Computed tomographic (CT) imaging allows new accessibility to shells of gastropod fossil taxa and their extant relatives, providing new data for interpreting former systematic assignments. The highly questionable ellobiid assignment of the nonmarine gastropod genus Protocarychium Pan, 1982 from the Lower Jurassic of Hunan, China, is reevaluated using CT imaging to assess internal aspects of the shell. By comparing these new data to those of stylommatophoran, ellobiid, and caenogastropod clades in the literature, this work reveals that Protocarychium bears no affinity to the Carychiidae, which are otherwise known only from the Cenozoic, but rather to the Paleozoic land snail family Anthracopupidae Wenz, 1938. This finding constitutes the first Asian appearance of anthracopupid snails beyond their known North American and European range. Contrary to the current opinion, we suggest the Anthracopupidae to be a basal stylommatophoran clade, which places the origin of Stylommatophora at least in the late Carboniferous.

The hurdiid radiodont Cambroraster is characterized by a prominent horseshoe-shaped head carapace, which bears conspicuous posterolateral spinous processes and covers half of the body length, and a pair of frontal appendages possessing rake-like endites. Here we report the discovery of characteristic central and lateral elements as well as a possible appendage endite for Cambroraster from the Cambrian Wuliuan Mantou Formation of North China. This discovery suggests that Cambroraster had dispersed over a wide biogeographic range and expands the geographic range of Hurdiidae. The increasing discoveries of exceptionally preserved fossils in the Upper Shale Member of the Mantou Formation indicate that this horizon is a promising Burgess Shale–type Lagerstätte.

The small, Lower Cretaceous, iron ore deposit at Zengővárkony (Mecsek Mountains, southern Hungary, Europe) contains new brachiopod taxa of kingenoid relationships. Dictyothyropsis vogli, Zittelina hofmanni, and Smirnovina ferraria are described as new species from late Valanginian to earliest Hauterivian strata. The new taxa strengthen the presence of Early Cretaceous biogeographical connections with the Western Carpathians and the Pieniny Klippen Belt of southern Poland. The newly described taxa have significantly larger dimensions than their closest relatives from the type localities, which is in line with previous research on brachiopods from this environment. These brachiopods lived in a nutrient-rich, unique environment related to iron-ore deposition linked to former hydrothermal activity on the seafloor that might have contributed to the large size of these brachiopods. Larger than normal rhynchonellide and terebratulidine brachiopods have previously been recorded from this locality.

UUID: http://zoobank.org/353882-2838-4eb7-b21a-8cc665d13408

The first fossil rove beetle (Coleoptera, Staphylinidae) from New Zealand is described from the earliest Miocene Foulden Maar fossil-Lagerstätte, Otago. The new species, Sphingoquedius meto n. sp., is attributable to the tribe Amblyopinini of the subfamily Staphylininae based on the scutellum with anterior scutellar ridge only; isodiametric microsculpture on the pronotum; multidirectional arranged setae on the elytra; and presence of radiating setae on the fourth abdominal tergite. Sphingoquedius meto n. sp. is the first Southern Hemisphere fossil record of Amblyopinini and its affinity to the extant fauna as well as biogeographic and paleocological implications are discussed.

UUID: http://zoobank.org/52275b83-d5ff-48af-A66F-2d62c3461677

The Miocene Solemyidae of Chile are revised. The holotype of Solemya antarctica Philippi, 1887, originally described as Solenomya, is lost. Due to the lack of information on internal characters, its systematic position is considered as uncertain. A new species, Solemya lucifuga n. sp., is described from the lower Miocene Ranquil Formation of south-central Chile. Its dense radial external ornamentation shows that it is clearly different from S. antarctica. It is currently the only confirmed and described Miocene solemyid bivalve from the Southern Hemisphere.

UUID: http://zoobank.org/a35ee02e-600b-4e9f-ad06-1887e9493ff8

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 order Intejocerida is an enigmatic, short-lived cephalopod taxon known previously only from Early–Middle Ordovician beds of Siberia and the United States. Here we report a new genus, Cabaneroceras, and a new species, C. aznari, from Middle Ordovician strata of central Spain. This finding widens the paleogeographic range of the order toward high-paleolatitudinal areas of peri-Gondwana. A curved conch, characteristic for the new genus, was previously unknown from members of the Intejocerida.

UUID: http://zoobank.org/21f0a09c-5265-4d29-824b-6b105d36b791

Nectocaridids are soft-bodied Cambrian organisms that have been controversially interpreted as primitive cephalopods, at odds with the long-held belief that these mollusks evolved from a shell-bearing ancestor. Here, I document a new nectocaridid from the Whetstone Gulf Formation, extending the group's range into the Late Ordovician. Nectocotis rusmithi n. gen. n. sp. possesses a robust internal element that resembles a non-mineralized phragmocone or gladius. Nectocaridids can be accommodated in the cephalopod total group if the earliest cephalopods (1) inherited a non-mineralized shell field from the ancestral mollusk; and (2) internalized this shell field. This evolutionary scenario would overturn the traditional ectocochleate, Nautilus-like reconstruction of the ancestral cephalopod, and indicate a trend towards increased metabolic efficiency through the course of Cambrian–Ordovician evolution.

UUID: http://zoobank.org/ed594200-37b9-4642-bd8f-4fb72dc544eb

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

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

The Veneridae are the most speciose modern family of bivalves, and one of the most morphologically conservative and homoplastic, making subfamily- and sometimes even genus-level classification difficult. The widespread Cretaceous genus Legumen Conrad, 1858 is currently placed in the subfamily Tapetinae of the Veneridae, although it more closely resembles the Solenoida (razor clams, Pharidae and Solenidae) in general shell form. Here we provide high-resolution images of the Legumen hinge for the first time. We confirm from hinge morphology that Legumen belongs in Veneridae, but it should be referred to incertae subfamiliae, rather than retained in the Tapetinae, particularly in light of the incomplete and unstable understanding of venerid systematics. Legumen represents a unique hinge dentition and a shell form—and associated life habit—that is absent in the modern Veneridae despite their taxonomic diversity. Veneridae are hyperdiverse in the modern fauna, but strikingly ‘under-disparate,’ having lost forms while gaining species in the long recovery from the end-Cretaceous extinction.

The Brechin Lagerstätte of southern Ontario contains an exceptionally diverse and well-preserved Late Ordovician (Katian) crinoid fauna. We describe four genera and eight species of camerate crinoids from the Brechin Lagerstätte, including six new species. Consequently, the total diversity of the fauna now stands at 27 genera and 39 nominal species, thereby making it the most taxonomically diverse Ordovician crinoid fauna known. Taxa described include the diplobathrid Pararchaeocrinus kiddi new species and the monobathrids Glyptocrinus ramulosus Billings, 1856, Periglyptocrinus priscus (Billings, 1857a), Periglyptocrinus astricus new species, Periglyptocrinus kevinbretti new species, Periglyptocrinus mcdonaldi new species, Periglyptocrinus silvosus new species, and Abludoglyptocrinus steinheimerae new species. We summarize the taxonomic composition, diversity, and abundance distribution of all known crinoids from the Brechin Lagerstätte to better characterize the paleoecological structure and complexity of the community. We establish that the fauna is dominated by the subclass Pentacrinoidea, both in terms of abundance and species richness. In addition, we analyze species-level abundance data using Relative Abundance Distribution (RAD) models to evaluate the ecological complexity of the paleocommunity. We found that community structure of the Brechin Lagerstätte is best explained by an ecologically ‘complex’ RAD model, which suggests that species partitioned niches along multiple resource axes and/or the presence of multiple ecological ways of life. These results indicate that the Brechin Lagerstätte is significant not only for being the most taxonomically diverse Katian crinoid assemblage, but also for being an early ecologically complex fauna that developed in the wake of the Great Ordovician Biodiversification Event.

UUID: http://zoobank.org/f86582ed-5db6-469E-befe-34b801f9a113

Gonatocaris decora (Clarke, 1901) is an unusual, highly ornamented Silurian phyllocarid from the Pittsford Shale Member of the Vernon Formation, Salina Group of New York, that has been allied with Emmelezoe in the past due to the possession of an ‘optic tubercle.’ The systematic positions of Gonatocaris and Emmelezoe have been difficult to assess due to the lack of complete remains, attributions based on spurious characters, and missing and unfigured specimens. Relatively recent collections of Gonatocaris decora, along with a relocation of the original type material of Emmelezoe, have allowed a reexamination of all relevant genera and a more robust systematic placement for G. decora. Characters of generic importance for Gonatocaris include a greatly elongated abdomen; an acute angle of the ventral carapace margin separating distinct anteroventral and posteroventral margins from each other; and distinctive, multiscale horizontal ridge-like ornament of the carapace valves. Growth was found to be isometric. Experimental work done using a model of the scale-like ornament and multiple grain sizes of substrate indicates that cuticle ornament may have served to increase friction between the animal and a fine sand substrate, although resistance to predation is the currently favored hypothesis. Because Gonatocaris species are currently known only from the eastern margin of Laurentia and northern South China, it is likely that this genus was nektonic. Accumulations of similarly sized carapaces and other disarticulated sclerites in shallow-water settings may indicate spawning events following molting en masse.

The presence of a new taxon, Duraznovis gallegoi new genus new species is reported and described from an early Late Triassic (Carnian) deposit in Argentina. Two specimens, recovered from the Quebrada del Durazno locality, uppermost levels of the Potrerillos Formation, Cuyana Basin (Mendoza), are represented by the molds of their shield and imprints of soft parts. The identity of these specimens appears enigmatic but closely resembles in the possession of a generalized arthropod morphology and a distinctive combination of characters, to living and fossil representatives of xiphosurans (Chelicerata) and notostracans (Branchiopoda). The new fossils are associated with a rich biota comprising abundant insects, spinicaudatans, plants, and scarce fish remains living in semipermanent swamps and/or ponds within a delta plain environment with intermittent episodes of flooding, in a warm temperate and humid megamonsoonal climate during Triassic times. In this context, we analyze the taphonomic and ecological implications of their presence. Lastly, these unique specimens at the Quebrada del Durazno locality adds to the diversity of the biota, revealing the importance of this site as an exceptional paleontological Triassic deposit.

UUID: http://zoobank.org/8d1194c2-170c-4b7a-b0c6-5c08b0974d95

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.

Two new isocrinids are described from the Lower Cretaceous Agrio Formation of the Neuquén Basin, west-central Argentina. Isocrinus (Chladocrinus) covuncoensis new species is based on several beautifully preserved specimens from Valanginian beds of the Pilmatué Member. It is characterized by a small size, multiramose crown with six arm divisions, 240 arm tips, mostly isotomous branching, seven (or rarely eight) secundibrachials, smooth and stout column, short noditaxis, and pentalobate columnals. The species occurs in a 30 m thick interval of cross-bedded sandstones and mixed clastic-carbonate sediments that represent the migration of large, tidally influenced, subaqueous dunes developed in the offshore. Sudden burial of crinoids that dwelled on the dune toes and interdunes, possibly by the acceleration of the lee face migration, provided the exceptional preservation of specimens and thus this finding can be considered as a local crinoid Konservat Lagerstätte. Isocrinus (Chladocrinus) pehuenchensis new species is described from a single articulated specimen preserved in a silty calcareous concretion collected from a late Hauterivian concretion level of the Agua de la Mula Member. It is characterized by isotomous branching, eight or nine secundibrachials (IIBr), slender column ornamented with medial ridge of fine tubercles, interradius acuminated with fine tubercles on its tip, short noditaxis, and pentastellate columnals. It is associated with low-energy fall-out deposits in the offshore. The excellent state of preservation was due to an early cementation process by carbonate that enhanced lithification around the specimen.

UUID: http://zoobank.org/4763ae70-9d0a-4015-b1c9-6a7ceeedfb1a

The homalonotid trilobite Burmeisteria Salter, 1865 is revised from material from the Lower Devonian of central western Argentina (the Talacasto Formation in the Precordillera Basin). In contrast to other closely related Devonian marine basins from South America (mainly Bolivia, Brazil and Uruguay), the only species recognized in Argentina include Burmeisteria herschelii (Murchison, 1839) and B. notica (Clarke, 1913). New observations on the structure of the carapace indicate that structures sometimes interpreted as granules with taxonomic meaning are, in fact, taphonomic characters that represent the fillings of pore canals exposed by decortication. The antero-ventral process of the rostral plate may be a locking device during enrollment, which allowed long-term defensive behavior with a minimum of muscular energy. Burmeisteria is an upper Silurian to Middle Devonian endemic representative of southwestern Gondwanan (Malvinokaffric) basins. In the Argentine Precordillera Basin, this genus is virtually restricted to a sandy, Pragian, stratigraphic interval capped by a marker bed first reported by Keidel in 1921.

Family-level taxonomic concepts for Paleozoic Asteroidea are problematic; data availability for the family Urasterellidae, however, is judged sufficient for familial reappraisal. The survey cannot be fully comprehensive because certain available names are founded on very limited material, with specimens widely scattered among collections. Recognized urasterellid genera include Urasterella Schuchert, 1914; Anthroosasterias Blake, 2017; Meturaster Blake, Guensburg, and Lefebvre, 2016; Salteraster Stürtz, 1893; Staurasterias Blake, 2017; Ulrichaster Spencer, 1950; and Pegoasterella pompom n. gen. n. sp. Other available generic names are held in abeyance. The name Urasterella has approached form-genus status, an outcome exacerbated by incomplete understanding of the type species. Urasterella huxleyi (Billings, 1865) is reassigned to Anthroosasterias. Relatively large specimen suites for Anthroosasterias huxleyi (Billings, 1865) and Salteraster grandis (Meek, 1872) enable some interpretation of urasterellid intrataxon variation, ontogenetic change, and life mode. Morphologic changes accompanying ontogeny of A. huxleyi are argued as favoring a shift from substrate surface feeding to feeding through extension of arms into the water column in a manner comparable to that seen among extant Forcipulatida and ophiurans.

UUID: http://zoobank.org/82d2eca7-55e0-42ed-a164-c666652e3972

Aerliceaster nexosus n. gen. n. sp. (Echinodermata), one of the oldest of known asteroids, is based on six specimens from the Floian (Early Ordovician) Garden City Formation of Idaho, and Kolataster perplexus n. gen. n. sp. is based on two specimens from the Sandian (Late Ordovician) Mifflin Formation of Illinois. Although the asterozoan skeleton is subdivided into few ossicular categories, evolutionary derivations of all the categories are not fully established, and therefore published evaluations differ. Beginning with phylogenetic work placing asteroid ancestry within the Somasteroidea together with the new taxa described herein, aspects of early asteroid morphology are evaluated and ambiguities in need of further study identified. Uncertainties are considered to be founded in rapid early asterozoan diversification and the scanty fossil record.

UUID: http://zoobank.org/b43d07cc-c8fb-4a84-bc6f-40aa6e0daea2

A new trace fossil, Feoichnus martini new ichnospecies, from the Two Medicine Formation is here described. This ichnotaxon is reported from the upper Campanian deposits of the Egg Mountain locality (Montana) and consists of a hemispherical to hemiellipsoidal structure with a truncated upper edge, and a regular, rounded lower edge marked by a lined border composed of stained layers. The trace maker likely impregnated the border using organic fluids. The simple lined wall observed in F. martini n. isp. suggests that the structure was produced by an invertebrate soil-dwelling organism, likely an insect. Specimens are preserved as casts in calcitic Inseptisols alongside an abundant vertebrate fossil record composed by dinosaurian and nondinosaurian fossil remains, dinosaur eggs and nest structures, and pervasive insect bioturbation. Feoichnus martini n. isp. represents an additional, minor component of the impoverished Celliforma ichnofacies reported at Egg Mountain and expands the paleogeographical distribution of the ichnogenus Feoichnus Krause et al., 2008 to the Upper Cretaceous deposits of Montana, USA.

UUID: http://zoobank.org/7c1a5026-7f27-4f12-a9fb-1eb8daa93baf