Tetrapod Footprint Biostratigraphy and Biochronology

Tetrapod footprints have a fossil record in rocks of Devonian-Neogene age. Three principal factors limit their use in biostratigraphy and biochronology (palichnostratigraphy): invalid ichnotaxa based on extramorphological variants, slow apparent evolutionary turnover rates and facies restrictions. The ichnotaxonomy of tetrapod footprints has generally been oversplit, largely due to a failure to appreciate extramorphological variation. Thus, many tetrapod footprint ichnogenera and most ichnospecies are useless phantom taxa that confound biostratigraphic correlation and biochronological subdivision. Tracks rarely allow identification of a genus or species known from the body fossil record. Indeed, almost all tetrapod footprint ichnogenera are equivalent to a family or a higher taxon (order, superorder, etc.) based on body fossils. This means that ichnogenera necessarily have much longer temporal ranges and therefore slower apparent evolutionary turnover rates than do body fossil genera. Because of this, footprints cannot provide as refined a subdivision of geological time as do body fossils. The tetrapod footprint record is much more facies controlled than the tetrapod body fossil record. The relatively narrow facies window for track preservation, and the fact that tracks are almost never transported, redeposited or reworked, limits the facies that can be correlated with any track-based biostratigraphy.

A Devonian-Neogene global biochronology based on tetrapod footprints generally resolves geologic time about 20 to 50 percent as well as does the tetrapod body fossil record. The following globally recognizable time intervals can be based on the track record: (1) Late Devonian; (2) Mississippian; (3) Early-Middle Pennsylvanian; (4) Late Pennsylvanian; (5) Early Permian; (6) Late Permian; (7) Early-Middle Triassic; (8) late Middle Triassic; (9) Late Triassic; (10) Early Jurassic; (11) Middle-Late Jurassic; (12) Early Cretaceous; (13) Late Cretaceous; (14) Paleogene; (15) Neogene. Tetrapod footprints are most valuable in establishing biostratigraphic datum points, and this is their primary value to understanding the stratigraphic (temporal) dimension of tetrapod evolution.     

Late Smithian microbial deposits and their lateral marine fossiliferous limestones (Early Triassic,Hurricane Cliffs,Utah, USA)

Recurrent microbialite proliferations during the Early Triassic are usually explained by ecological relaxation and abnormal oceanic conditions. Most Early Triassic microbialites are described as single or multiple lithological units without detailed ecological information about lateral and coeval fossiliferous deposits. Exposed rocks along Workman Wash in the Hurricane Cliffs (southwestern Utah, USA) provide an opportunity to reconstruct the spatial relationships of late Smithian microbialites with adjacent and contemporaneous fossiliferous sediments. Microbialites deposited in an intertidal to subtidal interior platform are intercalated between inner tidal flat dolosiltstones and subtidal bioturbated fossiliferous limestones. Facies variations along these fossiliferous deposits and microbialites can be traced laterally over a few hundreds of meters. Preserved organisms reflect a moderately diversified assemblage, contemporaneous to the microbialite formation. The presence of such a fauna, including some stenohaline organisms (echinoderms), indicates that the development of these late Smithian microbial deposits occurred in normal-marine waters as a simple facies belt subject to relative sea-level changes. Based on this case study, the proliferation of microbialites cannot be considered as direct evidence for presumed harsh environmental conditions.     

Late Triassic (Late Norian) gastropods from the Wallowa Terrane (Idaho,USA)

A diverse Late Triassic (Late Norian) gastropod fauna is described from the Mission Creek Limestone of the Wallowa terrane (Idaho, USA). Sample standardization by rarefaction analysis indicates that the fauna is even more diverse than the Late Triassic gastropod fauna from the Pucara Formation (Peru) which represents the most diverse gastropod fauna from South America. The gastropod fauna consists of 66 species; several genera are reported for the first time from North America. A high percentage of the species are highly ornamented and several have distinct siphonal canals. This suggests that the appearance of truly Mesozoic elements among the gastropods began before the Mesozoic Marine Revolution in other clades. The fauna is dominated by high-spired strongly ornamented procerithiids, a group more characteristic for the Jurassic. Comparison of the present fauna and the Iranian Nayband Formation gastropod fauna show that the procerithiids underwent a first global radiation in the Late Triassic. The high number of new species in this fauna suggests that sampling of Late Triassic gastropod faunas is still incomplete and hinders palaeobiogeographic considerations. Previous suggesions that gastropod faunas from the Wallowa and Wrangellia terranes resemble each other and are distinct from those of Alexander, Chulitna, and Farewell terranes are basically corroborated. The gastropod fauna of the Mission Creek Limestone differs considerably from that of the western and central Tethys but shares several taxa with the Late Triassic gastropod fauna of the Pucara Formation in Peru. Thus, the Hispanic corridor was probably not present in the Norian but opened only in the Early Jurassic. The subfamily Andangulariinae is introduced and placed in the Zygopleuridae. The generaSpiniomphalus, Nodoconus, Gudrunella, Blodgettella, Idahospira, andSiphonilda and the subgenusCryptaulax (Wallowax) are introduced. 27 species are erected. A lectotype is designated forCryptaulax rhabdocolpoides Haas, 1953.      

Dispersal and diversity in the earliest North American sauropodomorph dinosaurs, with a description of a new taxon

Sauropodomorph dinosaurs originated in the Southern Hemisphere in the Middle or Late Triassic and are commonly portrayed as spreading rapidly to all corners of Pangaea as part of a uniform Late Triassic to Early Jurassic cosmopolitan dinosaur fauna. Under this model, dispersal allegedly inhibited dinosaurian diversification, while vicariance and local extinction enhanced it. However, apomorphy-based analyses of the known fossil record indicate that sauropodomorphs were absent in North America until the Early Jurassic, reframing the temporal context of their arrival. We describe a new taxon from the Kayenta Formation of Arizona that comprises the third diagnosable sauropodomorph from the Early Jurassic of North America. We analysed its relationships to test whether sauropodomorphs reached North America in a single sweepstakes event or in separate dispersals. Our finding of separate arrivals by all three taxa suggests dispersal as a chief factor in dinosaurian diversification during at least the early Mesozoic. It questions whether a 'cosmopolitan' dinosaur fauna ever existed, and corroborates that vicariance, extinction and dispersal did not operate uniformly in time or under uniform conditions during the Mesozoic. Their relative importance is best measured in narrow time slices and circumscribed geographical regions.     

Biogeography of Triassic tetrapods: evidence for provincialism and driven sympatric cladogenesis in the early evolution of modern tetrapod lineages

Triassic tetrapods are of key importance in understanding their evolutionary history, because several tetrapod clades, including most of their modern lineages, first appeared or experienced their initial evolutionary radiation during this Period. In order to test previous palaeobiogeographical hypotheses of Triassic tetrapod faunas, tree reconciliation analyses (TRA) were performed with the aim of recovering biogeographical patterns based on phylogenetic signals provided by a composite tree of Middle and Late Triassic tetrapods. The TRA found significant evidence for the presence of different palaeobiogeographical patterns during the analysed time spans. First, a Pangaean distribution is observed during the Middle Triassic, in which several cosmopolitan tetrapod groups are found. During the early Late Triassic a strongly palaeolatitudinally influenced pattern is recovered, with some tetrapod lineages restricted to palaeolatitudinal belts. During the latest Triassic, Gondwanan territories were more closely related to each other than to Laurasian ones, with a distinct tetrapod fauna at low palaeolatitudes. Finally, more than 75 per cent of the cladogenetic events recorded in the tetrapod phylogeny occurred as sympatric splits or within-area vicariance, indicating that evolutionary processes at the regional level were the main drivers in the radiation of Middle and Late Triassic tetrapods and the early evolution of several modern tetrapod lineages.     

Large-Diameter Burrows of the Triassic Ischigualasto Basin,NW Argentina: Paleoecological and Paleoenvironmental Implications

Large-diameter ichnofossils comprising three morphotypes have been identified in the Upper Triassic Ischigualasto and Los Colorados formations of northwestern Argentina. These burrows add to the global record of the early appearance of fossorial behavior during early Mesozoic time. Morphotypes 1 and 2 are characterized by a network of tunnels and shafts that can be assigned to tetrapod burrows given similarities with previously described forms. However, differences in diameter, overall morphology, and stratigraphic occurrence allow their independent classification. Morphotype 3 forms a complex network of straight branches that intersect at oblique angles. Their calcareous composition and surface morphology indicate these structures have a composite biogenic origin likely developed due to combined plant/animal interactions. The association of Morphotypes 1 and 2 with fluvial overbank lithologies deposited under an extremely seasonal arid climate confirms interpretations that the early appearance of burrowing behavior was employed by vertebrates in response to both temperature and moisture-stress associated with seasonally or perpetually dry Pangean paleoclimates. Comparisons of burrow morphology and biomechanical attributes of the abundant paleovertebrate fauna preserved in both formations permit interpretations regarding the possible burrow architects for Morphotypes 1 and 2. In the case of the Morphotype 1, the burrow constructor could be one of the small carnivorous cynodonts, Ecteninion or Probelesodon. Assigning an architect for Morphotype 2 is more problematic due to mismatches between the observed burrow morphology and the size of the known Los Colorados vertebrates.     

Early Triassic Archosaur-Dominated Footprint Assemblage from the Argana Basin (Western High Atlas,Morocco)

An assemblage of abundant and well-preserved tetrapod footprints has been discovered in the Tanamert Member (T3) of the Triassic Timezgadiouine Formation (Argana basin, western High Atlas, Morocco). It is the first fossil record from T3. Surfaces from different localities show a uniform tetrapod ichnofauna that consists of chirotherian and small lacertoid forms. The chirotherians are assigned to the plexus Protochirotherium—Synaptichnium, their trackmakers interpreted as basal archosaurs. The lacertoid imprints show close affinities with Rhynchosauroides and may reflect archosauromorphs or lepidosauromorphs. Protochirotherium—Synaptichnium assemblages are characteristic of the Early Triassic and were known previously only from units of this age in central Europe. Biostratigraphically, the European record implies a wide-spread pre-Anisian Protochirotherium—Synaptichnium dominated assemblage preceding the first appearance of Chirotherium barthii near the Olenekian-Anisian boundary. The stratigraphic position of T3 between Late Permian (uppermost T2) and Middle Triassic (T4) and the European correlatives suggest an Early Triassic age of this unit. It is the first record of Early Triassic continental deposits in Morocco. The surfaces from T3 open up perspectives for further contributions to ecology, biogeography and locomotion of early archosaurs. Furthermore, excellent outcrops and quality of footprint preservation in the Argana basin offer a potential for clarification of ichnotaxonomic and biostratigraphic issues.     

A multivariate taxonomic analysis of the Late Carboniferous vertebrate ichnofauna of Alveley, southern Shropshire, England

A diverse Late Carboniferous (Westphalian D; Moscovian) vertebrate ichnofaunal assemblage from the Alveley Member (Salop Formation, Warwickshire Group) of Alveley, southern Shropshire, UK, is a significant example of an early, transitional tetrapod community. Positive tetrapod footprint casts (convex hyporeliefs) within fine- and medium-grained sandstone red-beds were formed subaqueously in an alluvial floodplain setting. A statistical analysis of this material, the first using multivariate techniques on Late Palaeozoic trackways, has been undertaken to determine the ichnospecific diversity and morphological variation within the ichno-assemblage. Nine ichnotaxa have been identified, following a study of more than 200 trackways. These include the amniote ichnogenera Dimetropus Romer and Price, 1940, Ichniotherium Pohlig, 1885 and Hyloidichnus Gilmore, 1927, but there is a dominance of trackways of the ichnogenus Limnopus Marsh, 1894, which represent stem-lissamphibian 'temnospondyls'. Following statistical analysis, the ichnogenera Limnopus and Batrachichnus are subsumed as ichnosubgenera under the senior available name Limnopus . The Alveley ichnofaunal assemblage provides significant range extensions for a number of amniote and stem-lissamphibian trackmakers from the latest Carboniferous or Permian down into the mid-Late Carboniferous. It also marks a key transitional stage in the evolution of tetrapod communities, from the 'amphibian' grade assemblages of the Carboniferous to the more terrestrial, amniote-dominated communities of the Early Permian.     

The palaeoecology of the dinosaurs of the Wessex Formation (Wealden Group, Early Cretaceous), Isle of Wight, Southern England

The Wessex Formation on the Isle of Wight yields an Early Cretaceous dinosaur fauna. Sedimentological evidence shows that this represents a mosaic of fluvial, floodplain and lacustrine environments within a relatively narrow east-west oriented valley. The vegetational cover on the alluvial plain had a savannah-or chaparral-like aspect, probably of low productivity. The relative scarcity of small aquatic vertebrates, absence of coals, abundance of oxidixed sediments and the presence of immature calcretes indicate seasonal water supply. The dinosaur taxa compising the Wessex Formation faunal assemblage represent a single palaeocommunity which inhabited the local alluvial plain, although some species may have been transient. The fauna had a relatively low diversity and this is attributed to the low productivity of the local vegetation. Iguanodontids and Hypsilophodon were the dominant elements in the fauna. In contrast to Late Jurassic dinosaur faunas, sauropods are less abundant in the Wessex Formation, although they remain taxonomically diverse. It is concluded that climatic changes which took place in the Late Jurassic or Early Cretaceous resulted in the appearance of low productivity vegetation and that this was incapable of supporting large sauropod populations.     

A basal thunnosaurian from Iraq reveals disparate phylogenetic origins for Cretaceous ichthyosaurs

Cretaceous ichthyosaurs have typically been considered a small, homogeneous assemblage sharing a common Late Jurassic ancestor. Their low diversity and disparity have been interpreted as indicative of a decline leading to their Cenomanian extinction. We describe the first post-Triassic ichthyosaur from the Middle East, Malawania anachronus gen. et sp. nov. from the Early Cretaceous of Iraq, and re-evaluate the evolutionary history of parvipelvian ichthyosaurs via phylogenetic and cladogenesis rate analyses. Malawania represents a basal grade in thunnosaurian evolution that arose during a major Late Triassic radiation event and was previously thought to have gone extinct during the Early Jurassic. Its pectoral morphology appears surprisingly archaic, retaining a forefin architecture similar to that of its Early Jurassic relatives. After the initial latest Triassic radiation of early thunnosaurians, two subsequent large radiations produced lineages with Cretaceous representatives, but the radiation events themselves are pre-Cretaceous. Cretaceous ichthyosaurs therefore include distantly related lineages, with contrasting evolutionary histories, and appear more diverse and disparate than previously supposed.     

First glimpse into Lower Jurassic deep-sea biodiversity: in situ diversification and resilience against extinction

Owing to the assumed lack of deep-sea macrofossils older than the Late Cretaceous, very little is known about the geological history of deep-sea communities, and most inference-based hypotheses argue for repeated recolonizations of the deep sea from shelf habitats following major palaeoceanographic perturbations. We present a fossil deep-sea assemblage of echinoderms, gastropods, brachiopods and ostracods, from the Early Jurassic of the Glasenbach Gorge, Austria, which includes the oldest known representatives of a number of extant deep-sea groups, and thus implies that in situ diversification, in contrast to immigration from shelf habitats, played a much greater role in shaping modern deep-sea biodiversity than previously thought. A comparison with coeval shelf assemblages reveals that, at least in some of the analysed groups, significantly more extant families/superfamilies have endured in the deep sea since the Early Jurassic than in the shelf seas, which suggests that deep-sea biota are more resilient against extinction than shallow-water ones. In addition, a number of extant deep-sea families/superfamilies found in the Glasenbach assemblage lack post-Jurassic shelf occurrences, implying that if there was a complete extinction of the deep-sea fauna followed by replacement from the shelf, it must have happened before the Late Jurassic.     

贵州毕节麻窝口洞与智人伴生的长鼻类化石

最近在贵州毕节麻窝口洞发现了3枚古人类牙齿化石和伴生的哺乳动物群。其中,古人类牙齿经初步研究可归入解剖学上的现代人,而与古人类相伴的大、小哺乳动物化石,经初步鉴定共计8目20科43属53种。本文系统记述了该动物群中大哺乳动物的典型代表——长鼻类化石,共2属2种:东方剑齿象(Stegodon orientalis)和亚洲象(Elephas maximus)。麻窝口洞的长鼻类缺失我国南方早更新世的典型种类——中华乳齿象(Sinomastodon)和华南剑齿象(Stegodon huananensis),具有从典型的中更新世大熊猫-剑齿象动物群(Ailuropoda-Stegodon fauna)向晚更新世亚洲象动物群(Asian elephant fauna)过渡的特征。依动物群的性质和地貌地层的特征,毕节麻窝口洞的智人及伴生动物群的地质时代很可能为中更新世晚期或晚更新世早期,这与堆积物的光释光年代测定的初步结果(距今约11.2-17.8万年)基本吻合。麻窝口洞东方剑齿象与亚洲象的组合明显具有东洋界亚热带动物群的特点,指示温暖潮湿的气候,这些长鼻类与智人等生存于近水的森林和灌丛中,并镶嵌了一些草地。     

Evolution of ecospace occupancy by Mesozoic marine tetrapods

Ecology and morphology are different, and yet in comparative studies of fossil vertebrates the two are often conflated. The macroevolution of Mesozoic marine tetrapods has been explored in terms of morphological disparity, but less commonly using ecological‐functional categories. Here we use ecospace modelling to quantify ecological disparity across all Mesozoic marine tetrapods. We document the explosive radiation of marine tetrapod groups in the Triassic and their rapid attainment of high ecological disparity. Late Triassic extinctions led to a marked decline in ecological disparity, and the recovery of ecospace and ecological disparity was sluggish in the Early Jurassic. High levels of ecological disparity were again achieved by the Late Jurassic and maintained during the Cretaceous, when the ecospace became saturated by the Late Cretaceous. Sauropterygians, turtles and ichthyosauromorphs were the largest contributors to ecological disparity. Throughout the Mesozoic, we find that established groups remained ecologically conservative and did not explore occupied or vacant niches. Several parts of the ecospace remained vacant for long spans of time. Newly evolved, radiating taxa almost exclusively explored unoccupied ecospace, suggesting that abiotic releases are needed to empty niches and initiate diversification. In the balance of evolutionary drivers in Mesozoic marine tetrapods, abiotic factors were key to initiating diversification events, but biotic factors dominated the subsequent generation of ecological diversity.     

Early and Middle Jurassic Sporo-Pollen Assemblages of Qiquanhu Coal-Field of Turpan Xinjiang

One hundred and thirteen species of spores and pollens belonging to 58 genera were found from 66 core samples taken from two boreholes of the Early and Middle Jurassic deposits in the Qiquanhu coal-field of the Turpan Basin, Xinjiang have been investigated in this paper. Two of the species are described as new. Three sporo-pollen zones from lower to upper are described as follows: Sporo-pollen zone Ⅰ (equivalent bed: Badaowan Formation) . The pteridophytic spores are slightly more abundant than the gymnospermous pollen in this assemblage. The most abundant spores are Cyathidites minor, C. australis and Undulatisporites pflugii, U. taenus and so on. The second important fern spores are Osmundacidites, Lycopodiumsporites, Apiculatisporis and Duplexisporites. The most important gymnospermous genus is Cycadopites which ranks highest in the assemblage and includes C. nitidus and C. typicus.The other gymnospermous pollen include Classopollis, Pinuspollenites, Protoconiferus Piceites. Besides, there are a few pollen grains of Taeniaesporites and Chordasporites surviving from the Late Triassic in this assemblage. The present assemblage may be compared with the Early Jurassic sporopollen assemblage from the Daling Formation of the North-East Guangxi. Therefore, the Badaowan Formtion may be referred to Early Jurassic in age, probably Early Lias. Sporo-pollen zone Ⅱ (equivalent bed: Sangonghe Formation) In this assemblage the gymnosperms are predominant.Of them, Podocarpidites and Pinuspollenites increase obviously, and Quadraeculina and Protoconiferus are common. Few surviving elements are observed. The most abundant spores are those of Cyathidites, and the particularly abundant pollens are those of Cycadopites, This assemblage may be compared with the Late Lias sporo-pollen assemblage from the Fuxian Formation of the Northern Shangganning Basin. Therefore, the Sangonghe Formation should belong to Early Jurassic, probably Late Lias in age. Sporo-pollen zone Ⅲ (equivalent bed: Xishanyao Formation). The contents of the spores and pollens are almost equal in this assemblage. Cyathidites and Cycadopites are the most important spores and pollen. Osmundacidites and Quadraeculina are common. Neoraistrickia, Lycopodiumsporites and Eucommiidites troedssonii are present. Ancient striate bisaccate pollen disappears. The characteristics of this assemblage show some resemblances to those of the Middle Jurassic Yanan Formation in Chongxin county of Gansu province. So the Xishanyao Formation should belong to Middle Jurassic.     

Evolution of the Permian and Triassic tetrapod communities of Eastern Europe

The Permo-Triassic terrestrial and freshwater tetrapod communities of Eastern Europe are reconstructed as food-webs. The Late Permian theriodont-dinocephalian community (Ocher, Mezen, Isheyevo) changes to a latest Permian theriodont-pareiasaur community (North Dvina, Vyazniki). After a major extinction, the Triassic thecodontian-dicynodont communities appear, a lystrosaurid one in the Early Triassic (Lower and ?Upper Vetluga), and a kannemeyerid one in the later Early Triassic (?Yarenga) and the Mid Triassic (Donguz, Bukobay). Similar stages are represented in the evolution of aquatic communities: the Late Permian temnospondyl community (Ocher, Isheyevo), the latest Permian chroniosuchian one (North Dvina, Vyazniki), the Lower and Middle Triassic new temnospondyl one (from Vetluga to Bukobay). The faunal changes in Eastern Europe are mirrored in other parts of the world, although there are some endemic Russian forms.     

Small Theropod Track Assemblages from Middle Jurassic Eolianites of Eastern Utah: Paleoecological Insights from Dune Ichnofacies in a Transgressive Sequence

New discoveries show that very small theropod tracks (cf. Wildeichnus) are abundant in the upper part of the Moab Member, recently assigned to the Curtis Formation (formerly considered part of the Entrada Formation) in the Mid-?Late Jurassic of eastern Utah. The tracks represent a distinct small-theropod ichnofacies associated with eolian dune deposits that is easily differentiated from the water-lain beds of the overlying Megalosauripus-Therangospodus ichnofacies, which comprises the single-surface Moab megatracksite. Pterosaur track assemblages, representing the Pteriachnus ichnofacies, are found a few meters above the megatracksite surface in the upper tongue of the Summerville Formation.

The small theropod ichnofacies is reminiscent of other early Mesozoic dune facies ichnofaunas from the Wingate and Navajo formations (Late Triassic and Early Jurassic) where small theropod tracks occur in association with other small tetrapod footprints. All such examples evidently represent a recurrent dune facies ecosystem dominated by diminutive vertebrates. Because the small theropod ichnofacies is one of three ichnofacies found in a thin stratigraphic sequence (<20 m) that contains no body fossils, it is clear that vertebrate tracks play an important role in providing insight into the paleoecology of units previously considered devoid of any useful fossil evidence. The three successive ichnofacies represent a transgressive transition from sand dunes, through sandy shoreline to shallow marine environments, each with its quite distinct vertebrate fauna.     

Rhynchocephalians (Diapsida: Lepidosauria) from the Jurassic Kota Formation of India

Despite their rarity today, rhynchocephalians formed a diverse Early Mesozoic clade with a comparatively good fossil record. They had a Pangaean distribution in the Late Triassic and Early Jurassic, although the Gondwanan record remains more limited than the Laurasian one. We report here on new sphenodontian material from the Jurassic Kota Formation of peninsular India. Two taxa are represented, and both are attributed to new genera. One is a relatively derived sphenodontian with a premaxillary morphology similar to that of the Late Triassic/ Early Jurassic genus Clevosaurus. The other is somewhat more primitive in its morphology, although clearly a crown-group sphenodontian. In addition, three dentary fragments and a partial maxilla signal the presence of a primitive pleurodont lepidosauromorph similar to the basal rhynchocephalians Gephyrosaurus and Diphydonto-saurus from Britain.     

A REVIEW OF TRIASSIC TETRAPOD TRACK ASSEMBLAGES FROM ARGENTINA

Abstract:  This paper contains the first comprehensive ichnotaxonomic review of the Triassic tetrapod track record in Argentina, including previous accounts and new material recently discovered, and an analysis of its composition and stratigraphic distribution. Triassic footprints have been recorded from three basins: the Ischigualasto-Villa Unión and Cuyo basins in north-west Argentina, and the Los Menucos depocentre in northern Patagonia. Most are in successions of Middle Triassic age; a lower number are from the Late Triassic, and there are two records from Early Triassic rocks. The known track types include: Brachychirotherium isp., cf. Brachychirotherium isp., Chirotherium barthii , Dicynodontipus ispp., Grallator isp., Rhynchosauroides isp., Rigalites ischigualastianus , Rigalites isp., Tetrasauropus isp., and bird-like, chirotheriid and unidentified tridactyl footprints. The ichnogenera Gallegosichnus Casamiquela, 1964, Calibarichnus Casamiquela, 1964, Palaciosichnus Casamiquela, 1964 and Stipanicichnus Casamiquela, 1975 are considered to be synonyms of Dicynodontipus (Hornstein, 1876). In addition, the abandonment of the following ichnogenera (and single ichnospecies) that are based on poorly preserved material is suggested: Ingenierichnus sierrai Casamiquela, 1964, Rogerbaletichnus aguilerai Casamiquela, 1964 and Shimmelia chirotheroides Casamiquela, 1964. At least eight Triassic ichnofaunas can be recognized. The most peculiar is that of the Late Triassic Los Menucos depocentre, which is characterized by the dominance of therapsid footprints ( Dicynodontipus ispp.). The track assemblages from the Cuyo Basin display the highest ichnodiversity, with five footprint types.     

Arizonasaurus and its implications for archosaur divergence

The Moenkopi Formation has yielded partial and isolated remains of important archosaurs including rauisuchian skull fragments and isolated poposaur centra and pelvic girdle elements. A recently discovered skeleton referable to Arizonasaurus babbitti shows that most of these archosaurian remains belong to one taxon. Characteristics of the skeleton of Arizonasaurus show that it belongs to a poorly known group of Middle Triassic (240-230 Myr ago) archosaurs called the ctenosauriscids, and that ctenosauriscids are or are closely related to poposaurs. Furthermore, many characteristics of Arizonasaurus provide evidence that poposaurids and ctenosauriscids are derived rauisuchians. The presence of a poposaurid in the early Middle Triassic suggests that the divergence of birds and crocodiles occurred earlier than previously thought. Middle Triassic ctenosauriscids also allow the correlation of Triassic faunas in Europe, Asia, North America and Africa. The Moenkopi Formation fauna represents a transitional fauna between Early and Late Triassic faunas.     

Diversity variation and tempo-spatial distributions of the Dipteridaceae ferns in the Mesozoic of China

The extant family Dipteridaceae is a remarkable leptosporangiate fern because it includes only one genus with a restricted distribution to tropical regions. The fossil record of this family has been widely reported from the Mesozoic strata in Eurasia, America, Australia, and Greenland. In China, numerous fossils of the Dipteridaceae have been documented, in total, about 74 species of 6 genera. Geographically, they are distributed both in the Southern and Northern Floristic Provinces, and were particularly well developed in the Southern Floristic Province during the Late Triassic and the Early Jurassic intervals. Fossil diversity of Dipteridaceae varies in the different episodes of the Mesozoic in China. It is shown that Dipteridaceae has undergone a diversity development process and a distinct turnover during the Mesozoic. They appear to have diversified in the warm and humid Late Triassic–Early Jurassic, but declined sharply as aridity developed in the Middle Jurassic, and became extinct at the end of the Early Cretaceous. The diversity variation and tempo-spatial distribution pattern is suggested to be linked with paleoclimatic variations during the Mesozoic.