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Theropod dinosaurs are one of the most remarkable lineages of terrestrial vertebrates in the Mesozoic, showing high taxonomic and ecological diversity. We investigate the cranial diversity of non-avian theropods and some basal birds, using geometric morphometrics to obtain insights into the evolutionary modifications of the skull. Theropod skulls mostly vary in the shape of the snout and length of the postorbital region (principal component [PC] 1), with further variation in orbit shape, depth of the postorbital region, and position of the jaw joint (PC 2 and PC 3). These results indicate that the cranial shape of theropods is closely correlated with phylogeny and dietary preference. Skull shapes of non-carnivorous taxa differ significantly from carnivorous taxa, suggesting that dietary preference affects skull shape. Furthermore, we found a significant correlation between the first three PC axes and functional proxies (average maximum stress and an indicator of skull strength). Interestingly, basal birds occupy a large area within the morphospace, indicating a high cranial, and thus also ecological, diversity. However, we could include only a small number of basal avialan species, because their skulls are fragile and there are few good skull reconstructions. Taking the known diversity of basal birds from the Jehol biota into account, the present result might even underestimate the morphological diversity of basal avialans.
Paranyctoides aralensis, based on a partially known set of lower postcanines from the Turonian Bissekty Formation of Uzbekistan, is proposed to be a junior subjective synonym of Sailestes quadrans, itself based on an M1 from the same stratigraphic unit. As a result, the latter taxon is recombined as Paranyctoides quadrans comb. nov. Based on newly collected or recognized specimens, we show that this species has four lower incisors, including a procumbent il and a relatively small i4, as well as five double-rooted premolars, the smallest of which (p3) can be lost ontogenetically. The p5 of this species is premolariform, rather than molariform as previously thought based on an erroneous identification, and resembles North American species of Paranyctoides in the presence of a small, cingulid-like paraconid and a distal talonid cusp, as well as the absence of a metaconid. The lower molars of Paranyctoides are unique among Late Cretaceous eutherians in having a larger, lingually placed paraconid. Paranycoides probably is the sister taxon of the Late Cretaceous Zhelestidae.
The alvarezsauroid theropod Linhenykus monodactylus from the Upper Cretaceous of Inner Mongolia, China is the first known monodactyl non-avian dinosaur, providing important information on the complex patterns of manual evolution seen in alvarezsauroids. Here we provide a detailed description of the osteology of this taxon. Linhenykus shows a number of features that are transitional between parvicursorine and non-parvicursorine alvarezsauroids, but detailed comparisons also reveal that some characters had a more complex distribution. We also use event-based tree-fitting to perform a quantitative analysis of alvarezsauroid biogeography incorporating several recently discovered taxa. The results suggest that there is no statistical support for previous biogeographic hypotheses that favour pure vicariance or pure dispersal scenarios as explanations for the distributions of alvarezsauroids across South America, North America and Asia. Instead, statistically significant biogeographic reconstructions suggest a dominant role for sympatric (or “within area”) events, combined with a mix of vicariance, dispersal and regional extinction. At present the alvarezsauroid data set is too small to completely resolve the biogeographic history of this group: future studies will need to create larger data sets that encompass additional clades.
Recent discoveries of more than ten new species of tyrannosauroid theropods are helping to understand the origin and evolution of colossal body size and other characteristic features of Tyrannosaurus rex and its terminal Cretaceous relatives. Particularly important has been the discovery and reinterpretation of Late Jurassic tyrannosauroids from Europe and North America, which are intermediate in size and phylogenetic position between small basal tyrannosauroids and the largest Late Cretaceous species. The fragmentary nature of these Jurassic specimens, however, has frustrated attempts to understand their systematics and phylogeny. A new specimen from the Late Jurassic of England was recently named as a new species (Stokesosaurus langhami) of the genus Stokesosaurus, which is known from several fragmentary fossils from North America. We review the systematics and phylogeny of these European and North American specimens and show that there are no unequivocal synapomorphies uniting them. Furthermore, a revised phylogenetic analysis does not recover them as sister taxa. This necessitates a taxonomic revision of this material, and we name a new genus (Juratyrant) for the British specimen.
A Mongolian ankylosaurid specimen identified as Tarchia gigantea is an articulated skeleton including dorsal ribs, the sacrum, a nearly complete caudal series, and in situ osteoderms. The tail is the longest complete tail of any known ankylosaurid. Remarkably, the specimen is also the first Mongolian ankylosaurid that preserves impressions of the keratinous scales overlying the bony osteoderms. This specimen provides new information on the shape, texture, and arrangement of osteoderms. Large flat, keeled osteoderms are found over the pelvis, and osteoderms along the tail include large keeled osteoderms, elongate osteoderms lacking distinct apices, and medium-sized, oval osteoderms. The specimen differs in some respects from other Tarchia gigantea specimens, including the morphology of the neural spines of the tail club handle and several of the largest osteoderms.
The Early Triassic record of the large capitosaurid amphibian genus Parotosuchus is supplemented by new material from fluvial deposits of Wióry, southern Poland, corresponding in age to the Detfurth Formation (Spathian, Late Olenekian) of the Germanic Basin. The skull of the new capitosaurid shows an “intermediate” morphology between that of Parotosuchus helgolandicus from the Volpriehausen-Detfurth Formation (Smithian, Early Olenekian) of Germany and the slightly younger Parotosuchus orenburgensis from European Russia. These three species may represent an evolutionary lineage that underwent a progressive shifting of the jaw articulation anteriorly. The morphology of the Polish form is distinct enough from other species of Parotosuchus to warrant erection of a new species. The very large mandible of Parotosuchus ptaszynskii sp. nov. indicates that this was one of the largest tetrapod of the Early Triassic. Its prominent anatomical features include a triangular retro articular process and an elongated base of the hamate process.
Fossil catshark egg capsules, Scyliorhinotheca goederti gen. et sp. nov., are reported from a Late Eocene deep-water methane-seep calcareous deposit in western Washington State, USA. The capsules are preserved three-dimensionally and some show mineralized remnants of the ribbed capsule wall consisting of small globular crystals that are embedded in a microsparitic matrix. The globules are calcitic, but a strontium content of 2400–3000 ppm suggests that they were originally aragonitic. The carbonate enclosing the egg capsules, and the capsule wall itself, show δ13C values as low as -36.5‰, suggesting that formation was induced by the anaerobic oxidation of methane and hence in an anoxic environment. We put forward the following scenario for the mineralization of the capsule wall: (i) the collagenous capsules experienced a sudden change from oxic to anoxic conditions favouring an increase of alkalinity; (ii) this led to the precipitation of aragonitic globules within the collagenous capsule wall; (iii) subsequently the remaining capsule wall was mineralized by calcite or aragonite; (iv) finally the aragonitic parts of the wall recrystallized to calcite. The unusual globular habit of the early carbonate precipitates apparently represents a taphonomic feature, resulting from mineralization mediated by an organic matrix. Taphonomic processes, however, are at best contributed to an increase of alkalinity, which was mostly driven by methane oxidation at the ancient seep site.
The new retiolitid species, Spinograptus tubothecalis, is described from the Colonograptus praedeubeli and C. deubeli biozones from two localities in Poland: a borehole on the East European Platform and the Holy Cross Mountains. This was a recovery phase after the severe Silurian Cyrtograptus lundgreni Event. The new species has a unique, previously undescribed form of finite rhabdosome. Unlike the species Spinograptus reticulolawsoni and S. lawsoni, in which the finite rhabdosomes taper distally, its rhabdosome is parallel-sided with the two distal thecae developed as isolated tubes without genicular processes, with a small appendix between them. The new species also has preserved membranes of the sicula, thecae and ancora sleeve, similar to a few species of Spinograptus from the lower Homerian. Spinograptus tubothecalis, like Spinograptus clathrospinosus and S. spinosus, has paired reticulofusellar genicular processes on the pre-thecal ventral orifices, similar to but shorter than thecal processes. Transverse rods, a rare character in post-Cyrtograptus lundgreni Event retiolitids occur in the new species in rudimentary form.
Timothy P. Topper, Lars E. Holmer, Christian B. Skovsted, Glenn A. Brock, Uwe Balthasar, Cecilia M. Larsson, Sandra Pettersson Stolk, David A.T. Harper
The morphology and organophosphatic shell structure of the paterinate brachiopod Askepasma is documented using new and previously collected specimens from the lower Cambrian of South Australia. Lack of adequately preserved material has seen the majority of paterinate specimens previously reported from South Australia referred to the genus Askepasma and treated under open nomenclature. Large collections of paterinates from the lower Cambrian Wilkawillina, Ajax, and Wirrapowie limestones in the Arrowie Basin, South Australia have prompted redescription of the type species Askepasma toddense and the erection of a new species, Askepasma saproconcha sp. nov. Askepasma saproconcha sp. nov. currently represents the oldest known brachiopod from the lower Cambrian successions in South Australia with a FAD in pre-trilobitic (Terreneuvian, Cambrian Stage 2, lower Atdabanian) strata in the basal part of the Wilkawillina and Wirrapowie limestones. Askepasma toddense predominantly occurs in Abadiella huoi Zone equivalent strata (Unnamed Cambrian Series 2, Stage 3, middle-upper Atdabanian) in the upper part of the lower Wilkawillina, Wirrapowie, and Ajax limestones. The shell microstructure of Askepasma suggests a proximal stem group position within the Brachiopoda and similarities with tommotiid taxa provides further evidence that the ancestry of crown group brachiopods is firmly entrenched within the Tommotiida.
In a letter to Nature (February, 2011), Xunlai Yuan and collaborators recorded carbon compression fossils from black shales of the Lantian Formation (Ediacaran), southern Anhui Province, South China. The new fossils, described under five morphological types (Types A to E), exhibit degrees of morphological differentiation suggesting that they were multicellular eukaryotes. Some of the Lantian macrofossils were interpreted as algae, but others are of unknown affinities. For reasons noted in this discussion, Type A fossils attracted our particular attention, and we suggest an alternative interpretation of their affinities. According to our view, some of them (at least those with three faces and no globose holdfast at their base) may represent conulariid cnidarians or close medusozoan relatives. The undistorted organism probably was a three-sided cone in life. We believe that our suggested alternative interpretations of the anatomy and affinities of the fossils in question can be useful in guiding future research on the oldest currently known fossil assemblage of multicellular organisms.
The findings of preserved soft body parts including pedicle in the linguloid brachiopods are extremely rare in the fossil record of which the early Cambrian Chengjiang (southern China) and Burgess Shale (British Columbia) faunas are the most important. However, these characteristic Cambrian soft-bodied faunas largely disappeared from the fossil record well before the end of the Cambrian. Here we describe the first record of the pedicle in a linguloid brachiopod from the post-Cambrian strata, preserved with remarkable fidelity. Contrary to the Chengjiang and Burgess Shale-type faunas which are commonly preserved as essentially two-dimensional aluminosilicate or degraded organic carbon films or pyritized compressed fossils, the specimens now recovered from the Ordovician of China show a three-dimensional, pyritized pedicle with preserved external morphology, in detail. The presence of streamlined shell shape and burrowing shell sculpture in our specimens supports an assumption for infaunal mode of life of the genus. Since all linguloid brachiopods of the early Cambrian are interpreted as epifaunal or semi-infaunal, it seems that the here described Ordovician linguloid is the oldest representative of fully infaunal brachiopods. Apparently, the long vermiform and flexible linguloid pedicle has appeared as a functionally optimized construction.
The fossil dragonfly Eumorbaeschna adriankini sp. nov. is decribed as first fossil insect from the Upper Jurassic of Central Poland (Owadów-Brzezinki quarry, Tomaszów Mazowiecki area), and as first record of the family Eumorbaeschnidae (Odonata, Anisoptera, Aeshnoptera) outside the Solnhofen lithographic limestone.
Investigations of the Upper Permian strata in the Iran-Transcaucasia resulted in identification of 32 ammonoid genera. The majority of ammonoids in this collection belong to the order Ceratitida (75%). Among Dzhulfian ceratitid ammonoids representatives of the family Araxoceratidae (Otoceratoidea) are most abundant. The assemblage structure changed radically during latest Permian (Dorashamian) time, bringing a domination of the family Dzhulfitidae. The Induan (Lower Triassic) succession in the Verkhoyansk area provided a few groups of ammonoids which are Palaeozoic in type: families Episageceratidae (Episageceras), Xenodiscidae (Aldanuceras and Metuphiceras), and Dzhulfitidae (Tumpuphiceras) and superfamily Otoceratoidea (Otuceras and Vaviluvites). It demonstrates the survival of ammonoids belonging to these groups the Permian—Triassic (P—T) boundary extinction event and their quick migration to the vast areas of higher latitudes (together with some representatives of the Mesozoic-type families). Induan—Olenekian ammonoid successions in South Primorye, Mangyshlak, and Arctic Siberia illustrate the high rate of Early Triassic ammonoid recovery in both the Tethys and the Boreal realm. New ammonoid taxa are described: Proptychitina subordo nov., Ussuritina subordo nov., Subbalhaeceras shigetai gen. and sp. nov. (Flemingitidae), Mesohedenstruemia ulgae sp. nov. (Hedenstrormiidae), and Inyuites sedini sp. nov. (Inyoitidae).
The Spathian (late Early Triassic) Virgin Formation of south-western Utah (USA) yields a comparatively diverse benthic fauna that flourished ∼2 Ma after the end-Permian mass extinction. In this study, we present quantitative palaeoecological data, which are analysed in the context of depositional environments. This integrated approach helps to discriminate between effects of the end-Permian mass extinction event and local environmental factors on alpha diversity and ecological structure of the Virgin Fauna. Shallow subtidal environments yield the highest species richness and lowest dominance values as recorded in two benthic associations, the Eumorphotis ericius Association and the Protogusarella smithi Association, both of which contain 20 benthic species (bivalves, gastropods, brachiopods, echinoderms, and porifers). Tidal inlet deposits yield a low diverse fauna (Piarorhynchella triassica Association) with a very high dominance of filter feeders adapted to high energy conditions. Another comparably low diverse fauna is recorded by the Bakevellia exporrecta Association, which occurs in deposits of the offshore transition zone, most likely reflecting unconsolidated substrates. A single sample containing five bivalve species (Bakevellia costata Assemblage) is recorded from a marginal-marine setting. The Virgin fauna yields a bulk diversity of 30 benthic species (22 genera) of body fossils and 14 ichnogenera and, thus, represents the most diverse marine bottom fauna known so far from the Early Triassic. Our results suggest that oceanographic conditions during the early Spathian enabled ecosystems to rediversify without major abiotic limitations. However, taxonomical differentiation between habitats was still low, indicating a time lag between increasing within-habitat diversity (alpha diversity) and the onset of taxonomical differentiation between habitats (beta diversity). We suggest that taxonomical habitat differentiation after mass extinction events starts only when within-habitat competition exceeds a certain threshold, which was not yet reached in the Spathian of the investigated area. This interpretation is an alternative to previous suggestions that the prevalence of generalistic taxa in the aftermath of mass extinction events reflects protracted environmental stress. The onset of increasing beta diversity is a potential criterion for distinguishing two major recovery phases, the first ending with habitat saturation and the second ending with the completion of ecosystem differentiation.
The fossil record of coelacanths is patchy, with very few taxa known from the Triassic of Asia. We report here two new genera and species of coelacanths from the Luoping Biota, a recently found site of exceptional fossil preservation from Yunnan, South China. The first new taxon, Luopingcoelacanthus eurylacrimalis, is based on four specimens, which together show most aspects of the anatomy. One specimen shows two small coelacanths inside the ventral portion of the abdominal cavity, and these are interpreted as intrauterine embryos, close to birth size, based on comparisons with previously reported embryos of the fossil coelacanths Rhabdoderma and Undina, and the extant genus Latimeria. Our new find extends the evidence for ovoviviparity in coelacanths back from the Late Jurassic to the Middle Triassic. The second new taxon, Yunnancoelacanthus acrotuberculatus, is based on one specimen, and differs from Luopingcoelacanthus in the dentary, lachrymojugal, number of rays of the first dorsal fin, and especially in the ornament on dermal bones and scales. A cladistic analysis shows that the new taxa are closest relatives to the derived clade Latimerioidei. The relatively high diversity of coelacanths in the Early Triassic, and adaptations of living Latimeria to low-oxygen conditions, suggests that the group may have included ‘disaster taxa’ that benefited from anoxic and dysoxic ocean conditions in the aftermath of the end-Permian mass extinction.
One of the most complete Permian-Triassic boundary sections located in the Bükk Mountains (Hungary) was sampled for ostracod study. Seventy-six species are recognized, belonging to twenty genera. Fifteen new species are described and figured: Acratia? jeanvannieri Forel sp. nov., Acratia nagyvisnyoensis Forel sp. nov., Bairdia anisongae Forel sp. nov., Bairdia davehornei Forel sp. nov., Callicythere? balvanyseptentrioensis Forel sp. nov., Cytherellina? magyarorszagensis Forel sp. nov., Eumiraculum desmaresae Forel sp. nov., Hollinella fengqinglaii Crasquin sp. nov., Hungarella gerennavarensis Crasquin sp. nov., Langdaia bullabalvanyensis Crasquin sp. nov., Liuzhinia venninae Forel sp. nov., Liuzhinia bankutensis Forel sp. nov., Microcheilinella egerensis Forel sp. nov., Reviya praecurukensis Forel sp. nov., Shemonaella? olempskaella Forel sp. nov. One species is renamed: Bairdia baudini Crasquin nom. nov. Comparison of the Bálvány North section with the Meishan section (Zhejiang Province, South China), Global Boundary Stratotype Section and Point (GSSP) of the Permian-Triassic Boundary (PTB), reveals discrepancies linked to the environmental setting and particularly to bathymetry. The stratigraphical distribution of all the species is given and diversity variations are discussed. The Bálvány North section exhibits the lowest extinction rate of all PTB sections studied for ostracods analysis associated with a high level of endemism.
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