Miocene mammalian faunas from Wushan,China and their evolutionary,biochronological, and biogeographic significances

We report Miocene mammalian faunas from three nearby localities: Kangping, Nanyu, and Yangping, in the Wushan Subbasin, Gansu Province, China. From the Kangping locality, we identified four species, Platybelodon grangeri, Hispanotherium wushanense n. sp., Kubanochoerus sp., and Turcocerus cf. kekemaidengensis, which indicate that this fauna can be correlated with MN7/8. From the Nanyu locality, we identified Platybelodon aff. tongxinensis, which, together with the previously reported specimens of Gomphotherium wimani and Micromeryx cf. flourensianus, enables us to correlate this fauna with MN6. The Yangping locality is found to be early Miocene in age based on the presence of cf. Gomphotherium sp. This occurrence is the earliest record of proboscideans in China, and thus represents the “Proboscidean Datum Event”. Evolutionary trends of Platybelodon and Hispanotherium are figured and discussed. The possibility of a geographic boundary associated with the eastern and western differentiation of species of Turcocerus and bunodont listriodonts in China is also evaluated. This boundary was consistent with the major division of the eastern and western mammalian faunas of the middle Miocene in China. The localities in Inner Mongolia suggest a large area with transitional habitats.     

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 species of Amphilagus (Lagomorpha, Mammalia) from the Miocene of the Valley of Lakes, central Mongolia

New lagomorph species, Amphilagus orientalis sp. nov., A. plicadentis sp. nov., and A. magnus sp. nov. from Lower Miocene localities (Biozone D) of the Valley of Lakes in Central Mongolia are described.     

Lagomorphs from the Miocene of Sandelzhausen (southern Germany)

Three genera of lagomorphs, Prolagus, Lagopsis, and “Amphilagus,” were identified during a revision of the lagomorph material from Sandelzhausen (MN5, Early/Middle Miocene boundary, southern Germany). Evidence of two morphological and dimensional classes were observed at some tooth positions in Prolagus (some p3 show an unmistakable P. oeningensis morphology, others closely resemble P. crusafonti), but not at other tooth positions (e.g., M1–2). Insufficient data from Sandelzhausen precludes identification of two different species of Prolagus from this locality, and to define the characteristics of the possible P. crusafonti-like species. Thus, all Prolagus specimens have been classified as P. aff. oeningensis. The genus Lagopsis is represented by L. cf. penai, whose presence is compatible with a MN5 age. The relative abundance of Lagopsis to Prolagus may indicate relatively cool and wet palaeoclimatic conditions. The largest primitive lagomorph species from continental Europe is present at Sandelzhausen. Morphological and dimensional comparisons with other European primitive lagomorphs exclude any affinity with the genera Eurolagus and Titanomys and with the species included in “Amphilagus ulmensis”. Some common features with “Amphilagus antiquus” were observed, although they are not sufficient for the attribution to this taxon. Until there is a general revision of European primitive lagomorphs, the Sandelzhausen giant lagomorph is classified as “Amphilagus” sp. Its origins, whether from evolution within Europe or migration from Asia, remain unknown.      

Re-interpreting the biochronology of the La Celia and Los Gargantones mammal sites (Late Miocene,Murcia, Spain)

The biochronological age of the small-mammal populations of Los Gargantones 1, 2 and La Celia (upper Miocene, La Celia sub-basin, Murcia, Spain) is re-interpreted. The presence in Los Gargantones of Occitanomys adroveri, Parapodemus barbarae, Parapodemus cf. gaudryi, Huerzelerimys turoliensis, Atlantoxerus cf. adroveri, and Alilepus evidences a correlation to MN12 (∼7.5–7 Ma) rather than to MN11 (∼9–7.5 Ma), as inferred previously. The assemblage corresponds to that of the more eastern, near-coast sites of Crevillente 8 and 15, situated in the Alicante area. The stratigraphically highest site of La Celia contains Hispanomys adroveri, a species also indicative of MN12. The presence of Castromys cf. littoralis together with an advanced O. adroveri points to a slightly younger age than that of Los Gargantones, approaching that of MN13 sites. The assemblage best matches that of Crevillente 17. Other species described in this paper are Prolagus crusafonti, Prolagus sp., Parasorex cf. ibericus, Panelimnoecus cf. repenningi, and Blarinella aut Petenyia sp. indet.     

Les Megacricetodon du gisement karstique miocène moyen de Blanquatère 1 (Pyrénées-Orientales, Sud de la France) : nouvelles espèces, implications biochronologique et phylogénique

Three Megacricetodon species are recognized in the Middle Miocene locality of Blanquatère 1 (Languedoc-Roussillon province, France). Two are new, the small-sized M. tautavelensis nov. sp. the medium to large-sized M. aunayi nov. sp., the third one being the already known M. “collongensis-gersii”, a medium-sized species found in other Miocene localities of the area. Size and morphology differentiate these species. Such a high number of contemporaneous species of the genus Megacricetodon is found for the first time in a Miocene locality in the region and indicate that the evolution of this genus in western Europe is likely more complicated. It may testify both of a biogeographical differentiation and of migration events. According to our calibration charts, the deposit is dated ca. 16 Ma ago. The position of Vieux-Collonges (MN 5) and Sansan (MN 6) on the biochronological scale is also discussed, as well as the estimated age of the biozonal boundaries MN 3/MN 4, MN 4/MN 5 and MN 5/MN 6.     

Eomyidae fauna turnover at the Early-Middle Miocene boundary in the Morteral section (Magro Basin,Valencia, Spain)

The material of Eomyidae from three localities (Morteral 1, Morteral 20A and Morteral 22) of the Morteral section in the Magro Basin, eastern Spain, is described. These localities contain remains of Ligerimys and Eomyops. Morteral 1 is situated in the lowest part of the section and contains Ligerimys ellipticus, whereas Morteral 20A and Morteral 22 are situated in the middle part of the section and contain, among other rodent species, Eomyops noeliae. The youngest localities in the Morteral section (Morteral 41 and 44) record Megacricetodon collongensis, first representative of Megacricetodon in the Middle Miocene (MN5). The locality Morteral 20A contains Megacricetodon primitivus; Morteral 1 and Morteral 22 contain a form of Megacricetodon different from both M. primitivus and M. collongensis, therefore, these three localities are of lower Aragonian age (MN4). In the Aragonian type area, L. ellipticus is rare in zone B (lower part of MN4) and frequent in zone C (upper part of MN4). The first record of Eomyops in the Aragonian type area occurs in MN7/8 and in MN5 in Central Europe. The turnover of eomyid faunas in MN4 localities of the Morteral section clearly confirms the hypothesis of Engesser (1999) that Eomyops may have appeared earlier than supposed so far (MN5).     

Messinian Lago-Mare mollusc fauna from the Gorgona Island slope, Tyrrhenian Sea

Late Miocene Lago-Mare macrofossiliferous sediments were recovered in the northeastern Tyrrhenian Sea by dredging the continental slope off Gorgona Island, Tuscan Archipelago, at 300-470 m depth. The fossil assemblage consists of a rich lymnocardiid bivalve fauna dominated by Pontalmyra ex gr. P. incerta (Deshayes), associated with Dreissena ex gr. D. rostriformis (Deshayes), Pontalmyra cf. partschi (Mayer), “Limnocardium” sp., the gastropods Melanopsis narzolina (D’Archiac), Melanopsis sp. and cf. Saccoia sp. All bivalve taxa recognized at species level are of Paratethyan (Pontian) affinity and widespread in the Late Miocene of the Mediterranean Basin while M. narzolina has so far only recorded from the Mediterranean Basin. This finding represents the most diverse Lago-Mare macrofauna reported thus far from any submerged location in the Mediterranean Basin and documents that the post-evaporitic Cusercoli Formation contributes to the syn-rift neoautochthonous units of this sector of the Northern Tyrrhenian Sea.     

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.     

Une nouvelle espèce du genre Bohlinia (Mammalia,Giraffidae) du Miocène supérieur de Toros-Menalla,Tchad

New Giraffidae specimens discovered from the Late Miocene fossiliferous sector of Toros-Menalla (northern Chad) are described. These specimens are assigned to Bohlinia, a genus known so far only in some Late Miocene localities of Eurasia. This is the first occurrence of the genus in Africa. Analysis of the material allows us to describe a new species of giraffid, Bohlinia adoumi n. sp., probably close to the Eurasian forms.     

New insights on Cricetodontini (Rodentia,Mammalia) from the Duero Basin,Spain

Abstract

The tribe Cricetodontini is a common cricetid group found in several European basins from the Miocene. Here we present a study of the fossils of this group found in the Duero Basin. We updated the biostratigraphical assignation of some of the localities in which several species of Cricetodontini have been found. Cricetodontini remains from eight localities from the central sector of the Duero Basin have been described, measured and assigned to a species. The presence in these localities of Hispanomys aguirrei, H. lavocati, H. nombrevillae and H. aragonensis has allowed correlating them to the biostratigraphic scale built for the Miocene in Calatayud-Daroca Basin, identifying biozones G3, H and I (MN7/8 – MN10, Late Aragonian – Early Vallesian, Middle – Late Miocene). Furthermore, this study constitutes the first citation of this species in this basin, except H. aguirrei, previously described in the Duero Basin. After this work, the biostratigraphical assignation of the studied sites is now well known. We evidenced the resemblance of Duero and Calatayud-Daroca basins.     

Italian Cenozoic crocodilians: taxa, timing and palaeobiogeographic implications

Crocodylian remains are collected in 39 fossil-bearing localities but only in seven localities specimens with reliable taxonomic attributions, at least to genus level have been collected. Three species have been reported from the early Lutetian Purga di Bolca site: Pristichampsus cf. Pristichampsus rollinati, Asiatosuchus sp., Hassiacosuchus sp. (=Allognathosuchus sp.). The three crocodilians discovered at Purga di Bolca have been reported also from Geiseltal and Messel (Middle Eocene, Germany). Bolca at that time was part of a Tethysian archipelago and no mammals have been found there till now. Crocodilians and turtles clearly arrived from the European mainland across a marine water barrier. Among the other fossiliferous localities of Veneto, very interesting is the Monte Zuello site, of late Middle Eocene age, yielding a longirostrine crocodilian, Megadontosuchus arduini, a tomistomine species. Tomistomines are known in contemporaneous sediments of both Europe and Africa, but the European forms Dollosuchus and Kentisuchus seem the closest taxa. Remains of Oligocene age have been collected in Veneto and Liguria, but the fossils discovered in the second region are teeth or fragmented bones. The fossil crocodilians of Monteviale (Veneto), of Early Oligocene age, have been assigned to two species but they have been recently all identified as Diplocynodon ratelii, known from several European sites of Late Eocene, Oligocene and Miocene age. This species arrived in the Monteviale area from the European mainland across a narrow sea. Several crocodilian fossils of Miocene age are very fragmentary or represented by isolated teeth. In the Middle and Late Miocene of Sardinia, a well-established species, Tomistoma calaritanum is present. Remains of Tomistoma of the same age have been reported in some localities in Tuscany, Apulia, Sicily and Malta. In the Mediterranean area, the genus is known from European and African sites (of older age). The colonisation of Europe by this genus is the result of a dispersion from Africa (or less probably from Asia). During Late Miocene Sardinia and Tuscany belong to the same palaeobioprovince characterized by the Oreopithecus-Maremmia fauna. In Tuscany, a crocodilian identified as Crocodylus bambolii is present in the late Miocene site of Monte Bamboli. If the generic attribution of this form is correct, its ancestors must have arrived from Africa. Another fossil assemblage of Late Miocene age characterizes the Apulia-Abruzzi palaeobioprovince (Hoplitomeryx-Microtia fauna) and testifies complete isolation between the two palaeobioprovinces. In this last area, remains of Crocodylus sp. have been collected in coastal sandstones at Scontrone (Abruzzi) and in several fissure fillings of Gargano of slightly younger age. The ancestors of this species arrived from Africa while no African elements are present among the mammalian fauna. The dispersion of the genus Crocodylus in the Italian palaeoislands may have taken place once, with allopatric differentiation of the two populations (Tuscany-Sardinia and Apulia-Abruzzi) or twice with independent colonisation of each area.     

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.     

Snake fauna (Reptilia: Serpentes) from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 (Germany)

The Early/Middle Miocene (European Land Mammal Zone MN5) localities of Sandelzhausen and Rothenstein 13 in southern Germany have yielded remains of about 13 ophidian taxa: Eoanilius sp. (Aniliidae), Bavarioboa ultima (Boidae), “Coluber” sp., ?Telescopus sp., Natrix sp., cf. Natrix sp., cf. “Neonatrix” sp., unidentified “colubrines” and “natricines” (Colubridae), Naja sp., an unidentified elapid (Elapidae), Vipera sp. (“Oriental viper”), Vipera sp. (“aspis complex”) (Viperidae). Both faunas document a transitional phase from those reported from several late Early and Middle Miocene sites of Central and Western Europe. The climates of Sandelzhausen and Rothenstein 13, as indicated by ophidian fossils, were warm, although not tropical or subtropical.     

Ostracodes et stratigraphie du néogène et du quaternaire méditerranéens

A stratigraphical chart of marine ostracoda from Lower Miocene to Recent is established. Selected species (approximatively 220) are those morphologicaly well characterized and known from different parts of the Mediterranean area. It appears that: • lower Miocene ostracodes are still poorly known; • specific diversity is high during the Tortonian and the Lower Messinian before the complete disappearance of marine Mediterranean species during the Upper Messinian évaporitic episodes; • during the early Pliocene, about half of the Upper Miocene marine species are reintroduced with the Atlantic waters; other species migrate for the first time in the Mediterranean Sea by the same way; • at the end of the Pliocene or at the beginning of the Pleistocene several species known in Mediterranean since the Middle Miocene or before, such as Cytherella sp. gr. transversa and Ruggieria tetraptera, as well some “nordic guests” such as Hemicythere villosa and Cythere lutea, appear. This work is an opportunity to confirme a Late Miocene age for the Neogene of Skyros (Aegean Sea), to assign the “Upper Pliocene” of Terquem to the Lower Pleistocene and to refute the existence of a pliocene psychrosphere.     

New Late Miocene murids from China (Mammalia, Rodentia)

New murids of Late Miocene (medial Baodean Chinese Mammal Unit) age from Inner Mongolia, northern China, and from Yunnan Province, southern China, are described. Hansdebruijnia perpusilla nov. sp. represents the earliest known and morphologically most primitive species of the genus, which is known from the latest Miocene of Europe and western Asia. The new species suggests an eastern Palaearctic origin of Hansdebruijnia. “Progonomys” yunnanensisQIU and STORCH, 1990 from Lufeng, Yunnan Province, is referred to Linomys nov. gen. New samples from Leilao, Yuanmou County, Yunnan Province, are included in this species, although this population is somewhat more primitive than that from Lufeng. Leilaomys zhudingi nov. gen. and sp. from Leilao shows a unique combination of apomorphic and plesiomorphic characters, which suggest an early divergence from the murid stem. The new findings indicate that our knowledge of the early radiation of murids in southeast Asia is still in its infancy.     

Miocene hyaenids (Hyaenidae; Carnivora) from Siwaliks of Pakistan

Newly discovered Miocene hyaenid specimens, recently collected from the Siwalik Group, are described and discussed. A careful comparison with the known material reveals that these specimens belong to the early hyaenid species Thalassictis cf. T. proava, Ictitherine indet. and Lepthyaena sivalensis. The stratigraphic range of T. proava extended up to the Dhok Pathan Formation (Middle Miocene to Early Pliocene). The stratigraphic range of T. proava comprises the Middle to Late Miocene, with the youngest record in Hasnot, Potwar Plateau in the Siwalik Group. The material is of great interest because Siwalik carnivoran material is rare.     

Vertebrates from the Pontian of the Shkodova Gora locality (Northwestern Black Sea Region,Upper Miocene)

Remains of fishes, turtles, birds, and mammals are described from the Upper Miocene (MN13) Shkodova Gora locality (Ukraine, Odessa Region), the only representative locality of Pontian large vertebrates in the northwestern Black Sea Region. Asiatic fish and avian taxa are recorded for the first time in the Upper Miocene of Europe. The Shkodova Gora ichthyofauna includes eight freshwater fish species (belonging to Acipenseridae, Cyprinidae, Siluridae, and Percidae). Taxa that have previously been recorded only in Western Siberia (Abramis bliccoides, Perca lepidopoma), eastern Kazakhstan, and the Altai Mountain (Rutilus tungurukensis) are revealed. Acipenser gueldenstaedtii is recorded for the first time in the Upper Miocene of Eastern Europe. The occurrence of the genus Ctenopharyngodon, a member of the Chinese plain faunal assemblage is of great interest. The Shkodova Gora avifauna (seven taxa) is mostly composed of waterfowl, including Phalacrocorax mongoliensis, which has previously been known only from the Lower Pliocene of Mongolia. The finding of Proanser major is the youngest record of this species in the fossil record. Mammals of the Shkodova Gora association include a hare resembling Trischizolagus dumitrescuae, Palaeoryx cf. pallasi, and Hipparion cf. moldavicum, which have not been mentioned among Pontian taxa of the northern Black Sea Region. Pontian paleolandscapes in the area of the Shkodova Gora locality are reconstructed.