A late-surviving basal theropod dinosaur from the latest Triassic of North America

The oldest theropod dinosaurs are known from the Carnian of Argentina and Brazil. However, the evolutionary diversification of this group after its initial radiation but prior to the Triassic-Jurassic boundary is still poorly understood because of a sparse fossil record near that boundary. Here, we report on a new basal theropod, Daemonosaurus chauliodus gen. et sp. nov., from the latest Triassic 'siltstone member' of the Chinle Formation of the Coelophysis Quarry at Ghost Ranch, New Mexico. Based on a comprehensive phylogenetic analysis, Daemonosaurus is more closely related to coeval neotheropods (e.g. Coelophysis bauri) than to Herrerasauridae and Eoraptor. The skeletal structure of Daemonosaurus and the recently discovered Tawa bridge a morphological gap between Eoraptor and Herrerasauridae on one hand and neotheropods on the other, providing additional support for the theropod affinities of both Eoraptor and Herrerasauridae and demonstrating that lineages from the initial radiation of Dinosauria persisted until the end of the Triassic. Various features of the skull of Daemonosaurus, including the procumbent dentary and premaxillary teeth and greatly enlarged premaxillary and anterior maxillary teeth, clearly set this taxon apart from coeval neotheropods and demonstrate unexpected disparity in cranial shape among theropod dinosaurs just prior to the end of the Triassic.     

Palaeoenvironmental implications of the ichnology and geochemistry of the Westbury Formation (Rhaetian), Westbury‐on‐Severn,south‐west England

Abstract: The Westbury Formation (Rhaetian) beds of Westbury Garden Cliff, Westbury‐on‐Severn, west of Gloucester, Britain, show an unusual combination of features. Both deep water and emergent characteristics are present within the sediments and the trace fossils. The ichnoassemblage consists of abundant Selenichnites, Planolites beverlyensis and Lockeia with rarer Oniscoidichnus, Chondrites, Rhizocorallium irregulare, Taenidium serpentium, an unusual form of Walcottia and Merostomichnites‐like traces. These trace fossils display an interesting relationship with the sediments: low‐energy Cruziana ichnofacies is found within high‐energy sandstones. The sandstones are interbedded with laminated mudstones, apparently deposited in deep water, but some aspects of the ichnoassemblage, preservation and sedimentation indicate shallower water. One new trace fossil, Radichnus allingtona igen. et isp. nov., closely resembles the traces of modern fiddler crabs and imply emergence, by analogy. This ichnofauna is similar to early stage disaster colonisation in recent experiments in Long Island Sound (south of Connecticut, USA) and with storm‐influenced deposits within the Cardium Formation (Seebe, Alberta, Canada). This indicates a lagoonal environment with influxes of sand and oxygen. Total organic carbon levels were found to fluctuate greatly between stratigraphic layers but remained relatively high. This implies low oxygen conditions. The abundance of sulphur (in pyrite) also supports an interpretation of anoxic conditions, and low sedimentation rates within the shale layers. A restricted shallow basin or lagoonal environment is proposed for the palaeoenvironment, with fluctuating oxygen influencing diversity.     

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

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

Fossil moonseeds from the Paleogene of West Gondwana (Patagonia,Argentina)

Premise of the Study

The fossil record is critical for testing biogeographic hypotheses. Menispermaceae (moonseeds) are a widespread family with a rich fossil record and alternative hypotheses related to their origin and diversification. The family is well‐represented in Cenozoic deposits of the northern hemisphere, but the record in the southern hemisphere is sparse. Filling in the southern record of moonseeds will improve our ability to evaluate alternative biogeographic hypotheses.

Methods

Fossils were collected from the Salamanca (early Paleocene, Danian) and the Huitrera (early Eocene, Ypresian) formations in Chubut Province, Argentina. We photographed them using light microscopy, epifluorescence, and scanning electron microscopy and compared the fossils with similar extant and fossil Menispermaceae using herbarium specimens and published literature.

Key Results

We describe fossil leaves and endocarps attributed to Menispermaceae from Argentinean Patagonia. The leaves are identified to the family, and the endocarps are further identified to the tribe Cissampelideae. The Salamancan endocarp is assigned to the extant genus Stephania. These fossils significantly expand the known range of Menispermaceae in South America, and they include the oldest (ca. 64 Ma) unequivocal evidence of the family worldwide.

Conclusions

Our findings highlight the importance of West Gondwana in the evolution of Menispermaceae during the Paleogene. Currently, the fossil record does not discern between a Laurasian or Gondwanan origin; however, it does demonstrate that Menispermaceae grew well outside the tropics by the early Paleocene. The endocarps’ affinity with Cissampelideae suggests that diversification of the family was well underway by the earliest Paleocene.     

Variation in the Late Triassic Canjilon quarry (Upper Chinle Group, New Mexico) phytosaur skulls: a case for sexual dimorphism

The Canjilon quarry, located in north-central New Mexico near Ghost Ranch, contains a death assemblage of phytosaurs located stratigraphically high within the Petrified Forest Formation of the Chinle Group (Revueltian = early-mid Norian). The site has yielded numerous fossils ofPseudopalatus-grade phytosaurs, including at least 10 skulls collected from the locality byCharles Camp in 1928 and 1933 and another collected more recently byAlex Downs, curator of paleontology at Ghost Ranch. A re-examination of these skulls reveals two morphotypes that differ only in the relative lengths and relative robustness of their premaxillae. In these two morphotypes, the premaxillae define the shape and length of the rostral crest, the dimensions of which are independent of skull size. In one morphotype, the premaxillae are long, thin bones that lead to an abrupt, volcano-like narial crest. In the second morphotype, the premaxillae are of approximately the same length, but expand dorsoventrally halfway along their lengths, creating a longer and more robust crest. The most probable explanation of these two variants in rostral crest morphology in a Single, catastrophic death assemblage is thatPseudopalatus-grade phytosaurs are sexually dimorphic. Thus, the larger, more robust crest of the first morphotype is probably a display feature, most likely of the male animal. The more gracile snout änderest characterize the female morph. In the sample of phytosaurs examined, there are three individuals of the more robust (male) morphotype, five individuals of the gracile (female) morphotype, and three individuals (2 adult, 1 juvenile) that cannot be assigned to either morphotype because the skulls are too damaged to make an aecurate assessment. This is the first clear evidence of sexual dimorphism in phytosaurs, and has important implications for phytosaur species-level taxonomy, as well as for understanding aspects of their paleobiology (e.g., population dynamics).     

Re‐evaluation of the ontogeny and reproductive biology of the Triassic fish Saurichthys (Actinopterygii,Saurichthyidae)

Viviparity has evolved independently at least 12 times in ray‐finned fishes. However, the fossil record of actinopterygian viviparity is poor, with only two documented occurrences. Both of these are from the non‐teleost actinopterygian Saurichthys, and include S. curionii and S. macrocephalus from the Middle Triassic Meride Limestone (Monte San Giorgio, Switzerland). Here, we present new data on the reproductive biology of these species, giving unprecedented insights into their life‐history. Based on positional and preservational criteria, six specimens were identified as unambiguously gravid. Embryos were positioned dorsal to the gastrointestinal tract, parallel to the axial skeleton and to each other, in the posterior two‐thirds of the abdominal region. A minimum of 16 embryos are preserved in the most fecund females and, based on the largest preserved embryos and smallest preserved neonates, birth must have occurred at 7–12% of maternal fork length. Embryonic crania and teeth are relatively well‐ossified, however ossification of the parietal region is delayed. In the postcranium, the median scale rows and lepidotrichia are ossified, but not the lateral scale rows. Ossified squamation and gradual allometric growth suggests that neonates did not undergo metamorphosis and were relatively precocial. When considered in a phylogenetic context, neither live birth nor internal fertilization appears to represent the primitive state for saurichthyid fishes.     

Depositional environment and biofacies characterisation of the Triassic (Carnian to Rhaetian) carbonate succession of Punta Bassano (Marettimo Island,Sicily)

The aims of this study are to reconstruct the geological setting of the Punta Bassano series (Marettimo Island, Egadi Archipelago, western Sicily) and its palaeogeographic evolution. The reference section for the Upper Triassic of Marettimo shows an alternation of marl and limestone beds together with brecciated levels. The limestones are both homogeneous mudstones with evaporite pseudomorphs and laminated with fenestrae. Foraminiferal, palynomorph, and ostracod associations constrain the Punta Bassano sequence to the Carnian-Rhaetian interval. The Punta Bassano succession represents a shallow inner ramp, ranging from open-marine environment with good water circulation to lagoonal and peritidal protected environments. Freshwater input from rivers or groundwater on the carbonate ramp is indicated by the ostracod microfauna. The comparison of facies and microfauna with those from other sequences of the Mediterranean Upper Triassic (Pyrenees, Corsica, Sardinia, and Tunisia) allows us to confine the Punta Bassano sedimentation to the northern margin of the Tethys, between the Corsican and the Pyrenean depositional setting. These new results indicate that Marettimo Island, which is considered a single structural element being formed by four tectonic units, is a piece of the southern margin of the European Plate, displaced over a longer distance to become part of the other Egadi Islands, when the Corso-Sarde block made its rotation and successive collision with the North African Margin.     

The presence of Compsocerops (Brachyopoidea: Chigutisauridae) (Late Triassic) in southern Brazil with comments on chigutisaurid palaeobiogeography

Abstract: Chigutisauridae is the longest‐lived trematosaurian clade (from early Triassic to early Cretaceous). They were reported in Argentina, Australia, India and South Africa. This contribution reports a putative chigutisaurid specimen in the Carnian of southern Brazil (Santa Maria Formation, Paraná Basin). The material comprises two skull fragments, a mandibular fragment, a clavicular blade and a humerus. Ontogenetic features point to an early development stage of the specimen. The presence of a long, straight and pointed tabular horn, which runs parallel to the skull midline towards its tip, and a distinctive projection in the posterior border of the postparietal indicates a close relationship of the Brazilian chigutisaurid with the Indian Compsocerops cosgriffi. Three distinctive and combined characters suggest that the Brazilian chigutisaurid is a distinctive specimen: the presence of an alar process of the jugal in the ventral margin of the orbit; jugal does not extend well beyond the anterior margin of the orbit; and tabular does not contact the parietal. These characters could justify the erection of a new taxon; however, they might reflect its immature ontogenetic stage as well. Accordingly, we attribute this new specimen to Compsocerops sp. Argentinean and Indian occurrences are dated as Norian, so the presence of a Carnian chigutisaurid in southern Brazil indicates that western Gondwana chigutisaurids have first occupied the Paraná Basin and later migrated towards west (to Argentina) and east (India). However, the presence of ghost chigutisaurid taxa cannot be dismissed, because their long temporal range contrasts with their still short (in comparison with other temnospondyl groups) geographic distribution. Hence, they might have been more geographically widespread than their fossil record suggests.     

Fossil Flowers and Associated Plant Fossils from the Kamikitaba Locality (Ashizawa Formation, Futaba Group, Lower Coniacian, Upper Cretaceous) of Northeast Japan

in situ spores, and megaspores also document the presence of Selaginellaceae and Schizaeaceae. Received 8 February 1999/ Accepted in revised form 7 April 1999     

Fossil fruits of Illigera (Hernandiaceae) from the Eocene of central Tibetan Plateau

Illigera (Hernandiaceae) is a liana genus distributed mainly in the tropical Asia and Africa. Previous fossil records suggested that Illigera was restricted in western North America during the Eocene. Recent paleobotanical investigation has unveiled a Paleogene flora that is totally different from today's vegetation in central Tibet. This provides novel insights into the paleoenvironmental change during the evolution of the Tibetan Plateau (TP). Here, we investigated 10 fruit impressions of Illigera from the early middle Eocene Niubao Formation in the Bangor Basin, central TP. The fossil winged fruits are characterized by their eroded fruit wings, well preserved fusiform locular areas with a median ridge bisecting the fruit, and short veins fanning radially outward. These features allow us to assign these fossils to Illigera eocenica, a species originally discovered in the Eocene of western North America. This is the second fossil occurrence of Illigera worldwide, and the first in Asia. Our finding suggests a warm and humid climate in the central TP during the early middle Eocene, and a close floristic link between Asia and North America during the Paleogene. We also propose a Northern Hemisphere origin and a Paleogene dispersal event from Northern Hemisphere to Africa for Illigera.     

Geological significance of the discovery of Middle Triassic (Ladinian) radiolarians from the Hong Hoi Formation of the Lampang Group,Sukhothai Zone,northern Thailand

This paper presents a newly found fauna of Ladinian (Middle Triassic) radiolarians from the Hong Hoi Formation of the Lampang Group, Sukhothai Zone, northern Thailand. The Ladinian age determined by the radiolarian fauna concords with the age previously determined by mollusks. The study section consists of intercalated sandstone and shale in the lower part and a thick conglomerate in the upper part. The radiolarian-bearing siliceous beds are intercalated within the lower unit. The lithic sandstone of the lower unit consists mainly of volcanic rock fragments, quartz, and feldspar, whereas the thickly bedded conglomerate of the upper unit is characterized by abundant gravel-sized clasts of volcanic rock and limestone. These lithic features of the study section suggest that during deposition the Hong Hoi Formation was located near a supply of volcanic materials. A forearc basin close to the Sukhothai Arc would be the most suitable environment for the deposition. The Ladinian age determined by the radiolarian fauna supports the occurrence of intensive volcanic activity within the Sukhothai Arc during the Middle Triassic.     

Plants of Leptostrobus Heer （Czekanowskiales） from the Early Cretaceous and Late Triassic of China, with Discussion of the Genus

Investigation of the Mesozoic seed plant Leptostrobus Heer from the Yangcaogou Formation of the Late Triassic and the Yixian Formation of the Early Cretaceous, Liaoning Province, China, provides new Insight Into Its general morphology and geographical distribution. The materials of L. cancer from the Yixian Formation described herein are later than all the past findings of this species and add to the record of L. cancer during the Early Cretaceous. Based on well-preserved specimens, the specific diagnosis Is slightly emended and the reconstructlon of L. cancer Is perfected. The materials from the Yangcaogou Formation of the Late Triassic are placed in L. spheericus, in addition, we review the history of investigation of the genus Leptostrobus since its establishment in 1876 and discuss the main characteristics of each species.     

Strontium isotope stratigraphy of the Gabbs Formation (Nevada): implications for global Norian–Rhaetian correlations and faunal turnover

The Luning and Gabbs formations in west‐central Nevada, USA represent a Late Triassic shallow marine sedimentary succession with global significance (the Gabbs Formation was a candidate for the basal Jurassic GSSP). Typically, the Norian–Rhaetian stage boundary is placed at the contact between the formations, and the Rhaetian–Hettangian boundary (the Triassic–Jurassic boundary) is within the Müller Canyon Member of the Gabbs Formation. However, the use of different biostratigraphical index‐species schemes in Norian–Rhaetian successions between Tethys and Panthalassa, the two largest ocean basins at the time, makes precise correlation problematic. Here, we compare ⁸⁷Sr/⁸⁶Sr measurements of well‐preserved carbonate shell material from Nevada to the well‐known and biostratigraphically constrained ⁸⁷Sr/⁸⁶Sr record from Tethys, where a negative excursion in ⁸⁷Sr/⁸⁶Sr is noted across the Norian–Rhaetian boundary. Our new ⁸⁷Sr/⁸⁶Sr data from the Luning and Gabbs formations reveal a comparable trend, with a sharp drop in ⁸⁷Sr/⁸⁶Sr within the Nun Mine Member of the Gabbs Formation, suggesting the position of the Norian–Rhaetian boundary is higher in the succession, and not between the Luning and Gabbs formations as previously defined. Relating the stage boundary using global isotopic signals is a useful tool for biostratigraphical correlation of successions between Tethys and Panthalassa, and for estimating the rate of faunal turnover at the Norian–Rhaetian stage boundary in comparison with the succeeding Late Triassic mass extinction. If correct, this biostratigraphical–chemostratigraphical correlation suggests that the current index groups for the Panthalassic stage boundary should be changed.     

Additional thylacocephalans (Arthropoda) from the Lower Triassic (upper Olenekian) Osawa Formation of the South Kitakami Belt,Northeast Japan

The Lower Triassic Osawa Formation in the South Kitakami Belt, Northeast Japan, consisting mostly of mudstone of shallow-marine environment, was deposited during the late Olenekian (ca. 250 Ma), and is an important unit through which to examine the biotic recovery process after the end-Permian mass extinction. The Osawa Formation is the only unit in Japan that yields thylacocephalans (Arthropoda). Three species belonging to three genera have been reported before: Ankitokazocaris bandoi, Kitakamicaris utatsuensis and Ostenocaris sp. In addition to the known species, some thylacocephalans, including one new genus and three new species, are described in the present paper: Ankitokazocaris tatensis n. sp., Concavicaris parva n. sp., Miyagicaris costata n. gen. n. sp. and Ostenocaris? sp. Although Thylacocephala have a rather long stratigraphic range (from Silurian to Cretaceous) and are known from a wide geographical region, there are only about thirty genera in this group. The Osawa thylacocephalan fauna comprises at least five genera, making it one of the most diverse in the world at the generic level. During the Triassic Period, the Thylacocephala diversified and spread widely throughout low-latitude regions.     

发情交配期贺兰山岩羊的集群特征

2003 年和2004 年的11 ～12 月,采用样线法和全事件取样法研究了贺兰山岩羊的集群特征。共观察到岩羊715 群,共计2 473只。发情前期平均群大小3.5±2.8 只(n = 241),发情期平均群大小3.4±3.2 只(n =297)。发情前期与发情期雄性群、雌性群、雌雄群、混合群及母子群大小间均无显著差异。发情前期到发情期群类型组成差异极显著,发情期雌雄群、混合群和独羊比例显著上升;雄性群、雌性群和母子群比例显著下降。发情前期到发情期,雄性岩羊在雄性群、雌雄群、混合群和独羊中的分布差异极显著,成年雌性岩羊在雌性群、雌雄群、混合群、母子群和独羊中的分布差异极显著。发情期4 个年龄等级雄性在雄性群、雌雄群、混合群和独羊中分布差异极显著,4 个年龄等级雄性岩羊都倾向分布于混合群。
     

A new Hyperodapedon (Archosauromorpha,Rhynchosauria) from the Upper Triassic of India: implications for rhynchosaur phylogeny

A new species of the rhynchosaur genus Hyperodapedon, namely H. tikiensis, is described from well‐preserved skeletal elements that were collected from the Upper Triassic Tiki Formation of India. Hyperodapedon tikiensis is diagnosed on the basis of several cranial and postcranial features including longer than wide basipterygoid process, crest‐shaped maxillary cross section lateral to the main longitudinal groove, deeply excavated neural arches of mid‐dorsal vertebrae, long scapular blade, a pronounced deltopectoral crest, proximal humeral end much broader than distal end, iliac length greater than iliac height, equal pre‐ and postacetabular iliac lengths and circular femoral cross section. Two distinct morphotypes of the maxillary tooth plates can be discerned, which are attributed to ontogenetic variations. A maximum‐parsimony analysis was carried out to show that the order Rhynchosauria is characterized by nine cranial and one postcranial character states. The analysis reveals that Otischalkia elderae is invalid and the basal forms, Howesia and Mesosuchus, are closely related. The Mid‐Triassic genus Ammorhynchus is more derived and forms a sister group to the Late Triassic subfamily Hyperodapedontinae. Isalorhynchus and Teyumbaita are basal to the pandemic genus Hyperodapedon. Twenty‐four characters that are not homoplasious document major patterns of skeletal evolution in rhynchosaurs. From laterally oriented scapula and slender propodials, the postcranial skeleton evolved into a more robust form as is evident from nearly vertical scapula and increase in the robustness of the propodials. Shortening of the femur is noted in the derived Late Triassic forms as exemplified in Hyperodapedon gordoni, Hyperodapedon huxleyi and H. tikiensis.     

The occurrence and preservation of ammonites in the Blue Lias Formation (lower Jurassic) of Devon and Dorset, England and their palaeoecological, sedimentological and diagenetic significance

More than two thirds of beds in the lowest Jurassic, Blue Lias Formation lack ammonites, which are commonly preserved in irregular or planar-bedded, bioturbated limestones, very rarely in laminated limestones and almost never in laminated black shales. Ammonites are preserved in 3D in nodular and planar-bedded limestones and at any orientation to bedding. Co-occurrence with macrobenthos and absence from beds without benthos suggest that Blue Lias ammonites were nektobenthonic. Scour structures and imbrication of ammonites in the Best Bed imply presence of traction currents. Lack of epifauna on large cephalopod shells (and other fossils) implies rapid deposition in event beds. Blue Lias deposition was episodic, not slow and continuous as the fine grain size implies. Undistorted trace fossils, uncrushed ammonites and stable isotope values all suggest early cementation of limestone beds from pore waters of a similar composition to contemporary Jurassic sea water. A clear diagenetic trend exists, with limestones having least, and laminated black shales most, modified stable isotope values. Contrast between trace fossil fills and host sediment demonstrates that Blue Lias rhythms are primary, but limestone beds have been diagenetically cemented.