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.     

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.     

The fossil Bovidae (Artiodactyla, Mammalia) from Gesher Benot Ya'aqov, Israel: Out of Africa during the Early-Middle Pleistocene transition

We report the study of the collection of fossil bovid specimens from the Early-Middle Pleistocene Acheulian site of Gesher Benot Ya‘aqov. This locality, situated in the Levantine Corridor (the bottleneck that connects Africa and Eurasia) is a key site to explain the faunal and human dispersals out of Africa during the Matuyama/Brunhes boundary around 0.8 Ma. Two species of bovine (Bos sp., and Bovini gen. et sp. indet. cf. Bison sp.), one antelope (Gazella sp. cf. G. Gazella), and another indeterminate Bovidae gen. et sp. indet., have been recorded. The largest species, Bos sp., is an African immigrant related to the species from the Eritrean site of Buia, Bos buiaensis, which evolved from the buffalo of Olduvai Pelorovis oldowayensis, and colonized the Eurasian continent in parallel with the dispersal of the Acheulian culture into the northern continent. Numerous important species first recorded in several localities of Early-Middle Pleistocene transition from Eurasia are included in this dispersal out of Africa, including the megaherbivore, Palaeoloxodon antiquus, and the carnivores Crocuta crocuta, and later, Panthera leo and Panthera pardus. This faunal turnover is coincident with the change to colder climates that dominated the Middle Pleistocene.     

Presence de Gazella deperdita (Gervais) 1847 du gisement du Miocene superieur (Turolien) des Mistrals (Vaucluse,Sud-Est de la France)

Among the two bovid genera represented in the turolian locality Mistrals I, Gazella and Miotragocerus, the most abundant is a gazelle which, after comparison with Gazella deperdita from the Mont-Luberon, turns out to belong to this species. Resemblances between the two faunas, already noticed in the study of the Hipparions, are thus comfirmed.     

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.     

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.     

Discovery of an Upper Miocene Vertebrate fauna near Tizi N’Tadderht,Skoura, Ouarzazate Basin (Central High Atlas,Morocco)

The discovery of Upper Miocene vertebrates at Tizi N’Tadderht in the Ouarzazate basin (Morocco) helps to fill a gap in our knowledge of Neogene faunas in North Africa. The new fauna includes an ostrich cf. Struthio sp, a turtle cf. Centrochelys sp., Crocodylus cf. niloticus, and a relatively diverse fauna of large mammals. The mammal assemblage probably includes three hipparion species, including a very small form not previously reported from Africa, aff. Cremohipparion periafricanum, two species of rhinoceros cf. Ceratotherium sp. and aff. Chilotherium sp., a Proboscidean cf. Tetralophodon sp., a large member of the Giraffidae similar to “Palaeotragus” germaini and two bovids of which one is likely related to Prostrepsiceros, while the other is a new medium-sized antelope with spiral horns, certainly a representative of the Caprinae, a group that is rare in Africa. A late Miocene age, corresponding to the European Turolian Mammal age, is most likely for this fauna.     

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.     

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 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.     

Early Miocene Amphibians (Caudata, Salientia) from the Mokrá-Western Quarry (Czech Republic) with comments on the evolution of Early Miocene amphibian assemblages in Central Europe

The Mokrá-Western Quarry exhibits the rare occurrence of Early Miocene (MN 4) vertebrate fauna within the area of the eastern part of Central Europe. In addition to a rich fauna of reptiles and mammals, two fossiliferous karst joints (Mokrá-Western Quarry, 1/2001 Turtle Joint and Mokrá-Western Quarry, 2/2003 Reptile Joint) yielded a rich fauna of amphibians including 13 amphibian taxa: Salamandridae: Mioproteus sp., Chelotriton sp., type I, Chelotriton sp., type II, Triturus aff. roehrsi, Triturus cf. marmoratus, Triturus sp. (T. cristatus species group), Chioglossa meini, Mertensiella mera, Salamandridae gen. and sp. indet.; Pelobatidae: Pelobates sanchizi; Ranidae: Rana sp. (synklepton Rana esculenta); Bufonidae: Bufo sp. The first records of the West European species Triturus cf. marmoratus and Chioglossa meini are reported from the eastern part of Central Europe indicating the wide distribution of those taxa throughout the whole of Europe as early as MN 4. The oldest known record of Pelobates sanchizi documents the Early Miocene presence of representatives closely related to the extinct Late Oligocene representatives of Pelobates. The slow evolution of amphibian species is documented by the presence of Triturus cf. marmoratus and the oldest known occurrence of the extinct salamander Mertensiella mera.     

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.     

Ammonites du Toarcien du Haut Atlas central (Maroc)

In the central High Atlas, the Toarcian Stage is represented by a marl and marly limestone series of variable thickness. Two sections have been studied: Amellago (500 m) and Aït Athmane (100 m). Ammonites from the two sections allowed to characterize the Polymorphum, Levisoni, Bifrons, Gradata, and Speciosum zones. The Polymorphum Zone provided, among others: Dactylioceras (Eodactylites) mirabile, D. (E.) pseudocommune, Dactylioceras (Orthodactylites) aff. crosbeyi and Neolioceratoides aff. hoffmanni; the Levisoni Zone provided a rich ammonite fauna composed of, Calliphylloceras nilssoni, Dactylioceras (Orthodactylites) cf. semiannulatum, Eleganticeras exaratum, Harpoceras falciferum, H. pseudoserpentinum, H. serpentinum, H. subplanatum, Hildaites cf. forte, H. levisoni, H. cf. serpentiniformis, H. cf. subserpentinus, H. striatus, H. wrighti, Lytoceras siemensi, Lytoceras sp., Maconiceras soloniacense, Phylloceras sp., Polyplectus pluricostatum et Polyplectus sp.; the Bifrons Zone provided Eleganticeras sp., Harpoceras subplanatum?, Hildoceras bifrons, H. lusitanicum, H. semipolitum, H. sublevisoni, Hildoceras sp., Porpoceras gr. vortex – verticosum; the Gradata Zone was characterized by Pseudocrassiceras bayani, P. frantzi, Pseudocrassiceras sp., Pseudocrassiceras sp. indet.; and the Speciosum Zone provided an Hammatoceras aff. insigne. These species have been described and illustrated for the first time for the central High Atlas. Most of them are common to several basins of the north Tethyan margin and the Subboreal Domain of NW Europe.     

Aubenas-les-Alpes (S-E France). Part III – Last and final part of the mammalian assemblage with some comments on the palaeoenvironment and palaeobiogeography

The study of the new material from Aubenas-les-Alpes makes it possible to add to the faunal list taxa hitherto unknown in this locality (Neurogymnurus cayluxi, Issiodoromys minor, Eomys minor, Pseudocricetodon cf. hausi, Pseudocricetodon aff. philippi). New specimens of the shrew Srinitium marteli increase our knowledge of this rare insectivore. The characteristics of the theridomorph I. minor confirm, as does the whole rodent fauna, the correlation with the MP25 level (Late Rupelian). The assemblage of Aubenas-les-Alpes indicates a rather open and dry environment, in accordance with the Early Oligocene climatic context, but some differences compared to other localities of the same age are also noticed. Finally the mammalian assemblage suggests an ongoing biogeographic differentiation at European scale during the Late Rupelian.     

Microfaunal remains at Middle Pleistocene Qesem Cave, Israel: Preliminary results on small vertebrates, environment and biostratigraphy

Preliminary results of the investigation of the microfauna at the Acheulo-Yabrudian Middle Pleistocene site of Qesem Cave, Israel, are presented. Thus far the assemblage includes ca. 10,000 bone and tooth fragments, of which 50% could be identified to the generic and some hundreds to the species level. Based on the current material, the fauna includes the following squamate reptiles: Laudakia sp., Chamaeleo sp., Gekkonidae indet., Lacertidae indet., Scincidae indet., Pseudopus sp., Varanus sp., Colubroidea indet. (at least three species) and micromammals: Suncus etruscus, Crocidura cf. leucodon, Crocidurinae indet. (large form), Chiroptera indet., Sciurus cf. anomalus, Cricetulus cf. migratorius, Microtus guentheri, Nannospalax ehrenbergi, Dipodillus cf. dasyurus, Meriones cf. tristrami, Gerbillidae indet., Mus cf. musculus, Apodemus cf. flavicollis. These results suggest that the fauna includes only taxa that occur recently in the territory of Israel. The ecological preferences of the nearest living relatives of the recorded taxa allow us to infer a paleoenvironment with a mosaic of open and woodland habitats. However, comparing the lower with the upper levels of the microfauna-bearing profile, a slight shift towards more wooded conditions might be detectable. Biostratigraphical inferences from the recorded micromammal taxa cover a rather wide age range, whereas the radiometric (U-series and preliminary TL) dating enable a provisionally estimated date for the microfauna-bearing levels at 360-300 ka. Detailed morphometric comparisons with material from other sites in the region are necessary and may yet provide further insights.     

Early Jurassic (latest Toarcian) brachiopods from the northeastern margin of the Western Tethys (Central Iran) and their paleobiogeographical significance

Brachiopod fauna from central Iran, recorded in the upper part of the Shemshak Group and attributed to the upper Toarcian (Pseudoradiosa-Aalensis zones), are reported for the first time in Iran. The assemblage recognized includes six different taxa: Homoeorhynchia sepahanensis nov. sp., formally described in this paper, Globirhynchia subobsoleta, Pseudogibbirhynchia sp., Tetrarhynchiidae sp. indet., Monsardithyris? aff. haresfieldensis, and Zeilleria cf. leckenbyi. Analysis of faunal affinities with other paleobiogeographical regions shows a free connection of the central Iranian brachiopod fauna and wide areas of the northern shelf margin of the Tethys Ocean. This is due to an apparent disruption of bioprovinciality inferred for the late Toarcian-earliest Aalenian, congruent with a connection through the northern seaway across the peri-Laurasian epicontinental platforms.     

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 well-preserved ostracod fauna from the middle Burdigalian (lower Miocene) of the North Alpine Foreland Basin

The marine middle Burdigalian sediments of the North Alpine Foreland Basin (NAFB) or Molasse Basin are well known for abundant occurrences of benthic foraminifers and ostracods. However, taxonomic studies of ostracod assemblages are comparatively rare and, given its abundance, the group remains heavily understudied. Here we report a new, rich and well-preserved ostracod fauna from the middle Burdigalian Neuhofen Formation at Mitterdorf, in the northeastern part of the NAFB. The material comprises a total of 3029 ostracod specimens. We identified 44 species; all species are described and, with one exception, illustrated. Our material reveals the presence of 17 ostracod species that were previously unknown from the middle Burdigalian in the NAFB, i.e., Cytherella aff. vulgatella, Cytherella aff. méhesi, Bairdoppilata aff. subdeltoidea, Paranesidea? sp., Paracypris aff. aerodynamica, Parakrithe dactylomorpha, Pseudopsammocythere sp., Callistocythere daedalea, Costa aff. reticulata, Ruggieria aff. longecarenata, Heliocythere aff. vejhonensis, Heliocythere sp., Loxocorniculum hastatum, Neocytherideis cypria, Cytheretta cf. tenuipunctata dentata, Cytheropteron cf. ruggierii, and Semicytherura resecta. The palaeobiogeography of the ostracods indicates a relatively open marine Burdigalian connection between the NAFB and the Mediterranean Sea, in agreement with previous work. Our data also suggest that a significant degree of endemism existed in the middle Burdigalian sea of the NAFB, of which previous studies yielded only sparse signs. Moreover, our results indicate that some ostracod species survived the late Burdigalian sea retreat from the NAFB by dispersing eastward into the adjacent regions (Vienna Basin, Carpathian Foredeep), where marine conditions still prevailed. A compilation of all known middle Burdigalian ostracod faunas from the NAFB documents a high species diversity, with over 120 species. Their spatial distribution, with many species being restricted to a single site, suggests heterogenous microhabitats, possibly influenced by small-scale differences in water depth, nutrient availability and oxygenation.     

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.     

Les grands mammifères du Villafranchien supérieur des travertins du Bassin de Denizli (Sud-Ouest Anatolie,Turquie)

The large mammals from travertine deposits in the Denizli basin include the following species: Archidiskodon meridionalis meridionalis, Equus cf. altidens s. l., E. cf. apolloniensis, Stephanorhinus cf. etruscus, Metacervoceros rhenanus, Cervalces (Libralces) ex gr. minor-gallicus, Palaeotragus sp., Bovinae gen. and sp. indet. This association resembles those from the late Villafranchian of Southern and Eastern Europe, and, to some extent, fromWestern Asia, and could be older than 1.2 Ma.