New basal procolophonid reptile from the Katberg formation (Lower Triassic) of the South African Karoo

A new procolophonid reptile, Kitchingnathus untabeni n. gen. et n. sp., is described from the uppermost strata of the Lystrosaurus Assemblage Zone of the Karoo Basin, South Africa. The new taxon co-occurs with the well-known Procolophon trigoniceps. The most distinctive feature of the new taxon is the presence of numerous small bicuspid molariforms in both the maxilla and the dentary. A phylogenetic analysis indicates that Kitchingnathus occupies a basal position among procolophonids. Character optimisation suggests that bicuspid teeth were acquired independently by the new taxon, and originated twice in procolophonid evolution.     

The first ctenosauriscid (Reptilia: Archosauromorpha) from the Lower Triassic Of Eastern Europe

A new Early Triassic thecodont from the Donskaya Luka locality is described. A new species and genus of Rauisuchidae, Bystrowisuchus flerovi gen. et sp. nov., the first East European and earliest known member of the family Ctenosauriscidae is established. The taxonomy and phylogeny of Rauisuchidae and their stratigraphical and geographical distribution in connection with new finds are discussed.     

The First Articulated Trematosaur 'amphibian' from the Lower Triassic of Madagascar: Implications for the Phylogeny of the Group

The temnospondyl Wantzosaurus elongatus Lehman, 1961, from the Lower Triassic of Madagascar, is redescribed on the basis of a new specimen, which is the most complete trematosaur ever found. Detailed osteological observations and comparisons provide new data on the anatomy, ontogeny, palaeobiology and palaeoecology of this peculiar marine 'amphibian'. The morphology of this aquatically readapted taxon is compared to that of marine 'reptiles': Wantzosaurus might have been able to swim by undulation. A phylogenetic analysis of the trematosaurs is performed for the first time and suggests that Wantzosaurus is a derived taxon within the clade Trematosauridae. The family is defined on the basis of derived character states and is shown to be monophyletic.     

New tanystropheids (Reptilia: Archosauromorpha) from the Triassic of Europe

A new prolacertilian species and genus, Augustaburiania vatagini gen. et sp. nov. (Reptilia: Archosauromorpha), from the Lower Triassic of the Don River Basin is described. It is the first representative of the Tanystropheidae in the Eastern Europe and the world oldest member of this family. Another new genus (Protanystropheus gen. nov.) from Central and Western Europe is also established. The diversity, systematics, phylogeny, evolution, and stratigraphic and geographical distribution of prolacertilians are discussed. The ecological role of prolacertilians in Early Triassic communities and adaptation to marine environments are analyzed.     

New beetles (Insecta,Coleoptera) from the Lower Triassic of European Russia

Fossil remains of beetles are described from two Lower Triassic localities: Entala (Induan) and Tikhvinskoe (Olenekian). Only one beetle fossil was previously known from the Lower Triassic of Tikhvinskoe. The fossils are rather few and poorly preserved, but they are worth describing as finds rare for the Lower Triassic. Five fossils from Entala most probably belong to beetles of the same species of the formal genus Pseudochrysomelites. Beetles of this genus are especially abundant in deposits close to the Permian–Triassic boundary and can be considered “disaster taxa.” There are no known cases, either in the Permian or in the Middle–Upper Triassic, of a random sample of five specimens belonging to a single species. This suggests that in the Entala oryctocenosis the species diversity of beetles is extremely low. All three beetle fossils found in Tikhvinskoe belong to beetles of different species, showing that diversity had already started to increase. However, it remained low, and all fossils belong to the formal family Schizocoleidae, and two of the three belong to the same genus, Pseudochrysomelites. The Khei-Yaga locality, which immediately follows Tikhvinskoe in time (topmost Olenekian or early Anisian), already contains beetles of the families Asiocoleidae and Permosynidae. In the Lower Anisian of the Buntsandstein, such typical Mesozoic beetles as Cupedidae and Coptoclavidae have been recorded. The appearance of such advanced beetles as early as the Lower Anisian suggests that the famous Permian–Triassic crisis was not as deep as it is usually believed, and many beetles survived it, disappearing, however, from the fossil record in the Early Triassic.     

The anatomy of the palatoquadrate in the Lower Triassic Proterosuchus fergusi (Reptilia,Archosauromorpha) and its morphological transformation within the archosauriform clade

The anatomy of the palatoquadrate ossifications of the Lower Triassic archosauromorph Proterosuchus fergusi from South Africa is described. It consists of two ossifications, the epipterygoid and the quadrate, which were joined by cartilage in life. The margins of the cartilage are clearly indicated by ridges and grooves on the dorsal surface of the pterygoid. The epipterygoid ossification consists of two structures: the anteroposteriorly expanded basal portion and, dorsally from it, an extending, slender, ascending process. From the anterior margin of the basal portion of the epipterygoid, a plate‐like structure, herein called the lamina epipterygoidea anteromedialis, extends anteromedially to form the anterolateral wall of the cavum epiptericum. Comparisons with the similarly constructed embryonal and adult epipterygoid components of Sphenodon punctatus show that the anteromedial lamina of the epipterygoid of P. fergusi is an additional component of the epipterygoid of this species and that this lamina is absent in the former species. However, a structure in a topologically similar position to the anteromedial lamina of the epipterygoid of P. fergusi is present in the palatoquadrate of Alligator mississippiensis. In the latter species, the structure is called the lamina palatoquadrati anterior; it ossifies in membrane and forms the dorsolateral cover of the huge trigeminal ganglion. It is hypothesized here that the anteromedial lamina of the epipterygoid of P. fergusi and the anterior lamina of the palatoquadrate of A. mississippiensis are most probably homologous structures and are present in both the basal and one of the crown taxa of the archosauromorph clade, respectively.     

Skeletal microstructure of Stenopterygius quadriscissus (Reptilia,Ichthyosauria) from the Posidonienschiefer (Posidonia Shale,Lower Jurassic) of Germany

Ichthyosaurians (Ichthyosauria) are a major clade of secondarily aquatic marine tetrapods that occupied several major predatory niches during the Mesozoic Era. Multiple lines of evidence including isotopic, body shape and swimming modality analyses suggest they exhibited elevated growth and metabolic rates, and body temperatures. However, applications of osteohistological methods to test hypotheses regarding their physiology are few. Previous studies focused on the humeri, vertebrae and ribs from a small number of taxa. Here, we use osteohistological methods to describe the bone microstructure of over 30 cranial and post‐cranial elements from a nearly complete, articulated individual of Stenopterygius quadriscissus from the Posidonienschiefer Formation (Posidonia Shale, Lower Jurassic) of Germany. The specimen shows highly vascularized primary bone and spongious secondary bone in its limbs, suggesting an overall shift to a lighter spongious‐structured skeleton was achieved through multiple developmental mechanisms. The modified perichondral ossification in elements of the limbs distal to the stylopodium informs our understanding of functional morphology, including hydrodynamic forces on the paddles. The ribs show variation in cortical thickness and trabecular organization along their length. Cyclical growth is inferred from changes in vascularization and osteocyte density as well as the presence of annuli in primary fibrolamellar bone. Cranial elements, due to their relative density and better preservation of growth marks, may prove to be of particular importance in future skeletochronological studies of post‐Triassic ichthyosaurians. We infer and corroborate hypotheses of elevated growth rates and metabolic rates in ichthyosaurians, and the potential for thermoregulation similar to extant homeothermic ectotherms.     

A new genus of pliosaur (Reptilia: Sauropterygia) from the Lower Jurassic of Holzmaden,Germany

Abstract: The exquisitely preserved holotype of the pliosaur ‘Rhomaleosaurus’victor (SMNS 12478) is described from the Toarcian Posidonien‐Schiefer (Upper Lias, Lower Jurassic) of Holzmaden (Baden‐Württemberg), Germany. The specimen presents a novel combination of synapomorphies and unique morphometric proportions separating it from Rhomaleosaurus sensu stricto and warranting the erection of a new genus, Meyerasaurus gen. nov. Historically, the name ‘Thaumatosaurus’ has been interchangeable with Rhomaleosaurus and is frequently associated with SMNS 12478 in the literature. However, this is an invalid taxon and cannot be reinstated. The anatomy of Meyerasaurus victor is compared in detail with other pliosaurs, and its taxonomic affinity is reviewed. M. victor belongs to the family Rhomaleosauridae and shares several anatomical characters with Rhomaleosaurus including a short and robust premaxillary rostrum (length‐to‐width ratio c. 1.0), parallel premaxilla–maxilla sutures anterior to the nares, vomers contacting the maxillae posterior to the internal nares, and c. 28 cervical vertebrae minus the atlas–axis. The known geographical distribution of Rhomaleosaurus, which previously extended across the German and English palaeobiogeographical zones, is reduced to the English zone as a consequence of the referral of SMNS 12478 to a new genus. This is significant because it contributes to an ongoing trend of increasing generic separation between the German and English zones, while increasing the generic diversity within the German zone itself.     

A new heterodontosaurid dinosaur (Reptilia: Ornithischia) from the Upper Triassic Red Beds of Lesotho

A new species of ornithischian dinosaur ( Lycorhinus consors sp. nov.) is established on a skull from the Upper Triassic Red Beds of Lesotho. This ornithischian is assigned to the family Heterodontosauridae of the suborder Ornithopoda. The dinosaurs of the family Heterodonto-sauridae are reviewed: Geranosaurus atavus Broom (1911) is considered a nomendubium and the genus name Heterodontosaurus Crompton & Charig (1962) is held to be a junior synonym for Lycorhinus Haughton (1924).
Functional and palaeoecological implications of the heterodontosaurid dentition are discussed. The pattern of tooth wear may reflect a highly specialized jaw action which involved protraction and retraction of the mandible to produce a grinding effect between upper and lower cheek teeth. Lycorhinus consors is presumed to be a female heterodontosaurid because it differs from all other heterodontosaurids in lacking caniniform tusks. It is suggested that the tusks of heterodontosaurids were functionally analogous to those of tayassuids and tragulids and that they were employed as weapons for intra-specific combat and defence. Dental peculiarities indicate that tooth replacement processes were suppressed in heterodontosaurids; replacement of the teeth seems to have been restricted to a brief period each year (presumably when heterodontosaurids underwent aestivation or hibernation).
A new diagnosis is formulated for the family Heterodontosauridae. The relationships of early ornithopod dinosaurs are briefly reviewed and a new classification is proposed. Ten families of ornithopod dinosaurs are recognized; these are ranked in two grades-one (named Dolichopoda) representing the conservative main stem of the ornithischian phylogenetic tree and the other (named Brachypoda) comprising the several more advanced lines of ornithopod evolution.     

The genus Paragehyra (Reptilia: Sauria: Gekkonidae) in southern Madagascar

Paragehyra Angel is a gekkonine genus endemic to Madagascar. From 1929 until the present, the genus was represented only by the holotype of the single known species. Paragehyra petiti. The type locality of P. petiti is in the driest part of Madagascar (south-west) where rainless periods may exceed 11 months and the original vegetation consists of thorn scrub or dry deciduous forest. No new individuals of P. petiti were observed. However, a second, undescribed species of the genus was discovered in October 1990 on the opposite side of the island (south-east) in primary-low montane rainforest. The new form was observed at four localities at the southernmost extremity of rainforest in Madagascar.     

Cranial morphology of Platypterygius longmani Wade, 1990 (Reptilia: Ichthyosauria) from the Lower Cretaceous of Australia

Exceptionally well-preserved remains belonging to the Australian Early Cretaceous (Albian) ichthyosaur Platypterygius longmani Wade, 1990 are described in detail. The material is used to reconstruct some of the cranial musculature and provide a brief functional analysis of the skull and mandible. Preparation of specimens using acetic acid and application of high-resolution X-ray computed tomographic analyses has revealed many previously unrecorded anatomical details, including the absence of a coronoid and squamosal, and the presence of well-developed epipterygoid facets on the pterygoid and parietal (possibly indicating retention of an unossified epipterygoid element). Reconstruction of the jaw musculature suggests a well-developed m. adductor mandibulae internus pterygoideus (serving to close the jaws against inertia and drag of the surrounding water), and possibly an m. intramandibularis (acting to accentuate static pressure along the elongate mandible when the jaws were closed). Despite its large size (maximum total body length of around 7 m), the long, narrow snout of P. longmani (together with preserved gut contents) indicates selective feeding on relatively small prey such as fish, small tetrapods and probably cephalopods. Large orbits are indicative of a visual hunter, but extensive external passages for nerves and blood vessels might suggest the presence of a dermal sensory system. The massive stapes appears to have lost all sound conductive function, serving primarily as structural support in the basicranial region.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 583–622.     

Evidence of Spondyloarthropathy in the Spine of a Phytosaur (Reptilia: Archosauriformes) from the Late Triassic of Halberstadt,Germany

Pathologies in the skeleton of phytosaurs, extinct archosauriform reptiles restricted to the Late Triassic, have only been rarely described. The only known postcranial pathologies of a phytosaur are two pairs of fused vertebrae of “Angistorhinopsis ruetimeyeri” from Halberstadt, Germany, as initially described by the paleontologist Friedrich von Huene. These pathologic vertebrae are redescribed in more detail in this study in the light of modern paleopathologic methods. Four different pathologic observations can be made in the vertebral column of this individual: 1) fusion of two thoracic vertebral bodies by new bone formation within the synovial membrane and articular capsule of the intervertebral joint; 2) fusion and conspicuous antero-posterior shortening of last presacral and first sacral vertebral bodies; 3) destruction and erosion of the anterior articular surface of the last presacral vertebra; and 4) a smooth depression on the ventral surface of the fused last presacral and first sacral vertebral bodies. Observations 1–3 can most plausibly and parsimoniously be attributed to one disease: spondyloarthropathy, an aseptic inflammatory process in which affected vertebrae show typical types of reactive new bone formation and erosion of subchondral bone. The kind of vertebral shortening present in the fused lumbosacral vertebrae suggests that the phytosaur acquired this disease in its early life. Observation 4, the smooth ventral depression in the fused lumbosacral vertebrae, is most probably not connected to the spondyloarthropathy, and can be regarded as a separate abnormality. It remains of uncertain origin, but may be the result of pressure, perhaps caused by a benign mass such as an aneurysm or cyst of unknown type. Reports of spondyloarthropathy in Paleozoic and Mesozoic reptiles are still exceptional, and our report of spondyloarthropathy in fossil material from Halberstadt is the first unequivocal occurrence of this disease in a Triassic tetrapod and in a phytosaur.     

Systematics, speciation and biogeography of the dwarf chameleons (Brookesia; Reptilia, Squamata, Chamaeleontidae) of northern Madagascar

Brookesia dwarf chameleons, endemic to Madagascar, were surveyed at the following localities in northern Madagascar (north of 16°S): Montagne d'Ambre, Ankarana, Manongarivo, Tsaratanana, Marojejy and Masoala. A total of 15 species occur in this region. Six new species are described and five new synonyms are identified. The genus Brookesia , the most speciose chamaeleontid genus in Madagascar, contains 23 species.
Almost all the northern Brookesia species are restricted to rainforest and occupy a relatively narrow elevational range. Although the northern rainforests represent just one-third of the total rainforest and about 5% of the total island area, 65% of the Brookesia species occur in this region, and 52% are endemic to the northern rainforest. Five new biogeographic regions of the northern rainforest are identified based on centres of Brookesia endemicity: Montagne d'Ambre, Northwest, Tsaratanana, Northeast and East. Speciation is thought to have been facilitated in the north through geographic isolation, with the Tsaratanana mountain range and the dry forests south of Montagne d'Ambre forming barriers to dispersal, and the Tsaratanana mountains acting as a centre of isolation. The fragmented distribution of several Brookesia species of low altitude rainforest suggests a period in Madagascar's history when the climate was wetter and low altitude rainforest much more widespread.     

Elasmobranchs from the Lower Triassic Sulphur Mountain Formation near Wapiti Lake (BC, Canada)

Hybodontoid and nonhybodontoid sharks are described from the Lower Triassic Vega‐Phroso Siltstone Member of Sulphur Mountain Formation on the basis of newly discovered material. The age of the classic fossil site ‘Wapiti Lake’ in the Canadian Rocky Mountains is discussed on the basis of new field data and one conodont found in association. Preliminary results suggest that these elasmobranch remains are between early Smithian and Spathian in age. Apart from the enigmatic genus Listracanthus and previously reported edestoids, the shark fauna consists of at least one hybodont, at least two questionable hybodontoid genera and an elasmobranch of enigmatic affinities, represented by peculiar denticles only and described as ‘genus A’incertae sedis. The presence of the only previously reported hybodont genus, cf. Palaeobates, is erroneous. The largest specimen represents the most complete Early Mesozoic shark known. The heterodonty of its dentition, fin spine morphology and the short, robust body shape imply it represents a member of a new family of shark, Wapitiodidae fam. nov. , and is described here as Wapitiodus aplopagus gen. et sp. nov. The unique dental morphology shows affinities to Polyacrodus but clearly differs in the complete lack of side cusps. Wapitiodus gen. nov. possesses a primitive fin spine structure. The tooth crowns are entirely blunt in the distal (posterior) tooth files, and are acuminate‐unicuspid in several anterior files. Tooth morphology, the shape of the basal cartilages, the proximal insertion of the fin spines and the pectoral fin structure are interpreted as diagnostic characters for this new genus, and possibly for the Wapitiodidae fam. nov. The majority of observed characters appear to be primitive and are reminiscent of Palaeozoic sharks, however, and these features include dorsal fin spine morphology and gross skull anatomy. A second species, provisionally placed in the same genus, is described as Wapitiodus homalorhizo sp. nov. Wapitiodus homalorhizo sp. nov. can be distinguished from W. aplopagus gen. et sp. nov. by the proportions of the fin spines, tooth morphology and possibly the body shape. Several isolated teeth and other fragmentary material are referred to either Wapitiodus gen. nov. sp. indet. or to ?Polyacrodus sp. (Polyacrodontidae gen. et sp. indet.). A third genus of elasmobranch (incertae sedis) is described as ‘Genus A’ and is recognized by its peculiar scales. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149 , 309–337.     

New species of Thylacocephala (Arthropoda) from the Spathian (Lower Triassic) of Chaohu,Anhui Province of China

We describe a new species of Thylacocephala, Ankitokazocaris chaohuensis sp. nov., from the Upper Spathian (Early Triassic) of Chaohu, Anhui Province, China. It is diagnosed by its unique outline of the carapace, small size, and the narrow and asymmetrical anterior notch. Fine preservation reveals at least 14 posterior appendages, traces of gills and raptorial appendages, and remains of trunk segments and muscles in Ankitokazocaris for the first time. This is the first report of Thylacocephala from the Early Triassic of China, confirming the wide distribution of this group in the Tethys. The new species is closer to the type species of Ankitokazocaris, which is stratigraphically somewhat older, than to a recently described almost coeval species from Japan.     

A partial systematic revision of the day geckos, Phelsuma Gray, of Madagascar (Reptilia: Squamata: Gekkonidae)

Recent survey work in Madagascar has led to significant changes in the systematics of Madagascan Phelsuma. A new species from the Masoala Peninsula in northeastern Madagascar is described, which has a nostril position typical of Phelsuma from Mauritius, Reunion and Rodriguez islands. Phelsuma breviceps was rediscovered in extreme southern Madagascar and is resurrected from the synonymy of P. mutabilis. Phelsuma breviceps has fragile skin, a probable predator escape mechanism, similar to the condition found in geckos of the genera Ailuronyx and Geckokpis. Three new synonyms are recognized. Phelsuma befotahensis and P. checki are junior synonyms of P. abbotti , and P. minuthi is a junior synonym of P. lineata. Twenty species of Phelsuma occur in Madagascar, of which 16 are endemic. A checklist and identification key are provided. Conservation problems of Phelsuma in the coastal regions of Madagascar are briefly discussed.     

A new specimen of the Triassic pistosauroid Yunguisaurus,with implications for the origin of Plesiosauria (Reptilia,Sauropterygia)

An adult skeleton of the pistosauroid sauropterygian Yunguisaurus liae reveals a number of morphological features not observed in the holotype, such as the complete morphology of the skull roof, stapes, atlas and axis, ventral view of the postcranium, and nearly complete limbs and tail. Size and morphological differences between the two specimens are mostly regarded as ontogenetic variation, and newly added data did not affect the phylogenetic relationships with other pistosauroids significantly. The number of mesopodia (11 carpals and 8 tarsals) exceeds that known in any other Triassic marine reptiles and does not serve as a precursor of the plesiosaurian pattern with fewer mesopodia of different topology; it demonstrates variability of the limb morphology among the Triassic pistosauroids. The pectoral girdles of Corosaurus, Augustasaurus and Yunguisaurus may indicate early stages of the adaptation towards the plesiosaurian style of paraxial limb movements with ventroposterior power stroke.