Three new species of Cyphaspides are proposed: C. ammari, C. nicoleae, and C. pankowskiorum. These species are based on specimens obtained from Middle Devonian (Eifelian) strata of the Bou Tchrafine Group, near Erfoud, in the Province of Errachidia, southeastern Morocco. The present contribution enhances our knowledge of Cyphaspides by providing details of three new species that are based on well-preserved, complete, and articulated types. The genus Cyphaspides is discussed, and an emended diagnosis is provided. The paleobiogeography, ontogeny, and relationships of the genus are discussed.

UUID: http://zoobank.org/4a7aab8f-8c8e-4498-9cc2-6f8c69b85213

The giant carnivorous phorusrhacid bird Phorusrhacos longissimus (Aves, Cariamiformes) was first described in 1887 by Florentino Ameghino on the basis of a jaw fragment. The majority of a skull of the species still encased in crumbling rock was preserved only long enough for illustrations to be made by Carlos Ameghino in the field and for a brief description to be written. Skull remains of this species have remained scarce, and few postcranial remains have been figured. Here, we reassess the cranial anatomy of this outstanding ‘terror bird’ species taking into account data from a newly discovered skull. An additional specimen of a well-preserved dorsal vertebra referable to Phorusrhacinae is also described from a separate locality within the Miocene Santa Cruz Formation (late early Miocene) from Santa Cruz Province in Argentina. The skull includes most of the rostrum, skull roof, and mandible and is compared with material from other members of the Phorusrhacinae. The new data from the skull and vertebra provide morphological features of this clade that benefit future taxonomic and phylogenetic analyses of this iconic group of birds.

The lower-middle Hetang Formation (Cambrian Stage 2–3) deposited in slope-basinal facies in South China is well known for its preservation of the earliest articulated sponge fossils, providing an important taphonomic window into the Cambrian Explosion. However, the Hetang Formation also hosts a number of problematic animal fossils that have not been systematically described. This omission results in an incomplete picture of the Hetang biota and limits its contribution to the understanding of the early evolution of animals. Here we describe a new animal taxon, Cambrowania ovata Tang and Xiao, new genus new species, from the middle Hetang Formation in the Lantian area of southern Anhui Province, South China. Specimens are preserved as carbonaceous compressions, although some are secondarily mineralized. A comprehensive analysis using reflected light microscopy, scanning electron microscopy, energy-dispersive X-ray spectroscopy, and micro-CT reveals that the new species is characterized by a spheroidal to fusoidal truss-like structure consisting of rafter-like crossbars, some of which are secondarily baritized and may have been internally hollow. Some specimens have aperture-like structures that are broadly similar to oscula of sponges, whereas others show evidence of a medial split reminiscent of gaping carapaces. While the phylogenetic affinity of Cambrowania ovata Tang and Xiao, n. gen. n. sp. remains problematic, we propose that it may represent carapaces of bivalved arthropods or more likely sponges in early life stages. Along with other problematic metazoan fossils such as hyolithids and sphenothallids, Cambrowania ovata Tang and Xiao, n. gen. n. sp. adds to the diversity of the sponge-dominated Hetang biota in an early Cambrian deepwater slope-basinal environment.

UUID: http://zoobank.org/44de9472-7e3f-42d1-9554-7b3434df91d9

The recent 2014–2017 controlled excavations carried out in quarries of Sverdlovsk and Tyumen regions, Ural and Western Siberia, are contributing to the knowledge of the middle to late Eocene marine fauna from the Turgay Strait, mainly composed of invertebrates and cartilaginous and bony fishes. Here we present a preliminary report of the bony fishes collected during the campaigns carried out in two Eocene Trans-Urals localities. The sediments of these localities were deposited in a large epicontinental marine basin during the middle to late Eocene, when the Turgay Strait connected the Arctic waters to the North and the Peritethys to the South. Most of the bony fish material examined can be assigned to codfishes (Gadiformes) according to morphological comparison with living and fossil taxa. In particular, nearly complete three-dimensionally preserved lower jaws are confidently assigned to the Merlucciidae, whereas isolated vertebrae and bones are referred to as Gadiformes incertae sedis. This report documents the easternmost occurrence of this group of fishes in the northern hemisphere during the early phases of their radiation, revealing their presence into the Turgay Strait before its closure.

Large body sizes among nonavian theropod dinosaurs is a major feature in the evolution of this clade, with theropods reaching greater sizes than any other terrestrial carnivores. However, the early evolution of large body sizes among theropods is obscured by an incomplete fossil record, with the largest Triassic theropods represented by only a few individuals of uncertain ontogenetic stage. Here I describe two neotheropod specimens from the Upper Triassic Bull Canyon Formation of New Mexico and place them in a broader comparative context of early theropod anatomy. These specimens possess morphologies indicative of ontogenetic immaturity (e.g., absence of femoral bone scars, lack of co-ossification between the astragalus and calcaneum), and phylogenetic analyses recover these specimens as early-diverging neotheropods in a polytomy with other early neotheropods at the base of the clade. Ancestral state reconstruction for body size suggests that the ancestral theropod condition was small (~240 mm femur length), but the ancestral neotheropod was larger (~300–340 mm femur length), with coelophysoids experiencing secondary body size reduction, although this is highly dependent on the phylogenetic position of a few key taxa. Theropods evolved large body sizes before the Triassic–Jurassic extinction, as hypothesized in most other ancestral state reconstructions of theropod body sizes, but remained rare relative to smaller theropods until the Jurassic.

The Flat Rocks locality in the Wonthaggi Formation (Strzelecki Group) of the Gippsland Basin, southeastern Australia, hosts fossils of a late Barremian vertebrate fauna that inhabited the ancient rift between Australia and Antarctica. Known from its dentary, Qantassaurus intrepidus Rich and Vickers-Rich, 1999 has been the only dinosaur named from this locality. However, the plethora of vertebrate fossils collected from Flat Rocks suggests that further dinosaurs await discovery. From this locality, we name a new small-bodied ornithopod, Galleonosaurus dorisae n. gen. n. sp. from craniodental remains. Five ornithopodan genera are now named from Victoria. Galleonosaurus dorisae n. gen. n. sp. is known from five maxillae, from which the first description of jaw growth in an Australian dinosaur is provided. The holotype of Galleonosaurus dorisae n. gen. n. sp. is the most complete dinosaur maxilla known from Victoria. Micro-CT imagery of the holotype reveals the complex internal anatomy of the neurovascular tract and antorbital fossa. We confirm that Q. intrepidus is uniquely characterized by a deep foreshortened dentary. Two dentaries originally referred to Q. intrepidus are reassigned to Q. ?intrepidus and a further maxilla is referred to cf. Atlascopcosaurus loadsi Rich and Rich, 1989. A further ornithopod dentary morphotype is identified, more elongate than those of Q. intrepidus and Q. ?intrepidus and with three more tooth positions. This dentary might pertain to Galleonosaurus dorisae n. gen. n. sp. Phylogenetic analysis recovered Cretaceous Victorian and Argentinian nonstyracosternan ornithopods within the exclusively Gondwanan clade Elasmaria. However, the large-bodied taxon Muttaburrasaurus langdoni Bartholomai and Molnar, 1981 is hypothesised as a basal iguanodontian with closer affinities to dryomorphans than to rhabdodontids.

UUID: http://zoobank.org/4af87bb4-b687-42f3-9622-aa806a6b4116

Thirteen bryozoan species are described from the Brewer Dock (Hickory Corners) Member of the Reynales Formation (lower Silurian, Aeronian) at the locality Hickory Corners in western New York, USA. Three species are new: trepostomes Homotrypa niagarensis n. sp. and Leioclema adsuetum n. sp. and the rhabdomesine cryptostome Moyerella parva n. sp. Only one species, Hennigopora apta Perry and Hattin, 1960, developed obligatory encrusting colonies whereas the others produced erect ramose colonies of various thicknesses and shapes: cylindrical, branched, and lenticular. Bryozoans display high abundance and richness within the rock. This fauna is characteristic of a moderately agitated environment with a stable substrate. The identified species reveal paleobiogeographic connections to other Silurian localities of New York as well as Ohio and Indiana (USA) and Anticosti (Canada).

UUID: http://zoobank.org/c24f3184-4a8e-44c4-b6c0-d30433e841c9

An assemblage of Cambrian Series 2, Stages 3–4, conchiferan mollusks from the Shackleton Limestone, Transantarctic Mountains, East Antarctica, is formally described and illustrated. The fauna includes one bivalve, one macromollusk, and 10 micromollusks, including the first description of the species Xinjispira simplex Zhou and Xiao, 1984 outside North China. The new fauna shows some similarity to previously described micromollusks from lower Cambrian glacial erratics from the Antarctic Peninsula. The fauna, mainly composed of steinkerns, is relatively low diversity, but the presence of diagnostic taxa, including helcionelloid Davidonia rostrata (Zhou and Xiao, 1984), bivalve Pojetaia runnegari Jell, 1980, cambroclavid Cambroclavus absonus Conway Morris in Bengtson et al., 1990, and bradoriid Spinospitella coronata Skovsted et al., 2006, as well as the botsfordiid brachiopod Schizopholis yorkensis (Ushatinskaya and Holmer in Gravestock et al., 2001), in the overlying Holyoake Formation correlates the succession to the Dailyatia odyssei Zone (Cambrian Stages 3–4) in South Australia.

Two chondrichthyan assemblages of Late Mississippian/Early Pennsylvanian age are now recognized from the western Grand Canyon of northern Arizona. The latest Serpukhovian Surprise Canyon Formation has yielded thirty-one taxa from teeth and dermal elements, which include members of the Phoebodontiformes, Symmoriiformes, Bransonelliformes, Ctenacanthiformes, Protacrodontoidea, Hybodontiformes, Neoselachii (Anachronistidae), Paraselachii (Gregoriidae, Deeberiidae, Orodontiformes, and Eugeneodontiformes), Petalodontiformes, and Holocephali. The euselachian grade taxa are remarkably diverse with four new taxa recognized here; the Protacrodontidae: Microklomax carrieae new genus new species and Novaculodus billingsleyi new genus new species, and the Anchronistidae: Cooleyella platera new species and Amaradontus santucii new genus new species The Surprise Canyon assemblage also has the youngest occurrence of the elasmobranch Clairina, previously only known from the Upper Devonian. The Surprise Canyon Formation represents a nearshore fluvial infilling of karstic channels, followed by a shallow marine bioherm reef, and finally deeper open water deposition. The early Bashkirian Watahomigi Formation represents open marine deposition and contains only two taxa: a new xenacanthiform, Hokomata parva new genus new species, and the holocephalan Deltodus. The relationship between the Surprise Canyon and Watahomigi chondrichthyan assemblages and other significant coeval chondrichthyan assemblages suggests that there may have been eastern and western distinctions among the Euamerican assemblages during the Serpukhovian due to geographic separation by the formation of Pangea.

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Sertulaster keslingi new genus new species (Palaeasteridae) and Delicaster hotchkissi new species (Permasteridae) are asteroid echinoderms described, respectively, from the Ordovician and Carboniferous of eastern North America. The new genus and species help to document diversity within taxa of lower rank. S. keslingi is similar to the Early Ordovician Eriaster Blake and Guensburg, 2005 but exhibits less differentiation of the skeletal elements from beyond the ambulacral column, that of the so-called extraxial skeleton, whereas the comparatively robust construction of Delicaster hotchkissi clearly departs from that of the type species, D. enigmaticus (Kesling, 1967). Small sample sizes and incomplete exposure of available specimens illustrate ambiguities typically encountered in the study of fossil asteroids.

UUID: http://zoobank.org/fb44075c-396b-4f95-b997-29cdf3c4f3c1

Dissorophoid temnospondyls are widely considered to have given rise to some or all modern amphibians (Lissamphibia), but their ingroup relationships still bear major unresolved questions. An inclusive phylogenetic analysis of dissorophoids gives new insights into the large-scale topology of relationships. Based on a TNT 1.5 analysis (33 taxa, 108 characters), the enigmatic taxon Perryella is found to nest just outside Dissorophoidea (phylogenetic defintion), but shares a range of synapomorphies with this clade. The dissorophoids proper are found to encompass a first dichotomy between the largely paedomorphic Micromelerpetidae and all other taxa (Xerodromes). Within the latter, there is a basal dichotomy between the large, heavily ossified Olsoniformes (Dissorophidae + Trematopidae) and the small salamander-like Amphibamiformes (new taxon), which include four clades: (1) Micropholidae (Tersomius, Pasawioops, Micropholis); (2) Amphibamidae sensu stricto (Doleserpeton, Amphibamus); (3) Branchiosauridae (Branchiosaurus, Apateon, Leptorophus, Schoenfelderpeton); and (4) Lissamphibia. The genera Platyrhinops and Eoscopus are here found to nest at the base of Amphibamiformes. Represented by their basal-most stem-taxa (Triadobatrachus, Karaurus, Eocaecilia), lissamphibians nest with Gerobatrachus rather than Amphibamidae, as repeatedly found by former analyses.

UUID: http://zoobank.org/dadf36db-e003-4af7-bfa7-44d79bc04450

Phosphatic sclerites of the problematic Tarimspira Yue and Gao, 1992 (Cambrian Series 2) recovered by weak acid maceration of limestones display a unique range of mainly strongly coiled morphologies. They were likely organized into multielement scleritomes, but the nature of these is poorly known; some sclerites may have had a grasping function. Tarimspira sclerites grew by basal accretion in an analogous fashion to younger paraconodonts (Cambrian Series 3–4) but lack a basal cavity. Based on proposed homologies, Tarimspira may provide an extension of the early vertebrate paraconodont–euconodont clade back into the early Cambrian. Tarimspira is described for the first time from Laurentia (North Greenland), extending its known range from China and Siberia in Cambrian Series 2. In addition to the type species, Tarimspira plana Yue and Gao, 1992, the Greenland record of Tarimspira includes two morphotypes of a new species, Tarimspira artemi.

UUID: http://zoobank.org/c7c536c8-cdaf-49a9-ae1d-77c392f553fc.

A new specimen of a theropod dinosaur found in the Upper Jurassic (Freixial Formation, late Tithonian) of the Lusitanian Basin is described. It corresponds to a single individual and includes a sequence of articulated caudal vertebrae, an almost complete right pes, and other fragments of the appendicular skeleton. The specimen includes the most complete pes of a theropod dinosaur currently known in the Lusitanian Basin and represents one of the youngest skeletal records of theropod dinosaurs currently known in the Portuguese Upper Jurassic.

A systematic analysis of this specimen is performed and it shows a combination of characters that allows us to interpret it as belonging to an allosauroid taxon. Within this clade, the material from Cambelas shares a few features with some carcharodontosaurids, including the presence of a lateral lamina extending along the anterior end of the centrum in the caudal vertebra and of a low vertical crest on the lateral surface of the femoral lesser trochanter. The set of remains described here shares some unusual features with another specimen previously described in the Portuguese fossil record, which also presents some synapomorphies of Carcharodontosauria. However, no autapomorphy or exclusive character combination can be recognized in the specimen here described in order to describe it as a new form. The presence of this specimen suggests a greater diversity in the allosauroid theropod fauna from the Late Jurassic of the Lusitanian Basin than currently known and probably expands the temporal record of Carcharodontosauria up to upper Tithonian of south-western Europe.

The Cretaceous landmass of Appalachia has preserved an understudied but nevertheless important record of dinosaurs that has recently come under some attention. In the past few years, the vertebrate faunas of several Appalachian sites have been described. One such locality, the Ellisdale site of the Cretaceous Marshalltown Formation of New Jersey, has produced hundreds of remains assignable to dinosaurs, including those of hadrosauroids of several size classes, indeterminate ornithopods, indeterminate theropods, the teeth, cranial, and appendicular elements of dromaeosaurids, ornithomimosaurians, and tyrannosauroids, and an extensive microvertebrate assemblage. The theropod dinosaur record of the Ellisdale site is currently the most extensive and diverse known from the Campanian of Appalachia. Study of the Ellisdale theropod specimens suggests that at least four or more non-avian theropod taxa are represented at the site, including tyrannosauroids, ornithomimosaurs, several different morphotypes of dromaeosaurids that are the first of that clade described from New Jersey, and indeterminate theropods. The specimens are important for increasing current knowledge about the theropod diversity of the Atlantic Coastal Plain (ACP) during the Campanian by representing the most speciose assemblage of the group during the time in the ACP as well as for shedding light on Appalachian dinosaur ecology and biogeography generally.

An ostracode fauna is described from lacustrine sediments of the Hettangian, Lower Jurassic, Whitmore Point Member of the Moenave Formation. The Moenave is well known for its rich, Late Triassic?–Early Jurassic fossil record, which includes fossil fishes, stromatolites, ostracodes, spinicaudatans, and a diverse ichnofauna of invertebrates and vertebrates. Four ostracode species, all belonging to the suborder Darwinulocopina, were recovered from these sediments: Suchonellina globosa, S. stricta, Whipplella? sp. 1, and W.? sp. 2. The diversity and composition of the Whitmore Point Member ostracode fauna agree with previous interpretations about Lake Dixie and nearby paleoenvironments as shallow lakes inhabited by darwinulocopine species that survived the effects of the Central Atlantic Magmatic Province and the subsequent end-Triassic extinction and quickly recolonized these areas, thanks to asexual reproduction by parthenogenesis. The Lake Dixie region, in its geographical isolation, could represent the last episode of darwinulocopine dominance in nonmarine environments before the Late Jurassic diversification of the cypridocopine/cytherocopine modern ostracodes.

Fossil insects from the Triassic-Jurassic boundary of England could provide an important resource for investigating the severity of extinction events in the terrestrial realm of the uppermost Triassic. However, the fossil record is poorly understood for this period even though there are abundant historical collections. Many of these collections are still in need of taxonomic revision before they can be used to reconstruct past entomofaunas and make inferences about diversity change through time. This paper is part of a larger project to revise the taxonomy of insects across the Triassic-Jurassic boundary of England to better understand changes in insect diversity through the Triassic-Jurassic boundary and associated extinction period. Herein, the damsel-dragonfly family Liassophlebiidae Tillyard, 1925 is revised and an additional specimen from the Early Jurassic of Antarctica is included. Rossiphlebia new genus is erected for Liassophlebia jacksoni Zeuner, 1962; L. batheri Tillyard, 1925 is considered nomen dubium and another specimen originally attributed to L. batheri is identified as L. withersi Tillyard, 1925. Liassophlebia (?) clavigaster Tillyard, 1925 and L. (?) hopei (Brodie, 1845) are considered incertae sedis at the generic level. Liassophlebia gigantea Zeuner, 1962 is based on a fragmentary specimen but has several unique key characteristics. We redescribe it in Anglophlebia new genus and tentatively in Anglophlebiidae new family in Heterophlebioptera. Also discussed are L. magnifica Tillyard, 1925, L. withersi, and L. pseudomagnifica Whalley, 1985, which are redescribed with updated figures. Caraphlebia antarctica Carpenter, 1969 was originally described from the Early Jurassic of Antarctica as being closely related to Liassophlebia; it is herein confirmed in Selenothemistidae Handlirsch, 1939.

UUID: http://zoobank.org/8fe9a39c-1c3b-4bda-92a6-92c1a0fc95b8

The Late Triassic rhynchocephalian Clevosaurus latidens Fraser, 1993 is known from the fissure deposits of Cromhall Quarry, England. Many studies have questioned its referral to the genus Clevosaurus Swinton, 1939 and some phylogenetic analyses suggest a close relationship with herbivorous rhynchocephalians. We re-examine the type specimens and referred material of C. latidens to elucidate its taxonomic identity. Additionally, we provide new phylogenetic analyses of the Rhynchocephalia using both parsimony and Bayesian approaches. Our taxonomic review and both phylogenetic analyses reveal that C. latidens is not referable to Clevosaurus, but represents a new genus. We reassess C. latidens and provide an amended diagnosis for Fraserosphenodon new genus. Both parsimony and Bayesian analyses recover similar topologies and we propose formal names for two higher clades within Rhynchocephalia: Eusphenodontia new infraorder and Neosphenodontia new clade.

UUID: http://zoobank.org/65f29bd1-47e3-4a73-af8c-9181c19319e4

Despite 70 years of study, Dickinsonia remains one of the Ediacara biota’s most enigmatic taxa with both morphological characters and phylogenetic affinities still debated. A large population of relatively small Dickinsonia costata present on a semi-contiguous surface from the Crisp Gorge fossil locality in the Flinders Ranges (South Australia) provides an opportunity to investigate this taxon in its juvenile form. This population supports earlier findings that suggest D. costata’s early growth was isometric, based on the relationship between measured variables of length and width. The number of body units increases with length, but at a decreasing rate. A correlation between a previously described physical feature, present as a shrinkage rim partially surrounding some specimens and a novel, raised lip in some specimens, suggests that both features may have been the result of a physical contraction in response to the burial process, rather than due to a gradual loss of mass during early diagenesis. A marked protuberance in 15% of the population is also noted in limited specimens within the South Australian Museum collections and appears to be present only in juvenile D. costata. Both the abundance and narrow size range of this population support the notion that Dickinsonia was a hardy opportunist, capable of rapid establishment and growth on relatively immature textured organic-mat substrates.

The Ediacaran to lower Cambrian Chilhowee Group of the southern and central Appalachians records the rift-to-drift transition of the newly formed Iapetan margin of Laurentia. Body fossils are rare within the Chilhowee Group, and correlations are based almost exclusively on lithological similarities. A critical review of previous work highlights the relatively weak biostratigraphic and radiometric age constraints on the various units within the succession. Herein, we document a newly discovered fossil-bearing locality within the Murray Shale (upper Chilhowee Group) on Chilhowee Mountain, eastern Tennessee, and formally describe a nevadioid trilobite, Buenellus chilhoweensis n. sp., from that site. This trilobite indicates that the Murray Shale is of Montezuman age (provisional Cambrian Stage 3), which is older than the Dyeran (provisional late Stage 3 to early Stage 4) age suggested by the historical (mis)identification of “Olenellus sp.” from within the unit as reported by workers more than a century ago. Buenellus chilhoweensis n. sp. represents only the second known species of Buenellus, and demonstrates that the genus occupied both the Innuitian and Iapetan margins of Laurentia during the Montezuman. It is the oldest known trilobite from the Iapetan margin, and proves that the hitherto apparent absence of trilobites from that margin during the Montezuman was an artifact of inadequate sampling rather than a paleobiogeographic curiosity. The species offers a valuable biostratigraphic calibration point within a rock succession that has otherwise proven recalcitrant to refined dating.

UUID: http://zoobank.org/30af790b-e7b1-44c3-b1d5-55cdf579bde2

Exquisitely preserved three-dimensional examples of the classic Ediacaran (late Neoproterozoic; 570–541 Ma) frond Charniodiscus arboreus Jenkins and Gehling, 1978 (herein referred to as Arborea arborea Glaessner in Glaessner and Daily, 1959) are reported from the Ediacara Member, Rawnsley Quartzite of South Australia, and allow for a detailed reinterpretation of its functional morphology and taxonomy. New specimens cast in three dimensions within sandy event beds showcase detailed branching morphology that highlights possible internal features that are strikingly different from rangeomorph and erniettomorph fronds. Combined with dozens of well-preserved two-dimensional impressions from the Flinders Ranges of South Australia, morphological variations within the traditional Arborea morphotype are interpreted as representing various stages of external molding. In rare cases, taphomorphs (morphological variants attributable to preservation) represent composite molding of internal features consisting of structural supports or anchoring sites for branching structures. Each primary branch consists of a central primary branching stalk from which emerge several oval secondary branches, which likely correspond to similar structures found in rare two-dimensional specimens. Considering this new evidence, previous synonymies within the Arboreomorpha are no longer justified, and we suggest that the taxonomy of the group be revised.