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.     

Miocene skinks and geckos reveal long-term conservatism of New Zealand's lizard fauna

The New Zealand (NZ) lizard fossil record is currently limited to late Quaternary remains of modern taxa. The St Bathans Fauna (early Miocene, southern South Island) extends this record to 19–16 million years ago (Myr ago). Skull and postcranial elements are similar to extant Oligosoma (Lygosominae) skinks and Hoplodactylus (Diplodactylinae) geckos. There is no evidence of other squamate groups. These fossils, along with coeval sphenodontines, demonstrate a long conservative history for the NZ lepidosaurian fauna, provide new molecular clock calibrations and contradict inferences of a very recent (less than 8 Myr ago) arrival of skinks in NZ.     

At the feet of the dinosaurs: the early history and radiation of lizards

Lizards, snakes and amphisbaenians together constitute the Squamata, the largest and most diverse group of living reptiles. Despite their current success, the early squamate fossil record is extremely patchy. The last major survey of squamate palaeontology and evolution was published 20 years ago. Since then, there have been major changes in systematic theory and methodology, as well as a steady trickle of new fossil finds. This review examines our current understanding of the first 150 million years of squamate evolution in the light of the new data and changing ideas. Contrary to previous reports, no squamate fossils are currently documented before the Jurassic. Nonetheless, indirect evidence predicts that squamates had evolved by at least the middle Triassic, and had diversified into existing major lineages before the end of this period. There is thus a major gap in the squamate record at a time when key morphological features were evolving. With the exception of fragmentary remains from Africa and India, Jurassic squamates are known only from localities in northern continents (Laurasia). The situation improves in the Early Cretaceous, but the southern (Gondwanan) record remains extremely poor. This constrains palaeobiogeographic discussion and makes it difficult to predict centres of origin for major squamate clades on the basis of fossil evidence alone. Preliminary mapping of morphological characters onto a consensus tree demonstrates stages in the sequence of acquisition for some characters of the skull and postcranial skeleton, but many crucial stages--most notably those relating to the acquisition of squamate skull kinesis--remain unclear.     

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.     

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

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 tiny lizard (Lepidosauria,Squamata) from the Lower Cretaceous of Spain

Abstract: The smallest living amniotes are all lizards, but the fossil history of this size trait in Squamata is difficult to follow because small skeletons have low preservation potential and are often hard to detect in the field. A new squamate taxon, Jucaraseps grandipes gen. et sp. nov., is here described on the basis of an articulated skeleton from the Early Cretaceous Spanish lagerstätten of Las Hoyas. It differs from other known Mesozoic lizards in combining very small body size with a short rostrum, low maxillary tooth count, a relatively slender and elongated body, and short limbs with large hind feet. Phylogenetic analysis using TNT places it on the stem of a clade encompassing scincomorphs, gekkotans, snakes, amphisbaenians and anguimorphs. Comparison with modern lizards suggests it was probably a cryptic surface or subsurface ground dweller but not a burrower.     

AN EARLY HERBIVOROUS LIZARD FROM THE LOWER CRETACEOUS OF JAPAN

Abstract:  The Lower Cretaceous Tetori Group of Japan has yielded diverse freshwater and terrestrial vertebrate assemblages. The most productive small vertebrate locality is the 'Kaseki-Kabe' or 'fossil-bluff' at Kuwajima, Hakusan City, Ishikawa Prefecture. These deposits have produced at least six distinct lizard taxa of which one, described and named here as Kuwajimalla kagaensis , has lanceolate denticulate teeth convergent on those of the living Iguana . This type of dentition is rare among living lizards and is usually considered indicative of herbivory and, more specifically, folivory. Kuwajimalla kagaensis provides the earliest unambiguous record of squamate herbivory. Comparisons with modern and fossil lizards suggest that Kuwajimalla may be an early relative of the macrocephalosaurines, a group of large herbivores well represented in the Upper Cretaceous of Mongolia.     

Stretching the time span of hominin evolution at Kromdraai (Gauteng,South Africa): Recent discoveries

The Plio-Pleistocene locality of Kromdraai B has yielded the type specimen of Paranthropus robustus, as well as 27 additional fossil hominin specimens. In a number of both cranial and dental features, the states shown by the Kromdraai Paranthropus are more conservative when compared to the more derived conditions displayed by both South African conspecifics and the post-2.3 Ma eastern African Paranthropus boisei. Since 2014, we excavated the earliest known infilling of the Kromdraai cave system in a previously unexplored area. This new locality provided as yet 2200 identifiable macrovertebrate fossils, including 22 hominins, all tied in the earliest part of the stratigraphic sequence, representing three distinct depositional periods. Since we report here, for the first time, the occurrence of fossil hominins in Members 1 and 2, our discoveries stretch the time span of hominin evolution at Kromdraai and contribute to a better understanding of the origin of Paranthropus in southern Africa.     

A LONG-BODIED LIZARD FROM THE LOWER CRETACEOUS OF JAPAN

Abstract:  Platynotan lizards underwent a dramatic Late Cretaceous radiation into marine habitats. Beginning with small-bodied forms, the lineage culminated with the mosasaurs, large predatory lizards with a world-wide distribution in the Santonian–Campanian. Moreover, the marine squamate radiations of the Cenomanian–Turonian are remarkable in having produced a range of long-bodied, reduced-limbed swimmers (dolichosaurs, adriosaurs, coniasaurs and limbed snakes) that seem to have thrived in the shallow coastal environments of the Western Tethys region. Until now, none of these long-bodied aquatic squamates has been recorded prior to the Cenomanian, none has been recovered from a non-marine locality and none is known from Asia. Here we describe a small, gracile, long-bodied mosasauroid lizard from a swampy continental deposit in the Lower Cretaceous of Japan.     

Flore du Jurassique basal de la ville de Mende (Lozère) : synthèse des gisements historiques,nouvelles données sédimentologiques,paléontologiques et paléoenvironnementales

During the 20th century, because of urbanization, most palaeontological sites yielding plants (Hettangian–Sinemurian in age) from the city of Mende have disappeared, or they have been forgotten. This article presents a synthesis of sites yielding plants, from their discoveries in the 19th century to recent prospecting. New lithostratigraphic and palaeobotanical data are brought for two historic sites (Ravin del Pouset and Petit Séminaire), as well as two new localities (Causse d’Auge road and Gardès road). This study reveals the diversity of plant-containing lithofacies (dolomudstone, marl, limestone/calcarenite and oo-bioclastic limestone) and the diversity of plant preservations (charcoalified compressions with or without cuticle, isolated cuticle, external casts/impressions). Microfaciological study of plant beds demonstrates the diversity of depositional environments. Protected, restricted and shallow environments with low wave exposure and probable euryhaline conditions are clearly the most favourable to the accumulation and the preservation of fossil plants. These early Liassic palaeofloras, among the southernmost yet reported for Laurasia, are of significance for the plant geography and global paleoecology at that time.     

A new pachyostotic squamate reptile from the Cenomanian of France

Cenomanian beds from western France have yielded a new pythonomorph squamate (Reptilia) that is described here as Carentonosaurus mineaui gen. et sp. nov. This fossil is referred to non-mosasaurid mosasauroid lizards, i.e. to 'aigialosaurs'. It has been found in sediments of marine origin and its anatomy confirms that it was an aquatic lizard. It is characterized by a combination of characters that has not been reported, thus far, for squamates. Moreover, its vertebral column includes non-pachyostotic cervical vertebrae and highly pachyostotic dorsal vertebrae as in several other 'aigialosaurs'. This new taxon is perhaps restricted to the upper Cenomanian. It lived in shallow and rather warm water of the inner shelf. It is worth mentioning that nearly all pachyostotic squamates are concentrated in the Cenomanian and/or lower Turonian deposits of the present European-North African-Middle East portion of the Tethys. A parallel is drawn between the high percentage of pachyostotic squamates and the fact that this period corresponds to both the largest transgression of the Phanerozoic and the warmest period in the whole of the Mesozoic and Caenozoic.     

Dental remains of Lower Miocene small rodents from Napak (Uganda): Afrocricetodontidae,Myophiomyidae, Kenyamyidae and Sciuridae

The Early Miocene sites of Napak (Uganda), which have been prospected by the Uganda Palaeontology Expedition since 1985, have yielded abundant fossil remains, including a rich and diverse rodent fauna. Ever since the work of Lavocat (1973) on the East African Miocene rodents, which focused mainly on the Kenyan remains (but included some material from Napak discovered by Bishop during the 1950s and 1960s), few studies have been made on the Ugandan specimens. This lack provides motivation for further study of Napak micro-mammals, especially the abundant rodents, which were collected at five sites, Napak IV, V, XV, XXX and XXXI. The fossils described here are attributed to six species representing four families: Afrocricetodontidae (Afrocricetodon songhori, Protarsomys macinnesi), Myophiomyidae (Myophiomys arambourgi), Kenyamyidae (Simonimys genovefae, Kenyamys mariae) and Sciuridae (Vulcanisciurus africanus). Comparison of the dentition of extant and extinct taxa allows us to propose dietary hypotheses (granivory, herbivory and omnivory) for these rodents. In addition, the fossil rodents associated with other fauna from the same localities suggest a more-or-less dense forest environment with clearings, and the probable presence of a humid climate at the time of deposition.     

Observations on the Turin specimen ofMesopithecus pentelici (Wagner, 1839)

A fossil specimen ofMesopithecus pentelici Wagner, 1839, from the Pikermi paleontological assemblage and currently housed at the Geological Museum of the Earth Sciences Department of the University of Turin, is described herein. The left mandibular ramus displays dental features which permit us to assign it to a young female individual. Anatomical patterns are discussed and are compared with other similar remains of more recent discovery.     

记内蒙古宁城道虎沟上侏罗统/下白垩统一具有长肢的蜥蜴

道虎沟化石层分布于内蒙古宁城县道虎沟以及毗邻的河北北部和辽宁西部等地,已报道4种有尾类、1个无尾类蝌蚪、1个幼年蜥蜴、3种带原始羽毛的兽脚类恐龙、2种翼龙、3种哺乳动物以及双壳类、叶肢介、昆虫和植物等化石(有学者称之为"前热河生物群")。该层位比热河群义县组低,但地层划分存在争议:中侏罗统九龙山组、上侏罗统道虎沟组和下白垩统热河群均有报道。本文研究的道虎沟蜥蜴是该地点发现的第2个蜥蜴化石,其身体细长,为一幼年个体;前颌骨、顶骨和额骨均成对;上颌骨的面突高;后额骨和后眶骨不愈合;后额骨较大,但可能未进入上颞孔;后眶骨具宽的后突;牙齿小而尖锐,结构简单且可能为侧生;头骨的眶前区较长;具27个荐前椎;虽然未成年但个体较大;未见真皮骨板;手掌和四肢较长。上述特征组合区别于所有现生蜥蜴类,四肢比例也与其他蜥蜴有较大区别。欧美中侏罗世-早白垩世的小盗蜥(Parviraptor)和蒙古早白垩世的一个幼年个体(可能属于壁虎型类)与道虎沟标本比较相似。头骨特征和较大的荐前椎数目显示道虎沟标本可能属于硬舌蜥类(scleroglossan)。但因标本为幼年个体,本文暂将它归入有鳞类属种未定(Squamata gen.et sp.indet.)。测量数据显示,道虎沟标本的手掌和前肢相对较长;该特征常见于适应攀爬的蜥蜴种类,似可显示攀爬习性。但肢体比例可能会随身体增长而发生变化,所以此生活习性并非定论。     

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

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.     

Musteloids and feloids (Mammalia, Carnivora) from the late Miocene locality of Pikermi (Attica, Greece)

Some relatively rare carnivore remains from Pikermi are studied. The present specimens include Simocyon primigenius(ROTH and WAGNER, 1854), Promeles palaeatticus(WEITHOFER, 1888), Martes woodwardiPILGRIM, 1931, Felis atticaWAGNER, 1857 and Machairodus giganteus(WAGNER, 1848), as well as some postcranial remains that resemble Enhydriodon? latipesPILGRIM, 1931 and are assigned to Mustelidae gen. et spec. indet. Some of these species were not known until now but from their holotypes as M. woodwardi and E.? latipes. Other species such as F. attica and M. giganteus are better known but from relatively scant material. Since the material of the earlier mentioned species comes from Pikermi, their type locality, it is of especial importance and improves our knowledge for the Pikermi carnivores and their relationships to other species. In some cases where the material comes from ossiferous blocks, the accompanying faunal context is discussed.     

Squamate phylogeny, taxon sampling, and data congruence

To investigate the affinities of snakes, amphisbaenians and dibamids, the phylogenetic relationships among the major lineages (families) of extinct and extant squamates are assessed through a combined analysis of 248 osteological, 133 soft anatomical, and 18 ecological traits. The osteological data set represents a revision of previous data, taking into account recent criticism; the ecological data set is new. In addition, potentially critical fossil taxa (polyglyphanodontids and macrocephalosaurs) are included for the first time. The osteological and soft anatomical data sets each place snakes within anguimorphs, with dibamids and amphisbaenians near gekkotans. The putative primitive fossil amphisbaenian Sineoamphisbaena groups with macrocephalosaurs and polyglyphanodontids, together the sister group to scleroglossans. All three data sets are congruent, and these results are reinforced by combined analyses. In these, as in the osteological analyses, snakes are nested within marine lizards. However, exclusion of fossil taxa from the osteological data set results in a ‘limbless clade’ consisting of snakes, amphisbaenians and dibamids, and introduces significant conflict between osteology and soft anatomy. Also, deletion tests and character weighting reveal that the signal in the reduced osteological data set is internally contradictory. These results increase confidence in the arrangement supported by the all-taxon osteological, the soft anatomical, and the combined data, and suggest that exclusion of fossils confounds the signal in the osteological data set. Finally, the morphological data support the nesting of snakes within marine lizards, and thus a marine origin of snakes. This result still holds when relationships between living forms are constrained to the topology suggested by molecular sequences: if marine lizards are allowed to ‘float’ within this molecular framework, they form the stem group to snakes, and do not group with varanids as previously suggested.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-04.htm.     

Middle to Late Pleistocene herpetofauna from Scladina and Sous-Saint-Paul caves (Namur,Belgium)

For the first time, the fossil herpetofauna from the Middle and Late Pleistocene of Scladina and Sous-Saint-Paul caves (Sclayn, Belgium) is described. The amphibians and squamate reptiles are represented by one salamander (Salamandra salamandra), three anurans (Pelodytes punctatus, Bufo bufo and Rana temporaria), two lizards (Lacerta cf. agilis and Anguis fragilis) and two snakes (Zamenis longissimus and Vipera cf. berus). The occurrence of the Parsley Frog (Pelodytes punctatus) and the Aesculapian Snake (Zamenis longissimus), which are not currently represented in Belgium, is of particular interest. Scladina also represents one of the northernmost fossil mentions for the Fire Salamander (Salamandra salamandra) although it is within its current distribution in Europe. Finally, the presence of the Adder (Vipera cf. berus) is very probably attested in Scladina whereas today this snake is infrequent and classified as endangered in Belgium.     

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.     

The uppermost Oligocene of Aix-en-Provence (Bouches-du-Rhône,Southern France): A Cenozoic brackish subtropical Konservat-Lagerstätte,with fishes,insects and plants

Aix-en-Provence is one of the most famous localities for Cenozoic fossils in Europe, known since the end of the 18th century for the exquisitely preserved fishes, insects and plant remains extracted from laminated deposits exposed in the underground quarries developed for the exploitation of the gypsum, and in some sites located inside and around the city. The Aix-en-Provence fossil assemblage includes a considerable variety of organisms that provide a well-documented evidence of the existence of a large subtropical brackish (primarily oligohaline) lagoon temporarily connected to the sea in southern France during the latest Oligocene.