Seven species of cornulitids, one unidentified tubicolous shell, and the problematic bryozoan Lagenosypho Spandel, 1898 are here described from the Katian of Baltica. Three new species—Cornulites lindae new species, Cornulites meidlai new species, and Conchicolites kroegeri new species—are described. The unidentified tubicolous organism has punctate shell structure and setae-like structures that can best be affiliated with lophophorates. The Hirmuse fauna indicates that the diversity and number of cornulitids in the Ordovician of Baltica has been underestimated and it is likely that the Baltic cornulitid fauna was as diverse and abundant as the fauna of Laurentia. Clay mud-bottom environments supported the highest cornulitid diversity in the Late Ordovician of Baltica. The occurrence of intermediate forms indicates that some tentaculitid characters, e.g., regular annulation and a nearly straight shell, which were thought to be apomorphies of free-living tentaculitids, were actually inherited from ancestral cornulitids. The cornulitid fauna of the Late Ordovician of Laurentia somewhat resembles the cornulitid fauna of the Late Ordovician of Baltica, but there are fewer common faunal elements between Gondwana and Baltica.

UUID: http://zoobank.org/cc704237-b578-4d06-a8ed-8457ca3ce972

Zygocycloides? foerstei n. sp. is described from the Llandovery (Aeronian) Brassfield Formation of southwestern Ohio. This is among the oldest reported Silurian cyclocystoids from North America and is the only North American Llandovery cyclocystoid that is preserved with a complete rim of marginal ossicles. Zygocycloides Smith and Paul, 1982 is most similar to Nicholsodiscus Glass et al., 2003 (Katian) and Perforocycloides Ewin et al., 2019 (Llandovery, Telychian), both from Anticosti Island, Québec. Cyclocystoids (Ordovician to Mississippian) survived Late Ordovician extinctions, and this discovery documents that this echinoderm clade was part of shallow-water, marine paleocommunities during the initial post-extinction transgression onto the Laurentian platform.

UUID: http://zoobank.org/3d3e092e-267b-4841-9245-6275d1eb79a1

Ray-finned fishes comprise nearly half of extant vertebrate species and include several ancient lineages with fossil records that stretch over 200 Myr in time. One of these old clades, the sturgeons and paddlefishes, is distributed across the Northern Hemisphere and includes some of the largest known freshwater fishes. Yet, the fossil record of this lineage (Acipenseriformes) is poor compared to similarly ancient ray-finned fish clades. Here, I describe sturgeon fossils from two geological units in North America < 10 Myr younger than the Cretaceous-Paleogene boundary. Both come from individuals of ~1.5 m in length. These Paleogene forms establish the long history of large body size in Acipenseriforms and reveal sturgeons were some of the largest inhabitants of freshwater ecosystems that were still recovering from the Cretaceous-Paleogene extinction.

A new hibbertopterid eurypterid, Cyrtoctenus bambachi n. sp., is described from the Early Mississippian (Tournaisian) Price Formation of western Virginia. The same unit yields an unidentifiable stylonurine eurypterid. These are the first eurypterids documented from the Mississippian of North America, and only the fourth locality of this age anywhere in the world to yield eurypterids.

UUID: http://zoobank.org/5c84d3a2-ea5a-402c-970d-9b0d867f52c7

Anticosti Island, eastern Canada, records an exceptionally thick and well-exposed Ordovician/Silurian boundary section that hosts a series of diverse marine invertebrate faunas across the Late Ordovician mass extinction. However, the base of the terminal Ordovician stage, the Hirnantian, has been difficult to identify on Anticosti due to the lack of a traditional Hirnantia fauna within the Upper Ordovician Ellis Bay Formation. Previously, the eponymous taxon of the Hirnantia fauna, and type species of the genus Hirnantia, H. sagittifera (M'Coy, 1851) has been reported only from the uppermost Ellis Bay Formation, leading to uncertainty as to the age of the lower Ellis Bay Formation. Here we report Hirnantia notiskuani n. sp. from the lowermost Ellis Bay Formation. This new species is similar to the type species, H. sagittifera, but is distinguished by its strongly dorsibiconvex shell in mature forms and variously developed uniplicate anterior commissure. Occurrences of these two similar species of Hirnantia, H. notiskuani and H. sagittifera, within the lower and uppermost Ellis Bay Formation, respectively, indicate a Hirnantian age for the entire Ellis Bay Formation, a finding that is supported by recent palynological and chemostratigraphic studies. Brachiopod assemblages within the Ellis Bay Formation therefore are best characterized as a unique and diverse Hirnantia fauna, consisting of genera from both the typical Hirnantia fauna and the epeiric seas of Laurentia.

UUID: http://zoobank.org/1c1dff6a-ee38-4cd3-b8bd-3d803a2774ef

Although the Cretaceous is widely regarded as a time of great evolutionary transition for the freshwater fish fauna of North America, the fossil record of this period is notoriously poor, consisting mostly of fragments and isolated skeletal elements. Exceptions include the acipenseriforms, discussed in this paper, and some exceedingly rare teleosts. Here we describe two new species of well-preserved sturgeons (Acipenseridae) from the Tanis site in the Late Cretaceous Hell Creek Formation of North Dakota. The type and referred materials were preserved in a loosely consolidated matrix. †Acipenser praeparatorum n. sp. is represented by multiple body fossils (including the head and relatively complete postcranial remains) and a specimen of an intact, three dimensionally preserved skull and pectoral girdle. This taxon can be diagnosed based on features of the opercular elements (exceptionally tall and narrow branchiostegal). The second species, †Acipenser anisinferos n. sp., is represented by a partially preserved skull, and can be diagnosed by a relatively elongate preorbital region (i.e., snout) and the absence of thorn-like spines on the skull roofing bones. Most known sturgeon fossils from the Cretaceous are represented only by undiagnosable fragmentary remains (i.e., scutes and pectoral-fin spines) or poorly preserved partial skeletons (e.g., †Protoscaphirhynchus), with †Priscosturion and †Anchiacipenser (both monotypic) being rare exceptions. Therefore, the newly discovered Tanis fossils give a rare glimpse into the evolution of Acipenseridae at a critical time in the phylogenetic history of acipenseriforms, and suggest significant morphological and taxonomic diversity early in the evolution of this group.

UUID: http://zoobank.org/375b586a-2dd8-4a31-b6c4-42151e6e8fc8

Non-archosaur archosauromorphs are a paraphyletic group of diapsid reptiles that were important members of global Middle and Late Triassic continental ecosystems. Included in this group are the azendohsaurids, a clade of allokotosaurians (kuehneosaurids and Azendohsauridae + Trilophosauridae) that retain the plesiomorphic archosauromorph postcranial body plan but evolved disparate cranial features that converge on later dinosaurian anatomy, including sauropodomorph-like marginal dentition and ceratopsian-like postorbital horns. Here we describe a new malerisaurine azendohsaurid from two monodominant bonebeds in the Blue Mesa Member, Chinle Formation (Late Triassic, ca. 218–220 Ma); the first occurs at Petrified Forest National Park and preserves a minimum of eight individuals of varying sizes, and the second occurs near St. Johns, Arizona. Puercosuchus traverorum n. gen. n. sp. is a carnivorous malerisaurine that is closely related to Malerisaurus robinsonae from the Maleri Formation of India and to Malerisaurus langstoni from the Dockum Group of western Texas. Dentigerous elements from Puercosuchus traverorum n. gen. n. sp. confirm that some Late Triassic tooth morphotypes thought to represent early dinosaurs cannot be differentiated from, and likely pertain to, Puercosuchus-like malerisaurine taxa. These bonebeds from northern Arizona support the hypothesis that non-archosauriform archosauromorphs were locally diverse near the middle Norian and experienced an extinction event prior to the end-Triassic mass extinction coincidental with the Adamanian-Revueltian boundary recognized at Petrified Forest National Park. The relatively late age of this early-diverging taxon (Norian) suggests that the diversity of azendohsaurids is underrepresented in Middle and Late Triassic fossil records around the world.

UUID: http://zoobank.org/e6eeefd2-a0ae-47fc-8604-9f45af8c1147.

Platymerella from the lower Red Mountain Formation in Georgia and Tennessee, the Bowling Green Dolomite of west-central Illinois, and the Elwood Dolomite of northeastern Illinois represents a paleosubtropical, cool-water occurrence of virgianid brachiopods in Laurentia during the early Silurian (middle–late Rhuddanian). These occurrences were located in the southern Appalachian foreland basin and the distal end of the Sebree Trough, likely subjected to frequent cool-water current and upwelling from Gondwana. Compared with broadly coeval species of Virgiana from lower paleotropical to equatorial latitudes, Platymerella has significantly smaller, dorsoventrally flattened shells, with subequal ventral and dorsal umbones and beaks that extend only slightly above the hinge line. Relative to its shell size, however, Platymerella has more prominent thickening of the shell wall, median septum, spondylium, and hinge plates than Virgiana, resembling more closely the extravagant shell thickening of Tcherskidium and Proconchidium from the Late Ordovician (late Katian) equatorial regions. The thickening of hinge plates resulted in the formation of a pseudocruralium, which separates Platymerella from Virgiana. In latest Ordovician–earliest Silurian virgianids, there was a general morphological gradient toward a smaller shell, reduction in the ventral-valve convexity, and reduction in the size and height of the ventral umbo from paleoequatorial to southern subtropical regions, with Platymerella representing the most southerly forms.

The early geologic history of family Pinnidae, a diverse and abundant component of mud-dominated shallow marine environments of the late Paleozoic, is obscured by ill-defined genus and species concepts. Mistakes in descriptions and reconstructions have provided a basis for suggestions that older taxa in the family differ from younger taxa and may not belong in family Pinnidae. This report provides improved documentation that removes the basis for the concept of divergence. Late Paleozoic genera and species have definite pinnid bivalve characters, including a dorsomarginal fold that holds the ligament, a thick outer columnar prismatic shell layer, and an equivalved, triangular shell, all of which provide evidence for confident assignment of the genus to family Pinnidae. The suggested synonymy of Aviculopinna with Pteronites is invalid, and genus Aviculopinna is limited to occurrence in Permian strata. The two genera belong in different families. A neotype is designated for genus Aviculopinna type species Aviculopinna pinnaeformis Geinitz from Gera, Germany, and a lectotype for Aviculopinna neukirchensis Langenhan from Nowy Kościół, Poland. Inferences on the life habits of Aviculopinna based on its occurrence in Poland are presented. An evaluation of the subterminal beak versus terminal beak concept in pinnids is presented and conclusion presented that there are few data available to support the concept or to test it. In its present form, the concept has no utility in the study of pinnid bivalves.

This work describes a rhynchonellid and spire-bearing brachiopod fauna from the Ixtaltepec Formation of Oaxaca, Mexico. Leiorhynchoidea perrilliatae, Allorhynchus scientiana, and Anthracospirifer oaxacaensis are new species. The specific determination, along with information of previously described taxa, allowed the establishment of precise relative ages of the different fossiliferous intervals (API-1 to API-8) of the formation. The occurrence of Serpukhovian taxa in API-1 to API-3 allowed assignment of the strata to the Upper Mississippian. The presence of Bashkirian species allowed the assignment of the rocks of API-5 and API-6 to the Lower Pennsylvanian. Likewise, Middle Pennsylvanian brachiopods in API-7 and API-8 enabled correlation of the strata with the Moscovian stage. This study shows that the Ixtaltepec Formation is represented by a succession of well-delimited Serpukhovian, Bashkirian, and Moscovian rocks. Regarding paleogeography, the brachiopod fauna displays clear taxonomic variations that concur with global geological changes that occurred between the Serpukhovian to Moscovian. In the Serpukhovian intervals, we recorded numerous cosmopolitan taxa of tropical waters, coinciding with the migration pathway of the Rheic Ocean. For the Bashkirian, we observed a North American provincialism; however, because of the presence of Australian and South American species, it is proposed that the Austropanthalassic-Rheic corridor had a close connection with Oaxaca. The main provincialism was observed in the Moscovian association because most of those taxa have been reported from different localities in the United States. This study supports that the main resemblance between Oaxacan and North American faunas continued until the Pennsylvanian and not the Mississippian, as was previously proposed.

UUID: http://zoobank.org/181e49cf-a08a-4e99-8200-65f4a90dafcd

A highly diversified fauna of hermit crabs associated with reef environments from the Serraduy Formation (lower Eocene) in the southern Pyrenees (Huesca, Spain) is described. Other European Eocene outcrops have yielded paguroids associated with a single environment; however, the studied association represents one of the highest paguroid diversities in a single Eocene outcrop worldwide. The new material increases the diversity of known fossil paguroids, including eight species from which six are new: Clibanarius isabenaensis n. sp., Parapetrochirus serratus n. sp., Dardanus balaitus n. sp., ?Petrochirus sp., Eocalcinus veteris n. sp., ?Pagurus sp., Paguristes perlatus n. sp., and Anisopagurus primigenius n. sp. We erected a new combination for Paguristes sossanensis De Angeli and Caporiondo, 2009 and Paguristes cecconi De Angeli and Caporiondo, 2017 and transfer them to the genus Clibanarius. This association contains the oldest record of the genera Eocalcinus and Anisopagurus. Our data demonstrate that paguroids were diverse by the early Eocene in coral-reef environments and fill an important gap between the poorly known Paleocene assemblages and the more diverse mid- to late Eocene ones.

UUID: http://zoobank.org/091f7a79-946c-4cd9-bdad-f727695e8503

Claraia Bittner, 1901 is a prominent extinct bivalve genus of the Permian-Triassic transition (Newell and Boyd, 1995). Although it first appeared in the Wuchiapingian (Late Permian; Fang Zong-Jie, 2010), its massive proliferation in the immediate wake of the end-Permian mass extinction makes it the archetype of a cosmopolitan and eurytopic disaster species and a hallmark of the base of the Triassic (e.g., Schubert and Bottjer, 1995). Diener (1923, p. 38) fixed “Posidonomya Clarai Emmrich (1844)” as the type species of Claraia by subsequent designation, but spelling and authorship of this species have been controversial. This short contribution aims to clarify these issues.

Borophagus is the terminal genus of the highly diverse and successful subfamily, Borophaginae. Skeletal remains of this bone-crushing canid are most commonly found in transitional or grassland environments across North America between Late Miocene–Middle Pleistocene, but are rare or absent in forested habitats. Here, we describe a humerus from the Gray Fossil Site of Tennessee, which is the first occurrence of this genus in a heavily forested ecosystem. The distinct limb proportions of Borophagus suggest the genus may have been well suited for a closed habitat like the Gray Fossil Site, contrary to where a majority of their fossils have been previously found. This discovery documents the first pre-Pleistocene occurrence of a canid in the Appalachian region of the eastern United States.

The Late Ordovician (late Katian) Tcherskidium fauna consisted of large- and thick-shelled virgianid pentamerid brachiopods characterized by large and ribbed shells of Tcherskidium and Proconchidium and usually associated with Holorhynchus, Deloprosopus, and Eoconchidium. This unique fauna was widely distributed across several tectonic plates, largely confined to the paleoequatorial and especially the northern paleotropical zones, such as northern Laurentia, accretionary terranes of Alaska, Kolyma, Baltica, Siberia, Kazakh and adjacent terranes, and South China. In Laurentia, the eponymous genus Tcherskidium was predominant in regions north of the paleoequator and, in sharp contrast, was absent south of the paleoequator. In this study, Tcherskidium lonei n. sp. and Proconchidium schleyi n. sp. are described from Alaska and North Greenland, respectively, adding new data on the Tcherskidium fauna of the Late Ordovician Northern Hemisphere. Shell gigantism, together with the sharp paleobiogeographic division, suggests that the Late Ordovician (late Katian) Northern Hemisphere had a prevailing warm-water mass, probably due to the lack of large landmass beyond the northern tropics. This was in sharp contrast to the Southern Hemisphere, which was frequently influenced by cold-water invasions from the ice-bearing Gondwana supercontinent centered on the South Pole.

UUID: http://www.zoobank.org/25d9b772-bd7d-4ad6-bfc6-ba02b1567cf3

Lower and middle Cambrian strata of the eastern Mackenzie Mountains, Northwest Territories, Canada, were deposited in a semi-enclosed basin along the eastern flank of the Mackenzie Arch. The Mount Clark Formation is predominantly composed of nearshore sandstone and is overlain by deeper water siltstone, mudstone, and carbonates of the Mount Cap Formation. The contact between these formations is interpreted as a flooding surface. Trilobite biostratigraphy indicates the presence of the traditional upper Olenellus through Glossopleura zones (Cambrian Series 2, Stage 4–Miaolingian, Wuliuan) and suggests the flooding surface is diachronous, spanning the Dyeran-Delamaran boundary. Above the Olenellus Zone, the Amecephalus arrojosensis-Eokochaspis nodosa Zone, the new Aitkenaspis keelensis Zone, the new Albertelloides mischi Zone, and the Glossopleura walcotti Zone are recognized. Whereas the older zones are comparable to those in other areas of Laurentia, the trilobite faunas in the Albertelloides mischi and Glossopleura walcotti zones show a greater abundance of zacanthoidids and dolichometopids. They also have a lower diversity of ptychoparioids and oryctocephalids, and lack agnostoids, eodiscoids, dorypygids, and ogygopsidids. This suggests that zacanthoidids and dolichometopids were able to tolerate conditions that were unfavorable to the other groups, probably related to semi-restricted conditions in the basin. Four endemic species exhibit characteristics that are considered paedomorphic. This developmental process took place in three separate lineages, suggesting that heterochrony was also environmentally provoked.

New taxonomic names are authored by Handkamer and Pratt. New genera are Eobathyuriscus, Sahtuia, Mexicaspidella Aitkenaspis, Dodoella, and Mackenzieaspis. New species are Bolbolenellus dodoensis, Eobathyuriscus mackenziensis, E. macqueeni, Glossopleura youngi, Sahtuia carcajouensis, Aitkenaspis keelensis, Albertelloides eliasi, Dodoella kobayashii, Mackenzieaspis parallelispinosa, and M. divergens.

UUID: http://zoobank.org/a10a2470-853c-4ce8-bf62-7de1d2c99fab.

The diversity of mid-Cretaceous tyrannosauroids is poorly understood. We describe a partial tyrannosauroid femur from the Albian–Cenomanian Wayan Formation of eastern Idaho that helps to fill in an important spatiotemporal gap in the North American record of tyrannosaurs. This specimen, consisting of the proximal half of the bone, is morphologically similar to the femur of Moros intrepidus, a small-bodied tyrannosauroid from the Cenomanian Mussentuchit Member of the Cedar Mountain Formation of Utah, but not referable to this taxon. The Wayan femur lacks an autapomorphy diagnostic for Moros intrepidus, indicating the presence of a previously unrecognized tyrannosauroid taxon in the early Late Cretaceous of Laramidia. Histological results indicate that, at the time of death, this individual was at least five years old, skeletally immature, and undergoing growth at a moderate rate. The addition of this tyrannosauroid to the Wayan-Vaughn Assemblage provides additional evidence for the widespread distribution of various tyrannosauroid taxa in Laramidia during the early Late Cretaceous.

Two rare, phosphatized, tubular microfossils of uncertain affinity, probably algal, are described from the Portfjeld biota (Ediacaran) of North Greenland. Portfjeldia aestatis n. gen. n. sp. preserves two tubules, one of which branches, within an annulated outer sheath. Its morphology can be compared to that of the much larger Ramitubus from the Weng'an biota of South China. An unnamed long, parallel-sided tubule occupied the postmortal cavity formed within diagenetically mineralized specimens of the cyanobacterium Jiangispirellus.

The Cambrian successions at the Chihuarruita Hill outcrop, Sonora, Mexico, have yielded two successive linguliform brachiopod assemblages that are transitional between Cambrian Stage 4 and the newly recognized global Wuliuan Stage. The lowermost assemblage includes Dictyonina sp., Paterina sp., Eothele sp., Hadrotreta rara? (Cooper), and Linnarssonia arellanoi? (Cooper), coming from the upper part of the Buelna Formation. The younger, recently named El Gavilán Formation contains a more diverse linguliform brachiopod assemblage, including Acrothele concava Cooper, Batenevotreta? mexicana n. sp., Dictyonina minutipuncta Cooper, Eothele sp., Eoobolus sp., Hadrotreta rara? (Cooper), Linnarssonia arellanoi? (Cooper), Micromitra sp., Paterina sp., and Prototreta sp. The El Gavilán Formation contains a diverse trilobite fauna suggesting Delamaran age in terms of the Laurentian regional stratigraphical scheme. The base of the global Wuliuan Stage and Miaolingian Series is defined by the first occurrence of Oryctocephalus indicus; in the absence of the index species, the base should be provisionally placed at the base of the El Gavilán Formation. The Wuliuan age of the brachiopod assemblage recovered from the El Gavilán Formation is supported by the occurrence of Acrothele in the Cambrian biostratigraphical succession of Himalaya, where the genus makes its first appearance in the Kaotaia prachina Zone. In addition, the co-occurrences of Acrothele and Eothele can be taken as an indication of the Wuliuan age of the fauna. A new biogeographic analysis confirms that the Eothele Fauna first appeared at the end of Cambrian Stage 4, as a result of increased faunal migration within the southern tropical latitudes directed from Australasian Gondwana to Laurentia.

UUID: http://zoobank.org/6598f51f-cb07-4df3-bd1f-025a13e64edf

Two previously known species of stromatoporoids are reported from Nevada: Stromatoporella perannulata, from the Bartine Member of the McColley Canyon Formation (middle Emsian); and Stictostroma moosense, from the Oxyoke Canyon Sandstone (uppermost Emsian). A new species, Syringodictyon nevadense n. sp., is described from the Coils Creek Member of the McColley Canyon Formation (upper Emsian). Stromatoporella perannulata also occurs in Emsian-age strata on Ellesmere Island, arctic Canada, and in strata of Emsian and/or Eifelian age in Ontario, New York, Ohio, and Kentucky. Outside Nevada, Syringodictyon is confined to the Emsian of Ellesmere Island, and Emsian/Eifelian of Ontario and New York. Stictostroma moosense is known from only the Emsian/Eifelian of Ontario.

It is concluded that the three Nevada specimens are Emsian in age. This implies that in other localities where two of the species, S. perannulata and S. moosense, occur in strata where the placement of the Emsian-Eifelian boundary is in question, it might be higher than previously thought. The presence of the two species, plus Syringodictyon, in both the Old World Realm (Nevada, arctic Canada) and the Eastern Americas Realm (Ontario, New York, Ohio, Kentucky), infers a selective marine connection between the two realms that was not available to other taxa. The breach in the inter-realm barrier most likely occurred as a shallow seaway across the Canadian Shield.

UUID: http://zoobank.org/e73023c6-88ca-4b4d-9b01-f74a75b99c43

The Nasep and Huns members of the Urusis Formation (Nama Group), southern Namibia, preserve some of the most diverse trace-fossil assemblages known from the latest Ediacaran worldwide, including potentially the world's oldest “complex” vertical sediment-penetrating burrows. These sediments record relatively diverse communities of bilaterian metazoans existing before the base of the Cambrian and an increase in the intensity of metazoan ecosystem engineering behaviors that could eventually produce profound changes in the character of the Phanerozoic sedimentary record (the “agronomic revolution”). Despite this, relatively little about this trace-fossil assemblage is known. We explore the Nasep–Huns transition at two localities in the Witputs sub-basin and describe the trace- and body-fossil diversity present in these horizons alongside a paleoenvironmental reconstruction. We document eight unique ichnotaxa from these localities, including well-preserved “probes” potentially left by priapulids. We also report the first occurrence of Corumbella from Namibia, helping to establish a biostratigraphic link between Namibia, Brazil, Paraguay, Iran, and the southwestern United States. Last, we find that several ichnotaxa, in particular small treptichnids, appear to be preferentially preserved on the bases of gutter casts, hinting at the potential existence of an unusual late Ediacaran preservational window with possible implications for timing the first appearance of key bilaterian behaviors.