Late Ordovician extinctions of bryozoans

An analysis of the final stratigraphic appearances of byrozoan species and genera, compiled in a world-wide bryozoan data base, revealed three discrete Late Ordovician extinctions. A Late Carddoc (Onnian) extinction was most pronounced on the plates of Baltica and Siberia. Endemic species and genera, confined to one plate and one lithotope were most affected and the extinction was coincident with increased migrations of bryozoan genera to Baltica and Siberia. The Late Caradoc extinction may be related to decreasing provinciality and competition between migrant and stenotopic taxa. Two major extinctions occurred in the Late Ashgill. The greatest of the two is recognized at the end of the Rawtheyan. and affected primarily taxa on the North American plate. The extinction at the end of the Hirnantian affected primarily Baltic taxa. The exact timing of the end-Rawtheyan extinction in North America cannot be established owing to incompleteness of the stratigraphic record. The Rawtheyan extinction occurred during a major glaciation centered in North Africa and a regression of epeiric seas. The large majority of North American survivors of the extinction are represented by Faunas preserved on Anticosti Island. which remained submerged during the regression. This evidence supports regression as a cause of the Rawtheyan extinctions in North America. The end-Hirnantian extinctions may be related to the ensuing transgression or to a wave of faunal migrations associated with the transgression. * Bryozoa, extinctions, Ordovician, Rawtheyan, Hirnantian, North America, Baltica .     

Paleozoic bryozoans of Mongolia

Stratigraphic assemblages characteristic for the Ordovician, Silurian, Devonian, Carboniferous, and Permian are presented based on the analysis of newly refined and supplemented data on the composition and distribution of Paleozoic bryozoans in Mongolia. Thirty-four auxiliary biostratigraphic units ranked as beds with bryozoans are established in order to subdivide the Paleozoic strata of Mongolia.     

Upper Ordovician bryozoans from the Xiazhen Formation of Yushan,northeastern Jiangxi,East China

The bryozoan fauna from the Xiazhen Formation (Katian, Upper Ordovician) of northeast Jiangxi Province, southeast China is reported here. Seventeen species of bryozoans belonging to fifteen genera and four orders are identified: Homotrypa yushanensis, Homotrypa sp., Prasopora yushanensis, Trematopora sp., Monotrypella sp., Rhombotrypa sp., Orbignyella sp., Constellaria jiangxiensis, Constellaria sp., Stictopora nicholsoni, Trigonodictya parvula, Ptilodictya ensiformis, Stictoporella sp., Pseudopachydictya sp., Nematopora sp., Arthrostylidae sp. indet., and Chasmatoporidae sp. indet. Four of these genera have been reported previously but nine genera (Trematopora, Monotrypella, Rhombotrypa, Orbignyella, Trigonodictya, Ptilodictya, Stictoporella, Pseudopachydictya, and Nematopora), one rhabdomesine and one fenestrate are found for the first time in the Late Ordovician strata of South China. Our palaeogeographical analysis suggests that the bryozoan association is typical for the Katian, which is mostly widespread in Laurentia, Siberia, Baltica and Mediterranean, and displays palaeobiogeographical relationships to the Laurentia–Siberia Province.     

Horizon of the oldest known bryozoans (Ordovician)

Among the six species of fossil bryozoans described by Xia et al. [Xia, F.S., Zhang, S.G., Wang, Z.Z., 2007. The oldest bryozoans: new evidence from the late Tremadocian (Early Ordovician) of East Yangtze Gorges. Journal of Paleontology 81 (6), 1308–1326] from the Fenghsiang Formation of the Chenjiahe section of Yichang and the Guanzhuangping section, Liujiachang Town of Songzi, the Yangtze Gorges, Nekhorosheviella nodulifera occurs in the pristinus Conodont Subzone (lower Subzone of deltifer Conodont Zone) and the five other species are associated with the conodont Paltodus deltifer deltifer and the graptolites Acanthograptus sinensis and A. erectoramus. Based on the conodonts and graptolites, this bryofauna stratigraphically corresponds to the A. sinensis Graptolite Zone or P. deltifer Conodont Zone. The bryofauna is of Tremadocian Age (Early Ordovician) and represents the oldest bryozoans so far as known.     

Zooidal anatomy in Ordovician and Carboniferous trepostome bryozoans

Fossil bryozoans sometimes contain fossilised brown bodies which remain after polypide degeneration. Position and shape of brown bodies as well as different skeletal diaphragms within living chambers allow outlining of autozooid anatomy in Palaeozoic trepostome bryozoans. In some trepostome bryozoans short autozooids were restricted by complete basal diaphragms to distal parts of autozooecia. In species with alternating hemiphragms, as in the specimen ofHemiphragma sp. described here, long autozooids occupied the entire autozooecial chambers. Their short polypides were positioned in distal parts of the autozooid. Both anatomic types correspond to the progressive polypide cycle afterBoardman. For species with ring septa, non-alternating hemiphragms (studied in an example ofNipponostenopora karatauensis) as well as without any diaphragms the stationary type of polypide cycle seems also to be possible. In that case, the polypides should be as long as the cystid.      

Stratigraphic distribution of selected acritarchs in the Ordovician subsurface,Canning Basin,Western Australia

Palynological investigation of 80 core samples from four boreholes, drilled through the Lower to Middle Ordovician marine sections of the central-northeastern Canning Basin, northwestern Australia, reveals diverse acritarch associations including a number of stratigraphically significant species. The stratigraphic succession studied, embracing the Willara Formation and conformably overlying Goldwyer Formation, has been independently dated as early Arenig to Llanvirn by conodont faunas (Oepikodus communis through Phragmodus-Plectodina zonal interval). Nine morphologically distinctive acritarch species have relatively narrow vertical ranges within the study interval and likely constitute serviceable palynostratigraphic indices for the Ordovician in and possibly beyond the Western Australian study area. Three new species, Comasphaeridium setaricum, Polyancistrodorus kunzeanensis, and Stelomorpha calix, are established; and one new combination, Baltisphaeridium variocavatum (Playford and Martin, 1984), is proposed. Other stratigraphically significant species include Petaloferidium comptum Playford and Martin, 1984 and Peteinosphaeridium sp. cf. P. exornatum Tongiorgi et al., 1995; together with Dorsennidium symmetricum (Lu, 1987) Sarjeant and Stancliffe, 1994, Peteinosphaeridium coronula Yin et al., 1998, and Striatotheca rarirrugulata (Cramer et al., 1974) Eisenack et al., 1976, which are found for the first time in Australia. The vertical distribution, and hence the stratigraphic utility, of each of the above species is documented through the studied sections.     

Upper Ordovician microphytoplankton of the Bill's Creek Shale and Stonington Formation,Upper Peninsula of Michigan,U.S.A.: Biostratigraphy and paleogeographic significance

An abundant, diverse, and well-preserved organic-walled microphytoplankton assemblage is described from the Upper Ordovician Bill's Creek Shale and the lower Stonington Formation (Bay de Noc Member) in the Upper Peninsula of Michigan, U.S.A. Based on graptolite and conodont evidence, the Bill's Creek Shale and Stonington Formation are Richmondian (=Ashgill) in age. The assemblage is dominated by acritarchs, which comprise 29 species (including the enigmatic palynomorph Gloeocapsomorpha prisca) assigned to 20 genera. The prasinophyte phycomata are represented by undifferentiated species of Leiosphaeridia and Tasmanites. In addition, chitinozoans are abundant, and scolecodonts and graptolite fragments are common. Paleontologic-palynologic and sedimentologic evidence indicates that the Bill's Creek Shale was deposited in a low-energy, shallow, nearshore marine environment. The overlying Bay de Noc Member of the Stonington Formation also accumulated in a low-energy, normal marine environment, but in a more offshore, somewhat deeper water setting. Both formations experienced minor transgressive and regressive episodes as indicated by fluctuations in the composition of the palynoflora. The combined Bill's Creek/Stonington acritarch assemblage closely resembles those described from the Richmondian-aged Maquoketa Shale (Missouri and Kansas), Sylvan Shale (Oklahoma), and Vauréal Formation (Anticosti Island, Québec, Canada). The overall composition of the acritarch assemblage from these four formations reflects a distinctive, recognizably Laurentian character. Nonetheless, many of the Bill's Creek/Stonington acritarchs have been reported from Upper Ordovician localities elsewhere, providing additional evidence for Late Ordovician cosmopolitanism of the marine microphytoplankton community. Additionally, the restricted stratigraphic range of many of the taxa further enhances their biostratigraphic application, both regionally and globally, and reaffirms the Richmondian (=Ashgill) age of the Bill's Creek Shale and Stonington Formation.     

Symbiosis between gastropods and bryozoans in the late Ordovician of Cumbria, England

Five examples of symbiosis between gastropods and trepostome bryozoans are described from the Ashgill (late Ordovician) of Cumbria (England). The gastropods are invariably found associated with the bryozoans. whereas the bryozoans may be free-living. Encrustation is considered to have taken place, in most cases, on mature, living gastropod shells, resulting in the trepostome utilizing the shell as a surrogate basal disc. Three specimens show encrustation on an empty gastropod shell, causing the trepostome to develop a basal disc similar to non-encrusting forms. The bryozoan colony probably benefited from the symbiotic relationship by increased water flow over the colony, caused by gastropod locomotion, whereas the gastropod was afforded protection from predators. The new species Spiroecus nidhoeggi. Diplotrypa hvergelmi and Monotrypa fontinalis are described.     

Upper Ordovician sequences of western Estonia

The Upper Ordovician (uppermost Caradoc-Ashgill) section of western Estonia consists of a series of seven open-shelf carbonate sequences. Depositional facies grade laterally through a series of shelf-to-basin facies belts: grain-supported facies (shallow shelf), mixed facies (middle shelf), mud-supported facies (deep shelf and slope) and black shale facies (basin). Locally, a stromatactis mud mound occurs in a middle-to-deep shelf position. Shallow-to-deep shelf facies occur widely across the Estonian Shelf and grade laterally through a transitional (slope) belt into the basinal deposits of the Livonian Basin.

Each sequence consists of a shallowing-upward, prograding facies succession. Sequences 1 (Upper Nabala Stage) and 2 (Vormsi Stage) record step-wise drowning of underlying shelf units (lower Nabala) that culminated in the deposition of the most basinal facies (Fjäcka Shale) in the Livonian Basin. Sequences 3–6 comprise the overlying Pirgu Stage and record the gradual expansion of shallow and middle-shelf facies across the Estonian Shelf. The Porkuni Stage (sequence 7) is bracketed by erosional surfaces and contains the shallowest-water facies of the preserved strata. The uppermost part of the section (Normalograptus persculptus biozone) is restricted to the Livonian Basin, and includes redeposited carbonate and siliciclastic grains; it is the lowstand systems tract of the lowest Silurian sequence 8. Sequence 7 and the overlying basinal redeposited material (i.e., the lowstand of sequence 8) correspond to the latest Ordovician (Hirnantian) glacial interval, and the bracketing unconformities are interpreted as the widely recognized early and late Hirnantian glacial maximums.

The sequences appear correlative to Upper Ordovician sequences in Laurentia. Graptolite biozones indicated that the Estonian sequences are equivalent to carbonate ramp sequences in the western United States (Great Basin) and mixed carbonate-siliciclastic sequences in the eastern United States (Appalachian Basin–Cincinnati Arch region). These correlations indicate a strong eustatic control over sequence development despite the contrasting tectonic settings of these basins.     

Systematics of Upper Cretaceous calloporid bryozoans with primitive spinose ovicells

The majority of fossil and Recent cheilostome bryozoans brood their larvae in ovicells. These double-walled, hood-like skeletal structures are thought to have arisen through modification of spines belonging to the zooid distal of the maternal zooid. Support for this hypothesis comes from the existence of ovicells constructed of multiple spines in a few Upper Cretaceous species belonging to two groups, microporids and cribrimorphs. Here we report the discovery of similar multispinose ovicells in a third group, calloporids, which are closely related to primitive cheilostomes that do not brood their larvae. The genus Distelopora Lang, 1915 from the Cenomanian ('Chalk Marl') of Cambridge is taken out of synonymy and shown to comprise the type species ( D. bipilata ) and two new species ( D. langi and D. spinifera ) of multiserial calloporids. Between 5 and 15 spine bases are arranged in a crescent on the gymnocyst of the zooid distal of each maternal (egg-producing) zooid in Distelopora . This indicates the presence of an ovicell formed by a cage of basally articulated spines. Similar ovicells represented by 18–19 spine bases occur in a uniserial calloporid from the German Campanian Allantopora krauseae Voigt and Schneemilch, 1986, which is made the type species of the new genus Unidistelopora . Another calloporid from the Cambridge Cenomanian has ovicells constructed by two claw-like, flattened, non-articulated and laterally juxtaposed spines. Described as Gilbertopora larwoodi gen. et sp. nov., this multiserial species provides a link between Distelopora and more typical cheilostome ovicells. The spines forming primitive ovicells provide a good example of exaptations, co-opted from their original function protecting the polypide of the distal zooid.     

New Famennian bryozoans (Upper Devonian) from Azerbaijan and Armenia

New bryozoans are described from the Famennian of Southern Transcaucasia: Petalotrypa myunkhbalaensis sp. nov. and Eridotrypella danzikensis sp. nov. (order Trepostomata) and Rectifenestella kadrluiensis sp. nov. and R. famenniensis sp. nov. (order Fenestrata). The paleobiogeographic relations that existed between Nakhichevan (Azerbaijan) and Armenia and other regions of Eurasia and the United States at the end of the Devonian are investigated.     

新疆柯坪和甘肃平凉上奥陶统底部附近的牙形刺

新疆柯坪大湾沟和甘肃平凉官司庄剖面是Nemagraptusb gracilis带及上奥陶统底界的重要剖面,研究这两剖面的牙形刺具有十分重要的意义。本文描述了采自新疆柯坪大湾沟和甘肃平凉官司庄剖面上奥陶统底界附近的牙形刺共13属18种。     

Functional morphologies of the columns of Upper Ordovician Xenocrinus and Dendrocrinus

Two common Upper Ordovician crinoids, Xenocrinus and Dendrocrinus , had distinctive columns that showed marked contrasts of both form and function. Xenocrinus baeri (Meek) had a tetragonal facet geometry but functioned in the normal manner for a symplectially articulated column. Column flexure was by bending, the four directions of curvature being limited by the tetragonal arrangement of crenulae. In contrast, columns of Dendrocrinus casei Meek, with a characteristic pentastellate symmetry, were able to twist, a functional adaptation not previously reported from crinoids; this was controlled by the unusual geometry of the crenularium. This suggests that some or all crinoid columns may be subject to twisting stresses, perhaps associated with changes in the current direction, that crenulae may resist passively. Functional morphology, column, crinoids, X enocrinus , D endrocrinus     

The trace fossil Thalassinoides from the Upper Ordovician of Tuva

Traces of Thalassinoides (the tunnels of unknown burrowing organisms) are described from carbonates of the Khondelensky layers of the Upper Ordovician of Tuva. Hitherto, this fossil was unknown in the Ordovician of the USSR. They demonstrate great similarity with Thalassinoides from coeval deposits of the Great Basin, USA. The traces are assumed to have been made in terrigenous-carbonate sediments deposited on the areas of a gently sloping shelf in quiet water, below wave base. □ Upper Ordovician, terrigenous-carbonate sediments, burrows.     

The graptolite correlation of the North American Upper Ordovician Standard

The North American Upper Ordovician reference standard, the Cincinnatian Series, contains rich shelly microfossil faunas in its type area in the Cincinnati Region but graptolites are uncommon in most of its shallow-water calcareous sediments. Consequently, the graptolite correlation of this key sequence has remained uncertain, in part, even controversial. A review of both previously published recently discovered graptolite Occurrences in the type Cincinnatian, combined with data from the important graptolite successions in Oklahoma New York-Quebec, has not only clarified the graptolite correlation of the Cincinnatian but also added new data on the morphology taxonomy, the vertical horizontal distribution, of several taxa. The information now at hand indicates that the Edenian Stage correlates with the C. spiniferus Zone, the Maysvillian Stage with the C. pygmaeus lower middle P. manitoulinensis Zone, the Richmondian Stage with the upper P. manitoulinensis , the D. complanatus , possibly part of the C. inuiti Zone. Comparison between graptolite conodont biostratigraphic evidence reveals no apparent conflict. Correlations are proposed between Upper Ordovician North American stages, graptolite conodont zones, successions in Texas, Oklahoma, Sweden, European graptolite zones, British series.     

Stratigraphic distribution and ecology of European Jurassic bivalves

Documentation of bivalve generic and species diversity and times of first and last appearance through successive Jurassic stages in Europe, together with data on turnover and changes in taxonomic and ecological composition of the faunas, indicate an approximation to the establishment of an equilibrium fauna by early Middle Jurassic times. Subsequently faunal change was slight compared with the early Jurassic. A diversity increase through the Lower into the Middle Jurassic correlates with an increase in the area of epicontinental seas, while a major species extinction in the early Toarcian is bound up with the onset of widespread stagnation associated with a rise of sea level. An increase of the generic extinction rate at the end of the period correlates with a regional marine regression. The mean species longevity is estimated at 15×10⁶ years. The ecological factors thought to control bivalve distribution are reviewed and four ecological associations distinguished: the reefal, lagoonal and nearshore and basinal marine.     

Paleoecology of Sphenothallus on an Upper Ordovician hardground

A hardground from the Upper Ordovician Dillsboro Formation near Dillsboro, Indiana, U.S.A., preserves an assemblage of encrusting and boring fossils on both top and bottom surfaces. The slab is inferred to have been an undercut ledge, and the dominant fossils of the assemblage, holdfasts of the tube-building worm Sphenothallus and trepostome bryozoans, are prevalent on both sides. The clumping of Sphenothallus holdfasts has been statistically demonstrated using a nearest-neighbor technique. Sphenothallus has also been shown to withstand overgrowth in interactions with bryozoans.     

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.