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.     

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.     

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.     

Epizoans on late Ordovician brachiopods from Southeastern Indiana

The frequency of epizoans (cornulitids, inarticulate brachiopods, bryozoans, solitary and colonial rugosan corals) on over 8000 specimens of articulate brachiopods (four strophomenids, five orthids, one rhynchonellid) was calculated for four stratigraphic horizons in the Dillsboro Formation of southeastern Indiana. Frequency of shells encrusted correlates significantly with the surface area of the valves. Punctae in brachiopod shells (Onniella meeki) may have deterred larval settlement of epizoans. Coarse ribbing on articulates deterred encrustation by the inarticulate brachiopod. The horn coral shows a preference for attachment to the anterior of Hiscobeccus capax. Bryozoans show a preference for the incurrent lateral margins of inferred living hosts, suggesting rheotropic behavior by settling larvae. Inarticulate brachiopods are concentrated around the sloping commissure of the brachial valve of strophomenids, suggesting geotropic behavior and/or selective survival of settling larvae. Inarticulates deterred overgrowth by bryozoans. High frequencies of encrustations of the medial region of pedicle valves of orthids and strophomenids probably reflect post‐mortem encrustations. Alternating intervals of slow sediment accumulation punctuated by tropical storms and rapid shell burial may account for the high frequency of shells with either their entire surface veneered or only a very small area encrusted by bryozoans.     

New brachiopods from the Ordovician of northeastern Russia

New brachiopod taxa, Starnikoviella gen. nov., with the type species S. settedabanica sp. nov. (family Clitambonitidae), and Avdeevella gen. nov., with the type species A. mica sp. nov. (subfamily Glyptomeninae), from the Middle–Upper Ordovician boundary deposits of northeastern Russia are described.     

Studies of fossil brachiopods of Mongolia: Ordovician,Silurian, Devonian,Carboniferous, and Permian

Fossil brachiopods from the Ordovician, Silurian, Devonian, Carboniferous, and Permian deposits of Mongolia have been studied for the last forty-five years by the Joint Soviet-Mongolian (later RussianMongolian) Paleontological and Geological Expeditions. New data on the taxonomic composition, stratigraphic and geographic distribution of the brachiopod assemblages have been obtained. The brachiopod systematics has been further refined and detailed, and the stratigraphic and correlation scales and biogeographic reconstructions have been elaborated for the Paleozoic of Mongolia.     

Epithelial moulds from some Upper Ordovician acrotretide brachiopods of Ireland

Three new genera of acrotretid brachiopods are of general morphological interest in that they display polygonal mosaics on the internal shell surfaces, features which are believed to be moulds of the outer epithelial cells. The mosaic is further noted in a specimen of Eoconulus , a form which may also belong to the Acrotretacea. Such mosaics have not previously been recorded from the Acrotretacea. Polygonal mosaics, epithelial moulds, outer epithelium , Eoconulus, Acrotretacea, Ordovician.     

Late Silurian brachiopods from Darhan Mumingan Joint Banner, Inner Mongolia

Seven brachiopod species (including three new species) and three undetermined forms are described and figured here from an unnamed stratigraphic unit which mainly consists of argillaceous and bioclastic limestone near the stratotype of the Late Silurian Xibiehe Formation of the Xibiehe section in the Darhan Mumingan Joint Banner, north of the Bateaobao area, Inner Mongolia. Three new species, Eoschizophoria minuta, Navispira navicula and Howellella asiatica are established with transverse serial sections for their interiors. All brachiopod taxa described here show that this unnamed stratigraphic unit is higher than the Late Silurian Xibiehe Formation in the Xibiehe section and is possibly transitional between the Late Silurian and the Early Devonian. Authors temporarily regard this unit as the upper part of the Xibiehe Formation in this paper.     

Miospore evolution from the Ordovician to the Silurian

The definition of the term cryptospore is amended to include only spores thought to be produced by embryophytes and to exclude all enigmatic palynomorphs. Cryptospores are included here in the miospore group.The oldest published assemblage of typical cryptospores is Llanvirn in age. The evolution of cryptospore assemblages is very slow from the Llanvirn to the early Llandovery. The Telychian and Sheinwoodian stages are periods of drastic impoverishment in cryptospore biodiversity, followed by an important modification in miospore assemblages. This step in the evolution of the vegetation is correlated with the rapid transgression of seas on the continent during the early Llandovery. On the other hand, the Hirnantian glaciation does not affect diversity of the miospore assemblages.     

Ordovician and Silurian coralline red algae

Solenopora gotlandica Rothpletz, hitherto placed in the Solenoporaceae Pia, is locally common in the mid-Silurian of Gotland and Wales. It possesses small cells, radial monomerous thallus organization, trichocytes, and sporangia arranged in irregular sori. Solenopora gotlandica is not congeneric with S. spongioides , the type species of Solenopora. It closely resembles the extant coralline red alga Sporolithon Heydrich. S. gotlandica is here transferred to Craticula gen. nov., and Craticula is placed in the Craticulaceae fam. nov., which morphologically closely resembles the Mesozoic-Cenozoic family Sporolithaceae. The Craticulaceae is placed in the extant rhodophyte order Corallinales. Middle-Late Ordovician Petrophyton kiaeri Høeg resembles Late Ordovician Solenopora richmondense Blackwell et al. , which has sporangia in calcified compartments arranged in sori. These fossils may be craticulaceans. The apparent gap in the fossil record between Silurian Craticula and Cretaceous Sporolithon favours separation of these genera at family level. However, future work may demonstrate that the Craticulaceae is a junior synonym of the Sporolithaceae. Recognition of the Craticulaceae extends the earliest record of the Corallinales from Cretaceous to Silurian, and possibly Ordovician.     

Silurian reef-dwelling pentamerid brachiopods, Wisconsin and Illinois, USA

Eight species of pentamerid brachiopods, assigned toApopentamerus, Capelliniella, Kirkidium, Lissocoelina, Rhipidium, Supertrilobus andStenopentamerus, occur in stromatoporoid-coral reefs of the Silurian (Wenlockian) Racine Formation, Wisconsin and Illinois, USA. One new taxon,Rhipidium kuglitschi n.sp., is described. In contrast to level-bottom communities, where single pentamerid species occur in great numbers over large areas, pentamerids were generally rare members of the Racine reef fauna, and most species have localized distributions. An increase in diversity from one pentamerid species in the north to six species in the south correlates with a southward increase in reef size. Four morphologic groups of pentamerid species differed in the shape of the mantle cavity, which suggests different dynamics of suspension-feeding. Where two or more species are present in single reefs, co-occurrence of these groups represents a partitioning of the pentamerid niche. The relatively high diversity, local distributions, and morphologic distinctions among the Racine pentamerid fauna represent an evolutionary radiation and functional specialization of pentamerids in Wenlock reefs.     

Algal borings and framboidal pyrite in Upper Ordovician brachiopods

Algal borings in the shells of the articulate brachiopods Plaesiomys subquadrata (Hall) and Hebertella sinuata (Hall) from the Richmond Formation of Ohio are empty, or partially to entirely filled with pyrite. The pyrite occurs as single framboids and other crystal forms, or in chains filling the bores. The borings provide some insight into the early diagenetic history of the Richmond sediments near Cincinnati. Pyritization probably occurred within a few years, only a short distance (a few centimeters) below the sediment surface through the activities of sulfur-reducing bacteria. Pyrite precipitated around a nucleus such as a bacterium or algal cell, or developed within an organic structure such as an algal cell or organic membrane.     

Global diversity and endemism in Early Silurian (Aeronian) brachiopods

Silurian brachiopods of Middle Llandovery (Aeronian) ages are reviewed, and 215 genera are identified here, compared with 109 in the preceding Early Llandovery (Rhuddanian), indicating a recovery‐radiation interval after the end‐Ordovician mass extinction. The chief regions in which they are found are the continents of South China, Avalonia‐Baltica, Laurentia and Siberia, which were all at tropical latitudes with the exception of Avalonia. In addition, the very large superterrane of Gondwana, although with patchy brachiopod distribution, included temperate faunas, as well as subtropical faunas (in Iran and Afghanistan). Aeronian brachiopods greatly increased in diversity, with dominance of four major groups: orthides and strophomenides (which had flourished previously in the Rhuddanian), pentamerides and atrypides (which became dominant in the Aeronian), and many newly evolved taxa, and occupied deeper water and wider ecological niches (level bottom and reef) than those in the Rhuddanian. Each of the continents has some endemic genera, but there is a greater proportion of them in South China, where some groups (such as the pentamerides and atrypides) are more diverse and others, such as the orthides, are much less common than elsewhere. Affinity indices (AI) show that two megaunits can be recognized: South China and Avalonia‐Baltica‐Laurentia (ABL); Siberia seems to have been loosely connected with ABL and even more loosely to South China presumably because of its geographical separation in the Northern Hemisphere. The separation of South China from the other megaunits is further supported by cluster analysis.     

Upper Ordovician and Lower Silurian chitinozoans from central Nevada and Arctic Canada

Late Ordovician and Early Silurian chitinozoans from the uppermost Vinini Formation, and the Hanson Creek Formation in central Nevada and the lower Cape Phillips Formation, Cape Manning section, in Arctic Canada have been re-investigated and a new chitinozoan biozonation is proposed. The Upper Ordovician of central Nevada can easily be correlated to that of Arctic Canada through the common occurrence of the Ordochitina nevadensis biozone in both regions. No such correlation, however, is possible with the Late Ordovician of Anticosti Island in eastern Canada because of the absence of the index Upper Ordovician chitinozoan zonal species of central Nevada such as Belonechitina tenuispinata sp. nov, Ordochitina nevadensis sp. nov. and Nevadachitina vininica gen. nov., sp. nov. in the former area.One new genus, Nevadachitina, and nine new species, Eisenackitina ripae, Belonechitina martinica, Nevadachitina vininica, Nevadachitina praevininica, Ordochitina nevadensis, Belonechitina tenuispinata, Belonechitina parvispinata, Tanuchitina laurentiana, Angochitina hansonica are described and illustrated in this paper and four species are left in open nomenclature.     

Predation in the Ordovician and Silurian of Baltica

Signs of predation appear in the Middle Ordovician of Baltica. Shell repair dominates over the predatory borings in the Ordovician and Silurian. Predators attacked molluscs, brachiopods and tentaculitoids in the Ordovician and molluscs, tentaculitoids, brachiopods and ostracods in the Silurian. There is an increase in the number of prey species in the Late Ordovician, which could be related to the Great Ordovician Biodiversification Event. Molluscs are the favourite prey taxon in the Ordovician, but in the Silurian, molluscs became less dominant as the prey. This is probably not an artefact of preservation as Ordovician and Silurian molluscs are equally well preserved.     

Cementing strophomenide brachiopods from the Silurian of Gotland (Sweden): Morphology and life habits

Dorsal valves of the cementing strophomenide brachiopods Leptaenoidea silurica Hedström and Liljevallia gotlandica Hedström are described for the first time, and it is shown that both these species could also live ambitopically. The lower Wenlock Scamnomena rugata (Lindström) represents young individuals of the ambitopic variant of Leptaenoidea silurica, and is placed in synonymy, resulting in the valid name for the taxon being Leptaenoidea rugata. The range of this species now spans the whole Wenlock, from the upper Visby Formation to the Klinteberg Formation, and possibly even into the Ludlow. Ambitopic gerontic specimens of L. rugata develop very thick shells, in which the ventral valves have strong curvature, and become deeper not by geniculation but by successive mantle retractions and subsequent re-growth, in a way similar to that of atrypides. This shape was probably an adaptation to “floating” on softer substrates. The thickened gerontic dorsal valves have well-developed lophophoral support, showing the shape of the lophophore, which comprises two branches that curve inwards and then backwards; the lophophore was probably ptycholophous and similar to that in living members of the Thecideidina. Ambitopic specimens of Liljevallia could grow to a much larger size than cementing forms, where the dorsal valves have very large, posterior-facing cardinal process lobes and deeply impressed muscle fields and anterior scars. The presence of a ventral process and long, posteroventrally elongated cardinal process lobes, and the absence of dental plates reveals that Liljevallia was probably an early member of the Douvillinidae and is thus removed from the Leptaenoideidae.     

The first report of epipunctae in non-plaesiomyid brachiopods from the lowest Silurian of southeastern China

Epipunctae are microscopic perforations that do not penetrate the shell but are confined to the outer layers of the shell. They have been known previously only in the orthidine family Plaesiomyidae. The striated walls of epipunctae are considered unique to brachiopods and thus of significance for the study of stem-group brachiopod. In this study, we report the first occurrence of epipunctae in a non-plaesiomyid brachiopod based on a well-preserved external mould of Cathaysiorthis yushanensis from the lower Silurian Shiyang Formation of Jiangxi, southeastern China. The species belongs to the Family Cathaysiorthidae which differs from the Family Plaesiomyidae in many aspects especially in dorsal internal structures. The new data imply that epipunctae may be more widespread in orthide brachiopods than previously thought, strengthening the notion that this is a plesiomorphic character of the brachiopod shell. The discovery of epipunctae, however, depends on excellent preservation of the outer surface of the shells as well as careful cleaning and observation.     

Taxonomy and palaeoecology of the mollusc Pterotheca from the Ordovician and Silurian of Scotland

A large sample of the bizarre bellerophontiform mollusc Pterotheca from the Upper Llandovery Wether Law Linn Formation in the Scottish Pentland Hills shows marked polymorphic asymmetry. A survival strategy that presents to a predator a mosaic of different search images is suggested to account for the marked variability of the Pterotheca morphology. Multivariate comparisons with Pterotheca mullochensis from the Llandovery of Girvan, P. simplex from the Caradoc of Girvan, P. cf. simplex from the Caradoc of the Northern Belt of the Southern Uplands and P. aff. mullochensis from the Llandovery in the Pentland Hills have helped clarify the morphological differences between these groups. A new species, Pterotheca trimerelloides , is established for the specimens from the Pterotheca beds in the Wether Law Linn Formation. □ Mollusca, Silurian, Ordovician , Pterotheca, taxonomy, palaeoecology.     

Late Ordovician and early Silurian Proetida (Trilobita) from north-western and central Europe

Species belonging to genera of three families of the Proetida (Proetidae, Rorringtoniidae and Toernquistiidae) are described from the upper Ashgill of southern Sweden, and the same or related species noted from the adjacent parts of northern and central Europe. Proetidae are represented by Paraproetus brevifrons (Angelin), Ogmocnemis irregularis Kielan, Ogmocnemis jaanussoni sp. nov. and Parvigena putealis sp. nov.; it is argued that all belong to the Cornuproetinae, not Tropidocoryphinae where they were previously classified. Ogmocnemis is resurrected for some Upper Ordovician species included previously in Decoroproetus . The rorringtoniid genera Rorringtonia and Madygenia are recognised for the first time in southern Sweden, and it is argued that these are probably closely related, and both are included in the same family; both are recorded for the first time from Silurian (Llandovery) strata. Zetaproetus is claimed to be a junior subjective synonym of Rorringtonia. Solariproetus , first described from north-west China, is assigned to the Toernquistiidae as a late, derived member, and S. prosthemesos sp. nov. is described from late Rawtheyan strata in the Holy Cross Mountains, Skåne and the English Lake District. Comments are given on the stratigraphical and palaeogeographical distribution of these taxa.