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Traditionally, rarefaction curves are used to test sampling sufficiency; however, such measures are not always robust. Similarly, the Pareto Principle and “90/70”rule, commonly applied to biological sampling, are also inappropriate for palaeontological data. Here, we argue that the sufficiency of sampling could be determined by the consistency between a calculated diversity and the “actual” diversity. We have applied a method introduced from biology to estimate the species richness at four localities with the early Silurian brachiopod Cathaysiorthis fauna in South China and to estimate their actual diversity using the software EstimateS. Sample size corresponding to a given diversity was calculated by means of curve fitting, which can provide important information on future sampling strategies. Diversity estimation is a relatively immature field in palaeontology that promises accurate diversity comparisons. Here, we suggest that rarefaction should be accompanied by estimation assessment to extract from sa...

The great ordovician biodiversification event (GOBE) is one of the major biotic revolutions in Early Paleozoic marine ecosystems. Through three internationally coordinated IGCP projects (IGCP 410, 503 and 591) in the past two decades, an enormous amount of paleontological, stratigraphical, paleoecological, and paleobiogeographical data have been accumulated, leading to a number of significant discoveries and common understandings regarding the GOBE. First, the multiple radiation pulses spanned nearly the entire Ordovician Period (more than 40 Ma), with several diversity acmes. Second, the biotic radiations were highly diachronous, characterized by different timing and rates in different regions. Third, the radiations were not associated with fundamental changes in the marine environments, but were rather controlled by both intrinsic factors (macroevolutionary regularities) and extrinsic forces (gradual environmental changes). Case studies conducted in South China have contributed greatly to the understanding of GOBE at the global scale. Future work on the GOBE includes more in-depth case studies to reveal the exact process and pattern of the radiations by combining systematic paleontology, high-resolution biostratigraphy, sedimentology, plate tectonics and geochemistry. This will help improve our understanding of the dynamics of the radiation and co-evolution between the organic and inorganic realms of Ordovician marine ecosystems.

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.

The Geobiodiversity Database (GBDB – www.geobiodiversity.com), an integrated system for the management and analysis of stratigraphic and paleontological information, was started in 2006 and became available online in 2007. Its goal is to facilitate regional and global scientific collaborations focused on regional and global correlation, quantitative stratigraphy, systematics, biodiversity dynamics, paleogeography and paleoecology. It is unique among global, public access databases in that it is a section-based online database system, incorporating data from a wide range of disciplines of stratigraphy and paleontology, with inherent interrelationship between different kinds of data sets. It provides the capability of completely digitizing raw data, as well as integrating of different interpretations to the same paleontological and stratigraphic content. Several Windows-based visualization and analysis applications, either fully integrated with the database or supported by subset-export functions, have been developed to make the database more useful as a scientific and educational tool. The GBDB became the formal database of the International Commission on Stratigraphy (ICS) in August 2012 at the 34th International Geological Congress in Brisbane, and will produce comprehensive and authoritative web-based stratigraphic information service for global geoscientists, educators and the public.

Taxonomic identification of Paleozoic solitary rugose corals relies heavily on serial sections to determine the ontogenetic development of skeletal structures. Previously, small rugose corals have been difficult to study because their size and curvature make it difficult to serially section them at a right angle to the longitudinal growth axis. In this paper, we describe a newly designed apparatus, with a rotatable clamp to hold the fossil coral. This guarantees easy adjustment of the cutting angle so that serial sections can be made at changing angles that are always at a right angle to the curved longitudinal axis of a rugose coral. In addition, the apparatus allows the distance between consecutive sections to be controlled precisely to mm scales.

Corals are comparatively abundant in the upper member of the Baiyun’an Formation (upper Aeronian to lowerest Telychian, Llandovery, Silurian) of Huaying, eastern Sichuan Province, Southwest China. Three coral associations have been recognized, i.e., in ascending order, the Mesofavosites Association, the Dentilasma Association, and the Paraceriaster Association. The Mesofavosites Association occurs at horizons indicating a shallow and high-energy environment. It is dominated by large favositids accompanied by a few broken pieces of halysitids and small solitary rugose corals. The Dentilasma Association has a pretty high coral diversity, dominated by the solitary rugose coral Dentilasma meitanense, together with some subordinative heliolitids and favositids. The sedimentary evidence shows that this association might live in a quiet and nutritive shallow water environment. The Paraceriaster Association yields in the grayish green calcareous mudstone indicating a comparatively deep water environment. Systematically it contains abundant fasciculate Paraceriaster micropora and a few favositids. The distinction among these three coral associations shows that corals could adopt different adaptive strategies under different environments, which provide some new data for the investigation on the ecology of Silurian corals and further study on the “Lower Red Beds”.

The phylogeographical evolution and the consequent changing distribution and diversity of rhynchonelliform brachiopods through the Ordovician are linked to the dynamic palaeogeography of the period. The Early Ordovician (Tremadocian and Floian) is characterized by globally low-diversity faunas with local biodiversity epicentres, notably on the South China Palaeoplate; low-latitude porambonitoid-dominated faunas with early plectambonitoid and clitambonitoid representatives, as well as high-latitude assemblages mostly dominated by orthoids, can be recognized, but many taxa are rooted in Late Cambrian stocks. The Early Ordovician displays a steady increase in rhynchonelliformean biodiversity, which was mostly driven by the increasing success of the Porambonitoidea and Orthoidea, but the billingsellids and early plectambonitoids also contributed to this expansion. During the Early to Mid Ordovician (Dapingian–Darriwilian), marine life experienced an unprecedented hike in diversity at the species, genus and family levels that firmly installed the suspension-feeding benthos as the main component of the Palaeozoic fauna. However, this may have occurred in response to an early Darriwilian annihilation of existing clades, some of which had been most successful during the Early Ordovician. New clades rapidly expanded. The continents were widely dispersed together with a large number of microcontinents and volcanic arcs related to intense magmatic and tectonic activity. Climates were warm and sea-levels were high. Pivotal to the entire diversification is the role of gamma (inter-provincial) diversity and by implication the spread of the continents and frequency of island arcs and microcontinents. The phylogeographical analysis demonstrates that this new palaeogeographical configuration was particularly well explored and utilized by the strophomenides, especially the Plectambonitoidea, which radiated rapidly during this interval. The porambonitoids, on the other hand, were still in recovery following the early Darriwilian extinctions. Orthides remained dominant, particularly at high latitudes. Biodiversity epicentres were located on most of the larger palaeoplates, as well as within the Iapetus Ocean. Provincial patterns were disrupted during the Sandbian and early Katian with the migration of many elements of the benthos into deeper-water regimes, enjoying a more cosmopolitan distribution. Later Katian faunas exhibit a partition between carbonate and clastic environments. During the latest Katian, biogeographical patterns were disrupted by polewards migrations of warm-water taxa in response to the changing climate; possibly as a consequence of low-latitude cradles being developed in, for instance, carbonate reef settings. Many clades were well established with especially the strophomenides beginning to outnumber the previously successful orthides, although this process had already begun, regionally, in the mid to late Darriwilian. At the same time, atrypoid and pentameroid clades also began to radiate in low-latitude faunas, anticipating their dominance in Silurian faunas. The Hirnantian was marked by severe extinctions particularly across orthide-strophomenide clades within the context of few, but well-defined, climatically controlled provincial belts. Supplementary material: The individual localities and a reference list for the data sources are provided at: http://www.geolsoc.org.uk/SUP18667

Enigmatic millimetre-scale micro-concretions with pseudocrystal faces and dominated by green clay minerals occur in unfossiliferous siliciclastic mudstone of the Lower Ordovician (479.0-466.0 Ma) Tonggao Formation, South China. The fossil-free mudstone unit is associated with local biodiversity decline. The mineralogy and mineral chemistry of these concretions were unknown previously, and this study comprises a preliminary investigation. The concretions are dominated by Fe-rich phyllosilicate minerals including glauconite and clinochlore, with minor quartz and traces of magnetite. The tex- tural relations between the micro-concretions and the surrounding matrix, and the preservation of original mudstone lamina- tions within the concretions, point to an origin during early diagenesis. The mineralogy and chemistry of these concretions are consistent with an origin in a restricted, hypersaline, relatively deep-water environment, in accordance with stratigraphi- cal and paleonotological data. These micro-concretions provide clues for a stressed environment with poor water circulation and anomalies of salinity and oxygen. Resume : Des micro-concretions enigmatiques d'echelle millimetrique, avec des faces pseudocristallines et dominees par des mineraux argileux verts, se trouvent dans des mudstones siliclastiques non fossiliferes de la Formation de Tonggao de l'Ordovicien inferieur (479,0-466,0 Ma), dans le sud de la Chine. L'unite mudstone sans fossile est associee a un declin de la biodiversite locale. La mineralogie et la chimie des mineraux de ces concretions n'etaient pas connues anterieurement et la presente etude comprend une investigation preliminaire. Les concretions sont dominees par des mineraux phyllosilicates riches en Fe, incluant la glauconite et le clinochlore, avec un peu de quartz et des traces de magnetite. Les relations textura- les entre les micro-concretions et la matrice environnante ainsi que la preservation des laminations originales du mudstone a l'interieur des concretions signalent une origine au debut d'une diagenese. La mineralogie et la chimie de ces concretions correspondent a une origine dans un environnement aqueux limite, hypersalin et relativement profond, ce qui concorde avec les donnees stratigraphiques et paleontologiques. Ces micro-concretions fournissent des indices qu'il s'agissait d'un environ- nement perturbe avec une mauvaise circulation d'eau et des teneurs anormales en sel et en oxygene. (Traduit par la Redaction)

Piperock, a kind of characteristic ichnofabrics in Phanerozoic, was thought to decline gradually from Cambrian to Ordovician. A new compilation on the occurrences of the Cambrian and Ordovician piperocks of China and the world shows that piperocks generally flourished in Cambrian and declined in Ordovician, but a sharp decrease occurred during Middle and Late Cambrian. The case-study on the piperocks from the Lower-Middle Ordovician Hongshiya Formation at Dabaochang of Qiaojia, northern Yunnan Province, Southwest China indicates that the forming and preservation of piperocks were controlled by the depositional environment, the intensities of predation, competition, bioturbation, and the contents of nutrition and oxygen in watermass. A careful study on the development of the Cambrian and Ordovician piperocks suggests that the decrease of nearshore siliciclastic sediments and the low contents of oxygen and nutrition in the watermass may have contributed to the decline of piperocks in Middle and Late Cambrian.

Abundant information on drilling predation upon fossil bivalves, gastropods, and brachiopods has been accumulated during the past several decades, but very little is known about the predation on marine, infaunal carnivorous scaphopods. A survey of over 440 specimens of the scaphopod Dentalium gracile collected from the Upper Cretaceous Millwood Member (Campanian) of the Pierre Shale at a site near Russell, Manitoba displays a drilling percentage of approximately 35%. This figure is higher than previously reported for the scaphopods of similar age elsewhere, but is comparable to or lower than that of the sub-Holocene (34–61%). The difference in drilling percentage among different collections may be taxon-related or affected by the composition and structure of the fossil community. Morphologically, the drill-holes, produced by predatory drilling, are beveled with a countersunk profile of clean sharp edges. The drill-hole inner margin is ovate whereas the outer margin is nearly circular. Among the Manitoba specimens, drill-holes seem to be more common on the lateral side. Presumably due to the lack of effective defense strategy, the prey effectiveness is low (∼3.2%). There is no correlation between drill-hole dimension and scaphopod prey size, indicating that predator size does not correlate with the prey size. Nearly 480 specimens of cephalopods, pelecypods, and gastropods were also collected from the same community. There were only a total of 16 drill-holes on this additional material. Over 400 specimens of the only naticid Euspira obliquata were recovered from the same site and are believed to be the predators of the scaphopods and other molluscs. The low percentage (∼2.5%) of drilling on the abundant gastropod E. obliquata may indicate mistaken or opportunistic attacks by the predator, or that the cannibalism is not common in this community.

A diverse brachiopod fauna from a relatively deep water carbonate facies of the Upper Ordovician Beiguoshan Formation (uppermost Caradoc – lower Ashgill, middle Katian) is characterized by small shells and contains the oldest known Dicoelosia and Epitomyonia, two diagnostic taxa of deep water brachiopod palaeocommunities during the Late Ordovician and Silurian. Three new species are recognized: Dicoelosia cordiformis sp. nov., Dicoelosia perbrevis sp. nov. and Epitomyonia fui sp. nov. These pioneer forms of the family Dicoelosiidae show a relatively high degree of morphological plasticity. The shells of Dicoelosia from the Beiguoshan Formation range from the typical slender-lobed form with a concavoconvex profile to the strongly equibiconvex, fat-lobed morphotype that was not known previously until the late Silurian. The Beiguoshan dicoelosiids point to an important attribute of the deep water brachiopods: small generalists with high morphological plasticity, which make them ideal candidates as progenitors for the evolution of shallow water brachiopod faunas in shelf and platform depositional environments.

The Hirnantia Fauna is a globally-represented, cool water brachiopod fauna that originated and flourished in shallow marine environments at intermediate to low latitudes after the first episode of the end-Ordovician mass extinction. It was well-developed, widely distributed, long in stratigraphical range, and complex in palaeoecological differentiation on the Upper Yangtze Platform of the South China paleoplate, where it has been extensively studied. Qualitative and quantitative analyses show that the FAD of the Hirnantia Fauna in South China becomes stratigraphically higher from nearshore to offshore localities on the Yangtze Platform, while the range of the fauna becomes shorter whereas the taxonomic diversity increases. Over its stratigraphical range the diversity of the Hirnantia Fauna at nearshore localities consistently decreases, but trends in the opposite direction at offshore, deeper water localities. The fauna was represented by different communities, subcommunities or associations with changing environmental factors (such as water depth and substrate) at different localities or horizons.

The Lower–Middle Ordovician Zitai Formation of the South China palaeoplate consists of a succession of purple red, nodular argillaceous limestones. Palaeogeographically, it is distributed along the southeastern margin of the Yangtze Platform, and is of late Floian to Dapingian age, correlative with the Dawan Formation of the Middle and Lower Yangtze Platform. In Shitai County, Anhui Province, East China, the Zitai Formation is rich in conodonts, enabling the recognition of four biozones based on first appearance data. Detailed palaeontological and biostratigraphical study of these conodonts reveals that the Ordovician conodont radiation in the Lower Yangtze Platform attained its first diversity peak low in the Oepikodus evae Biozone. This diversification is generally consistent with macroevolutionary trends of brachiopods of South China and graptolites of the Upper Yangtze Platform, but was earlier than that of trilobites and acritarchs of the same palaeoplate. Correlation with the sea-level curve for Sou...

The Ordovician radiation is among the major bioevents in earth history that have drawn great attention from geologists all over the world, and there have been hundreds of publications treating various aspects of this radiation. The study of the event involves a broad spectrum of research fields and concerns many scientific questions. Many opinions and hypotheses have been proposed on the triggering and controlling factors of the radiation. Herein the authors critically review the study of the Ordovician radiation and dissect four principal aspects by synthesizing the studies over the past years: (1) relationships of the radiation to fossil clades, ecotypes, and environments, such as water depth, latitude, and climate; (2) the ecological structure and food webs of Ordovician marine organisms; (3) fossil records and true biodiversities; and (4) likely triggering and controlling factors of the Ordovician radiation. The authors also recommend approaches to furthering studies of these aspects.

Located northwest of the Jiangshan-Shaoxing Fault Zone, western and northwestern Zhejiang and northeastern Jiangxi provinces experienced major changes in bio- and litho-facies and paleogeography through the Ordovician-Silurian transition (late Katian, Hirnantian, and early Rhuddanian), as manifested by stratigraphic, paleontologic and synecologic records. Three geographic units under consideration are South, Central, and North areas. The western margin of the South area was occupied by the Huaiyu Mountains, whereas the other two parts were covered by the Zhe-Gan Sea during late Katian (Late Ordovician) time. In the early stage of the Changwu Formation deposition (late Katian), the sea was deepening northeastward, but with shallower conditions in the southwest and deeper conditions in the northeast. During mid to late stages of Changwu Formation deposition (latest Katian), the sea became much shallower and the sea bottom was uplifted substantially, which occurred somewhat prior to the global trend. During the Hirnantian (latest Ordovician) and early Rhuddanian (earliest Silurian), the study region became a shallow bay under expansion of the Cathaysian Oldland. There occurred a major drop of sea level and great changes in benthic biota with the occurrence of many new immigrants through the Ordovician-Silurian transition, which are closely related to a unique regional orogeny. The Yangtze and Cathaysian blocks may have amalgamated within the South China Paleoplate during this interval to cause the continuous uplifting and northwestward expansion of the Cathaysian Oldland.

In the immediate aftermath of global extinctions, organisms were normally much smaller than those prior to these events. This ‘Lilliput Effect’ can be subdivided into two types: 1) a specific type, following the original definition of the effect which targets species-level taxa associated with inhospitable environments, and 2) a more general type, related to the reactions of higher-rank taxa above the species-level. The body sizes of brachiopods from South China through the Ordovician and Silurian transition (Late Katian, Hirnantian, and earliest Rhuddanian) are compared at generic, superfamilial, ordinal, and class levels. The results indicate that the body sizes of the taxa of lower rank (e.g. genus-level) are highly variable within these different intervals. The type of evidence for the Lilliput Effect through the end Ordovician mass extinction is thus quite different from that of the end Permian mass extinction probably reflecting differences in the intensity of these two major bioevents. However, the relationships between the contrasting trends in body-size change of some taxa of higher rank (e.g. at the ordinal-level) and the relative dominance of these taxa in the latest Ordovician and earliest Silurian suggest that the brachiopods of the two major Ordovician groups, the strophomenoids and orthoids, adopted different survival strategies during and immediately after the crisis from those of the pentamerides and rhynchonellides, that were common in Silurian assemblages.

High-resolution δ 13 C data of organic carbon from a continuous section of the Late Ordovician–Early Silurian reveal two positive δ 13 C excursions that are associated with the mass extinction in South China. The fi rst stratigraphic δ 34 S measurements on pyrite tied to well-established biostratigraphy indicate a large perturbation of the sulfur cycle, consistent with major sea-level changes related to the glaciation. The elevated δ 34 S values of pyrites and a large, short-lived negative δ 34 S excursion of ~20‰ associated with the decay of the glaciation suggest deep-water anoxia during the Hirnantian Stage, in contrast to the conventional view that the global oceans were oxygenated. We suggest that deep-water anoxia may have contributed to the Late Ordovician mass extinction in South China and possibly elsewhere.

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 cri...

Late Ordovician strophomenide brachiopods (superfamilies Strophomenoidea and Plectambonitoidea) from the upper Changwu Formation (mid Ashgill, late Katian) of Jianglutang, Chun’an County, western Zhejiang Province, consist of ten genera and 12 species. Five new species of three new genera are recognized: Chunanomena triporcata, Chunanomena sembellina, Cheramomena subsolana, Lateriseptomena modesta, and Lateriseptomena rugosa. The strophomenide brachiopods from the upper Katian strata described in this study and those from the border region of Zhejiang and Jiangxi provinces reported in previous work contain 16 strophomenoid and 12 plectambonitoid genera, and most of the strophomenoids are endemic to South China. Numerical analysis of well-documented late Katian strophomenide brachiopod faunas indicates a strong provincialism, characterized by the highly distinct North American province (Laurentia), the South China-Kazakhstan province, and the Avalonia-Baltica province (Wales, Belgium and Sweden). Surprisingly, the Girvan district of Scotland, which was a peri-Laurentian terrane during the Ordovician, contains a late Katian brachiopod fauna that is more closely related to the contemporaneous brachiopods of Avalonia-Baltica than to those of North America.

Stegerhynchus eoborealis sp. nov. from the Dalmanitina Beds (Hirnantian, uppermost Ordovician), Borenshult, Ostergotland, Sweden, is described in this study as the earliest known species of Stegerhynchus, within the family Rhynchotrematidae. This implies that Stegerhynchus originated during the latest Ordovician mass extinction event, and radiated in the Silurian Period to become one of most common rhynchonellide brachiopod genera in shallow tropical seas. In the early evolutionary stage of the genus, Stegerhynchus eoborealis already exhibited a notable reduction in the size of septalium compared to the closely related genus Rhynchotrema. Compared to its Silurian counterparts (e.g. Stegerhynchus borealis and S. peneborealis), Stegerhynchus eoborealis has some variable characters within single populations, particularly in the number of costae in the sulcus and on the fold, implying a morphological plasticity during the early evolutionary stage of the genus. During the Hirnantian mass extinction events, five rhynchonellide families were represented by small taxonomic groups, but only the Rhynchotrematidae and Trigonirhynchiidae diversified into many highly successful clades of rhynchonellides during the Silurian and Devonian. However, the Ancistrorhynchidae, Orthorhynchulidae, and Sphenotretidae largely disappeared by the end of the Ordovician, with a few species (such as Plectothyrella sp.) surviving into the earliest Silurian but shortly becoming extinct.

The continuous and richly fossiliferous Ordovician succession of China (particularly South China) comprises a heterogeneous suite of litho- and bio-facies, which has been a main focus of stratigraphical and palaeontological research in recent years. Among the seven GSSPs established in China, three are within the Ordovician System, and the GSSP of the Darriwilian Stage at Huangnitang, Changshan County, western Zhejiang Province, was the first “golden spike” in China and the first for the Ordovician System. A series of case studies have revealed that: (1) the Ordovician radiation of some fossil groups on the Upper Yangtze Platform (e.g., brachiopods and graptolites) reached their first α-diversity acme in the Didymograptellus eobifidus Biozone, four zones earlier than the global trend; (2) the β-diversity peak was attained 3–4 zones later than the α-diversity peak; (3) many brachiopod communities or faunas first occurred in the central part of the Upper Yangtze Platform and subsequently expanded to both more offshore and near-shore facies; (4) diachroneity existed in many aspects of the radiation. The end-Ordovician mass extinction was a severe event in South China. Two pulses of the extinction are recognized for a number of major fossil groups, some being most strongly affected during the first pulse whereas the others suffered during the second pulse. Macroevolution during the Ordovician–Silurian transition has been investigated in detail, and the role of the Lazarus effect has been found to be less important than previously believed.

The Ordovician radiation is the earliest and most important biodiversification event in the evolution of the Paleozoic Evolutionary Fauna (PEF), when the basic framework of PEF was established. The radiation underwent a gradual, protracted process spanning more than 40 million years and was marked by several diversity maxima of the PEF. Case studies conducted on the Upper Yangtze Platform (South China Palaeoplate) showed that the Ordovician radiation was characterized by drastic increases in α- and β-diversity in various groups of organisms. During the radiation, brachiopods, trilobites, and graptolites of the PEF became more diverse to dominate over the Cambrian Evolutionary Fauna (CEF) in all marine environments. At either global or regional scales, however, the Ordovician radiation was highly heterogeneous in time and space, and the rate and pattern of radiation exhibited by different major fossil groups were also variable.

The lower Meitan Formation (Floian, upper Lower Ordovician) at the Dajiaosi section, Zunyi District, northern Guizhou Province of South China, contains a moderately rich and diverse Sinorthis Fauna, with 22 species attributable separately to 15 families and seven orders of brachiopods. The fauna can be differentiated into three associations: the Paralenorthis serica, the Sinorthis typica, and the Tarfaya intercalare associations. These occupied a relatively wide palaeoecological range from lower BA3 to upper BA2 settings and from silty to clay substrate conditions. In South China, the Early Ordovician brachiopod radiation was marked by the diversification of the orthide-dominated Sinorthis Fauna, which first appeared and diversified in the middle part (relatively deep water) of the Upper Yangtze Platform in the Didymograptellus eobifidus Biozone, but rapidly declined in the succeeding Corymbograptus deflexus and basal Azygograptus suecicus biozones. During the latest Floian and early Dapingian, it expanded into comparatively shallow-water settings in the onshore direction (Changning area, southern Sichuan), offshore carbonate platform (Yichang, Hubei), and areas adjacent to the submergent Qianzhong Arch (such as the Zunyi area). The first appearance datum of the Sinorthis Fauna in shallower-water settings generally postdates that in the deeper-water environment in the central Upper Yangtze Platform, probably as a result of the fauna tracking a favoured BA3 setting during a gradual marine transgression.

Epipunctae, a new type of shell perforation, are well developed in typical taxa of the family Plaesiomyidae, a group of common orthide brachiopods from Laurentia and some other tropically located tectonic plates of Late Ordovician age. These minute, prominently elongate, tubular structures are similar to endopunctae in size and density, but differ in being oblique, intersecting the shell surface at a relatively low angle, and being confined largely to the outer portion of the shell wall. The tubules are similar in orientation to aditicules within the same shells but are much smaller and denser, usually aligned along fine growth lines and arranged in crude longitudinal columns. Exceptionally preserved phosphatic molds of bundled setal canals inside epipunctae and aditicules, described for the first time in this paper, are direct evidence that these two types of tubular structures of different sizes had the same function of housing sensory setae along the shell margin, but both the setae and the tu...

Sixteen species from the Lower Silurian (mostly Aeronian and rarely Telychian) of South China have been assigned previously to the brachiopod family Pentameridae, mainly to the genus Pentamerus. Study of intrapopulation variation suggests that 13 species from the upper Aeronian Lojoping Formation of the Yichang area, western Hubei Province, can be synonymized with Sulcipentamerus dorsoplanus, the most abundant pentamerid species of South China but confined to the Yichang area. The conspicuously high and obtuse ventral umbo, strong ventribiconvexity and a sulcate anterior commissure warrant genus status for Sulcipentamerus, independent of Pentamerus. True Pentamerus is rare in South China, known only from upper Aeronian rocks of Guizhou Province. Multivariate analyses indicate that Aeronian pentameride brachiopods of South China are characterized by a high level of provincialism, distinct from those of Laurentia. Also, the Aeronian pentamerides of South China or Laurentia have a low degree of similarity to those of Baltica, Siberia, Avalonia, Kazakhstan and other palaeotropically located plates or terranes. Besides South China and Laurentia, the pentamerides of other palaeotropical regions constituted a relatively tight cluster. This suggests that the palaeogeographical positions of South China and Laurentia only allowed limited or periodic faunal exchange of these generally large-shelled brachiopods adapted to living in relatively shallow, warm waters.

Systematic and statistical study of brachiopods from the Dawan Formation of the Yichang area, western Hubei Province, central China, indicates that, at generic and specific levels, the α diversity of Early to Mid Ordovician brachiopods in the study area reached the first acme at the beginning of the Corymbograptus deflexus Biozone, marked by the diverse Sinorthis typica Community. This diversity maximum of brachiopods occurred approximately one biozone later than that of the Upper Yangtze Platform. In taxonomic constituents, the main contributors to the diversity maximum were orthoids, which experienced a major increase in the number of genera in the upper Didymograptellus eobifidus Biozone and the lower C. deflexus Biozone. In brachiopod communities or associations, orthoids and pentameroids were the dominant elements. Analyses of Early–Mid Ordovician brachiopods from the Upper Yangtze Platform indicate that a high level of brachiopod community organization, with temporal and spatial stability, was achieved by the Sinorthis Community, which persisted from the D. eobifidus to the Azygograptus suecicus biozones (early Arenig). In the D. eobifidus Biozone, the Sinorthis Community thrived across a wide ecological range (BA3 to upper BA4) in the siliciclastic facies of the central part of the Upper Yangtze Platform and, by the latest D. eobifidus interval, invaded more carbonate-rich substrates in more offshore settings in the northeastern part of the platform (the present Yichang area).

Zhan R. & Harper, D.A.T. 2006 09 12: Biotic diachroneity during the OrdovicianRadiation: evidence from South China. Lethaia, Vol. 39, pp. 211 226. Oslo. ISSN 0024-1164.The Ordovician radiation was one of the most marked and sustained increases inPhanerozoic biodiversification; nevertheless it occurred against a background of minimalglobal climatic and environmental perturbations. Detailed investigations of theOrdovician successions on the Yangtze Platform of the South China palaeoplate indicatethat: (1) the brachiopod a- and b-diversity changes are diachronous; (2) macroevolu-tionary patterns were different across the South China palaeoplate, with the EarlyOrdovician brachiopod radiation first occurring in normal marine, shallow-waterenvironments and then moving gradually to both nearer-shore and offshore locations;(3) the main contributors to the initial Ordovician brachiopod radiation were theOrthida and Pentamerida; the typical Ordovician brachiopod fauna, dominated by theOrthida and Strophomenida, did not appear until the late Mid Ordovician (Undulo-graptus austrodentatus Biozone) when the Strophomenida apparently replaced thedominant position of the Pentamerida within the fauna; (4) different ecotypes (e.g.,sessile benthos, mobile benthos together with pelagic and planktonic organisms)demonstrate substantially different macroevolutionary patterns. The Ordovician bra-chiopod radiation of South China was apparently earlier than that suggested by globaltrends together with the data available from other palaeoplates or terranes, which may berelated to its unique palaeogeographic position (peri-Gondwanan terrane graduallymoving to equatorial latitudes). I Brachiopods, diachroneity, a-diversity, b-diversity,Ordovician radiation, South China.Renbin Zhan, State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute ofGeology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China,[rbzhan@nigpas.ac.cn]; David A. T. Harper, Geological Museum, Oster Voldgade 5 7,DK-1350 Copenhagen K, Denmark, [dharper@snm.ku.dk]; 14th November 2005, revised6th March 2006.

Lower to Middle Ordovician transitional strata (Acrograptus filiformis Biozone to Exigraptus clavus Biozone) of the Upper Yangtze Platform contain rich and diverse brachiopods, particularly in the Daguanshan Formation of Sichuan Province, the Meitan Formation of Guizhou Province, and the Yingpan Formation of Houping, Chongqing City. On the basis of multivariate analyses of 207 samples (about 15 000 brachiopod specimens) and 61 species from these strata, 23 brachiopod-dominated communities and associations are delineated through six graptolitic biozones (Acrograptusfiliformis to Exigraptus clavus biozones). The disparity of brachiopod associations or beta-diversity, as measured by the number of contemporaneous associations across variable ecological settings, was low in the A. filiformis Biozone and increased through the overlying graptolitic biozones to reach an acme in the Azygograptus suecisus and Expansograptus hirundo biozones, where as many as nine different brachiopod associations thrived in mid-shelf and also established in inner- and outer-shelf environments. Despite a sharp increase in a-diversity within the Didymograptellus eobifidus Biozone, beta-diversity remained relatively low, with only three distinct brachiopod communities. The diversity of brachiopod associations suffered an abrupt drop in the E. clavus Biozone, as a result of a major faunal turnover event, with only a single Metorthis Community present in an upper mid-shelf setting. Temporally, brachiopod associations appear to be most stable in mid-shelf settings, with the Paralenorthis, Sinorthis, Desmorthis, and Euorthisina communities having a relatively high a-diversity and lasting through two to five graptolitic biozones. In both species composition and temporal duration, the brachiopod associations at the shallow- (BA1 to upper BA2) and deep-shelf (BA4) settings are unstable, characterized by low a-diversity and rapid vertical changes. This study implies that, during the Early and Mid Ordovician, brachiopod communities already attained a high level of organization in mid-shelf settings, but remained poorly established in high-stress environments of very shallow or very deep shelf settings. Copyright (c) 2006 John Wiley & Sons, Ltd.

Earliest Rhuddanian (Silurian) brachiopods are recorded from the basal part of the Lower Llandovery Shiyang and Anji formations in western Zhejiang and northeastern Jiangxi provinces, East China. Associated graptolites including Normalograptus jerini indicate the lowest Rhuddanian Akidograptus ascensus Biozone. The surviving brachiopod fauna includes 19 genera dominated by orthids and strophomenids, whereas pentamerids and atrypids that inhabited mainly warmer water regimes, and were almost absent in the cool/cold Hirnantia Fauna, occur rarely in the studied fauna. Each family is represented by a single genus that seeded their recovery. The predominance of these long-ranging and widely distributed genera is one of major characters of the brachiopod survival in east China. From qualitative and quantitative analysis of faunal composition, diversity and abundance, with evidences from palaeoecology and palaeogeography, the Levenea qianbeiensis Association, Katastrophomena-Leptaena-Levenea Association, and Glyptorthis-Epitomyonia-Levenea Association are recognized and assigned to BA (Benthic Assemblage) 2, BA3, and an ecozone close to the BA3-4 boundary respectively. No Lazarus genera are recorded in this study. Skenidioides and Epitomyonia were chiefly regarded as deeper-water taxa in the Ordovician and Silurian, but are recorded from shallow-water in east China during the early Rhuddanian, indicating an ecologic experiment with these taxa migrating from deep into shallower, better-oxygenated sites at the crisis time and during the subsequent survival interval. This study further demonstrates that the brachiopod faunal turnover after the end-Ordovician extinctions may not have been completed until the late Rhuddanian in South China.

Reexamination of type and topotype material revealed the presence of well-developed ventral and dorsal interareas in two virgianid brachiopods from South China, Eoconchidium jiangshanensis Liang (in Liu et al., 1983) and Paraconchidium shiqianensis Rong, Xu, and Yang, 1974, of Late Ordovician and Early Silurian ages, respectively. A cladistic analysis of the common virgianid taxa, incorporating new data on the development of interareas, confirms Paraconchidium Rong, Xu, and Yang, 1974 as a valid genus (not a junior synonym of Pseudoconchidium Nikiforova and Sapelnikov, 1971) and warrants E. jiangshanensis as the type species of Deloprosopus new genus (not allied to either Eoconchidium or Tcherskidium as previously believed). Our preliminary survey on the suborder Pentameridina, based on available material or illustrations of well-preserved, disarticulated (typically silicified) valves, indicates that 1) the interareas are more commonly developed in the superfamily Pentameroidea than was reported previously, especially in the families Virgianidae and Subrianidae; 2) the ventral and dorsal interareas commonly do not occur as paired planar surfaces in the Pentameroidea, as they do in the superfamily Stricklandioidea; and 3) despite the common absence of a ventral interarea, the development of a sharply delimited dorsal interarea appears to be ubiquitous in the Pentameroidea and possibly in the suborder Pentameridina. In the currently adopted classification, the presence of matching ventral and dorsal interareas is treated as one of the diagnostic characters that separate Stricklandioidea from other superfamilies of the Pentameridina. The new data presented herein on the development of interareas imply the need to reevaluate the taxonomic and evolutionary significance of the ventral and dorsal interareas, pending a thorough survey on their distribution in the suborder Pentameridina.

Global analyses of 88 families and 284 genera of brachiopods from middle Ashgill, Late Ordovician, to early–middle Rhuddanian, Early Silurian, indicate that 18.6% and 12.5% of families and 51.0% and 41.3% of genera were eliminated in the first and second phases of the end-Ordovician mass extinction, respectively, with the total loss of 28.4% of families and 69.0% of genera in the crisis. New investigation demonstrates that brachiopods, at both generic and familial levels, suffered greater during the first phase than during the second phase. Four groups (victims, relicts, survivors, and new arrivals) are distinguished by their stratigraphical ranges. Generic survivors, occurring in the Kosov Province during the Hirnantian, can be split into three types with respect to their changing abundance: increasing, declining, and Lazarus taxa. Among the 88 genera that survived, numerous declining genera occurred in the Hirnantian: 16 Lazarus families and 18 Lazarus genera are provisionally known and may be regarded as end members of the declining type. Comparison of the abundance, population size, and distribution patterns of declining and Lazarus taxa shows important similarities between these two types which contribute to a better understanding of the nature of Lazarus taxa. In addition to these biological attributes, taphonomic failure and generally poor preservation, together with collecting bias and inadequate systematic data, are clearly involved. More collections will undoubtedly globally reduce the number of Lazarus taxa. A single, common refugium for end-Ordovician brachiopods probably did not exist; rather, these taxa used paleogeographically scattered locations in a range of environments for survival.

Restudy of two Early Silurian (Aeronian) stricklandioid species, Stricklandinia transversa Grabau, 1925 and Stricklandiella robusta Rong and Yang, 1981 from the Yichang area of South China, leads to the recognition of two new genera, Sinokulumbella and Sinostricklandiella. Two types of spondylia and four types of cardinalia are recognized for the Stricklandioidea and can be used for classification of the superfamily at the generic or even familial level. On the basis of these characters, five groups (typified by Stricklandia, Stricklandiella, Kulumbella, Microcardinalia, and Aenigmastrophia) can be recognized within the superfamily. Sinokulumbella n. gen., with a small, shallow, bowl-shaped spondylium and a pair of outer hinge plates that are discrete from the crura, is regarded to be affiliated with the Kulumbella group. Sinostricklandiella n. gen. has superimposed plicae and costae, although its internal structures are similar to the Stricklandiella group. A revised classification of stricklandioids implies that the Chinese pentamerides were characterized by strong provincialism during the early and middle Llandovery. This interpretation is supported further by the complete lack of true Stricklandia and Microcardinalia lineages in South China. Different stocks of stricklandioids in Baltica, Laurentia, South China, Siberia, and Kazakhstan may have experienced allopatric evolution, punctuated by several pulses of faunal migration or exchange between these paleoplates during the Llandovery. Evolution of the Stricklandia lineage is significantly different from that of Kulumbella and Sinokulumbella, particularly in that the outer plates disappeared earlier in the kulumbellids than in the stricklandiids.

A newly discovered Late Ordovician (early Ashgill) brachiopod assemblage from the Linhsiang Formation in the middle part of the Yangtze Platform, South China, bridges the paleobiogeographical gap between the early Ashgill Foliomena-bearing associations known previously from the upper and the lower parts of the Yangtze Platform. Characterized by minute shells in calcareous to siliciclastic mudstones, the fauna contains 13 brachiopod genera, of which two plectambonitoids are new: Hadroskolos and Jingshanella. Cluster and principal component analyses, based on 29 global occurrences of the Foliomena fauna in Laurentia, Avalonia, Kazakhstan, Baltica, Sardinia, Bohemia, Sibumasu, North China, and South China, revealed broad trends of spatial and temporal faunal differentiation in terms of taxonomic compositions. The analyses demonstrate for the first time that early Foliomena-bearing associations of Caradoc age occupied primarily deepwater (distal shelf) environments with a siliciclastic or calcareous mud substrate. The fauna attained its widest paleogeographical distribution and paleoecological range (midshelf to shelf margin settings) during the early Ashgill.

Affected by paleogeographic position, paleoclimatic condition and depositional environments, the increase of the Early-Mid Ordovician brachiopod diversity of South China commenced at the beginning of the Ordovician (early Tremadoc), accelerated from the Tetragraptus approximatus Biozone (base of Arenig), and reached its first acme in the Didymograptus eobifidus Biozone (mid early Arenig) when the number of brachiopod genera was over 7 times as great as that at the start of the Ordovician. This was the first radiation in the history of brachiopod macroevolution in South China, which occurred nearly 5 graptolitic biozones earlier than the global trend of the great Ordovician biodiversification (in the lower part of the Undulograptus austrodentatus Biozone). It is also characterized by (1) the origination or first occurrences of some major groups, such as the punctate dalmanelloids and the pseudopunctate strophomenoids including Plectambonitoidea (cardinal process simple or absent) and Strophomenoidea (cardinal process bilobed) in South China; (2) niche expansion, particularly in the first occupation of deeper water benthic regimes by the Euorthisina-Nocturnellia Association developed at Houping, Chengkou, northern Chongqing; and (3) the differentiation of brachiopod faunas under different environmental conditions. The gradual and increasing separation from Gondwana may have been one of the factors responsible for the radiation in South China.

Holorhynchus giganteus Kiaer, 1902, a common Late Ordovician (mid-Ashgill) pentameride brachiopod in the Baltic region, Kazakhstan, and southern Tien Shan, is documented for the first time from the Badanjilin Formation (mid-Ashgill) of western Inner Mongolia (Alxa block), North China. Serial sections of the Chinese material confirm the presence of a vestigial ventral median septum in the early growth stage of H. giganteus, but the septum becomes embedded in the secondary shell thickening at the adult growth stage. A survey of the type material from Norway and additional material from other regions indicates that the incipient ventral median septum is a much more commonly developed structure than was previously believed. The presence of a well-developed pseudodeltidium in the Tien Shan material of H. giganteus and the absence of such a structure in conspecific material from many other regions require a systematic revision of the generic group. Holorhynchus has rodlike crura (=brachial processes) t...

The Upper Ordovician (mid-Ashgill) Xiazhen and Changwu formations in the Zhejiang-Jiangxi border region of southeastern China contain an abundant and diverse suite of brachiopods. A cluster analysis of 59 key samples from 180 collections (over 23 000 specimens) allowed delineation of eight distinct brachiopod communities, herein named the Ectenoglossa minor Community, the Eospirifer praecursor Community, the Ovalospira dichotoma Community, the Antizygospira liquanensis – Sowerbyella sinensis Community, the Kassinella shiyangensis Community, the Altaethyrella zhejiangensis Community, the Tcherskidium jiangshanensis Community and the Foliomena folium Community. These communities occur predominantly in limey, clayey, to silty mudstone facies, similar to Ziegler's Early Silurian brachiopod communities in the Welsh Borderland. On the basis of lithological, faunal, and paleogeographical evidence, the eight communities are interpreted to have distributed along an onshore–offshore gradient from BA1 to BA6 settings on the northeastern margin of the Zhe-Gan Platform. In a broad paleo-embayment and mixed carbonate–siliciclastic shelf setting, the distribution of the eight brachiopod communities appears to have been controlled primarily by water depth because a given brachiopod community can be found in different lithofacies. Substrate types, however, must have played an important role, as some of the communities show preferred lithofacies. This study shows that the ecological zonation and community organization of the Late Ordovician pre-extinction brachiopods were similar to those of Early Silurian post-extinction brachiopods, although pronounced brachiopod provincialism during the Ashgill makes it difficult to directly correlate these communities with coeval brachiopod communities reported from other paleo-plates.

The Central Guizhou and Yichang Uplifts are present in central Guizhou to western Hubei. Biostratigraphic evidences from more than 20 sections in 14 counties of this region provide the data about the duration of these uplifts. Although this duration differs from locality to locality, it is mainly from Ashgillian to Rhuddanian. The uplifts result from a horizontally driven movement to the South China Paleoplate from an uncertain block in southeast. Global sea-level drop during the end of the Ordovician made the uplifts obvious, particularly the Central Guizhou Uplift. It might have emerged above sea level in the short interval between the Ordovician and Silurian.

Gnamptorhynchos, a rhynchonellid-like orthid brachiopod, evolved from Platystrophia in Maysvillian (early Ashgill) time, and survived both the end-Richmondian and end-Hirnantian episodes of the latest Ordovician mass extinction. The name of the type species of Gnamptorhynchos, G. inversum Jin, 1989, is rejected and replaced by Gnamptorhynchos globatum (Twenhofel, 1928), which is a senior synonym. Gnamptorhynchos manitobensis new species is described here from the Selkirk Member (Maysvillian) of the Red River Formation, southern Manitoba. The new species is characterized by a transversely extended, strongly biconvex to globular shell with prominent umbones, relatively numerous costae and a notothyrial cavity supported dorsally by a short median ridge. It constituted part of a Late Ordovician epicontinental fauna that once spread widely in shallow, equatorial seas of North America. The new species is a morphological intermediate between Platystrophia and Gnamptorhynchos, with Platystrophia-like int...

Gnamptorhynchos, a rhynchonellid-like orthid brachiopod, evolved from Platystrophia in Maysvillian (early Ashgill) time, and survived both the end-Richmondian and end-Hirnantian episodes of the latest Ordovician mass extinction. The name of the type species of Gnamptorhynchos, G. inversum Jin, 1989, is rejected and replaced by Gnamptorhynchos globatum (Twenhofel, 1928), which is a senior synonym. Gnamptorhynchos manitobensis new species is described here from the Selkirk Member (Maysvillian) of the Red River Formation, southern Manitoba. The new species is characterized by a transversely extended, strongly biconvex to globular shell with prominent umbones, relatively numerous costae and a notothyrial cavity supported dorsally by a short median ridge. It constituted part of a Late Ordovician epicontinental fauna that once spread widely in shallow, equatorial seas of North America. The new species is a morphological intermediate between Platystrophia and Gnamptorhynchos, with Platystrophia-like interareas, hingeline, and cardinal process, but Gnamptorhynchos-like shell posterior and notothyrial platform. Previously, Gnamptorhynchos was known from Hirnantian to mid-Aeronian rocks. The new species extends the lower range of the genus to Maysvillian.

Survivor, Lazarus and progenitor taxa are sources of biotic recovery following mass extinction. Investigations of the benthic brachiopods through the latest Ordovician mass extinction shows that progenitors developed many evolutionary novelties and successful surviving mechanisms. They are superior to survivors and Lazarus taxa in their ability to adapt to environmental changes. They are the primary source of macroevolution and the ancestors of a number of new taxa. Three kinds of progenitors are recognized based on the Ordovician-Silurian brachiopods from South China: survivor-progenitors, crisis-progenitors and Lazarus-progenitors; the last has the strongest ability to resist adverse environments, and is the most diverse and abundant.

A new halysitid species, Halysites gaojiangensis, is systematically described from the middle Rhuddanian (Llandovery, Silurian) Wulipo Bed at Gaojiang, Meitan County, northern Guizhou Province, Southwest China. It provides new data for further investigation of the tabulate faunal change across the Ordovician–Silurian transition. Halysites across the Ordovician–Silurian transition is reviewed and briefly discussed.

The Saucrorthis Fauna is a brachiopod-dominated shelly fauna developed in relatively deeper-water benthic regimes of a few peri-Gondwana terranes (e.g. northern Iran) and palaeoplates (South China and Sibumasu) during the Darriwilian (late Middle Ordovician). It has its longest geological range, widest palaeogeographic distribution, most abundant and diversified brachiopod taxonomic composition and most complex palaeoecological differentiation in South China, where it concentrated in the centre of the Upper Yangtze Platform. The generally low similarity coefficients (< 0.3) between any two representatives of the Saucrorthis Fauna firmly support a statement about the Great Ordovician Biodiversification Event (GOBE) that strong endemism or provincialism made a great contribution to the second pulse of the Ordovician brachiopod radiation in South China, which might also be true for the international trend.