Eight bryozoan species are described from the Hanchiatien Formation (lower Silurian, Telychian) of southern Chongqing, South China. Four species are new: the trepostomes Asperopora sinensis n. sp., Trematopora jiebeiensis n. sp., and Trematopora tenuis n. sp., and the fenestrate Moorephylloporina parvula n. sp. One species, the cystoporate Hennigopora sp. indet., is described in open nomenclature. Moorephylloporina parvula n. sp. is eurytopic, occurring in all types of facies within the bioherms. Erect Moorephylloporina Bassler, 1952, Trematopora Hall, 1852, and Leioclema Ulrich, 1882 formed pioneering communities on weakly cemented substrata, whereas encrusting Fistulipora M‘Coy, 1849, Hennigopora Bassler, 1952, and Asperopora Owen, 1969 occurred on hardgrounds and formed densely compact framestones. Robust branched Trematopora and Leioclema tend to occur out of the reef core (framework) where they could have formed reef-flank thickets in more agitated conditions. The generic composition of the studied fauna correlates with other localities in South China, and they show general paleobiogeographic relations to Siberia and Indiana, USA.

UUID http://zoobank.org/3326dd2f-7c9e-43bc-9dab-84047b274f89

In order to maximize the utility of future studies of trilobite ontogeny, we propose a set of standard practices that relate to the collection, nomenclature, description, depiction, and interpretation of ontogenetic series inferred from articulated specimens belonging to individual species. In some cases, these suggestions may also apply to ontogenetic studies of other fossilized taxa.

A new sinistrally coiled univalved mollusk Catalanispira n. gen. is described with two species; Catalanispira reinwaldti (Öpik, 1930) from the Middle Ordovician Kõgekallas Formation (Darriwilian) of Estonia and Catalanispira plattevillensis n. gen. n. sp. from the Upper Ordovician Platteville Formation (Sandbian) of northern Illinois, USA. Morphological features include a large, low-trochiform shell, a narrow lenticular aperture, a deep funnel-like umbilicus, a falcate inner lip and a large (1.4 mm wide) protoconch. Ornamentation consists of fine commarginal growth lines or ribs but superimposed on a slightly irregular shell surface. Catalanispira n. gen. is placed within the sinistrally coiled order Mimospirida and the family Onychochilidae, and Catalanispirinae n. subfam. is proposed. The large Lower Ordovician (Tremadocian) Pelecyogyra Ebbestad and Lefebvre, 2015 from Morocco and France is transferred to this new subfamily. The well-preserved initial growth stage of Catalanispira plattevillensis n. gen. n. sp. is cap-shaped, slightly asymmetrical, unusually large, and smooth, and represents either an unusually large embryonic shell (protoconch 1) or a larval shell (protoconch 2). It differs from the smaller protoconch described for the clisospirine Mimospira Koken in Koken and Perner, 1925, which might include a multiwhorled larval shell (protoconch 2). Mimospirids are dominantly Ordovician, and have been classified as untorted mollusks (only distantly related to gastropods), dextral hyperstrophic gastropods, or sinistral orthostrophic gastropods. Sinistral asymmetry already in the embryonic shell and lack of conclusive evidence for coiling direction, e.g., an operculum, could suggest that Catalanispira n. gen. or similar mimosprids were sinistral orthostrophic gastropods. Currently the group is therefore classified as a group of sinistral orthostrophic gastropods, unranked within the Gastropoda.

UUID: http://zoobank.org/affc8dcf-4c0f-493d-bee5-75a457996e84

The Lakeview Limestone is one of the westernmost Cambrian exposures in the northwestern United States and occurs on the western edge of the Montania paleotopographic high. These deposits occur between the deeper water deposits to the west and carbonate banks and intracratonic basins to the east and provide critical link between the regions. A re-investigation of the Cambrian trilobite faunas from the lower portion of the Lakeview Limestone, Pend Oreille Lake, Idaho, is undertaken due to the inadequate illustrations and descriptions provided by Resser (1938a). Resser's type specimens and additional material are figured and described. The trilobite assemblages represent the Ptychagnostus praecurrens Zone, Wuliuan Stage, Miaolingian Series and including two new taxa: Itagnostus idahoensis n. sp., and Utia debra n. sp. Because of the similarity between some species of Amecephalus from the Lakeview Limestone to specimens from the Chisholm Shale, Nevada, the type specimens of Amecephalus piochensis (Walcott, 1886) and Am. packi (Resser, 1935), Walcott's and Resser's type specimens are re-illustrate and their taxonomic problems are discussed. Utia curio Walcott, 1924 from the Spence Shale, Utah, have never been re-illustrated since Walcott (1925), these type specimens are also re-illustrated and compared to Utia debra n. sp.

UUID: http://zoobank.org/5ef6ee5e-bddc-4395-becf-9bcea54c679c

Bituminous limestone of the Ediacaran Shibantan Member of the Dengying Formation (551–539 Ma) in the Yangtze Gorges area contains a rare carbonate-hosted Ediacara-type macrofossil assemblage. This assemblage is dominated by the tubular fossil Wutubus Chen et al., 2014 and discoidal fossils, e.g., Hiemalora Fedonkin, 1982 and Aspidella Billings, 1872, but frondose organisms such as Charnia Ford, 1958, Rangea Gürich, 1929, and Arborea Glaessner and Wade, 1966 are also present. Herein, we report four species of Arborea from the Shibantan assemblage, including the type species Arborea arborea (Glaessner in Glaessner and Daily, 1959) Glaessner and Wade, 1966, Arborea denticulata new species, and two unnamed species, Arborea sp. A and Arborea sp. B. Arborea arborea is the most abundant frond in the Shibantan assemblage. Arborea denticulata n. sp. resembles Arborea arborea in general morphology but differs in its fewer primary branches and lower length/width ratio of primary branches. Arborea sp. A and Arborea sp. B are fronds with a Hiemalora-type basal attachment. Sealing by microbial mats and authigenic cementation may have played an important role in the preservation of Arborea in the Shibantan assemblage. The Shibantan material of Arborea extends the stratigraphic, ecological, and taphonomic ranges of this genus.

UUID: http://zoobank.org/554f21da-5f09-4891-9deb-cbc00c41e5f1

Dilophosaurus wetherilli was the largest animal known to have lived on land in North America during the Early Jurassic. Despite its charismatic presence in pop culture and dinosaurian phylogenetic analyses, major aspects of the skeletal anatomy, taxonomy, ontogeny, and evolutionary relationships of this dinosaur remain unknown. Skeletons of this species were collected from the middle and lower part of the Kayenta Formation in the Navajo Nation in northern Arizona. Redescription of the holotype, referred, and previously undescribed specimens of Dilophosaurus wetherilli supports the existence of a single species of crested, large-bodied theropod in the Kayenta Formation. The parasagittal nasolacrimal crests are uniquely constructed by a small ridge on the nasal process of the premaxilla, dorsoventrally expanded nasal, and tall lacrimal that includes a posterior process behind the eye. The cervical vertebrae exhibit serial variation within the posterior centrodiapophyseal lamina, which bifurcates and reunites down the neck. Iterative specimen-based phylogenetic analyses result in each of the additional specimens recovered as the sister taxon to the holotype. When all five specimens are included in an analysis, they form a monophyletic clade that supports the monotypy of the genus. Dilophosaurus wetherilli is not recovered as a ceratosaur or coelophysoid, but is instead a non-averostran neotheropod in a grade with other stem-averostrans such as Cryolophosaurus ellioti and Zupaysaurus rougieri. We did not recover a monophyletic ‘Dilophosauridae.’ Instead of being apomorphic for a small clade of early theropods, it is more likely that elaboration of the nasals and lacrimals of stem-averostrans is plesiomorphically present in early ceratosaurs and tetanurans that share those features. Many characters of the axial skeleton of Dilophosaurus wetherilli are derived compared to Late Triassic theropods and may be associated with macropredation and an increase in body size in Theropoda across the Triassic-Jurassic boundary.

A new species of microconchid tubeworm, Microconchus cravenensis is described from the Mississippian Cracoean reefs of North Yorkshire, United Kingdom. Despite the fact that some other microconchid species could have attained large tube length, the new species possesses the largest recorded diameter (to 7.7 mm) of the planispirally-coiled (attachment) tube and the largest recorded aperture diameter (8.3 mm) in the helically uncoiled portion. Thus, with respect to these features, Microconchus cravenensis n. sp. is the largest and most robust microconchid species recognized so far. At present, it is only known from the Craven Reef Belt of North Yorkshire, where it attached to corals and possibly bivalve shells, and was preyed upon by small durophagous animals, as indicated by repaired injuries preserved on one of the tubes.

UUID: http://zoobank.org/2ba8f87b-ec1c-4bb3-8615-115e7a527376

Lovén's Law described the position of larger basicoronal ambulacral plates in echinoids. The smaller basicoronal plates form first in ontogeny. We restate Lovén's Law to describe the position of first ambulacral plates using Carpenter's ambulacra and left (L) or right (R) as: AR, BL, CR, DL, ER, with EA the pair of ambulacra both identical and adjacent. First ambulacral plates of the Cambrian edrioasteroid, Walcottidiscus, code identically. The transition from a tri-radiate 1-1-1 pattern to the 2-1-2 ambulacral pattern of Walcottidiscus and other pentaradiate Paleozoic echinoderms results in Lovén's Law. This provides an hypothesis for the origin of Lovén's Law and predicts its widespread occurrence among echinoderm classes.

The ‘BD different’ pattern of primary brachioles in pentaradiate glyptocystitoid Rhombifera results from subterminal branching of the ambulacra. The ontogenetic sequence was triradiate, then lateral ambulacra bifurcated, and finally second brachioles developed. Positions of second brachioles of pentaradiate glyptocystitoids code as AR, BL, CR, DL, ER. Other examples of Lovén's Law occur in Diploporita, Ophiuroidea, Edrioasteroidea, Edrioblastoidea, Holothuroidea, Ophiocistioidea, and Bothriocidaris. Our working hypothesis is that Lovén's Law arose with pentaradiality. An objective test requires a reliable independent landmark across all classes, which does not exist currently. Our hypothesis is only parsimonious if Lovén's Law arose just once in echinoderm evolution and is widespread within the phylum. So far, both conditions appear to be met, but edrioasteroid first ambulacral cover plates commonly exhibit an alternative Lovén-like pattern of AL, BL, CR, DL, ER, with AB the identical adjacent ambulacra.

UUID: http://zoobank.org/4c40d338-8bfa-4515-9037-934f3bbab153

A locality in the Upper Cretaceous Two Medicine Formation of Montana preserves abundant and variable horseshoe crab tracks and trails of the ichnotaxon Kouphichnium isp. These specimens span six morphologies differing in track form and trail configuration. These differences likely reflect variations in track-maker locomotion and behavior, substrate consistency, epichnial versus hypichnial preservation, and undertrack versus true tracks. Several tracks preserve the first clear appendage impressions for an extinct horseshoe crab. This discovery adds new information to the fossil horseshoe crab diversity in the Cretaceous Period. Trackway dimensions, such as the external width across the pusher legs or of the prosomal drag mark, provide information on the track-maker size. Most trackways correspond with crabs 9–14 cm wide; the abundance but limited size range of the traces suggests the large assemblage corresponds to a mating aggregation. The trace fossil record of xiphosurids indicates that throughout their history, horseshoe crabs inhabited both marine and nonmarine settings. They were definitively present in freshwater habitats from the lower Carboniferous through at least the Paleogene. Horseshoe crab trace abundance is highest from the upper Carboniferous through the Jurassic and likely reflects two factors: true upper Carboniferous taxonomic diversity and a preponderance of suitable sites for trackway preservation in the Late Triassic and Jurassic. Cretaceous traces are uncommon, and this Two Medicine locality is the first occurrence of horseshoe crab traces in the Late Cretaceous worldwide. Overall, track abundance and diversity would seem to correspond well with the reported horseshoe crab body fossil record.

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.