The locality of Gerakarou is a new Pleistocene one in Macedonia (Greece) found in 1978. Between the material recovered there is a suid skull, which is studied in this article. Its morphological characters and dimensions allow us to determine this asSus strozzii Forsyth Major. It seems to be similar to the Upper Valdarno and Olivola suids and different from the other Villafranchian suid,Sus minor. Thus it is dated to Late Villafranchian (Villanyian). The certain presence of the species in Greece, where it was known only from some isolated specimens referred asSus cf.strozzii, completes the knowledge of the geographic distribution of the species in Southern Europe.     

Afrochoerodon nov. gen. kisumuensis (MacInnes) (Proboscidea,Mammalia) from Cheparawa,Middle Miocene,Kenya

A proboscidean skull from Cheparawa, (Muruyur Formation, Kenya), differs markedly from those of Eurasian Choerolophodon (C. pentelici, C. dhokpathanensis). It is morphologically and metrically close to the holotype of Choerolophodon kisumuensis (MacInnes) a partial skull from Maboko, much of which has been reconstructed in plaster of Paris. The more complete remains of this species now available indicate that it should be placed in a genus separate from Choerolophodon. The new genus Afrochoerodon is erected for it. Choerolophodon ngorora from Ngorora and Fort Ternan (Kenya), Choerolophodon zaltaniensis from Gebel Zelten (Libya) and Choerolophodon chioticus from Chios, Greece, should be transferred to the genus Afrochoerodon. Late Miocene specimens from Nakali, Kenya are probably referrable to the genus Choerolophodon. Fossils from Burji-Soyama (Ethiopia) hitherto assigned to Choerolophodon sp. are excluded from the subfamily Choerolophodontinae.     

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.