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 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 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.     

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.     

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.     

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.     

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.     

Palaeoecological interpretation of the late Miocene landscapes and vegetation of northern Greece: A comment to Merceron et al., 2016 (Geobios 49, 135–146)

Recent works on feeding habits of ungulates, isotopic composition of equid tooth enamel, and phytoliths from late Miocene localities of northern Greece suggested the presence of savannah and excluded dense forests. Furthermore, Mediterranean-like climates were advocated for the late Miocene of Greece. Here, I compare palaeoenvironments inferred for two mammal localities from Chalkidiki and the Axios Valley (Nikiti, upper Vallesian and lower Turolian; Dytiko, upper Turolian) with evidence from contemporaneous plant assemblages from adjacent areas in Greece and Bulgaria. I use vegetation units inferred from pollen and spore, fruit and seed, and leaf assemblages and compare them with the vegetation inferred from mammal and phytolith data. Open vegetation as inferred from mammal and phytolith data is also part of the range of vegetation units discovered from the pollen and spore, seed and fruit, and leaf record (here called the palaeobotanical record). Poaceae are consistently present in late Vallesian to late Turolian pollen records of northern Greece. Further, a number of vegetation units are indicative of forest vegetation ranging from lowland to upland forests dominated by woody angiosperms and mixed coniferous-angiosperm forests. The presence of such forests is not in conflict with the results from mammal and phytolith studies, but it broadens the view on landscapes present in the late Miocene of northern Greece. In addition to a generalized vegetation type commonly inferred by mammal studies, the palaeobotanical record demonstrates the presence of various complementary vegetation types. A comprehensive view on late Miocene landscapes in northern Greece shows that there is no conflict in inferring open herb dominated landscapes and light to dense forests and provides new opportunities for the ecological interpretation of late Miocene ungulates.     

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.     

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 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.     

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.     

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.     

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.     

The Late Miocene hominoids Ouranopithecus and Graecopithecus. Implications about their relationships and taxonomy

Two Late Miocene hominoids are known from Greece. The first, Graecopithecus freybergi, is known by a single mandible with the worn m2 from the locality of Pyrgos Vassilissis, near Athens. The other, Ouranopithecus macedoniensis, is known from Axios Valley and Chalkidiki (Macedonia, Greece) by a partial skull and a set of maxillary and mandibular remains. Some authors consider these two hominoids as synonyms and in the present article a detailed comparison of them is given. The morphology and size of the symphysis, the more robust mandible, the more open dental arcade of Ouranopithecus distinguishes it clearly from Graecopithecus. Moreover, the incompleteness of the mandible of Graecopithecus with the doubtful morphology and size, the limited material and the uncertain geological age of the locality cannot allow precise and clear comparisons with the rest extant and extinct hominoids. Thus in our opinion there are not enough data to support the similarity, and therefore, the synonymy of the two genera. The Pyrgos mandible must remain as a separate and isolated genus with one species, which only includes this sole mandible.     

Intraspecific variation and evolutionary trends of Alicornops simorrense (Rhinocerotidae) in Spain

The Spanish material of Alicornops simorrense constitutes the best representation of the species in Western Europe. It provides interesting data on the metrical and morphological intraspecific variation of A. simorrense and its evolutionary trends from late Middle Miocene to the early Late Miocene. From late Aragonian to early Vallesian, a slight increase in size is observed, but without clear limits among series. During the late Vallesian, A. simorrense evolved in central Spanish basins into a second species, A. alfambrense, greater in size and with more robust proportions. In the Vallés-Penedés basin, A. simorrense shows a noticeable increase in size, while maintaining its proportions, during the Vallesian. A. simorrense was a very abundant species, an open woodland dweller, with gregarious behaviour, whose extinction was probably linked to the climatic change that took place at the end of the Vallesian and the beginning of the Turolian.     

The importance of Late Miocene faunal exchanges between Eastern Mediterranean areas and Central Europe

Recent studies of mammal faunas from the Vienna and Pannonian Basins—in particular the assemblage from Kohfidisch in Burgenland (Austria)—provide new data on the faunal turnover at the Vallesian—Turolian transition. They demonstrated a considerable influence of the faunal exchanges between Greco-Iranian, Eastern European and Central European faunal provinces on renewal of mammal communities in Central Europe, particularly at MN10/MN11 boundary around 8.7 Ma. Five new comers from the Balkano-Iranian region (Gazella aff. pigrimi, ?Nisidorcas, Tragoportax gaudryi, Protoryx and Palaeoryx) coexisted in the Early Turolian of Central Europe with the Middle Miocene autochthonous (Orygotherium, Dorcatherium naui, Micromeryx, Euprox, Amphiprox anocerus and Miotragocerus pannoniae) and Late Miocene invaders from Eastern Europe (Procapreolus and Cervavitus). Dispersal events were close related to palaeoenvironmental and climatic changes.     

The fossil turtles of Greece: An overview of taxonomy and distribution

Turtle remains are common in the Miocene-Holocene deposits of Greece, and are a key focus of the growing research interest in Neogene herpetofaunas from the Aegean region. Some of the most important finds include one of Europe's stratigraphically youngest pleurodiran taxa, Nostimochelone lampra, from the Early Miocene of Macedonia, together with arguably the richest record of fossil tortoises from the Eastern Mediterranean. This incorporates the presently oldest definitive representatives of the quintessential genus Testudo sensu stricto from the Late Miocene of Attica and Macedonia, and numerous specimens of the colossal (carapace ∼2 m-length) testudinid Cheirogaster from Late Miocene-Late Pliocene sediments in southern and northern Greece, as well as on the eastern Aegean islands of Samos and Lesvos. Tantalising, but as yet unconfirmed Miocene accounts of the geoemydid Mauremys in Macedonia, and indeterminate emydid-like remains from Euboea, also provide potentially significant range extensions. Although hampered by a historically sparse documentation, the fossil turtles of Greece are a significant resource that record both assemblage changes and the origin of modern lineages.     

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).