A first possible chameleon from the late Miocene of India (the hominoid site of Haritalyangar): a tentative evidence for an Asian dispersal of chameleons.

Miocene rare fossils from India, tentatively attributed to chameleons, are described for the first time. The material consists of a fragment of the left squamosal and an element interpreted as a posterodorsal process of the parietal. The specimens come from a late Miocene site of the Nagri Formation (Middle Siwaliks, ~ 9 Mya) at Haritalyangar, North India. This material presents a possible evidence for a chameleon dispersal to Asia. Based on molecular data, the dispersion of an Asian chamaeleonid lineage from Africa to Arabia/Asia is dated at approximately 13 Mya and its diversification in situ at around 6-8 Mya. However, till now, no Miocene-age fossil record has been described to support crown chamaeleonid presence in this area. The material described herein is very fragmented. If correctly allocated, the Haritalyangar chameleons show the oldest known occurrence of this clade in India, at least approximately 9 Mya ago.

A new gecko from the earliest Eocene of Dormaal, Belgium: a thermophilic element of the 'greenhouse world'.

We here describe a new gekkotan lizard from the earliest Eocene (MP 7) of the Dormaal locality in Belgium, from the time of the warmest global climates of the past 66 million years (Myr). This new taxon, with an age of 56 Myr, together with indeterminate gekkotan material reported from Silveirinha (Portugal, MP 7) represent the oldest Cenozoic gekkotans known from Europe. Today gekkotan lizards are distributed worldwide in mainly warm temperate to tropical areas and the new gecko from Dormaal represents a thermophilic faunal element. Given the Palaeocene-Eocene thermal maximum at that time, the distribution of this group in such northern latitudes (above 50° North - the latitude of southern England) is not surprising. Although this new gekkotan is represented only by a frontal (further, dentaries and a mandibular fragment are described here as Gekkota indet. 1 and 2-at least two gekkotan species occurred in Dormaal), it provides a new record for squamate diversity from the earliest Eocene 'greenhouse world'. Together with the Baltic amber gekkotan Yantarogekko balticus, they document the northern distribution of gekkotans in Europe during the Eocene. The increase in temperature during the early Eocene led to a rise in sea level, and many areas of Eurasia were submerged. Thus, the importance of this period is magnified by understanding future global climate change.

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.

Anatomy of the neural endocranium, parasphenoid and stapes of Diadectes absitus (Diadectomorpha) from the early Permian of Germany based on the high-resolution X-ray microcomputed tomography.

A detailed anatomy of the braincase and stapes of the subadult specimen of Diadectes absitus from early Permian sediments of Germany are described for the first time based on the high-resolution X-ray microcomputed tomography. In contrast to previous studies of Diadectes, the bones of the braincase (opisthotic, prootic, supraoccipital, basioccipital, exoccipital, basisphenoid, and sphenethmoid), and parasphenoid of D. absitus are not co-ossified, but suturally defined. This has allowed for a reconstruction of a complete braincase with all sutures between the individual bones. The opisthotic, prootic, and supraocciptal contain a well-preserved endosseous labyrinth. The three-dimensional-reconstruction of its cavities shows a well-preserved vestibule, three semicircular canals, and well-developed cochlear recess. In addition, a shallow subarcuate fossa is present on the ventral surface of the supraoccipital, which lies medial to the anterior semicircular canal. A typical feature of the diadectid braincase is the presence of the otic tube leading from the fenestra vestibuli to the vestibule. A revision of the topology of this structure is presented here. Here, we describe new structures of the stapes, especially in its proximal portion, as well as its position to the fenestra vestibuli. These structures are described for the first time not only in D. absitus, but for the genus.

The only complete articulated early Miocene chameleon skull (Rusinga Island, Kenya) suggests an African origin for Madagascar's endemic chameleons.

We here present the first detailed study of the specimen KNM-RU 18340 from Rusinga Island (Kenya), the only known complete early Miocene chameleon skull, using micro-CT. This specimen represents one of the oldest chameleon fossils ever recovered. For the first time, the skull bone internal surfaces, their sutures, and elements contained inside the rocky matrix are observed. Our morphological comparisons and phylogenetic analyses place this specimen confidently in the genus Calumma and a new species, Calumma benovskyi sp. nov., is erected for it. Since all species of this genus are endemic to Madagascar, this fossil uniquely demonstrates the existence of Calumma on continental Africa in the past. Our results challenge the long-held view that chameleons originated on Madagascar and dispersed over water to Africa, and provide a strong evidence of an African origin for some Malagasy lineages. The Oligocene-early Miocene dispersal to Madagascar, using oceanic currents that favoured eastward dispersal at that time, is a highly supported scenario matching the suggested dispersal of lemurs to this island. This is consistent with a previously suggested hypothesis based on molecular data.

The Key to Understanding the European Miocene Chalcides (Squamata, Scincidae) Comes from Asia: The Lizards of the East Siberian Tagay Locality (Baikal Lake) in Russia.

The early middle Miocene (MN 5) lizards from the East Siberian Tagay locality (Baikal Lake, Russia) in Asia are described here. The lizard fauna consists of two clades, Lacertidae and Scincidae. The skink material is allocated to Chalcides. While this taxon was previously reported from Europe, it has rarely been observed in the Neogene record with only jaw fragments and frontal bones described. Its taxonomy was therefore enigmatic. The Tagay material is almost identical to the European fossils of Chalcides from Austria and Hungary, but it also contains the parietal bone. While the material is also similar to the extant Ch. ocellatus, it exhibits several morphological differences. A new species is therefore erected-Chalcides augei sp. nov. These findings further support the connection of the Baikal Lake area with central Europe during the first half of the Miocene. The comparative anatomy of the frontals, parietals and lower jaws was evaluated by micro-CT in selected skink taxa. This comparison highlights several important differences, for example, paired frontals are present in Broadleysaurus (an outgroup taxon), in Acontias and all studied members of Scincidae herein. The character optimization in Mesquite supports fused frontals as being the condition at the basal node of the Ateuchosauridae + Sphenomorphidae + Eugongylidae + Lygosomidae + Egerniidae + Mabuyidae clade. While the parapineal foramen is restricted to the parietal in most taxa studied herein, it is absent (or vestigial) in Acontias and Feylinia. In contrast to all other skinks, this foramen is located on the frontal in Ateuchosaurus chinensis. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:1901-1934, 2020. © 2019 American Association for Anatomy.

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.

The atlas-axis complex in Dibamidae (Reptilia: Squamata) and their potential relatives: The effect of a fossorial lifestyle on the morphology of this skeletal bridge.

We report on the first detailed study of the atlas-axis complex in the lizard clade Dibamidae, a family of poorly known fossorial squamates distributed in tropical or subtropical climates. This skeletal bridge is characterized by several features, such as the complete absence of the first intercentrum or the appearance of the first free cervical rib on the axis (usually less developed in Dibamus relative to that in Anelytropsis). Our study shows morphological differences of the atlas-axis complex in the Mexican blind lizard Anelytropsis relative to those of Asian Dibamus, the only two known extant genera of this clade. With regard to taxonomy and phylogenetic topology of the Dibamidae within Squamata, a huge conflict exists between morphology versus molecules. The morphology of the atlas-axis complex is therefore compared with several potential sister clades + Sphenodon. Dibamids share several features with limbless Gekkota, Scincoidea, and Amphisbaenia. The complete absence of the first intercentrum is observed in Rhineura floridana and in Ateuchosaurus chinensis as well, and the free rib associated with the synapophyses of the axis is also present in Acontias meleagris. However, some of these features may result from a limbless, burrowing ecology and thus could represent homoplastic characters. In any case, the morphology of the atlas-axis shows that dibamids share most character states with skinks. Although the atlas-axis complex forms only an additional source of information, this conclusion is consistent with most morphological rather than molecular tree topologies.

The first Miocene fossils of Lacerta cf. trilineata (Squamata, Lacertidae) with a comparative study of the main cranial osteological differences in green lizards and their relatives.

We here describe the first fossil remains of a green lizard of the Lacerta group from the late Miocene (MN 13) of the Solnechnodolsk locality in southern European Russia. This region of Europe is crucial for our understanding of the paleobiogeography and evolution of these middle-sized lizards. Although this clade has a broad geographical distribution across the continent today, its presence in the fossil record has only rarely been reported. In contrast to that, the material described here is abundant, consists of a premaxilla, maxillae, frontals, parietals, jugals, quadrate, pterygoids, dentaries and vertebrae. The comparison of these elements to all extant green lizard species shows that these fossils are indistinguishable from Lacerta trilineata. Thus, they form the first potential evidence of the occurrence of this species in the Miocene. This may be also used as a potential calibration point for further studies. Together with other lizard fossils, Solnechnodolsk shows an interesting combination of survivors and the dawn of modern species. This locality provides important evidence for the transition of an archaic Miocene world to the modern diversity of lizards in Europe. In addition, this article represents a contribution to the knowledge of the comparative osteological anatomy of the selected cranial elements in lacertids. This study gives special emphasis to the green lizards, but new data are also presented for related taxa, e.g., Timon lepidus, Podarcis muralis or Zootoca vivipara. Although the green lizards include several cryptic species for which determination based on isolated osteological material would be expected to be difficult, our comparisons show several important morphological differences, although a high degree of variability is present.

Vertebral Comparative Anatomy and Morphological Differences in Anguine Lizards With a Special Reference to Pseudopus apodus.

The article reports on the first detailed vertebral and rib morphology of anguine taxon Pseudopus apodus using micro-computed tomography. A comparison shows significant morphological differences of vertebrae of Pseudopus relative to those of Anguis and Ophisaurus. Usually, there are 55 presacral vertebrae, two sacral, and 95-97 caudal vertebrae. Pseudopus apodus can be defined by 23 diagnostic features concerning the vertebral column. Although zygapophyseal articulation between atlas and axis is well developed in limbed anguid gerrhonotine lizards like Abronia or Barisia, it is absent in the extant representatives of the clade Anguinae, which are limbless. Thus, our study brings further support to the hypothesis about the complete reduction of this articulation in forms with reduced or absent limbs. Comparison of adult and juvenile morphology of vertebrae of P. apodus was also analyzed. Heterochrony in the evolution of this taxon was previously confirmed by its skull morphology and it can be also documented on the basis of vertebrae. Our data suggest that a peramorphic heterochronic process played a role in the evolution of this largest extant anguine species. Geometric morphometric analyses revealed a pattern of high vertebral disparity among species. We found a clear separation of limbless forms in morphospace. Pseudopus apodus always clusters within Ophisaurus-species confirming molecular and some morphological phylogenies. Only the first tail vertebra shows a distinct difference to those of other anguids, which might be related to altered locomotion associated to the larger body size in this species. Anat Rec, 302:232-257, 2019. © 2018 Wiley Periodicals, Inc.

From lizard body form to serpentiform morphology: The atlas-axis complex in African cordyliformes and their relatives.

The comparative vertebral morphology of the atlas-axis complex in cordyliforms, xantusiid and several skinks is studied here. These lizards are particularly interesting because of their different ecological adaptations and anti-predation strategies, where conformation ranges from the lizard-like body to a snake-like body. This transition to serpentiform morphology shows several evolutionary patterns in the atlas-axis complex: 1) the zygapophyseal articulations are lost in the early stage of the transition. In contrast to mammals, the atlas is more or less locked to the axis in lepidosaurs, but the absence of zygapophyseal articulation releases this locking for rotation. However despite its serpentiform morphology, Chamaesaura is different, in possessing this articulation; 2) the first intercentrum of Chamaesaura and Tetradactylus africanus (serpentiform grass-swimmers) is fully curved anteriorly, underlying the occipital condyle. While this limits ventral skull rotation beyond a certain angle, it locks the skull, which is a crucial adaptation for a sit-and-wait position in grassland habitats that needs to keep the head stabilized; and 3) in Acontias, most of the atlas articular surface with the occipital condyle is formed by the lateral aspect of the articulation area relative to the area located in the dorsal region of the slightly reduced intercentrum. A similar state occurs in amphisbaenians, most likely reflecting a fossorial lifestyle of the limbless lizards. Although Chamaesaura and Tetradactylus live sympatrically in grasslands, Chamaesaura differs in several ways in atlas-axis complex: for example, aforementioned presence of the atlas-axis zygapophyseal articulation, and long posterodorsal processes. Its occipital condyle protrudes further posteriorly, placing the atlas-axis complex further from the endocranium than in Tetradactylus. Hence, adaptation in the same niche, even among sister clades, can lead to different atlas-axis morphology due to different lifestyle strategies, for example, different foraging mode, while similar atlas-axis morphology can evolve in two lineages occupying different niches, as in Ablepharus and Scelotes.

The Quadratojugal of Eryops studied by computed tomography and the morphological variability of foramina and canals in the quadratojugal of basal tetrapods.

With respect to its large size and abundance, Eryops is an important representative of Permo-Carboniferous basal tetrapods and one of the best-known large temnospondyl amphibians of this period. This taxon forms a significant component of the Early Permian tetrapod fauna of Texas and New Mexico and here we describe a new record of skull remains, the first one from Brushy Creek (30 km northeast of Seymour) in Texas (Petrolia Formation, Wichita Group; Lower Permian - lower Artinskian). Our material, found in 2015, consists of a left nasal, a jaw fragment (premaxilla or maxilla), left quadratojugal fragments, and a partial left mandible. We used computed tomography methods (micro-CT) for imaging both internal and external structures, for the first time, for Eryops. The quadratojugal presented here is exceptional compared to all known basal tetrapods in having four different internal foramina. CT data show that these foramina are interconnected by canals within the bone. This indicates that the morphology of the foramina and the course of the canals in the quadratojugal of basal tetrapods are more variable than hitherto thought. Anat Rec, 299:1073-1079, 2016. © 2016 Wiley Periodicals, Inc.

Variation in the position of the jugal medial ridge among lizards (reptilia: squamata): its functional and taxonomic significance.

The course of the medial ridge in the lizard jugal shows considerable morphological variation. There are four basic configurations: (1) the medial ridge is located ventral to mid-height on the suborbital process and anterior to mid-length on the postorbital process; (2) the medial ridge is located ventrally on the suborbital process (as above), but posteriorly on the postorbital process; (3) the medial ridge is located dorsally on the suborbital process and anteriorly on the postorbital process; and (4) the medial ridge is centrally located along the entire length of the jugal. Ancestral character state reconstruction shows that type 1 is plesiomorphic for Squamata regardless of the broad-scale phylogenetic topology. Type 3 is present in chamaeleonids and convergently in Anolis barbatus. Type 3 is a synapomorphy of the chamaeleonids. Type 2 is considered plesiomorphic for Anguidae, Heloderma and Xenosaurus, although it is independently modified in some extant members. These taxa form a clade in molecular phylogenies of Squamata, and the course of the medial ridge of the jugal therefore provides some measure of morphological support for this arrangement. The course of the medial ridge may be best explained by the position of the eye and by the angle of the jugal; its relations with other bony orbital structures (supraocular osteoderms, palpebral, supraorbital flanges) and the posterior extent of the maxilla are also discussed.

The atlas-axis complex in chamaeleonids (Squamata: Chamaeleonidae), with description of a new anatomical structure of the skull.

The comparative vertebral morphology of different chamaeleonid genera has been generally neglected and some aspects such as the comparative anatomy of the neck region remain poorly known. The atlas and axis of all chamaeleonid genera (Brookesia, Rieppeleon, Archaius, Rhampholeon, Nadzikambia, Bradypodion, Chamaeleo, Calumma, Furcifer, Kinyongia, and Trioceros) are studied here. Considerable morphological differences are revealed. Additionally, some taxa exhibit sexual dimorphism in the atlas and axis. An extremely long, divided posterodorsal process is present in males of the Trioceros johnstoni + Trioceros jacksonii clade. The solid and well-developed morphology of the posterodorsal process in males of this taxon could reflect its competitive behavior-males fight with their horns and attempt to dislodge one another from branches during encounters. An additional area of insertion for the cervical musculature may indicate an incremental cervical musculature mass and cross sectional area that can add extra support and stability to the head and assist during combat involving lateral pushing. This character is not present in females. Heterochronic processes have played a role in the evolution of chamaeleonids, as evidenced in many characters of the atlas-axis complex. A new hypothesis of an anterior shifting of synapophyses of the axis is erected and a new derived anatomical structure of the parietal of Chamaeleo calyptratus is described (the processus parietalis inferior). The presence of the processus parietalis inferior is associated with the evolution of the dorsally elevated parietal crest.