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

New material of Laophis crotaloides, an enigmatic giant snake from Greece, with an overview of the largest fossil European vipers

Laophis crotaloides was described by Richard Owen as a new and very large fossil viperid snake species from Greece. The type material is apparently lost and the taxon was mostly neglected for more than a century. We here describe a new partial viperid vertebra, collected from the same locality and of equivalent size to the type material. This vertebra indicates that at least one of the three morphological characters that could be used to diagnose L. crotaloides is probably an artifact of the lithographer who prepared the illustration supporting the original description. A revised diagnosis of L. crotaloides is provided on the basis of the new specimen. Despite the fragmentary nature of the new vertebra, it confirms the validity of L. crotaloides, although its exact relationships within Viperidae remain unknown. The new find supports the presence of a large viperid snake in the early Pliocene of northern Greece, adding further data to the diversity of giant vipers from Europe

AN OVERVIEW ON THE LATE MIOCENE VERTEBRATES FROM THE FISSURE FILLINGS OF MONTICINO QUARRY (BRISIGHELLA, ITALY), WITH NEW DATA ON NON-MAMMALIAN TAXA

The latest Messinian Monticino Quarry fissure fillings, near Brisighella in Italy, are well known for their diverse mammal fauna. Conversely, little is known about other vertebrates from this rich site. Beside presenting an overview of the mammals, here we describe fish, amphibian, reptile, and bird remains, identifying three, eight, at least 18, and five taxa, respectively. Some of these represent the oldest occurrences either worldwide (an erycine snake related to Eryx jaculus, Malpolon, and otidids birds) or locally (common toads in the Apennine Peninsula and eremiadine lacertids in Europe, the latter being also the only known occurrence of these lizards in Italy). Altogether, the vertebrate fauna from Monticino Quarry is indicative of a complex palaeoenvironment, which was warm and somehow dry. Water bodies were indeed present though, as well as a patchy set of habitats with loose or sandy soils, rocky outcrops, open biotopes, and (even though maybe to a lesser extent) wooded areas. Furthermore, estuarine or lagoonal conditions were likely present near the depositional site. From a palaeobiogeographical point of view, the non-mammal vertebrate assemblage from Monticino Quarry shows considerable Afro-Asian affinities at least for some components. In the context of the Italian late Miocene continental vertebrate assemblages, that of Monticino is remarkably similar to those from the Piedmont Basin, especially if compared with the Tusco-Sardinian area, Apulian Platform, Calabrian Arc and Sicily. This is consistent with the palaeobiogeographic scenario defined by mammal assemblages, with northcentral Italy, Tuscany and Sardinia, Apulian Platform, and Calabria and Sicily representing separated bioprovinces

The impact of major warming at 14.7 ka on environmental changes and activity of Final Palaeolithic hunters at a local scale (Orawa-Nowy Targ Basin, Western Carpathians, Poland

There is a widespread belief that the abrupt warming at 14.7 ka had a profound impact on the environment. However, the direct correlation between the global climatic event and changes in local environments is not obvious.We examined faunal succession in an intra-mountain basin of theWestern Carpathians to assess the potential influence of the climatic change between Greenland Stadial-2a and Greenland Interstadial-1e on the local environment.We investigated three vertebrate assemblages (total number of identified specimens = 18,745; minimumnumber of individuals = 7515; 138 taxa) from Obłazowa Cave (western entrance) and a Rock overhang in Cisowa Rock, radiocarbon dated to the period before and after the global warming, between ca. 17.0 and 14.0 ka. Our data revealed that the major abrupt warming that occurred 14.7 ka had little impact on the local environment, which could suggest that ecosystems in Central Europe were resilient to the abrupt global climate changes. The increase in fauna population sizes and species diversities in local biotopes was gradual and began long before the temperature increase. This was supported by the analysis of ancient DNA of Microtus arvalis, which showed a gradual increase in effective population size after 19.0 ka. The results of palaeoclimatic reconstruction pointed out that the compared sites were characterized by similar climatic conditions. According to our calculations, the differences in the annual mean temperatures did not exceed 0.5 °C and mean annual thermal amplitude changed from 22.9 to 22.4 °C. The environmental changes before 14.7 ka had no impact on the activity of Final Palaeolithic hunters in the studied area

Upper Pleistocene herpetofauna from Cueva de Cobrante

8 páginas, 12 láminas, 1 tabla.[EN]: Herpetological samples from the late Pleistocene wurmian site Cueva de Cobrante (San Miguel de Aras, Cantabria, Spain) are studied. The herpetological remains are associated with different Upper Palaeolithic levels. The predominant species in all excavated levels is Rana temporaria, accompanied by scant remains of Alytes obstetricans and Bufo bufo. Sporadic isolated remains of Lacerta sp., new-born snakes and perhaps Bufo calamita have also been detected. Within the osteological anomalies, two remarkable Rana temporaria opisthocoelous vertebrae have been found. Probably most of the herpetological material comes from raptor pellets different from Tyto alba, plus a minor component derived from carnivore mammal predation. Based on its availability as prey, the sedimentation of these remains seems to have occurred between April and October, probably at a higher rate during the early summer period.[ES]: Se estudian los restos herpetológicos wurmienses del yacimiento del Pleistoceno tardío de la cueva de Cobrante (San Miguel de Aras, Cantabria, España), asociados a niveles correspondientes a diversas fases del Paleolítico Superior. La especie predominante en todos los niveles excavados es Rana temporaria, detectándose también escasos restos de Alytes obstetricans y Bufo bufo. De manera esporádica han aparecido restos aislados de Lacerta sp., serpientes neonatas, y un resto dudoso de Bufo calamita. Entre las anomalías óseas detectadas destacan dos vértebras opistocélicas de Rana temporaria. El material parece proceder de egagrópilas de rapaces distintas a Tyto alba, además de un componente menor de alimentación de mamíferos carnívoros. Dada su disponibilidad como presas, la sedimentación de los restos parece haberse dado entre abril y octubre, probablemente en mayor medida a principios de verano.Esta investigación se ha realizado mediante el proyecto CGL 2008-03881, Ministerio de Ciencia e Innovación.Peer reviewe

First occurrence of <i>Psammophis</i> (Serpentes) from Europe witnesses another Messinian herpetofaunal dispersal from Africa – biogeographic implications and a discussion of the vertebral morphology of psammophiid snakes

We here describe abundant new snake material from the late Miocene (MN 13) of Salobreña, Spain. Vertebral morphology suggests a referral of the specimens to the extant psammophiid Psammophis, documenting the first occurrence of this genus in Europe. The diversity and disparity across the vertebral morphology of different psammophiid genera are discussed. We identify vertebral features that could diagnose Psammophis and therefore enable the recognition of the genus in the fossil record. A comparison of the new Spanish form with other taxa is conducted. We provide a detailed review of the psammophiid fossil record. Material previously described from the middle Miocene of Beni Mellal, Morocco is here tentatively referred to as ?Psammophis sp., an action that renders that occurrence as the oldest (probable) record of the genus and Psammophiidae as a whole, providing thus a potential calibration point. On the other hand, Eastern European Pliocene material that had been previously supposedly referred to Psammophis is here discarded as being rather fragmentary, not affording any more precise determination. The two psammophiid genera Psammophis and Malpolon appear almost simultaneously in the European fossil record (MN 13), with the former achieving only a short-lived and apparently geographically limited distribution in the continent, while the latter still exists in its modern herpetofauna. We assess biogeographic implications of the new find, suggesting a direct dispersal event from northwestern Africa to the Iberian Peninsula during the late Miocene, facilitated by the Messinian Salinity Cri

Oligocène terminal–Miocène inférieur en Europe : l&#039;énigme des serpents booïdes

En Europe, la diversité des serpents booïdes a fortement chuté pendant l\u27Oligocène terminal (zones MP 29-30) et le début du Miocène (MN 1-2, et peut-être MN 3). Les quelques booïdes qui subsistaient étaient petits et de tendances fouisseuses. Cette crise a été qualifiée de Dark Period des booïdes. Elle a probablement été déclenchée par l\u27aridification et le refroidissement qui se sont renforcés vers la fin de l\u27Oligocène. Cette période de crise s\u27est terminée avec le retour de l\u27humidité et de la chaleur ; en plus d\u27un effet direct, ces changements ont aussi entraîné l\u27arrivée d\u27immigrants, qui ont certainement contribué à l\u27élimination des booïdes de la Dark Period.The diversity of booid snakes strongly decreased during the Latest Oligocene (zones MP 29-30) and Early Miocene (zones MN 1-2, and perhaps MN 3) in Europe. Moreover, the few remaining taxa were all small and displayed fossorial trends. This crisis is referred to as the Dark Period of booid snakes. The outset of this event probably resulted from cooling and aridization that were prevailing during the Late Oligocene. The period ended when a warm and humid climate was restored; aside from direct effect, the latter change caused the arrival of immigrants that probably contributed to the elimination of the booids of the Dark Period.</p

Amphibian and reptilian fauna from the early Miocene of Echzell, Germany

The present study describes a rich amphibian and reptilian assemblage from the early Miocene locality Echzell, Germany. It consists of one allocaudate, five salamander, five frog, one gecko, chamaeleonids, anguine lizards, one lacertid, one skink and five snake taxa. The entire herpetofauna of Echzell is represented by genera and/or families very broadly known from the early Miocene of Europe. Contrary to other early Miocene herpetofaunas, the Echzell assemblage includes surprisingly only one form of crocodile-newts (Chelotriton). The Echzell Palaeobatrachus robustus represents the youngest record of the species and extends its stratigraphic range to the late early Miocene. Regarding chameleons, the frontal is partly preserved, but represents the first described frontal of the extinct species Chamaeleo andrusovi. The only anguine lizard that can be identified in the assemblage is represented by a new genus and species Smithosaurus echzellensis. Our phylogenetic analyses consistently recovered it as the sister taxon to either [Ophisauriscus quadrupes + Ophisaurus holeci] + [Anguis + Ophisaurus] (in the first analysis) or [Anguis + Ophisaurus] (in the second analysis). However, the results are based on limited fossil material – the parietal – and the support for the clade is very low. Thus, the interpretation of the Smithosaurus relationship among anguines needs to be taken with caution and has to be tested in further studies. Among snakes, Natrix longivertebrata represents the oldest record of the species and extends the stratigraphic range of this fossil snake back to the early Miocene. In addition, we provide here a broader comparison of the Echzell amphibian and reptilian assemblage with their European records for the MN3 and MN4 biostratigraphical units. Besides that, the entire herpetofauna of Echzell includes very broadly known early Miocene European forms. Remains of other groups of the same period such as Bufonidae, Hylidae, Pelodytidae, Amphisbaenia, Varanidae, Cordylidae, Pseudopus, are not found in the material available to us. We also conclude that the amphibian and reptilian fossil record across MN3–MN4 is significantly biased by taphonomic and/or environmental conditions. The amphibian and reptilian assemblage of Echzell is rich in forms living in humid and warm environments with forested areas, permanent water bodies and also some open habitats. The following climatic parameters can be reconstructed based on the herpetofauna: a mean annual temperature of 17.4–28.8 °C, minimal warm month temperature 18–28.3 °C, minimal cold month temperature 8–22.2 °C, and mean annual precipitation with a value of 791±254 mm.A.Č. acknowledges financial support from the Scientific Grant Agency of the Ministry of Education of the Slovak Republic and Slovak Academy of Sciences, Grant Nr. 1/0191/21. T.M. acknowledges financial support from the Bolin Center for Climate Research, Stockholm University (RA6 grant).</p