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A major macroevolutionary question concerns how long-term patterns of body-size evolution are underpinned by smaller scale processes along lineages. One outstanding long-term transition is the replacement of basal therapsids (stem-group mammals) by archosauromorphs, including dinosaurs, as the dominant large-bodied terrestrial fauna during the Triassic (approx. 252–201 million years ago). This landmark event preceded more than 150 million years of archosauromorph dominance. We analyse a new body-size dataset of more than 400 therapsid and archosauromorph species spanning the Late Permian–Middle Jurassic. Maximum-likelihood analyses indicate that Cope's rule (an active within-lineage trend of body-size increase) is extremely rare, despite conspicuous patterns of body-size turnover, and contrary to proposals that Cope's rule is central to vertebrate evolution. Instead, passive processes predominate in taxonomically and ecomorphologically more inclusive clades, with stasis common in less inclusive clades. Body-size limits are clade-dependent, suggesting intrinsic, biological factors are more important than the external environment. This clade-dependence is exemplified by maximum size of Middle–early Late Triassic archosauromorph predators exceeding that of contemporary herbivores, breaking a widely-accepted ‘rule’ that herbivore maximum size greatly exceeds carnivore maximum size. Archosauromorph and dinosaur dominance occurred via opportunistic replacement of therapsids following extinction, but were facilitated by higher archosauromorph growth rates.

The early evolution of archosauromorphs during the Permo-Triassic constitutes an excellent empirical case study to shed light on evolutionary radiations in deep time and the timing and processes of recovery of terrestrial faunas after a mass extinction. However, macroevolutionary studies of early archosauromorphs are currently limited by poor knowledge of their phylogenetic relationships. In particular, one of the main early archosauromorph groups that need an exhaustive phylogenetic study is “Proterosuchia,” which as historically conceived includes members of both Proterosuchidae and Erythrosuchidae. A new data matrix composed of 96 separate taxa (several of them not included in a quantitative phylogenetic analysis before) and 600 osteological characters was assembled and analysed to generate a comprehensive higher-level phylogenetic hypothesis of basal archosauromorphs and shed light on the species-level interrelationships of taxa historically identified as proterosuchian archosauriforms. The results of the analysis using maximum parsimony include a polyphyletic “Prolacertiformes” and “Protorosauria,” in which the Permian Aenigmastropheus and Protorosaurus are the most basal archosauromorphs. The enigmatic choristoderans are either found as the sister-taxa of all other lepidosauromorphs or archosauromorphs, but consistently placed within Sauria. Prolacertids, rhynchosaurs, allokotosaurians and tanystropheids are the major successive sister clades of Archosauriformes. The Early Triassic Tasmaniosaurus is recovered as the sister-taxon of Archosauriformes. Proterosuchidae is unambiguosly restricted to five species that occur immediately after and before the Permo-Triassic boundary, thus implying that they are a short-lived “disaster” clade. Erythrosuchidae is composed of eight nominal species that occur during the Early and Middle Triassic. “Proterosuchia” is polyphyletic, in which erythrosuchids are more closely related to Euparkeria and more crownward archosauriforms than to proterosuchids, and several species are found widespread along the archosauromorph tree, some being nested within Archosauria (e.g., “Chasmatosaurus ultimus,” Youngosuchus). Doswelliids and proterochampsids are recovered as more closely related to each other than to other archosauromorphs, forming a large clade (Proterochampsia) of semi-aquatic to aquatic forms that includes the bizarre genus Vancleavea. Euparkeria is one of the sister-taxa of the clade composed of proterochampsians and archosaurs. The putative Indian archosaur Yarasuchus is recovered in a polytomy with Euparkeria and more crownward archosauriforms, and as more closely related to the Russian Dongusuchus than to other species. Phytosaurs are recovered as the sister-taxa of all other pseudosuchians, thus being nested within Archosauria. Fil: Ezcurra, Martin Daniel. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina

CAPES Apresentamos uma descrição detalhada da morfologia e osteohistologia de pequenos ossos de Arcosauromorpha encontrados da Zona de Associação de Hyperodapedon, referente à Sequência Santa Maria II, Supersequência Santa Maria, Triássico Superior do Rio Grande do Sul, Brasil. O material é composto por ossos de tamanho reduzido, com dimensões menores do que 57mm, encontrados associados in situ. O espécime é composto por um úmero e um metatarsal III direitos, extremidade proximal de uma falange ungueal, um arco neural quase completo de uma vértebra pré-sacral e um centro vertebral pré-sacral. Comparações morfológicas entre UFSM11326 e outros organismos do Triássico mostraram se tratar de um dinossauriano, com possível associação ao grupo Sauropodomorpha. São elas: morfologia da crista deltopeitoral do úmero apresentando prolongamento do cume na crista deltoide; eixo de torção das porções proximal e distal do úmero; morfologia do metatarsal III, apresentando ângulo de torção entre as extremidades proximal e distal superior a 60° e presença do ombro medial; posições da diapófise, parapófise e das infracavidades diapofisiais restantes no arco neural. Este grupo de arcossauros já foi registrado anteriormente em estratos triássicos da Supersequência Santa Maria. As análises do fechamento das suturas do arco neural indicaram um indivíduo jovem, com suturas abertas em forma de zíper. As sessões finas da diáfise do úmero e do metatarsal III, corroboram este estado ontogenético, indicando se tratar de um espécime em fase inicial de desenvolvimento. Exibiu um complexo ósseo fibrolamelar (comum em espécimes dinossaurianos), composto por ósteons primários, sem ocorrência de marcas de crescimento (LAGs ou annuli), áreas extensas de reabsorção ou a presença de lamelas circunferenciais externas (external fundamental system = EFS). Este padrão evidencia uma estratégia de crescimento rápido, sustentado por elevadas taxas metabólicas, superiores a dos répteis modernos, e comparáveis com aquelas já encontradas para este grupo de arcossauros. Definimos portanto que UFSM11326 corresponde a um Sauropodomorpha jovem, sem a conclusão de seu crescimento assintótico, com elevados níveis de deposição óssea e consequente crescimento acelerado até o momento de sua morte. Here we present a detailed description of the morphology and osteohistology of small bones of Arcosauromorpha found at the Hyperodapedon Association Zone, referring to the Santa Maria II Sequence, Santa Maria Supersequence, Upper Triassic of Rio Grande do Sul States, Brazil. The material consists of small sized bone (dimensions smaller than 57mm) found associated in situ. The specimen consists of a right humerus and metatarsal III, proximal extremity of an ungueal phalanx, an almost complete neural arch of a pre-sacral vertebra, and a pre-sacral vertebral center. Morphological comparisons between UFSM11326 and other organisms from Triassic revealed that it is a dinosaurian, possibly related to Sauropodomorpha group. The similarities include the morphology of the deltopectoral humerus crest that has an extension of the deltoid crest; twist axis from proximal and distal portions of the humerus; morphology of metatarsal III with torsion angle between the proximal and distal extremities greater than 60° and medial shoulder; positions of the infracavities of the apophysis, parapophysis and diapophysal remaining at neural arch. This archosaurs group has already been registered previously in Triassic strata of the Santa Maria Supersequence. The analyses of the closure of the sutures of the neural arch indicate it to be a young individual with open sutures zipper-shaped. The final sessions of the humeral diaphysis and metarsal III corroborate this ontogenetic state, showing it to be a specimen at an early stage of development. A fibrolamellar bone complex that is common in dinosaurian specimens is present, composed of primary osteons, without occurrence of growth marks (LAGs or annuli), extensive reabsorption areas or external circumferential lamellas (external fundamental system = EFS). This pattern shows a rapid growth strategy, supported by high metabolic rates that are greater than in modern reptiles and comparable with those already found to this archosaurs group. We conclude that UFSM11326 corresponds to a young Sauropodomorpha without completing its asymptotic growth and with high levels of bone deposition and subsequent rapid growth until the moment of his death.

Made available in DSpace on 2019-10-04T12:33:09Z (GMT). No. of bitstreams: 0 Previous issue date: 2016-01-01 Emmy Noether Programme Award from the Deutsche Forschungsgemeinschaft Marie Curie Career Integration Grant Brazil Visiting Fellows scheme of the University of Birmingham The rhynchosaurian archosauromorphs are an important and diverse group of fossil tetrapods that first appeared during the Early Triassic and probably became extinct during the early Late Triassic (early Norian). Here, the early evolution of rhynchosaurs during the Early and early Middle Triassic (Induan-Anisian: 252.2-242 Mya) is reviewed based on new anatomical observations and their implications for the taxonomy, phylogenetic relationships and macroevolutionary history of the group. A quantitative phylogenetic analysis recovered a paraphyletic genus Rhynchosaurus, with Rhynchosaurus brodiei more closely related to hyperodapedontines than to Rhynchosaurus articeps. Therefore, a new genus is erected, resulting in the new combination Langeronyx brodiei. A body size analysis found two independent increases in size in the evolutionary history of rhynchosaurs, one among stenaulorhynchines and the other in the hyperodapedontine lineage. Maximum likelihood fitting of phenotypic evolution moths to body size data found ambiguous results, with body size evolution potentially interpreted as fitting either a non-directional Brownian motion model or a stasis model. A Dispersal-Extinction-Cladogenesis analysis reconstructed the areas that are now South Africa and Europe as the ancestral areas of Rhynchosauria and Rhynchosauridae, respectively. The reconstruction of dispersal events between geographic areas that are broadly separated paleolatitudinally implies that barriers to the dispersal of rhynchosaurs from either side of the paleo-Equator during the Middle Triassic were either absent or permeable. Museo Argentino Ciencias Nat Bernardino Rivadavia, Secc Paleontol, Buenos Aires, DF, Argentina Univ Birmingham, Sch Geog Earth & Environm Sci, Birmingham, W Midlands, England Univ Estadual Paulista, Dept Biol & Zootecnia, FEIS, Ilha Solteira, Brazil Univ Estadual Paulista, Dept Biol & Zootecnia, FEIS, Ilha Solteira, Brazil Emmy Noether Programme Award from the Deutsche Forschungsgemeinschaft: BU 2587/3-1 Marie Curie Career Integration Grant: PCIG14-GA-2013-630123 ARCHOSAUR RISE

© 2016, © 2014 Taylor & Francis. European Russia has yielded several fragmentary but potentially important archosauriform specimens from the Middle Triassic, but these have been only briefly described in the literature. One of these puzzling taxa is Dongusuchus efremovi Sennikov, 1988, described from the Donguz Svita. We present a redescription of Dongusuchus efremovi, which includes the first photographic atlas and thorough anatomical description of the holotype and referred specimens. This taxon is shown to be a gracile, probably fast-running species with elongate and slender limbs. A phylogenetic analysis recovers Dongusuchus efremovi as an early-diverging, non-archosaurian archosauriform. Previous work had suggested that this taxon was a ‘rauisuchid’. The gracile proportions of the femur and somewhat wedge-shaped head, however, are unusual for basal archosauriforms and are similar to the plesiomorphic state in crocodile and avian-line crown archosaurs. Several Early-Middle Triassic basal archosauriforms and early members of the crocodile and avian lineages were gracile with elongate, slender limbs. This suggests that the limb morphology of Dongusuchus efremovi may be plesiomorphic for Archosauria and proximal clades.

Doswellia sixmilensis is a new species of the dos-welliid archosauromorph genus Doswellia named for an incomplete skeleton from the Upper Triassic Bluewater Creek Formation of the Chinle Group in west-central New Mexico, USA. D. sixmilensis differs from D. kaltenbachi Weems, the type and only other known species of Doswellia, in its larger size, higher tooth count and greater heterodonty, possession of keels on the cervical centra and the presence of discrete knobs or spikes on some osteoderms. The holotype of D. six-milensis is the fourth occurrence of Doswellia and only the second occurrence of a Doswellia skull, which includes the previously unknown premaxilla and maxilla (and therefore the best dentition) and has the best-preserved cervical vertebrae. Although it adds to our knowledge of the anatomy of Doswellia, this new information does not alter previous concepts of the phylogenetic relationships of the doswelliid gen-era, largely because they are so poorly known anatomically. The genus Doswellia is known from the Newark Supergroup in Virginia, and the Chinle Group in Texas, New Mexico and Utah, in strata of Otischalkian–Adamanian age. The type locality of D. sixmilensis is c. 43 m stratigraphically below a bed from which U-Pb dating of detrital zircons yields a maximum depositional age of c. 220 Ma, so this is a reasonable approximate numerical age for D. sixmilensis.

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 Upper Triassic Tiki Formation of India is a rich storehouse of varied vertebrate fossil assemblages. So far, there is no information on taphonomic signatures of the Tiki vertebrate assemblages in comparison to that of other fossiliferous horizons of India. We report a new, low diversity, mono-dominant, multitaxic vertebrate accumulation where rhynchosaurs constitute the dominant component. The formation of the rhynchosaur bonebed is attributed to biological aggregation with a hydraulic overprint resulting in a mixed-origin concentration. Other taxa include rauisuchid, phytosaur, small indeterminate archosaur and cynodonts. Taphonomic study based on 617 skeletal specimens of rhynchosaurs collected from nine sites within an area of about 250 × 217 m 2 shows that most of the specimens are disarticulated and disassociated but in close spatial proximity to one another; some are associated specimens and few are articulated. About 13 to 20 individuals of rhynchosaur at different ontogenetic stages are estimated from the specimens collected, suggesting gregarious behavior, possibly herding. These specimens show varying degree of weathering, breakage, encrustation, abrasion and deformation. The bonebed is preserved within the Tiki red mudstone unit and is found in association with paleosol profiles, suggesting prolonged subaerial exposure. Spatial distribution and relative bone frequencies show differential susceptibility of the skeletal specimens to fluvial transport. 55.4% of the collected skeletal specimens belonged to Voorhies Group I, whereas 12.4% and 24% constituted Voorhies Groups II and III respectively, and 8.2% of the collected specimens belonged to the intermediate Groups I and II. It appears that the animals concentrated in the vicinity of the water sources during prolonged period of aridity and died possibly during high seasonal rainfall that resulted in a major flood event. Subsequently, the soft tissues decomposed, and the skeletons suffered prolonged subaerial exposure when the water receded leading to disarticulation and fragmentation followed by minor dispersion by low velocity water currents. This resulted in segregation of skeletal specimens, which were gradually covered by mud deposited during later flooding events. Based on the known flora and fauna, the Tiki Upper Triassic ecosystem is reconstructed for the first time. In the aquatic ecosystem, the metoposaurid labyrinthodonts occupied the top of the food pyramid together with the semi-aquatic parasuchids, which occupied an ecological niche similar to that of the present day crocodilians. The abundance of herbivorous rhynchosaurs at the base with few large and carnivorous rauisuchids and parasuchids at the top suggest a trophic structure similar to that of a modern day terrestrial ecosystem.

A new reptile, Kyrgyzsaurus bukhanchenkoi gen. et sp. nov., from the Triassic (Madygen Formation) of southwestern Kyrgyzstan is described based on the anterior part of the skeleton (skull, cervical and anterior dorsal vertebrae, ribs, pectoral girdle) and skin imprints. This is the most archaic representative of the family Drepanosauridae (Archosauromorpha, Diapsida). The most prominent features of the new form are the shortened lower jaw, numerous teeth, granular body osteoderms, large supraorbital shelflike skin folds, and thick and extensive throat sac.

During the end-Permian mass extinction, marine ecosystems suffered a major drop in diversity, which was maintained throughout the Early Triassic until delayed recovery during the Middle Triassic. This depressed diversity in the Early Triassic correlates with multiple major perturbations to the global carbon cycle, interpreted as either intrinsic ecosystem or external palaeoenvironmental effects. In contrast, the terrestrial record of extinction and recovery is less clear; the effects and magnitude of the end-Permian extinction on non-marine vertebrates are particularly controversial. We use specimen-level data from southern Africa and Russia to investigate the palaeodiversity dynamics of non-marine tetrapods across the Permo-Triassic boundary by analysing sample-standardized generic richness, evenness and relative abundance. In addition, we investigate the potential effects of sampling, geological and taxonomic biases on these data. Our analyses demonstrate that non-marine tetrapods were severely affected by the end-Permian mass extinction, and that these assemblages did not begin to recover until the Middle Triassic. These data are congruent with those from land plants and marine invertebrates. Furthermore, they are consistent with the idea that unstable low-diversity post-extinction ecosystems were subject to boom–bust cycles, reflected in multiple Early Triassic perturbations of the carbon cycle.

Until now the Doswelliidae was considered a monospecific family including Doswellia kaltenbachi from the Late Triassic of North America. The phylogenetic position of this taxon remained enigmatic until recently, when a sister-group relationship with the Proterochampsidae was suggested. In the present contribution we describe the new doswelliid species Archeopelta arborensis gen. et sp. nov. from the Middle–Late Triassic of Brazil. A cladistic analysis recovered Archeopelta, Doswellia, and Tarjadia within a monophyletic group of basal archosauriforms, the Doswelliidae. The monophyly of this family is supported by the presence of osteoderm ornamentation that is coarse, incised, and composed of regular pits and the presence of an unornamented anterior articular lamina. Archeopelta is more closely related to Doswellia than to other archosauriforms by the presence of basipterygoid processes anterolaterally orientated, dorsal centra with a convex surface, width of the neural arch plus ribs of the first primordial sacral that are three times the length of the neural arch, and iliac blade laterally deflected, with strongly convex dorsal margin, and a length less than three times its height. The phylogenetic analysis indicates that Doswellidae is the closest large monophyletic entity to Archosauria, which achieved a wide palaeolatitudinal distribution during the late Middle and Late Triassic time span. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161, 839–871.

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.

Detailed description of the cranial anatomy of the rhynchosaur previously known as Scaphonyx sulcognathus allows its assignment to a new genus Teyumbaita. Two nearly complete skulls and a partial skull have been referred to the taxon, all of which come from the lower part of the Caturrita Formation, Upper Triassic of Rio Grande do Sul, southern Brazil. Cranial autapomorphies of Teyumbaita sulcognathus include anterior margin of nasal concave at midline, prefrontal separated from the ascending process of the maxilla, palatal ramus of pterygoid expanded laterally within palatines, dorsal surface of exoccipital markedly depressed, a single tooth lingually displaced from the main medial tooth-bearing area of the maxilla, and a number of other characters (such as skull broader than long; a protruding orbital anterior margin; anguli oris extending to anterior ramus of the jugal; bar between the orbit and the lower temporal fenestra wider than 0·4 of the total orbital opening; mandibular depth reaching more than 25 of the total length) support its inclusion in Hyperodapedontinae. T. sulcognathus is the only potential Norian rhynchosaur, suggesting that the group survived the end-Carnian extinction event.

The best-known South American Early Triassic archosauriform belongs to a putative proterosuchid briefly reported by Jose Bonaparte in 1981, collected from the Quebrada de los Fosiles Formation (Puesto Viejo Group, Argentina). This specimen consists of well-preserved natural external molds of a partial postcranium that preserve dorsal vertebrae, osteoderms, a dorsal rib, a possible gastralium, a chevron, a humerus, an ilium, two metapodials, and an ungual. We re-describe this specimen and identify autapomorphies that allow us to recognize Koilamasuchus gonzalezdiazi, gen. et sp. nov. The presence of an iliac blade with a slightly convex dorsal margin and with a maximum length more than 3 times its maximum height places Koilamasuchus within Archosauriformes. A cladistic analysis of basal Archosauriformes positions Koilamasuchus more crownwards than Proterosuchus, Sarmatosuchus, Fugusuchus, and Osmolskina, as the sister taxon of the clade that includes Erythrosuchidae and Archosauria. Proterosuchida...

Here, we describe a new species of Azendohsaurus from the Middle–Late Triassic of Madagascar, extending the geographical range of a taxon known otherwise only by a single species from Morocco. Although Azendohsaurus has consistently been regarded as an early dinosaur (based on various advanced dental and gnathic features resembling those characterizing certain dinosaur subgroups), the relatively complete skeletal material, now available from Madagascar, argues strongly against its dinosaurian affinities. Rather, the retention of numerous primitive cranial and postcranial features indicates a surprisingly early divergence of Azendohsaurus within Archosauromorpha and an unusual mosaic of characters in this taxon. Features considered diagnostic of Sauropodomorpha thus are inferred to occur homoplastically in at least one clade of nondinosaurian archosauromorphs, indicating a complex evolution and distribution of features traditionally thought to be derived within archosaurs. Azendohsaurus has teeth resembling those of both early sauropodomorph and ornithischian dinosaurs, yet also possesses numerous inarguable basal archosauromorph cranial and postcranial attributes. This highlights the risk of uncritically referring isolated, Middle–Late Triassic (or even later), ‘leaf-shaped’ teeth with denticles to the Dinosauria. Similarly, the occurrence of such teeth in an early diverging archosauromorph indicates that specializations for herbivory originated more frequently within this clade than conventionally assumed. For example, Azendohsaurus and numerous basal sauropodomorph dinosaur taxa share an array of convergently acquired features associated with herbivory, including tooth denticles, expanded tooth crowns, a downturned dentary and the articular located at the ventral margin of the mandible. Some of these features (denticles, expanded crowns and the ventrally deflected articular) are even more widespread among archosauromorphs, including aetosaurs, silesaurs and ornithischian dinosaurs. A downturned dentary also occurs in Trilophosaurus, a taxon further marked by unique specializations for herbivory, including transversely lophate, tricuspid teeth. An array of features associated with herbivory also occurs in rhynchosaurs and certain crocodilians (e.g. Simosuchus). This distribution suggests that craniodental features associated with herbivory were much more pervasive across the archosauromorph clade than previously recognized, possibly evolving at least six to eight times independently.

Flexible, or soft-shelled, eggs are almost unknown in the fossil record, leaving large gaps in our knowledge of the reproductive biology of many tetrapod clades. Here, we report two flexible-shelled eggs of the hyphalosaurid choristodere Hyphalosaurus baitaigouensis from the Early Cretaceous of China, one containing an embryo and the second associated with a neonate. Choristoderes are an enigmatic group of aquatic reptiles that survived the K–T extinction but died out in the Miocene. Hyphalosaurids, a specialized clade of Choristodera, resemble miniature plesiosaurs and are considered to be primarily aquatic in habit. Scanning electron microscopy of samples from the eggs reveals a thin, non-columnar external mineralized layer characterized by rounded nodes and tentatively identified poorly structured irregular pores, with an underlying amorphous layer presumably representing decomposed protein fibrils. While the relationships of Choristodera remain controversial, eggshell microstructure more closely resembles that of Lepidosauromorpha (the lineage including lizards) as opposed to that of Archosauromorpha (the lineage including birds and crocodiles).

Vancleavea campi Long & Murry, 1995, from the Late Triassic of western North America, represents the latest surviving non-archosaurian archosauriform known to date. We present here a detailed comparative description based on a nearly complete, articulated skeleton from the Coelophysis Quarry in north-central New Mexico and other fragmentary specimens. The unique combination of morphological features of Vancleavea is unparalleled within Reptilia; it has four unique morphologies of imbricated osteoderms covering the entire body, a short, highly ossified skull, relatively small limbs and morphological features consistent with a semi-aquatic lifestyle. Vancleavea is placed in a rigorous phylogenetic analysis examining the relationships of non-archosaurian archosauriforms, and is found to be more closely related to Archosauria than both Erythrosuchus and Proterosuchus, but outside of the crown group. The analysis confirms previously hypothesized relationships, which found Euparkeria to be the closest sister taxon of Archosauria. It is not clear whether specimens referred to Vancleavea campi represent a single species-level taxon or a clade of closely related taxa that lived through much of the Late Triassic of North America, given the poor fossil record of the taxon.

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.

A detailed redescription of the Late Triassic archosauromorph reptile Doswellia kaltenbachi Weems, 1980 from the Poor Farm Member of the Falling Creek Formation in the Taylorsville basin (Newark Supergroup) in Virginia is presented based upon additional preparation of the holotype. The euryapsid skull has a distinctive occiput with a prominent supraoccipital process that is flanked by posterior “horn-like” projections of the squamosals. Postfrontals, tabulars, and postparietals are absent. Plesiomorphic features of the palate and braincase include a plate-like horizontal parabasisphenoid, a pair of foramina for the internal carotid arteries on the ventral surface of the basisphenoid, and two fields of teeth on the palatal surface of the pterygoid. A sharp angle along the cervical and anterior dorsal ribs clearly separates the dorsal and lateral sides of the neck and anterior thoracic region. The posterior thoracic region has shorter ribs that project laterally with only a slight curvature. The il...

Reptiles (sauropsids) represent the sister group to mammals, and the basal members of Reptilia may provide a good model for the condition of the common ancestor of both groups. Sex-determining mechanisms (SDM) and organizations of sex chromosomes among genotypically sex-determining (GSD) species vary widely across reptiles. Birds and snakes, for example, are entirely GSD whereas other reptiles, like all crocodilians, exhibit temperature-dependent sex determination (TSD). Here we explore the evolution of sex chromosomes and SDM within reptiles, using family-level analyses of character evolution and applying parsimony, likelihood, Bayesian, and stochastic methods. We find support for the common ancestor of amphisbaenians and whiptail lizards (Laterata) possessing the XY (male heterogametic) GSD mechanism, while the ancestors of Testudines and Crocodylia, as well as the larger group Archosauromorpha (here containing turtles) are inferred to have exhibited TSD. We also find evidence consistent with the hypothesis that the XY system is more labile and evolves faster than does the ZW (female heterogametic) system. Phylogenetic-based speciation tests do not support an association between GSD and speciation, and reject the hypothesis that the presence of the XY system is associated with speciation in reptiles.

We present a description of new cranial and postcranial material representing a new genus of rhynchosaur (Diapsida, Archosauromorpha) from the Otter Sandstone Formation (Mid Triassic) of Devon, south-west England. The taxon had been named Rhynchosaurus spenceriBenton, 1990, but cladistic analysis of the clade, and one autapomorphy, show that it does not belong to Rhynchosaurus, and a new generic name is required. We propose the name Fodonyx for this genus. A cladistic analysis of the Rhynchosauria confirms the main discoveries of previous analyses, and that Fodonyx is sister group to the Hyperodapedontinae, the clade of Late Triassic rhynchosaurs. The new cladistic analysis, for which many more characters were coded for Fodonyx than before (a rise from 39 to 75 per cent), counter-intuitively produced less well-resolved results: the new codings of previously uncoded characters introduced conflict so that Fodonyx turns out to be less like the Late Triassic rhynchosaur clade than had been assumed before.

A new tetrapod taxon from the Upper Triassic Cow Branch Formation of Virginia is described solely on the basis of computed tomography (CT) scans of 2 individuals. The new form is characterized by the presence of extremely elongate thoracolumbar ribs that presumably supported a gliding membrane in life. It differs from all other known gliding tetrapods in possessing a very pronounced elongate neck. The grasping hindfoot is consistent with an arboreal habit. A gliding habit has been reported in a handful of fossil reptiles, with the oldest occurrence in Coelurosauravus (Carroll, 1978; Evans, 1982: Evans and Haubold, 1987) from the Permian of Europe and Africa. Elongate ribs were originally described in this form and these were thought to have supported a gliding membrane in life, but it has since been shown that the membrane-supporting structures are not true ribs, but separate bundles of rodlike neomorph ossifications (Frey et al., 1997). However, 3 closely related forms (Icarosaurus, Kuehneosaurus and Kuehneosuchus) from the Upper Triassic of Europe and North America do have exceptionally elongate thoracolumbar ribs, and all have been referred to a single family, the Kuehneosauridae (Robinson, 1962; Colbert, 1970). They are further characterized by the ribs forming hinge joints with the markedly elongate transverse processes on the dorsal vertebrae. This contrasts with the living gliding agamid Draco (Colbert, 1970), in which the elongate thoracolumbar ribs are flexible and lack the hinge-joint with the vertebral transverse processes. A fourth Triassic tetrapod, Sharovipteryx from the Triassic of Kirghizia, also possessed a gliding membrane (Gans et al., 1987) but in this form it is stretched between the hind limbs. Here we describe a new Triassic tetrapod with elongate ribs that is comparable in size to the contemporaneous Icarosaurus, but differs significantly from kuehneosaurs in having a long neck, a character that is potentially very unstable in a gliding animal. The 2 specimens of the new tetrapod were recovered from

The Agua de la Peña Group of the Ischigualasto-Villa Unión Basin (northwestern Argentina) documents the evolution of archosauromorph assemblages in western Gondwana during the late Middle and Late Triassic. However, the South American archosauromorph record in the aftermath of the Permo/Triassic mass extinction (Early–early Middle Triassic) is remarkably scarce and restricted to isolated bones. Here, we describe a recently collected isolated second sacral vertebra and rib that represents one of the few fossils known from the Lower–Middle Triassic Tarjados Formation, the unit that underlies the Agua de la Peña Group. This specimen is identified as an archosauromorph because of the presence of a non-notochordal vertebra and a bifurcated distal end of the second sacral rib. A quantitative phylogenetic analysis places the new specimen as an archosauromorph more derived than protorosaurs. In particular, this specimen resembles Pamelaria, Prolacerta and early rhynchosaurs in the presence of a squared posterior projection of the bifurcated second sacral rib. The new specimen represents the first body fossil of a diapsid formally described in the Tarjados Formation and the oldest member of the group in the Ischigualasto-Villa Unión Basin. As a result, this specimen increases the high-level taxonomic richness of Archosauromorpha in South America in the aftermath of the Permo/ Triassic mass extinction. El Grupo Agua de la Peña de la Cuenca de Ischigualasto-Villa Unión (noroeste de Argentina) documenta la evolución de las faunas de arcosauromorfos en el oeste de Gondwana durante el Triásico Medio tardío y Triásico Tardío. Sin embargo, el registro de arcosauromorfos sudamericanos luego de la extinción masiva permo/triásica (Triásico Temprano–Medio temprano) es particularmente escaso y restringido a huesos aislados. Describimos aquí una vértebra y costilla sacra colectada recientemente, que representa uno de los pocos fósiles conocidos de la Formación Tarjados (Triásico Inferior–Medio), unidad que subyace al grupo Agua de la Peña. Este espécimen es identificado como un arcosauromorfo debido a la presencia de una vertebra no notocordal y un extremo distal birfurcado de la segunda costilla sacra. Un análisis filogenético cuantitativo ubica al nuevo espécimen como un arcosauromorfo más derivado que los protorosaurios. En particular, este ejemplar se asemeja a Pamelaria, Prolacerta y rincosaurios basales en la presencia de una proyección posterior cuadrangular del extremo distal de la segunda costilla sacra. El nuevo espécimen representa el primer registro fósil corpóreo formalmente descripto de un diápsido en la Formación Tarjados y el más antiguo para el grupo en la Cuenca de Ischigualasto-Villa Unión. Como resultado, este ejemplar incrementa la riqueza taxonómica de Archosauromorpha en América del Sur inmediatamente después de la extinción masiva Permo/Triásica. Fil: Ezcurra, Martin Daniel. University of Birmingham. School of Geography, Earth and Environmental Sciences; Reino Unido. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Martinelli, Agustín G.. Universidade Federal Do Rio Grande Do Sul; Brasil Fil: Fiorelli, Lucas Ernesto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Regional de Investigaciones Cientificas y Transferencia Tecnológica de Anillaco; Argentina Fil: Da Rosa, Atila A. S.. Universidade Federal de Santa Maria; Brasil Fil: Desojo, Julia Brenda. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales; Argentina

An unusual new reptile, Teraterpeton hrynewichorum, is described based on a partial skeleton. The holotype comes from the Upper Triassic Wolfville Formation (Newark Supergroup) of Nova Scotia. The temporal region of the skull is of the euryapsid type. The premaxillary rostrum and dentary symphysis are elongate and edentulous. The dentition comprises distinctive, occluding upper and lower teeth. A phylogenetic analysis shows that Teraterpeton hrynewichorum is an archosauromorph reptile. Trilophosaurus buettneri from the Upper Triassic Dockum Group of Texas may represent a closely related species.

A new genus and species Pamelaria dolichotrachela, belonging to the family Prolacertidae is reported for the first time from the Middle Triassic of India. The material consisting of associated skeletons and isolated bones has been excavated from the red mudstone of Yerrapalli Formation of Pranhita–Godavari Basin, South India. The nature of disposition of bones strongly suggests a catastrophic death and quick burial. The reconstructed skeleton of P. dolichotrachela strongly suggests that it was a quadrupedal sprawler. A detailed description of the new taxon and its comparison with other better known prolacertids of the world reveals a very close resemblance with Prolacerta broomi [Ann. Mag. Nat. History 16 (1935) 197], the type genus and species of the family. A number of characters like a relatively large skeleton; confluent nostril, lower jaw with a prominent coronoid and accessory neural spines in the posteriorcaudals with a sudden shift of neural spine position establish its distinctiveness.

Vertebrate genomes are mosaics of isochores. On the assumption that marked differences exist in the isochore structure between warm-blooded and cold-blooded animals, variations among vertebrates were previously attributed to adaptation to homeothermy. However, based on the data of coding regions from representatives of extant vertebrates, including a turtle, a crocodile (Archosauromorpha) and a few kinds of snakes (Lepidosauromorpha), it was recently hypothesized that the common ancestors of mammals, birds and extant reptiles already had the "warm-blooded" isochore structure. To test this hypothesis, the nucleotide sequences of alpha-globin genes including non-coding regions (introns) from two snakes, N. kaouthia and E. climacophora, were determined (accession number: AB104824, AB104825). The correlation between the GC contents in the introns and exons of alpha-globin genes from snakes and those from other vertebrates supports the above hypothesis. Similar analysis using data for exons and introns of other genes obtained from the GenBank (Release 131) also support the above hypothesis.

A former clay quarry near Schoningen in Lower Saxony exposes deposits dating from the upper half of the Ladinian. There are several bone beds in this quarry, which differ partly in their faunal composition and in the preservation of the fossils. One of these bone beds contains many morphologically different teeth of mainly terrestrial tetrapods and a variety of remains of actinopterygians, chondrichthyans and dipnoans. The tetrapod teeth are described in this paper. A precise taxonomic determination is not possible, but the material appears to contain two temnospondyls, a synapsid and several species of archosaurs. Some tooth morphotypes can not be assigned, even at a high taxonomic level. The tooth assemblage, which is described in this paper, is briefly compared with published and unpublished data on teeth from the Middle and Upper Keuper of Central Europe.

Codon usages are known to vary among vertebrates chiefly due to variations in isochore structure. Under the assumption that marked differences exist in isochore structure between warm-blooded and cold-blooded animals, the variations among vertebrates were previously attributed to an adaptation to homeothermy. However, based on data from a turtle species and a crocodile (Archosauromorpha), it was recently proposed that the common ancestors of mammals, birds and extent reptiles already had the "warm-blooded" isochore structure. We determined the nucleotide sequences of alpha-globin genes from two species of heterotherms, cuckoo (Cuculus canorus) and bat (Pipistrellus abramus), and three species of snakes (Lepidosauromorpha), Naja kaouthia from a tropical terrestrial habitat, Elaphe climacophora from a temperate terrestrial habitat, and Hydrophis melanocephalus from a tropical marine habitat. Our purposes were to assess the influence of differential body temperature patterns on codon usage and GC content at the third position of a codon (GC3), and to test the hypothesis concerning the phylogenetic position at which GC contents had increased in vertebrates. The results of principal component analysis (PCA) using the present data and data for other taxa from GenBank indicate that the primary difference in codon usage in globin genes among amniotes and other vertebrates lies in GC3. The codon usages (and GC3) in alpha-globin genes from two heterotherms and three snakes are similar to those in alpha-globin genes from warm-blooded vertebrates. These results refute the influence of body temperature pattern upon codon usages (and GC3) in alpha-globin genes, and support the hypothesis that the increase in GC content in the genome occurred in the common ancestor of amniotes.

The Permo-Triassic archosauromorph record is crucial to understand the impact of the Permo-Triassic mass extinction on the early evolution of the group and its subsequent dominance in Mesozoic terrestrial ecosystems. However, the Permo-Triassic archosauromorph record is still very poor in most continents and hampers the identification of global macroevolutionary patterns. Here we describe cranial and postcranial bones from the Permo-Triassic Buena Vista Formation of northeastern Uruguay that contribute to increase the meagre early archosauromorph record from South America. A basioccipital fused to both partial exoccipitals and three cervical vertebrae are assigned to Archosauromorpha based on apomorphies or a unique combination of characters. The archosauromorph remains of the Buena Vista Formation probably represent a multi-taxonomic assemblage composed of non-archosauriform archosauromorphs and a ‘proterosuchid-grade’ animal. This assemblage does not contribute in the discussion of a Late Permian or Early Triassic age for the Buena Vista Formation, but reinforces the broad palaeobiogeographic distribution of ‘proterosuchid grade’ diapsids in Permo-Triassic beds worldwide.

The first osteohistological study focused exclusively on rhynchosaurs (non-archosauriform archosauromorphs), based on the hyperodapedontines Teyumbaita sulcognathus and Hyperodapedon sp., from the Upper Triassic of Southern Brazil, indicates a relatively rapid growth rate in early ontogeny shown by the fibrolamellar complex, with a change to slow intermittent growth during late ontogeny represented by parallel-fibred bone with several growth marks. Contrary to previous studies, which described a typical non-archosaur reptilian bone tissue pattern for rhynchosaurs, with growth marks extending across the entire cortex, we demonstrate that, in both studied taxa, the initial growth rate was faster in comparison to the later. This suggests that the ability of rapid growth at high rates was already present in basal non-archosauriform archosauromorphs.

Sauria is the crown-group of Diapsida and is subdivided into Lepidosauromorpha and Archosauromorpha, comprising a high percentage of the diversity of living and fossil tetrapods. The split between lepidosauromorphs and archosauromorphs (the crocodile-lizard, or bird-lizard, divergence) is considered one of the key calibration points for molecular analyses of tetrapod phylogeny. Saurians have a very rich Mesozoic and Cenozoic fossil record, but their late Paleozoic (Permian) record is problematic. Several Permian specimens have been referred to Sauria, but the phylogenetic affinity of some of these records remains questionable. We reexamine and review all of these specimens here, providing new data on early saurian evolution including osteohistology, and present a new morphological phylogenetic dataset. We support previous studies that find that no valid Permian record for Lepidosauromorpha, and we also reject some of the previous referrals of Permian specimens to Archosauromorpha. The most informative Permian archosauromorph is Protorosaurus speneri from the middle Late Permian of Western Europe. A historically problematic specimen from the Late Permian of Tanzania is redescribed and reidentified as a new genus and species of basal archosauromorph: Aenigmastropheus parringtoni. The supposed protorosaur Eorasaurus olsoni from the Late Permian of Russia is recovered among Archosauriformes and may be the oldest known member of the group but the phylogenetic support for this position is low. The assignment of Archosaurus rossicus from the latest Permian of Russia to the archosauromorph clade Proterosuchidae is supported. Our revision suggests a minimum fossil calibration date for the crocodile-lizard split of 254.7 Ma. The occurrences of basal archosauromorphs in the northern (30°N) and southern (55°S) parts of Pangea imply a wider paleobiogeographic distribution for the group during the Late Permian than previously appreciated. Early archosauromorph growth strategies appear to be more diverse than previously suggested based on new data on the osteohistology of Aenigmastropheus.

A new rhynchosaur, Hyperodapedon huenei sp. nov., is described from the Upper Triassic Santa Maria Formation of the Parana Basin, Brazil. The holotype is an almost complete skull and mandible, collected at Inhamanda, in the state of Rio Grande do Sul, Brazil. The diagnosis of the genus Hyperodapedon Huxley is revised to include not only H. huxleyi Lydekker and H. gordoni Huxley (as generally accepted), but also the new species described here, various specimens usually assigned to ‘Scaphonyx fischeri’ Woodward, and ‘S’. sanjuanensis Sill. H. huenei sp. nov. exhibits a number of plesiomorphic features and appears to be the least derived species of Hyperodapedon, forming a sister taxon to the remaining members of the genus. A preliminary phylogenetic analysis for the more derived rhynchosaurs is presented. ‘Scaphonyx’sulcognathus Azevedo and Schultz represents the sister taxon of Hyperodapedon, while ‘Rhynchosaurus’spenceri Benton is considered to be a more derived Middle Triassic rhynchosaur. key words: Rhynchosauria, Hyperodapedon, Triassic, Brazil.

New specimens of the rhynchosaur Zsalorhynchus genovefae Buffetaut, 1983, have been found in the Makay Formation (Late Triassic of the Morondava Basin), Malio River area, southwest Madagascar. The material includes several maxillae as well as the first known rhynchosaur dentaries from the region. The maxilla bears a single longitudinal groove, with fewer than three longitudinal tooth rows both medial and lateral to it. The dentary possesses a single blade, with an additional row of teeth on its lingual surface. A phylagenetic study, based on the morphological features possible to measure for Zsalorhynchus, as well as on previous cladistic analysis of Rhynchosauria, showed that the Malagasian rhynchosaur is nested within the genus Hyperodapedon, and a new combination - H. genovefae - is proposed. The phyloge- netic study also depicts clear trends in the tooth evolution of Late Triassic rhynchosaurs, with the reduction of the medial structures in relation to the lateral ones, in both maxilla and dentaty. 0 2000 Editions scientitiques et medicales Elsevier SAS

New choristoderan fossils from the Late Cretaceous and Palaeocene of Alberta and Saskatchewan, Canada, are described: incomplete maxillae and dentaries from the Upper Cretaceous Oldman Formation, Alberta, extend the range of the primitive Cteniogenys from the Jurassic in the North American Western Interior; an incomplete dentary from the Palaeocene Ravenscrag Formation, Saskatchewan, comprises the first occurrence of the crocodile-like Simoedosaurus in Canada and the earliest record of the genus; well-preserved skulls and mandibles from the Oldman and Horseshoe Canyon formations, Alberta, document a new species of Champsosaurus and clarify the status of previously known species of the genus. New information about Asian choristoderes supports a Tchoiria –( Ikechosaurus + Simoedosaurus ) relationship, contrary to previous work. Choristoderes share no convincing synapomorphies with either Lepidosauromorpha or Archosauromorpha, but occupy a more basal position within Diapsida, possibly as a sister-taxon with Neodiapsida (Younginiformes+Sauria).

Four different species belonging to three genera of South African proterosuchids have been described: Proterosuchus fergusi (Broom, 1903), Chasmatosaurus vanhoepeni (Haughton, 1924), Elaphrosuchus rubidgei (Broom, 1946), and Chasmatosaurus alexandri (Hoffman, 1965). The specimens and descriptions suggest that all four are subjective synonyms for the same concept. The type specimen is specifically determinable and the senior synonym, Proterosuchus fergusi Broom, 1903, is selected as the valid name for the South African proterosuchid species. The diagnosis for the species includes features such as: the dentigerous borders of the premaxilla and maxilla form an angle of about 120 degrees; the palatal teeth are arranged in a specific pattern; the laterosphenoid is not reduced in its anterior development; and the epipterygoid is axe-shaped and has an anteroventral medial process.

A new diapsid reptile, Jesairosaurus lehmani, gen. et sp. nov., is described on the basis of cranial and postcranial remains from the base of the Zarzaitine Series (Triassic) of Algeria. Jesairosaurus is a small prolacertiform with a low skull and an elongated neck; the snout is narrow and elongated and the post-orbital region short and narrow; the quadratojugal is lost; the posterior process of the jugal is reduced and spur-shaped; the hind-limb is large in comparison with the forelimb and the rest of the postcranial skeleton. The relationships of basal archosauromorphs and the interrelationships of the prolacertiforms are examined. The analysis of 71 binary characters leads to the identification of Jesairosaurus lehmani as a prolacertiform and suggests that Prolacertiformes forms a monophyletic group within the Archosauromorpha. The analysis indicates that the Rhynchosauria is the sister group of a clade comprising Trilophosaurus, Prolacertiformes, and Proterosuchus. The Choristodera is the mos...

A detailed description of the early archosaur Sarmatosuchus otschevi Sennikov, 1994, from the Middle Triassic Donguz Formation of southern European Russia, is presented. New morphological data forms the basis of a preliminary investigation of the phylogenetic relationships between a selection of the earliest archosaurs. A parsimony analysis based on 9 taxa and 35 characters recovers two most parsimonious trees that support the referral of Sarmatosuchus and Fugusuchus to a proterosuchid clade, as well as the paraphyly of the Proterosuchia. Characters from the structure of the braincase are more homoplastic than those from the rest of the skeleton. The presence of Sarmatosuchus in the Middle Triassic affords evidence that the proterosuchids were a more diverse and persistent clade than has been previously envisaged. Evidence suggesting that proterosuchids may have existed with erythrosuchids and early crown-group archosaurs in the Middle Triassic requires a revision of our understanding of the earl...

Rhynchosaurs first appear in the Early Triassic fossil record and flourished during the late Carnian as the dominant members of several assemblages worldwide. In Argentina, the rhynchosaur record is currently restricted to the Ischigualasto Formation of late Carnian–earliest Norian age. Recent fieldwork in the new locality of Brazo del Puma, in the lowermost levels of the Chañares Formation, yielded three rhynchosaur tooth-bearing bones, which were collected five metres above the contact with the underlying Tarjados Formation. The most complete specimen is the posterior end of the alveolar region of a left dentary. The dentary possesses densely packed tooth rows on the lingual surface and medial half of the occlusal surface, showing longitudinal Zahnreihen. The teeth of the occlusal surface are worn flat and those of the lingual surface are organized in multiple rows, supporting the referral of the specimen to Rhynchosauridae. In addition, the dentary teeth are conical to mesiodistally compressed, resembling the condition observed in hyperodapedontines. The rhynchosaur remains reported here are the oldest collected in Argentina and among the oldest in South America, together with an unnamed form from Brazil. The new rhynchosaur specimens come from levels in which dicynodonts are numerically dominant, whereas cynodonts are considerably less abundant. Accordingly, the specimens reported here bolster faunal differences within the Chañares Formation and add a new faunal component to this already diverse vertebrate assemblage. Fil: Ezcurra, Martin Daniel. University of Birmingham; Reino Unido. Universität München; Alemania Fil: Trotteyn, Maria Jimena. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Bahía Blanca. Instituto Geológico del Sur; Argentina Fil: Fiorelli, Lucas Ernesto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Regional de Investigaciones Cientificas y Transferencia Tecnológica de Anillaco; Argentina Fil: Von Baczko, Belen. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales; Argentina Fil: Taborda, Jeremías Ramón Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales; Argentina Fil: Iberlucea, Maximiliano Javier. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales; Argentina Fil: Desojo, Julia Brenda. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales; Argentina

Rhynchosaurs were a group of archosauromorphs that dominated the guild of herbivores during the early Late Triassic. Despite the large number of specimens available, paleobiological studies are rare in the literature, especially concerning the South American species. The present study analyzes the paleoneurology of Teyumbaita sulcognathus, a Brazilian hyperodapedontine rhynchosaur, along with its nasal cavity, based on tomographic images of the specimen UFRGS-PV-0232-T. Although the endocast only reveals the morphology of the posterior half of the encephalon due to the incompletely ossified braincase, it is possible to infer the presence of great olfactory bulbs because of their impressions left on the ventral surface of the frontals. Although the snout is relatively short, the areas of the nasal cavity probably devoted to olfaction were also large and, along with the size of the olfactory bulbs, it is possible to infer that olfaction was important for the behavior and ecology of T. sulcognathus, as previously proposed for Hyperodapedon. Marcos A.F. Sales. Departamento de Paleontologia e Estratigrafia, Instituto de Geociencias, Universidade Federal do Rio Grande do Sul, Av. Bento Goncalves, 9500, Agronomia, CEP 91501-970, Porto Alegre, Rio Grande do Sul, Brazil. marcos.paleo@yahoo.com Cesar L. Schultz. Departamento de Paleontologia e Estratigrafia, Instituto de Geociencias, Universidade Federal do Rio Grande do Sul, Av. Bento Goncalves, 9500, Agronomia, CEP 91501-970, Porto Alegre, Rio Grande do Sul, Brazil. cesar.schultz@ufrgs.br

Teyumbaita sulcognathus is a peculiar endemic Brazilian rhynchosaur that remained somewhat obscure until recently, when its skull anatomy was described and a new generic name was assigned to this highly autapomorphic taxon. Here, the postcranial skeleton of Teyumbaita sulcognathus is for the first time fully described based on the holotype and the two more complete referred specimens. Rhynchosaur postcranial anatomy has usually been considered to be rather conservative, but T. sulcognathus shows unforeseen morphological variation. Autapomorphic traits were added to the diagnosis of T. sulcognathus and intraspecific variation was also identified. In addition, six new phylogeneticaly informative postcranial characters were recognized. Some of these represent apomorphies of clades such as Rhynchosauridae (axis with ventral keel, crest on the anteromedial surface of tibial shaft) and Hyperodapedontinae (postaxial cervical vertebrae with ventral keel, supinator process composed of a low supinator ridg...

In recent years the hypothesis that pterosaurs were the major sister-group of dinosaurs and a closely-linked hypothesis that pterosaurs evolved flight from the ground up have gained general acceptance. A cladistic analysis of the Archosauromorpha using characters presented by previous workers results in a single most parsimonious tree with the Pterosauria as the major sister-group of the Dinosauria. However, that sister-group relationship is supported only by a suite of hindlimb characters that are correlated with bipedal digitigrade locomotion in dinosaurs. In pterosaurs the characters have been interpreted as correlates of bipedal cursorial locomotion, arboreal leaping, or involvement of the hindlimb in the wing. The homology of those characters in dinosaurs and pterosaurs cannot be supported. Reanalysis of the data after exclusion of those hindlimb characters results in most parsimonious trees with the Pterosauria as the sister-group of the Erythrosuchidae + Proterochampsidae +Euparkeria+ Archosauria, in that order. This sister-group relationship is supported by a diverse assemblage of functionally independent skeletal characters from all regions of the skeleton. The results of the analysis cast doubt on the hypothesis that pterosaurs evolved flight from the ground up.

The morphology of the erythrosuchid ankle joint is reassessed. Two specimens, recently thought to have been incorrectly referred toErythrosuchus africanus, are shown without doubt to belong to this taxon. Furthermore, the morphology is essentially similar to that of other early archosaurs. The tarsus ofErythrosuchusis poorly ossified and consists of a calcaneum, astragalus, and two distal tarsals. The calcanea ofErythrosuchus, Vjushkovia triplicostata, andShansisuchus shansisuchusare all similar in being dorsoventrally compressed, possessing a lateral tuber, and lacking a perforating foramen. The astragalus ofV. triplicostatais currently unknown. The astragalus ofShansisuchusis apparently unique in form. The erythrosuchid pes is therefore more derived than has been recently proposed. The tarsal morphology of several other archosauromorph taxa is reviewed and many details are found to be at variance with the literature. The plesiomorphic condition for the Archosauromorpha consists of four distal tarsals and a proximal row of three elements; two of which articulate with the tibia. These proximal elements are interpreted as the astragalus, calcaneum, and a centrale, and the same pattern is retained in the earliest archosaurs. This reassessed tarsal morphology has implications for the homology of the centrale and reconstruction of early diapsid phylogeny.

A reinvestigation of the osteology of the holotype of Drepanosaurus unguicaudatus Pinna, 1980 suggests that in earlier descriptions some osteological features were misinterpreted, owing to the crushing of the bones and because taphonomic aspects were not considered. The pattern of the shoulder girdle and fore-limb was misunderstood: the supposed interclavicle is in fact the right scapula, and the bones previously identified as coracoid and scapula belong to the anterior limb. The new reconstruction of the shoulder girdle, along with the morphology of the phalanges and caudal vertebrae, leads to a new hypothesis about the mode of life of this reptile. Drepanosaurus was probably an arboreal reptile which used its enormous claws to scrape the bark from trees, perhaps in search of insects, just as the modern pigmy anteater (Cyclopes) does. Available diagnostic characters place Drepanosaurus within the Neodiapsida Benton, but it is impossible to ascribe this genus to one or other of the two major neodiapsid lineages, the Archosauromorpha and the Lepidosauromorpha.

BackgroundArchosaurs (birds, crocodilians and their extinct relatives including dinosaurs) dominated Mesozoic continental ecosystems from the Late Triassic onwards, and still form a major component of modern ecosystems (>10,000 species). The earliest diverse archosaur faunal assemblages are known from the Middle Triassic (c. 244 Ma), implying that the archosaur radiation began in the Early Triassic (252.3-247.2 Ma). Understanding of this radiation is currently limited by the poor early fossil record of the group in terms of skeletal remains.Methodology/principal findingsWe redescribe the anatomy and stratigraphic position of the type specimen of Ctenosauriscus koeneni (Huene), a sail-backed reptile from the Early Triassic (late Olenekian) Solling Formation of northern Germany that potentially represents the oldest known archosaur. We critically discuss previous biomechanical work on the 'sail' of Ctenosauriscus, which is formed by a series of elongated neural spines. In addition, we describe Ctenosauriscus-like postcranial material from the earliest Middle Triassic (early Anisian) Röt Formation of Waldhaus, southwestern Germany. Finally, we review the spatial and temporal distribution of the earliest archosaur fossils and their implications for understanding the dynamics of the archosaur radiation.Conclusions/significanceComprehensive numerical phylogenetic analyses demonstrate that both Ctenosauriscus and the Waldhaus taxon are members of a monophyletic grouping of poposauroid archosaurs, Ctenosauriscidae, characterised by greatly elongated neural spines in the posterior cervical to anterior caudal vertebrae. The earliest archosaurs, including Ctenosauriscus, appear in the body fossil record just prior to the Olenekian/Anisian boundary (c. 248 Ma), less than 5 million years after the Permian-Triassic mass extinction. These earliest archosaur assemblages are dominated by ctenosauriscids, which were broadly distributed across northern Pangea and which appear to have been the first global radiation of archosaurs.

Archosaurs have a nearly 250 million year record that originated shortly after the Permian-Triassic extinction event and is continued today by two extant clades, the crocodylians and the avians. The two extant lineages exemplify two bauplan extremes among a diverse and complex evolutionary history, but little is known about the common ancestor of these lineages. Renewed interest in early archosaurs has led to nearly a doubling of the known taxa in the last 20 years. This study presents a thorough phylogenetic analysis of 80 species-level taxa ranging from the latest Permian to the early part of the Jurassic using a dataset of 412 characters. Each terminal taxon is explicitly described and all specimens used in the analysis are clearly stated. Additionally, each character is discussed in detail and nearly all of the character states are illustrated in either a drawing or highlighted on a specimen photograph. A combination of novel characters and comprehensive character sampling has bridged previou...

The Crocodylotarsi are a group erected by Benton and Clark (1988) for archosaurs that share the “crocodile-normal” ankle structure. In this study, the phylogeny of the Crocodylotarsi was re-examined based on study of most relevant fossil material of the early non–crocodyliform members of the clade. Relationships among the major archosaurian taxa (Ornithodira, Ornithosuchidae, and Crocodylotarsi) and their proximal outgroups (Proterochampsidae, Euparkeria) were also considered. The monophyly of the Archosauria, Crurotarsi, and Crocodylotarsi is supported by the current analysis. The Parasuchia are the most plesiomorphic clade within the Crocodylotarsi. The Suchia comprise the crocodylotarsans excluding the phytosaurs. The taxon Prestosuchidae is placed near the base of this clade, as are Lewisuchus and Turfanosuchus dabanensis. The new taxon Rauisuchiformes includes the last common ancestor of Aetosauria and Crocodylia and its descendants. Rauisuchia (new combination) includes Rauisuchidae, Gracil...