Changhsingian brachiopod communities along a marine depth gradient in South China and their ecological significance in the end-Permian mass extinction

Diversity indices (dominance and evenness) and ecological spatial structure (lifestyles and relative abundances) are important features of Changhsingian brachiopod communities prior to the end-Permian mass extinction (EPME) and could predict temporal and spatial extinction patterns during the EPME. In South China, Changhsingian brachiopod communities show higher diversity than other contemporaneous brachiopod communities in the world and have been reported from a variety of sedimentary environments. In this paper, brachiopods from 18 sections in South China were selected to divide communities and compare their ecological structure. Based on the results of network analysis, cluster analysis and quantitative data from the selected sections, we show that Changhsingian brachiopod communities in South China can be categorized into three assemblages along a marine depth gradient: the Neochonetes–Fusichonetes–Paryphella Assemblage from the shallow-water clastic-rock facies, Spinomarginifera–Peltichia–Oldhamina Assemblage from the shallow-water carbonate platform facies and Fusichonetes–Crurithyris Assemblage from the deep-water siliciclastic intracontinental basin facies. Compared with communities from carbonate platform facies, the communities from siliciclastic facies were characterized by high dominance, low evenness and low lifestyle diversity, which might be important biotic factors leading to earlier extinctions. After the extinction began in all environments, the whole earliest Triassic brachiopod community was first dominated by Fusichonetes and then by Crurithyris. These patterns of domination and replacement could be explained by morphological and ecological advantages. The domination of these two genera, which were already adapted to the oxygen and food-limited deep-water habitat, indicates that the cooler deep-water environment might have been a relatively less stressed habitat after the beginning of the EPME. This suggests that global warming might be the main trigger among the previously proposed synergistic environmental stresses, while anoxia might not, at least for the beginning of EPME.     

Cryptospore Assemblages from the Ordovician/silurian Boundary in the Puna Region, North-west Argentina

Palynomorph assemblages have been recovered from deposits believed to straddle the Ordovician/Silurian boundary, from the upper member of the Salar del Rincón Formation, in the Puna region of north-west Argentina. The palynomorph assemblages are dominated by terrestrial cryptospores, but also contain marine elements (acritarchs, prasinophycean algae and chitinozoans). The cryptospore assemblages are similar in composition to those described from coeval deposits worldwide, suggesting that the producers were cosmopolitan and tolerated a wide range of climatic conditions. They are correlated with the Imperfectotriletes spp. Interval Biozone (sub-biozone α) of the Imperfectotriletes spp.- Laevolancis divellomedia Assemblage Biozone, indicating a Hirnantian (latest Ordovician) age. Acritarchs include late Ordovician species such as Eupoikilosusa striata and Villosacapsula cf. setosapellicula , that coexist with the Llandovery species Dactylofusa estillis. Based on lithological and palynological evidence, an early Llandovery, or a late Hirnantian (post-glacial) age is proposed for the basal part of the upper member of the Salar del Rincón Formation.     

Foliomena Fauna (Brachiopoda) from the Upper Ordovician of Sardinia

The late Ordovician brachiopod assemblage from Sardinia is one of the youngest members of the deep-water Foliomena fauna and is characterized by the following core taxa: Christiania , Cyclospira , Dedzetina and Foliomena . The fauna also contains Epitomyonia , Leangella , Glyptorthis and Skenidioides , which are more typical of shallower-water environments during the late Ordovician but occupied deeper-water niches during the Silurian following the termination of the Foliomena fauna. The suprafamilial placement of the family Chrustenoporidae is discussed and the new species Dedzetina serpaglii and Leangella ( Leangella ) fecunda are established. In common with many mid-Ashgill Foliomena faunas the Sardinian assemblage shows significant differences from other faunal developments of this type, reflecting its geographical position and shallower water conditions than those of the classic early Ashgill Foliomena faunas. The brachiopods occur with abundant trilobites belonging to a variant of the cyclopygid fauna. The faunas developed on part of a complex of microcontinents derived from peri-Gondwana during the Ordovician.     

Ecological disruption close to the Ordovician‐Silurian boundary

Following a long period of stability there was substantial ecological disruption in the late Ordovician, accompanied by widespread extinction. The ecological upheaval was spread over a period of time (1 My?) and varied in intensity between different groups.

The greatest change in brachiopod faunas occurred in the tropics, where there were widespread extinctions. In temperate regions the community structure, although disrupted, was not destroyed. The deep shelf associations (Benthic Assemblages 4–5) of the Rawtheyan persisted in an attenuated form into the Hirnantian when they were supplemented by genera of the ”Hirnantia fauna” and then became strongly re‐established in the Llandovery.

Trilobites, in contrast to brachiopods, suffered a worldwide extinction in the late Ashgill. Approximately 40% of trilobite genera became extinct at, or near, the Rawtheyan‐Hirnantian boundary.

Analysis of the distribution of Rawtheyan trilobite and brachiopod genera supports a correlation between geographical distribution and survival. Global range charts for brachiopod and trilobite genera, incorporating ecological and palaeogeographical data, have been compiled for this study.     

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

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

Palaeoenvironmental aspects of Late Ordovician Sericoidea shell concentrations in an impact crater,Tvären,Sweden

Frisk, Å.M. & Harper, D.A.T. 2010: Palaeoenvironmental aspects of Late Ordovician Sericoidea shell concentrations in an impact crater, Tvären, Sweden. Lethaia, Vol. 44, pp. 383–396. Numerous studies have reported the presence of the small, thin‐shelled cosmopolitan rhynchonelliformean Sericoidea as being environmentally controlled and, together with its close relatives, characteristic of deep‐water, distal, clastic Ordovician and Silurian settings. Assemblages of Sericoidea have been analysed from post‐impact strata in a newly formed Late Ordovician impact crater. In the crater succession, colonization of benthic faunas can be monitored through the post‐impact limestone, demonstrating a number of environmental preferences. Consequently, the crater, as a result of its restricted area, provides an experimental arena for faunal distributions to be correlated with specific environments. The continuous infilling of the crater following its formation reveals a transition from argillaceous mudstones to carbonates deposited in deeper‐water environments to shallower regimes. Rhynchonelliformean brachiopods inhabited the crater depression very late after the impact and are entirely represented by the genus Sericoidea, occurring abundantly in the upper third of the existing crater infill. The deep‐water regime that existed in the depression during the initial interval of crater formation had been substantially reduced. Clearly Sericoidea‐bearing associations associated with shaly substrates did not merely favour and occupy deep‐water environments as previously suggested. The unfavourable conditions triggered by the impact and the inhospitable aftermath allowed Sericoidea to exploit a less‐crowded ecospace. This reorganization, following the catastrophe, from a deep‐water related ecological niche to considerable shallower settings suggests that Sericoidea was a pioneer colonist displaying an opportunist r‐strategy. The shell beds analysed are related to shallower water and this may, moreover, help unravel the dilemma of the general absence of Sericoidea in the deeper‐water Foliomena fauna. □Dalby Limestone, impact crater, Late Ordovician, opportunists, Sericoidea, Tvären.     

Discovery of Anticostia uniformis from the Xiazhen Formation at Zhuzhai,South China and its stratigraphic implication

The Xiazhen Formation is an Upper Ordovician lithostratigraphic unit in the Jiangshan-Changshan-Yushan (JCY) area, which contains series of Late Ordovician reef successions. The reef successions of the Xiazhen Formation at Zhuzhai are critical for evaluation of the Late Ordovician marine diversity and palaeoecology. However, their age has long been uncertain and generally is regarded as of upper Katian, based on the occurrences of shelly fossils and correlation with the stratigraphic equivalent Changwu Formation. The newly discovered graptolite species Anticostia uniformis, in the Xiazhen Formation, together with the combined evidence of brachiopods and sedimentology, indicates an age range for the graptolite locality from the Dicellograptus complanatus Biozone to the Diceratograptus mirus Subzone of late Katian, but the graptolites do not rule out the possibility that it is Hirnantian.     

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 .     

Evolution,palaeoecology, and palaeobiogeography of the Late Ordovician–Early Silurian brachiopod Epitomyonia

Epitomyonia is characterized by various types of dorsal ridges, which may be transverse, longitudinal, or highly convoluted and probably served as skeletal supports for lophophores of various complexity. Multivariate analyses suggest that the Epitomyonia-bearing brachiopod associations lived in relatively shallow-water environment in the Late Ordovician, and inhabited mainly deep-water environments in the early Wenlock. The temporal and spatial change in the faunal distribution may be explained by three alternative scenarios: (1) Epitomyonia followed the broad evolutionary trend of the Palaeozoic Evolutionary Fauna to shift from shallow- to deeper-water settings over time; (2) the dicoelosiid communities could not compete with the large-shelled pentameride communities in continental shelf settings during the Early Silurian; or (3) only the shallow-water Epitomyonia died out in the Late Ordovician mass extinction event, whereas some poorly known deep-water Late Ordovician forms survived into the Early Silurian. Epitomyonia paucitropida n. sp. from the lower Whittaker Formation (late Katian) of the Mackenzie Mountains, northwestern Canada, is reported as the first known Ordovician species of Epitomyonia from the palaeocontinent of Laurentia, characterized by a small shell with weak, transverse dorsal ridges that are most primitive for the genus.     

Molecular,morphological, and ecological differences between the terrestrial and aquatic forms of Oxyrrhynchium speciosum (Brid.) Warnst. (Brachytheciaceae)

Oxyrrhynchium speciosum (Brid.) Warnst. is a relatively uncommon moss that can be found in wetland habitats across Europe and New Zealand. In this paper, we address the morphological, molecular and ecological relationships between the aquatic form of O. speciosum discovered in a deep, Chara-dominated lake (Budzis?awskie, NW Poland) and its terrestrial relatives. Morphological assessments revealed a considerable difference between terretrial and deep-water samples of O.speciosum. The aquatic form of O. speciosum was, on average, 12 times larger, with more branched stems (4–30 times) and longer leaves (1–2 times).

Investigation of ITS1-5·8S-ITS2-26S region sequences, however, revealed no distinction between the terrestrial and aquatic forms. An analysis of the ecological data suggests that the presence of O. speciosum in deep lake habitats is not incidental (as with many land species developing in aquatic environments). On the contrary, aquatic forms have a distinct ecological niche. Our results confirm that both forms of O. speciosum are closely related to O. hians (Hedw.) Loeske (a terrestrial species). The data indicate that O. speciosum is nested within a well-supported O. hians clade and separation of these species may be connected with polyploidization that occurred during their evolution.     

The Biology of Lophelia pertusa (Linnaeus 1758) and Other Deep-Water Reef-Forming Corals and Impacts from Human Activities.

Over the last twenty years, human exploitation has begun to have an impact in the deep sea, especially in the upper bathyal zone. This has mainly taken the form of deep-sea fishing but more recently oil exploration has extended beyond the continental shelf. Deep-water coral reefs occur in the upper bathyal zone throughout the world. These structures, however, are poorly studied with respect to their occurrence, biology and the diversity of the communities associated with them. In the North-East Atlantic the coral Lophelia pertusa has frequently been recorded. The present review examines the current knowledge on L. pertusa and discusses similarities between its biology and that of other deep-water, reef-forming, corals. It is concluded that L. pertusa is a reef-forming coral that has a highly diverse associated fauna. Associated diversity is compared with that of tropical shallow-water reefs. Such a highly diverse fauna may be shared with other deep-water, reef-forming, corals though as yet many of these are poorly studied. The main potential threats to L. pertusa in the North-East Atlantic are considered to be natural phenomena, such as slope failures and changes in ocean circulation and anthropogenic impacts such as deep-sea fishing and oil exploration. The existing and potential impacts of these activities on L. pertusa are discussed. Deep-sea fishing is also known to have had a significant impact on deep-water reefs in other parts of the world.     

Reef to basin sediment transport using Halimeda as a sediment tracer,Grand Cayman Island,West Indies

Fragments of the calcareous green alga Halimeda form a large part of the sediment in the fringing reef system and adjacent deep marine environments of Grand Cayman Island, West Indies. Nine species combine to form three depth-related assemblages that are characteristic of the major reef-related environments (lagoonpatch reef, reef terraces, and deep reef). These modern plant assemblages form the basis of the use of Halimeda as a sediment tracer. Halimeda-based tracer studies of Holocene sediments indicate that only sediments containing deep reef species of Halimeda are presently being transported through the reef system by sediment creep and being deposited at the juncture of the upper and lower island slope. Sediments containing shallow reef Halimeda are retained within the reef and lithified by marine carbonate cements. Tracer studies of Pleistocene sediment indicate large amounts of reef-derived carbonate sand containing deep water Halimeda were produced during interglacial high stands of sea level. Much of this material was removed by turbidity currents moving out of the reef system to the island slope down submarine channels perpendicular to the reef trend. These channels may still be identified on bathymetric profiles, but are no longer receiving coarse reef debris and are veneered with a blanket of pelagic carbonate mud.     

Can the Lilliput Effect be detected in the brachiopod faunas of South China following the terminal Ordovician mass extinction?

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.     

华南与缅甸奥陶纪末赫南特贝动物群中的无铰类腕足动物

在奥陶纪末生物大灭绝的两幕之间,海洋底域繁盛着赫南特贝腕足动物群（Hirnantia Fauna）。它数量丰富,分布广泛,历程短暂。以往国内外学者研究这个动物群时,常把有铰类腕足动物作为重点,而无铰类只被简单描述或列出名单,整体面貌不明。无铰类化石尽管材料有限、研究基础薄弱,但只就属的数目而言,占据了全球赫南特贝动物群总属数的近1/5,其群落、演化和环境意义不可小觑。文中专门记述上扬子区（华南古板块）观音桥层（赫南特早中期）与缅甸曼德勒地区（滇缅马苏古地体）Hwe Mawng紫色页岩段（赫南特中期）所产赫南特贝动物群的无铰类化石,计有3目、4超科[Linguloidea（舌形贝超科）、Discinoidea（平圆贝超科）、Craniopsoidea（似髑髅贝超科）和Cranioidea（髑髅贝超科）]的10属、12种,包括5个命名属种[Plectoglossa cf. davidsoni (Barrande),Schizotretinia cf. euxina (Havlí?ek), Pseudopholidops partibilis (Rong),Petrocrania cribrum (Temple),Xenocrania haimei (Reed)]和7个未定种（Trematis sp.,Paracraniops sp.,Acanthocrania sp.,Petrocrania? sp. 1,Petrocrania? sp. 2, Pseudolingula? sp. 和Orbiculoidea sp.）,其中,Pseudolingula? sp.和Orbiculoidea sp.两个未定名种因标本不佳,未正式描述。其中,以Pseudopholidops最为常见,其次是Xenocrania和Petrocrania。前两属是华南、滇缅马苏、波罗的卡、阿瓦隆尼亚、佩鲁尼卡等古板块或地体赫南特贝动物群的常见分子。它们已知限于南、北纬30°之间的低纬度地区,这可能与化石采集和研究程度有关,但更可能反映奥陶纪末全球气候仍存在分异现象。研究识别了奥陶纪晚期无铰类腕足动物7个目,可归为3个类群：灭绝目（仅神父贝目Paterinida）、消减目（多样性与丰度大幅衰减,尤其是三分贝目Trimerellida和乳孔贝目Acrotretida在Hirnantian销声匿迹）和延续目（适应能力强、忍耐阈值高,成功穿越大灾难的首幕）。这些分类单元对研究奥陶纪-志留纪交界期腕足动物群的多样性、群落生态、生物地理及宏演化等有重要意义。     

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

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

Island benthic assemblages: With examples from the Late Ordovician of Eastern Australia

The Benthic Assemblage (B.A.) concept, developed by A.J. Boucot two decades ago principally for continental marine margins, is extended to offshore island settings. Characteristics of modern island biotas, including ecological displacement, and the effects of r and K selection, can be identified in Late Ordovician volcanic islands of central New South Wales. Two variants of B.A.1 are represented, a quiet‐water lingulide biofacies, and a rough‐water rhynchonellide biofacies. The quiet‐water Eodinobolus biofacies occupied a B.A.1–2 position, onshore of and sheltered by a Tetradium wave‐baffle (B.A.2). The offshore shelfal high diversity strophomenide biofacies is equivalent to B.A.3. Remnants of periplatformal B.A.4–5 communities are recognised in allochthonous limestone breccias which were displaced downslope into graptolitic B.A.6 sediments. Steeper offshore gradients, typical of islands, laterally compress the B.A. profiles, and also contribute to downslope slumping. Ecological displacement in island environments results in extension of the habitat range of species into adjacent B.A.s. Reefs complicate the usual B.A. profile by introducing distinct sheltered and turbulent water environments. These characteristics may have applicability in interpretation of islands throughout the Palaeozoic record.     

Conodonts from the upper Ordovician siliceous rocks of Central Kazakhstan

Several conodont localities of the upper Sandbian Stage are known in siliceous deposits of Central Kazakhstan. All of them produced similar assemblages overwhelmingly dominated by Periodon grandis with insignificant admixture of Scabbardella altipes, Hamarodus europaeus, Pygodus anserinus, Protopanderodus sp., and Drepanodus sp. The main feature of this fauna is in the co-occurrence of H. europaeus and P. grandis, forms characteristic for deep-water facies at shelf or microcontinents margins of temperate and warm-water paleobiogeographic provinces. The Ordovician paleo-oceanic basin of Kazakhstan and southern Urals were parts of the uniform biogeographic area as indicated by similarity of Ordovician conodont assemblages in siliceous deposits of these regions.     

The trace fossil Sinusichnus from the Upper Ordovician of Anticosti Island,eastern Canada

Branched, sinusoidal burrows assigned to Sinusichnus cf. seilacheri Knaust et al. 2016; occur in Upper Ordovician (Hirnantian) carbonates on Anticosti Island. These are abundant and monospecific at the base of the Oncolite Bed of the uppermost Ellis Bay Formation, marking a regional discontinuity surface during the multiphase Ordovician mass-extinction event. In contrast to Mesozoic and Cenozoic records of Sinusichnus, commonly attributed to the activity of decapod and isopod crustaceans, the burrow systems from Anticosti Island were probably produced by other arthropods of unknown affinity. A combined dwelling, locomotion and feeding behaviour of their trace maker is assumed. This is the oldest record of the ichnogenus Sinusichnus, previously only known since the Triassic, and its first evidence from North America.     

Restudy of some Ordovician–Silurian boundary graptolites from Anticosti Island, Canada, and their biostratigraphic significance

The Ellis Bay Formation on Anticosti Island has long been recognized for the biostratigraphic importance of its latest Ordovician conodont, palynomorph, and shelly fossil assemblages. However, a sparse record of graptolites has made it difficult to correlate these assemblages with the graptolite biozonation. Restudy of some previously described and examination of newly collected normalograptid specimens from the Ellis Bay and lower Becscie formations, however, shows that biostratigraphically important taxa are present and these species provide important constraints on the age of the strata. Normalograptus parvulus and N. minor have been identified in the upper half of the Ellis Bay Formation, suggesting that these strata are late Hirnantian in age ( N. persculptus Biozone). Normalograptus imperfectus and Normalograptus sp. aff. N. acceptus occur in the basal beds of the Becscie Formation, indicating that these strata are earliest Silurian. These graptolite data support the hypothesis that the positive carbon isotope excursion seen in the uppermost strata of the Ellis Bay Formation is isochronous with that seen in the Hirnantian strata at Dob's Linn, Scotland, and does not span much of the lower to mid-Hirnantian, as is the case in Arctic Canada and Nevada, USA.     Anticosti Island , Hirnantian, graptolites, Ordovician, Silurian.