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.     

Late Miocene (Turolian,MN13) squirrels from Moncucco Torinese,NW Italy

Squirrel remains are rare in the fossil record, which hampers a properly documented interpretation of their evolutionary history. The Late Miocene sciurids from Moncucco Torinese (NW Italy) are described based on a diverse tooth assemblage. The data presented herein confirm the presence of Sciurus warthae and Pliopetaurista pliocaenica, and also include the first unquestionable occurrence of Hylopetes hungaricus plus an indeterminate Sciurinae. The report of Sciurus warthae from Moncucco Torinese represents the earliest record of the genus Sciurus in Europe. Moreover, Moncucco Torinese is the first European locality recording the co-occurrence of Sciurini and Pteromyini in the Late Miocene. Comparative analyses support a close relationship between S. warthae and Sciurus vulgaris corroborating the hypothesis of an earliest divergence of the “Sciurus vulgaris clade” in the Late Miocene. Biochronological, paleobiogeographical and paleoecological implications are also discussed.     

Kretzoiarctos gen. nov., the Oldest Member of the Giant Panda Clade

The phylogenetic position of the giant panda, Ailuropoda melanoleuca (Carnivora: Ursidae: Ailuropodinae), has been one of the most hotly debated topics by mammalian biologists and paleontologists during the last century. Based on molecular data, it is currently recognized as a true ursid, sister-taxon of the remaining extant bears, from which it would have diverged by the Early Miocene. However, from a paleobiogeographic and chronological perspective, the origin of the giant panda lineage has remained elusive due to the scarcity of the available Miocene fossil record. Until recently, the genus Ailurarctos from the Late Miocene of China (ca. 8–7 mya) was recognized as the oldest undoubted member of the Ailuropodinae, suggesting that the panda lineage might have originated from an Ursavus ancestor. The role of the purported ailuropodine Agriarctos, from the Miocene of Europe, in the origins of this clade has been generally dismissed due to the paucity of the available material. Here, we describe a new ailuropodine genus, Kretzoiarctos gen. nov., based on remains from two Middle Miocene (ca. 12–11 Ma) Spanish localities. A cladistic analysis of fossil and extant members of the Ursoidea confirms the inclusion of the new genus into the Ailuropodinae. Moreover, Kretzoiarctos precedes in time the previously-known, Late Miocene members of the giant panda clade from Eurasia (Agriarctos and Ailurarctos). The former can be therefore considered the oldest recorded member of the giant panda lineage, which has significant implications for understanding the origins of this clade from a paleobiogeographic viewpoint.     

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

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

First fossil record of the catfish genus Sorubim (Siluriformes,Pimelodidae), from the Late Miocene of Paraná, Northeastern Argentina

The aim of the present contribution is to describe a partial fossil skull belonging to the catfish genus Sorubim. The specimen originates from the Late Miocene Ituzaingó Formation, at Paraná locality, Entre Ríos Province, Argentina. The material described here (MACN Pv-14224) was determined just to genus because the living species of Sorubim are morphologically similar. Presence of Sorubim is in concordance with the hypothesis indicating that most extant pimelodid genera were already present by the Late Miocene. Present finding constitutes the first fossil record for the genus.     

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.     

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.     

Fossil fruits of Firmiana and Tilia from the middle Miocene of South Korea and the efficacy of the Bering land bridge for the migration of mesothermal plants

Determining whether the high-latitude Bering land bridge (BLB) was ecologically suitable for the migration of mesothermal plants is significant for Holarctic phytogeographic inferences. Paleobotanical studies provide a critical source of data on the latitudinal positions of different plant lineages at different times, permitting assessment of the efficacy of the BLB for migration. Here we report exceptionally preserved fossils of Firmiana and Tilia endochrysea from the middle Miocene of South Korea. This represents a new reliable record of Firmiana and the first discovery of the T. endochrysea lineage in the fossil record of Asia. The occurrence of these fossils in South Korea indicates that the two lineages had a distribution that extended much farther north during the middle Miocene, but they were still geographically remote from the BLB. In light of the broader fossil record of Asia, our study shows that, in the middle Miocene, some mesothermal plants apparently inhabited the territory adjacent to the BLB and thus they were possibly capable of utilizing the BLB as a migratory corridor. Some other mesothermal plants, such as Firmiana and the T. endochrysea lineages, however, are restricted to more southern regions relative to the BLB based on current fossil evidence. These lineages may have been ecologically unable to traverse the BLB, which raises questions about the efficacy of the BLB as a universal exchange route for mesothermal plants between Asia and North America during the middle Miocene.     

Amphibians and reptiles from the late Miocene and early Pliocene of the Ptolemais area (Western Macedonia,Greece)

We herein describe the amphibians and reptiles from the Ptolemais fossil assemblage, originating from 12 nearby localities in northwestern Greece, spanning from the late Miocene (MN 13) to the early Pliocene (MN 15). Amphibians are known exclusively of anurans, with the genera Latonia and Rana being identified, the latter constituting the oldest so far known record of that lineage in Greece. Turtles are represented by the testudinid cf. Testudo, whereas numerous other indeterminate testudinoids are known. Lizards include scincids, lacertids, and at least two anguids. Among them, the material referred to scincids and the anguid Anguis constitute one of only rather few such occurrences described in the fossil record of the Eastern Mediterranean. Snakes are represented by indeterminate “colubrines” and the genus Natrix. The different ages of the Ptolemais fossiliferous localities, along with their close geographic vicinity offer the opportunity to study potential survival/extinction patterns of its amphibians and reptiles across the Mio-Pliocene boundary, a time interval that has been crucial for European herpetofaunas.     

A Late Miocene Paratethyan mollusc fauna from the Denizli Basin (southwestern Anatolia, Turkey) and its regional palaeobiogeographic implications

We report on a Late Miocene mollusc fauna from a single locality in the Denizli Basin in southwestern Turkey that is composed of ten presumably euryhaline species and three freshwater species. The fauna is remarkably distinct from faunas of the adjacent Late Miocene Aegean and Euxinian Basins, and has elements in common with the modern Caspian Sea fauna in the form of Didacna species. The suggested age of the fauna (Late Miocene) would extend the stratigraphic range of the lymnocardiid genus Didacna (hitherto Calabrian-Extant) considerably. The palaeobiogeographic significance of the Denizli fauna is discussed.     

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.     

Evolution of bird communities in the Neogene of Central Asia,with a review of the Neogene fossil record of Asian birds

A complete taxonomic review of Neogene birds of continental Asia is provided. To date, avifauna from the latter half of the Miocene and Pliocene of Central Asia (Mongolia and adjacent regions of Inner Asia) are most thoroughly investigated. Available data enable a reconstruction of successive replacement of Early and Middle Miocene avifaunas by communities of the Recent type. Middle Miocene avifaunas of Mongolia include a great number of extinct genera and species, many of which were widespread in Eurasia. Extant genera became dominant in the Late Miocene and taxa close to living species appear in the Late Pliocene fossil record. Late Pliocene communities of birds of Central Asia were complex in genesis, composed of Miocene relicts (Struthio), immigrants from the European regions of the Palearctic (phasianid Plioperdix), North American immigrants (Calcarius), and also autochthonous elements, the origin of which is apparently connected with the arid belt of Central Asia (diverse passerines).     

Şerefköy-2, a new Late Miocene mammal locality from the Yatağan Formation,Muğla,SW Turkey

Here we report on a new fossil locality, ?erefköy-2, from the Yata?an Basin of southwestern Turkey that preserves a well-sampled, abundant, and diverse mammal fauna. Indeed, after three field seasons, more than 1200 catalogued specimens representing 26 mammal species belonging to 14 genera make the ?erefköy-2 mammalian assemblage one of the richest Late Miocene fauna from Anatolia. Five hipparionines, six bovids, including the rare and enigmatic Urmiatherium rugosifrons and the presence of Pliohyrax graecus, strongly support affinities with Late Miocene faunas from Samos Island, Greece. Through a consideration of the identified material and the subsequent comparison with material from well-known Balkan and Anatolian faunas, a Middle Turolian (MN12) age for ?erefköy-2 is indicated.     

The southernmost sirenian record in the eastern Pacific Ocean, from the Late Miocene of Chile

A tooth of a sirenian from the Late Miocene sediments of the Bahia Inglesa Formation (Chile) is described and referred to the Dugongidae. The fossil represents the first sirenian record from Chile and the southernmost record of the Sirenia in the eastern Pacific Ocean (latitude 27° S). The Chilean record extends the already wide geographical distribution of fossil sirenians along the Eastern Pacific coast. The presence of a sirenian during the Miocene on the Chilean coast is related to a globally warmer climatic condition and a still limited northern extension of the cold Humboldt Current. To cite this article: G. Bianucci, C. R. Palevol 5 (2006).     

Scolecophidia (Serpentes) of the Late Oligocene and Early Miocene,North America,and a fossil history overview

An abundant fossil record of the snake clade Scolecophidia exists in Europe; however, the minute snake is noticeably absent in reports about the North American Paleogene and Neogene. Presented here are four localities from Florida, USA, that contain scolecophidian remains older than the Pleistocene: Thomas Farm (late Early Miocene, Hemingfordian Land Mammal Age, LMA), Live Oak (Oligocene-Miocene transition, latest Arikareean LMA), White Springs 3B (late Arikareean LMA), and Brooksville 2 (Late Oligocene, middle Arikareean LMA). These remains extend their known existence by about 26 m.y. and are now the oldest reported scolecophidian remains in North America. Molecular evidence on extant scolecophidians concludes that these tiny snakes have a Gondwanan origin. Interestingly, the oldest record of a scolecophidian is from Europe (Belgium) and dates back to the middle Paleocene (MP 1–5). The earliest African record of the snake clade comes from the Paleocene-Eocene boundary in Morocco. The clade is apparently absent from Europe and Middle East deposits dating from the latest Eocene through to the latest Oligocene (MP 19–30) and to the Early Miocene (MN 4). A portion of this time is known as the booid ‘Dark Period’ which represents an apparent response to global aridization and cooling. Scolecophidians appear to re-emerge into the southern Eurasian record in the Early Miocene (MN 4) and become widely dispersed throughout Europe and Middle East. The fossil record of these minute snakes is largely absent in southern Asia and South America. It is possible that the current lack of a decent fossil scolecophidian record outside of Europe and Middle East is due mainly to a bias in the methodology to recover fossils; wet sieving sediments through < 1.0 mm mesh is needed to recover the minuscule vertebrae.     

Révision du genreProstrepsiceros Major 1891 (Mammalia,Bovidae)

The study of new fossils from the Miocene of Macedonia (Greece) has led us to a revision of the genusProstrepsiceros, which we propose to divide into two subgenera:P. (Prostrepsiceros) andP. (Helicotragus), these subgenera comprising respectively two and three species.     

The westernmost tarsier: A new genus and species from the Miocene of Pakistan

As the closest living sister group of anthropoids, tarsiers (Family Tarsiidae) are an important group in primate evolution. However, their fossil record is poor: only four species have been described, two from the Eocene of China and two from the Miocene of Thailand. All are from outside the range of the living species, which occur only on islands off Southeast Asia. Here, we describe a new fossil tarsier from Pakistan, a significant range extension. This record consists of two lower molars, an upper molar, and a lower premolar found in the Miocene Manchar Formation (∼18–16 Ma [millions of years ago]) of Sindh Province, southern Pakistan. The Pakistani tarsier is morphologically distinct from all living and fossil tarsiers, but most similar to the middle Miocene Thai species Tarsius thailandicus. Though living tarsiers have traditionally been classified in a single genus, a recent revision proposed a division into three genera, which is strongly supported by molecular data. The Pakistani species is not referable to any of these genera, and we create for it and T. thailandicus a new tarsiid genus. This discovery broadens our understanding of the geographic range and morphological diversity of Miocene tarsiers and helps to put the living tarsiers into their evolutionary context.