The Ao Mo Lae Formation of the Tarutao Group crops out on Thailand's Tarutao Island and contains a diverse assemblage of late Furongian trilobite taxa, including several endemic forms. This study presents a new genus and species, Satunarcus molaensis, discovered at two locations on the island. A cladistic analysis of the kaolishaniid subfamily Mansuyiinae in light of Satunarcus and similar genera known from across upper Cambrian equatorial Gondwanan rocks suggests that the subfamily is polyphyletic in its current definition, and thus is not a natural group. Separating Mansuyia Sun, 1924 from the other taxa conventionally placed in Mansuyiinae permits recognition of a previously unrecognized monophyletic subfamily Ceronocarinae new subfamily. As established herein, this kaolishaniid subfamily contains Satunarcus n. gen. and all genera previously recognized as Mansuyiinae. with the exception of Mansuyia itself. Ceronocarinae n. subfam. occur in middle Jiangshanian to middle Cambrian Stage 10 sedimentary rocks from Australia, South China, North China, and Sibumasu, with most genera endemic to Australia.

UUID: http://zoobank.org/618c5136-73f0-4912-a7d3-e56559d2a76c

Ptychites is among the most widely distributed ammonoid genera of the Triassic and is namesake of a family and superfamily. However, representatives of the genus mostly show low-level phenotypic disparity. Furthermore, a large number of taxa are based on only a few poorly preserved specimens, creating challenges to determine ptychitid taxonomy. Consequently, a novel approach is needed to improve ptychitid diversity studies. Here, we investigate Ptychites spp. from the middle and late Anisian of Nevada. The species recorded include Ptychites embreei n. sp., which is distinguished by an average conch diameter that is much smaller and shows a more evolute coiling than most of its relatives. The new species ranges from the Gymnotoceras mimetus to the Gymnotoceras rotelliformis zones, which makes it the longest-ranging species of the genus. For the first time, the ontogenetic development of Ptychites was obtained from cross sections where possible. Cross-sectioning highlights unique ontogenetic trajectories in ptychitids. This demonstrates that, despite showing little phenotypic disparity, Ptychites was highly ontogenetically differentiated, and thus the high taxonomic diversity at the species level is justified for the species investigated.

UUID: http://zoobank.org/5abe2487-8a00-4b48-adc7-ec7db7a097f7

The fossil record of the family Camerobiidae has been represented by only one species, Neophyllobius succineus Bolland and Magowski, 1990, described from Eocene Baltic amber. These prostigmatan mites are distinguishable by their distinctly long and slender stilt-like legs, and they are associated with aboveground vegetation where they hunt for other small invertebrates. This paper enhances the knowledge of fossil stilt-legged mites. Two new fossil species, N. electrus new species and N. glaesus new species, are described from samples of Baltic amber, and remarks on their morphology and taphonomy are provided. The discovery is complemented with a discussion on morphological singularities (the shape of the prodorsum, the location of setae h1 and h2 in living specimens, and lengths of genual setae), an anomaly of hypertrophied seta (found in the N. glaesus holotype), and some biogeographical issues.

UUID: http://zoobank.org/d1602384-ae4f-4f90-b4a1-6cdedd77c9e1

The recent description of the nevadioid trilobite Buenellus chilhoweensis Webster and Hageman, 2018 established the presence of early Cambrian Montezuman Stage (Cambrian Series 2, Stage 3) faunas in the Murray Shale of Chilhowee Mountain, Tennessee. The description recognized the oldest known age-diagnostic Cambrian trilobite from the Laurentian margin of the former Iapetus Ocean since Buenellus Blaker, 1988 is known otherwise only from the Sirius Passet Lagerstätte on the Innuitian margin of North Greenland. The bivalved arthropods Isoxys chilhoweanus Walcott, 1890 and Indota tennesseensis (Resser, 1938a) have also been described from the Murray Shale, but hyolithids appear to be the dominant body fossils in terms of diversity and abundance. Although poorly preserved, the hyolithids occurring together with Buenellus chilhoweensis are described to improve understanding of the Murray Shale biota. The hyolith assemblages of the Murray Shale and Sirius Passet Lagerstätte are not closely similar, although the poor preservation of both hinders comparison.

A new, previously undescribed Middle Ordovician (middle Darriwilian: Dw2) radiolarian assemblage has been recovered from the Table Cove Formation at Piccadilly Quarry, western Newfoundland. Constituents of the fauna described herein are both distinctive and exceptionally well preserved. Three-dimensional X-ray micro-computed tomography (μ-CT) is used to make a detailed examination of four key spumellarian specimens. This technology enables visualization of hitherto ambiguous details of the internal morphologies of key lower Paleozoic taxonomic groups, among which a lack of knowledge has impeded resolution of higher taxonomic rankings.

In contrast to almost all other invertebrate phyla that constructed biomineralized skeletons during the “Cambrian explosion” and maintained them during the entire fossil record, ascidian tunicates evolved this protective and stabilizing advantage only during the Permian, although soft-bodied representatives of this subphylum made their first appearance already in the early Cambrian. It remains enigmatic why these compound calcareous skeletons persisted only until the Late Triassic, subsequently followed by less-rigid internal skeletons from the Lower Jurassic onwards, which consist of scattered isolated spicules only. In addition to recently described aragonitic ascidian exoskeletons from the Permian and Triassic, new discoveries of similar, but colonial ascidian compound endoskeletons in the lower Carnian exhibit a short-living branch of this group, which moreover contain the first indubitable calcareous spicules. The latter are embedded in the solid endoskeleton, which is composed of polygonal aragonitic plates with smooth outer and zigzag lined inner boundaries. They consist of irregular, parallel (orthogonal), or fan-shaped (clinogonal) arrangements of acicular aragonite crystals. The following taxa are described as new: order Cassianomorpha new order with the family Cassianosomidae new family and the genus Toscanisoma new genus with the species T. multipartitum new species and T. triplicatum new species.

UUID: http://zoobank.org/03555353-cdab-42e8-8e99-9bfce15fa249

Cardichelyon rogerwoodi is an enigmatic fossil turtle from the late Paleocene to early Eocene of North America. Previous analyses suggested affiliation with Testudinoidea, in particular the big-headed turtle Platysternon megacephalum, based on the presence of multiple musk-duct foramina and a large head. We here highlight previously undocumented characteristics for this turtle, notably the presence of short costiform processes, a rib-like axillary process, and a posterior plastral hinge. Phylogenetic analysis places Cardichelyon rogerwoodi within Testudinoidea, but the exclusion of testudinoids suggest an affiliation with Dermatemydidae. Using consilience with external data we favor placement within Kinosternoidea. Cardichelyon rogerwoodi is therefore an aberrant, hinged kinosternoid that developed in situ in North America during the Paleocene long before the arrival of testudinoids on this continent in the early Eocene.

Intermediate morphologies of a new fossil crinoid shed light on the pathway by which crinoids acquired their distinctive arms. Apomorphies originating deep in echinoderm history among early nonblastozoan pentaradiate echinoderms distinguish Tremadocian (earliest Ordovician) crinoid arms from later taxa. The brachial series is separated from the ambulacra, part of the axial skeleton, by lateral plate fields. Cover plates are arrayed in two tiers, and floor plates expressed podial basins and pores. Later during the Early Ordovician, floor plates contacted and nestled into brachials, then were unexpressed as stereom elements entirely and cover plates were reduced to a single tier. Incorporation of these events into a parsimony analysis supports crinoid origin deep in echinoderm history separate from blastozoans (eocrinoids, ‘cystoids’). Arm morphology is exceptionally well-preserved in the late Tremadocian to early Floian Athenacrinus broweri new genus new species. Character analysis supports a hypothesis that this taxon originated early within in the disparid clade. Athenacrinus n. gen. (in Athenacrinidae new family) is the earliest-known crinoid to express what is commonly referred to as ‘compound’ or ‘biradial’ morphology. This terminology is misleading in that no evidence for implied fusion or fission of radials exists, rather it is suggested that this condition arose through disproportionate growth.

UUID: http://zoobank.org/b383e039-3298-4472-a7e3-e81684f87cfe

Oryctocephalid trilobites are seldom abundant and often tectonically deformed, creating problems for robust species delimitation and compromising their utility in biostratigraphic and evolutionary studies. By studying more than 140 specimens recovered from the upper portion of the Combined Metals Member (Pioche Formation, Nevada; Cambrian Stage 4, Series 2), we exploit a rare opportunity to explore how morphological variation among oryctocephalid specimens is partitioned into intraspecific variation versus interspecific disparity. Qualitative and quantitative analyses reveal that two species are represented: Oryctocephalites palmeri Sundberg and McCollum, 1997 and Oryctocephalites sp. A, the latter known from a single cranidium stratigraphically below all occurrences of the former. In contrast to the conclusions of a previous study, there is no evidence of cranidial dimorphism in O. palmeri. However, that species exhibits considerable variation in cranidial shape and pygidial spine arrangement and number. Cranidial shape variation within O. palmeri is approximately one-half of the among-species disparity within the genus. Comparison of cranidial shape between noncompacted and compacted samples reveals that compaction causes significant change in mean shape and an increase in shape variation; such changes are interpretable in terms of observed fracture patterns. Nontaphonomic variation is partitioned into ontogenetic and nonallometric components. Those components share similar structure with each other and with interspecific disparity, suggesting that ontogenetic shape change might be an important source of variation available for selection. This highlights the importance of ontogenetic and taphonomic sources of variation with respect to species delimitation, morphospace occupation, and investigation of evolutionary patterns and processes.

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.

UUID: http://zoobank.org/54a906b6-4873-4f84-92b5-ca0752de01aa

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