Stratigraphic and paleogeographic distribution of the Upper Ordovician bryozoans of Mongolia

The taxonomic diversity and distribution of the Upper Ordovician bryozoans of Mongolia are analyzed. Five heterochronous assemblages of bryozoans are established: Chigertei and Tsagaan del in the Sandbian Stage and Bairim Ovoo, Uuregnur, and Sairin in the Katian Stage. The bryozoans are shown to be important for solving complex problems of stratigraphy of the Ordovician of Mongolia and for paleogeographic reconstructions.     

Late Paleozoic and Triassic bryozoans from the Tethys Himalaya (N India, Nepal and S Tibet)

New findings of Late Paleozoic and Triassic bryozoans from the Tethys Himalaya are reported. Fifteen species, including three undetermined, were described, while determinations at order, family or genus level were obtained on fragmentary material. The studied bryozoans were framed in a solid lithostratigraphic scheme spanning across the Tethys Himalaya for over 1500 km, from NW India (Zanskar and Spiti) through Nepal (Dolpo and Manang) to South Tibet; they are concentrated in correlatable stratigraphic intervals corresponding to the Upper Devonian-Tournaisian, Bashkirian, Sakmarian, Midian and Lower Norian. This study allowed us to integrate bryozoan occurrences reported from the literature, mostly referring to the Tournaisian, Bashkirian and Early Norian, and to refine the available biostratigraphic dates. Bryozoan-rich intervals are interpreted as associated to long-term transgressions, in turn driven by major climatic and/or tectonic events. Cosmopolitan genera which prevail in the Paleozoic, are replaced by provincial forms after the Triassic crisis.     

Ecologic stability of the dysaerobic biofacies during the Late Paleozoic

Late Paleozoic faunas found in sediments of dysaerobic origin represent a unique community type that remained essentially unchanged from the Middle Devonian to the Early Permian. This dysaerobic community had the following unique characteristics: (1) dominance by vagile molluscs, with suspension-feeding brachiopods, bryozoans, corals and echinoderms usually subordinate; (2) a trophic structure dominated by deposit feeders and carnivorous or scavenging cephalopods; (3) small body size as the result of high juvenile mortality and possibly stunting; and (4) preservation of most individuals as pyritic stein-kerns. All these characteristics are directly linked to the low oxygen levels of the dysaerobic environment. The relative stability of the deeper water, dysaerobic environment is consistent with Sanders' Stability-Time Hypothesis. The dysaerobic environment apparently offered a refuge for more archaic biotas that evolved in nearshore environments of the early Paleozoic.     

Zooid orientation structures and water flow patterns in Paleozoic bryozoan colonies

Anstey, Robert L. 1981 12 15: Zooid orientation structures and water flow patterns in Paleozoic bryotoan colonies. Lethaia . vol. 14, pp. 287–302. Oslo. ISSN 0024–1164.
By means of direct physical evidence provided by zooecial orientation structures, active water flow systems in Paleozoic bryozoans are inferred to be variously centripetal, centrifugal, or basipetal. Monticules, previously assessed as excurrent water outlets, fall into three additional functional types: incurrent, bypassed, and funnel. In one species circular zoarial fenestrations served as excurrent water outlets. Water flow patterns are strongly correlated with zoarial growth form, which vanes in a general way with inferred habitat conditions in ancient environments. Monticular astogeny and phylogeny include a graded series of sizes, types, and functions. Analogy with zooidal polarities in extant stenolaemates suggests that colony bases and centripetal monticules in the Paleozoic orders were anally budded, but that erect branches and centrifugal monticules were orally budded, a character shared only by the freshwater Phylactolaemata. * Bryozoa, Stenolaemata, functional morphology, monticule function, hydrodynamics, feeding currents, Palaeozoic .     

Studies of fossil brachiopods of Mongolia: Ordovician,Silurian, Devonian,Carboniferous, and Permian

Fossil brachiopods from the Ordovician, Silurian, Devonian, Carboniferous, and Permian deposits of Mongolia have been studied for the last forty-five years by the Joint Soviet-Mongolian (later RussianMongolian) Paleontological and Geological Expeditions. New data on the taxonomic composition, stratigraphic and geographic distribution of the brachiopod assemblages have been obtained. The brachiopod systematics has been further refined and detailed, and the stratigraphic and correlation scales and biogeographic reconstructions have been elaborated for the Paleozoic of Mongolia.     

The role of bryozoans in fossil reefs—an example from the Middle Devonian of the Western Sahara

Stenolaemate bryozoans with their stable calcitic skeletons play a significant role in reef building. In the Middle Devonian Sabkhat Lafayrina reef complex (Western Sahara), bryozoans are abundant and diverse. Although they do not form part of the principal framework of reefs, bryozoans are involved significantly in reef growth, especially in the initial stage. In this way, bryozoans are important with respect to initiating reef growth. They contribute greatly to sediment stabilization, making it possible for principal reef builders to grow on hardened and stabilized substrates, and also play sediment-baffling and sediment-filling roles. The aim of this study is to document the diversity of bryozoans in a Middle Devonian reef complex and to estimate their potential for initiation and contribution to reef structures.     

Bryozoans of the Latorp and Volkhov horizons (Lower-Middle Ordovician) of the Leningrad Region

This monograph deals with the morphology of bryozoans of the orders Cystoporida and Trepostomida from the Latorp and Volkhov horizons (Lower-Middle Ordovician, Floian-Darriwilian stages) of the Leningrad Region. Diagnoses of some families belonging to these orders are revised. Based on morphological features of the bryozoans studied and on the succession of their taxa, four assemblages and three stages in their development are established. In addition, the stratigraphic and paleogeographic distributional patterns of bryozoans in the Lower and Middle Ordovician of the world are analyzed.     

New Devonian and Carboniferous bryozoans of the Holy Cross Mountains (central Poland)

New bryozoans from the Middle-Upper Devonian (Eifelian-Frasnian) and the Lower Carboniferous (Tournaisian) of the Holy Cross Mountains in central Poland are described: Bigeyella sparsa gen. et sp. nov., B. separata gen. et sp. nov., Eridopora singula sp. nov., Leptotrypa pulchra sp. nov., Kysylschinipora klarae sp. nov., Coelotubulipora rara sp. nov., Alternifenestella genuina sp. nov., Exfenestella polonica sp. nov., and Rectifenestella localis sp. nov. Some paleogeographic and stratigraphic aspects of the Paleozoic deposits of this region are discussed and main bryozoan localities are described.     

Bryozoans and microbial communities of cool-temperate to subtropical latitudes—paleoecological implications

The environmental distribution of encrusting bryozoans settling on disarticulated and living bivalve shells has been recorded from five stations in Japan and New Zealand. Some insight into the observed distribution patterns emerges from information on the interaction of bryozoans with microbial mats. Advancing existing classifications, we have subdivided the encrusting bryozoan morphotypes into seven different growth types that largely reflect the biological potentials of bryozoans in competition for space on substrate surfaces. The frequency distribution of these types (s-/c-/m-/z-laminae, runners, spots, bryostromatolites) reveals the influence of microbial mats as a control factor of bryozoan substrate coverage. Microbial mats in turn are correlated with latitudinal gradients in Japan and New Zealand from cool-temperate to subtropical and tropical waters. Unlike erect bryozoans, laminar ones are probably underrated as facies fossils. Accordingly, laminar bryozoan growth types are reconsidered as a tool for paleoecological interpretation of marine hard substrate communities.     

Effects of two marinas on the composition of fouling assemblages

The composition of fouling assemblages was surveyed inside and near two fully enclosed marinas using settlement plates. The location of a plate, inside or outside the marina, influenced the abundances of four functional groups of fouling organisms (solitary ascidians, arborescent bryozoans, encrusting bryozoans and colonial ascidians). Transplantation of mature assemblages revealed that reductions in the abundance of arborescent bryozoans inside marinas might be explained by increased growth and recruitment of these bryozoans outside the marina. Surveys of settlement revealed decreased recruitment of encrusting bryozoans inside the marinas, a result consistent with patterns of adult abundance. It is proposed that an increased abundance of solitary ascidians inside marinas may be due to decreased competition. A second survey of adult assemblages was performed with multiple ‘Outside’ sites per marina. Effects of location consistent with the first survey were found for arborescent bryozoans, and in one marina area for solitary ascidians and encrusting bryozoans, but not in the other. Although mechanisms can be proposed to explain the effects of the marina (inside or outside) on the abundances of solitary ascidians, arborescent bryozoans and encrusting bryozoans, the second survey revealed that the effects may vary among marinas.     

Bryozoaires et Brachiopodes des “Calcaires bajociens a Bryozoaires” des Causses (France) et leur paléoécologie

Two Lower Bajocian new localities, with many silicified bryozoans and brachiopods have been discovered in Causses (France): Saint-Rome-de-Cernon (Aveyron) and Mende (Lozère). The study of the faunas allows to increase the anterior knowledges supplied by the study of localities in the South of Causses (neighbourhood of Alzon and Sumène = North of the Seuil sud-caussenard).The interest is, at first, the distance between the localities: Mende in the North, Saint-Rome-de-Cernon in the West. Moreover, in the second, well-preserved faunas were fossilized without transport. Bryozoans and brachiopods are very numerous. Large and complete bryozoan colonies have been get out by action of HCl. Fine internal structures like rhynchonellid Nannirhynchia or like thecidean transversarium and hemispondylium have been studied. At last, observations and paleoecological hypothesis are drawn, likewise comparison between three localities or locality-groups known in the Causses basin.The fauna from Saint-Rome-de-Cernon is biocoenosis, the substratum was soft, the hydrodynamism low and the depth, according to lamellibranchs, about 200 meters. The beds with bryozoans and small brachiopods contain a fauna with two predominant bryozoan species: Ceriocava straminea and Mesenteripora wrighti. These two species could built large colonies, in spite of fine granulometry of the sedimentary bottom. These colonies provided supports to less adapted bryozoans, to thecidids and to lamellibranch Lopha. The beds with terebratulids and bryozoans contain lesser bryozoans and thecidids. Terebratulids are the main group and are fixed with their peduncle to organic support above the irregular deposits. Beside these benthic animals, the rhynchonellid Nannirhynchia campestriensis is epibiontic on gorgonids or epiphytic on floating sea-weeds.On the contrary, all fossils from Mende have been more or less carried, excepted large sponges perhaps. The most numerous bryozoans (Ceriocava straminea, Mesenteripora wrighti) and numerous Lenticulines (Foraminifera) lived on hard bottom with moving water in community with crinoids. It is possible that a smaller part of bryozoans, requiring lower hydrodynamism, lived, either in sheltered parts of the crinoid community or closer to the sedimentation zone. Thecidids were fixed either on bryozoans in crinoid communities or on shelly remains and on bryozoans nearer to the sedimentation zone. Like in Saint-Rome-de-Cernon, Nannirhynchia campestriensis was epibiontic, or epiphytic, or both.In comparison with the whole localities from the Seuil sud-caussenard, we notice the simplicity of Saint-Rome-de-Cernon and Mende. The localities from the Seuil sud-caussenard were assemblages proceeding from 2 or 3 biotops. On the contrary, at Saint-Rome-de-Cernon we have a biocoenosis and at Mende a thanatocoenosis supplied by one biotop almost.     

Evaluation of malacosporean life cycles through transmission studies

Myxozoans, belonging to the recently described Class Malacosporea, parasitise freshwater bryozoans during at least part of their life cycle, but no complete malacosporean life cycle is known to date. One of the 2 described malacosporeans is Tetracapsuloides bryosalmonae, the causative agent of salmonid proliferative kidney disease. The other is Buddenbrockia plumatellae, so far only found in freshwater bryozoans. Our investigations evaluated malacosporean life cycles, focusing on transmission from fish to bryozoan and from bryozoan to bryozoan. We exposed bryozoans to possible infection from: stages of T. bryosalmonae in fish kidney and released in fish urine; spores of T. bryosalmonae that had developed in bryozoan hosts; and spores and sac stages of B. plumatellae that had developed in bryozoans. Infections were never observed by microscopic examination of post-exposure, cultured bryozoans and none were detected by PCR after culture. Our consistent negative results are compelling: trials incorporated a broad range of parasite stages and potential hosts, and failure of transmission across trials cannot be ascribed to low spore concentrations or immature infective stages. The absence of evidence for bryozoan to bryozoan transmissions for both malacosporeans strongly indicates that such transmission is precluded in malacosporean life cycles. Overall, our results imply that there may be another malacosporean host which remains unidentified, although transmission from fish to bryozoans requires further investigation. However, the highly clonal life history of freshwater bryozoans is likely to allow both long-term persistence and spread of infection within bryozoan populations, precluding the requirement for regular transmission from an alternate host.     

Bryozoans as taphonomic engineers,with examples from the Upper Ordovician (Katian) of Midwestern North America

A combination of encrusting calcitic bryozoans and early seafloor dissolution of aragonitic shells recorded in the Cincinnatian Series of the upper Midwest of North America allowed the preservation of abundant moulds of mollusc fossils bioimmured beneath the attachment surfaces of the bryozoans. We here call this preservational process ‘bryoimmuration’, defined as a bryozoan‐mediated subset of bioimmuration. The bryozoans moulded very fine details of the mollusc shells, usually with more accuracy than inorganic sediment moulds. Most of the bryozoans are heterotrypid trepostomes with robust low‐Mg calcite skeletons. The molluscs are primarily bivalves, gastropods, nautiloids and monoplacophorans with their originally aragonitic shells now dissolved. Many of the encrusting bryozoans are so thin and broad that they give the illusion of calcitic mollusc shells clinging to the moulds. Some molluscs in the Cincinnatian, especially monoplacophorans and epifaunal bivalves, would be poorly known if they had not been bryoimmured. Unlike internal and external moulds in sediment, bryoimmured fossils could be transported and thus record aragonitic faunas in taphonomic assemblages (e.g. storm beds) in which they would otherwise be rare or absent. In addition, bryoimmurations of aragonitic shells often reveal the ecological succession of encrustation on the shells by exposing the earliest encrusters and borings that were later overgrown. Bryoimmuration was common during the Late Ordovician because the calcite sea at the time quickly dissolved aragonitic shells on the seafloor before final burial, and large calcitic bryozoans very commonly used molluscs as substrates. Bryoimmuration is an important taphonomic process for preserving aragonitic faunas, and it reveals critical information about sclerobiont palaeoecology. Several Cincinnatian mollusc holotypes are bryoimmured specimens. Bryozoans involved in bryoimmuration enhance the preservation of aragonitic fauna and thus act as taphonomic engineers.     

奥陶纪苔藓动物的多样性演变——兼论苔藓动物的起源

苔藓动物是一类多为海生、滤食性的群体生物。奥陶纪是苔藓动物发生、演化辐射和灭绝的重要时期,也是苔虫礁形成的最早时期。已知最老的化石苔藓动物发现于中国特马豆克晚期。构成苔藓动物基本分类框架的狭唇纲(包括变口目、隐口目、泡孔目和管孔目)和宽唇纲(包括窗孔目和栉口目)也都是在奥陶纪时期逐步形成的,其中,变口目出现于特马豆克期Tr2时间段,在弗洛期和大坪期,多样性较低,但从达瑞威尔期开始,经桑比期至凯迪期,多样性不断增高,并出现辐射。隐口目(特别是"双叶类隐口目苔虫")也经历了与变口目相类似的发展过程,但它首次出现的时间要相对略迟于变口目。这两个目在整个奥陶纪苔藓动物群中一直占据主导地位。泡孔目、管孔目和窗孔目,先后首次出现在弗洛期Fl2时间段、大坪期Dp1和Dp2时间段,但它们在整个奥陶纪期间一直处于低多样性态势。至于栉口目,它首次出现的时间可能更迟,在凯迪期Ka4时间段,犹如昙花一现。苔藓动物的演化在接近奥陶纪末时呈两幕式灭绝,一次发生在凯迪期Ka2时间段(可能相当于塔凯和安斯蒂的"拉夫塞伊灭绝"),另一次发生在赫南特期Hi2时间段(可能相当于塔凯和安斯蒂的"赫南特灭绝")。分子生物学和形态学证据表明,苔藓动物属原口动物,而不是以前长期认为的后口动物,或介于原口动物和后口动物之间的过渡类型;而且,苔藓动物与腕足动物、帚形动物之间没有直接的亲缘关系。苔藓动物可能起源于一种叫原内肛动物的生物,它们的目一级分类单元之间的系统发育关系目前尚未形成共识,本文绘制的谱系图还有待于化石记录的不断补充和分子生物学研究的逐步介入以使其日趋完善。     

Faunal migration into the Late Permian Zechstein Basin - Evidence from bryozoan palaeobiogeography

Late Permian bryozoans from the Wegener Halvø, Ravnefjeld and Schuchert Formations in East Greenland have been investigated. 14 genera are recognised.Integration of the new bryozoan data from the Upper Permian of East Greenland with data on the distribution of Permian bryozoans along the northern margin of Pangea is used to test hypotheses concerning Late Palaeozoic evolution of the North Atlantic region. During the Permian, the Atlantic rift system formed a seaway between Norway and Greenland from the boreal Barents Shelf to the warm and arid Zechstein Basin. This seaway is considered to be the only marine connection to the Zechstein Basin and therefore the only possible migration route for bryozoans to enter the basin. The distribution of Permian bryozoans is largely in keeping with such a connection from the cool Barents Shelf past the East Greenland Basin to the warm Zechstein Basin and also corroborates the change in temperature through this connection.     

塔里木盆地奥陶系苔虫多样性演变

本文研究了塔里木盆地奥陶系苔虫的分布延限和演化规律。苔虫的演化分为四个阶段:第一阶段为鹰山组—一间房组中下部沉积期,苔虫多样性和丰度均很低;第二阶段为一间房组上部—吐木休克组沉积期,苔虫的多样性处在一种波动上升的状态;第三阶段包括良里塔格组良五段—良四段沉积期,苔虫多样性和丰度均有一定程度的增加,而且绝大部分属种在接下来的第四阶段都有出现;第四阶段在良里塔格组良三段—良一段沉积期,是苔虫总体多样性明显增长的时期,也是本区奥陶纪苔虫的重要辐射期。随后,展开苔虫的多样性分析的研究。研究发现塔里木盆地奥陶系苔虫的演化趋势与世界范围的演化趋势基本一致,说明是受苔虫宏演化规律控制的。     

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

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

新疆西准噶尔晚泥盆世法门期的苔藓虫动物群及其形态特异性和古环境意义*

本文重新研究了产自新疆西准噶尔晚泥盆世法门期洪古勒楞组和铁列克提组的苔藓虫动物群, 重点关注其形态学特征和古生态学意义。该苔藓虫动物群由隐口目(Cryptostomata)、窗格苔藓虫目(Fenestrata), 变口目 (Trepostomata)和泡孔目(Cystoporata)组成。优势类群隐口目苔藓虫是具有中轴或中板的细枝状和双叶状群体。 它们的虫室具有加厚的外区体壁, 弯曲强烈的自虫室, 数量众多的大虫室(metazooecia)或隐蔽虫室(tectizooecia), 以及径向刺突和/或棘状刺突。多样化的异虫室、刺突、自虫室等结构, 以及不同生长环境中的不同生态组合是新疆西准噶尔法门期苔藓虫成功演化的基础。洪古勒楞组下段和铁列克提组下部的苔藓虫动物群可以划分为三类生态组合(即: No. 1, No. 2和No. 3), 代表前滨—近岸—远岸的沉积环境。其中, 近岸环境的枝状—双叶状—网状苔藓虫组合的属种丰度、分异度最高。