Lizards and amphisbaenians (Reptilia, Squamata) from the middle Eocene of Mazaterón (Soria, Spain).

The assemblage of lizards and amphisbaenians (Reptilia, Squamata) from the middle Eocene locality of Mazaterón (Spain) is described. Considering the rather limited material available for the study, the assemblage shows a moderate diversity with eight taxa corresponding to five different families. In most cases the scarcity and fragmentary nature of squamate specimens precludes a precise identification, but provides insights on identity of the groups represented. Mazaterón fills the gap between early and late Eocene Iberian localities, showing the persistence of iguanids (possibly Geiseltaliellus), lacertids (possibly Dormaalisaurus), and glyptosaur (tribes glyptosaurini and "melanosaurini") and anguine anguids through most of the Iberian Eocene. It also records the return of amphisbaenians (Blanidae) after their temporary retrieval from Europe during most of the middle Eocene, and the presence of two scincids, one of them possibly corresponding to a new taxon. The information provided by squamates complements what is already known from mammals, crocodylians, and turtles in what is arguably one of the most important vertebrate Paleogene localities of the Iberian Peninsula.

The Jurassic rise of squamates as supported by lepidosaur disparity and evolutionary rates.

The squamates (lizards, snakes, and relatives) today comprise more than 10,000 species, and yet their sister group, the Rhynchocephalia, is represented by a single species today, the tuatara. The explosion in squamate diversity has been tracked back to the Cretaceous Terrestrial Revolution, 100 million years ago (Ma), the time when flowering plants began their takeover of terrestrial ecosystems, associated with diversification of coevolving insects and insect-eating predators such as lizards, birds, and mammals. Squamates arose much earlier, but their long pre-Cretaceous history of some 150 million years (Myr) is documented by sparse fossils. Here, we provide evidence for an initial radiation of squamate morphology in the Middle and Late Jurassic (174-145 Ma), and show that they established their key ecological roles much earlier than had been assumed, and they have not changed them much since.

A new Early Cretaceous lizard in Myanmar amber with exceptionally preserved integument.

We here report on a well-preserved juvenile lizard specimen in Albian amber (ca. 110 mya) from the Hkamti site (Myanmar). This new taxon is represented by an articulated skull and the anterior portion of the trunk, including the pectoral girdle and forelimbs. The scleral ossicles and eyelid are also visible, and the specimen exhibits pristine detail of the integument (of both head and body). In a combined molecular and morphological analysis, it was consistently recovered as a scincoid lizard (Scinciformata), as sister to Tepexisaurus + Xantusiidae. However, the phylogenetic position of the new taxon should be interpreted with caution as the holotype is an immature individual. We explored the possibility of miscoding ontogenetically variable characters by running alternative analyses in which these characters were scored as missing data for our taxon. With the exception of one tree, in which it was sister to Amphisbaenia, the specimen was recovered as a Pan-xantusiid. Moreover, we cannot rule out the possibility that it represents a separate lineage of uncertain phylogenetic position, as it is the case for many Jurassic and Cretaceous taxa. Nonetheless, this fossil offers a rare opportunity to glimpse the external appearance of one group of lizards during the Early Cretaceous.

Unusual morphology in the mid-Cretaceous lizard Oculudentavis.

Oculudentavis khaungraae was described based on a tiny skull trapped in amber. The slender tapering rostrum with retracted narial openings, large eyes, and short vaulted braincase led to its identification as the smallest avian dinosaur on record, comparable to the smallest living hummingbirds. Despite its bird-like appearance, Oculudentavis showed several features inconsistent with its original phylogenetic placement. Here, we describe a more complete specimen that demonstrates Oculudentavis is actually a bizarre lizard of uncertain position. The new specimen is described as a new species within the genus Oculudentavis. The new interpretation and phylogenetic placement highlight a rare case of convergent evolution in skull proportions but apparently not in morphological characters. Our results re-affirm the importance of Myanmar amber in yielding unusual taxa from a forest ecosystem rarely represented in the fossil record.

Enigmatic amphibians in mid-Cretaceous amber were chameleon-like ballistic feeders.

Albanerpetontids are tiny, enigmatic fossil amphibians with a distinctive suite of characteristics, including scales and specialized jaw and neck joints. Here we describe a new genus and species of albanerpetontid, represented by fully articulated and three-dimensional specimens preserved in amber. These specimens preserve skeletal and soft tissues, including an elongated median hyoid element, the tip of which remains embedded in a distal tongue pad. This arrangement is very similar to the long, rapidly projecting tongue of chameleons. Our results thus suggest that albanerpetontids were sit-and-wait ballistic tongue feeders, extending the record of this specialized feeding mode by around 100 million years.

First nearly complete skull of Gallotia auaritae (lower-middle Pleistocene, Squamata, Gallotiinae) and a morphological phylogenetic analysis of the genus Gallotia.

The Canary Islands are an Atlantic archipelago known for its high number of endemic species. Among the most known endemic vertebrate species are the giant lizards of the genus Gallotia. We describe the cranial osteology of the first almost complete and articulated fossil skull of the taxon Gallotia auaritae, recovered from the lower-middle Pleistocene of the La Palma island. In this work, X-ray computed microtomography images were used to perform an exhaustive phylogenetic analysis where most of the extant and fossil species of the genus Gallotia were included for first time. This analysis recovered a monophyletic Gallotia clade with similar topology to that of molecular analyses. The newly described specimen shares some characters with the group formed by G. bravoana, G. intermedia and G. simonyi, G. auaritae, and its position is compatible with a referral to the latter. Our study adds new important data to the poorly known cranial morphology of G. auaritae, and the phylogenetic analysis reveals an unexpected power of resolution to obtain a morphology-based phylogeny for the genus Gallotia, for inferring the phylogenetic position of extinct species and for helping in the identification of fossil specimens.

Revision of Varanus marathonensis (Squamata, Varanidae) based on historical and new material: morphology, systematics, and paleobiogeography of the European monitor lizards.

Monitor lizards (genus Varanus) inhabited Europe at least from the early Miocene to the Pleistocene. Their fossil record is limited to about 40 localities that have provided mostly isolated vertebrae. Due to the poor diagnostic value of these fossils, it was recently claimed that all the European species described prior to the 21st century are not taxonomically valid and a new species, Varanus amnhophilis, was erected on the basis of fragmentary material including cranial elements, from the late Miocene of Samos (Greece). We re-examined the type material of Varanus marathonensis Weithofer, 1888, based on material from the late Miocene of Pikermi (Greece), and concluded that it is a valid, diagnosable species. Previously unpublished Iberian material from the Aragonian (middle Miocene) of Abocador de Can Mata (Vallès-Penedès Basin, Barcelona) and the Vallesian (late Miocene) of Batallones (Madrid Basin) is clearly referable to the same species on a morphological basis, further enabling to provide an emended diagnosis for this species. Varanus amnhophilis appears to be a junior subjective synonym of V. marathonensis. On the basis of the most complete fossil Varanus skeleton ever described, it has been possible to further resolve the internal phylogeny of this genus by cladistically analyzing 80 taxa coded for 495 morphological and 5729 molecular characters. Varanus marathonensis was a large-sized species distributed at relatively low latitudes in both southwestern and southeastern Europe from at least MN7+8 to MN12. Our cladistic analysis nests V. marathonensis into an eastern clade of Varanus instead of the African clade comprising Varanus griseus, to which it had been related in the past. At least two different Varanus lineages were present in Europe during the Neogene, represented by Varanus mokrensis (early Miocene) and V. marathonensis (middle to late Miocene), respectively.

An archosauromorph dominated ichnoassemblage in fluvial settings from the late Early Triassic of the Catalan Pyrenees (NE Iberian Peninsula).

The vertebrate recovery after the end-Permian mass extinction can be approached through the ichnological record, which is much more abundant than body fossils. The late Olenekian (Early Triassic) tetrapod ichnoassemblage of the Catalan Pyrenean Basin is the most complete and diverse of this age from Western Tethys. This extensional basin, composed of several depocenters, was formed in the latest phases of the Variscan orogeny (Pangea breakup) and was infilled by braided and meandering fluvial systems of the red-beds Buntsandstein facies. Abundant and diverse tetrapod ichnites are recorded in these facies, including Prorotodactylus mesaxonichnus isp. nov. (tracks possibly produced by euparkeriids), cf. Rotodactylus, at least two large chirotheriid morphotypes (archosauriform trackmakers), Rhynchosauroides cf. schochardti, two other undetermined Rhynchosauroides forms, an undetermined Morphotype A (archosauromorph trackmakers) and two types of Characichnos isp. (swimming traces, here associated to archosauromorph trackmakers). The Pyrenean ichnoassemblage suggests a relatively homogeneous ichnofaunal composition through the late Early Triassic of Central Pangea, characterized by the presence of Prorotodactylus and Rotodactylus. Small archosauromorph tracks dominate and present a wide distribution through the different fluviatile facies of the Triassic Pyrenean Basin, with large archosaurian footprints being present in a lesser degree. Archosauromorphs radiated and diversified through the Triassic vertebrate recovery, which ultimately lead to the archosaur and dinosaur dominance of the Mesozoic.

New genus and species names for the Eocene lizard Cadurcogekko rugosus Augé, 2005.

Cadurcogekko rugosus Augé, 2005 was described as a gekkotan lizard from the Eocene of France. A revision of the material has revealed that the holotype, a nearly complete dentary, actually belongs to a scincid lizard, for which we erect the new genus Gekkomimus. The rest of material originally referred to C. rugosus is of undoubted gekkotan nature and is included in the new species Cadurcogekko verus, with the exception of a partial left dentary belonging to the iguanid lizard Cadurciguana hoffstetteri.

An amphisbaenian skull from the European miocene and the evolution of Mediterranean worm lizards.

The evolution of blanid amphisbaenians (Mediterranean worm lizards) is mainly inferred based on molecular studies, despite their fossils are common in Cenozoic European localities. This is because the fossil record exclusively consists in isolated elements of limited taxonomic value. We describe the only known fossil amphisbaenian skull from Europe - attributed to Blanus mendezi sp. nov. (Amphisbaenia, Blanidae) - which represents the most informative fossil blanid material ever described. This specimen, from the Middle Miocene of Abocador de Can Mata (11.6 Ma, MN7+8) in the Vallès-Penedès Basin (Catalonia, NE Iberian Peninsula), unambiguously asserts the presence of Blanus in the Miocene of Europe. This reinforces the referral to this genus of the previously-known, much more incomplete and poorly-diagnostic material from other localities of the European Neogene. Our analysis - integrating the available molecular, paleontological and biogeographic data - suggests that the new species postdates the divergence between the two main (Eastern and Western Mediterranean) extant clades of blanids, and probably precedes the split between the Iberian and North-Western African subclades. This supports previous paleobiogeographic scenarios for blanid evolution and provides a significant minimum divergence time for calibrating molecular analyses of blanid phylogeny.

A new miniaturized lizard from the late Eocene of France and Spain.

We report here a new lizard genus and species shared by two late Eocene localities situated at both versants of the present Pyrenees (South-Western Europe), one located in France (Escamps, MP19), and the other in Catalonia, Spain (Sossís, MP17a). The recovered specimens are remarkable because of their small size and peculiar morphology. Features of the dentary are interpreted as adaptations to a fossorial or semi-fossorial lifestyle, although such modifications obscure the exact phylogenetic relationships of the new taxon. We suggest that it might represent a further example of scincoid lizard that independently achieved adaptations for burrowing or surface-dwelling. This taxon reinforces the hypotheses that link the Southern Pyrenean assemblages to those from France rather than to those of the rest of the Iberian Peninsula, which are supposed to be somehow isolated and endemic to a certain degree during the middle and late Eocene, forming part of the so-called Western Iberian Bioprovince.