Cryptostomid Bryozoans From The Sassito Formation,Upper Ordovician Cool‐Water Carbonates of the Argentinean Precordillera

Abstract: Two new bryozoan species are described from the Upper Ordovician Sassito Formation of the Argentinean Precordillera: Moyerella  spinata sp. nov. and Phylloporina  sassitoensis sp. nov. The bryozoans are found in cool‐water carbonates. The Silurian genus Moyerella is reported the first time in the Ordovician, showing palaeobiogeographic connections with Estonia and Siberia.     

STARFISH DIVERSITY IN THE WENLOCK OF ENGLAND

Abstract: Although their record extends back to the Early Ordovician, the occurrence of fossil starfish (Echinodermata: Asteroidea) is dependent almost exclusively upon horizons of exceptional preservation. Thus, asteroids found in Silurian obrution deposits of the English Midlands and Welsh Borderlands are particularly significant to an understanding of the early diversity of the group. Six species are described here: Hudsonaster? carectum sp. nov. (Hudsonasteridae), from the lower part of the Lower Elton Formation; and, from the Much Wenlock Limestone Formation, the hudsonasterids Doliaster brachyactis gen. et sp. nov. and Siluraster? ketleyi (Spencer, 1916), the lepidasterids Lepidaster grayi Forbes, 1850 and Lepidactis wenlocki Spencer, 1918, and the palasterinid Palasterina orchilocalia sp. nov. Though few in number, they show a diverse range of body morphologies when compared with Ordovician taxa: L. wenlocki had long, slender rays when fully grown whereas D. brachyactis is the first asteroid with the short‐rayed body form of extant cushion stars. Most distinctive of all is L. grayi, the earliest multiradiate taxon known, all complete specimens of which have 13 rays. This morphological variety is interpreted as indicating that by the Early Silurian starfish were exploiting a wide range of feeding habits and ecological niches.     

New crinoids from the Baltic region (Estonia): fossil tip‐dating phylogenetics constrains the origin and Ordovician–Silurian diversification of the Flexibilia (Echinodermata)

This study documents previously unknown taxonomic and morphological diversity among early Palaeozoic crinoids. Based on highly complete, well preserved crown material, we describe two new genera from the Ordovician and Silurian of the Baltic region (Estonia) that provide insight into two major features of the geological history of crinoids: the early evolution of the flexible clade during the Great Ordovician Biodiversification Event (GOBE), and their diversification history surrounding the end‐Ordovician mass extinction. The unexpected occurrence of a highly derived sagenocrinid, Tintinnabulicrinus estoniensis gen. et. sp. nov., from Upper Ordovician (lower Katian) rocks of the Baltic palaeocontinent provides high‐resolution temporal, taxonomic and palaeobiogeographical constraints on the origin and early evolution of the Flexibilia. The Silurian (lower Rhuddanian, Llandovery) Paerticrinus arvosus gen. et sp. nov. is the oldest known Silurian crinoid from Baltica and thus provides the earliest Baltic record of crinoids following the aftermath of the end‐Ordovician mass extinction. A Bayesian ‘fossil tip‐dating’ analysis implementing the fossilized birth–death process and a relaxed morphological clock model suggests that flexibles evolved c. 3 million years prior to their oldest fossil record, potentially involving an ancestor–descendant relationship (via ‘budding’ cladogenesis or anagenesis) with the paraphyletic cladid Cupulocrinus. The sagenocrinid subclade rapidly diverged from ‘taxocrinid’ grade crinoids during the final stages of the GOBE, culminating in maximal diversity among Ordovician crinoid faunas on a global scale. Remarkably, diversification patterns indicate little taxonomic turnover among flexibles across the Late Ordovician mass extinction. However, the elimination of closely related clades may have helped pave the way for their subsequent Silurian diversification and increased ecological role in post‐Ordovician Palaeozoic marine communities. This study highlights the significance of studies reporting faunas from undersampled palaeogeographical regions for clade‐based phylogenetic studies and improving estimates of global biodiversity through geological time.     

Late Ordovician brachiopods from Peru and their palaeobiogeographical relationships

A Sandbian brachiopod association from the Calapuja Formation, in the Peruvian Altiplano, north‐west of Lake Titicaca, has allowed a re‐examination of the palaeobiogeographical relationships between Gondwana and Avalonia during the Late Ordovician, when the palaeocontinents are considered to be already very distant from one another. The brachiopod fauna includes the new species Onnizetina calapujensis sp. nov., Horderleyella chacaltanai sp. nov., Drabovinella minuscula sp. nov. and Tasmanella curtiseptata sp. nov., as well as Caeroplecia sp., Dinorthis cf. flabellulum and Tunariorthis cardocanalis. In addition, Colaptomena expansa expansa and Heterorthis retrorsistria, known from the British Burrellian Stage of the Caradoc Series (late Sandbian) in Wales and the Welsh Borderlands, have also been identified. The brachiopod collection is the most diverse known from a single locality in the whole Central Andean Basin. Within it, forms with clear Gondwanan links occur, such as the new species of Onnizetina, Drabovinella and Horderleyella, and typical representatives of the Avalonian faunas, such as the Welsh Colaptomena expansa expansa and Heterorthis retrorsistria. The brachiopod species exchange between the Proto‐Andean margin of Gondwana and Avalonia, now believed to be possible during the late Sandbian, allows a reconsideration of the global taxonomic affinities of both regions. With this in mind, detrended correspondence analysis (DCA) and cluster analysis have been applied to an updated rhynchonelliformean brachiopod matrix consisting of presence/absence data. The scatter plot resulting from the DCA allows a vivid visualization of the grouping and geographical trends of the South American localities with respect to Avalonia–Baltica and the Mediterranean margin of Gondwana during the Sandbian. Our results agree with previous palaeogeographical reconstructions, depicting Avalonia very close to Baltica and already distant from Gondwana. As a few brachiopod species, with low dispersal potential, would have been able to migrate between those distant palaeocontinents, the existence of intermediate islands in the Rheic Ocean, permitting the transit by island hopping of eurythermal species, must be considered.     

Evolution,palaeoecology, and palaeobiogeography of the Late Ordovician–Early Silurian brachiopod Epitomyonia

Epitomyonia is characterized by various types of dorsal ridges, which may be transverse, longitudinal, or highly convoluted and probably served as skeletal supports for lophophores of various complexity. Multivariate analyses suggest that the Epitomyonia-bearing brachiopod associations lived in relatively shallow-water environment in the Late Ordovician, and inhabited mainly deep-water environments in the early Wenlock. The temporal and spatial change in the faunal distribution may be explained by three alternative scenarios: (1) Epitomyonia followed the broad evolutionary trend of the Palaeozoic Evolutionary Fauna to shift from shallow- to deeper-water settings over time; (2) the dicoelosiid communities could not compete with the large-shelled pentameride communities in continental shelf settings during the Early Silurian; or (3) only the shallow-water Epitomyonia died out in the Late Ordovician mass extinction event, whereas some poorly known deep-water Late Ordovician forms survived into the Early Silurian. Epitomyonia paucitropida n. sp. from the lower Whittaker Formation (late Katian) of the Mackenzie Mountains, northwestern Canada, is reported as the first known Ordovician species of Epitomyonia from the palaeocontinent of Laurentia, characterized by a small shell with weak, transverse dorsal ridges that are most primitive for the genus.     

New brachiopod species from the order acrotretida from the Cambrian of central Kazakhstan

Four new brachiopod species from the order Acrotretida (class Lingulata): Picnotreta saryarkensis sp. nov., Stilpnotreta propria sp. nov., Anabolotreta firma sp. nov., and Batenevotreta variabilis sp. nov. are described from the Agyrek Mountains, northeastern Central Kazakhstan from two olistoliths of limestones, which contain the uppermost Middle and lowermost Upper Cambrian fauna and are located in the Upper Ordovician olistostrome. These new species supplement the described earlier brachiopod assemblage from this region (Koneva and Ushatinskaya, 2008). Strong age and intraspecific variability of some Kazakh acrotretids is shown.     

Middle Ordovician Aporthophyla brachiopod fauna from the roof of the World,southern Tibet

A Darriwilian (late Middle Ordovician) brachiopod fauna from the Lower Formation of the Chiatsun Group at Jiacun, northern Nyalam, southern Tibet, consists of ten brachiopod species, forming a distinct Aporthophyla–Paralenorthis Association. Its taxonomic composition is typical of the Aporthophyla Fauna that occupied lower BA2 to upper BA3 benthic environments on sandy lime mud substrates. The occurrence of Paralenorthis in southern Tibet is confirmed for the first time, represented by P. costata sp. nov. Numerical analyses (PCA and CA) of 18 Darriwilian brachiopod faunas from ten palaeoplates or terranes indicate that: (1) the Aporthophyla Fauna was confined to a specific latitudinal belt although it had a wide lateral distribution from the large palaeocontinents of Gondwana to Laurentia; (2) the Saucrorthis Fauna, a typical late Middle Ordovician regional fauna, is limited to a much smaller area, marginal to the Gondwana supercontinent; (3) the strong provincialism persistent in the late Middle Ordovician contributed to increased gamma biodiversity during the Great Ordovician Biodiversification Event.     

First report of craniide brachiopods in the Palaeozoic of Iran (Pseudocrania,Ordovician), and Early to Mid‐Ordovician biogeography of the Craniida

Abstract: Lower Ordovician faunas of Bohemia (Perunica), Baltica and North China include the oldest known representatives of the Order Craniida, but otherwise in Gondwana and associated terranes, the record of craniides is sparse. Pseudocrania insperata sp. nov. from the Lashkarak Formation of the Eastern Alborz Mountains is the first and as yet only record of the occurrence of craniides in the Middle Ordovician (Darriwilian) of Iran and temperate to high latitude peri‐Gondwana. Pseudocrania was known hitherto only from the Middle Ordovician of Baltoscandia and the Chu‐Ili terrane of Kazakhstan.     

Ontogenic study of the brachiopod Dicoelosia by geometric morphometrics and morphing techniques

The distinctive brachiopod Dicoelosia King 1850 is characterized by a strongly bilobed outline. To date, studies have concentrated on its functional morphology, taxonomy and evolution; little attention has been paid to its ontogeny. Here, we map population variation by principal component analysis for over 80 specimens distributed across five species of Dicoelosia. Using geometric morphometrics with landmarks for some 40 specimens, the ontogenic trends in D. sp. nov. are compared with those of Dicoelosia biloba. In addition, the ontogenic pathway in D. sp. nov. is investigated by morphing with control points, a new technique introduced here to palaeontology. Combining the results above, the ontogeny of the key character of the genus, emargination, is modelled. Within single populations, taxa may develop from broad weakly emarginate forms into those that are elongate and deeply emarginate. As the identification of the genus and its species depends on external morphological characters, the definition of ontogenetic trends in each species is essential for taxonomic discrimination. Substantial population variation exists in many of its species; however, the morphing technique provides a method of simulation, predicting the full range of ontogenetic variation in given populations.     

Latest Ordovician brachiopod and trilobite assemblage from Yuhang,northern Zhejiang,East China: a window on Hirnantian deep-water benthos

A moderately diverse brachiopod and trilobite assemblage, the Leangella–Dalmanitina (Songxites) Assemblage, occurs in the upper Yankou Formation (Hirnantian, probably equivalent to the Normalograptus persculptus Biozone) at Shizi Hill, Yuhang, west of Hangzhou, northern Zhejiang, E China. The brachiopods are rare, characterised by minute, thin shells with very small body cavities, preserved in mudstones as moulds. They may have inhabited quiet, deep-water and dysaerobic slope environments with low levels of nutrients, equivalent to Benthic Assemblage 5. Most genera were adapted for life in deep water and either remained there or alternatively migrated into relatively shallower habitats to evade perturbations during the first phase of the end Ordovician extinctions. The slope environments were recolonised from outer shelf and upper slope communities during the early Hirnantian, but isolated biotas may also have survived in deeper-water habitats by reducing their population size and diversity during the crisis. The Leangella–Dalmanitina (Songxites) Assemblage provides an unique Hirnantian window through which we can monitor the changes in the deep-water biofacies following the first phase of the extinctions. Significantly, parts of the deep water marine environment may have survived intact, the end Ordovician extinctions.     

SILICIFIED RHYNCHONELLIFORM BRACHIOPODS FROM THE KUNIUTAN FORMATION (DARRIWILIAN: MIDDLE ORDOVICIAN), GUIYANG, SOUTH CHINA

Abstract: A silicified brachiopod fauna from the Middle Ordovician Kuniutan Formation (lower–middle Darriwilian, i.e. uppermost Arenig to lower Llanvirn) at Wudang, Guiyang, central Guizhou, South China, includes abundant specimens of Yangtzeella, Orthambonites and Leptellina together with common Parisorthis, Saucrorthis, rare Anomalorthis?, Hemipronites?, Leptestia? and, significantly, Aporthophyla; associated are rare trilobites, gastropods, crinoids and nautiloids. The Yangtzeella kueiyangensis‐Orthambonites delicata Association is defined for this shallow‐water, Benthic Assemblage 3, association. This first record of Aporthophyla in South China may indicate a link between South China and the Toquima‐Table Head Province, where the Aporthophyla fauna is more typically developed. However, this association is significantly different from the latter in having three endemic genera, Yangtzeella, Saucrorthis and Parisorthis, rare numbers of Aporthophyla and Anomalorthis?, and the absence of many other typical elements of the Aporthophyla fauna. The occurrence of Orthambonites, Hemipronites? and Leptestia? suggests some relationships between South China and the Baltic Platform during the Mid Ordovician. The various brachiopod associations bearing Aporthophyla may be quite different in nature, composition and diversity, and may possess different background palaeobiogeographical signatures. The assemblages containing Aporthophyla in South China, Qaidam, Malaysia, Australia and possibly Tibet are clearly different biogeographically from those associated with the Toquima‐Table Head and the Celtic provinces. Two new species, Aporthophyla sinensis sp. nov. and Leptellina orientalis sp. nov. are described.     

The biological affinity of Amsassia: new evidence from the Ordovician of North China

Amsassia shaanxiensis sp. nov. occurs in the Middle Ordovician part of the Jinghe Formation in Yongshou and the lower part of the Upper Ordovician Beiguoshan Formation in Longxian, Shaanxi Province, north‐central China. In addition to module increase by bipartite longitudinal fission, which is also known in other species of Amsassia, tripartite and rare quadripartite fission are recognized in A. shaanxiensis. All species previously assigned to Lichenaria from the Middle to Upper Ordovician of Shaanxi probably belong to Amsassia. Therefore, Amsassia, rather than the tabulate coral Lichenaria, should be credited as an important contributor to reef‐building in this area. Reports of Lichenaria from elsewhere in the North China Platform require confirmation in the light of the present study. Some morphological characteristics of Amsassia are comparable to those of tabulate corals, tetradiids and chaetetid sponges. Consequently, various authors have assigned Amsassia to the Lichenariida, Tetradiida (now Prismostylales; florideophycean rhodophyte algae) and Chaetetida. Other important characters, however, seem to exclude Amsassia from those taxonomic groups. The phacelocerioid organization of modules having separate walls would not be expected in sponges. The basic symmetry of individuals may have been radial, unlike the tetramerous symmetry of tetradiids. Module increase by longitudinal fission, involving infoldings of the wall, is fundamentally different from modes of increase in corals, tetradiids and chaetetids. The skeleton was probably aragonitic, whereas that of tabulates was calcitic. The affinity of Amsassia remains unresolved, but it is unlikely to have been a coral, tetradiid or sponge. Perhaps, like the tetradiids, Amsassia was an alga.     

Early Silurian (Telychian) rugose coral fauna of Daguan area, northeast Yunnan province, China

Study on rugose coral fauna of the Sifengya Formation (early Telychian) and Daluzhai Formation (mid-late Telychian) in Daguan area, northeast Yunnan Province, China was carried out. Rugose coral fauna of the Sifengya Formation included 18 genera and 34 species, while Daluzhai Formation with nine genera, ten species. We described rugose coral fauna (12 genera, 19 species) including one new genus and five new species, i.e. Protoketophyllum daguanense gen. et sp. nov., Crassilasma huanggexiense sp. nov., Pseudophaulactis heae sp. nov., P. convolutus sp. nov., and Shensiphyllum minor sp. nov. The characteristics and geological significance of rugose coral fauna of Sifengya Formation and Daluzhai Formation were analyzed. Particularly, rugose coral fauna of the Sifengya Formation represent early Telychian rugosan fauna in the Upper Yangtze region and improve the sequences of early Silurian (Llandovery) rugose coral assemblages in Yangtze region. It is therefore very meaningful to further analyze radiation period of rugose coral fauna in such epoch. __________ Translated from Acta Palaeontologica Sinica, 2005, 44(2): 229–246 [译自: 古生物学报, 2005, 44(2): 229–246]     

UPPER ORDOVICIAN BRYOZOANS OF THE PIN FORMATION (SPITI VALLEY, NORTHERN INDIA)

Abstract: Twenty‐nine species of bryozoans from the Upper Ordovician–Lower Silurian Pin Formation (Spiti, India) have been identified. Eight of these are new: Trematopora minima, Ulrichostylus bhargavai, Ptilodictya exiliformis, Phaenopora ordinarius, Oanduellina himalayaica, Pesnastylus? vesiculosum, Ralfina? originalis and Pinocladia triangulata. The fossil record and facies analyses of the area investigated indicate shallow‐water conditions within the subtropical–tropical realm. The distribution pattern of fossils among the Ordovician/Silurian succession on the Northern Gondwana shelf and the influence of the Late Ordovician cooling phases on marine organisms are distinctive owing to a dramatic reduction in diversity globally. As far as the bryozoan taxa of Spiti are concerned, only one (Helopora fragilis) of the 29 species was recorded above the Ordovician/Silurian boundary. Observed bryozoan communities are very similar to faunas of Laurentia, the Baltic, Siberia and southern China of early–late Ordovician age.     

The earliest atrypides and athyridides (Brachiopoda) from the Ordovician of Kazakhstan

The middle Ordovician brachiopod faunas of Kazakhstan provide one of the most complete records of the evolution and radiation of some of the oldest known spire-bearing brachiopods. By contrast with North American faunas, Kazakhstanian atrypide taxa mostly belong to the suborders Atrypidina and Lissatrypidina, whereas the suborder Anazygidina is completely absent. Kazakhstanian species referred previously to ZygospiraKuzgunia are reassigned to Sulcatospira, which appeared in the Caradoc Diplograptus multidensClimacograptus clingani biozones (Sulcatospira? praecursor and Sulcatospira prima sp. nov.). Primitive, and possibly the oldest known Athyridida also appeared in Kazakhstan sometime during the Caradoc (Kellerella misiusi sp. nov.) and became widespread in brachiopod assemblages developed in carbonate mud mounds. Phylogenetic analysis suggests the early divergence of the Anazygidina, Atrypidina and Athyridida, which probably evolved independently from various primitive smooth Lissatrypidina. The new atrypide subfamily Pectenospirinae and two new atrypide genera (Rozmanospira gen. nov. and Pectenospira gen. nov. with P. pectenata sp. nov. as type species) are erected.     

Probing the deep shelf: a Lagerstätte from the Upper Ordovician of Girvan, southwestern Scotland

A deep‐water Konservat Lagerstätte from the lower Caradoc (Sandbian) at Girvan is dominated by the trilobite Diacanthaspis trippi, the carpoids Anatifopsis n. sp.? and a new genus of ctenocystoid together with the polyplacophoran Solenocaris solenoides and the brachiopod Onniella williamsi. Most of these are multi‐element organisms, with many specimens preserved in an articulated state in finely laminated rocks, indicating minimal disturbance and suggesting that the fauna is largely an in situ association. It contains few of the species known from other deep‐water sites of similar age at Girvan which contain diverse assemblages of trilobites and brachiopods absent from the Lagerstätte. The taphonomy of the site indicates preservation by rapid burial followed by early diagenesis under dysaerobic conditions. It provides a ‘taphonomic window’ on otherwise unknown faunas from distal shelf facies on the Ordovician Laurentian margin, and, moreover, is an important reminder of the hidden biodiversity that resided in thin‐shelled, multi‐element organisms.     

Systematics, shell structure and affinities of the Palaeozoic Problematicum Cornulites

The genus Cornulites, with the type species C. serpularius Schlotheim, 1820, from the Silurian of Gotland, comprises annulated, conical or tubular calcite shells, often found attached to the hard parts of other organisms. No consensus has ever been reached over the zoological affinities of the taxon, and no examples of soft‐part preservation are known: detailed examination of shell structures and growth patterns provide the only means of assessing its systematic position. Using transverse and longitudinal thin sections of C. serpularius Vine, 1882, and C. cellulosus sp. nov. , from the Much Wenlock Limestone Formation of England, the shell structure of Cornulites is shown to be lamellar, but with conspicuous internal chambers (camerae) at the apical end of the shell and, particularly in C. cellulosus, numerous smaller vacuities (cellulae) between the lamellae in the apertural shell region. Growth of the shell was by the secretion of low‐magnesian calcite increments within one another, giving a cone‐in‐cone structure, with the prominent development of cellulae in C. cellulosus probably a constructional feature relating to an upright life position. By comparison of morphology and shell structure with other taxa, the zoological affinities of Cornulites are re‐examined; previously suggested affinities with annelids, foraminifers, molluscs and poriferans can be ruled out. Specific shell structures, most notably pseudopuncta similar to those of bryozoans and brachiopods, have led some recent workers to interpret cornulitids as lophophorates. However, it is shown that they can be interpreted alternatively as solitary, aseptate members of the stem‐Zoantharia (Cnidaria: Anthozoa). Four cornulitid species are recognized in the Much Wenlock Limestone Formation: C. cellulosus sp. nov. , C. gremialis sp. nov. , C. scalariformis and C. serpularius. In the absence of the type material, C. serpularius is here restricted to cornulitids closely resembling the specimens originally figured by Schlotheim. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 150 , 681–699.     

Silurian craniide brachiopods from Gotland

Seven craniide brachiopod genera are described from the Silurian (Wenlock–Ludlow) of Gotland, including one new genus and five new species. The new genus and species Thulecrania septicostata is unique among Silurian craniides as it possesses solid spines. The new species Lepidocrania multilamellosa is the first Silurian record of this poorly known Permian genus. The problematic North American Propatella Grubbs, 1939 , was originally described as a gastropod, but the new species Propatella palmaria from Gotland shows that it is a craniide with sutured hollow spines of a type not previously recorded from Silurian craniides. The dorsal valves of the new species Valdiviathyris? bicornis are remarkably similar to those of the type species and represent the first possible Palaeozoic record of this poorly known extant craniide. This first systematic study of craniide brachiopods from the Silurian of Gotland shows that the diversity is relatively high as compared to other known Silurian craniide faunas, but a more thorough comparison is not possible due to the lack of data from most parts of the world. The new data from Gotland support the view that the craniides were not affected by the end‐Ordovician extinction.     

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.