THE COMPOSITION AND STRUCTURE OF LOWER SILURIAN MARINE COMMUNITIES

Five benthic communities occupied the shelf regions of the British Isles, Norway, and North America in Upper Llandovery times. The communities are listed below in order of increasing distance from shore.

• 1 The Lingula Community is the least diverse; it has both infaunal elements, including a protobranch, and two lingulids, and epifaunal elements, including a rhynchonellid, a pterioid, and a cornulitid. A restricted and protected near-shore environment, such as a bay or estuary, is postulated.
• 2 The Eocoelia Community shares elements in common with the former community, but is more diverse and is dominated by epifaunal forms; the many small pedunculate brachiopods probably lived attached to the large leptostrophiid brachiopod.
• 3 The Pentamerus Community is dominated by this genus which lived free and upright on the bottom; smaller pedunculate brachiopods probably attached to this large neighbor.
• 4 The Costistricklandia Community was similar in structure to the former community with the many small pedunculate brachiopods being attached to the large Costistricklandia.
• 5 The Clorinda Community is the most diverse, with a great variety of small brachiopods which were probably able to attach to small objects in this quiet off-shore environment, or to some moderately sized brachiopods, such as Clorinda and Cyrtia, which apparently lived free on the bottom.

The brachiopod dominated communities of the Silurian clearly inhabited the ‘level bottom’, an area now occupied mainly by infaunal forms. The main attachment surfaces for the epifaunal elements of the Silurian communities were disarticulated, convex-upward shells.     

Late Cretaceous gastropod dominated communities of the Western Tarim Basin, Southern Xinjiang, China

Late Cretaceous strata of the Western Tarim Basin, Southern Xinjiang, China are divided into the Kukebai, Wuyitake. Yigeziya and Tuyilouk Formations. The gastropod faunas can be arranged into six communities corresponding to broad depositional environments. Two communities commonly occur in the nearshore shoal environments: the Vernedia pusillina Community from the Lower Member of the Kukebai Formation and the Trochactaeon (Neocylindrites)-Actaeonella Community from the Lower Member of the Yigeziya Formation. Three communities occur in the shelf environments: these are the Helicaulax and Ascensovoluta-Gyrodes Communities of the lower and upper parts of the Upper Member of the Kukebai Formation respectively and the Ampullina-Cantharus Community of the Middle-Upper Member of the Yigeziya Formation. A hypersaline environment is indicated by the Haustator-Corbula Community from the Middle Member of the Kukebai Formation.     

Marine benthic communities in the Upper Devonian of New York

Four distinctive benthic communities occur in the mid-shelf deltaic deposits of the Upper Devonian Sonyea Group: the Bellerophon Community, dominated by herbivorous gastropods and deposit-feeding bivalves; the Prodrtctella Community, dominated by brachiopods with adaptations for life on muddy substrata; the Cypricardella Community, dominated by bivalves and brachiopods with a variety of adaptations; and the Rhipidomella Community, dominated by brachiopods adapted for life on sandy substrata. The geographic and stratigraphic occurrence of the communities suggest that their distribution was controlled by two principal sets of environmental variables: (1) onshore to offshore gradients in salinity, dissolved oxygen concentrations, and sedimentary substrata, and (2) rate of delta progradation. A change in the rate of delta build-out during mid-Sonyea time led to a dramatic change in onshore to offshore community patterns. The change suggests that great care must be used in constructing generalized community models for longer intervals of Paleozoic time.     

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 .     

A ‘Problematic fossil’ revealed:Pycnoporidium ? eomesozoicum Flügel, 1972 (Late Triassic,Tethys)—not an enigmatic alga but a strophomenid brachiopod (Gosaukammerella n.g.)

Summary The microproblematicumPycnoporidium ? eomesozoicum Flügel, 1972, from Upper Triassic reefs of the Alpine-Mediterranean region, Turkey Oman and Iran (originally interpreted as possible alga) represents the type species of a new strophomenid brachiopod genus (Gosaukammerella n.g.). The genus is characterized by a very small, millimeter-sized plano-convex shell, whose ventral valve is attached to the substratum (mainly sponges) by symmetrically arranged outgrowths developing from a pseudopunctate, lamellose foliated shell wall and composed of densely spaced subparallel ‘tubes’ comparable with productide spines secreted by papillose extensions of the mantle.Gosaukammerella seems to be the only reliable candidate for the existence of post-Paleozoic strophomenid (productid ?) brachiopods. Gosaukammerella eomesozoica is restricted to possibly cryptic, shaded reef environments inhabited predominantly by sponges serving as substrates for micromorphic brachiopods.     

Marine faunal assemblages in the Silurian-Devonian Keyser Limestone of the central Appalachians

Sedimentological analysis of the Keyser Limestone (Upper Silurian - Lower Devonian of the central Appalachians) indicates that its sediments were deposited in a range of marginal and shallow marine environments. Major depositional environments include: tidal flat, lagoon, barrier bar and island, and open marine shelf. Each major environment is represented by a lithofacies which is lithologically and faunally distinct. Tidal flat lithofacies are characterized by eurytopic organisms, including ostracodes, gastropods, stromatoporoids and blue-green algae. Lagoon lithofacies are dominated by bryozoans, brachiopods, ostracodes and stromatoporoids. Barrier lithofacies are characterized by rooted crinoids, encrusting bryozoans and robust brachiopods. Open shelf lithofacies contain a diverse fauna of cystoids, crinoids, bryozoans and brachiopods.
The distributions of faunal assemblages in the Keyser show no simple relationship to either water depth or distance from shore. They are, in general, related to the distributions of depositional environments.
Recurring associations of brachiopod genera were not found in the Keyser. With few exceptions, any genus may be found in any subtidal environment. Abundance of brachiopods is related to the abundance of local hard substrates (usually bryozoans).     

关岭新铺卡尼期腕足类古生态及对创孔海百合假浮游生活方式的质疑

关岭新铺晚三叠世早期卡尼期(Carnian)龙层及其下伏竹杆坡组上、中部的腕足类从上到下可划分为Linguloid,Laballa,Pseudokoninckina等3个化石群落。根据各群落特征分子的固着方式、生活习性、贝体形态和大小,以及与其他生物共同埋藏情况,并考虑围岩性质等方面的综合因素,得出Pseudokoninckina群落的古生态底域位于BA4内侧至BA3外侧,Laballa群落的古生态位置相当于BA2,而Linguloid群落则为BA1。在此基础上进一步阐明关岭新铺从竹杆坡组中部至龙层呈现明显海退过程,其古生态底域位置从BA4内侧至BA3外侧一直退到BA1。依据此变化绘出新铺卡尼期海平面下降示意图。对创孔海百合被一些学者推断为假浮游的生活方式提出质疑和讨论,得出当时创孔海百合不可能为假浮游的生活方式,它们应是营底栖的。     

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)。     

DISTRIBUTION AND DIVERSITY OF PENNSYLVANIAN MARINE FAUNAS RELATIVE TO WATER DEPTH AND DISTANCE FROM SHORE

Variability of marine faunas in Middle Pennsylvanian rocks in central Colorado, as generally of modern marine faunas, increases with increasing depth and distance from shore in shallow water. Less than 3 genera of brachiopods lived nearshore, whereas 10 were present in water computed to have been 22 meters deep, about 5 kilometers offshore. Foraminifers increase from 1 or 2 irregularly formed types nearshore, to 6–8 types in offshore areas, and foraminiferal density increases sharply 3^l/2 kilometers offshore at a depth of about 15 meters. Several genera of brachiopods, 3 types of foraminifers, and phylloid algae show a distribution closely related to depth of water and distance from shore. On the basis of significant changes in genera of brachiopods, types of foraminifers, general faunal diversity, and foraminiferal density, three distinct biotopes can be recognized. These biotopes were occupied by the euphemitid, dictyoclostid- Composita , and fusulinid fossil communities.     

Thecal designs in isorophinid edrioasteroids

A study of functional morphology in the edrioasteroid suborder Isorophina reveals three major thecal designs: domal, pyrgate, and clavate. Pyrgate forms are characterized by specialized morphologic features that facilitate extension of the theca well above the substrate and contraction of the theca to a low, streamlined, domal profile. Domal forms have a low, domal thecal profile capable of little shape change. Clavate forms are characterized by modifications to the pyrgate theca to facilitate higher-level suspension feeding. Pyrgate species are generalized edrioasteroids that inhabited a variety of nearshore open-marine environments. Domal species are small specialized edrioasteroids that inhabited shallow-marine to intertidal environments. Clavate species inhabited offshore, open-marine environments. The size reduction of the peripheral rim and the modifications to the pedunculate zone are adaptations allowing clavate edrioasteroids to compete for small attachment sites while maintaining the ability to extend the theca for higher-level suspension feeding and respiration. A phylogenetic analysis of well-known isorophinid edrioasteroids to determine the relationships between the three designs indicates that domal species are a monophyletic group within the Isorophinidae, clavate species are a monophyletic group within the Agelacrinitidae, and isorophinid pyrgate species are a paraphyletic group ancestral to both domal and clavate forms. □ Edrioasteroidea, Echinodermata, functional morphology, phylogeny.     

A southward extension of the Silurian Tuvaella brachiopod fauna

The co-occurrence of Tuvaella gigantea Chernyshev, Tannuspirifer pedaschenkoi (Chernyshev), Leptos-trophia elegestica Chernyshev and Stegerhynchus sp. in the Upper Hongliuxia Formation, Barkol, NE Xinjiang, China, shows that this is a representative of the well-known Tuvaella Fauna in China. The correlation of the Tuvaella gigantea Assemblage Zone of the Upper Hongliuxia Formation at its type locality is discussed to document the correlation of this formation with the Pichishui Beds of the Upper upper Chergak Formation in Tuva, the Woduhe Formation m the Lesser Khingan Mountains, and other equivalents based on the presence in some abundance of T. gigantea , and without T. rackovskii. The Tuvaella gigantea-bearing strata are tentatively assigned to the Ludlow. With regard to the paleozoogeographic affinity of the Tuvaella Fauna it is at present suspected by some that the area inhabited by Tuvaella represents a peculiar zoogeographic province or region during the Silurian. However, the information available indicates that the Tuvaella occurrences in east-central Asia represent nothing more than a specialized environmental situation in the light of the presence of a single endemic genus, i.e. Tuvaella itself, among 25 articulate brachiopod genera examined. The evidence of both lithology and brachiopods investigated has important implications for suggesting a shallow, commonly rough-water environment between low intertidal and high subtidal areas.     

Bouchardia rosea,a vanishing brachiopod species of the Brazilian platform: taphonomy,historical ecology and conservation paleobiology

Dead-live faunal comparisons can offer powerful data to detect natural or human-induced population changes in the late Holocene. Here, we document dead–live comparisons for death assemblages of the brachiopod Bouchardia rosea in nearshore (0–45 m) environments along the northern coast of São Paulo State, Brazil. The sampling programme included 30 stations (14 at Ubatuba, 16 at Picinguaba bay). The bottom was sampled via Van Veen grab sampler, and also dredged. Out of 30 stations, 22 yielded brachiopods. The fidelity estimates were obtained by direct comparisons of live biota with dead shells. A total of 6627 brachiopods were recovered, 5339 (80.6%) from Ubatuba and 1288 (19.4%) from Picinguaba. Out of these, 6621 (99.9%) were empty, dead shells, while only six individuals (0.1%) were found alive, all in the Picinguaba Bay. These results suggest extremely poor dead–live compositional fidelity for B. rosea assemblages. The spatial data suggest that the distribution of B. rosea accumulations has been highly patchy in the region, whereas the great scarcity of live brachiopods may point to a recent decline in local populations. Several lines of evidences indicate that changes in water temperature, nutrient availability, population history and even pollution, may have all affected spatio-temporal dynamics of B. rosea populations.     

Boring Late Cambrian organisms

Specimens of inarticulate brachiopods (family Acrotrctidae) with boreholes were found in Upper Cambrian carbonates in the southern Great Basin of the United States. Some morphologic features and preferred orientation of the borings are similar to those made by predators and previously reported in the fossil and Holocene record. Such predatory activity on brachiopods is previously unknown in Cambrian rocks. Taxa associated with these specimens are not known to have been predators and identity of the predatory organism is unknown. Cambrian brachiopods, predation, paleoecology.     

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.     

Spatio‐temporal transitions in Paleozoic Bivalvia: A field comparison of upper Ordovician and upper Paleozoic bivalve‐dominated fossil assemblages

Because of the uncertain relationship between relative abundance and taxonomic richness, abundance data are potentially critical components in any analysis of faunal dominance within fossil assemblages. However, such data are often not available in the literature. In this paper, relative abundance data collected during field work at 25 Upper Ordovician and 22 Upper Pennsylvanian to mid‐Permian localities are used to evaluate Paleozoic transitions in the environmental distributions of bivalve molluscs. On the basis of multivariate analyses of these data, it is shown that even early in their history, during the Ordovician, bivalves were generally not restricted in abundance to just marginal environments. They apparently thrived in terrigenous—dominated regions, both nearshore and offshore, but were less abundant in carbonate settings. However, by the late Paleozoic, bivalve abundances had increased in some carbonate environments, in part because of the growing abundances of free‐burrowing suspension‐feeders, which had been substantially less abundant even in terrigenous environments earlier in the Paleozoic. The results of this study corroborate those of a companion analysis, using literature‐derived data, of paleoenvironmental diversity transitions exhibited by Paleozoic Bivalvia.     

Origin of brachiopod-bryozoan assemblages in an Upper Carboniferous limestone: importance of physical and ecological controls

Skeletal accumulations in the Upper Carboniferous Bird Spring Formation of southern Nevada have yielded new information regarding ecological succession in the fossil record and the development of shell concentrations. Through hydraulic baffling processes, a community dominated by cndobyssate bivalves and strophomcnid brachiopods was locally supplanted by a community composed primarily of bryozoans, compositid brachiopods, and various epizoans. The general trophic and diversity changes resemble those expected during 'short-term ecological succession' but are due for the most part to the sedimentological and hydraulic regime. In view of this evidence, previous paleontologic examples of 'autogenic' succession arc re-interpreted as having been primarily controlled by the physical environment. Upper Carboniferous, Brachiopoda, Bryozoa, paleoecology, ecological succession, epifauna.     

湘西北地区志留,泥盆系遗迹化石再研究

记述了湘西北地区志留、泥盆系的遗迹化石１４属,２８种、亚种（包括１５新种,２新亚种）。这些遗迹化石主要集中于Ｔｒａｕｅｕｍｉｃｈｉｓ－Ａｒｔｈｒｏｐｈｙｃｕｓ群落（代表前滨下部或近滨上部）,Ｔｈａｌａｓｓｉｎｏｉｄｅｓ－Ｌｏｃｋｅｉａ群落（代表近滨上部或前滨下部环境）和Ｃｒｕｚｉａｎａ－Ｄｉｍｏｒｐｈｉｃｈｎｕｓ群落（代表近滨环境）,并对上述３个遗迹群落的古生态作了分析。     

Les Rhynchonelles du Bathonien du domaineprovencal: Liaison entre l'installation des peuplements et les modalités de la transgression bathonienne dans les Alpes-Maritimes

In the Maritime Alps, the Middle and Upper Bathoniancorresponds to a transgressive episode on the Provençal platform. The advancement of the marine invasion is very noticeable between Grasse and Antibes. The sediment shows the existence of a shallow marine environment in which benthic organisms were predominant.The deposits are at first varied: marly layers withbivalves (burrowers for the most part) and calcareous layers with monospecific populations of brachiopods (Burmirhynchia mediterranea nov. sp. in a calm environment, and B. turgidaBuckman in a more turbulent environment).The deposits then become uniform and show theexistence of very shallows environments: B. decorata kiliani nov. subsp. is the only representative of macrofauna in the micritic limestones that can be attributed to the Upper Bathonian.The Rhynchonellids, as on the other Bathonian carbonateplatforms of Western Europe, seem to be the only organisms likely to stay alive and proliferate in the specific environments in which entirely carbonate sediments are deposited in shallow water. Due to their relationships to those existing in other platforms, the species described in this article permit the clarification of biostratigraphic correlations.     

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。前人已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岭统乌溜阶腕足动物的研究程度仍然有限,相关生物地层学也有待进一步探索。本文首次系统描述了山东潍坊寒武系苗岭统乌溜阶馒头组上部碎屑岩中的腕足动物,共鉴定出舌形贝型亚门4属5种,包括馒头组洪河段的舌孔贝未定种Lingulellotreta sp.indet.以及馒头组上页岩段的山西小舌形贝Lingulella shansiensia(Walcott,1905)、德姆斯小舌形贝Lingulella damesi (Walcott,1905)、伊瑞克乳房贝Acrothele eryx (Walcott,1905)和东方原孔贝Prototreta orientalis(Resser and Endo,1937)。值得注意的是,在馒头组上页岩段的3层岩性略有差异的泥岩中,分布着3层完全不同的腕足动物,共有4种,但这3个组合都位于Bailiella lantenoisi带这一三叶虫生物带。腕足动物组合在极短时间内的快速交替现象,表明了腕足动物对环境背景的微弱变化具有较强的响应。     

Gradient versus cluster analysis of fossil assemblages: a comparison from the Ordovician of southwestern Virginia

Studies in modern ecology indicate that most species are distributed independently along environmental gradients according to their individual requirements. Steep gradients often produce species associations separated by discontinuities; gradual gradients produce broadly-overlapping distributions. Approaching the distribution of species populations as a continuum, using gradient analysis, avoids artificial subdivision of totally intergrading distributions, yet permits discontinuities to emerge where present. Faunas of the Martinsburg Formation (Ordovician) in southwestern Virginia offer an excellent opportunity to test the applicability of gradient analysis in a paleoecological setting. A broad spectrum of environments, from nearshore to open-marine, clastic to carbonate-dominated facies, provide both temporal and geographic variation against which to evaluate changes in species distributions. Variations of five classical, Petersen-type communities were recognized in the Martinsburg using cluster analysis: (1) Lingula, (2) bivalve, (3) Rafinesquina, (4) Onniella, and (5) Sowerbyella-dominated communities. Two gradient analysis techniques, ordination and Markov analysis, revealed the same basic associations. However, ordination and Markov analysis permit arrangement of these associations along one or more interpreted environmental gradients. Factors related to water depth and distance from clastic source areas, particularly bottom stability and disturbance frequency, appear to have been the most important of a complex of interrelated physical parameters. The high-stress, nearshore end of the Martinsburg gradient complex was occupied by a Lingula association, followed seaward by an association of bivalves adapted to less-stressed environments. Low-stress, open-shelf environments were occupied by Rafines-quina, Onniella, or Sowerbyella-dominated associations. Broad overlaps among these articulate brachio-pod associations reflect variations in the open-shelf habitat.