The Early Silurian brachiopod Eocoelia from the Hudson Bay Basin, Canada

Eocoelia akimiskii sp. nov. from the Lower Silurian (upper Telychian) Attawapiskat Formation of Akimiski Island, James Bay, Nunavut, is the first and oldest Eocoelia to be described from the Hudson Bay Basin, one of three largest inland basins of North America. The new species lacks dental plates, dental cavities, and marginal deflection or lip, which indicates a post-Aeronian form of the well-known Eocoelia lineage. In rib numbers, Eocoelia akimiskii falls between E. curtisi and E. sulcata , being closer to E. sulcata . In rib strength, however, the new species is more closely allied to E. curtisi . Although Eocoelia is currently assigned to the Rhynchonellida because of its lack of spiralia, several features (particularly the lack of a septalium, the presence of a unique notothyrial platform and cardinal process, and dense, free-hanging fibrous growth frills) of the genus and other leptocoeliids are distinctly atypical of the rhynchonellides. The new species occurs in an inter-reef, shelly packstone facies within the Attawapiskat Formation, which is characterized by coral-stromatoporoid reefs with abundant, diverse, reef-dwelling brachiopods and other shelly organisms. The close association of Eocoelia akimiskii with the Attawapiskat reefs supports a shallow subtidal (BA2) setting generally assigned to the Eocoelia Community. The reefs themselves, however, host an extremely abundant brachiopod fauna dominated by Pentameroides , Trimerella , Septatrypa , and Gypidula . Four species of Clorinda are also common elements of the reef-dwelling brachiopods. This demonstrates that the concept of the classic Early Silurian level-bottom brachiopod communities cannot be directly applied to reef-dwelling brachiopod communities.     

Earliest ontogeny of the Silurian orthotetide brachiopod Coolinia and its significance for interpreting strophomenate phylogeny

Well‐preserved juvenile specimens of the orthotetide brachiopod Coolinia pecten (Linnaeus, 1758 ) from the Silurian of Gotland, Sweden, demonstrate evidence of a planktotrophic larval habit. Larval shell morphology indicates the absence of a pedicle sheath: this character is otherwise typical of derived billingsellides, strophomenides and productides, which form the conventional strophomenide clade. The presence of a rudimentary colleplax structure in the larval shell of Colinia suggests instead a phylogenetic link to chiliate brachiopods and the enigmatic genus Salanygolina. This relationship suggests an early divergence of rhynchonellate and strophomenate brachiopods.     

Predatory borings in the inarticulate brachiopod Artiotreta from the Silurian of Oklahoma

A sample containing inarticulate acrotretid brachiopods, from the Silurian Clarita Formation of Oklahoma, was studied to determine the occurrence, nature and frequency of boring in the shells, and the possible identity of the boring organism. Specimens of Artiotreta parva Ireland, 1961 are abundant and specimens of Opsiconidion n. sp. and Acrotretella siluriana Ireland, 1961 are rare in the samples studied. All of the borings discovered are in A. parva except for one, which is in a specimen of O. n. sp. The boreholes are considered to be of predatory origin, and are similar in morphology to borings made by Recent gastropods. The small countersunk boreholes are equally frequent on pedicle and brachial valves, but preferentially located in particular regions of each valve. Boring frequency is high. The identity of the boring organism has not been determined, but is considered to be a soft bodied predator or an archaeo-gastropod.     

Cincinnetina,a new Late Ordovician dalmanellid brachiopod from the Cincinnati type area,USA: implications for the evolution and palaeogeography of the epicontinental fauna of Laurentia

Abstract: The most common forms of Late Ordovician dalmanellid brachiopods from the Cincinnatian type area, previously treated as either Dalmanella or Onniella, are assigned to Cincinnetina gen. nov. The new genus differs from Dalmanella and Onniella in having a consistently developed primary medial costa in the dorsal valve, a larger cardinal process that tends to develop a trilobed myophore, strongly differentiated fine and coarse punctae, and sparse aditicules. Cincinnetina can be distinguished from the closely related Paucicrura and Diceromyonia in its smaller trilobed cardinal process (when developed) that does not have a dominant medial lobe and does not extend into the delthyrial cavity of the ventral valve. Globally, Dalmanella and Onniella occur most commonly in deposits of relatively deep‐ or cool‐water palaeoecological settings, whereas in North America, Cincinnetina, Paucicrura and Diceromyonia are found mainly in carbonate‐rich deposits in warm‐water depositional environments, with Cincinnetina and Paucicrura most common in pericratonic settings and Diceromyonia in palaeoequatorial inland seas.     

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.     

The functional morphology and palaeoecology of the Dinantian brachiopod Levitusia

The functional morphology of Levitusia , based principally upon the type-species L. humerosa (J. Sowerby), is described and compared with living brachiopods. It is concluded that Levitusia species lived in tropical Dinantian seas with agitated waters and at shallow depths, perhaps no more than 50 m. The lithologies in which specimens are found support this view and indicate the need for some revision in our interpretations of some Dinantian sedimentary environments. Further evidence for the synonymy of L. christiani (de Koninck) in L. humerosa is presented.     

A biogeographically mixed, Middle Permian brachiopod fauna from the Baoshan Block, western Yunnan, China

A small brachiopod fauna is described from the carbonate rocks of the basal Shazipo Formation of the Baoshan Block, western Yunnan, south-west China, including significant new ventral and dorsal internal morphological features of Cryptospirifer omeishanensis Huang. This fauna is regarded as Wordian (Middle Guadalupian, Middle Permian) because of the presence of Cryptospirifer omeishanensis Huang and associated fusulinids ( Neoschwagerina craticulifera Zone). Palaeobiogeographically, the brachiopod fauna is of considerable interest because of its admixed nature characterized by typical warm-water Cathaysian elements intermingled with temperate Peri-Gondwanan taxa. This in turn is interpreted to indicate that the Baoshan Block may have been situated in an intermediate palaeogeographical position between Gondwanaland to the south and Cathaysia to the north during the Mid Permian and, as such, it probably furnished an important 'stepping stone' for the dispersal of Mid Permian eastern Tethyan marine invertebrate taxa (e.g. Cryptospirifer ) to the western Tethys.     

Ecological aspects of a silicified bivalve fauna from the Silurian of Gotland

The silicified Wenlockian (Silurian) bivalve fauna from MÖllbos, Gotland, is part of a life assemblage. The vast number of shells show unusual phenomena, e.g. shell repair, pearl and tumour formation, etc. A number of shells contain epibionts and bored, round holes. Presumptive predators of the bivalve community are discussed. Size-frequency distribution of the two most abundant species possibly reflects age classes. The fauna, comprising eleven species, is dominated by deposit-feeders (90 %). They exhibit niche diversification, including at least three different feeding levels within the sediment.     

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.     

Geochemical parameters of sedimentation and the distribution of Silurian brachiopod communities in Lithuania

Silurian rocks from borehole cores in the Baltic Basic were analyzed for Si, Al, Ti, Fe, Ca and Mg oxide content and chemical data were related to sedimentary environment, lithology and brachiopod community distribution. Evaluation of these relationships enables precise positioning of closely related communities within Boucot's (1975) brachiopod Benthic Assemblages. Preliminary attempts are made to evaluate climatic change based on Al/Ti oxide ratios.     

Regular mode of increase, and constrained but variable growth, in the Silurian rugose coral Stauria favosa

Stauria favosa exhibits a typical pattern of axial division (tripartite or quadripartite). The four protosepta first appear with a definite polarity in offset corallites, and metasepta are inserted serially, following Kunth's rule. There are no variations in essential modes of asexual reproduction. However, at the corallum periphery, up to four offset corallites appear to have been derived by discrete offsetting within the parent calices. Detailed observations of vertical morphological changes, however, indicate that this apparently unique mode of increase is derived from axial division at the sites of the four protosepta, as usual in this species. Just after the initiation of division, the contraction of polyps occurred in response to somewhat accidental, deteriorating habitat conditions. The combination of asexually divided corallites and rejuvenescence resulted in phaceloid daughter corallites, which subsequently underwent partial mortality at the corallum periphery. Stauria favosa always exhibits not only regular modes of asexual reproduction and septal insertion under the strict control of phylogenetic and developmental constraints, but also variable growth forms of corallites and coralla. This variability, though within limited options, is palaeoecologically significant for successfully coping with unpredictable habitats.     

The root of the problem: palaeoecology of distinctive crinoid attachment structures from the Silurian (Wenlock) of Gotland

The holdfast (attachment structure) is the most understudied aspect of the palaeoecology of the endoskeleton of fossil crinoids. A new collection of well-preserved holdfasts from a recently reopened quarry at Hunninge, Gotland, in Homerian (upper Wenlock) strata includes several morphologies. The most common are terminal dendritic radicular holdfasts (TDRHs) that may be derived from the cladid Ennallocrinus d'Orbigny. These have a consistent morphology of five, equally spaced, long radices that spread across the sea floor. These crinoids were gregarious, and TDRHs in a group commonly show the same radice orientations. The radices have a large axial canal compared with those of modern crinoids; each included, at least, nervous tissues. Taken together, these features suggest that, apart from attachment, these distinctive TDRHs may have served a sensory function. Other holdfasts in this assemblage also show monospecific aggregations, perhaps suggesting biochemical attraction such as that shown by certain other sessile invertebrates such as barnacles.     

Ribbing patterns in the brachiopod Diceromyonia

Ribbing patterns in two collections of the Ordovician brachiopod Diceromyonia from central North America are compared using the system invented by Bancroft (1928; Manchester Memoirs 72 ) for naming ribs. Simple and nearly symmetric arrangements of frequently occurring branches characterize both collections, with much of the complexity evident in larger valves due to irregular occurrence of low-frequency branches on either side of major ribs. Examination of median sectors reveals little support for Bancroft's derivation of the rhipidomelloid midrib from isorthoid forms, and an alternative non-evolutionary interpretation is proposed in which median patterns appear as predictable outcomes of a simple space-filling process. The approximately 2500 ribs identified in this study are treated as a population in their own right: orderly patterns which emerge are not inferred but are simple summaries of complete counts, obtained by hand. This approach is quick and painless compared to methods using order relations to assess ribbing differences in dalmanellid brachiopods. For this reason, it may encourage systematic study of ribbing patterns among these taxa and lead ultimately to better understanding of the utility of ribbing in their taxonomy.     

The Cambrian brachiopod fauna from the first-trilobite age Shuijingtuo Formation in the Three Gorges area of China

The Yangtze platform of South China offers evidence within its Ediacaran–Cambrian geological record of the Cambrian explosion and diversification events in metazoan history. To understand the explosive radiation of animals and the environments in which it took place, the basal Cambrian fauna succession of the Aijiahe section in the Three Gorges area, western Hubei Province, has been studied, revealing the earliest brachiopod fauna (Tsunyidiscus trilobite Zone) in this region, which was dominated numerically by acrotretoids. This is accompanied by abundant skeletal fossils including minute well-preserved phosphatized archaeocyath cups and an assortment of abundant sponge spicules, chancelloriids, mollusks, hyoliths, and bradoriids, retrieved by acid-etching limestone interbeds in the black shale-dominated Shuijingtuo Formation (Series 2). The brachiopods comprise two species of acrotretoids, two types of botsfordiids (Botsfordiidae gen. et sp. indet. A and B), and four species of linguloids. Of the latter, Spinobolus popovi n. gen. n. sp. is strikingly distinctive and typified by spine-like ornamentation seen for the first time in the lower Cambrian; the remaining three linguloid genera, Palaeobolus, Eoobolus, and Lingulellotreta, have a trans-paleocontinental distribution. The Three Gorges Shuijingtuo brachiopod assemblage differs from that of the upper Atdabanian Stage (Cambrian Stage 3) in Siberia and South China, but shows great similarities with those discovered in the Tsanglangpuan (equivalent to Botoman or Stage 4) Stage of eastern Yunnan Province, Siberia, and South Australia, suggesting a much more prolonged sedimentary hiatus in basalmost Shuijingtuo Formation of the Three Gorges area than previously expected. The presence of such unconformities provides a caveat to stable isotope-based correlations that involve a number of discussions of global ocean geochemical changes across the time interval that witnessed Cambrian explosion of metazoans.     

A soft-bodied lophophorate from the Silurian of England

Soft-bodied taxa comprise an important component of the extant lophophorate fauna, but convincing fossils of soft-bodied lophophorates are extremely rare. A small fossil lophophorate, attached to a brachiopod dorsal valve, is described from the Silurian (Wenlock Series) Herefordshire Lagerstätte of England. This unmineralized organism was bilaterally symmetrical and comprised a subconical body attached basally to the host and partially enclosed by a broad ‘hood’; the body bore a small, coiled lophophore. Where the hood attached laterally, there is a series of transverse ridges and furrows. The affinities of this organism probably lie with Brachiopoda; the hood is interpreted as the homologue of a dorsal valve/mantle lobe, and the attachment as the homologue of the ventral valve and/or pedicle. The ridges are arranged in a manner that suggests constructional serial repetition, indicating that they are unlikely to represent mantle canals. Extant brachiopods are not serially structured, but morphological and molecular evidence suggests that their ancestors were. The new organism may belong to the brachiopod stem group, and might also represent a significant element of the Palaeozoic lophophorate fauna.     

Silurian bolbozoids and cypridinids (Myodocopa) from Europe: pioneer pelagic ostracods

Abstract: Bolbozoid and cypridinid myodocope ostracods from France, the Czech Republic and Sardinia consist of four genera (one new: Silurocypridina) and at least ten species (eight new: Bolbozoe acuta, B. rugosa, B. parvafraga, Parabolbozoe armoricana, Silurocypridina retroreticulata, S. variostriata, S. calva and Calocaria robusta). They have a functional design, palaeogeographical distribution, depositional setting and faunal associates that suggest that they were swimmers in the water column, living in dim light, relatively deep environments, above near bottom dysoxic/anoxic conditions. These myodocope faunas include some possible pelagic species sensu stricto, with widespread palaeogeographical (transoceanic) distribution, and some species that occupied hyperbenthic niches and were capable of making short incursions to the sediment to scavenge on carrion. These data support the model that Silurian myodocopes were pioneer pelagic ostracods.     

Evolution of the Rhynchotrema–Hiscobeccus lineage: implications for the diversification of the Late Ordovician epicontinental brachiopod fauna of Laurentia

Multivariate analysis based on nine biometric characters of 171 Late Ordovician rhynchonellide specimens from nine upper Sandbian–upper Katian localities in North America supports the hypothesis that one of the diagnostic taxa of the North American epicontinental brachiopod fauna, Hiscobeccus, evolved from Rhynchotrema, which lived predominantly in peri‐cratonic settings. The oldest known Hiscobeccus, H. mackenziensis of early Katian age, exhibits transitional characteristics between Rhynchotrema and Hiscobeccus, and it clusters more closely with Rhynchotrema than with younger species of Hiscobeccus of mid–late Katian (Maysvillian–Richmondian) age. Diversification of the Hiscobeccus lineage in epicontinental seas was characterized by drastic increase in shell size, globosity and lamellosity, especially in palaeoequatorially located inland basins. Such morphological trends are interpreted as adaptation to relatively shallow, muddy substrates, moderate water turbulence, relatively low oxygen content and unstable supply of nutrients in generally overheated epicontinental seas with sluggish circulation.     

A route to recovery: The early Silurian shallow-water shelly fauna in the northern Oslo basin

The shelly fauna of the lowest part of the Sælabonn Formation in the Hadeland district of the Oslo Region provides a rare insight into the shallow-water biota of the earliest Silurian and hence the start of biotic recovery after the end Ordovician extinction event. It is dominated by the brachiopods Dalmanella cf. pectinoides Bergström, Coolinia cf. columbana (Reed), Leptaena cf. haverfordensis Bancroft and Zygospiraella scotica (Salter) together with the trilobite Acernaspis elliptifrons (Esmark). The first three species in this unique association belong to eurytopic Ordovician survivor genera which were also present in the underlying unit in Hadeland and continued to thrive during the Silurian. Significantly, Zygospiraella and Acernaspis have no unequivocal Ordovician record but diversified rapidly and became common during the early Silurian (Rhuddanian) in many parts of the world. Moreover, Acernaspis and Zygospiraella were highly eurytopic, pioneer taxa that were among the first elements of the Rhuddanian shelly fauna to appear in many parts of the world especially around the margins of the remnant Iapetus Ocean, following the drowning of areas previously emergent during the late Ordovician sea-level lowstand. The Hadeland fauna provides evidence of a previously undocumented route (among several) by which life ultimately returned to the global diversity plateau established in the mid-Ordovician.     

A relict Ordovician brachiopod fauna from the Parakidograptus acuminatus Biozone (lower Silurian) of the English Lake District

At Yewdale Beck in northern England, a brachiopod fauna dominated by species of Hindella, Kinnella, Mirorthis, Paromalomena and Plectothyrella , occurs interbedded with a graptolite fauna which includes Akidograptus ascensus , Atavograptus ceryx , Persculptograptus parvulus and Normalograptus spp. (including probable examples of N. normalis , N. angustus and N. medius ) within the lower part of the Skelgill Formation. This suggests that taxa of the terminal Ordovician Hirnantia fauna occur within the lower P. acuminatus Biozone, representing the youngest documented occurrence of the Hirnantia brachiopod fauna. Biostratigraphy, brachiopods, graptolites, Ordovician-Silurian boundary.     

Endolithic micro-organisms silicification of a bivalve fauna from the Silurian of Gotland

The silicified Wenlockian bivalve shells at Möllbos have been fragmented to a considerable extent. Shells which were broken prior to silicification exhibit possible original shell layers while those which were fragmented during laboratory treatment show no primary structures. The fauna at Möllbos was attacked by endolithic micro-organisms. The borings of these were then coated with a carbonate envelope. After burial the unattacked original shell material was dissolved the envelope silicified. Later, empty moulds were subsequently filled with drusy calcite occasionally with quartz crystals. A third silicification went occurred at a later diagenetic stage when matrix had become lithified.