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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.
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.
Gennaeocrinus tariatensis new species is an Emsian (Devonian) monobathrid crinoid described from the Tarvagatay Terrane of Mongolia and part of the Central Asian Orogenic Belt. The Tarvagatay Terrane is an arc terrane that accreted to the southern margin of the Siberian Craton. Gennaeocrinus tariatensis was collected from the Emsian Tariat Formation, a terrigenous sequence of conglomerates, sandstones, and siltstones. Associated faunas include brachiopods, molluscs, and rare tabulate corals. Although Gennaeocrinus is well known from the Emsian–Givetian of North America, this is the first occurrence of the genus outside Laurussia. Mongolia is a large country with many terranes having varied paleogeographic, sedimentological, and tectonic histories; but reports of Paleozoic echinoderms are rare. The crinoid occurrence from the Tariat Formation is from the same age as previously described Emsian crinoids from the Chuluum Formation but differs significantly in sedimentology, paleogeography, and paleolatitude.
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.
The youngest species of Amphoracrinus, A. tenax new species, is described from the Muldraugh Member of the Borden Formation (early Viséan) of north-central Kentucky. With this new occurrence, both the oldest and youngest named species of Amphoracrinus are from North America. Numerous Tournaisian and Viséan crinoid faunas are documented in the United States, but only four are known to contain Amphoracrinus. Morphological analysis indicates that A. tenax is more closely aligned with species from China than with species from Western Europe or other species from North America, where Amphoracrinus was most diverse and abundant, which has implications for understanding paleogeographic dispersal. The holotype of A. tenax was partially disarticulated on the seafloor before burial, and final burial occurred early during disarticulation. The relative state of disarticulation from pinnules to columnals suggests that plates bound only with ligaments disarticulated as a function of surface area of ligaments binding an articulation.
The Cincinnatian (Katian) of the Cincinnati Tri-State area is widely regarded as one of the most fossiliferous sections known (Meyer and Davis, 2009). Echinoderms from these strata include well-described asteroids, crinoids, cyclocystoids, edrioasteroids, glyptocystoids, mitrates, and ophiuroids. John Pope discovered a partially articulated echinoderm in float from the Fairview Formation that does not correspond to any known Cincinnatian echinoderm. Although mentioned in Ubaghs (1966, as a presumable personal communication from Pope, 1960), Haude and Langenstrassen (1976), Reich (2001), and Reich and Haude (2004), this specimen at the Cincinnati Museum Center (CMCPIP 51316) has neither been described nor illustrated; yet, these authors attributed it to Volchovia Hecker, 1938 in the Class Ophiocistioidea. Questions swirl around this fossil: what is its complete morphology; does it belong to Volchovia; whether or not it can be assigned to Volchovia, is it an ophiocistioid? The first step to understand this enigmatic echinoderm is to illustrate and describe the specimen, which is the objective of this note.
The history of life on earth is largely reconstructed from time-averaged accumulations of fossils. A glimpse at ecologic-time attributes and processes is relatively rare. However, the time-sensitive and predictability of echinoderm disarticulation makes them model organisms to determine post-mortem transportation and allows recognition of ecological-time data within paleocommunity accumulations. Unlike many other fossil groups, this has allowed research on many aspects of echinoderms and their paleocommunities, such as the distribution of soft tissues, assessment of the amount of fossil transportation prior to burial, determination of intraspecific variation, paleocommunity composition, estimation of relative abundance of taxa in paleocommunities, determination of attributes of niche differentiation, etc. Crinoids and echinoids have received the most amount of taphonomic research, and the patterns present in these two groups can be used to develop a more thorough understanding of all echinoderm clades.
The Brechin Lagerstätte of southern Ontario contains an exceptionally diverse and well-preserved Late Ordovician (Katian) crinoid fauna. We describe four genera and eight species of camerate crinoids from the Brechin Lagerstätte, including six new species. Consequently, the total diversity of the fauna now stands at 27 genera and 39 nominal species, thereby making it the most taxonomically diverse Ordovician crinoid fauna known. Taxa described include the diplobathrid Pararchaeocrinus kiddi new species and the monobathrids Glyptocrinus ramulosus Billings, 1856, Periglyptocrinus priscus (Billings, 1857a), Periglyptocrinus astricus new species, Periglyptocrinus kevinbretti new species, Periglyptocrinus mcdonaldi new species, Periglyptocrinus silvosus new species, and Abludoglyptocrinus steinheimerae new species. We summarize the taxonomic composition, diversity, and abundance distribution of all known crinoids from the Brechin Lagerstätte to better characterize the paleoecological structure and complexity of the community. We establish that the fauna is dominated by the subclass Pentacrinoidea, both in terms of abundance and species richness. In addition, we analyze species-level abundance data using Relative Abundance Distribution (RAD) models to evaluate the ecological complexity of the paleocommunity. We found that community structure of the Brechin Lagerstätte is best explained by an ecologically ‘complex’ RAD model, which suggests that species partitioned niches along multiple resource axes and/or the presence of multiple ecological ways of life. These results indicate that the Brechin Lagerstätte is significant not only for being the most taxonomically diverse Katian crinoid assemblage, but also for being an early ecologically complex fauna that developed in the wake of the Great Ordovician Biodiversification Event.
Upper Ordovician (Katian) strata of the Lake Simcoe region of Ontario record a spectacularly diverse and abundant echinoderm fauna known as the Brechin Lagerstätte. Despite recognition as the most taxonomically diverse Katian crinoid paleocommunity, the Brechin Lagerstätte has received relatively little taxonomic study since Frank Springer published his classic monograph on the “Kirkfield fauna” in 1911.
Using a new collection of exceptionally preserved material, we evaluate all dicyclic inadunate crinoids occurring in the Brechin Lagerstätte, which is predominantly comprised of cladids (Eucladida and Flexibilia). We document 15 species across 11 genera, including descriptions of two new genera and four new species. New taxa include Konieckicrinus brechinensis n. gen. n. sp., K. josephi n. gen. n. sp., Simcoecrinus mahalaki n. gen. n. sp., and Dendrocrinus simcoensis n. sp.
Although cladids are not commonly considered major components of the Early Paleozoic Crinoid Macroevolutionary Fauna, which is traditionally conceived as dominated by disparids and diplobathrid camerates, they are the most diverse major lineage of crinoids occurring in the Brechin Lagerstätte. This unexpected result highlights the important roles of specimen-based taxonomy and systematic revisions in the study of large-scale diversity patterns.
Three new Llandovery (early Silurian) crinoids from Estonia provide an improved understanding of the paleogeographic aspects of the crinoid diversification following the end-Ordovician extinctions. The new taxa are Euspirocrinus hintsae new species (Rhuddanian eucladid), Oepikicrinus perensae new genus new species (Aeronian camerate), and Rozhnovicrinus isakarae new genus new species (Aeronian eucladid). This brings the total of described Llandovery crinoids in Estonia to eight nominal species and a further three taxa in open nomenclature. The Rhuddanian radiation in Baltica mirrored that on Laurentia and Avalonia and was dominated by Ordovician clades that continued to diversify during the Silurian. Known Aeronian crinoids from Estonia continue these clades, whereas new clades diversified on Laurentia and Avalonia. However, by the Wenlock, a largely cosmopolitan fauna existed on Laurentia, Avalonia, and Baltica.
Living crinoids are exclusively passive suspension feeders and benthic as adults. However, in the past they adapted to a broad range of ecological niches. For instance, the stratigraphically important middle Paleozoic scyphocrinoids are hypothesized to have been planktonic, employing their inferred gas-filled globular, chambered structure at the distal end of the stem, the so-called lobolith, as a buoyancy device with the crinoid calyx suspended below it. Here, we evaluate this hypothesis using evidence from skeletal micromorphology and theoretical biomechanical modeling. Lobolith walls are typically composed of ossicles, which are exclusively composed of constructional labyrinthic stereom. In plates from the distal side of the lobolith, this stereom extends into microperforate stereom layer, forming wavy ridges and spines. No microscale adaptations for preventing gas leaks and/or ingress of water (such as internal and external imperforate stereom layers) are known. Furthermore, theoretical calculations suggest that the scyphocrinoid tow-net mode of feeding would have resulted in small relative velocities between the towed filter and the ambient water, thus making it an ineffective passive filter feeder. We suggest that the lobolith of these crinoids acted as a modified holdfast rather than as a floating buoy. Its globular shape and distally positioned microspines served as adaptations for living in unconsolidated sediments, analogous to iceberg- and snowshoe-like strategies used by some mollusks and brachiopods. Like modern isocrinids, scyphocrinoids could have maintained an upright feeding posture by extending the distal portion of the stalk along the bottom. In this recumbent posture, the distal part of the stalk with the lobolith might have functioned as a drag anchor. As a consequence of the ~3-m-long stem, even with this posture, the benthic scyphocrinoids could have risen to the highest epifaunal tier in the Paleozoic.
End-Ordovician extinctions had a profound effect on shallow-water benthic communities, including the Crinoidea. Further, recovery after the extinctions resulted in a macroevolutionary turnover in crinoid faunas. Anticosti Island is the most complete Ordovician-Silurian boundary section recording shallow-water habitats. Both new taxa and changes in Anticosti Island stratigraphic nomenclature are addressed herein. New taxa include Becsciecrinus groulxi n. sp., Bucucrinus isotaloi n. sp., Jovacrinus clarki n. sp., Plicodendrocrinus petryki n. sp., Plicodendrocrinus martini n. sp., Thalamocrinus daoustae n. sp., and Lateranicrinus saintlaurenti n. gen. n. sp. The status of Xenocrinus rubus as a boundary-crossing taxon is confirmed, range extensions of several taxa are documented, and the distribution of crinoids with the revised stratigraphic nomenclature is documented.
A new Lower Devonian fauna from the Iberian Chains (NE Spain) is described. Specimens have been collected from the shaley intervals of the Mariposas Formation dated as early Emsian. These include the camerates Acanthocrinus carsli n. sp., Platyhexacrinus santacruzensis n. sp., Culicocrinus breimeri n. sp., Camerata indeterminate, and an indeterminate eucladid. Compared with other faunas from Spain, this represents a low diversity crinoid assemblage that was probably concentrated in shallow, turbid environments. A summary of crinoids previously described from the Spanish Devonian is reported, which indicates that crinoid faunas become progressively more cosmopolitan during the Devonian.
Using polarized light microscopy, the large, triangular or cylindrical second brachial plate of the Petalocrinidae is demonstrated to be a compound brachial formed through fusion of brachial plates along the distal margin of the growing arms. Based on the number of ambulacral bifurcations, brachials from the primibrachitaxis through at least the quintibrachitaxis may have been fused to form this large plate. In Petalocrinus, fused brachials form a second brachial that assumed the same crystallographic orientation, but in Spirocrinus, multidirectional extinctions preserve some of the original multiplate arrangement.
The Kalana Lagerstätte of early Aeronian (Llandovery, Silurian) age in central Estonia preserves a diverse shallow marine biota dominated by non-calcified algae. This soft-tissue flora and decalcified and calcified crinoids are preserved in situ, in a lens of microlaminated, dolomitized micrite interbedded in a sequence of dolomitized packstones and wackestones. Although the Lagerstätte is dominated by non-calcified algae, crinoids (together with brachiopods and gastropods) are among the most common organisms that were originally comprised of a carbonate skeleton. Two new crinoids are described from this unit, Kalanacrinus mastikae n. gen. n. sp. (large camerate) and Tartucrinus kalanaensis n. gen. n. sp. (small disparid). Interestingly, these two crinoids display contrasting preservation, with the more common large camerate preserved primarily as a decalcified organic residue, whereas the smaller disparid is preserved primarily in calcite. Preservation was assessed using elemental mapping of C, Ca, S, and Si. Columns have the highest portion of Ca, once living soft tissue is indicated by C, S was dispersed as pyrite or associated with organics, and Si is probably associated with clay minerals in the matrix. This new fauna increases our understanding of the crinoid radiation on Baltica following Late Ordovician extinctions.
The Brechin Lagerstätte (Katian, Ordovician) from the Lake Simcoe region of Ontario, Canada contains a diverse array of echinoderms. Here, we describe seven disparid and two hybocrinid crinoids (subclass Pentacrinoidea, infraclass Inadunata), including a new disparid species belonging to the Anomalocrinidae (order Homocrinida). In total, the disparids include Anomalocrinus astrictus n. sp.; Cremacrinus guttenbergensis Kolata, 1975; C. inaequalis Billings, 1859; Daedalocrinus bellevillensis Billings, 1883; Eustenocrinus springeri Ulrich, 1925; Iocrinus trentonensis Walcott, 1883; and Isotomocrinus tenuis Billings, 1857b. The hybocrinids include Hybocrinus tumidus Billings, 1857a and Hybocystites problematicus Wetherby, 1880. Previously known from only the holotype, three additional specimens of E. springeri expand our understanding of this unusual crinoid. Nomenclatural acts include: (1) the recommended designation of D. kirki Ulrich, 1925 as a junior synonym of D. bellevillensis is followed; (2) Hybocrinus pristinus Billings, 1858 is designated as a junior synonym of H. tumidus, and previous decisions are followed to retain Hybocystites eldonensis (Parks, 1908) as a junior synonym of H. problematicus; (3) although probably assignable to Anomalocrinus Meek and Worthen, 1865, the aberrant crinoid Glaucocrinus falconeri Parks and Alcock, 1912, and its genus Glaucocrinus Parks and Alcock, 1912, are designated as nomena dubia; (4) Iocrinus similis (Billings, 1857) is also designated as a nomen dubium; and (5) Iocrinus subcrassus torontoensis Fritz, 1925 is designated a junior synonym of I. subcrassus Meek and Worthen, 1865.
A diverse Permian crinoid fauna is reported from the Taiyuan Formation, Dajian Member (Asselian) at Anyang, northeastern Henan Province of the North China Craton. The specimens are well preserved, including articulated crowns and cups. The fauna contains representatives of each of the major Paleozoic crinoid clades: Cladida (including the Flexibilia), Disparida, and Camerata. Identified genera suggest a greater affinity with North American faunas than with Tethyan faunas. Four new species, Neoprotencrinus anyangensis, Ulocrinus qiaoi, Artichthyocrinus limani, and Synbathocrinus chenae, are proposed herein.
The Batocrinidae are characteristic faunal elements in Lower Mississippian shallow-marine settings in North America. Recent delineation of objectively defined genera allows a reexamination of batocrinid species and their distribution in the Fort Payne Formation (early Viséan, late Osagean), a well-studied array of carbonate and siliciclastic facies. The Fort Payne batocrinid fauna has 14 species assigned to six genera, plus hybrid specimens. Magnuscrinus spinosus (Miller and Gurley, 1895a) is reassigned to its original placement in Eretmocrinus. Hybrid specimens (Ausich and Meyer, 1994) are regarded as Eretmocrinus magnificus×Eretmocrinus spinosus. Macrocrinus casualis is the dominant species of Macrocrinus in the Fort Payne, and M. mundulus and M. strotobasilaris are recognized in the Fort Payne Formation for the first time. Magnuscrinus cumberlandensis n. sp. is named, 13 species are designated as junior synonyms, the name for the hybrid specimens is changed to Eretmocrinus magnificus×Eretmocrinus spinosus, and the previous occurrences of two species in the Fort Payne are rejected. The Eastern Interior Seaway was a mixed carbonate-siliciclastic setting with both shallow- and deep-water epicontinental sea facies ranging from relatively shallow autochthonous green shales to deep-water turbidite facies. Dizygocrinus was restricted to shallow-water carbonate and siliciclastic facies, Eutrochocrinus was restricted to shallow-water carbonate facies, and Magnuscrinus was restricted to deep-water facies. Species distributions varied from Abatocrinus steropes, Alloprosallocrinus conicus, Macrocrinus mundulus, and Uperocrinus nashvillae, which occurred throughout the Eastern Interior Seaway, to species that were restricted to a single facies. Eretmocrinus magnificus, Alloprosallocrinus conicus, and Uperocrinus robustus were the dominant batocrinids in the Fort Payne Formation.
The Upper Ordovician (lower Katian) Bobcaygeon and Verulam formations from the Lake Simcoe region of Ontario contain a highly diverse echinoderm assemblage that is herein recognized as a Konservat-Lagerstätte. Although fossil crinoids have long been recognized from these formations, the fauna has not received a comprehensive taxonomic evaluation since Springer’s classic 1911 monograph. Recent extensive collection and preparation of new material from the Bobcaygeon and Verulam formations near Brechin, Ontario recovered numerous exceptionally preserved crinoid specimens with arms, stems, and attachment structures intact. The Brechin Lagerstätte is the most taxonomically diverse Katian crinoid fauna, with more than 20 crinoid genera represented in this collection.
Here, all dicyclic crinoids belonging to subclass Camerata from the Brechin Lagerstätte are evaluated. The following four genera and seven species are described from the fauna, including one new genus and four new species: Reteocrinus stellaris, Reteocrinus alveolatus, Archaeocrinus sundayae n. sp., Archaeocrinus maraensis n. sp., Priscillacrinus elegans n. gen. n. sp., Cleiocrinus regius, and Cleiocrinus lepidotus n. sp. The exceptional preservation of this collection provides an opportunity to describe more fully the morphologic and ontogenetic details of known Ordovician crinoid taxa, to conduct a taxonomic re-evaluation of many species, to describe new taxa, and to provide a framework for subsequent studies of crinoid community paleoecology.