Modular coral-like fossils from Lower Ordovician (Tremadocian) thrombolitic mounds in the St. George Group of western Newfoundland were initially identified as Lichenaria and thought to include the earliest tabulate corals. They are here assigned to Amsassia terranovensis n. sp. and Amsassia? sp. A from the Watts Bight Formation, and A. diversa n. sp. and Amsassia? sp. B from the overlying Boat Harbour Formation. Amsassia terranovensis n. sp. and A. argentina from the Argentine Precordillera are the earliest representatives of the genus. Amsassia is considered to be a calcareous alga, possibly representing an extinct group of green algae. The genus originated and began to disperse in the Tremadocian, during the onset of the Great Ordovician Biodiversification Event, on the southern margin of Laurentia and the Cuyania Terrane. It inhabited small, shallow-marine reefal mounds constructed in association with microbes. The paleogeographic range of Amsassia expanded in the Middle Ordovician (Darriwilian) to include the Sino-Korean Block, as well as Laurentia, and its environmental range expanded to include non-reefal, open- and restricted-marine settings. Amsassia attained its greatest diversity and paleogeographic extent in the Late Ordovician (Sandbian–Katian), during the culmination of the Great Ordovician Biodiversification Event. Its range included the South China Block, Tarim Block, Kazakhstan, and Siberia, as well as the Sino-Korean Block and Laurentia, and its affinity for small microbial mounds continued during that time. In the latest Ordovician (Hirnantian), the diversity of Amsassia was reduced, its distribution was restricted to non-reefal environments in South China, and it finally disappeared during the end-Ordovician mass extinction.

UUID: http://zoobank.org/ef0abb69-10a6-46de-8c78-d6ec7de185fe

A diverse, well-preserved radiolarian assemblage is reported from the Sandbian age Climacograptus bicornis Graptolite Biozone. This new assemblage, recovered from the Pingliang Formation in the Guanzhuang section, China, includes six new species along with 13 other previously described taxa. Geminusphaera new genus incorporates G. grandis n. sp. and G. kongtongensis n. sp. and is proposed for inaniguttids constructed from two distinct porous spheres bearing seven or more primary spines. Protopylentonema new genus is introduced to incorporate pylomate entactinarians with five-rayed initial spicules. It includes P. ordosensis n. sp. as well as P. aperta, P. rimata, and P. insueta that were formerly assigned to Kalimnasphaera. Micro-computed tomography investigation of skeletal microstructure supports establishment of the new genera together with other new spumellarian species: Haplotaeniatum implexa n. sp., Inanigutta quadrispinosa n. sp., and Kalimnasphaera pingliangensis n. sp. It also indicates that family-level reassignment of Etymalbaillella from the Proventocitidae to the Ceratoikiscidae is appropriate.

Global distribution of Late Ordovician radiolarian occurrences highlights a strong preference for areas with equatorial to tropical sea surface temperatures. This must have led to ecological stress among radiolarian communities in adapting to global cooling in the Hirnantian.

UUID: http://zoobank.org/3d3f55b8-0e70-4f9f-9738-265750d8ec3a.

Lagenochitina esthonica is a globally distributed chitinozoan in Early to Middle Ordovician rocks. It is regarded as an index species for the early Floian in North America and has a stratigraphically constrained range in other regions. Lagenochitina esthonica is distinguished from other chitinozoans by a distinct flexure, a nearly rounded-square chamber, and a cylindrical neck with a flaring collar. However, since the first description of the species in the 1950s, it has included two varieties: a relatively short form with a test length ~400 μm, and a slender form usually longer than 600 μm. In order to revise the taxonomy of the L. esthonica group, we carried out a statistical morphometric study of a large collection of well-preserved specimens from the Baltic region where the taxon was first established. Additionally, the stratigraphic and geographic distribution of both forms was analyzed based on available occurrence data. The results show that the short form occurs in the upper Tremadocian to lower Dapingian, whereas the slender form is mostly reported from the lower and middle Darriwilian. Both forms are identified on Baltica; the short form has also been reported from Laurentia and South China, whereas the other is known also from Avalonia and Gondwana. The morphological distinction, together with differences in stratigraphic and spatial ranges, suggest that the two forms represent separate species: the original stout L. esthonica, based on the morphology of the holotype, and the slender L. megaesthonica n. sp., described herein. The updated taxonomy enhances the stratigraphic and biogeographic usefulness of lagenochitinids globally.

UUID: http://zoobank.org/ec49166e-2a8d-4941-8723-f023853c5a7e.

A moderately diverse assemblage of brachiopods from the Latham Shale Lagerstätte (Cambrian Series 2, upper Stage 4) and the upper Cadiz Formation (Miaolingian, Wuliuan), California is described in detail for the first time. The fauna includes both linguliform and rhynchonelliform brachiopods—Hadrotreta primaea (Walcott, 1902), Paterina prospectensis (Walcott, 1884), Dictyonina pannula (White, 1874), and Mickwitzia occidens Walcott, 1908; and Nisusia fulleri Mount, 1981 and Wimanella highlandensis (Walcott, 1886), respectively—together with olenellid trilobites and hyolithids. The fauna differs from other Cambrian Konservat-Lagerstätten (notably Cambrian Series 2 Chengjiang and Guanshan Lagerstätten, eastern Yunnan) in that the brachiopod shell valves in many cases are still preserved with their original mineralization. Moreover, the excellently preserved shale-hosted valves even include cases with exquisite epithelial cell molds, otherwise only seen in acid-etched material from carbonate rocks. The pitted ornamentation in D. pannula closely resembles that described from Ordovician linguliforms. The unusual preservation of N. fulleri provides important clues for ancestral composition of the brachiopod shell. The two articulated rhynchonelliform species probably represent the oldest records of this group from the west Laurentia. The fauna could also represent the earliest onset of the transition from the Cambrian Evolutionary Fauna (CEF) to the Paleozoic Evolutionary Fauna (PEF).

Strata of the Solvik Formation in the central Oslo region (upper Hirnantian to upper Aeronian) are rich in atrypides that elucidate the evolution of this group. A few athyridide brachiopods are also present. Eleven taxa of smooth spire-bearers are described taxonomically using fossils and peels. Among these occur one new genus, Eisaella, and five new species: Eisaella uniplicata, Thulatrypa huangi, T. vikenensis, ?Becscia pentagona, and Shelvothyris bivittata. Atrypides make up 30% (24 genera) of the total brachiopod genera in the Solvik Formation, more than one third of the known global atrypide fauna during that interval. Most are members of the family Lissatrypidae, illustrating the evolution of this group from upper Hirnantian and lower Silurian strata. These show close affinity to each other attesting to ongoing radiation. The new genus Eisaella is a likely ancestor of Lissatrypa, while Meifodia displays a gradual evolution through the formation. Atrypides were most diverse at the base and top of the Solvik Formation due to heterogony in environments and water depth, but were less diverse in the comparatively constant environmental setting of the late Rhuddanian time. The basal assemblages with few atrypides from shallow facies may be compared to the shallow Cathay Fauna of China. Most of the abundant Norwegian atrypide fauna is found in deeper water with few comparable faunas known globally. The Oslo region may have served as a center for the evolution and spread of atrypides immediately following the end-Ordovician glacial drawdown in sea level.

UUID: http://zoobank.org/508dba95-5501-403d-9a4d-deeb324847c9

A new genus and species of microconchid tubeworm, Aculeiconchus sandbergi n. gen. n. sp., is described from the Givetian (Devonian) Maywood Formation of Cottonwood Canyon, Wyoming, USA. It possesses unique hollow spines of various lengths on the tube underside, a position previously undocumented for these fossils. Like some cyclostome bryozoans possessing basal tubular extensions, the basal spines of Aculeiconchus n. gen. were presumably also used for fixation to flexible substrata, e.g., algal thalli, which is a previously undocumented adaptive strategy in microconchids. Together with other skeletal features, such basal spines could suggest that ‘lophophorate’ microconchids, unlike the other tentaculitoids, might be phylogenetically not as distant from bryozoans as previously thought. The Maywood Formation, which contains a few-millimeters thick, monospecific shell accumulation of the microconchids described herein, records deposition in an estuarine brackish setting within narrow channels that were cut into underlying strata. The microconchids were opportunistic taxa that repeatedly colonized these salinity-stressed estuarine channels, leading to a series of adaptive innovations, including colonization of plant stems during the Early Devonian (Beartooth Butte Formation) and possibly flexible, soft-algal substrata during the Middle Devonian (Maywood Formation, this study). Tectonic quiescence during the Early and Middle Devonian indicates that erosion and subsequent deposition of the Maywood and the underlying Beartooth Butte Formation channels were responses to major eustatic events. Over a span of nearly 30 Myr, channels were cut successively during lowstand conditions and a distinctive faunal assemblage with microconchids tracked marine transgressions into the channels.

UUID: http://zoobank.org/394c8b32-d5e7-411e-8e56-6fb9f55bbb8a

The record of Middle Ordovician (Oretanian, ca. Darriwilian 2) hyolithids from the Ossa Morena Zone of the Iberian Massif in southwestern Spain is increased with the recognition of Robardetlites sevillanus n. gen. n. sp., Andalucilites parvulus n. gen. n. sp., Pauxillites desolatus n. sp., Leolites malinkyi Marek and Gutiérrez-Marco n. sp., and Cavernolites sp. in that region. Andalucilites n. gen. and Robardetlites n. gen. are endemic whereas the other genera are known from coeval strata in the Barrandian region in central Europe, France, and Morocco, giving this assemblage a decidedly “Mediterranean province” character. The discovery of Pauxillites, Leolites, and Cavernolites in Iberia extends their geographic ranges to that region, and Leolites from this area extends its stratigraphic range downward, making its occurrence in Iberia the oldest known for that genus.

UUID: http://zoobank.org/f5b29223-6dac-4c7e-8851-551e634da667

The cancellarid genus Sveltia Jousseaume, 1887, is widespread in western European and North African Neogene marine fossil assemblages. In Pliocene deposits it is commonly represented by Sveltia varicosa (Brocchi, 1814), which until recently was considered a widely distributed taxon in the Mediterranean Sea and adjacent Atlantic faunas. A recent review of the species from the Pliocene of Italy and Spain (Guadalquivir Basin), leading to the erection of S. confusa, prompted the reassessment of the Sveltia material from the Atlantic Pliocene of the Portuguese Mondego Basin and the subsequent description of Sveltia sofiae n. sp. Consequently, a mosaic of species has emerged from what was previously viewed as the broad Atlanto-Mediterranean range of the widespread and quite variable S. varicosa. From a biogeographic standpoint, it is now clear that S. varicosa was a Mediterranean species, occurring east of the Alboran Sea. Sveltia confusa had a mainly Atlantic distribution, from the French Pliocene Ligerian Gulf to the Gulf of Cadiz, at least, and straddling the Strait of Gibraltar into the Alboran Sea. Sveltia sofiae n. sp. was endemic to western Iberia, represented today only in the western Portuguese Mondego Basin. Sveltia is a thermophilic genus. Since early Pliocene times, because of northeastern Atlantic sea surface temperature decline, it underwent a southward range contraction, occurring today—in the eastern Atlantic—from Cape Blanc, Mauritania, south. This range reduction was coupled with the post mid-Piacenzian southward contraction of the Pliocene Mediterranean-West African tropical molluscan province and the consequent rise of the present-day Mediterranean-Moroccan subtropical province.

UUID: http://zoobank.org/0cf3c73a-8d57-472e-87e4-f1ad065e5fb6.

Exceptionally preserved fossil eggs and embryos provide critical information regarding paleoembryogenesis, reproductive strategies, and the early ontogeny of early arthropods, but the rarity of preservation of both eggs and egg-bearing organisms in situ limits their use in detailed evolutionary developmental (evo-devo) studies. Burgess Shale-type deposits preserve rare instances of egg-bearing arthropods as carbonaceous compressions; however, the eggs are usually poorly preserved with no compelling evidence of embryos. We describe the first record of a brooding specimen of Waptia cf. W. fieldensis from the Spence Shale, a Cambrian (Wuliuan Stage) Burgess Shale-type deposit in northeastern Utah and southeastern Idaho. This is the first record of an egg-bearing arthropod from the Spence Shale and it exhibits two distinct modes of preservation among eggs within the single clutch: carbonization and phosphatization. Unlike the egg-bearing Burgess Shale specimens, many eggs of this Utah specimen are also preserved three-dimensionally. In addition, synchrotron radiation X-ray tomographic microscopy reveals internal distributions of mineral phases, along with potential remnants of the egg membrane and attachment structures, but, as in the Burgess Shale, no explicit traces of developing embryos. The distinct modes of preservation highlight the existence of diagenetic microenvironments within some eggs, but not in others during fossilization.

The Devonian saw the emergence and subsequent diversification of the clam shrimp. To date, there are no credible records of any clam shrimp prior to the Devonian. Therefore, discoveries and taxonomic studies of early forms, especially the Early Devonian species, are essential to broaden our knowledge of the origin and early evolution of clam shrimp. An Early Devonian clam shrimp community that consists of four species, Cornia cheni n. sp., Pseudestheria cf. P. diensti, Palaeolimnadiopsis zhangi n. sp., and ?Palaeolimnadia sp. is described and discussed in detail in this study. These species are the oldest clam shrimp in China and among the oldest worldwide. This discovery indicates that clam shrimp communities with high diversity existed since the late Early Devonian and that their early forms may have had a profound effect on the origin and evolutionary trends of the following forms.

UUID: http://zoobank.org/ae9d0ab8-0b0e-4103-ad71-601d11a024fc.

Despite the abundant literature on limnic Cretaceous ostracode faunas, the database on mid-Late Cretaceous taxa is still scarce. The Songliao Basin in northeastern China preserves a diverse assemblage of ostracode fossils from the Santonian–Campanian Nenjiang and Sifangtai formations. This rather unique material is of major importance to comprehend Early to mid-Late Cretaceous (Cenomanian–Campanian) limnic ostracode faunas and therefore the evolution of Late Cretaceous basins in China. A study of this fauna was conducted to detail the taxonomy of ostracode species from members 1 and 2 of the Nenjiang and Sifangtai formations in the Songliao Basin. Well-preserved specimens of 14 ostracode species were recovered from samples of the ZKY2-1 well in southwestern Songliao Basin: Scabriculocypris liaukhenensis Liu in Netchaeva et al., 1959, Ilyocyprimorpha netchaevae Su in Netchaeva et al., 1959, Cypridea acclinia Netchaeva in Netchaeva et al., 1959, Cypridea cavernosa Galeeva, 1955, Cypridea gracile Netchaeva in Netchaeva et al., 1959, Cypridea lepida Ye in DOFEAD, 1976, Cypridea squalida Sou in Netchaeva et al., 1959, Fabaeformiscandona? Disjuncta (Hao in Hao et al., 1974), Lycopterocypris profunda Lübimova, 1956, Mongolocypris magna (Hou, 1958), Mongolocypris tera (Su in Netchaeva et al., 1959), Talicypridea obliquecostae (Szczechura and Błaszyk, 1970), Talicypridea reticulata (Szczechura, 1978) and Renicypris renalata (Su in Hao et al., 1974). Four of the species identified received emended descriptions and diagnoses. Other important results include the first discussion on sexual dimorphism in Cypridea acclinia and Fabaeformiscandona? disjuncta, the first study on the ontogeny of Ilyocyprimorpha netchaevae, and the reassignment of Cypridea liaukhenensis to the genus Scabriculocypris. Several species recorded herein are also found in other continental far-eastern Asian basins, while genera range from worldwide to far-eastern Asian distribution; these results support strong affinities among faunas of Mongolia, China, and Japan.

Two cupressocrinitids (Crinoidea, Eucladida) from the famous crinoid collection of Dr. Ludwig J.T. Schultze deposited in the Museum of Comparative Zoology (MCZ) are revised in the present study: Cupressocrinites hybridus n. comb. and Halocrinites minor n. comb. The rare C. hybridus has a stratigraphically and regionally restricted occurrence at the Eifelian–Givetian boundary of the Hillesheim Syncline (Eifel, Rhenish Massif, Germany), but the more common H. minor occurs supraregionally from the Eifelian to the Frasnian (Middle to Upper Devonian). Both cupressocrinitids are redescribed on the basis of the type material stored in the MCZ. In addition, the rare Halocrinites heinorum n. sp. from the Eifelian–Givetian boundary beds and lower Givetian of the Eifel (Rhenish Massif, Germany) is described, and Encrinus townsendi (König, 1825) is questionably reassigned to Robustocrinites: R.(?) townsendi n. comb. Pre- and postmortem ossicular modifications of the studied cupressocrinitid skeletons are also discussed.

UUID: http://zoobank.org/4598e1fa-8ca5-4160-8fee-6e8c6d444115

The Miocene beds of Kutch in India are well known for their mammalian assemblages, including the extinct ape Sivapithecus, but far less is known about the fossil squamates from this area. Although India with its over 800 reptile species is recognized as one of the global biodiversity hotspots, knowledge of past diversity and paleobiogeography of squamates on this subcontinent is very limited. We here report on new lizard finds, which have been recovered from two stratigraphic levels: the older Palasava locality (dated to the middle Miocene, ca. 14 Ma) and the younger Tapar site (late Miocene, ca. 11–10 Ma). Although fragmentarily preserved, the material described here sheds important light on the composition and paleobiogeography of squamates during the Miocene in South Asia. The older Palasava locality contains cf. Uromastyx s.l. and Varanus sp., the latter representing the oldest record of this taxon in the region of India south of the Himalayas and its occurrence here suggests a mean annual temperature not less than 15°C. The material from the younger Tapar locality consists of an unidentified acrodontan lizard, here questionably placed in agamids, and a skink. The latter shows a resemblance to mabuyines, however, the fragmentary nature of the material does not allow a precise allocation without doubts. The cosmopolitan mabuyines have been suggested to have their origin in Asia, so the potential presence of mabuyines in the Tapar locality might represent the first, but putative, Asian evidence of the occurrence of this group in the Miocene.

We report new specimens of the Plotopteridae from Washington State (USA), an area where these flightless seabirds underwent significant diversification during the late Eocene and Oligocene. To date, five plotopterid species from western Washington have been formally named. Specimens previously assigned to Tonsala buchanani Dyke, Wang, and Habib, 2011 belong to at least two, but probably even three, different species. One of these, the large-sized “Whiskey Creek specimen” from late Eocene deposits mapped as the Makah Formation, is the oldest known plotopterid and is here tentatively assigned to ?Klallamornis clarki Mayr and Goedert, 2016. Another specimen originally referred to T. buchanani is also likely to belong to a different species and is among the most substantial records for North American plotopterids. We formally transfer T. buchanani to the taxon Klallamornis and show that the only unambiguously identified specimen of the species—the holotype—is currently poorly diagnosed from Klallamornis abyssa Mayr and Goedert, 2016, which is from coeval strata of the Pysht Formation. Although the holotype of K. abyssa is larger than that of K. buchanani, there remains a possibility that plotopterids were sexually dimorphic in size. We describe the first ungual phalanx of a plotopterid, which is referred to K. buchanani, and report previously unknown elements of the large ?K. clarki and the first records of this species from the Lincoln Creek Formation. Current data indicate that plotopterids originated in the middle or late Eocene on islands off western North America, and we hypothesize that the radiation of these birds in the North Pacific Basin may have been related to the evolution of kelp forests.

One of the oldest fossil horseshoe crabs figured in the literature is Entomolithus lunatus Martin, 1809, a Carboniferous species included in his Petrificata Derbiensia. While the species has generally been included within the genus Belinurus Bronn, 1839, it was recently used as the type species of the new genus Parabelinurus Lamsdell, 2020. However, recent investigation as to the appropriate authority for Belinurus (see Lamsdell and Clapham, 2021) revealed that all the names in Petrificata Derbiensia were suppressed in Opinion 231 of the International Commission on Zoological Nomenclature (1954) for being consistently nonbinomial under Article 11.4 of the International Code of Zoological Nomenclature (ICZN) (International Commission on Zoological Nomenclature, 1999). Despite the validation of several species names for anthozoans, brachiopods, and cephalopods described in Petrificata Derbiensia in subsequent rulings (International Commission on Zoological Nomenclature, 1956a, b), Belinurus lunatus has not been the subject of any subsequent Commission ruling or opinion, and so its use in Petrificata Derbiensia remains suppressed. The Belinurus lunatus species name was used in several subsequent publications during the 1800s, none of which made the name available under ICZN article 11.5; Parkinson (1811) is also suppressed for being nonbinomial, while Woodward (1830), Buckland (1837), Bronn (1839), and Baily (1859) refer to the species only as a synonym of Belinurus trilobitoides (Buckland, 1837) through citation to the suppressed Pretificata Derbiensia. The first author to make Belinurus lunatus an available name was Baldwin (1905), who used the name in reference to a new figured specimen from Sparth Bottoms, Rochdale, UK, but again as an explicit junior synonym of Belinurus trilobitoides (Buckland, 1837). Therefore, it was not until Eller (1938) treated B. lunatus as a distinct species from B. trilobitoides that B. lunatus became an available name as per ICZN Article 11.6.1 under the authorship of Baldwin (1905) following ICZN Article 50.7.