Known antennae of trilobites are all flagelliform, in marked contrast to the varied first antennae of marine pancrustaceans. An exceptionally preserved specimen of the Early Ordovician (late Tremadocian) asaphid Asaphellus tataensis Vidal, 1998, from the Fezouata Shale in Morocco, exhibits both antennae in situ; they are relatively short, widen distally, and bear a series of round, dome-shaped organs along both their dorsal and ventral surfaces. These organs are vastly larger than chemo- or mechanosensory sensilla on the antennae of other arthropods, rendering their homology and function uncertain. The clavate antennae of Asaphellus reveal the furthest deviation from the inferred ancestral state of the trilobite antenna known to date. This could be an adaptation for detection of prey, because most Asaphidae have been claimed as predators and scavengers on the basis of specialized features of the calcified exoskeleton.

Toyotamaphimeia is an extinct crocodylian lineage whose name is derived from a mythological Japanese princess. Here, we re-examine the type specimens of a long-forgotten species: Tomistoma taiwanicus from the Pleistocene of Tainan (Taiwan) and revise its taxonomic status to Toyotamaphimeia taiwanicus n. comb., leading to the first recognized species of Toyotamaphimeia outside Japan. Our phylogenetic analyses also support this taxonomic assignment and, more interestingly, further suggest an East Asian lineage. In addition, Toyotamaphimeia taiwanicus n. comb. represents a tropical species, resolving a long-standing puzzle of why Toyotamaphimeia only inhabited a much higher latitudinal area (Japan). Given the large body size of Toyotamaphimeia taiwanicus n. comb. (~7 m) and the fact that it is geologically older than Toyotamaphimeia machikanensis from Japan, we propose a novel evolutionary scenario: the genus Toyotamaphimeia originated in Taiwan and evolved to a large body size with gigantothermic physiology, which resulted in migration out of Taiwan and dispersal farther north to Japan. Our taxonomic identification shows the presence of an extinct endemic crocodylian species from the Pleistocene of Taiwan with large-scale paleogeographic implications. This study, with our recent progress in vertebrate paleontology in Taiwan, should provoke more in-depth paleontological research on the Pleistocene extinction.

Graptolites from the latest Ordovician to the earliest Silurian rocks of northwest Peninsular Malaysia are described and reviewed. The fossils were collected previously by C. R. Jones and presently by the authors inside the black mudstones from the basal section of Tanjung Dendang Formation in Pulau Langgun, Langkawi, which comprises assemblages from the Hirnantian Metabolograptus extraordinarius Biozone to the Rhuddanian Akidograptus ascensus–Parakidograptus acuminatus Biozone. The latest Ordovician strata also include a Hirnantia fauna bed between the Metabolograptus extraordinarius and Metabolograptus persculptus biozones, in which shelly fossils such as Mucronaspis sp. could be recovered. A revised graptolite biozonation is proposed for the latest Ordovician to the earliest Silurian succession of northwest Peninsular Malaysia. This interval is significant for understanding the extent of mass extinction events happening right at the end of the Ordovician period and subsequent faunal change in the region.

New specimens of Cyclocystoides scammaphoris Smith and Paul, 1982, are here reported from the Upper Ordovician Platteville Formation of northern Illinois, Plattin and Decorah groups of east-central Missouri, and Lebanon Limestone of central Tennessee. These fossils reveal skeletal details that provide insight into the anatomy of cyclocystoids. Of particular significance is a network of channels that likely originate near the center of the central disk and extend along the oral side of the radial plates, bifurcating distally two or three times before entering the radial facets on the proximal surface of each marginal ossicle. From here, the network enters a series of facet canals that extend upward through each marginal ossicle, exiting in a linear row of pores. The canals are very similar in size and distribution to the nerve canals in living echinoderms. The axes of the canals, which number up to 500 in some specimens, and those of the radial ducts project proximally away from the oral surface at an elevation angle of about 25°, apparently forming a network that could have converged within the upper part of the body cavity. This origin and function are made clear by the connection between the channel on each radial plate and the radial facet canal pores within each marginal ossicle.

Nemavermes mackeei Schram, 1973, found in the Mazon Creek fossil site and the Bear Gulch Limestone, was described initially as a free-living marine nematode. Here we investigate 13 specimens of N. mackeei from the Mazon Creek to reassess its morphology and identity, and also two specimens originally identified as Gilpichthys greenei Bardack and Richardson, 1977. Based on the extensive morphological variation among these specimens, N. mackeei encompasses multiple species that are only distantly related. The holotype of N. mackeei is a proboscis of Tullimonstrum gregarium Richardson, 1966, making N. mackeei a junior synonym of T. gregarium. However, the other specimens that we investigated could not be attributed to T. gregarium. We name a new species from these specimens: Squirmarius testai new genus new species, a cyclostome. One specimen is likely a juvenile G. greenei. Other specimens were not identified during this study but represent a variety of vermiform bilaterians.

UUID: http://zoobank.org/d8c63f6a-0ef4-4a34-8dbd-c1d9099589cc

Nine crinoids are described from the Wooster Shale Member of the Cuyahoga Formation from Wayne and Ashland counties, Ohio, USA. Identifiable elements of the fauna include five camerate crinoids, one flexible crinoid, and three other eucladid crinoids. Five new species are described, including Cactocrinus woosterensis n. sp., Cusacrinus brushi n. sp., Agaricocrinus murphyi n. sp., Decadocrinus laevis n. sp., and Decadocrinus inordinatus n. sp. Overall, the distribution of crinoid clades in the Wooster Shale is similar to that of the stratigraphically lower Meadville Shale Member of the Cuyahoga Formation, although less diverse and with only one species (Cyathocrinites simplex) in common. Many of the Wooster Shale Member crinoids are completely or partially preserved with siderite either in nodules or within siderite beds. These crinoids are commonly preserved in trauma postures, which is characteristic of burial in episodic high turbulence events. The paleoenvironments and taxa of the two Cuyahoga Formation crinoid faunas more closely resemble Viséan faunas in siliciclastic settings than typical carbonate faunas of the Tournaisian.

UUID: http://zoobank.org/2d2678e1-2367-4429-bea0-a64cd020e98c

We report charadriiform and charadriiform-like birds from the early Eocene London Clay of Walton-on-the-Naze (Essex, UK). A partial skeleton of a small modern-type charadriiform is described as a new species, Charadriisimilis essexensis n. gen. n. sp., and most closely resembles taxa of the Charadrii (plovers, stilts, oystercatchers, and other “wader-like” shorebirds). Affinities to this clade were also supported by phylogenetic analyses, which placed the fossil as the sister taxon of either the Burhinidae or all crown group Charadrii. In addition, we identify specimens of the charadriiform-like taxon Scandiavis, which was before known only from the early Eocene Fur Formation in Denmark. Associated limb elements of two individuals are classified as Scandiavis cf. mikkelseni Bertelli et al., 2013, and remains of two further individuals are tentatively assigned to Scandiavis. The presence of a processus supracondylaris dorsalis on the previously unknown humerus corroborates charadriiform affinities of Scandiavis, whereas a plesiomorphic hypotarsus morphology indicates a position outside crown group Charadriiformes. Charadriisimilis essexensis is one of the earliest modern-type charadriiforms, and the holotype of the species is the most substantial early Paleogene fossil record of a charadriiform bird. Together with Scandiavis, as the best-represented taxon to be considered as a stem group charadriiform, it provides the basis for an improved understanding of the evolutionary history of charadriiform birds.

UUID: http://zoobank.org/ca15ee81-09e8-4577-8beb-a362debf6636

Palmichnium gallowayi (Sharpe, 1932) new combination from the Middle Ordovician Martinsburg Formation (proximal deltaic facies) of Rondout, near Kingston, New York State, is redescribed. It consists of opposing series of five tracks, the outer two large and pear-shaped, the inner three smaller and elliptical, arranged in a chevron converging in the direction of travel, on either side of a wide medial impression. It is attributed to a medium-sized stylonurid eurypterid using a decapodous gait, crawling onto the shoreline, traversing the intertidal zone, a behavior interpreted as part of its reproductive life cycle. This provides the earliest ichnological evidence for the ‘mass-molt-mate’ hypothesis, which proposes that eurypterids migrated en masse into nearshore environments to molt and mate.

The Lochkovian, Pragian, and basal part of the Emsian, which represent the post-Kwangsian Orogeny strata in the South China Block, are mainly composed of siliciclastic rocks. This lithological composition impedes investigation of Pragian and Lochkovian conodont biostratigraphy in the South China Block, which results in a persistent controversy on the age of relevant lithological units. The present study provides new evidence by reporting for the first time Lochkovian conodonts obtained from the South China Block, specifically the Gaoling Member of the Nahkaoling Formation at the Lingli section, central Guangxi. The conodont fauna, consisting of Pandorinellina exigua lingliensis Lu n. subsp., Pandorinellina exigua exigua, Zieglerodina? tuojiangensis Lu n. sp., Amydrotaxis praejohnsoni, and Eognathodus cf. E. irregularis, places the studied interval of the Gaoling Member in the lower or middle Lochkovian (contingent upon varying definitions for the base of the middle Lochkovian) to lower Pragian. Moreover, Amydrotaxis praejohnsoni, which was reported previously only in North America and eastern Australia, is herein also recorded in the South China Block, and thus may play an important role in intercontinental biostratigraphical correlation. By shedding light on the age of the upper limit of the underlying Lianhuashan Formation at the Lingli section, the present study indicates that the Kwangsian Orogeny ended before the late Lochkovian. This date is slightly earlier than the previously estimated late Lochkovian based on studies of fossil plants from the siliciclastic rocks deposited after the Kwangsian Orogeny.

UUID: http://zoobank.org/8010f983-4f6f-43dd-949f-de4653e53d9d

Zokors (Myospalacinae) continue to be the center of systematics discussions. Phylogenetic schemes based on molecular data do not always agree with each other, nor can phylogenetic schemes based on paleontological material be complete due to the only-partial description of West Siberian zokors. This paper tries to fill this gap and presents a description of the West Siberian lineage from the late early Pleistocene to the present, together with an analysis of molar development in other zokor lineages. We describe two new species and one subspecies (Myospalax myospalax krukoveri n. ssp., Myospalax convexus n. sp., Prosiphneus razdoleanensis n. sp.) ancestral to the extant Myospalax myospalax Laxmann, 1769. We also reveal differences in the ontogeny of molars of modern species that were not previously detailed. These differences, together with paleontological data, indicate that in the West Siberian lineage, peramorphosis occurred in the structure of the chewing surface (with the exception of the lower m1), whereas in all other zokor lineages, there was pedomorphosis. On the basis of these results, we suggest a new view on the systematics of Myospalacinae.

UUID: http://zoobank.org/b06d6c99-1648-454b-9b95-4d869bfe8bdc

The recently discovered mass mortality of fishes from the Tanis Site in the North Dakota portion of the Late Cretaceous Hell Creek Formation contains many well-preserved, three-dimensional skeletons. Among these are representatives of two acipenseriform families, Acipenseridae (sturgeons) and Polyodontidae (paddlefishes). This paper describes two new monotypic polyodontid genera, expanding our knowledge of polyodontid diversity. The first of the new species described here is †Parapsephurus willybemisi n. gen. n. sp. It is distinguished from all other known species by having a combination of posteriorly elongate parietals, the middle fenestra longitudinalis bordered medially by the parietal and frontal and laterally by the dermopterotic, slender and numerous dorsal caudal fulcra, an elongate hyomandibula that is not hourglass shaped, and gill rakers that are short and widely spaced. The second polyodontid species described here is †Pugiopsephurus inundatus n. gen. n. sp. It is diagnosed by a combination of having stellate bones that are exceptionally poorly developed or absent and having a dermopalatine with a medial expansion and lacking an ectopterygoid process. The two species of paddlefishes described in this paper add to the morphological and taxonomic diversity of Polyodontidae. The presence of these taxa within the Hell Creek Formation hints at substantial diversity of polyodontids at this stage of their evolutionary history.

UUID: http://zoobank.org/08cbf8c5-59ba-41fa-9265-8cb762cf7b13

Recent fossil discoveries from New Zealand have revealed a remarkably diverse assemblage of Paleocene stem group penguins. Here, we add to this growing record by describing nine new penguin specimens from the late Paleocene (upper Teurian local stage; 55.5–59.5 Ma) Moeraki Formation of the South Island, New Zealand. The largest specimen is assigned to a new species, Kumimanu fordycei n. sp., which may have been the largest penguin ever to have lived. Allometric regressions based on humerus length and humerus proximal width of extant penguins yield mean estimates of a live body mass in the range of 148.0 kg (95% CI: 132.5 kg–165.3 kg) and 159.7 kg (95% CI: 142.6 kg–178.8 kg), respectively, for Kumimanu fordycei. A second new species, Petradyptes stonehousei n. gen. n. sp., is represented by five specimens and was slightly larger than the extant emperor penguin Aptenodytes forsteri. Two small humeri represent an additional smaller unnamed penguin species. Parsimony and Bayesian phylogenetic analyses recover Kumimanu and Petradyptes crownward of the early Paleocene mainland NZ taxa Waimanu and Muriwaimanu, but stemward of the Chatham Island taxon Kupoupou. These analyses differ, however, in the placement of these two taxa relative to Sequiwaimanu, Crossvallia, and Kaiika. The massive size and placement of Kumimanu fordycei close to the root of the penguin tree provide additional support for a scenario in which penguins reached the upper limit of sphenisciform body size very early in their evolutionary history, while still retaining numerous plesiomorphic features of the flipper.

UUID: https://zoobank.org/15b1d5b2-a5a0-4aa5-ba0a-8ef3b8461730

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

The study of the cranial endocast provides valuable information to understand the behavior of an organism because it coordinates sensory information and motor functions. In this work, we describe for the first time the anatomy of the encephalon of an early Miocene pan-octodontoid caviomorph rodent (Prospaniomys priscus Ameghino, 1902) found in the Argentinean Patagonia, based on a virtual 3D endocast. This fossil rodent has an endocast morphology here considered ancestral for Pan-Octodontoidea and other South American caviomorph lineages, i.e., an encephalon with anteroposteriorly aligned elements, mesencephalon dorsally exposed, well-developed vermis of the cerebellum, and rhombic cerebral hemispheres with well-developed temporal lobes. Prospaniomys Ameghino, 1902 also has relatively small olfactory bulbs, large paraflocculi of the cerebellum, and low endocranial volume and degree of neocorticalization. Its encephalization quotient is low compared with Paleogene North American and European noncaviomorph rodents, but slightly higher than in several late early and late Miocene caviomorphs. Paleoneurological anatomical information supports the hypothesis that Prospaniomys was a generalist caviomorph rodent with terrestrial habits and enhanced low-frequency auditory specializations. The scarce paleoneurological information indicates that several endocast characters in caviomorph rodents could change with ecological pressures. This work sheds light on the anatomy and evolution of several paleoneurological aspects of this particular group of South American rodents.

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

Chiniquodontidae is a family of nonmammaliaform probainognathian cynodonts with occurrences in the Middle to Late Triassic of Africa and South America (Brazil and Argentina). The history of the family is marked by changes in composition and revisions due to the poor preservation of the first discovered specimens. Currently, Chiniquodontidae includes two genera and six species. Here, we apply the computed tomography (CT-scan) method to describe and revise the morphology of one of the best-preserved chiniquodontid specimens (MCP 1600 PV, holotype of ‘Probelesodon’ kitchingi Sá-Teixeira, 1982) from Brazil, which allowed the visualization of sutures and the segmentation of each bone. The CT-scan data also revealed some previously undescribed traits due to the presence of matrix covering structures of the skull. In addition, the morphological comparisons revealed differences of MCP 1600 PV in relation to the other chiniquodontids, and eight discrete cranial features not related to ontogeny indicate that ‘Probelesodon’ kitchingi might not be included within Chiniquodon theotonicus Huene, 1936. Therefore, we consider the species valid but propose the new combination Chiniquodon kitchingi. Moreover, the results obtained highlighted the necessity of a reassessment of the family, as well as the description of new specimens, morphometric analyses, and use of CT-scan, when feasible.