Chinese origin and radiation of the Palaeozoic crinoid family Petalocrinidae

The Ordovician to Devonian family Petalocrinidae includes 28 species belonging to 5 genera. This family is unique because the arm plates are fused into a large fan or cylinder. Paleobiogeographic occurrences of this family include Laurentia, Baltica, Avalonia, South China, Sibumasu, and Perunica blocks. The family has the oldest petalocrinids from China. An early radiation of this clade resulted in three genera during the Llandovery of China. Petalocrinus became cosmopolitan during the Llandovery and Wenlock, and the youngest genus is present in the Lower Devonian of the Czech Republic. Taxonomic determination for the Petalocrinidae is based on the fused arm plates instead of cup plates. The diverse morphology of these arm plates suggests a variety of aerosol suspension-feeding modes may have been used by petalocrinids.     

Did the Late Ordovician mass extinction event trigger the earliest evolution of ‘strophodontoid’ brachiopods?

‘Strophodontoid’ brachiopods represented the majority of strophomenide brachiopods in the Silurian and Devonian periods. They are characterized by denticles developed along the hinge line. The evolution of denticles correlated with the disappearance of dental plates and teeth and were already present when the clade originated in the Late Ordovician. Specimens of Eostropheodonta parvicostellata from the Kuanyinchiao Bed (early–middle Hirnantian, uppermost Ordovician) in the Hetaoba Section, Meitan, Guizhou Province, South China, display clear fossil population variation, during a process of loss of dental plates and the development of denticles. Three phenotypes of E. parvicostellata are recognized in a single fossil bed, likely heralding a speciation process. Non-metric multidimensional scaling (NMDS) based on five key characters of genera of the Family Leptostrophiidae shows a much wider morphospace for Silurian genera than for those in the Devonian. Phylogenetic analysis of the Family Leptostrophiidae supports the NMDS analysis and mostly tracks their geological history. The fossil population differentiation in E. parvicostellata discovered between the two phases of the Late Ordovician mass extinction event (LOME) linked to a major glaciation, suggests a Hirnantian origination of the ‘strophodontoid’ morphology, and links microevolutionary change to a macroevolutionary event.     

Family Lycophoriidae (Brachiopoda) from the Ordovician of Baltoscandia

The family Lycophoriidae from the Middle Ordovician of Baltoscandia with one monotypic genus and five species is revised. New inner structures were revealed and diagnoses of the family, genera, and species were specified in the study of cross sections and microstructure of the shell. The third prismatic layer, complex dental plates, and socket ridges with short pointed lateral outgrowths were recorded. The family Lycophoriidae cannot be assigned with confidence to any known brachiopod order.     

Aboral nervous system in two Ordovician crinoids: Reconstruction and comparison of Baltic <Emphasis Type="Italic">Pentamerocrinus</Emphasis> Jaekel and <Emphasis Type="Italic">Grammocrinus</Emphasis> Eichwald

Two unusual Middle Ordovician crinoid genera from the St. Petersburg Region, Pentamerocrinus Jaekel, 1918 and Grammocrinus Eichwald, 1860, are redescribed. A new species, Pentamerocrinus kosovi sp. nov., is described based on new material and the cup structure in the type species Grammocrinus lineatus, which has previously been known only from stem fragments, is reconstructed. The system of ridges and canals on the internal surface of the cup plates in both genera is described, allowing the reconstruction of the aboral nervous system. It is in general similar to extant crinoids, but the two Ordovician genera differ considerably in structural details. In Pentamerocrinus, ten nerve cords diverge interradially from the ganglionated nerve ring at the cup base and soon are fused in pairs inside the rays. In Grammocrinus, a similar nerve ring also gives rise to ten cords, but they are inside the rays and become fused in the interrays. This distinction in morphology of the nervous systems is correlated with the interradial arrangement of the axial canal angles of the stem in the first case and radial in the second. Therefore, despite general similarity in the structure and arrangement of calyceal plates, the genus Grammocrinus is assigned to the disparid crinoid family Iocrinidae, while the monotypic family Pentamerocrinidae Jaekel, 1918 is regarded as a fam. inc. sed. of inadunate crinoids.     

扬子区中、晚奥陶世和兰多维利世四射珊瑚起源、扩散及生物古地理关系

扬子区中奥陶世—志留纪兰多维利世四射珊瑚产出丰富 ,尤以兰多维利世最为繁盛。目前已报道的1 2 3属 (包括中奥陶世 4属 ,晚奥陶世 2 5属 ,兰多维利世 94属 )中 ,有 30属最早出现在扬子区 ,尔后扩散到欧洲、北美和其它邻区。例如 :Calostylis最早出现在川南的中奥陶世 (Llandeilo)地层中 ;Aphyllum和Cantrillia最早出现在浙西的晚奥陶世中Ashgill地层 ;扭心珊瑚类Briantelasma ,Pycnactis和Tunguselasma等 ,最早出现在黔东北晚Ruddanian地层 ;柱珊瑚类Ceriaster、Stauria、Amplexoides、Synamplexoides等 ,泡沫珊湖类Maikottia ,Rhizophyl lum等均最早出现在黔东北的中兰多维利统 (MiddleLlandovery)。根据这些资料 ,我们认为扬子区应该是奥陶纪和志留纪四射珊瑚起源中心之一。文中论述扬子区中奥陶世—兰多维利世四射珊瑚动物群特征及其生物古地理关系。该区中奥陶世珊瑚以穿孔珊瑚类的Calostylis和Yohophyllum为特征。下扬子区浙西晚奥陶世三衢山组 (中Ashgill)四射珊瑚群有某些澳大利亚分子Hillophyllum和Bowanophyllum ;而上扬子区晚Ashgill观音桥层的四射珊瑚动物群与北欧同期珊瑚群有高度的相似性。这表明扬子区当时与欧洲具有较密切的古生物地理关系。扬子区兰多维利世四射珊瑚群与西伯利亚。     

新疆塔里木柯坪地区上奥陶统凯迪阶串管海绵类化石及其滤食选择性探讨北大核心

奥陶纪是串管海绵早期演化的一个关键阶段,记录了该类群的首次大规模辐射演化事件(属级分类单元达15个)。然而,现有大部分奥陶系串管海绵的报道来自澳大利亚东部和北美等地,在中国至今仅发现了两个属,与前述地区相比研究程度偏低。早期研究主要基于形态学特征对串管海绵进行系统分类描述,对其滤食策略知之甚少,因此制约了对此类海绵在奥陶纪底栖生态系统作用的相关探讨。本文系新疆塔里木柯坪地区奥陶纪串管海绵的首次系统古生物学报道,详细描述凯迪阶(上奥陶统)印干组中保存完好的阔室丛花海绵(Corymbospongia amplia)化石的形态学特征,并通过研究标本外壁微孔孔径的大小分布,探讨其滤食选择性。海绵外壁微孔孔径的分析表明:小型和微型浮游生物可能是其食物的主要来源,而多细胞浮游动物因个体尺寸大于海绵外壁微孔,不能被阔室丛花海绵捕获。本研究为奥陶纪串管海绵古生态学及其地球生物学意义的探讨提供了进一步的依据。     

The structure and evolution of Ordovician conodont apparatuses

Multielement taxonomy was instituted for Ordovician conodonts over a decade ago, and probably a majority of the multielement genera have been defined or are well understood. The present systems of notation for elements within apparatuses are inadequate and cumbersome. A new notation scheme is proposed which applies a single-letter code to the position in the apparatus occupied by certain element morphotypes. The taxonomic status of all known Ordovician conodont genera is reviewed (appendix) using the new notation, and a new scheme to classify conodont apparatuses is presented. Five main apparatus types (I-V) and seventeen subtypes (IA-IC, etc.) are defined. Within these groups, all known Ordovician conodont genera can be accommodated, and probably few new groups are required to include all other conodont genera. The apparatus types and subtypes are defined on the basis of symmetry, curvature, and number of the element types, with a clear distinction being made between the first and second transition series. Certain homologous relationships, both between and within many apparatus types, are noted. The evolution of the five major types, and the subtypes, is traced through the Ordovician. The pattern of evolution suggests that the types and subtypes recognized are probably natural biologic groupings, largely reflecting phylogenetic change.     

A new look at age and area: the geographic and environmental expansion of genera during the Ordovician Radiation

Although available paleobiological data indicate that the geographic ranges of marine species are maintained throughout their entire observable durations, other evidence suggests, by contrast, that the ranges of higher taxa expand as they age, perhaps in association with increased species richness. Here, I utilize a database of Ordovician genus occurrences collected from the literature for several paleocontinents to demonstrate that a significant aging of the global biota during the Ordovician Radiation was accompanied by a geographic and environmental expansion of genus ranges. The proportion of genera occurring in two or more paleocontinents in the database, and two or more environmental zones within a six-zone onshore-offshore framework, increased significantly in the Caradocian and Ashgillian. Moreover, widespread genera tended to be significantly older than their endemic counterparts, suggesting a direct link between their ages and their environmental and geographic extents. Expansion in association with aging was corroborated further by demonstrating this pattern directly among genera that ranged from the Tremadocian through the Ashgillian. Taken together, these results are significant not only for what they reveal about the kinetics of a major, global-scale diversification, but also for what they suggest about the interpretation of relationships between diversity trends at the alpha (within-community) and beta (between-community) levels.     

Diverse middle Ordovician palaeoscolecidan worms from the Builth‐Llandrindod Inlier of central Wales

Abstract: Palaeoscolecidan worms are rare, Early Palaeozoic fossils with uncertain affinities within the Ecdysozoa. They are locally abundant in the Cambrian and scattered in the Early Ordovician, but very sparse thereafter. Forty‐four specimens have been collected from the Middle Ordovician of the Builth‐Llandrindod Inlier of Mid Wales and include well‐preserved material assigned to seven new genera, with four additional species in open nomenclature. An additional specimen from the Arenig Pontyfenni Formation of South Wales is also described in open nomenclature. The total demonstrates much greater palaeoscolecid diversity than hitherto suspected for this time. The specimens are preserved as cuticle fragments in shales and siltstones, often of submillimetre size but in many cases with excellent preservation. The level of detail preserved in some is equal to that found in Cambrian phosphatized faunas. The new approach to collecting, and the recognition that this material can yield taxonomically useful information, opens new avenues for palaeoscolecidan research in siliciclastic environments. The new taxa are the following: Radnorscolex bwlchi gen. et sp. nov., Aggerscolex murchisoni gen. et sp. nov., Bullascolex inserere gen. et sp. nov., Wernia eximia gen. et sp. nov., Ulexiscolex ormrodi gen. et sp. nov., Pluoscolex linearis gen. et sp. nov. and Loriciscolex cuspidus gen. et sp. nov. The high diversity, and the taxonomic separation from known groups described primarily from Cambrian carbonates, implies that palaeoscolecidans either diversified significantly during the Ordovician or were taxonomically segregated between carbonate and siliciclastic settings. Palaeobiological findings also include confirmation that some palaeoscolecid basal cuticles were solid and others reticulate, plates (and platelets) could form by lateral accretion, plates were in part primarily phosphatic and in part organic and that in at least some groups, platelet secretion occurred external to plate secretion.     

中国新疆塔里木盆地上奥陶统良里塔格组的钙藻化石

钙藻是可以发生生物钙化作用形成钙质"骨骼"的分属多个门类的藻类的俗称。钙藻最早出现于寒武纪,奥陶纪发生第一次辐射演化。本文系统讨论了绿藻门绒枝藻目(Dasycladales)、羽藻目钙扇藻科(Udoteaceae)的科和属级分类标准和红藻门珊瑚藻科、管孔藻科(Solenoporaceae)属级分类标准及其中各属分类中存在的问题。塔里木盆地晚奥陶世在塔中—巴楚台地和塔北台地发育了一套礁滩相良里塔格组碳酸盐岩,其中含有丰富的钙藻化石。本文系统描述了来自塔里木盆地塔中、巴楚、塔北地区取心井上奥陶统良里塔格组岩层中的钙藻化石11个属15个种。包括绿藻门绒枝藻目西莱特藻科(Seletonellaceae)的Dasyporella,Kazakhstanelia,Vermiporella,Aphroporella,Arthroporella,绿藻门的羽藻目Bryopsidales(siphonales)的钙扇藻科的Dimor phosiphon,Palaeo porella,红藻门珊瑚藻目(Corallinales)的管孔藻科的Solenopora,Parachaetetes及Corallinales incertus familiae的Petro-phyton,以及分类位置不明的Monili porella。其中绒枝藻Ajakmalajsoria被视作Kazakhstanelia的同义名。     

Phylogenetic Clustering of Origination and Extinction across the Late Ordovician Mass Extinction

Mass extinctions can have dramatic effects on the trajectory of life, but in some cases the effects can be relatively small even when extinction rates are high. For example, the Late Ordovician mass extinction is the second most severe in terms of the proportion of genera eliminated, yet is noted for the lack of ecological consequences and shifts in clade dominance. By comparison, the end-Cretaceous mass extinction was less severe but eliminated several major clades while some rare surviving clades diversified in the Paleogene. This disconnect may be better understood by incorporating the phylogenetic relatedness of taxa into studies of mass extinctions, as the factors driving extinction and recovery are thought to be phylogenetically conserved and should therefore promote both origination and extinction of closely related taxa. Here, we test whether there was phylogenetic selectivity in extinction and origination using brachiopod genera from the Middle Ordovician through the Devonian. Using an index of taxonomic clustering (R_CL) as a proxy for phylogenetic clustering, we find that A) both extinctions and originations shift from taxonomically random or weakly clustered within families in the Ordovician to strongly clustered in the Silurian and Devonian, beginning with the recovery following the Late Ordovician mass extinction, and B) the Late Ordovician mass extinction was itself only weakly clustered. Both results stand in stark contrast to Cretaceous-Cenozoic bivalves, which showed significant levels of taxonomic clustering of extinctions in the Cretaceous, including strong clustering in the mass extinction, but taxonomically random extinctions in the Cenozoic. The contrasting patterns between the Late Ordovician and end-Cretaceous events suggest a complex relationship between the phylogenetic selectivity of mass extinctions and the long-term phylogenetic signal in origination and extinction patterns.     

Provinces, communities, and the proposed nektobenthic habit of Ordovician conodontophorids

Ordovician conodontophorid distribution, paleogeography, paleoclimate, and global tectonics strongly support the existence of one biogeographic conodontophorid province, largely restricted to equatorial regions and characterized by raised salinity and temperature. This Midcontinent Province is recognized in North America, Australia, and Siberia. The same data indicate that in other areas the North Atlantic Province represents a normal-marine, virtually cosmopolitan fauna. In both provinces, communities are recognized that show a lateral segregation, strongly indicating that the majority of Ordovician conodontophorids were benthic or nektobenthic. The only Ordovician pelagic forms we recognize are the simple cone genera that either lack, or have a simple, symmetry transition series. It appears possible to correlate certain evolutionary changes in the conodont apparatus with changes in preferred habitat.     

Bryozoan provinces and patterns of generic evolution and extinction in the Late Ordovician of North America

The Late Ordovician bryozoan genera of central and southeastern North America are geographically distributed in three biotic provinces, separated by boundaries reflecting major lithofacies differences. The central Cincinnati Province contains most of the North American endemic genera, and represents a narrow ecological zone separating the clastic wedges of the marginal Reedsville-Lorraine Province from the cratonic carbonate platform of the Red River-Stony Mountain Province. The provinces provided major life zones, or biomes, for each of the five bryozoan orders. Genera comprising the provinces differed as well in morphologic complexity, geochronologic survivorship, tiering, endemism and eurytopy. Regions on either side of the Cincinnati Province were dominated by inferred immigrants from Baltoscandia. Al-logenic provincial succession produced time-averaged mixed faunas in regions near the provincial boundaries. Although most generic originations took place within the Cincinnati Province, evolutionary novelties are associated with the Reedsville-Lorraine Province. The loss of the diverse Cincinnati Province, connected with global cooling and a eustatic lowering of sea level, may have been a chief factor in the Late Ordovician extinction of bryozoan genera. Genera from the Red River-Stony Mountain Province differentially survived into the Silurian.     

Kissinella-Christiania Associations in the early Ashgill Foliomena brachiopod fauna of South China

A new ecological unit within the well-documented late Ordovician brachiopod Foliomena fauna is defined on the basis of assemblages from Northern Cuizhou and southern Sichuan. Southwest China. The Kassinella-Chrisriania Association, from the lower Ashgill Linhsiang and Chientsaokou formations, is dominated by the eponymous genera together with species of mainly Dedzetina, Sericoidea and Kozlowskites Foliomena is rare or absent. To date, the Foliomena fauna has been documented only from deep-water environments however, the new association apparently occupied shallower depths, probably in lowermost Benthic Assemblage Zone 3 and uppermost Benthic Assemblage Zone 4, presumably at the shallow end of the depth range of the Foliomere fauna. The incursion of the Foliomena fauna into relatively shallow-water environments may have been encouraged by abnormally low oxygen levels and sparse nutrients together with persistent soft substrates across this part of the Yangtze Platform during the early Ashgill. *** Ordovician, Ashgill, China, brachiopods. F oliomena fauna.     

Revision of <Emphasis Type="Italic">Scoliocystis</Emphasis> (Rhombifera: Echinoencrinitidae) and related cystoid genera

Species attributed to Scoliocystis Jaekel, 1899, including the type species S. pumila (Eichwald, 1860) and S. thersites Jaekel, 1899 from the Ordovician of the Leningrad Region, are reviewed. Scoliocystis sp. from the Upper Ordovician of Estonia, figured by Hecker (1964), is redescribed as Maennilocystis heckeri gen. et sp. nov. and it is attributed to the family Callocystitidae Bernard, 1895. The genus Scoliocystis is intermediate between the families Cheirocrinidae and Echinoencrinitidae in having five periproct border plates (as in cheirocrinids), but lacking plate R5 and having a reduced oral area, as in echinoencrinitids, but is retained in the Echinoencrinitidae. The similarity of Scoliocystis to the unusual North American cheirocrinid genus Sprinkleocystis is convergent. The genus Gonocrinites Eichwald, 1840, with two species, is restored among echinoencrinitids, as having four periproct border plates, whereas Echinoencrinites sensu stricto has three. The families Echinoencrinitidae and Callocystitidae arose from cheirocrinids with closed plate circlets by the loss of plate R5 and substitution of radial:lateral for radial:radial pectinirhombs. In addition, echinoencrinitids are characterized by a reduced oral area, whereas in callocystitids, the recumbent ambulacra are extensive. Scoliocystis is a stem-group genus to Echinoencrinitidae plus Callocystitidae. Several glyptocystitoid genera independently developed advanced pectinirhombs.     

Early Palaeozoic diversification of chitons (Polyplacophora, Mollusca) based on new data from the Silurian of Gotland, Sweden

Isolated, well-preserved silicified sclerites of an unusually diverse Silurian paleoloricate assemblage from Gotland preserve morphological features that are important in interpreting palaeobiology. The typically granular dorsal ornament is comparable with Recent chitons, and is hence possibly linked functionally with sensory aesthetes. Ventral structures particularly in thickened shells indicate major muscle attachment sites sub-apically or, equivalently, beneath the rim of the apical area, and also marginally. In early chitons such as Cambrian Matthevia, deep ventral cavities represent comparable sub-apical sites. Three Gotland genera with an unusual, holoperipheral shell growth style apparently represent plated aplacophorans (cf. Acaenoplax), which coexisted with paleoloricate chitons in shallow inshore carbonate shelf environments. Sclerite features of all the Gotland genera are discussed together since they share most characteristics. The new family Heloplacidae includes those genera and Acaenoplax, which in a preliminary cladistic analysis form a sister group to other Lower Palaeozoic paleoloricates. Multiplated skeletons in paleoloricates and this group of aplacophorans represent parallel evolution of dorsal armour, which in chitons resulted in overlapping, articulating sclerites. The diversity of the mid-Silurian Gotland assemblage is examined against early evolutionary diversification of polyplacophorans, aplacophorans, and in relation to the overall record of Palaeozoic paleoloricate and neoloricate chitons. Peaks in diversity in early Ordovician, mid-Silurian and early Carboniferous times correspond to periods with widespread development of low latitude carbonate shelves. Neoloricates, apparently with an additional shell layer that contributed articulatory plates, appeared in the Devonian from where the fossil record remains poorly known.     

An early Ordovician patelliform gastropod, Palaelophacmaea, reinterpreted as a coelenterate

Yochelson, Ellis L. & Stanley, George D., Jr. 1981 12 15: An early Ordovician patelliform gastropod, Palaelophacmaea , reinterpreted as a coelenterate. Lethaia , Vol. 15, pp. 323–330. Oslo. ISSN W24–1164.
The fossil Palaelophacmaea criola Donaldson, from the early Ordovician Stonehenge Formation of central Pennsylvania, was described as a patelliform gastropod. A reinterpretation of the type lot and study of a few additional specimens provide the basis for an alternative placement. Palaelophacmaea is here assigned to the Hydrozoa, as a possible chondrophore. It has an exceptionally thin shell or test and concentric but irregular corrugations. Cambrian univalve genera having a more or less circular outline that are currently assigned to the Gastropoda or Monoplacophora should be reexamined to see whether they have the features of fossil chondrophore coelenterates rather than those of molluscs. The late Cambrian Palaeoacmaea Hall & Whitfield is removed from the monoplacophoran Mollusca and left unassigned as to phylum. We judge that at least some early Cambrian species of Scenella are probably coelenterate remains. * Early Ordovician , Palaelophacmaea, Gastropoda, Monoplacophora, Hydrozoa .     

新疆晚奥陶世晚期管状体及其古植物学物意义

本文系统描述了产自我国新疆塔里木盆地晚奥陶世晚期的管状体,包括2属8种,其中一个新种。通过对比研究,本管状体组合与Burgess和Edwards(1991）的下组合相当。根据本组合的研究,对Burgess和Edwards的组合进行了补充,其组合由不同类型的光壁管、低分异度的内壁加厚管组成、未见外壁加厚管和丝状体,组合的时代从中奥陶世到早志留世（晚Llandovery)。根据植物学的对比研究,管状体主要产自Nematophytales Lang.     

A Tremadocian (Early Ordovician) palaeoscolecidan worm from graptolitic shales in Hunan Province,South China

A new locality exposing Tremadocian (Early Ordovician) graptolitic shales in Hunan Province, South China, has yielded an exceptionally well‐preserved annulated worm. This palaeoscolecidan is described as Waflascolex changdensis gen. et sp. nov. and reveals extremely fine detail of the cuticle organization and plating array. The new taxon is characterized by three critical characters: incomplete plate rows that occur only on the posterior end of the worm (except in the posterior‐most area) and do not extend over the entire circumference or along the entire trunk; a regular rhomboidal array of platelets around intercalations; and reduced cuticular organization at the posterior termination. The unique cuticular organization and platelet ornamentation in the new taxon offer insight into functional differentiation of plates in the scleritome. Palaeoscolecid distribution through the early Palaeozoic is reviewed, showing that the worms were widespread in the Cambrian and Ordovician, but became more restricted during the Silurian. Ordovician palaeoscolecidans are diverse in scleritome architecture, and strikingly different from Cambrian taxa, indicating that this group diversified as part of the Great Ordovician Biodiversification Event.