The Scontrone turtles – A new insular testudinoid fauna from the late Miocene of the Central Mediterranean

We here describe a small turtle assemblage originating from the early Tortonian (late Miocene) palaeois-land of Scontrone, central Italy, a locality previously known mostly for its endemic mammals and giantbirds, which were otherwise shared only with the Gargano localities, another fossiliferous area belongingto the same palaeobioprovince. The fossil turtle remains from Scontrone are referred to the geoemydidMauremyssp. and a so far unidentified large-sized testudinid. The biogeographic origins of theScontrone insular chelonians are discussed. The Scontrone geoemydid adds to the known occurrencesofMauremysin the late Miocene of the Mediterranean. The Scontrone large tortoise represents the oldestknown Mediterranean insular testudinid, predating significantly the well-known Quaternary endemicisland tortoises of the area

Crushed but not lost: a colubriform snake (Serpentes) from the Miocene Swiss Molasse, identified through the use of micro-CT scanning technology

An incomplete postcranial skeleton of a snake from the middle Miocene of the Swiss Molasse in Käpfnach mine, near Zurich, Switzerland, is described in this paper. The skeleton is rather crushed and resting on a block of coal, with only some articulated vertebrae partially discerned via visual microscopy. We conducted micro-CT scanning in the specimen and we digitally reconstructed the whole preserved vertebral column, allowing a direct and detailed observation of its vertebral morphology. Due to the flattened nature of the fossil specimen, several individual vertebral structures are deformed, not permitting thus a secure precise taxonomic identification. Accordingly, we only refer the specimen to as Colubriformes indet. Nevertheless, this occurrence adds to the exceedingly rare fossil record of snakes from Switzerland, which had so far been formally described solely from three other Eocene and Miocene localities

Necrosaurus or Palaeovaranus? Appropriate nomenclature and taxonomic content of an enigmatic fossil lizard clade (Squamata)

Necrosaurus cayluxi est un lézard énigmatique du Paléogène des Phosphorites du Quercy, France, qui a été mentionné pour la première fois au 19e siècle. Bien qu’il soit généralement admis que Filhol est l’auteur qui a établi ce taxon, je démontre ici que la paternité devrait en fait en être attribuée à Zittel, ce qui influence également non seulement la nomenclature générique, mais aussi le matériel type approprié. En tant que tel, le nom valide pour ce taxon devrait être Palaeovaranus cayluxi et son holotype est un maxillaire gauche. En outre, Ophisauriscus eucarinatus de l’Éocène Moyen de Geiseltal, Allemagne, autre taxon précédemment assigné à Necrosaurus, est considéré comme étant un nomen dubium, tandis que Melanosauroides giganteus de la même localité est considéré comme une espèce valable et est recombinée comme Palaeovaranus giganteus comb. nov. Les changements suggérés dans la nomenclature affectent également les « Necrosauridae », un clade mal défini de lézards du Crétacé–Paléogène d’Europe, d’Amérique du Nord et d’Asie. Afin de maintenir la stabilité de la nomenclature et de définir une lignée monophylétique, j’établis ici la nouvelle famille Palaeovaranidae fam. nov., qui comprend uniquement le genre Palaeovaranus. Les exemples connus de Palaeovaranus à travers le Paléogène d’Europe occidentale sont discutés.Necrosaurus cayluxi is an enigmatic lizard from the Paleogene of the Phosphorites of Quercy, France that was first mentioned in the 19th century. Although it is generally believed that Filhol was the author who established this taxon, I am herein demonstrating that authorship should in fact be attributed to Zittel, a fact that also influences not only its generic nomenclature, but also its appropriate type material. As such, the valid name for this taxon should be Palaeovaranus cayluxi and its holotype is a left maxilla. Additionally, Ophisauriscus eucarinatus from the middle Eocene of Geiseltal, Germany, another taxon that was previously assigned to Necrosaurus, is herein shown to be a nomen dubium, whereas Melanosauroides giganteus from the same locality, is considered a valid species and is recombined as Palaeovaranus giganteus comb. nov. The suggested changes in nomenclature also affect “Necrosauridae”, a poorly defined clade of lizards from the Cretaceous–Paleogene of Europe, North America, and Asia. In order to maintain nomenclatural stability and define a monophyletic lineage, I am here establishing the new family Palaeovaranidae fam. nov., which includes solely the genus Palaeovaranus. The known occurrences of Palaeovaranus across the Paleogene of Western Europe are discussed

A review of the fossil record of old world turtles of the clade pan-trionychidae

Turtles of the clade Pan-Trionychidae have a rich fossil record in the Old World, ranging from the Early Cretaceous (Hauterivian) to the Holocene. The clade most probably originated in Asia during the Early Cretaceous but spread from there to the Americas and Europe by the Late Cretaceous, to India and Australia by the Eocene, and to Afro-Arabia by the Neogene. The presence of a single pan-cyclanorbine in the Late Cretaceous (Maastrichtian) of Asia provides a minimum estimate for the age of the trionychid crown. As preserved, diversity was relatively high in Asia during the Late Cretaceous, but the subsequent, strong decline is likely a preservational bias, as extant faunas are relatively rich, especially throughout Asia. The range of trionychids contracted southward in Europe over the course of the Neogene, and the group is now locally extirpated. The group is now similarly absent from Arabia and Australia. A taxonomic review of the 180 named Old World taxa finds 42 nomina valida, 38 nomina invalida, 88 nomina dubia, 11 nomina nuda, and 1 nomen suppressum

Fossil lizards and snakes from Ano Metochi – a diverse squamate fauna from the latest Miocene of northern Greece

We here describe a new squamate fauna from the late Miocene (Messinian, MN 13) of Ano Metochi, northern Greece. The lizard fauna of Ano Metochi is here shown to be rather diverse, consisting of lacertids, anguids, and potential cordylids, while snakes are also abundant, consisting of scolecophidians, natricines and at least two different colubrines. If our identification is correct, the Ano Metochi cordylids are the first ones identified from Greece and they are also the youngest representatives of this group in Europe. A previously described scincoid from the adjacent locality of Maramena is here tentatively also referred to cordylids, strengthening a long term survival of this group until at least the latest Miocene. The scolecophidian from Ano Metochi cannot be attributed with certainty to either typhlopids or leptotyphlopids, which still inhabit the Mediterranean region. The find nevertheless adds further to the poor fossil record of these snakes. Comparison of the Ano Metochi herpetofauna with that of the adjacent locality of Maramena reveals similarities, but also striking differences among their squamate compositions

Late Paleogene herpetofaunas from the crossroads between two continents – new amphibian and reptile remains from the Oligocene of southern Balkans and Anatolia

We describe here new amphibian and reptile remains from three Oligocene localities of Turkey. Two of the localities (Kavakdere and Kocayarma) are situated in southeastern Europe and the other one (Kargi 2) in Anatolia, both areas where Oligocene herpetofauna is practically almost unknown. The material consists of albanerpetontids, pelobatid anurans, turtles, crocodylians, lacertids, scinciformatans, anguines, and “tropidophiids”. Albanerpetontids are for the first time identified in southeastern Europe, with the material being reminiscent of the younger species Albanerpeton inexpectatum Estes & Hoffstetter, 1976; the material potentially represents the oldest record of that species. Pelobatids, scinciformatans, and “tropidophiids” represent the oldest occurrences of these clades in the northeastern Mediterranean. The anguine genus Ophisaurus Daudin, 1803 is identified for the first time in the Paleogene of Eastern Europe. The “tropidophiids” are referred to two genera, Falseryx Szyndlar & Rage, 2003 and tentatively also to Platyspondylia Rage, 1974, with the latter having been so far exclusively known from western and central Europe. The role of a potential southern dispersal route of taxa among Asia and Europe, involving the area of southern Balkans and Anatolia, similarly to what has been recently demonstrated for mammals, is highlighted also for amphibians and reptiles

An illustrated atlas of the vertebral morphology of extant non-caenophidian snakes, with special emphasis on the cloacal and caudal portions of the column

Abstract We here present a thorough documentation of the vertebral morphology and intracolumnar variation across non-caenophidian snakes. Our studied sample of multiple individuals covers a large number of genera (67) and species (120), pertaining to almost all extant non-caenophidian families. Detailed figuring of multiple vertebrae across the trunk, cloacal, and caudal series for many different individuals / taxa documents the intracolumnar, intraspecific, and interspecific variation. An emphasis is given in the trunk-to-caudal transition and the pattern of the subcentral structures in that region of the column. Extant non-caenophidian snakes show an astonishing diversity of vertebral morphologies. Diagnostic vertebral features for extant families and many genera are given, though admittedly vertebral distinction among genera in certain groups remains a difficult task. A massive compilation of vertebral counts for 270 species, pertaining to 78 different genera (i.e., almost all known valid genera) and encompassing all extant non-caenophidian families, is provided based on our observations as well as an extensive literature overview. More particularly, for many taxa, detailed vertebral counts are explicitly given for the trunk, cloacal, and caudal portions of the column. Extant non-caenophidian snakes witness an extremely wide range of counts of vertebrae, ranging from 115 up to 546. A discussion on the diagnostic taxonomic utility and potential phylogenetic value of certain vertebral structures is provided. Comparisons of the subcentral structures of the cloacal and caudal vertebral series are also made with caenophidian lineages. We anticipate that this illustrative guide will set the stage for more vertebral descriptions in herpetological works but will also be of significant aid for taxonomic identifications in ophidian palaeontology and archaeozoology

Osteology, fossil record and palaeodiversity of the European lizards

he capability of palaeontologists to identify fossil remains of a particular group of vertebrates strongly depends on the knowledge they have of its comparative osteology and on the actual presence of diagnostic differences among the considered taxa. This could have a relevant influence on the study of palaeodiversity, since a low recognisability causes a loss of data when trying to reconstruct the history of taxa that lived on Earth in the past. Currently, more than 6000 extant species of lizards and worm lizards are known, and new ones continue to be discovered, mainly based on molecular data. But are we able to recognise this high diversity using osteology? As far as European taxa are concerned, the osteological recognisability of non-snake squamates is very low: only 31% of the extant European taxa can be identified based on their skeletal morphology. This is balanced partially by the fact that most recognisable taxa have been actually recognised in the fossil record, suggesting that the lost data are mainly due to the scarce knowledge of the comparative osteology of these reptiles and less influenced by other biases, such as taphonomic or collection biases. In this context, specimen-level phylogenetic analysis has proved to be a useful tool to identify diagnostic combinations of osteological features, at least for lacertid species, as evidenced by a case study focused on the genus Lacerta

A revision of the trionychid turtle Procyclanorbis sardus Portis, 1901 from the late Miocene of Sardinia (Italy)

Procyclanorbis sardus Portis, 1901 est la première tortue trionychidé fossile décrite en Sardaigne. Ce taxon du Miocène supérieur a été initialement considéré comme ayant des affinités avec les cyclanorbines d’Afrique et d’Asie du Sud. Nous décrivons ici en détail et figurons l’holotype de cette espèce, qui a subi une importante dégradation depuis sa publication originale. Une comparaison entre l’état originel du fossile et son état de conservation actuel est fournie. Sur la base de son anatomie, les affinités de Procyclanorbis sardus avec les cyclanorbinés sont rejetées car ce taxon s’avère être un pantrionychiné indéterminé. La distribution des trionychidés fossiles dans les îles de la Méditerranée est également discutée.Procyclanorbis sardus Portis, 1901 is the first fossil trionychid turtle described from Sardinia. This late Miocene taxon was originally considered to have affinities with the African and southern Asian cyclanorbines. We here redescribe in detail the holotype specimen of this species, which has suffered severe degradation since its original publication. A comparison between the original state of the fossil and its current state of preservation is provided. On the basis of its anatomy, affinities of Procyclanorbis sardus with cyclanorbines are discarded and this taxon is demonstrated to be an indeterminate pan-trionychine. The distribution of fossil trionychids in the Mediterranean Islands is also discussed

First fossil find of the Blanus strauchi complex (Amphisbaenia, Blanidae) from the Miocene of Anatolia

WOS: 000432213400008SYNTHESYS [SYNTHESYS ES-TAF-5910]; Ege University Research ProjectsEge University [TTM/001/2010, TTM/002/2011, TTM/001/2013, TTM/002/2014, 2015/Fen/17]; [TUBITAK-RFBR 111Y192]We would like to thank P. Lymberakis (NHMC), M. Calvo Revuelta (MNCN), and G. Gassner and C. Horweg (NHMW) for providing access to specimens under their care and photographs of extant specimens of Blanus spp. M. Delfino (University of Torino) provided insightful comments that enhanced the quality of the manuscript. A. Villa (University of Torino) helped with the literature. G.L.G. acknowledges travel support from SYNTHESYS (grant SYNTHESYS ES-TAF-5910) and the University of Torino. K.H., S.M., and T.K. were supported by TUBITAK-RFBR 111Y192 and Ege University Research Projects TTM/001/2010, TTM/002/2011, TTM/001/2013, TTM/002/2014, and 2015/Fen/17 during their field trips. We also thank Editor J. Head and the reviewers A. Bolet and J. Muller for their comments and help during the review process