Middle–Late Ordovician brachiopods from Ningnan,southern Sichuan Province,Southwest China: Implications for macroevolution and palaeogeography

Middle to Late Ordovician brachiopods from the Huadan Formation (upper Darriwilian–Sandbian) of Ningnan County, southern Sichuan Province, are systematically documented here for the first time. The locality belongs to the western margin of the Upper Yangtze Platform, South China palaeoplate, and the brachiopod fauna includes one new genus and three new species as well as five other constituents: Hingganoleptaena sp., Acculina zhongliangziensis n. sp., Ningnanmena longisepta n. gen. n. sp., Kassinella (Trimurellina) minuta n. sp., Lepidorthis typicalis Wang, 1955, Protoskenidioides weixinensis Zhan and Jin, 2005, Porambonites transversus Xu, Rong and Liu, 1974, and Psilocamerella sp. Taxonomically it is a typical representative of a Middle to early Late Ordovician brachiopod fauna, and, together with some other evidence from other fossil groups like trilobites, conodonts, chitinozoans, a late Darriwilian–Sandbian age could be inferred for the horizon yielding this fauna. According to the richness of each constituent, this fauna is suggested to be called the Acculina-Ningnanmena fauna (ANF). Numerical palaeogeographical analysis shows that two broad palaeobiogeographic provinces could be recognized during this particular time interval, and, although the ANF is grouped into the South China cluster, it shares very little similarity with other representatives of that group except for two cosmopolitans. It further confirms that the Great Ordovician Biodiversification Event (GOBE), in other words the Ordovician radiation, was actually manifested by the strong localization of major marine organisms such as brachiopods, trilobites, graptolites, etc.     

安徽石台奥陶纪弗洛期-大坪期牙形刺多样性的演变

基于安徽石台栗阳柳树亭下、中奥陶统紫台组牙形刺的系统研究,对该地区早奥陶世弗洛期一中奥陶世大坪期牙形刺多样性演变作了初步分析,发现牙形刺的简单分异度在弗洛晚期Oepikodusevae带达到峰值,有17属20种。之后,牙形刺分异度呈下降趋势。演变的总体趋势与湖北宜昌地区同期牙形刺的研究结果一致,其峰值出现的时间（Oepikodusevae带,相当于笔石Didymograptellus eobifidus带上部,Corymbograptusdeflexus带,及Azygograptussuecicus带下部）与华南上扬子区腕足动物宏演化趋势（首次峰值出现在Didymograptelluseob—ifidus带）大体相似,但比三叶虫多样性演变在奥陶纪的首次峰值（出现在Exigraptus clavus带）早,比扬子台地笔石多样性演变的首次峰值（在Acrograptusfiliformis带和Didymograptelluseobifidus带下部）晚。分析显示,该地区早、中奥陶世牙形刺多样性演变与海平面变化存在密切关联。     

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.     

SILICIFIED RHYNCHONELLIFORM BRACHIOPODS FROM THE KUNIUTAN FORMATION (DARRIWILIAN: MIDDLE ORDOVICIAN), GUIYANG, SOUTH CHINA

Abstract: A silicified brachiopod fauna from the Middle Ordovician Kuniutan Formation (lower–middle Darriwilian, i.e. uppermost Arenig to lower Llanvirn) at Wudang, Guiyang, central Guizhou, South China, includes abundant specimens of Yangtzeella, Orthambonites and Leptellina together with common Parisorthis, Saucrorthis, rare Anomalorthis?, Hemipronites?, Leptestia? and, significantly, Aporthophyla; associated are rare trilobites, gastropods, crinoids and nautiloids. The Yangtzeella kueiyangensis‐Orthambonites delicata Association is defined for this shallow‐water, Benthic Assemblage 3, association. This first record of Aporthophyla in South China may indicate a link between South China and the Toquima‐Table Head Province, where the Aporthophyla fauna is more typically developed. However, this association is significantly different from the latter in having three endemic genera, Yangtzeella, Saucrorthis and Parisorthis, rare numbers of Aporthophyla and Anomalorthis?, and the absence of many other typical elements of the Aporthophyla fauna. The occurrence of Orthambonites, Hemipronites? and Leptestia? suggests some relationships between South China and the Baltic Platform during the Mid Ordovician. The various brachiopod associations bearing Aporthophyla may be quite different in nature, composition and diversity, and may possess different background palaeobiogeographical signatures. The assemblages containing Aporthophyla in South China, Qaidam, Malaysia, Australia and possibly Tibet are clearly different biogeographically from those associated with the Toquima‐Table Head and the Celtic provinces. Two new species, Aporthophyla sinensis sp. nov. and Leptellina orientalis sp. nov. are described.     

贵州安顺募役剖面晚二叠世腕足动物群落及其生态特征*

本文对贵州募役剖面长兴期腕足动物群进行了系统分类鉴定和古生态分析,发现其与前人所报道的华南同时期腕足动物群存在着显著的差别,以Spiriferellina为绝对优势属,包含少量华南长兴期的常见分子,如Fusichonetes、Araxathyris和Peltichia等。募役剖面的腕足类化石主要产自碎屑岩层位,与华南同期碎屑岩相剖面的腕足动物群落相比,该动物群中的优势分子个体较大、壳体较厚且发育较粗大的壳疹,这可能与长兴期募役剖面动物群所处的特殊栖息地环境(浅水碳酸盐岩台地与深水硅质碎屑岩盆地之间的过渡地带)有关。二叠纪末期大规模的火山作用导致水体中碎屑物质含量增加,影响腕足类的滤食效率,而募役剖面清澈、水动力弱的水体环境,为滤食性的腕足动物提供了有利的环境条件。最后,通过与华南长兴期不同沉积相区腕足动物群落(六枝剖面、稻堆山剖面、中寨剖面、新民剖面、马家山剖面和仁村坪剖面群落)的对比分析,发现募役剖面腕足动物群与毗邻的六枝剖面腕足动物群在属级组成上的相似度较高,且过渡岩相栖息地环境下的腕足动物群在生物灭绝事件前也呈现出高优势度、低均匀度的群落结构特征,指示海洋底栖生物群落已经先于二叠纪末期生物集群灭绝事件出现了早期危机信号。     

An upper Kungurian/lower Guadalupian (Permian) brachiopod fauna from the South Qiangtang Block in Tibet and its palaeobiogeographical implications

A brachiopod fauna including 15 species belonging to 14 genera is described from a thin carbonate succession of the Lugu Formation at the Anmu section in the northern part of the South Qiangtang Block, which is about 15 km south to the Longmu Co-Shuanghu Suture. The brachiopod fauna is late Kungurian or early Guadalupian in age in terms of the presence of the brachiopod species Vediproductus punctatiformis, “Cryptospirifer” omeishanensis, and Paraplicatifera regularis. This is also generally consistent with the age indicated by the fusulines (e.g., Cancellina, Chusenella, Monodiexodina, Nankinella, Neofusulinella, Pseudofusulina, and Sphaerulina) from this formation at the nearby Mari and Duoma A (DMA) sections. Palaeobiogeographically, the brachiopod fauna exhibits a typical Cathaysian affinity and all the species are reported commonly from the equivalents in South China and Cimmerian blocks, which strongly suggest that the South Qiangtang Block had drifted to a position in the warm-water or temperate regime close to South China.     

湖北宜昌陈家河下、中奥陶统大湾组腕足动物多样性演变

在对宜昌陈家河大湾组(Arenig期)腕足动物化石进行逐层统计和详细研究的基础上,以笔石带为单位讨论腕足动物分类单元多样性的变化,发现该地区腕足动物属级分异度在Corymbograptellus deflexus带达到首次峰值,比上扬子区处于正常浅海底域地区(如贵州桐梓、沿河一带)的腕足动物宏演化趋势(在Didymograptelluseobifidus带达到首次峰值)晚了一个笔石带的时限,辐射主要缘于Sinorthis群落在本地区的出现和极度繁盛。华南早、中奥陶世腕足动物的辐射中心在相对近岸的上扬子区(特别是处于正常浅海底域的贵州桐梓一带)。     

The latest Ordovician Hirnantia Fauna (Brachiopoda) in time and space

The diachronous temporal and spatial distribution of the Hirnantia brachiopod fauna and the complicated pattern of terminal Ordovician events are documented through biostratigraphical analysis of the Ordovician-Silurian boundary strata in S China, Sibumasu, Xizang and elsewhere. The duration of these events (longer than the half Myr derived from isotopic excursions) indicates that they were not abrupt and instantaneous. The presence of some core taxa of the Hirnantia fauna in the upper P. pacificus Biozone (known from their earliest occurrence in China) signals the start of increased water ventilation due to the invasion of cool water across the Yangtze Basin. Low- and higher-diversity Hirnantia faunas related to onshore, shallow-water and to offshore, deeper-water environments, respectively, developed first in the basal and upper N. extraordinarius-N. ojsuensis Biozone. Disappearance of most of the fauna in the early N. persculptus Biozone suggests that the glacial maximum started to decline. The presence of the Hirnantia fauna in the upper N. persculptus to the lower P. acuminatus biozones indicates the continuation of cool water environments in some places. The diachronous disappearance of deteriorating environments (earlier in later Hirnantian and finally in the early Rhuddanian) is associated with geographical heterogeneity. Occurrences of atrypids, pentamerids and spiriferids along with key elements of the Hirnantia fauna in N Guizhou provide a link between the Late Ordovician radiation and Early Silurian recovery of these major brachiopod groups.     

STROPHOMENIDE BRACHIOPODS FROM THE CHANGWU FORMATION (LATE KATIAN, LATE ORDOVICIAN) OF CHUN'AN, WESTERN ZHEJIANG, SOUTH-EAST CHINA

Abstract:  Late Ordovician strophomenide brachiopods (superfamilies Strophomenoidea and Plectambonitoidea) from the upper Changwu Formation (mid Ashgill, late Katian) of Jianglütang, 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.     

The transition from planktotrophy to lecithotrophy in larvae of Lower Palaeozoic Rhynchonelliform brachiopods

Criteria are established for defining the presence of protegula formed on embryonic or larval mantle in representative genera of Lower Palaeozoic Obolellata, Strophomenata and Rhynchonellata. Width was used to define protegular type. Taxa with only an embryonic protegulum are inferred to have had lecithotrophic larvae while taxa with a larval protegulum or an embryonic protegulum surrounded by a larval protegulum are inferred to have had planktotrophic larvae. All or most of the taxa examined in the Obolellata, the Strophomenata and the orders Protorothida and Orthida in the Rhynchonellata had planktotrophic larvae. In the Pentamerida a minority of genera had only a larval, or an embryonic and a larval protegulum while a majority had protegular widths indicating lecithotrophy. In the orders Rhynchonellida, Atrypida, Athyrida and Spiriferida derived from the Pentamerida (with the exception of one species in the Atrypida) a number of the genera had protegular widths indicating lecithotrophy. It is suggested that the onset of lecithotrophy in the Pentamerida was associated with a developmental innovation in which the mantle lobe of the larva was reflected over the apical lobe during the process of metamorphosis. This evolutionary innovation probably occurred during the late Cambrian or early Ordovician and was subsequently inherited during the process of cladogenesis.     

The oldest known bryozoan: Prophyllodictya (Cryptostomata) from the lower Tremadocian (Lower Ordovician) of Liujiachang,south‐western Hubei,central China

A new cryptostome bryozoan, Prophyllodictya simplex sp. nov., is described from the Nantzinkuan Formation (Lower Ordovician, lower Tremadoc) of Liujiachang, central China. This antedates the previously oldest known bryozoan by several million years. Colony morphology and the phylogenetic position of Prophyllodictya within Cryptostomata are explored. Phylogenetic analysis suggests that Cryptostomata (except Prophyllodictya) can be divided into two major groups, and that Prophyllodictya occupies a basal position in the cryptostome tree, which accords with its simple morphology and antiquity. A close relationship is evident between bryozoans from the South China palaeoplate and those from Baltica.     

Aspects of recent advances in the Ordovician stratigraphy and palaeontology of China

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.     

Latest Guadalupian brachiopods from the Guadalupian/Lopingian boundary GSSP section at Penglaitan in Laibin, Guangxi, South China and implications for the timing of the pre-Lopingian crisis

A brachiopod fauna comprising nine species in eight genera from three closely spaced stratigraphic horizons of the same stratigraphic section is described for the first time from the Laibin Limestone in the uppermost part of the Maokou Formation in the Guadalupian/Lopingian (G/L) GSSP section at Penglaitan, Guangxi Autonomous Region, South China. The brachiopod assemblages are bracketed between two conodont zones: Jinogondolella xuanhanensis Zone below and Jinogondolella granti Zone above and, therefore, they can be safely assigned to the latest Capitanian in age. However, all but one of the nine brachiopod species from the Laibin Limestone carry strong early Lopingian (Wuchiapingian) aspect. Thus, the discovery of this brachiopod fauna not only suggests that some Lopingian brachiopod species had already appeared in the late Guadalupian (Capitanian); more importantly, it has also highlighted the fact that both the previously noted pre-Lopingian life crisis (or end-Guadalupian or Middle Permian mass extinction) and Lopingian recovery/radiation actually occurred in late Capitanian times, sometime before the G/L chronostratigraphic boundary. So far, the Penglaitan GSSP section provides the highest-resolution disappearance patterns of different fossil groups around the G/L boundary.     

简论相似性测度的选择——以奥陶纪末大灭绝后全球腕足动物古地理为例

灭绝事件对古生物地理格局的影响已引起关注,近期研究表明奥陶纪末大灭绝事件后多样性显著高于传统认识,而全球该时期腕足动物的古生物地理分布情况尚未见报道。本文基于已发表的和最新的资料及所掌握新数据的整理,建立全球腕足动物志留纪初鲁丹(Rhuddanian)早期（残存期）13个产地72属137个出现信息(occurrence...     

Systematic position and composition of the superfamily Notanoplioidea Gill, 1969 (Brachiopoda, Articulata)

The systematic position of several genera of articulate brachiopods grouped around Notanoplia Gill is controversial. Notanoplia was established in 1950 within the order Chonetida; this genus and several externally similar genera, which were established later, were referred by different authors to the orders Orthida, Strophomenida, Chonetida, Atrypida, Spiriferida, or considered as incertae sedis. The morphological analysis of the shell of Notanoplia and the related genera Boucotia and Costanoplia supports the placement of these genera in a separate superfamily, Notanoplioidea, within the order Strophomenida. Other genera similar in the shell outline and shape to Notanoplia and conditionally considered as related to it probably belong to other strophomenid superfamilies or other orders.     

Euryhalinity of Palaeozoic articulate brachiopods

OMonotypic and very low diversity virgianid shell beds from the Upper Ordovician to Lower Silurian dolomites of North Greenland were formed in marginal marine quiet-water hypersaline environments. In the light of this evidence the salinity tolerances of other Palaeozoic articulate brachiopods is evaluated. There are only a small number of species apparently invading hypersaline or brackish environments, but it is significant considering that previously all articulate brachiopods were thought to be fully marine. Two types of occurrence are noted, those species specifically related to marginally marine environments, disappearing with the introduction of fully marine faunas, and the majority of species which extend their normal marine range into marginal conditions. No brachiopod species appears to have invaded very hypersaline or truly brackish conditions. No single group of articulate brachiopods specifically specialised in colonising marginal marine environments, apart from possibly the virgianid pentamerids. Palaeozoic, Upper Ordovician, Lower Silurian, Brachiopoda. Pentamerida, Virgianidae, Greenland, palaeoecology. hypersaline environments, brackish environments .     

浙江淳安安吉组底部的几丁虫

浙江淳安县大坑坞剖面安吉组底部化石丰富,其中腕足动物组合属于BA3中上部,指示正常浅海底域环境。作者通过分析该层位腕足化石标本的围岩发现一个低分异度的几丁虫组合,包含3属4种,即Ancyrochitina sp.,Belonechitina cf.postrobusta sensu Butcher,Belonechitina sp.,Spinachitina verniersi。该组合具有强烈的奥陶-志留纪过渡色彩,其时代可定为奥陶纪末期至志留纪初期。这是首次在该地区获得奥陶-志留纪过渡地层的几丁虫,有助于开展国际奥陶-志留系界线精细划分对比研究。文中描述了Belonechinacf.postrobusta和Spinachiti-naverniersi。     

An ecologically mixed brachiopod fauna from Changhsingian deep-water basin of South China: consequence of end-Permian global warming

The Late Permian Shaiwa Group of the Ziyun area of Guizhou, South China is a deep-water facies succession characterized by deep-water assemblages of pelagic radiolarians, foraminifers, bivalves, ammonoids and brachiopods. Here we report 20 brachiopod species in 18 genera from the uppermost Shaiwa Group. This brachiopod fauna is latest Changhsingian in age and dominated by productides. The palaeoecologic and taphonomic analysis reveals that the brachiopod fauna is preserved in situ. The attachment modes and substratum preference demonstrate that the Shaiwa brachiopod fauna comprises admixed elements of deep-water and shallow-water assemblages. The presence of the shallow-water brachiopods in the Shaiwa faunas indicates the involuntary settlement of shallow-water brachiopods. The stressed ecologic pressure, triggered by warming surface waters, restricted ecospace and short food sources, may have forced some shallow-water elements to move to hospitable deep-water settings and others to modify their habiting behaviours and exploit new ecospace in deep-water environments. We infer that the end-Permian global warming and subsequent transgression event may have accounted for the stressed environmental pressure in the shallow-water communities prior to the end-Permian mass extinction.     

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.     

WUCHIAPINGIAN (LOPINGIAN, LATE PERMIAN) BRACHIOPODS FROM THE EPISKOPI FORMATION OF HYDRA ISLAND, GREECE

Abstract:  Twenty-three species of silicified brachiopods are described from four samples in the middle and upper parts of the Episkopi Formation from Hydra Island, Greece. These brachiopods are newly recorded from the region and together with previously described brachiopods from the same localities constitute the most diverse Lopingian (Late Permian) brachiopod fauna reported in southern Europe. The brachiopod fauna is Wuchiapingian as indicated by the associated conodonts. The fauna from Hydra exhibits strong palaeobiogeographical links with the faunas from South China. In addition, palaeobiogeographical affinities with the faunas of Thailand and the northern peri-Gondwanan region are also present, which implies a peri-Gondwanan origin for Hydra. Palaeoecologically, the brachiopod assemblage from sample EP in the middle part of the Episkopi Formation is dominated by pedically-attached and cementing genera and reflects moderate energy conditions above storm wave base and an abundance of hard substrates provided by sponges in the biohermal habitat. By contrast, the brachiopod assemblage in the other three samples from the upper part of the Episkopi Formation is dominated by spinose genera with a free-resting life habit, suggesting soft substrates in a quiet water environment below storm wave base on the outer part of the shelf. New taxa are Petinospiriferina gen. nov., Hustedia episkopiensis sp. nov., Waterhouseiella hydraensis sp. nov. and Xenosaria tenuis sp. nov.