Reappraisal of some Upper Devonian (Famennian) spiriferide brachiopods from the Band-e Bayan Domain (Afghanistan)

The upper and uppermost Famennian succession (Upper Devonian) of the Band-e Bayan Domain (also known as the Axial Zone), a Cimmerian terrane of western Afghanistan (northern margin of Gondwana), yielded diverse brachiopod faunas that are dominated by the spiriferides. Among the latter, six species belonging to four genera, previously erected by D. Brice in the seventies and also known from Iran, are reappraised on the basis of their type material and re-illustrated. The genera Enchondrospirifer and Eobrachythyris are transferred to the subfamilies Cyrtospiriferinae (Cyrtospiriferoidea) and Prospirinae (Spiriferoidea), respectively, whereas the ornament and micro-ornament of both Afghan species of Dichospirifer are documented. The genus Ghorispirifer nov. gen. (Cyrtiopsinae) is erected with Cyrtiopsis graciosa chakhaensis Brice as type species, and includes a second species, namely Cyrtiopsis lapparenti Brice. The new genus is characterized notably by the strong development of ventral apical callosities. Upper Devonian (Frasnian?) brachiopods from the Band-e Bayan Domain described by Maillieux in the first half of the 20th century are revised and photographically illustrated for the first time. The brachiopod generic diversity recognized in Afghanistan (Band-e Bayan Domain, Central Mountains) prior to the Hangenberg Crisis is very high in some sections of the Band-e Bayan Domain (Rhuk pass section, and to a lesser extent at Robat-e-Pai), as also observed in contemporaneous successions of South China and contrary to what is reported in Western Europe (Franco-Belgian Basin for instance).     

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.     

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。     

Microconchid tubeworms across the upper Frasnian – lower Famennian interval in the Central Devonian Field,Russia

Abstract: Microconchid tubeworms (Tentaculita) encrusting brachiopod shells have been investigated from the upper Frasnian – lower Famennian (Upper Devonian) deposits of the Central Devonian Field, Russia. The condition of microconchids and associated encrusting taxa is reported for the first time from the early Famennian recovery interval (crepida Chron) following the Frasnian–Famennian mass extinction. Two species, one new (Palaeoconchus variabilis sp. nov.) and the second one in open nomenclature (Palaeoconchus sp.), are described. Compared to lower Famennian specimens, they seem to be preferentially grouped on the anterior parts of the brachiopod host shells, which are interpreted as the most suitable sites away from the sea‐bottom and sediment. During the late Frasnian (Late rhenana Chron), microconchids, outnumbered by cornulitids and as abundant as foraminifers, were also associated with trepostome bryozoans, tabulates, rugose corals and various problematic encrusters. During the early Famennian recovery interval encompassing the crepida Chron, microconchids greatly outnumbered all associated encrusters, including the previously dominant cornulitids, while foraminifers, tabulates and rugose corals vanished. Early Famennian microconchids, represented by the single, albeit very abundant, species Palaeoconchus variabilis sp. nov., were opportunists that rapidly colonised the environment during the ongoing transgression following the regression‐driven biotic crisis in the area of the Central Devonian Field. In comparison to their late Frasnian predecessors and even other Middle Devonian specimens, no size reduction (the so‐called Lilliput effect) of early Famennian microconchid tubes was observed. It is probable that microconchids either rapidly attained their ‘normal’ sizes or they did not suffer any dwarfism following the Frasnian–Famennian event.     

Biostratigraphical value of some Strunian (Devonian, uppermost Famennian) Productidina, Rhynchonellida, Spiriferida, Spiriferinida brachiopods

The space of Devonian time considered in this paper corresponds to the uppermost part of the Famennian or the latest Famennian as a fourfold subdivision of the stage, usually called by many authors Strunian in neritic facies and Wocklumeria Stufe or Wocklumian in pelagic facies. Here, we examine the biostratigraphical value of certain brachiopod genera and species as bio-markers of the uppermost Famennian throughout the world (Europe, the former USSR, Middle East, Asia, North America, North Africa, and Australia). We have focused our study for species which stratigraphical range has been firmly established in correlation with the conodont biozones (Upper expansa, Early, Middle and Late praesulcata), the ammonoid biozones (do VI = “Wocklumeria Stufe”), or the foraminifera biozones (kobeitusana Zone). Other data for which the stratigraphical range is not based on standard biozonation are provided in the annex. The brachiopod bio-markers discussed in this paper belong to 14 Productidina genera (Acanthatia, Araksalosia, Ericiata, Hamlingella, Mesoplica, Nigerinoplica, Orbinaria, Ovatia, Rugauris, Semiproductus, Sentosia, Spinocarinifera, Steinhagella, Whidbornella), 7 Rhynchonellida genera (Araratella, Centrorhynchus, Hadyrhyncha, Megalopterorhynchus, Novaplatirostrum, Rozmanaria, Tchanakhtchirostrum), 13 Spiriferida genera (Brachythyris, Cyrtospirifer, Dichospirifer, Eochoristites, Imbrexia, Parallelora, Prospira, Rigauxia, Sphenospira, Tenisia, Toryniferella, Tylothyris, Voiseyella) and one Spiriferinida genus (Syringothyris). Other orders have not been studied in this paper. The main features of the uppermost Famennian brachiopod taxa represented in this paper are commented. All these taxa are listed as completely as possible throughout the world (with complements in the annex). A quick summary of the geographic distribution of the more represented taxa is given in conclusion.     

贵州南部独山地区泥盆纪弗拉期的石燕贝类腕足动物

弗拉期是了解泥盆纪弗拉-法门灭绝事件之前古代生物多样性的一个关键阶段.贵州南部的独山地区因其所具有的完整层序及丰富的底栖牛物化石成为我国主要泥盆系参考剖面之一.独山剖面弗拉期的石燕贝类腕足动物至少包括5种类型,其中4种在本文中为首次描述和图示.Conispirifer之前只发现于欧洲及北美的弗拉期地层中,目前也首次在华南独山剖面的同期地层中发现.大多先前从贵州南部其他地区弗拉期地层中描述的多种Cyrtospirifer可归于Cyrtospirifer subextensoides新种内.鉴于前人对Emanuella的模式种E.takwanensis 的内部构造描述的矛盾状况,本文暂把研究区具有抬离壳面的纤细铰窝、固着于壳面的主突起以及其前方下面为外铰板支撑的标本归于E.takwanensis.在华南弗拉期可初步识别出三个腕足动物群组合带:Yocrarhynchus-Phlogoiderhynchus动物群以小嘴贝类-无洞贝类-双腔贝类腕足动物为特征,其时代大致从弗拉阶底部至Pa.punctata带;之后为Cyrtospirifer动物群,以Cyrtospirifer的存在为特征,又可分为早晚两部分,前者(未命名)以conispiriferid类的Conispirifer和Pyramidaspirifer、弓石燕类、无洞贝类等为特征,其时代大致为Pa."hassi"带至下Pa.rhenana带;后者为Hunanotoechia动物群,以丰富的小嘴贝类-无洞贝类.弓石燕类腕足动物为特征,其时代大致为上Pa.rhenana带至Pa.1inguiformis带.     

First evidence of brachiopod diversification after the end-Triassic extinction from the pre-Pliensbachian Internal Subbetic platform (South-Iberian Paleomargin)

Lower Jurassic brachiopods are widely known in the External Betic Zone. Their occurrence was so far virtually restricted to the easternmost Subbetic Zone where they underwent a diversity burst and radiation event during the late Sinemurian–early Pliensbachian interval, leading to a bloom in brachiopod diversity from the early Pliensbachian onwards. Taxonomical and paleobiogeographical analyses performed in a newly recorded assemblage from the most offshore areas of the Subbetic Basin (Granada province, Spain) reveals that this diversification event occurred earlier than expected hitherto, probably in the Turneri–Obtusum chronozones, as similarly observed in the most intra-Tethyan basins such as the Northern Calcareous Alps and Transdanubian Ranges, illustrating the recovery of the background conditions for the establishment of diversified brachiopod communities after the end-Triassic extinction event. A new rhynchonellide species, Alebusirhynchia vorosi nov. sp., is formally described among the ten different taxa recorded for the first time in this area. The Mediterranean paleobiogeographical affinities revealed by the brachiopod assemblage emphasizes that the onset of the Mediterranean/Euro-Boreal bioprovinciality and the initial brachiopod diversification in the pre-Pliensbachian Internal Subbetic platform took place earlier in the Sinemurian as well, following the Euro-Boreal monotypic record previously reported in this region.     

Taxonomical and biostratigraphical remarks on the rhynchonellide “Paurorhyncha” (Brachiopoda) in the Yidade Formation of eastern Yunnan,China

Additional specimens of rhynchonellide brachiopods from the marly limestones in the Yidade Formation at the Panxi section in eastern Yunnan have been ascribed to the species “Paurorhyncha” squamosa Wang, 1956 and “P.” depressa Wang, 1956. The two species have been used frequently as index fossils for the Frasnian (Upper Devonian), but their taxonomic assignments are problematic because their internal structures remain unknown. In this study, detailed systematical examinations on both external characters and internal structures revealed by serial sections suggest that the two species are more appropriately assigned to the genus Hadrotatorhynchus Sartenaer, 1986. Based on the stratigraphical distributions of Hadrotatorhynchus and the conodonts in the upper unit of the Yidade Formation, the Hadrotatorhynchus-bearing horizons are re-considered as the uppermost Givetian (Middle Devonian) rather than Frasnian stage, although the precise position of the Middle/Upper Devonian boundary still depends on further investigations of high-resolution biostratigraphy.     

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.     

Famennian conodonts from the Hongguleleng Formation at the Bulongguoer stratotype section,western Junggar,Northwest China

The Hongguleleng Formation, the highest and most important Devonian marine carbonate horizon in western Junggar, contains an endemic shallow-water Icriodus–Polygnathus conodont fauna with rare palmatolepids and other genera. The conodont faunas from the Bulongguoer and the Wulankeshun sections are similar, with high abundance (about 40%) of endemic taxa (13 taxa), indicating isolation of the Junggar Basin during early Famennian. Non-endemic species in the faunas suggest that the Lower Member of the Hongguleleng Formation is assignable to the Pa. rhomboidea Zone to the Pa. marginifera marginifera Zone of early Famennian, not including the Frasnian–Famennian boundary. The Upper Member may be of late Famennian–early Tournaisian in age on the basis of our preliminary faunal analysis.     

Biostratigraphic control of the latest-Ordovician glaciogenic unconformity in Alnif (Eastern Anti-Atlas, Morocco), based on brachiopods

In the Alnif region of the Eastern Atlas (Morocco), seven fossiliferous horizons within the Lower-Ktaoua and Upper-Tiouririne formations (Ktaoua Group), as well as in the glaciomarine microconglomeratic shales of the Upper Formation of the Second-Bani Group have yielded biostratigraphically significant brachiopods and other taxa, such as trilobites and echinoderms. Several brachiopod species with short stratigraphic ranges, well-known in south-western Europe, allow a precise chronostratigraphic control of a succession that displays important lateral lithological and facies changes, when compared with the type sections in the Central Anti-Atlas. They have also permitted a better consensus between the macrofauna-determined age and that based on micropaleontological analyses. For the first time, the occurrence of a Hirnantia Fauna within the microconglomeratic shales of the Upper Formation of the Second Bani Group is reported. The biostratigraphic conclusions restrict the age of the Latest Ordovician glaciation to the early Hirnantian.     

Late Middle to Late Frasnian Atrypida, Pentamerida, and Terebratulida (Brachiopoda) from the Namur-Dinant Basin (Belgium)

In the Namur-Dinant Basin (Belgium), the last Atrypida and Pentamerida originate from the top of the Upper Palmatolepis rhenana Zone (Late Frasnian). Within this biozone, their representatives belong to the genera Costatrypa, Desquamatia (Desquamatia), Radiatrypa, Spinatrypa (Spinatrypa), Spinatrypina (Spinatrypina?), Spinatrypina (Exatrypa), Waiotrypa, Iowatrypa and Metabolipa. No representative of these orders occurs within the Palmatolepis linguiformis Zone. The disappearance of the last pentamerids, mostly confined to reefal ecosystems, is clearly related to the end of the edification of the carbonate mounds; it precedes shortly the atrypid one. This event, resulting from a transgressive episode, which induces a progressive and dramatic deterioration of the oxygenation conditions, takes place firstly in the most distal zones of the Namur-Dinant Basin (southern border of the Dinant Synclinorium; Lower P. rhenana Zone). It is only recorded within the Upper P. rhenana Zone in the Philippeville Anticlinorium, the Vesdre area, and the northern flank of the Dinant Synclinorium. It would seem that the terebratulids were absent during the Famennian in this basin, probably due to inappropriate facies. Among the 13 species described or briefly discussed (Palmatolepis hassi to Upper P. rhenana zones), Pseudoatrypa godefroidi nov. sp. and Spinatrypina (Exatrypa) marmoris nov. sp. are proposed as new.     

Permophricodothyris Pavlova, 1965 (Brachiopoda, Spiriferida) from the Permian of South China: its morphology, biostratigraphy and distribution

A study of both silicified and nonsilicified specimens of Permian reticularioid brachiopods from South China suggests thatPermophricodothyris, a genus previously rarely reported from China, is actually very common and abundant in the Middle and especially Upper Permian of South China. This study also clarifies, for the first time, that many of the reticularioid brachiopod species previously described asSquamularia in fact belong toPermophricodothyris. The new data presented in this paper also allows a critical evaluation ofPermophricodothyris in relation to its closest allies:Phricodothyris, Squamularia, Bullarina andNeophricodothyris. The revision reveals that a total of 18Permophricodothyris species are present in the Middle and Upper Permian of South China, with only one species,P. squamularioides, having survived the Permian-Triassic mass extinction. Two species,P. grandis (Chao) andP. guangxiensis Han, Zhou & Wang, are redescribed here, providing critical new information on the morphology and taxonomy of these species.     

Pennsylvanian brachiopod,fish and conodont faunas from the Caliza Masiva (San Emiliano Formation) at the Mina Profunda area,Cantabrian Zone,NW Spain

A rock sample obtained from the Caliza Masiva of the San Emiliano Formation (Bashkirian–early Moscovian) in the Mina Profunda area (NE Villamanín) of the Bodón Nappe (Cantabrian Zone, NW Spain) has yielded numerous brachiopods and fish remains not frequently represented in the fossil record. The brachiopod assemblage comprises 13 taxa and is characterized by phosphatic (Langella, Orbiculoidea) as well as exceptionally preserved silicified calcitic elements (a small chonetid, Composita, Crurithyris, Lambdarina, and two minute terebratulids) as the main faunal components. Of special importance is the record of the microbrachiopod Lambdarina winklerprinsi nov. sp., which reduces the large Viséan–Upper Permian gap in the stratigraphic record of this genus. Conodont elements recovered from the same insoluble residue are indicative of the upper Bashkirian Idiognathoides sulcatus parvus Zone. The accompanying fish remains consist of chondrichthyan teeth and scales, an acanthodian scale and osteichthyan tooth-bearing bones, isolated teeth and isolated scales, representing the first Pennsylvanian ichthyoliths analyzed from the Cantabrian Zone. The limestone beds with selective silicification in the San Emiliano Formation provide an exceptional opportunity to improve our knowledge on the patterns of life diversity over geological time.     

Early Cambrian organophosphatic brachiopods from the Xinji Formation,at Shuiyu section,Shanxi Province,North China

Abundant and diverse small shelly fossils have been reported from Cambrian Series 2 in North China, but the co-occurring brachiopods are still poorly known. Herein, we describe seven genera, five species and two undetermined species of organophosphatic brachiopods including one new genus and new species from the lower Cambrian Xinji Formation at Shuiyu section, located on the southern margin of North China Platform. The brachiopod assemblage comprises one mickwitziid (stem group brachiopoda), Paramickwitzia boreussinaensis n. gen. n. sp., a paterinide, Askepasma toddense Laurie, 1986, an acrotretoid, Eohadrotreta cf. zhenbaensis Li and Holmer, 2004, a botsfordiid, Schizopholis yorkensis (Holmer and Ushatinskaya in Gravestock et al., 2001) and three linguloids, Spinobolus sp., Eodicellomus cf. elkaniiformis Holmer and Ushatinskaya in Gravestock et al., 2001 and Eoobolus sp. This brachiopod assemblage suggests a late Age 3 to early Age 4 for the Xinji Formation and reveals a remarkably strong connection with coeval faunas from East Gondwana, particularly the Hawker Group in South Australia. The high degree of similarity (even at species level) further supports a close palaeogeographic position between the North China Platform and Australian East Gondwana during the early Cambrian as indicated by small shelly fossil data.     

山东潍坊寒武系苗岭统乌溜阶馒头组上部的腕足动物

寒武系腕足动物属种多样性高、个体数量丰富、形态差异明显、地理分布广泛,具有辅助寒武系三叶虫生物地层划分和对比的潜力.华北板块寒武系苗岭统沉积和化石记录发育良好,是中国苗岭统的经典研究区之一.前人己针对华北寒武系苗岭统乌溜阶腕足动物的系统古生物学开展了一系列基础工作,但这些相关研究主要集中于辽宁地区,目前对华北其他地区苗...     

Sedimentary and faunal changes across the frasnian/famennian boundary in the canning basin of Western Australia

The Canning Basin of northwestern Australia is a key area for understanding global changes at the “Kellwasser Events” and the Frasnian‐Famennian boundary. Frasnian stromatoporoid‐coral‐cyanobacterial reef platforms stretched out for enormous distances along the palaeoshelf but in the early Famennian they were completely replaced by cyanobacterial reef platforms. An iridium anomaly in the sequence was formerly believed to be at or close to the boundary and was interpreted as possible evidence for an asteroid impact. Recent field work and detailed biostratigraphy in the area east and southeast of Fitzroy Crossing has given dating relevant to the timing and extent of sea level changes, hypoxic incursions and reef backstepping. Goniatites and conodonts provide correlations with the international biostratigraphy.

In the Horse Spring area the stage boundary falls within the Virgin Hills Formation which normally has a rich pelagic goniatite, nautiloid and conodont fauna. In the latest Frasnian (Zone 13 of Klapper 1989; regional Sphaeromanticoceras lindneri Zone) large allochthonous reef blocks moved downslope into the open marine basin. A diverse gastropod fauna is associated with the last atrypid brachiopods. The faunal record at the immediate boundary is obscured by dolomitisation but manticoceratid goniatites range into this level. There is no evidence for the organic‐rich dark Kellwasser limestone facies.

In the McWhae Ridge area two Frasnian goniatite horizons with Beloceras trilobites and the giant Mamicoceras guppyi and Sphaeromanticoceras lindneri transgress over the reef slope. Stromatolitic cyanobacterial beds mark condensations. Again there is no trace of the oxygen‐depleted Kellwasser facies. However, as at Horse Spring, manticoceratids persisted into dolomitic marker beds that have no other preserved macrofauna. The iridium anomaly associated with Frutexites postdates the Frasnian‐Famennian boundary and was formed by cyanobacterial concentration.     

The late jurassic ‘Massenkalk fazies’ of Southern Germany: calcareous sand piles rather than organic reefs

Summary Upper Jurassic (Malm δ to ζ1) massive limestones (‘algal-sponge-reefs, sponge-reefs, reef-complexes, reefs, algal-sponge-bioherms, biolithites, Massenkalk, bioherms, Stillwasser-Mudmounds’) were analyzed in the Southern Swabian Alb, the Southern Franconian Alb and in drilling wells in the Molasse basin (Southern Bavaria). This analysis was carried out within the frame of a multidisciplinary DFG-study with the objective of decifering the controls on the development of Upper Jurassic spongiolites, their three-dimensional distribution, their characteristic faunal composition, and the diagenetic trends of the different primary facies. The data base consists of detailed facies mapping in the areas of the Eybtal and the Blautal (1300 samples) as well as comparative studies in the Upper Donautal (Swabian Alb) and the Southern Franconian Alb (400 samples). All together about 500 thin sections were studied. The distribution of the most important components (ooids, intraclasts, peloids, corals, sponges, sponge spicules, cyanobacterial crusts, brachiopods, molluscs, echinoids, bryozoans, serpulids,Terebella, Tubiphytes), and diagenetic features (dolomite, dedolomite, silicification, stylolites, clay flasers, hematite patches) results in a spatial distribution pattern of facies types. The largest part (70 %) of the massive limestones consists of a peloid-lithoclast-ooid sand facies rich in completely or partly micritized ooids. These ooids, especially in beds of the Malm δ to ε, might be the clue to a reinterpretation of the water depth. True biogenic constructions occur (about 30 % of the volume; sponge-algalmudmounds, algal-sponge-boundstones, and brachiopod-algal-sponge-mounds) within and at the margins of this facies and are interpreted as platform sands. The spatial distribution of the buildups in relation to the sand facies was probably controlled by hydrodynamic conditions. In addition, zoned sponge-algal-mounds occur in intraplatform channels and nodular sponge-algal-mudmounds in the marly basin sediments between platform sand areas. Breccias and slumpings in beds older than the Malm ζ have to be reinterpreted. Most of the breccias found originated from the flanks of the sand platforms, reflecting the faunal composition of the algal-sponge-boundstones which stabilized the flanks. Breccias of this composition occur throughout the Malm δ-ζ1 and differ markedly in their composition from the sand facies. The boundary breccia (Malm ε/ζ1) is interpreted as marking a regressive maximum. The increasing growth of buildups, rich in brachiopods in the Malm ζ1, is ascribed to an increase of reef growth at the beginning of a transgression. Detailed facies analyses necessary for the reconstruction of the spatial distribution of different facies types are in progress. Most of the older data on faunal distributions cannot be used for detailed facies analysis because they differentiated only between massive facies and bedded facies. Therefore Upper Jurassic limestones of Southern Germany should be restudied in order to recognize the volumetric importance of sand facies and buildups within massive limestones.     

Fluctuations in a late devonian-early Mississippian phytoplankton flora of Ohio,U.S.A

Fossil diversity and abundance fluctuations of Late Devonian (Famennian)—Early Mississippian (Tournaisian) phytoplankton assemblages were determined in a stratigraphic sequence of samples in Barberton test core 3, from northeast Ohio. Both diversity and abundance were relatively high during deposition of the lower Chagrin Shale (Upper Devonian) and both decreased during deposition of the upper Chagrin Shale. The overlying Cleveland Shale (Upper Devonian) is characterized by sharply increased abundance with a corresponding (although less marked) increase in diversity; in contrast, both abundance and diversity were at relatively low levels during deposition of the Lower Mississippian Bedford Shale, the uppermost unit of the sequence. These trends agree with the phytoplankton periodicity model presented by Tappan, which indicates large-scale extinction at the end of the Devonian.Many of the acritarch and prasinophycean algal taxa have restricted stratigraphic occurrences within the section studied, making them useful for local and perhaps worldwide correlation. The stratigraphically restricted taxa provide a basis for the proposal of five biostratigraphic zones within the Upper Famennian (Upper Devonian) and a portion of the Lower Mississippian. In ascending order occur: the Gorgonisphaeridium ohioense Assemblage Zone, the Acriora petala Assemblage Zone, the Muraticavea enteichia Range Zone, the Cymatiosphaera labyrinthica Range Zone, and the Navifusa drosera Assemblage Zone.     

Discovery of ventrally directed spiralia in a Permian spiriferellid brachiopod and implications for its feeding system

Spiralia are lophophore‐supporting, coiled internal structures developed in some extinct brachiopods. In spite of considerable variations in their orientation, the spiralia of most spiriferide and spiriferinide taxa are known to be laterally directed. Recent studies have shown that these brachiopods consistently have a median inhalant and lateral exhalant feeding system. Here, we report a Permian spiriferellid brachiopod fossil (Spiriferella protodraschei) bearing ventrally directed spiralia in its interior. Using the serial sections of the specimen, we have reconstructed the detailed morphology and orientation of the spiralia. Each spiralium in the specimen does not show the apically tapering pattern supposedly universal in all the known types of spiralia: instead it maintains a similar diameter even at its last whorl. The spiralia appear to have directly developed from strong and anteriorly extended crura, consisting of ten whorls in one side and 13 whorls in the other side. As the morphology and orientation of spiralia are immediately associated with the arrangement of spirolophous lophophore within the mantle cavity, the extraordinary orientation and form of the spiralia indicate that this brachiopod likely had developed a considerably modified feeding pattern with respect to most other spirolophous brachiopods. It is postulated that the inhalant/exhalant current circulation of the species (and its descendants) would be considerably different from that of other spiriferide taxa. In particular, the combination of the vertically oriented life posture (free‐lying with thickened ventral apex bottom) and ventrally directed spiralia resembles both fossil atrypide and modern rhynchonellide brachiopods in the orientation of spirolophe, suggesting that some spiriferellid brachiopods may have developed a lateral inhalant/median exhalant feeding current system. A few spiriferide and spiriferinide brachiopod taxa with a weakly transverse but strongly convex ventral valve are noted to exhibit similar modifications in their spiralia, possibly due to the spatial limitation of their mantle cavities.