Phylogeny and evolutionary history of diplobathrid crinoids (Echinodermata)

Order Diplobathrida is a major clade of camerate crinoids spanning the Ordovician–Mississippian, yet phylogenetic relationships have only been inferred for Ordovician taxa. This has hampered efforts to construct a comprehensive tree of life for crinoids and develop a classification scheme that adequately reflects diplobathrid evolutionary history. Here, I apply maximum parsimony and Bayesian phylogenetic approaches to the fossil record of diplobathrids to infer the largest tree of fossil crinoids to date, with over 100 genera included. Recovered trees provide a framework for evaluating the current classification of diplobathrids. Notably, previous suborder divisions are not supported, and superfamily divisions will require significant modification. Although numerous revisions are required for families, most can be retained through reassignment of genera. In addition, recovered trees were used to produce phylogeny‐based estimates of diplobathrid lineage diversity. By accounting for ghost lineages, phylogeny‐based richness estimates offer greater insight into diversification and extinction dynamics than traditional taxonomy‐based approaches alone and provide a detailed summary of the ~150 million‐year evolutionary history of Diplobathrida. This study constitutes a major step toward producing a phylogeny of the Crinoidea and documenting crinoid diversity dynamics. In addition, it will serve as a framework for subsequent phylogeny‐based investigations of macroevolutionary questions.     

Body-size trends of cyrtocrinids (Crinoidea,Cyrtocrinida)

Cyrtocrinids (Cyrtocrinida) are fully sessile post-Palaeozoic crinoids (Crinoidea) of unusual morphology exhibiting complex diversity dynamics and evolutionary history. To date, however, no study has ever examined the macro-evolutionary patterns of body-size trends in these crinoids. A compilation of a body-size dataset for cyrtocrinid genera revealed a trend of increasing size throughout their evolutionary history. A maximum-likelihood approach showed that the observed trend is best characterized by a general random walk. Recorded body-size pattern is thus consistent with the Cope-Depéret's rule implying the existence of active, directional selective pressures towards larger body-sizes. The case provides a rare example of directional body-size trend in the fossil record.     

Glycomics of a novel type-2 N-acetyllactosamine-specific lectin purified from the feather star, Oxycomanthus japonicus (Pelmatozoa: Crinoidea)

A lectin - designated OXYL for the purposes of this study that strongly recognizes complex-type oligosaccharides of serum glycoproteins - was purified from a crinoid, the feather star Oxycomanthus japonicus, the most basal group among extant echinoderms. OXYL was purified through a combination of anion-exchange and affinity chromatography using Q-sepharose and fetuin-sepharose gel, respectively. Lectin was determined to be a 14-kDa polypeptide by sodium dodecyl sulphate-polyacrylamide gel electrophoresis under reducing conditions. However, 14-kDa and 28-kDa bands appeared in the same proportion under non-reducing conditions. Gel permeation chromatography showed a 54-kDa peak, suggesting that lectin consists of four 14-kDa subunits. Divalent cations were not indicated, and stable haemagglutination activity was demonstrated at pH 4-12 and temperatures below 60°C. Surface plasmon resonance analysis of OXYL against fetuin showed k(ass) and k(diss) values of 1.4×10(-6)M(-1)s(-1) and 3.1×10(-3)s(-1), respectively, indicating that it has a strong binding affinity to the glycoprotein as lectin. Frontal affinity chromatography using 25 types of prydylamine-conjugated glycans indicated that OXYL specifically recognizes multi-antennary complex-type oligosaccharides containing type-2 N-acetyllactosamines (Galβ1-4GlcNAc) if α2-3-linked sialic acid is linked at the non-reducing terminal. However, type-1 N-acetyllactosamine (Galβ1-3GlcNAc) chains and α2-6-linked sialic acids were never recognized by OXYL. This profiling study showed that OXYL essentially recognizes β1-4-linkage at C-1 position and free OH group at C-6 position of Gal in addition to the conservation of N-acetyl groups at C-2 position and free OH groups at C-3 position of GlcNAc in N-acetyllactosamine. This is the first report on glycomics on a lectin purified from an echinoderm belonging to the subphylum Pelmatozoa.     

Doppellagige Kronenplatten: Ein neues anatomisches Merkmal paläozoischer Crinoiden und Revision der Familie Cupressocrinitidae (Devon)

Kurzfassung  Anhand von Material aus den mitteldevonischen Kalkmulden der Eifel (Rheinisches Schiefergebirge) wird eine auf Kelch- und Armsegmenten der Cupressocrinitidae C.F.Roemer, 1854 aufliegende und deutlich getrennte Skelettschicht als Exoplacoidschicht beschrieben. Aufbau, Morphologie, Ultrastruktur und geochemisches Signal werden vorgestellt, die Funktionsmorphologie diskutiert. Die Ausbildung der Exoplacoidschicht führt innerhalb der bisher monotypischen Familie Cupressocrinitidae C.F.Roemer, 1854 zur Abtrennung der neuen GattungenRobustocriniles n. gen. undAbbreviatocrinites n. gen. Die GattungRobustocrinhes n. gen. undCupressocrinites Goldfuss, 1831 besitzen eine monolamellare Exoplacoidschicht und unterscheiden sich durch weitere anatomische Merkmale von der durch eine multilamellare Exoplacoidschicht charakterisierten GattungAbbreviatocrinites n. gen. Dieser Gattung wird neben anderen, bisherCupressocrinites untergeordneten ArtenAbbreviatocrinites geminatus n. gen. n. sp. mit besonders deutlicher multilamellarer Exoplacoidschicht zugeordnet.      

NEW CRINOIDS (ECHINODERMATA) FROM THE LLANDOVERY (LOWER SILURIAN) OF THE BRITISH ISLES

Abstract:  British Llandovery crinoids remain poorly known. Three species are documented herein that were originally described, but not published, by W. H. C. Ramsbottom. Clematocrinus ramsbottomi Fearnhead sp. nov. (Tortworth Inlier, Gloucestershire; Telychian) has a heteromorphic column, N434243414342434, radices directed away from the crown, and ten long, uniserial arms with a pustular aboral sculpture and long, slender pinnules. Clematocrinus spp. are widely distributed in the Silurian of England. Ptychocrinus mullochillensis Fearnhead and Donovan sp. nov. (Girvan district, Strathclyde; Rhuddanian) is the second report of this Upper Ordovician–Lower Silurian genus from outside North America. Although incompletely known, this species is distinguished by its hidden infrabasals, 20 arms and sunken interbrachial plates with a strongly stellate sculpture. Petalocrinus bifidus (Bather MS) Donovan and Fearnhead sp. nov. (Woolhope Inlier, Herefordshire; Telychian) is locally common enough to give its name to a mappable lithostratigraphic unit, the Petalocrinus Limestone. Unlike other Petalocrinus species known from the Silurian of China, northern Europe and North America, the fused arms of P. bifidus are divided in two by a proximal adoral ridge and an associated distal notch. Unlike extant crinoids, Petalocrinus probably lived as a rheophobe; the fused arms may have acted to deter predators.     

Aspects of crinoid palaeontology of the North Esk Inlier,Scotland (Silurian,Llandovery, Telychian)

Abstract: The described fauna of well‐preserved Llandovery (Telychian) echinoderms from the North Esk Inlier, including six crinoids, one echinoid and seven starfish species, is mainly allochthonous. Most of these taxa are known only from starfish beds, channel fill deposits probably representing submarine mass flows and preserving a biota probably derived from elsewhere, presumably shallower water. Only one crinoid species, Pisocrinus cf. campana Miller, is recognized as a common fossil away from the starfish beds and is a biostratigraphic marker for the base of the Wether Law Linn Formation, forming part of the Skenidioides‐Cyrtia Association. Crinoid columnals preserved perpendicular to bedding (that is, in putative life position) in Lamont’s bivalve bed, Deerhope Formation, are tentatively interpreted as being in situ by comparison with a similar occurrence in the Silurian of Arisaig, Nova Scotia. Two new species of crinoid are described, the cladid Dendrocrinus? sp. and the columnal morphospecies Pentagonocyclicus (col.) lamonti sp. nov.     

A new holocrinid (Articulata) from the Paris Biota (Bear Lake County,Idaho, USA) highlights the high diversity of Early Triassic crinoids

After the end-Permian crisis and the extinction of their four Paleozoic subclasses, crinoids rapidly recovered. This group is classically believed to have radiated from a small surviving clade and to have diversified during the Middle and Upper Triassic from two lineages. Nevertheless, recent findings suggested that several lineages of crinoids had already diversified during the Early Triassic, and that their diversity has been overlooked. Here we describe a new form of holocrinid, Holocrinus nov. sp., from the earliest Spathian (Early Triassic) of southeastern Idaho (USA). So far, the exceptional completeness of sampled specimens, with skeletal elements of arms and stem in connection, is unique for the Early Triassic. They show that derived morphological features had already evolved ∼1.3 million years after the Permian–Triassic boundary, supporting the scenario of a rapid Early Triassic diversification of crinoids.     

The oldest illustration of Chinese crinoid calyx: Annotated translation of “Cupressocrinus abbreviatus” reported by Fritz Frech in 1911

The early history on the discovery of the first Chinese crinoid calyx of “Cupressocrinus abbreviatus” (now Halocrinites schlotheimii Steininger, 1831) remains unclear. The first record of “C. abbreviatus” was reported by Fritz Frech in 1911. It was included in the fifth and the last volume of the series titled China by the German traveler and explorer Ferdinand Freiherr von Richthofen. The cover page, title page, and prologue are informative; thus, they are explained and translated in detail. Richthofen's specimen is still the oldest documented cupressocrinitid calyx known in China today. Discovery of a second cupressocrinitid specimen confirms the existence of this family in China. Current taxonomic positions of the Chinese specimens are discussed.     

Testing reduced evolutionary rates during the Late Palaeozoic Ice Age using the crinoid fossil record

In 2003, Stanley & Powell reported depressed rates of origination and extinction in marine invertebrates during the Late Palaeozoic Ice Age (LPIA). Using a database of crinoid genera, rates of origination, extinction and genus duration were calculated at the stage level from the Early Devonian to the Late Permian. This 165 m.y. time span includes non‐glacial intervals before and after the LPIA, which spanned the Serpukhovian to Sakmarian, providing background rates for comparison. Data generated on crinoid evolutionary rates during the Middle to Late Palaeozoic were analysed and compared to Stanley & Powell's data to determine whether crinoid evolutionary patterns support their findings or suggest an alternative hypothesis. Rates of origination and extinction in all crinoid clades were reduced during the LPIA compared to the combined background intervals before and after the LPIA. However, crinoid diversity was higher during the LPIA than the surrounding time intervals. The difference in diversity trends between crinoids and other marine invertebrates is due to the advanced cladids clade. Unstable, fluctuating environmental conditions during the LPIA may have created habitats suitable for opportunistic crinoid genera that reduced both the probability of origination and extinction. The increased diversity of the advanced cladids is likely due to their unique adaptation of muscular arm articulations, which allowed them to thrive in marine settings with increased siliciclastic influx brought on by the Alleghenian orogeny. Despite the advanced cladids’ departure from the expected diversity count, the results of analyses performed on the updated crinoid database provide independent confirmation of Stanley & Powell's original hypothesis of depressed evolutionary rates in marine invertebrates during the LPIA.     

Roveacrinids (Crinoidea,Roveacrinida) from the Cenomanian-Turonian of southwest Algeria (Saharan Atlas and Guir Basin)

In the southwestern part of Algeria, the Cenomanian-Turonian marine deposits build up a prominent ledge in a perched syncline (Ksour Mountains, western Saharan Atlas) or at high radius of curvature (Guir Basin). The petrographical analysis of the Cenomanian-Turonian deposits of the Ksour Mountains and of the Guir Basin reveals unexpected assemblages of roveacrinoidal ossicles comparable with those formerly reported from the Tinrhert area. For the first time, isolated ossicles of genuine and undisputable Roveacrinidae are illustrated. Three sections, Djebel Rhoundjaia (western Saharan Atlas), Berridel and Kénadsa (Guir Basin), were scrutinized to recognize the microcrinoidal sections within the carbonate microfacies and to compile the successive occurrence of respective roveacrinid taxa (besides the classical search for standard index microfossils) in an attempt to pinpoint more precisely the position of the Cenomanian-Turonian boundary (C/T B). These assemblages are particularly morphologically and taxonomically diverse with three species of genus Roveacrinus and one of genus Orthogonocrinus. The presence of Saccocomidae (Applinocrinus) is especially unusual in such stratigraphic levels. The relative abundance and diversity of Roveacrinidae evidence a peak when approaching the C/T B. Such an event is recurring in the latest Cenomanian in various Tethyan and Atlantic areas. These fluctuations are consistent with a high surface-water productivity just before the C/T B.