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The family Actinocrinitidae was a significant contributor to the global biodiversity peak of crinoids that occurred during the Mississippian and is referred to as the “Age of Crinoids.” Although the actinocrinitids are a major component of that high diversity, they are also a source of much taxonomic confusion. Previously, generic concepts were not applied equally between Europe and North America creating disparity in the definition of genera. In this contribution, all genera are defined objectively by discrete characters, and the generic assignments of all species are reevaluated. Twenty genera are described. A total of 206 species were evaluated of which 56 species and one taxon in open nomenclature are reassigned to different genera, 21 species are designated as nomina dubia, and three species and one genus are now incertae sedis.
A phylogenetic hypothesis is presented for the relationships of the genera of Actinocrinitidae genera based on a parsimony-based analysis and plotted against stratigraphic ranges. Although groupings were revealed in this analysis, the Actinocrinitidae cannot be readily subdivided into subfamilies. Rapid diversification occurred during the Tournaisian following the Hangenberg Extinction of probable fish predators. The late Devonian (Famennian) occurrence of the highly derived genera Abactinocrinus and Physetocrinus suggests there is a more extensive, but undocumented, evolutionary history for the Actinocrinitidae during the Devonian.
Several Jurassic pterioid bivalve species have been referred to Parainoceramus Cox by different authors, yet this has proved inadequate because the meaning of such genus has been compounded by nomenclatural and idiomatic problems, as well as misinterpretations. Hence, the new genus Parainoceramya is here proposed to accommodate several species previously referred to Parainoceramus, with Crenatula ventricosa J. de C. Sowerby as its type. Permian species originally assigned to Parainoceramus, including the type species, are referred to the genus Kolymia Likharev. All species attributed to Parainoceramus s.l. are reviewed and the new genus is compared with related genera. As here understood, the new genus is first recorded in the Hettangian and attained a cosmopolitan distribution; its last occurrence is probably Berriasian.
Light microscope and scanning electron microscope observations on new material of unicellular microfossils Dictyosphaera macroreticulata and Shuiyousphaeridium macroreticulatum, from the Mesoproterozoic Ruyang Group in China, provide insights into the microorganisms' biological affinity, life cycle and cellular complexity. Gigantosphaeridium fibratum n. gen. et sp., is described and is one of the largest Mesoproterozoic microfossils recorded. Phenotypic characters of vesicle ornamentation and excystment structures, properties of resistance and cell wall structure in Dictyosphaera and Shuiyousphaeridium are all diagnostic of microalgal cysts. The wide size ranges of the various morphotypes indicate growth phases compatible with the development of reproductive cysts. Conspecific biologically, each morphotype represents an asexual (resting cyst) or sexual (zygotic cyst) stage in the life cycle, respectively. We reconstruct this hypothetical life cycle and infer that the organism demonstrates a reproductive strategy of alternation of heteromorphic generations. Similarly in Gigantosphaeridium, a metabolically expensive vesicle with processes suggests its protective role as a zygotic cyst. In combination with all these characters and from the resemblance to extant green algae, we propose the placement of these ancient microorganisms in the stem group of Chloroplastida (Viridiplantae). A cell wall composed of primary and secondary layers in Dictyosphaera and Shuiyouisphaeridium required a high cellular complexity for their synthesis and the presence of an endomembrane system and the Golgi apparatus. The plastid was also present, accepting the organism was photosynthetic. The biota reveals a high degree of morphological and cell structural complexity, and provides an insight into ongoing eukaryotic evolution and the development of complex life cycles with sexual reproduction by 1200 Ma.
A new Burgess Shale-type Lagerstätte is described from the middle Cambrian (Series 3, Drumian) Rockslide Formation of the Mackenzie Mountains, Northwest Territories, Canada. The Rockslide Formation is a unit of deeper water ramp to slope, mixed carbonate, and siliciclastic facies deposited on the northwestern margin of Laurentia. At the fossil-bearing locality, the unit onlaps a fault scarp cutting lower Cambrian sandstones. There it consists of a succession of shale and thick-laminated to thin-bedded lime mudstone, calcareous sandstone, and greenish-colored calcareous mudstone, overlain by shallower water dolostones of the Avalanche Formation, which is indicative of an overall progradational sequence. The Rockslide Formation is of similar age to the Wheeler and Marjum formations of Utah, belonging to the Bolaspidella Biozone. Only two 1 m thick units of greenish mudstone exhibit soft-bodied preservation, with most specimens coming from the lower interval. However, the biota is common but not as diverse as that of other Lagerstätten such as the Burgess Shale in its type area. The shelly fauna is dominated by the hyolith Haplophrentis carinatusMatthew, 1899 along with sparse linguliformean brachiopods, agnostoid arthropods, and ptychoparioid trilobites. The nonmineralized biota includes the macrophytic alga Margaretia dorus Walcott, 1911, priapulid worms, and the carapaces of a number of arthropods. The arthropods belong to Isoxys mackenziensis n. sp., Tuzoia cf. T. guntheriRobison and Richards, 1981; Branchiocaris? sp., Perspicaris? dilatusRobison and Richards, 1981; and bradoriids, along with fragments of arthropods of indeterminate affinities. The style of preservation indicates that most soft parts underwent complete biodegradation, leaving just the more resistant materials such as chitinous arthropod cuticles. The range of preservation and similarity to the coeval biotas preserved in Utah suggests that the composition of this Lagerstätte is probably representative of the community living on the relatively deep-water ramp or slope during middle Cambrian time in Laurentia. This would argue that the extraordinary diversity of the Burgess Shale at Mount Field is anomalous.
Study of new collections of the Wenlock and Pridoli, Silurian, crinoids from Saaremaa, western Estonia, result in taxonomic revision, expanded ranges, and new taxa. Eucalyptocrinites regularis and Periechocrinus laevis are recognized outside of Sweden for the first time. Desmidocrinus laevigatusAusich et al., 2012 is reassigned to Methabocrinus. Because the type species of Methabocrinus was previously known only from glacial sediments, the age and provenance of this genus are constrained for the first time. A new crotalocrinitid, Velocrinus coniculus new genus and species, is described. Although the Pridoli faunas of western Estonia contain only twelve species-level taxa assigned to nine genera, this fauna is one of the four richest Pridoli faunas known.
To investigate the phylogenetic affinity of Yuknessia simplexWalcott, 1919, scanning electron microscopy was applied to the Burgess Shale (Cambrian Series 3, Stage 5) type material and to new material from the Trilobite Beds (Yoho National Park) and specimens from the Cambrian of Utah. On the basis of fine-scale details observed using this approach, including banding structure interpreted as fusellae, YuknessiaWalcott, 1919 is transferred from the algae, where it resided for nearly a century, to the extant taxon Pterobranchia (Phylum Hemichordata). Considered as such, Yuknessia specimens from the Trilobite Beds and Spence Formation (Utah) are amongst the oldest known colonial pterobranchs. Two morphs regarded herein as two different species are recognized from the Trilobite Beds based on tubarium morphology. Yuknessia simplex has slender erect tubes whereas Yuknessia stephenensis n. sp., which is also known in Utah, has more robust erect tubes. The two paratypes of Y. simplex designated by Walcott (1919) are formally removed from Yuknessia and are reinterpreted respectively as an indeterminate alga and Dalyia racemataWalcott, 1919, a putative red alga.
Circum-Laurentian middle Cambrian (Cambrian Series 3) deposits in Greenland and British Columbia yield a new hipponicharionid bradoriid arthropod, Flumenoglacies n. gen., characterized by a comarginal, ramp-like structure which is crested by a continuous lobe. The narrow lobe is the result of the medial fusion of anterior and posterior lobes, seemingly a recurrent theme in hipponicharionid evolution. The type species, F. groenlandica n. sp., is described from the Ekspedition Brae Formation (Drumian Stage) of Peary Land but the description of two unnamed species from slightly older middle Cambrian strata of the Stephen Formation of British Columbia provides additional evidence for the wide distribution of Small Shelly Faunas during the Cambrian.
The earliest fossil gray whale (Eschrichtius) from the eastern North Pacific is reported from the Lower Pleistocene Rio Dell Formation of Humboldt County, Northern California. This specimen, a tympanic bulla and posterior process, is identical in morphology to extant Eschrichtius robustus and differs from Pliocene Eschrichtius sp. from the western North Pacific (Japan). Thus, it suggests that the modern bulla morphology of the gray whale had been acquired by the Early Pleistocene. The absence of fossil Eschrichtius in the Pliocene of the eastern North Pacific may indicate that the extant gray whale lineage originated in the western North Pacific during the Pliocene before invading the eastern North Pacific during the Early Pleistocene. Further discoveries of Plio-Pleistocene gray whale fossils will help test this hypothesis and properly interpret the evolutionary history of eschrichtiid clade.
The Upper Ordovician (Sandbian-Katian) bathyurid trilobite Raymondites Sinclair is revised using new collections from Missouri and Ontario, and archival material from Illinois, Wisconsin, New York, and Ontario. Phylogenetic analysis supports monophyly of Raymondites, but recognition of this genus renders Bathyurus Billings paraphyletic. We treat Raymondites as a subgenus of Bathyurus and label the paraphylum of species traditionally assigned to the latter as Bathyurus sensu lato. Bathyurus (Raymondites) is composed of five previously named species, B. (R.) spiniger (Hall), B. (R.) longispinus (Walcott), B. (R.) ingalli (Raymond), B. (R.) bandifer Sinclair, and B. (R.) trispinosus (Wilson), and two new species, B. (R.) clochensis, and B. (R.) missouriensis; an eighth species is placed in open nomenclature. All species share tuberculate sculpture on the glabella, a relatively short palpebral lobe whose length is less than half of preoccipital glabellar length, and a pygidial outline that is well rounded posteriorly. Aside from the most basal species, B. (R.) longispinus, they also possess occipital spines and, where the pygidium is known, axial pygidial spines.
Rhinoceroses were important in North American mammal faunas from the late middle Eocene to the Miocene, but the group's poor sampling outside the High Plains and eastern Rocky Mountain regions during their early evolution significantly hinders understanding of their biogeography. This limited geographic sampling is particularly true of early—middle Oligocene time, with the vast majority of Whitneyan localities occurring in the White River Badlands of South Dakota. Thus, any rhinocerotid from outside the High Plains during this period is significant. We describe two new rhinocerotid specimens from the middle Oligocene Steamboat Formation of the northeastern Warner Mountains of California. Although the Steamboat Formation is well known for fossil plants, this is the first report of mammalian fossils from the area: an isolated lower molar recovered in 1974 but not previously described or illustrated, and a mandibular fragment recovered approximately 20 years later and bearing two molar teeth, most likely pertaining to the same taxon and horizon. The lack of distinctive morphological characters suggests both fossils be conservatively referred to Rhinocerotidae incertae sedis. Based on published tooth measurement data, Trigonias osborni represents the closest size match, but that species is currently only known from the Chadronian. Similarly, the Whitneyan taxon Diceratherium tridactylum is approximately the right size, but is currently only known from the High Plains and its presence in California would expand its geographic range substantially. Of greatest importance here is that sediments of the eastern Warner Mountains may represent a largely unexplored locale for early—middle Oligocene fossil vertebrates, and may yield important future finds.
The bryozoan family Chlidoniopsidae Harmer, 1957 is reviewed in relation to a new Paleogene European fossil, Celiopsis vici new genus and new species. It differs from the type and only other genus of the family in having longer internodes with up to three zooids, shorter proximal caudae, and, more importantly, suture lines that unequally divided the umbonuloid frontal shield and basal (abfrontal) wall (and the hypostegal coelom in life) into sectors, analogous to the situation in the lepralioid-shielded Prostomariidae and Urceoliporidae. Unlike Prostomaria and Urceolipora, and like Chlidoniopsis, Celiopsis is uniserial. The suture lines in Celiopsis were lines of insertion (attachment) of epithecal membranes in life and each sector has its own longitudinal series of septular pores, sometimes doubled. Miocene to Recent Chlidoniopsis contains two species, and Eocene—Oligocene Celiopsis contains three species. The geographic distribution gives evidence of origination of the family in the Paratethys of Europe, with southeastwards migration to Australia and the tropical western Pacific. The temporal distribution suggests two macro-evolutionary trends—from multizooidal to unizooidal internodes, and from a broader area of basal wall, with a division into separate cryptocystal fields, to a narrower basal wall with no such division.
Gladius-bearing coleoids are rare in the fossil record. For the Cretaceous period, these cephalopods are mainly recorded in a few Lagerstätten in Lebanon (Haqel, Hajoula, En Nammoura, and Sahel Aalma). Here, we study 16 specimens of gladius-bearing coleoids from these Upper Cretaceous Lebanese Lagerstätten to investigate their taxonomic diversity. Besides two species that were already reported (Dorateuthis syriaca and Glyphiteuthis libanotica), one new species is identified in the Cenomanian site of Hajoula: Rachiteuthis acutali n. sp., as well as another form of Glyphiteuthis from En Nammoura. Several studied specimens exhibit well-preserved soft-part characters. Among them, we document for the first time two transverse rows of sessile suckers in D. syriaca and we confirm the absence of tentacles, as well as the presence of a crop in this species. This strongly supports the phylogenetic proximity of D. syriaca with modern vampyropods rather than with modern decabrachians. In turn, the similarity in gladius morphology between this taxon and modern squids is regarded as convergent.
The ankylosaurine Pinacosaurus is one of the best known ankylosaur to date in terms of the number and preservational quality of specimens. Juvenile to sub-adult postcrania collected by the Soviet-Mongolian Paleontological Expedition from the Upper Cretaceous Alagteeg Formation at Alag Teeg, Mongolia can be assigned to Pinacosaurus grangeri based on discrete cranial characters. One individual is significantly larger than the others and demonstrates delayed fusion of postcranial elements with the earliest occurring between dorsal ribs and vertebrae. The robustness of forelimb elements is positively allometric with respect to their length, indicating weight-bearing relationship. Such length-dependent correlations are not seen in the hind limbs. Finally, incipient cervical half rings suggest a developmental pathway of outgrowths from the underlying band combined with fusion of overlying osteoderms.
A rare diminutive belemnite, Pumiliobelus n. gen., is described based on the new species P. haigi and P. tumidus from the upper Gearle Siltstone (Cenomanian) of the Carnarvon Basin, Western Australia. No smaller belemnites than these species are known. That they are based on adult specimens rather than juveniles is supported by a large sample, some 80 specimens, of P. haigi which collectively show a proportional range in size typical of belemnite species in general, a group in which juveniles are conspicuously lacking in the fossil record. Rostral morphology places Pumiliobelus in the Dimitobelidae, a distinctively Austral family of Aptian—Maastrichtian age, which became progressively restricted to high latitudes through the late Cretaceous. Each species of Pumiliobelus is known from a single locality where it co-occurs with the long-ranging (Albian—Cenomanian) Dimitobelus diptychus Whitehouse of widespread distribution at mid to high latitudes in the Southern Hemisphere. Dwarfism in the Dimitobelidae, as expressed by Pumiliobelus, is considered to reflect adaptive evolution to engage opportunistic life strategies that favored small body size and rapid population turnover. The rare occurrence of dwarf Dimitobelidae indicates that such strategies were limited in both geographic range and duration. Dwarfism expressed by Pumiliobelus may relate to paedomorphosis induced by rising seawater temperatures in the mid Cretaceous.
Llanocystis wilbernsensis n. gen. n. sp. (Eocrinoidea, Echinodermata) is described based on three specimens from the Furongian Point Peak Shale Member of the Wilberns Formation in central Texas. It displays a unique morphology including a very long stem constructed with holomeric columnals, few feeding appendages, and a polyplated theca. The specimens occur in an intraformational conglomerate deposited in a proximal carbonate platform environment and represent an example of the poorly documented “pelmatozoan” radiation that occurred in proximal facies by the end of the Cambrian.