Tetrapod Fauna Research Articles

An Ornithopod-Dominated Tracksite from the Lower Cretaceous Jiaguan Formation (Barremian-Albian) of Qijiang, South-Central China: New Discoveries, Ichnotaxonomy, Preservation and Palaeoecology.

The historically-famous Lotus Fortress site, a deep 1.5–3.0-meter-high, 200-meter-long horizonal notch high up in near-vertical sandstone cliffs comprising the Cretaceous Jiaguan Formation, has been known since the 13th Century as an impregnable defensive position. The site is also extraordinary for having multiple tetrapod track-bearing levels, of which the lower two form the floor of part of the notch, and yield very well preserved asseamblages of ornithopod, bird (avian theropod) and pterosaur tracks. Trackway counts indicate that ornithopods dominate (69%) accounting for at least 165 trackmakers, followed by bird (18%), sauropod (10%), and pterosaur (3%). Previous studies designated Lotus Fortress as the type locality of Caririchnium lotus and Wupus agilis both of which are recognized here as valid ichnotaxa. On the basis of multiple parallel trackways both are interpreted as representing the trackways of gregarious species. C. lotus is redescribed here in detail and interpreted to indicate two age cohorts representing subadults that were sometimes bipedal and larger quadrupedal adults. Two other previously described dinosaurian ichnospecies, are here reinterpreted as underprints and considered nomina dubia. Like a growing number of significant tetrapod tracksites in China the Lotus Fortress site reveals new information about the composition of tetrapod faunas from formations in which the skeletal record is sparse. In particular, the site shows the relatively high abundance of Caririchium in a region where saurischian ichnofaunas are often dominant. It is also the only site known to have yielded Wupus agilis. In combination with information from other tracksites from the Jiaguan formation and other Cretaceous formations in the region, the track record is proving increasingly impotant as a major source of information on the vertebrate faunas of the region. The Lotus Fortress site has been developed as a spectacular, geologically-, paleontologically- and a culturally-significant destination within Qijiang National Geological Park.

A diverse Late Triassic tetrapod fauna from Manzanera (Teruel, Spain)

Several outcrops with vertebrate tetrapod remains have been located in the Keuper levels (Carnian, Upper Triassic) of the Spanish municipality of Manzanera (Teruel Province). The Manzanera remains include ribs and vertebrae of the sauropterygian nothosauroid Simosaurus, isolated osteoderms of cyamodontoid placodonts, partial vertebral centra of ichthyosaurs, and fragmentary dermal bones of temnospondyls. Therefore, a relatively high diversity is recognized in the Late Triassic of this region. The Simosaurus identification may represent the only occurrence of this genus in the Iberian Peninsula; it is one of the youngest in the fossil record. Several morphotypes of cyamodontoid placodont osteoderms are identified. The ichthyosaur elements from Manzanera represent the first record of this clade in the Iberian Keuper, constituting the second reference to this group in the Iberian Triassic after those from the Ladinian of Catalonia.

Sedimentology and vertebrate taphonomy of the Moradi Formation of northern Niger: A Permian wet desert in the tropics of Pangaea

Pangaean paleogeographic models place the Tim Mersoi basin of northern Niger in a 5000-km-wide corridor between Gondwana and Laurasia approximately 15 degrees south of the paleoequator. Late Permian paleoclimate models position this basin between tropical summer-wet to the north and desert to the south. Recent investigations of the fossil vertebrates and paleosols in the late Permian (Lopingian) Moradi Formation confirm that the climate was warm and hyperarid with highly seasonal monsoonal rainfall. Possibly as a result of these unusual “wet desert” conditions, the tetrapod fauna shows a high degree of endemism. This study tests existing paleoclimate models by providing additional data on sedimentary environments and vertebrate taphonomic processes. The Moradi red bed sequences accumulated in a gently subsiding sag basin to the west of the tectonically active Massif de l'Aïr. Low angle gravelly alluvial fans prograded westward from the massif and at times impinged on a large stable northward flowing meandering channel system. The interchannel mudrock sequences are over-thickened by the accumulation of loessic silts and preserve isolated skull and post crania of amphibians (Nigerpeton and Saharastega) as well as semi-articulated captorhinids (Moradisaurus). Detailed surface mapping of a fossil-rich exposure revealed an anastomosed network of loess-filled distributary channels incised into the floodplain mudrocks. This provided a locus for the accumulation and rapid burial of at least 15 associated skeletons of the pareiasaurian Bunostegos. Semi-permanent ponds are evidenced by patches of fissile red mudstone containing rare bivalves and spiral coprolites. In the distal floodplains away from the main river channels, the combination of a generally high groundwater table, warm mean annual temperatures, and deflation of fines from the floodplain surface promoted the formation of gypsiferous paleosols and end-point playa lakes. Carbonate-rich mud accumulated around the lake margins and provided ideal conditions for the imprinting and preservation of tetrapod trackways.

A Diverse Tetrapod Fauna at the Base of 'Romer's Gap'

The lack of fossil tetrapod bearing deposits in the earliest Carboniferous (‘Romer’s Gap’) has provoked some recent discussions regarding the proximal cause, with three explanations being offered: environmental, taphonomic, and collection failure. One of the few, and earliest, windows into this time is the locality of Blue Beach exposed in the Tournaisian deposits at Horton Bluff lying along the Avon River near Hantsport, Nova Scotia, Canada. This locality has long been known but, because the fossils were deposited in high energy settings they are almost always disarticulated, so the fauna has not been described in detail. Recent intensive collection has revealed a diverse assemblage of material, including for the first time associated elements, which permits an evaluation of the faunal constituents at the locality. Although not diagnosable to a fine taxonomic level, sufficient apomorphies are present to identify representatives from numerous clades known from more complete specimens elsewhere. The evidence suggests a diverse fauna was present, including whatcheeriids and embolomeres. A single humerus previously had been attributed to a colosteid, but there is some uncertainty with this identification. Additional elements suggest the presence of taxa otherwise only known from the late Devonian. Depositional biases at the locality favor tetrapod fossils from larger individuals, but indirect evidence from trackways and tantalizing isolated bones evidences the presence of small taxa that remain to be discovered. The fossils from Blue Beach demonstrate that when windows into the fauna of ‘Romer’s Gap’ are found a rich diversity of tetrapods will be shown to be present, contra arguments that suggested this hiatus in the fossil record was due to extrinsic factors such as atmospheric oxygen levels. They also show that the early tetrapod fauna is not easily divisible into Devonian and Carboniferous faunas, suggesting that some tetrapods passed through the end Devonian extinction event unaffected.

Youngest dinocephalian fossils extend the Tapinocephalus Zone, Karoo Basin, South Africa

The dinocephalians (Synapsida, Therapsida) were one of the dominant tetrapod groups of the Middle Permian (Guadalupian Epoch, ~270–260 million years ago) and are most abundantly recorded in the Tapinocephalus Assemblage Zone (AZ) of the Main Karoo Basin, South Africa. Dinocephalians are thought to have become extinct near the top of the Abrahamskraal Formation of the Beaufort Group and their disappearance is one criterion used to define the base of the overlying Pristerognathus AZ. Because of the abundance of fossils in the Karoo, the Beaufort Group biozones form the biostratigraphic standard for later Permian terrestrial tetrapod ecosystems, so their stratigraphic delineation is of great importance to Permian palaeobiology. We report two new specimens of the rare tapinocephalid dinocephalian Criocephalosaurus from the lowermost Poortjie Member, which makes them the youngest dinocephalians known from the Main Karoo Basin and extends the Tapinocephalus AZ from the Abrahamskraal Formation up into the Teekloof Formation. The extension of the Tapinocephalus AZ relative to the lithostratigraphy potentially affects the biozone or biozones to which a fossil species can be attributed; this extension has implications for biostratigraphic correlations within the Main Karoo Basin as well as with other basins across Gondwana. These discoveries also indicate that a population of herbivorous tapinocephalids survived as rare constituents of the tetrapod fauna after most generic richness within the clade had already been lost.

Isochirotherium trackways, their possible trackmakers (?Arizonasaurus): intercontinental giant archosaur migrations in the Middle Triassic tsunami-influenced carbonate intertidal mud flats of the European Germanic Basin

A remarkable Middle Triassic (Pelsonian, Anisian) fossil track site at Bernburg in Central Germany permits reconstruction of a tetrapod fauna at the point of recovery when early poposaur quadruped dinosaurs diversificated globally. The site has yielded numerous tetrapod and thousands of arthropod horseshoecrab tracks from several levels in intertidal biolaminites with intercalated storm and seismic-influenced carbonates. The tetrapod tracks, Procolophonichnium, Rhynchosauroides, Chirotherium, and Isochirotherium, are assigned to smaller Archosauromorpha and large Archosauria. The largest chirotheroid tracks Isochirotherium herculis (Egerton in Proc Geol Soc London 3:14–15, 1838) reach up to 350-mm-long pes sizes and 120-mm-long manus imprints. Those were most probably made by a 5-m-long poposauroid archosaur like Arizonasaurus, based on matching skeleton anatomy and trackways including a new three-dimensional model of those giants in locomotion. A first German poposauroid bone record is added herein with a scapula from shallow marine carbonates of similar Pelsonian ages. These crurotarsan archosaurs were at the top of their food chains globally, and fed most probably on smaller tetrapods (Hescheleria, Macrocnemus), the latter also represented well by long behavioural escaping Rhynchosauroides trackways. The smaller reptiles may have fed especially on “Millions” of horseshoecrab eggs in those arthropod reproduction seasons in the intertidal carbonate mud flats. Such tidal flats and sabkhas extended over hundreds of square kilometres surrounding the Germanic Basin in Central Europe in the Middle Triassic, representing a drastic change in the palaeoenvironment of central Pangaea, and a possible trigger for continuing diversification of archosauromorphs and eventually dinosaurs. The horseshoecrab reproduction seasons seem to have caused a food chain reaction along the southern Pangaean (northern Tethys) coast in carbonate mud flats between Northern America, Africa, Europe and China, with possibly long-distance migrations of poposauroids such as Arizonasaurus, whose bones were found in terrestrial to shallow marine carbonate deposits in terrest to coastal context.

Late Cretaceous reptilian biota of the La Colonia Formation, central Patagonia, Argentina: Occurrences, preservation and paleoenvironments

Cropping out on the southeastern margin of the Somún Curá Plateau, the La Colonia Formation (Campanian–Maastrichtian) has yielded, over the last several decades, a varied fossil tetrapod fauna (mammals, ophidians, turtles, dinosaurs, plesiosaurs, anurans). For this study several field trips were carried out specifically to recover reptile remains. Three sections, located on the southeastern slope of the Sierra de La Colonia and in the vicinity of Cerro Bayo, Chubut Province (Argentina), were selected for particular attention. The prospected sections are composed of massive, laminated or heterolithic siltstones and claystones, with scarce and thin intercalations of massive, heterolithic o cross-bedded fine sandstones and of fossiliferous conglomerates. The most abundantly recovered reptiles are terrestrial and freshwater turtles, followed by plesiosaurs and dinosaurs. Among the chelonians, more than 16 specimens of Patagoniaemys gasparinae (Meiolaniformes), three specimens of Yaminuechelys aff. Y. gasparinii (Chelidae), and the remains of a new genus of Chelidae were identified. Among the dinosaurs, theropod metatarsal fragments, an incomplete abelisaurid theropod skeleton, sauropod vertebrae, ankylosaur osteoderms and appendicular fragments of hadrosaurs were found. Among the plesiosaurs there are several well preserved elasmosaurids (including two with associated gastroliths) and a polycotylid (Sulcusuchus erraini). Except for the plesiosaurs, all the reptiles are terrestrial or freshwater taxa. However, analysis of the elasmosaurids indicates they are adult specimens of small body size, which could be related to forms that lived in restricted aquatic environments. Likewise, the polycotylid possesses deep rostral and mandibular grooves, and a conspicuous vascularization and/or innervation, that is consistent with the presence of some associated special sensory structures similar to those known in some cetaceans that inhabit modern rivers and estuaries. Sedimentological analysis suggests that deposition would have been mostly in low-energy restricted environments, like muddy plains, marshes and ponds cut by meandering channels, probably in the central mixed-energy zone within an estuary. This interpretation is consistent with the habitat inferred for the recovered reptiles, as well as with associated foraminifers and with the probable origin of gastroliths found associate with the plesiosaurs.

Adult morphology of the therocephalian Simorhinella baini from the middle Permian of South Africa and the taxonomy, paleobiogeography, and temporal distribution of the Lycosuchidae

The Middle Permian tetrapod fauna of the South African Beaufort Group is taxonomically diverse and includes representatives of all major therapsid groups, including the earliest records of Eutheriodontia. In the Middle Permian, eutheriodonts are represented mainly by large therocephalians, which made up a large proportion of the vertebrate predators in these faunas. Here we describe the skull and partial skeleton of a large therocephalian from the uppermost Tapinocephalus Assemblage Zone (AZ) of South Africa. A combination of features, including the short snout, presence of three to four upper postcanines and presence of teeth on the pterygoid processes, indicates that the new specimen belongs to the earliest-diverging therocephalian family, Lycosuchidae. The presence of a well-developed midline ridge on the ventral surface of the vomer indicates that the new specimen can be referred to Simorhinella baini, a species previously represented only by a tiny juvenile skull. The new specimen forms the basis for a taxonomic re-evaluation of the Lycosuchidae as well as of the geographic and stratigraphic range of the family. We recognize two valid species within the Lycosuchidae: the type species Lycosuchus vanderrieti represented by five specimens and Simorhinella baini represented by two specimens, with an additional 22 specimens currently identifiable as Lycosuchidae incertae sedis. Lycosuchid specimens range throughout the Tapinocephalus and Pristerognathus AZs; specimens of Simorhinella are restricted to the Tapinocephalus AZ, whereas Lycosuchus specimens are documented in both the Tapinocephalus and Pristerognathus AZs.

New dicynodonts (Therapsida, Anomodontia) and updated tetrapod stratigraphy of the Permian Ruhuhu Formation (Songea Group, Ruhuhu Basin) of southern Tanzania

ABSTRACTPermian tetrapod fossils were discovered in the Tanzanian Ruhuhu Formation in 1963, but they have received far less attention than the tetrapods of the overlying Usili (formerly Kawinga) Formation. Here, we describe two dicynodonts collected in the Ruhuhu Formation in 2008. Abajudon kaayai, gen. et sp. nov., is represented by a partial skull and mandible and is characterized by autapomorphic upper teeth that are triangular in cross-section, have procurved tips, and bear a deep groove on the mesial surface. Although it shows similarities to taxa such as Endothiodon and Chelydontops, the exact relationships between A. kaayai and other dicynodonts are uncertain. The second specimen also consists of a partial skull and mandible. We refer it to cf. Endothiodontia based on the medial placement of the long maxillary tooth rows, the presence of depressions on the palatine pad, a long posterior dentary sulcus, and similarities of the mandibular dentition. Tetrapods occur in three fossiliferous horizons in the Ruhuhu Formation. The likely Middle Permian lower horizon includes dinocephalians, temnospondyls, and the dicynodonts described here. The middle horizon includes a new, tusked species of Endothiodon and at least one other dicynodont. The upper horizon appears to sample an assemblage similar to that of the Usili Formation and therefore may be of Late Permian age. The discovery of Middle Permian fossils in Tanzania and Zambia provides the opportunity to test whether southern Gondwana was characterized by a cosmopolitan tetrapod fauna for an extended period of time before the biogeographic restructuring caused by the end-Permian mass extinction.

A new Late Triassic age for the Puesto Viejo Group (San Rafael depocenter, Argentina): SHRIMP U–Pb zircon dating and biostratigraphic correlations across southern Gondwana

The Puesto Viejo Group crops out in the San Rafael Block, southwest Mendoza, at approximately 35° S and 68°20′ W. It consists of the basal mainly grayish Quebrada de los Fósiles Formation (QF) overlying by the reddish Río Seco de la Quebrada Formation (RSQ). The basal unit includes both plant remains (pleuromeians and sphenopsids) and vertebrates (scattered fish scales, dicynodont synapsids and remains of an archosauriform). In contrast, the RSQ beds have yielded only tetrapods, although a more diverse fauna. The latter includes cynodonts as Cynognathus, Pascualognathus and Diademodon, and also dicynodonts (Vinceria and Kannemeyeria). Based on the assemblage of tetrapod taxa the bearing levels were correlated to the Cynognathus AZ of South Africa and thus referred to the Middle Triassic (Anisian). We obtained a SHRIMP 238U/206Pb age of 235.8 ± 2.0 Ma from a rhyolitic ignimbrite interdigitated between the QF and RSQ formations at the Quebrada de los Fósiles section. This new radiometric date for the Puesto Viejo Group suggests that the tetrapod fauna in the RSQ beds existed, instead, during the Late Triassic (early Carnian) some 10 Ma later than the currently accepted age. Two scenarios might explain our results: first, the Cynognathus AZ of South Africa is wrongly assigned to the lower Middle Triassic (Anisan) and should be considered younger in age, Late Triassic (Carnian); second, the relative age of the Cynognathus AZ of South Africa is correct but the inferred range of Cynognathus and Diademodon is incorrect as they were present during the Late Triassic (Carnian) at least in South America. In any case, this new date pose serious doubts about the validity of biostratigraphic correlations based solely on tetrapod taxa, a common practice for Triassic continental successions across Gondwana.

Revised chronostratigraphy of the Lower Chinle Formation strata in Arizona and New Mexico (USA): High-precision U-Pb geochronological constraints on the Late Triassic evolution of dinosaurs

The early history of dinosaurs in North America is obscured by an incomplete fossil record, taxonomic uncertainties and speculative correlations of tetrapod-bearing rocks, as well as poor calibration of the Late Triassic time scale. High-precision U-Pb geochronology provides a reliable means of correlating terrestrial rock formations independent of equivocal lithostratigraphy or vertebrate biostratigraphy, and hence the possibility of properly evaluating models for the early radiation and diversification of Dinosauria. Here we present new, high-precision, U-Pb ID-TIMS zircon geochronology from the presumed lowermost strata of the Upper Triassic Chinle Formation of the Colorado Plateau in Southwest United States, including a mean 206Pb/238U date of 219.39 ± 0.16 Ma from the renowned Placerias Quarry Bone Bed in eastern Arizona. The new results prompt revisions to the chronostratigraphy of the lower Chinle and provide a new temporal context for its rich tetrapod fauna. The oldest documented dinosaurs of North America coexisted with their non-dinosaurian near-relatives for a minimum of 12 m.y., from ca. 223 Ma to ca. 211 Ma, in the Norian. This early dinosauromorph record follows a ca. 6 m.y. period from which no tetrapod fossils have been documented and which was itself preceded by a ca. 10 m.y. depositional hiatus spanning nearly the entire Ladinian and Carnian stages of the terrestrial North America. The supposed late appearance of dinosauromorphs in North America compared to those in South America thus appears to be an artifact of incomplete preservation, as well as unsubstantiated age interpretations. This, together with the conspicuous biogeographic distinctions among the Triassic dinosauromorph assemblages, invalidates a simple diachronous model for the transcontinental radiation of early dinosaurs.

The Jiyuan tetrapod Fauna of the Upper Permian of China——1. New pareiasaur materials and the reestablishment of Honania complicidentata

Honania complicidentata and Tsiyuania simplicidentata are pareiasaur taxa based on material from the Shangshihezi Formation of Jiyuan, Henan Province, China that were earlier designated as nomina vana. Based on the study of new material, and the reexamination of old specimens, we determine that the pareiasaur material from Jiyuan represents a single species that differs from all known species from other localities. Thus, we resurrect the name H. complicidentata for the material from Jiyuan. H. complicidentata is characterized by maxillary teeth with high crowns, dentary teeth slightly posteriorly inclined compared to the dentary dorsal margin, nearly all preserved marginal teeth have a cusped cingulum on the lingual surface, and humerus without an ectepicondylar foramen. Phylogenetic analysis shows Honania is more basal than Shansisaurus and Shihtienfenia from the Sunjiagou Formation of China.

Bringing dicynodonts back to life: paleobiology and anatomy of a new emydopoid genus from the Upper Permian of Mozambique.

Dicynodontia represent the most diverse tetrapod group during the Late Permian. They survived the Permo-Triassic extinction and are central to understanding Permo-Triassic terrestrial ecosystems. Although extensively studied, several aspects of dicynodont paleobiology such as, neuroanatomy, inner ear morphology and internal cranial anatomy remain obscure. Here we describe a new dicynodont (Therapsida, Anomodontia) from northern Mozambique: Niassodon mfumukasi gen. et sp. nov. The holotype ML1620 was collected from the Late Permian K5 formation, Metangula Graben, Niassa Province northern Mozambique, an almost completely unexplored basin and country for vertebrate paleontology. Synchrotron radiation based micro-computed tomography (SRµCT), combined with a phylogenetic analysis, demonstrates a set of characters shared with Emydopoidea. All individual bones were digitally segmented allowing a 3D visualization of each element. In addition, we reconstructed the osseous labyrinth, endocast, cranial nerves and vasculature. The brain is narrow and the cerebellum is broader than the forebrain, resembling the conservative, “reptilian-grade” morphology of other non-mammalian therapsids, but the enlarged paraflocculi occupy the same relative volume as in birds. The orientation of the horizontal semicircular canals indicates a slightly more dorsally tilted head posture than previously assumed in other dicynodonts. In addition, synchrotron data shows a secondary center of ossification in the femur. Thus ML1620 represents, to our knowledge, the oldest fossil evidence of a secondary center of ossification, pushing back the evolutionary origins of this feature. The fact that the specimen represents a new species indicates that the Late Permian tetrapod fauna of east Africa is still incompletely known.

The vertebrate fauna of the Upper Permian of Niger—VII. Cranial anatomy and relationships of Bunostegos akokanensis (Pareiasauria)

ABSTRACT We describe newly recovered cranial material of Bunostegos akokanensis, a pareiasaurian reptile known from the Upper Permian Moradi Formation of northern Niger. Bunostegos is highly autapomorphic, with diagnostic cranial features including two or three hemispherical bosses located above and between the external nares; laterally projecting supraorbital ‘horn’ formed by an enlarged postfrontal; large foramen present on ventral surface of postfrontal; and hemispherical supratemporal boss located at posterolateral corner of skull roof. We addressed the phylogenetic position of Bunostegos by incorporating it into a cladistic analysis of 29 parareptilian taxa (including all 21 currently valid pareiasaurs) and 127 cranial and postcranial characters. The results of this analysis place Bunostegos as more derived than middle Permian forms such as Bradysaurus and as the sister taxon to the clade including Deltavjatia plus Velosauria. Certain characters, such as the pattern of cranial ornamentation and the size and placement of the tabulars, appear to be more similar to more derived pareiasaurs such as Elginia from Scotland and Arganaceras from Morocco, but the most parsimonious tree topology indicates that these features were evolved independently in the Nigerien form. The lack of both dicynodont herbivores and Glossopteris, combined with the presence of a giant herbivorous captorhinid, indicates a markedly different community structure in the Permian of Niger compared with those for contemporaneous southern Pangean basins (i.e., Karoo, Luangwa, Ruhuhu). The endemic tetrapod fauna of Niger supports the theory that central Pangea was biogeographically isolated from the rest of the supercontinent by desert-like conditions during Late Permian times. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Provincialization of terrestrial faunas following the end-Permian mass extinction

In addition to their devastating effects on global biodiversity, mass extinctions have had a long-term influence on the history of life by eliminating dominant lineages that suppressed ecological change. Here, we test whether the end-Permian mass extinction (252.3 Ma) affected the distribution of tetrapod faunas within the southern hemisphere and apply quantitative methods to analyze four components of biogeographic structure: connectedness, clustering, range size, and endemism. For all four components, we detected increased provincialism between our Permian and Triassic datasets. In southern Pangea, a more homogeneous and broadly distributed fauna in the Late Permian (Wuchiapingian, ∼257 Ma) was replaced by a provincial and biogeographically fragmented fauna by Middle Triassic times (Anisian, ∼242 Ma). Importantly in the Triassic, lower latitude basins in Tanzania and Zambia included dinosaur predecessors and other archosaurs unknown elsewhere. The recognition of heterogeneous tetrapod communities in the Triassic implies that the end-Permian mass extinction afforded ecologically marginalized lineages the ecospace to diversify, and that biotic controls (i.e., evolutionary incumbency) were fundamentally reset. Archosaurs, which began diversifying in the Early Triassic, were likely beneficiaries of this ecological release and remained dominant for much of the later Mesozoic.

Imperfect isolation: factors and filters shaping Madagascar's extant vertebrate fauna.

Analyses of phylogenetic topology and estimates of divergence timing have facilitated a reconstruction of Madagascar’s colonization events by vertebrate animals, but that information alone does not reveal the major factors shaping the island’s biogeographic history. Here, we examine profiles of Malagasy vertebrate clades through time within the context of the island’s paleogeographical evolution to determine how particular events influenced the arrival of the island’s extant groups. First we compare vertebrate profiles on Madagascar before and after selected events; then we compare tetrapod profiles on Madagascar to contemporary tetrapod compositions globally. We show that changes from the Mesozoic to the Cenozoic in the proportions of Madagascar’s tetrapod clades (particularly its increase in the representation of birds and mammals) are tied to changes in their relative proportions elsewhere on the globe. Differences in the representation of vertebrate classes from the Mesozoic to the Cenozoic reflect the effects of extinction (i.e., the non-random susceptibility of the different vertebrate clades to purported catastrophic global events 65 million years ago), and new evolutionary opportunities for a subset of vertebrates with the relatively high potential for transoceanic dispersal potential. In comparison, changes in vertebrate class representation during the Cenozoic are minor. Despite the fact that the island’s isolation has resulted in high vertebrate endemism and a unique and taxonomically imbalanced extant vertebrate assemblage (both hailed as testimony to its long isolation), that isolation was never complete. Indeed, Madagascar’s extant tetrapod fauna owes more to colonization during the Cenozoic than to earlier arrivals. Madagascar’s unusual vertebrate assemblage needs to be understood with reference to the basal character of clades originating prior to the K-T extinction, as well as to the differential transoceanic dispersal advantage of other, more recently arriving clades. Thus, the composition of Madagascar’s endemic vertebrate assemblage itself provides evidence of the island's paleogeographic history.

On the presence of the Late Permian dicynodont Endothiodon in Brazil

AbstractThe first dicynodont reported for the Permian of South America is described in detail here. The specimen consists of a partial skull and associated lower jaws, collected from the Serra do Cadeado locality, Rio do Rasto Formation, Brazil. The specimen was assigned to the genus Endothiodon Owen in a preliminarily description published in the 1970s. This identification implied a direct correlation with some of the well‐established biozones of the Beaufort Group, South Africa, where this taxon is known to occur. Although badly weathered, the Brazilian material preserves several features that allow us to confirm its assignment to Endothiodon, such as the pineal foramen situated on a boss, the bulbous swellings of the dentaries, a boss situated on the ventral margin of the jugal, the extensive number of dentary teeth and the upturned pointed beak of the lower jaw. However, this set of characters is not enough to establish a specific taxonomic assignment for the specimen, although it seems to be more closely related to E. bathystoma, E. uniseries and E. whaitsi than to E. mahalanobisi. The biostratigraphical correlations proposed for the tetrapod faunas of the Rio do Rasto Formation highlight the similarities between these associations and others from the Mid and Late Permian of South Africa and Eastern Europe. However, until now, these correlations are tentative because only a few faunal elements have been recovered from Brazil, and most of them lack stratigraphic data on the levels from where they were collected. Due to the presence of Endothiodon, at least part of the Rio do Rasto Formation can be correlated with deposits in India, Malawi, Mozambique, South Africa, Tanzania, Zambia and Zimbabwe.

Dinosaur remains from the ‘Botucaraí Hill’ (Caturrita Formation), Late Triassic of south Brazil, and their stratigraphic context

Vertebrate fossils recovered from sites nearby the Botucaraí Hill and Candelária (Caturrita Formation) depict a diverse Late Triassic tetrapod fauna from south Brazil. These records are of key importance to the biostratigraphy of the upper sections of the Rosario do Sul Group. A lithological and biostratigraphic survey on the main fossil localities of the Botucaraí Hill area confirms the occurrence of the lower Hyperodapedon and the upper Riograndia Assemblage Zones in the region, the latter yielding early saurischians. In this paper, three incomplete dinosaur specimens, an isolated sacral vertebra, an articulated left pubis–ischium and an isolated right ischium, from the ‘Botucaraí Hill’ site are described. A comparative survey suggests that these specimens have sauropodomorph affinities, but probably more primitive than typical ‘prosauropods’ from the Norian-Early Jurassic. Regardless of the phylogenetic position of Guaibasaurus as theropod or sauropodomorph, their occurrence in the Caturrita Formation, which also yielded ‘core prosauropods’ from the Santa Maria region, suggests either the survival of early members of the clade with more derived ‘prosauropods’ or that heterochronous faunas are sampled from that stratigraphic unit.

Strengthening Western Gondwanan correlations: A Brazilian Dicynodont (Synapsida, Anomodontia) in the Middle Triassic of Namibia

Terrestrial Middle Triassic strata occur throughout continental Africa and are particularly well exposed in South Africa, Tanzania, Zambia, and Namibia. The youngest age for all these African deposits is widely accepted as early Middle Triassic (Anisian). Fossils collected recently from the uppermost strata of the upper Omingonde Formation in Namibia highlighted the presence of Chiniquodon, a carnivorous cynodont previously only found in Ladinian-Carnian aged rocks of South America. In addition, work in progress indicates that a large archosaur, originally reported as Erythrosuchus, also discovered from levels close to the top of this unit, is in fact a rauisuchian, a group of archosaurs well known from Ladinian-Carnian beds of southern South America. Here we present the first record of the tuskless dicynodont Stahleckeria potens from the top of the upper Omingonde Formation in central Namibia. This taxon was up until now only known from the Ladinian Dinodontosaurus Assemblage Zone of the Santa Maria Formation in southern Brazil. Thus, compelling evidence for a Ladinian age for the upper levels of the upper Omingonde Formation is provided by two therapsid and one archosaur taxa. The tetrapod fauna of the upper Omingonde Formation partially fills the gap of the well-documented hiatus (Ladinian gap), prevalent throughout the Karoo basins of south and central Africa. The presence of the same therapsid taxa in the Namibian Waterberg Basin and the Paraná Basin of Brazil during Middle Triassic suggests that these basins were biogeographically linked through a series of interconnecting lowlands, with no major ecological, climatic and/or physical barriers.

The South African stereospondyl Microposaurus from the Middle Triassic of the Sydney Basin, Australia

ABSTRACT The second member of the Trematosaurinae (Temnospondyli, Stereospondyli) from Australia is described as Microposaurus averyi, n. sp., from the Anisian Rouse Hill Siltstone (Ashfield Shale, Wianamatta Group, Sydney Basin). This is the first occurrence of the genus from outside South Africa. M. averyi and the South African M. casei share an anterior emargination of the nostril that accommodated the dentary fangs (a condition unique among temnospondyls), but the emargination is much larger in the Australian taxon. The Anisian stereospondyls from the Sydney Basin occur in sediments younger than the early Anisian and may fill a worldwide gap in terrestrial tetrapod faunas from that time.