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.     

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

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

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 late Miocene bovids from Platania (Drama Basin,Greece), with description of a new species of Palaeoryx

This study presents the new fossil material of bovids from the recently discovered upper Miocene locality of Platania, Drama (Greece). The material was excavated from 2012 to 2016 and yielded approximately 760 specimens attributed to hipparions, rhinos, cervids, giraffids, suids, proboscideans, hyaenids, and turtles. The bovid material described here includes six taxa of Antilopinae and one of Bovinae. Antilopines are represented by Gazella sp., Gazella cf. ancyrensis, Prostrepsiceros aff. syridisi, cf. Palaeoreas, Palaeoryx minor nov. sp., and Tragoreas? aff. oryxoides. Bovines are recorded by a single boselaphine attributed to Miotragocerus sp. Departing from other Palaeoryx species, the new species P. minor has rather straight and weakly divergent horn-cores tilted backwards, obtuse facial and occipito-parietal angles, and smaller cranial and horn-core size, though associated with a proportionally large toothrow. Miotragocerus sp. from Platania seems to be conspecific to the Miotragocerus sp. from the end-Vallesian Nikiti-1 fauna (Greece). The bovid assemblage of Platania shows a mix of both Vallesian and Turolian taxa indicating a likely late Vallesian-early Turolian age.     

Rodents and insectivores from the late Miocene of Romerales (Fortuna Basin,Southern Spain)

Abstract

The Neogene Fortuna Basin (Murcia Region, SE Spain) is rich in microvertebrate sites. Its continental sections include localities extended from the middle Turolian (MN12) to the early Ruscinian (MN14). However, there are few works dealing with the taxonomy of these fossil assemblages. In this paper we provide a complete taxonomic study of the rodents and insectivores from three levels (ROM-2B, ROM-2C and ROM-3A) within the Romerales section. We infer a late Turolian age (late Messinian) for these levels, among which the richest and most diverse level is ROM-C, including at least 11 different taxa. In addition, the paleoecological analysis of these fossil assemblages suggests the dominance of open herbaceous meadows under temperate climate during the formation of these sites, with a slight decrease in temperature and humidity from ROM-2B to ROM-2C.     

The parrots (Aves: Psittaciformes) from the Middle Miocene of Sansan (Gers,Southern France)

The occurrence of Archaeopsittacus sp. (Psittaciformes) in the fossil deposits of Sansan (France) is reported, testifying to a survival of this genus in the Middle Miocene, as Archaeopsittacus verreauxi was described from the Early Miocene of Saint-Gérand-le-Puy (France) and was recorded only from its type locality. The data discussed here indicate the presence of two parrot species from Sansan, as it is the type locality of another parrot species, Pararallus dispar, only known from this locality. The differences between the humeri of these two taxa are described in detail, together with the differences from the other European fossil parrot species. The presence of more than one species of parrot in the same locality is not rare, but in Europe it is recorded in Sansan for the second time. Evidence for parrots also confirms the paleoenvironmental reconstruction of Sansan, as parrots are primarily arboreal species. Archaeopsittacus also represents one of the few common elements between the Early and Middle Miocene European bird assemblages.     

Remarks on some Eurasian pliocervines: Characteristics, evolution, and relationships with the tribe Cervini

In order to define the ancestors of the Eurasian genera Axis and Rusa, the representatives of the tribe Pliocervini have been reviewed, with the exclusion of the genus Pliocervus. The fossil remains referred to Pliocervini stored in the Palaeontological Museum of Mechnikov Odessa National University and studied by Khomenko [Khomenko, I., 1913. Meotian fauna of v. Tarakliya of Bendery district. Annuaire géologique et minéralogique de la Russie. 1. The ancestors of modern and fossil Cervinae. 2. Giraffidae and Cavicornia 15, 107-132 (in Russian)] are reappraised: these remains are attributed to Cervavitus novorossiae (Khomenko, 1913). The fossil deer collected from the locality of Novoelizavetovka (Odessa region, Ukraine) and observed in the same Museum of the Odessa National University, is referred to Cervavitus variabilis Alexeev (1915).The fossil material from Shanxi and Honan provinces (China), described by Zdansky [Zdansky, O., 1925. Fossil Hirsche Chinas. Palaeontologia Sinica C 2, 1-94] and Teilhard de Chardin and Trassaert [Teilhard de Chardin, P., Trassaert, M., 1937. The Pliocene Camelidae, Giraffidae and Cervidae of southeastern Shansi.] is referred to Cervavitus shanxius Dong and Hu, 1994. The phylogenetic relationships between Pliocervini from China and the European ones are investigated on the basis of the morphological, biochronological and paleogeographical data. These elements and the suggestions of the more recent genetical and biometrical data may indicate that the origin of the two genera Axis and Rusa could be recognized into the Chinese pliocervines.     

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.     

Percrocuta miocenica (Percrocutidae,Carnivora) from the middle Miocene of Brajkovac (Central Serbia)

This paper describes a well-preserved hemimandible of Percrocuta miocenica from Brajkovac (Lazarevac municipality, Central Serbia). Based on the faunal remains recovered so far, the site is attributed to Mammal Zone MN6 (Langhian, middle Miocene). Apart from Prebreza, this is only the second site where remains of P. miocenica have been found in Serbia, and the third one in the Balkans. The newly discovered fossil, along with other taxa (e.g., Bunolistriodon meidamon, Giraffokeryx punjabiensis, Alloptox sp.), underlines the similarities between the middle Miocene Balkan and Anatolian mammal faunas.     

The latest Miocene Rhinocerotidae from Sahabi (Libya)

The Rhinocerotidae material from the latest Miocene of Sahabi (Libya) is here revised in detail in order to clarify its systematic position and the paleobiogeographic implications. The family is represented by four specimens only at Sahabi, a phalanx, a mandible, a second upper molar (M2), and a second upper premolar (P2). Except for the phalanx, which can be only identified at the family level, the morphology and the dimensions of these specimens have revealed the presence of three taxa: Aceratheriini vel Teleoceratina, Brachypotherium lewisi and ‘Diceros’ sp. The presence of the large-sized B. lewisi has been suggested in several papers, but without a detailed comparison or critical revision. The Brachypotherium from Sahabi also resembles Brachypotherium heinzelini, suggesting a probable synonymy between this species and B. lewisi. A P2 from Sahabi differs from several species belonging to Ceratotherium, including Ceratotherium neumayri, and it resembles the genus Diceros. The rhinoceros association (Brachypotherium and a dicerotine) recognized at Sahabi has been recorded at Lothagam (Kenya), suggesting a biogeographic affinity with the eastern Africa assemblage.     

Rodents and Insectivores from the Upper Miocene of Molayan, Afghanistan

Although Molayan is the richest fossil mammal locality in Afghanistan, only a few species have been studied. This paper describes three rodents (Hystrix aryanensis sp. nov., Parapeolomys sp. and Pseudomeriones latidens sp. nov.) and one insectivore (Erinaceus sp.). Lithostratigraphical correlations and biostratigraphical studies in the Khurdkabul Basin provide evidence that the Molayan locality is younger than three other localities in this basin: Sherullah, Taghar and Ghazgay. In addition, the taxa described here, along with several species of large mammals from Molayan, permit correlations with the Siwalik succession in Pakistan to the east, and with European Neogene mammal ages to the west. The age of the Molayan fauna is concluded to be mid Turolian. Observations on the mammal faunas from the Khurdkabul Basin suggest that the altitude of the area, at present over 2000 m, was much lower during the late Miocene. Uplift of the basin probably occurred no earlier than the ?late Pliocene.     

Earliest example of a giant monitor lizard (varanus, varanidae, squamata)

Background

Varanidae is a clade of tiny (<20 mm pre-caudal length [PCL]) to giant (>600 mm PCL) lizards first appearing in the Cretaceous. True monitor lizards (Varanus) are known from diagnostic remains beginning in the early Miocene (Varanus rusingensis), although extremely fragmentary remains have been suggested as indicating earlier Varanus. The paleobiogeographic history of Varanus and timing for origin of its gigantism remain uncertain.

Methodology/Principal Findings

A new Varanus from the Mytilini Formation (Turolian, Miocene) of Samos, Greece is described. The holotype consists of a partial skull roof, right side of a braincase, partial posterior mandible, fragment of clavicle, and parts of six vertebrae. A cladistic analysis including 83 taxa coded for 5733 molecular and 489 morphological characters (71 previously unincluded) demonstrates that the new fossil is a nested member of an otherwise exclusively East Asian Varanus clade. The new species is the earliest-known giant (>600 mm PCL) terrestrial lizard. Importantly, this species co-existed with a diverse continental mammalian fauna.

Conclusions/Significance

The new monitor is larger (longer) than 99% of known fossil and living lizards. Varanus includes, by far, the largest limbed squamates today. The only extant non-snake squamates that approach monitors in maximum size are the glass-snake Pseudopus and the worm-lizard Amphisbaena. Mosasauroids were larger, but exclusively marine, and occurred only during the Late Cretaceous. Large, extant, non-Varanus, lizards are limbless and/or largely isolated from mammalian competitors. By contrast, our new Varanus achieved gigantism in a continental environment populated by diverse eutherian mammal competitors.     

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 Messinian rodents from Verduno (Piedmont,NW Italy): Biochronological,paleoecological and paleobiogeographic implications

The stratigraphic and paleoenvironmental context of the Verduno fossil vertebrate locality is discussed herein based on its rodent record. The Verduno section crops out in the southern part of the Tertiary Piedmont Basin (TPB), and can be included in the Messinian post-evaporitic Cassano Spinola Fm., chronologically corresponding to the so-called Lago-Mare event. Rodents are represented by a relatively rich assemblage. Murids are by far the most diverse and abundant, with at least four taxa, including the common Centralomys benericettii and Paraethomys meini, and the rare Apodemus gudrunae and Occitanomys sp. Cricetids are represented by a single species, Apocricetus cf. A. barrierei. Muscardinus aff. M. vireti appears to be the only glirid present at Verduno. The Verduno rodent assemblage shares some taxa with other Messinian post-evaporitic localities from Italy bearing continental vertebrate remains, such as Brisighella (central Italy) and Moncucco Torinese (NW Italy) (e.g., C. benericettii, P. meini) and, possibly, with Ciabòt Cagna (NW Italy). However, the general structure of these four Messinian assemblages displays substantial differences, which may reflect different palaeoenvironmental conditions.     

A synopsis of the late Miocene Mammal Fauna of Samos Island,Aegean Sea,Greece

As one of the oldest known Eurasian fossil vertebrate localities, Samos late Miocene fauna attracted the interest of specialists by its richness and overall importance. Nevertheless, crucial taxonomical questions and chronological problems obscured its value. The detailed study of the local stratigraphy, the collection of new fossil material and its study, the revision of the old collections and the updated magneto-chronology of the fossiliferous deposits permited to re-discuss most of the problems in a special volume edited in 2009 by Koufos and Nagel and to provide a clearer and more precise idea about the Samos fauna and its age. A synopsis of this work is given here. The systematic study of the new collection (～1200 identified specimens) allows the determination of 42 species from three fossil horizons, ranging from the upper part of early Turolian (MN11) to the end of middle Turolian (MN12). Taxonomic novelties are the presence of the carnivore genus Protictitherium found for the first time in Samos, the establishment of the new name Skoufotragus for Pachytragus Schlosser with the new species Skoufotragus zemalisorum, and the amended morphology of Pseudomeriones and Urmiatherium. Additionally six Hipparion and four Gazella species were recognized and a better morphometric distinction between Samotherium boissieri and Samotherium major was performed. This study also improved the correlation of the old fossiliferous sites with the new ones and with the local stratigraphy of the Mytilinii Basin, while precise ages have been obtained for the mammal localities. The new data together with the old collections indicate the presence in Samos of four chronologically successive mammal assemblages reflecting a “four stages-of-evolution” scheme. The Turolian palaeoenvironment of Samos is determined as an open bushland with thick grassy-floor of C3 graminoids with possible increase of the open and dry character from the beginning to the end of Middle Turolian. The Samos mammal faunas are palaeobiogeographically closer to the Asian ones than to those from the Greek mainland.     

A new species of Megacerini (Cervidae, Artiodactyla) from the Late Miocene of Taralyk-Cher, Tuva (Russia), and remarks on the relationships of the group

A new species of a megacerine deer, Praesinomegaceros venustus, from the Taralyk-Cher locality in the Tuva region on the south of Eastern Siberia, Russia, is described. P. venustus nov. sp., the oldest known megacerine, occurred in Asia in the Late Turolian, about 7 Ma. The early history of megacerines is discussed. The existence of the phyletic lineage Cervavitus-Praesinomegaceros-Sinomegaceros in Asia is confirmed.     

The family Phryganeidae (Insecta,Trichoptera) from the Mesozoic and Cenozoic of Asia (with a brief worldwide overview of the fossil caddisfly fauna)

The following new taxa of the family Phryganeidae are described: Proagrypnia sinitsae gen. et sp. nov. from the Upper Jurassic or Lower Cretaceous of Russia (Chernovskie Kopi locality) and Baissophryganoides popovi sp. nov. from the Lower Cretaceous of Mongolia (Anda-Khuduk locality). A larval case of Folindusia (Acrindusia) phryganoides sp. nov. from the Paleocene of Russia (Arkhara–Boguchan brown coal field) is described. A worldwide overview of the records of fossil adult insects and larval cases of Phryganeidae is presented.