Pharkidonotus megalius n. sp. is described from the Desmoinesian of the Mud Springs Mountains, New Mexico. This unusually large bellerophontid reaches a length of at least 90 mm and is also characterized by a prominent high median ridge bearing a narrow selenizone and subdued collabral ornamentation on the whorl surface at all stages of growth. Its size and subdued ornamentation are quite different from that of the two previously known North American Pennsylvanian species of Pharkidonotus. Although steinkerns of comparable size have been reported from several localities in North America, the specimens discussed here are the first relatively well preserved shells of very large Pennsylvanian bellerophontids to be described.

A single specimen of Hazelia palmata Walcott, 1920, was collected from the Middle Cambrian Marjum Formation near Marjum Pass, in the central House Range, western Utah. This is a first occurrence of the species outside the Burgess shale region of British Columbia, Canada. The flattened oval impression of the monaxonid demosponge shows characteristic tufts and spicule structures of the species.

A new species of Orthoptera, Parapleurites morrisonensis, is described from the upper Jurassic Morrison Formation of Colorado, USA. This is the first insect described from the Morrison Formation and the first orthopteran described from the Jurassic of North America. No other members of the family Locustopsidae have been described in North America previously, and the other species of Parapleurites are only known from Siberia. The lack of Jurassic Orthoptera in North America is likely due to a combination of taphonomic variables and collector bias. The discovery of Parapleurites morrisonensis and the potential for finding other Jurassic Orthoptera are important to understanding the evolution of this diverse and widely distributed group.

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.

A late Pliocene (3.36–3.06 Ma) exposure of the Tangahoe Formation on the North Island of New Zealand preserves close fossil relatives of many extant seabird clades. Here, we report an extinct member of the little penguin (Eudyptula Bonaparte, 1856) lineage from the Tangahoe Formation—the smallest extinct crown penguin yet known. Eudyptula wilsonae n. sp. is based on the nearly complete skulls of an adult and a fledged but immature individual. Both skulls show more slender proportions than modern little penguins and precede genome-derived estimates for the divergence between Eudyptula minor minor Forster, 1781 (endemic to New Zealand) and Eudyptula m. novaehollandiae Stephens, 1826 (native to Australia and recently established in New Zealand). This raises the possibility that the fossil taxon represents a lineage directly ancestral to extant little penguins. Our results support a Zealandian origin for little penguins, with subsequent Pleistocene dispersal to Australia and a more recent Holocene range expansion of Eudyptula m. novaehollandiae back into New Zealand.

UUID: https://zoobank.org/a415f70a-e441-4920-85e0-75f6457577ea

Evidence from the earliest-known crinoids (Tremadocian, Early Ordovician), called protocrinoids, is used to hypothesize initial steps by which elements of the calyx evolved. Protocrinoid calyces are composed of extraxial primary and surrounding secondary plates (both of which have epispires along their sutures) that are unlike those of more crownward fossil and extant crinoids in which equivalent calycinal plating is strongly organized. These reductions inspired several schemes by which to name the plates in these calyces. However, the primary-secondary systems seen in protocrinoids first appeared among Cambrian stem radial echinoderms, with primaries representing centers around which secondaries were sequentially added during ontogeny. Therefore, the protocrinoid calyx represents an intermediate condition between earliest echinoderms and crownward crinoids. Position and ontogeny indicate certain primaries remained as loss of secondaries occurred, resulting in abutting of primaries into the conjoined alternating circlets characteristic of crinoids. This transformative event included suppression of secondary plating and modification or, more commonly, elimination of respiratory structures. These data indicate subradial calyx plate terminology does not correspond with most common usage, but rather, supports an alternative redefinition of these traditional expressions. Extension and adoral growth of fixed rays during calyx ontogeny preceded conjoined primaries in earliest crinoids. Restriction with modification or elimination of calyx respiratory structures also accompanied this modification. Phylogenetic analyses strongly support crinoid origination from early pentaradiate echinoderms, separate from blastozoans. Accordingly, all Tremadocian crinoids express a distinctive aggregate of plesiomorphic and apomorphic commonalities; all branch early within the crinoid clade, separate from traditional subclass-level clades. Nevertheless, each taxon within this assemblage expresses at least one diagnostic apomorphy of camerate, cladid, or disparid clades.

The calcareous skeletons of 17 species of Triassic demosponges from the northeastern Dolomites have been analyzed for microstructure and diagenesis. The four microstructures recognized (irregular, spherulitic, penicillate aragonitic, and homogeneous granular Mg calcite) are described in terms of mineralogy; shape, dimension, and arrangement of microstructural elements; mode of growth; and possible biomineralization. The diagenesis in these sponge carbonate skeletons is of an aggrading type that occurred in diagenetic units, semi-closed systems, delineated by organic phragmas, which controlled the flux of diagenetic fluids. We tentatively interpret these phragmas as the remains of water-insoluble macromolecules for space delineation during the biomineralization process. In the aragonitic skeletons the preservation grade is correlated with Sr content, and the replacement of aragonite by calcite is marked by a Sr value around 4,000 p.p.m. Calcitized aragonite still retains a detectable amount of Sr. In Mg calcite skeletons the continuous and regular increase of grain size is inversely correlated with Mg content and directly with the distance from the organic phragmas.

Zigzagopora wigleyensis n. gen. n. sp. is an Upper Ordovician (Sandbian, early Caradoc) cyclostome bryozoan from the Arbuckle Mountains of Oklahoma, USA. It has runner-type colonies characterized by a mostly uniserial, geniculate arrangement of monomorphic zooids that bud alternately left and right, producing a zig-zag pattern of growth. This new genus has calcified interior walls and non-pseudoporous exterior walls. It is thus most likely affiliated with the paleotubuliporine family Sagenellidae, despite superficial similarities with the corynotrypid cyclostomes with which it co-occurs.

The new genus Tanycythere (Crustacea, Ostracoda) is described from the Jurassic (Hettangian–Bathonian) of western and central Europe based on T. caudata new species, T. praecaudata praecaudata new subspecies, T. praecaudata parallela new subspecies, T. duartei new species, T. posteroelongata new species, and T. wattonensis new species. Another new species, T. procera, is described from the Lower Cretaceous (?Barremian). Recognition of a new genus comprising six new species and subspecies indicates that overall Jurassic and Early Cretaceous ostracod biodiversity is underestimated.

A new crinoid association reported from the Kope Formation (Katian, Ordovician) of northern Kentucky and southwestern Ohio changes the model for facies distribution of crinoids along an Ordovician onshore-offshore depth gradient. Glyptocrinus nodosus n. sp., Plicodendrocrinus casei (Meek, 1871), Cincinnaticrinus varibrachialis (Warn and Strimple, 1977), and Ectenocrinus simplex (Hall, 1847) are reported from a suspension-feeding assemblage with 26 taxa. This assemblage developed above an argillaceous packstone with most of the fossils preserved in shale. The fauna was comprised principally of secondary epifaunally tiered suspension feeders, deposit feeders, and predators. This is the first reported occurrence of Glyptocrinus Hall, 1847 and Plicodendrocrinus Brower, 1995 from the Kope Formation (lower Cincinnatian), and Glyptocrinus is represented by a new species, G. nodosus. Also, this is the first report of pinnulate camerate crinoids from the deep-water facies of the Kope Formation. Thus, deep-water Cincinnatian crinoid assemblages were comprised of disparids, cladids, and camerates; and the assemblage was characterized by a variety of filtration fan types for acquisition of resources.

Bulk maceration of unconsolidated Potomac Group sediments collected from 12 samples of middle–late Albian and early Cenomanian age on the Elk Neck Peninsula, Maryland, yielded 14 genera and 18 species of megaspores. The identified megaspores have affinities to both heterosporous ferns, e.g., Arcellites and Molaspora, and to heterosporous lycopsids, e.g., Erlansonisporites, Minerisporites, and Paxillitriletes. Morphological variation within species is high, and most morphotypes differ from established taxa in one or more characters. Two new species—Dijkstraisporites praetextus and Verrutriletes diversus—are proposed. The composition of these middle–late Albian and early Cenomanian floras is a mixture of taxa previously recorded from Aptian and Santonian floras in North America. In addition, a fragmentary strobilus of selaginellalean affinity containing a megaspore and microspores is described.

Bryozoans from the Dapingian (Middle Ordovician) of the Baltic paleobasin remain poorly studied and their taxonomic composition is unclear. In this paper, three bryozoan taxa, a bifoliate cystoporate Planopora volkhovensis n. gen. n. sp., a trepostome Hemiphragma insolitum n. sp., and an esthonioporate Esthoniopora clara Koromyslova, are described from Dapingian deposits of an unusual clayey-calcareous Hecker-type mudmound on the right bank of the Volkhov River in Leningrad Oblast’, north-western Russia. Combined X-ray microtomography, scanning electron microscopy, and light microscopy of thin sections were used to characterize their morphology. Analysis of the stratigraphic distribution of early cystoporate bryozoans suggests that Planopora n. gen. is the oldest сystoporate bryozoan with an erect, bifoliate colony. The growth modes of these bryozoans are discussed. The colonies of P. volkhovensis n. gen. n. sp. and E. clara have an attachment structure, a holdfast, at their base, probably indicating their attachment to sponge spicules. The bryozoan H. insolitum n. sp. produced rod-like colonies, formed by overgrowing the problematic tubular fossil Sphenothallus Hall. It can be assumed that sponges with unfused siliceous spicules and individuals of Sphenothallus were numerous on the surface of the mudmound during its formation and provided a suitable substrate for settlement of bryozoan larvae.

UUID: http://zoobank.org/5715872a-8c61-446e-9413-d713fdef1a08

A new locality for low-latitudinal, Early Triassic fishes was discovered in the Candelaria Hills, southwestern Nevada (USA). The fossils are derived from the lower Candelaria Formation, which was deposited during the middle–late Dienerian (late Induan), ca. 500 ka after the Permian-Triassic boundary mass extinction event. The articulated and disarticulated Osteichthyes (bony fishes), encompassing both Actinistia (coelacanths) and Actinopterygii (ray-fins), are preserved in large, silicified concretions that also contain rare coprolites. We describe the first actinopterygians from the Candelaria Hills. The specimens are referred to Pteronisculus nevadanus new species (Turseoidae?), Ardoreosomus occidentalis new genus new species (Ptycholepidae), the stem neopterygian Candelarialepis argentus new genus new species (Parasemionotidae), and Actinopterygii indet. representing additional taxa. Ardoreosomus n. gen. resembles other ptycholepids, but differs in its more angulate hyomandibula and lack of an elongate opercular process. Candelarialepis n. gen. is one of the largest parasemionotids, distinguished by its bipartite preopercle and scale ornamentation. Presented new species belong to genera (Pteronisculus) or families (Ptycholepidae, Parasemionotidae) that radiated globally after the mass extinction, thus underlining the striking similarities between Early Triassic (pre-Spathian) osteichthyan assemblages. The current data suggest that the diversity of low-latitudinal, Early Triassic bony fishes may have been greater than indicated thus far by the fossil record, probably due to sampling or taphonomic failure. All 24 fossils from the Candelaria Hills represent mid-sized or large osteichthyans, confirming the obvious absence of very small species (≤ 10 cm adult body length) in the beginning of the Mesozoic Era—even in low latitudes.

UUID: http://zoobank.org/6a66ac96-d6b7-4617-94db-5a93cdb14215

Few bryozoans have been described from the Cretaceous Western Interior Seaway (WIS), which is consistent with the low diversity of other typically stenohaline groups in this large expanse of relatively shallow marine water. Here we describe a new cheilostome bryozoan, Conopeum flumineum n. sp., based on well-preserved material from the Campanian Judith River Formation of the Upper Missouri River Breaks National Monument in north-central Montana. The new species shows strong morphological similarities with Conopeum seurati, a Recent species that is often categorized as brackish, but which is euryhaline and can also be found in marine and stenohaline environments. The new Campanian bryozoan species was found in a locality also containing fragmentary remains of dinosaurs and other terrestrial vertebrates, as well freshwater mollusks and terrestrial plant debris. The sedimentology and facies associations of the fossil-bearing site suggest that the depositional setting was a swamp or tidally influenced fluvial backwater on the Judith River coastal plain. The proximity of the site to the western shoreline of the WIS presumably made it susceptible to occasional marine flooding during storms or extreme tides. Previous occurrences of Conopeum in the Cretaceous of the Western Interior have also been associated with dinosaur remains, corroborating the very nearshore and at times even ‘upstream’ distribution of this euryhaline genus.

UUID: http://zoobank.org/bb1fdc8a-5017-44c5-9251-9e24ef3995e3.

South Canyon, located in the New Pass Range in central Nevada (Fig. 1), is one of the most important localities for Upper Triassic marine invertebrates in North America. This site yields very rich ammonoid faunas, as well as cnidarians (Muller, 1936; Stanley, 1979; Roniewicz and Stanley, 1998), foraminifers (Gazdzicki and Stanley, 1983), bivalves (Waller and Stanley, 1998, 2005; Hopkin and McRoberts, 2003), and brachiopods.

Rugose and tabulate corals from the Lower Permian (Wolfcampian) part of the Bird Spring Group in the Providence Mountains, San Bernardino County, southeastern California, comprise eight species in eight genera. Heritschioides mckassoni n. sp. is the lowest stratigraphic record for this index genus on the undoubted shelf of western North America. Paraheritschioides applegatei n. sp. is the first record for the genus in southern California. Neomultithecopora providensis n. sp. is a second species for the genus in the southern Great Basin. The other five species provide close ties to previously described faunas from the Spring Mountains and the Arrow Canyon Range of southwestern and southeastern Nevada. The combined Wolfcampian coral faunas of these three areas are somewhat closer at the genus and species level to the McCloud Limestone Wolfcampian faunas of northern California than to the Wolfcampian shelf faunas in east-central Nevada. Additional species present in the combined faunas are known originally from the Wolfcampian of central Nevada and Kansas and a genus is not otherwise known south of British Columbia. The faunas suggest a subprovince of the Durhaminid Coral Province for the southern California and southern Nevada area and perhaps imply partial isolation from the more northerly parts of the province by land barriers such as the Antler Highlands.

New material of the poraspid Alainaspis platyrhina Elliott and Dineley has made it possible to provide a more extensive reconstruction of this animal. It is now clear that an elongated branchial opening was present below the lateral lamina and that a well-developed post-branchial lobe was present. No branchial plate has been identified and this may have been fused to the dorsal shield, forming the ventral lamina. The lateral lamina is now seen to terminate in a rounded angle before the posterior margin of the shield. Several ventral shields with similar ornament are identified as belonging to A. platyrhina. These are broad and deep and are consistent with the original interpretation of this animal as a filter feeder. One similar shield with a coarser ornament is tentatively assigned to the closely related species Boothiaspis ovata Broad.

Triassic lungfish (Dipnoi) have been extensively documented from the Gondwanan continental and marine shelf deposits of Africa and Madagascar (Teixeira, 1949; Lehman et al., 1959; Beltan, 1968; Martin, 1979, 1981; Kemp 1996), Australia (Kemp, 1993, 1994, 1997a, 1998), India (Jain et al, 1964; Jain, 1968), and Antarctica (Dziewa, 1980). Numerous records also exist from Laurasian landmasses including Europe (Agassiz, 1838; Schultze, 1981), North America (Case, 1921) and central and eastern Asia (Liu and Yeh, 1957; Vorobyeva, 1967; Martin and Ingavat, 1982). By comparison, nothing is known of contemporary lungfish fossils from the Middle East. Thus, the recent recovery of a single tooth plate representing a new geographic occurrence of the genus Ceratodus Agassiz, 1838 from paralic marine deposits of the Jilh Formation, a latest Anisian to lower Carnian unit that crops out along the eastern margin of the Proterozoic Arabian Shield in central Saudi Arabia (Fig. 1), is significant because it provides the stratigraphically oldest record of dipnoans from the Arabian Peninsula.

Relative differences between environmentally controlled variation and genetically controlled variation are important when investigating morphologic variation in general, especially when establishing species concepts. The colonial nature of bryozoans provides a means for distinguishing between the two sources; variation can be partitioned into within-colony (microenvironmental) and among-colony (environmental + genetic) components. For the Paleozoic order Cryptostomata, biologically and taxonomically significant morphologic characters are well defined and methods for recognizing morphotaxa are well established.

The importance of within-colony variation to the morphometric treatment of fenestrate species was assessed after the discovery of a large specimen of Hemitrypa sp. Variation within the colony was compared to variation among and within two congeneric species. The distribution of study segments across the colony allowed assessment of variation both along the growth axis and laterally between segments of approximately equivalent generational age. Repeatability of methods was assessed using data measured independently from identical segments by three workers.

Variation within the large colony is less than variation among congeneric species, indicating that genetic differences among species exceed variation resulting from combined phenotypic and genotypic sources within species. Neither astogenetic nor ontogenetic morphologic gradients are recognized. Variation between data collected from identical segments by pairs of workers falls within the range of variation for the entire colony. Thus, multiple workers can reproduce data to the finest level of meaningful resolution. Cryptostome morphospecies concepts are validated.

The potential for partitioning genotypic versus environmental variation in reduced, multidimensional morphospace is reinforced. Studies of microevolution and speciation may be designed that account for these factors.