A review of the Upper Permian genera Coelurosauravus, Weigeltisaurus and Gracilisaurus (Reptilia: Diapsida)

The holotype of the Upper Permian gliding reptile Weigeltisaurus jaekeli is redescribed. Comparison with the Madagascan reptile Coelurosauravus shows that the two forms are congeneric. Coelurosauravus Piveteau, 1926 has priority. The Upper Permian genus Gracilisaurus Weigelt, 1930 is reviewed and referred to Coelurosauravus.     

A Late Campanian sphenodontid (Reptilia, Diapsida) from northern Patagonia

The fossil record of sphenodontids in the Southern Hemisphere is much longer than in Laurasia, where they became extinct after Early Cretaceous times. Recent works demonstrated the persistence of at least eilenodontine sphenodontids until the ‘mid’-Cretaceous of Patagonia. Focused examination of the Los Alamitos Formation collections provided remains of Late Campanian sphenodontids. Although the phylogenetic position of these remains is still uncertain, they belong to a new, unrecorded species of sphenodontid perhaps related to opisthodontians or primitive, toothed sapheosaurs. To cite this article: S. Apesteguía, C. R. Palevol 4 (2005).     

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 Enigmatic Marine Reptile Nanchangosaurus from the Lower Triassic of Hubei,China and the Phylogenetic Affinities of Hupehsuchia

The study of the holotype and of a new specimen of Nanchangosaurus suni (Reptilia; Diapsida; Hupehsuchia) revealed a suite of hitherto unrecognized characters. For example, Nanchangosaurus has bipartite neural spines and its vertebral count is nearly identical to that of Hupehsuchus. It differs from the latter in having poorly developed forelimbs despite the advanced ossification in the rest of the skeleton. Other differences all pertain to hupehsuchian plesiomorphies retained in Nanchangosaurus, such as low neural spines. The relationship of Hupehsuchia within Diapsida was analyzed based on a data matrix containing 41 taxa coded for 213 characters, of which 18 were identified as aquatic adaptations from functional inferences. These aquatic adaptations may be vulnerable to the argumentation of character homology because expectation for homoplasy is high. There is an apparent incongruence between phylogenetic signals from aquatic adaptations and the rest of the data, with aquatic adaptations favoring all marine reptiles but Helveticosaurus to form a super-clade. However, this super-clade does not obtain when aquatic adaptations were deleted, whereas individual marine reptile clades are all derived without them. We examined all possible combinations of the 18 aquatic adaptations (n = 262143) and found that four lineages of marine reptiles are recognized almost regardless of which of these features were included in the analysis: Hupehsuchia-Ichthyopterygia clade, Sauropterygia-Saurosphargidae clade, Thalattosauria, and Helveticosaurus. The interrelationships among these four depended on the combination of aquatic adaptations to be included, i.e., assumed to be homologous a priori by bypassing character argumentation. Hupehsuchia always appeared as the sister taxon of Ichthyopterygia.     

On the anatomy and relationships ofEfraasia diagnostica (Huene) n. gen., a prosauropod dinosaur (Reptilia: Saurischia) from the Upper Triassic of Germany

Palaeosaurus (?) diagnosticus Huene from the Stubensandstone (Upper Triassic, Norian) of Nordwürttemberg, Germany is made the type species of a new genus of prosauropod dinosaurEfraasia, which is referred to the family Anchisauridae. The anatomy ofEfraasia diagnostica is conservative and it was an ideal ancestor for the later and more specializedAnchisaurus polyzelus of North America. The slenderness of the manus and pes ofEfraasia andAnchisaurus is matched amongst prosauropods only by the feet ofGyposaurus capensis andThecodontosaurus antiquus. Staurikosaurus pricei Colbert from the Upper Triassic of Brazil is probably a very primitive theropod dinosaur.     

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.     

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.     

First procolophonid (Reptilia,Parareptilia) from the Lower Triassic of Madagascar

Lasasaurus beltanae nov. gen. nov. sp., a new procolophonid (Reptilia, Parareptilia) from the Lower Triassic of Madagascar, is represented by a single partial skeleton preserved in a ferro-calcareous nodule from the Middle Sakamena Formation, in the North of the island. This new taxon is unique in possessing peculiar, fine and dendritic crests running along the posterolateral side of the squamosal, widely spaced maxillary teeth, subparallel mesiodistal ridges connecting maxillary teeth to the tooth row, and a strongly acute anterior margin of the copula (hyoid bone). This well-preserved specimen belongs to a juvenile individual. The inclusion of L. beltanae nov. gen. nov. sp. in a phylogenetic analysis suggests that it is close to Theledectinae, Procolophoninae, and Leptopleurinae, though their respective relationships are uncertain. This specimen is the first procolophonid described from Madagascar and represents a minor terrestrial component of a coastal vertebrate assemblage dominated by amphibious to fully-aquatic taxa.     

Braincase evolution in suchian archosaurs (Reptilia: Diapsida): evidence from the rauisuchian Batrachotomus kupferzellensis

The osteology of an almost complete braincase of the rauisuchian archosaur Batrachotomus kupferzellensis Gower from the Middle Triassic of Germany is described. There is a possibly discrete epiotic ossification, the metotic fissure is undivided by bone (i.e. there is a metotic foramen), the medial wall of the otic capsule is mostly ossified, the cerebral branch of the internal carotid artery entered the lateral surface of the parabasisphenoid, the ventral ramus of the opisthotic is more prominent laterally than a strong subvertical ridge on the exoccipital and basioccipital that lies posterior to the external foramen for the hypoglossal nerve, and the perilymphatic foramen faces away from the otic capsule in a posterior direction. Braincase morphology in the rauisuchians Saurosuchus galilei , Postosuchus kirkpatricki, and Tikisuchus romeri is reviewed. A matrix of 27 braincase characters for 12 archosaurian taxa is analysed. The most parsimonious hypothesis is consistent with the currently orthodox view of archosaurian phylogeny, except in that aetosaurians are more closely related to crocodylomorphs than is any rauisuchian. This phylogeny is used in a brief interpretation of the evolution of derived braincase features present in extant crocodilians. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 49–76.     

Grallator-Dominated Fossil Footprint Assemblages and Associated Enigmatic Footprints from the Chinle Group (Upper Triassic), Gateway Area,Colorado

The upper part of the Chinle Group (Late Triassic) of the Gateway area in western Colorado is extraordinarily rich in fossil footprint assemblages. Dominant track types include small Grallator tracks, generally attributed to Coelophysis-like theropods, which often occur in high densities of 50 to 100 per m². Other abundant ichnotaxa that are attributable to dinosaurs or dinosaur-like archosaurs include Pseudotetrasauropus and Tetrasauropus, attributed to prosauropods and sauropods, respectively. Several Pseudotetrasauropus-like tracks appear to be functionally didactyl and may indicate a new ichnotaxon that represents an animal that shows certain unusual features that are convergent with dromeosaurs and certain birds. Such convergence may reflect inherent growth programs as much as functional adaptations. Non-dinosaurian ichnotaxa include Brachychirotherium (probably of aetosaur affinity) and Rhynchosauroides, attributed to a sphenodontid/lizard-like form. Other ichnotaxa include probable therapsid (dicynodont) tracks labeled Pentasauropus sp., mammaloid (non-therian mammal and/or mammal-like reptile) tracks, and the trails of arthropods. Excellent preservation and high track densities mark the Gateway assemblages in a thin stratigraphic interval in the upper part of the Chinle Group (Rock Point Formation). The track assemblages are similar to those reported from the Chinle Group in other parts of the Colorado Plateau and Rocky Mountain region, extending over most of Colorado, Utah, northern Arizona and northern and eastern New Mexico. Some of the Chinle ichnotaxa (Grallator and Brachychirotherium) are found in the overlying Wingate Formation, indicating that it is also Late Triassic in age, at least in the lower part. However, overall the Chinle and Wingate assemblages are quite different, most notably in the rarity of mammaloid/mammal-like tracks in the Chinle Group.     

记云南罗平三叠纪小型始鳍龙类—新属种

根据一保存基本完整的骨架建立了始鳍龙类的一新属、种——纤细滇美龙(Dianmeisaurus gracilis gen.et sp.nov.)。标本采自云南省罗平县中三叠统关岭组Ⅱ段,为体态细长的小型物种。新属种与同产于该地区的利齿滇东龙(Diandongosaurus acutidentatus Shang et al.,2011)和丁氏滇肿龙(Dianopachysaurus dingi Liu et al.,2011a)身体各部位比例和个体大小接近,头骨均具有吻部两侧不收缩、眼眶显著大于上颞孔的特征。但新种眼眶间距要明显小于上颞孔间距,下颌关节与枕髁位于同一水平位置,锁骨前外侧缘的锁骨前突粗大而厚实,尺骨近端明显宽于远端,有比后两种更多的荐前椎(三属种荐前椎数分别为41、38、39枚)。此外,与滇东龙相对比,新种眶前区短于眶后区,前额骨和后额骨不相交。同时,新种下颌联合短,夹板骨参与下颌联合构成。新种前颌和下颌前部齿虽然也为獠齿(犬齿状齿),但未呈显著增大。头后骨骼对比,虽然新种和滇东龙均具有标志性的锁骨前外侧突,但后者的锁骨前外侧突尖细。与滇肿龙相对比,新种尾前部尾肋细长且未见明显加宽肿胀、仅具一个骨化的远侧腕骨、距骨为圆形且不具有近端凹入。去除数据不全的清镇龙、广西龙和三桥龙后的鳍龙类分支系统学分析表明,滇美龙与滇东龙互为姊妹群,同时它们与贵州龙、马家山龙和滇肿龙一起构成了一仅由在中国发现的属种组成的单系类群。这一单系类群与幻龙类的亲缘关系近于它们与欧洲肿肋龙类(Dactylosaurus,Anarosaurus,Serpianosaurus和Neusticosaurus)的亲缘关系。三种小型始鳍龙类的发现,指示云南罗平地区在三叠纪安尼期中晚期时鳍龙类是一类有较高分异度的海生爬行动物。     

Redescription and phylogenetic relationships of Megachirella wachtleri Renesto et Posenato, 2003 (Reptilia,Diapsida)

Megachirella wachtleri Renesto et Posenato, 2003, a well preserved partial reptile skeleton from the Middle Triassic of the Dolomites (N. Italy), was originally considered a lepidosauromorph, but no phylogenetic analysis was carried out. Consequently, the taxon was overlooked in later phylogenetic analyses of the Diapsida. Here, the holotype and only known specimen of M. wachtleri is redescribed, allowing an investigation of its phylogenetic relationships. Phylogenetic analyses confirm that Megachirella is a lepidosauromorph close to the crown group lepidosaurs (Squamata + Rhynchocephalia). Megachirella enhances our knowledge of the series of morphological modifications that led to the origin of the Lepidosauria, the most diverse clade of extant reptiles.     

Ultrastructural analysis of spermiogenesis in Iguana iguana (Reptilia: Sauria: Iguanidae)

Spermiogenesis in the lizard, Iguana iguana, was studied by transmission and scanning electron microscopy. During this process, structures such as the acrosomal complex in the spermatid head and the axonemal complex in the mid and principal pieces of the flagellum are formed. The nuclear content is initially compacted into thick, longitudinal chromatin filaments. Nuclear shape is determined by further compaction and by the manchette, a layer of microtubules surrounding the head. The acrosomal complex originates from Golgi vesicles and the interaction between the proacrosomal vesicle and the nucleus. The midpiece consists of a pair of centrioles, surrounded by a fibrous sheath and rings of simple and modified mitochondria. The centrioles sustain the axoneme that appears at the end of the midpiece. The axoneme extends throughout the principal piece of the flagellum with the 9 + 2 pattern, still surrounded by the fibrous sheath. In the endpiece, the axoneme continues, surrounded only by the plasma membrane. In the lumen of seminiferous tubules, immature spermatozoa retain abundant residual cytoplasm.     

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.     

A New Species of Garjainia Ochev, 1958 (Diapsida: Archosauriformes: Erythrosuchidae) from the Early Triassic of South Africa

A new species of the erythrosuchid archosauriform reptile Garjainia Ochev, 1958 is described on the basis of disarticulated but abundant and well-preserved cranial and postcranial material from the late Early Triassic (late Olenekian) Subzone A of the Cynognathus Assemblage Zone of the Burgersdorp Formation (Beaufort Group) of the Karoo Basin of South Africa. The new species, G. madiba, differs from its unique congener, G. prima from the late Olenekian of European Russia, most notably in having large bony bosses on the lateral surfaces of the jugals and postorbitals. The new species also has more teeth and a proportionately longer postacetabular process of the ilium than G. prima. Analysis of G. madiba bone histology reveals thick compact cortices comprised of highly vascularized, rapidly forming fibro-lamellar bone tissue, similar to Erythrosuchus africanus from Subzone B of the Cynognathus Assemblage Zone. The most notable differences between the two taxa are the predominance of a radiating vascular network and presence of annuli in the limb bones of G. madiba. These features indicate rapid growth rates, consistent with data for many other Triassic archosauriforms, but also a high degree of developmental plasticity as growth remained flexible. The diagnoses of Garjainia and of Erythrosuchidae are addressed and revised. Garjainia madiba is the geologically oldest erythrosuchid known from the Southern Hemisphere, and demonstrates that erythrosuchids achieved a cosmopolitan biogeographical distribution by the end of the Early Triassic, within five million years of the end-Permian mass extinction event. It provides new insights into the diversity of the Subzone A vertebrate assemblage, which partially fills a major gap between classic ‘faunal’ assemblages from the older Lystrosaurus Assemblage Zone (earliest Triassic) and the younger Subzone B of the Cynognathus Assemblage Zone (early Middle Triassic).     

Der Artbegriff and seine Bedeutung fur die Klassifikation der Echsen (Reptilia: Sauria)

Zusammenfassung Die einzelnen Artbegriffe werden zundchst in Grundzügen dargestellt und kritisiert. Dann wird die neue Definition, der phäno-genetische Artbegriff, entwickelt; sie lautet: eine Art ist die grötmögliche regionale Evolutionseinheit phäno-genetisch (in den typischen, artspezifischen Merkmalen) gleicher, sich selbst reproduzierender Deme. Sie ist von anderen sympatrischen Arten durch eine Fortpflanzungsisolation abgesondert, die ihr eine selbstandige, d.h. eine von anderen Arten unterschiedliche und hinreichend unbeeinflute, Evolution garantiert and die von einem anderen phäno-genetischen Merkmalssprung begleitet ist. Sie ist von anderen allopatrischen Arten durch einen phäno-genetischen Merkmalsprung abgesondert, der bei biparentalen Formen im Falle fehlender Fortpflanzungsisolation die Schlufolgerung erlaubt, da sich eine solche Isolation entwickelt hätte, wenn sie mit den allopatrischen Arten sympatrisch wäre. Diese Definition wird in verschiedenen evolutionären Situationen angewandt. Das jeweilige Ergebnis wird mit denen anderer Artbegriffe verglichen. Beispiele verdeutlichen die Auswirkung der verschiedenen Artbegriffe auf die Klassifikation der Echsen.

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Several species concepts are generally discussed and evaluated. Then the new definition, the pheno-genetic species concept, is developed; it reads: a species is the largest possible, regional evolutionary unit of pheno-genetically equal (in the typical, specific characters), identically reproducing demes. It is separated from sympatric species by a reproductive isolation that guarantees a unique evolution, an evolution different from that of other species and sufficiently uninfluenced, and that is accompanied by another distinctive pheno-genetic gap. It is separated from allopatric species by a distinctive pheno-genetic gap that - in the case of missing reproductive isolation in bisexual forms — allows the conclusion that such an isolation would have developed, if the species had been sympatric with the allopatric species. This definition is applied to several situations of evolution. The effects of the concepts on the classification of lizards are demonstrated by examples.

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Übersetzung eines Vortrages vor dem 11. Europäischen Kongress für Herpetologie, Toulouse, 1. - 6. September 1975, vgl. Bull.Soc.Zool. France 100(4), 676–677.     

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 large species of gecko of the genus Hemidactylus Oken, 1817 (Reptilia: Sauria: Gekkonidae) from the Eastern Ghats,India

A new large species of gecko superficially resembling Hemidactylus maculatus Duméril & Bibron and its allied large Hemidactylus is described from Eastern Ghats. The new species, Hemidactylus kangerensis sp. nov. can be easily distinguished from members of the H. maculatus complex based on the number of femoral pores, i.e. 18–21 on each side, separated medially by 4 non-pored scales. A phylogenetic analysis based on partial sequence of mitochondrial gene cytochrome b shows that the gecko is a member of the “H. prashadi” clade and is sister to H. maculatus, from which it differs in an uncorrected pairwise sequence divergence of 10%. The discovery of a new large gecko from Eastern Ghats advocates the need for biodiversity assessment across the neglected Eastern Ghats.