Late Devonian benthic marine communities of the central Appalachian Allegheny Front

Four benthic marine communities occur in the clastic facies of the prograding Upper Frasnian-Lower Famennian (Upper Devonian) Foreknobs Formation in the Central Appalachians along the Allegheny Front in Maryland, West Virginia, and Virginia. Deep-water, rapidly prograding environments were inhabited by the Ambocoelia-Chonetes Community, dominated by an epifauna of unattached brachiopods. Offshore bar environments were inhabited by the Cyrtospirifer-Camarotoechia Community, exhibiting adaptations to shallow-water, high-energy conditions and probably lowered salinities. Shallow-water, sublittoral environments were inhabited by the Atrypa-Cypricardella Community, a community in which existed a variety of life habitats and a diverse epifaunal and infaunal association of brachiopods and bivalve molluscs. The Leptodesma-Tylothyris Community flourished in nearshore bar-protected environments in the southern region of the study area, whereas in the north the Cyrtospirifer-Camarotoechia Community inhabited nearshore environments in conjunction with the onshore development of a large fluviodeltaic system.     

Marine paleoecology in the Upper Devonian of New York

In the clastic Genesee Group of the Catskill delta, lateral changes of the fauna are believed to reflect onshore-offshore physicochemical gradients. A shoreward increase of infauna is interpreted as adaptation to increased environmental stress. Free immobile taxa were concentrated offshore, while vagile forms, presumably able to cope with shifting substrata, are dominant nearshore. A shorewards replacement of brachiopods by bivalves reflects the eurytopy and infaunal habits of the bivalves.
In Gencsee time, progradation was first rapid, then slow. The sequence is reversed in the superjacent Sonyea Group and the accompanying reversal of faunal patterns is strong evidence of faunal control by the rate of progradation. This indicates the hazardous nature of attempts to trace 'community evolutioneditor' using only a few studies from each period.     

The Recent Terebratulina Community in the rocky subtidal zone of the Bay of Fundy, Canada

A distinctive rocky subtidal benthic community (the Terebratulina Community) is described from the Bay of Fundy, Canada. It is shown to consist of three closely intcrspersed sub-communities: a cavity sub-community, characterised by chitons, coelenterates, brachiopods, bryozoans, chordates and annelids; a rock-face subcommunity, similar in composition but enriched in brachiopods and sponges; and an upper-surface subcommunity, dominated by algae, chitons, bivalves and echinoids.
Comparison with recently described Jurassic, Cretaceous and Recent (Mediterranean) hard-ground communities suggests a relative constancy in composition, in terms of higher taxa, since the Mesozoic. Development of these (sub-)communities occurs wherever crevice or cavity systems on hard substrates lead to microenvironments differentiated mainly on the basis of water energy and light.
The composition and trophic structure of the community and the life habits of Terebratulina septentrionalis (Couthouy) are related to aspects of the environment. Marked differences in composition between the living cavity sub-community and the death assemblage in the sediments are shown to be due to differential preservation, fragmentation, population dynamics and limited local transportation.     

Middle gaspé limestones communities on the forillon Peninsula,Quebec, Canada (Siegenian,Lower Devonian)

Lower Devonian quiet-water marine benthic communities were examined to determine what parameters of community structure are maintained from locality to locality. The preserved elements of the communities are dominated by brachiopods. Each community has a characteristic taxonomic composition and diversity. From stratigraphically lowest to highest the communities include the newly proposed Dawsonelloides Community which has relatively low diversity, and is strongly dominated by a single species. The Beachia Community is less strongly dominated by a single species. The newly proposed Leptostrophia Community is very strongly dominated by a single species; less common species are rare or absent in the other communities. The Plicoplasia Community is slightly less diverse than the preceeding two communities, but unlike them has six species which dominate one or more collections and are commonly found in reduced abundance in the other collections. The gradual change in taxonomic composition and structure from one community to adjacent communities coupled with the similar structure of diversity and dominance within particular communities suggest that these characteristics are time-averaged and primary structural features of the communities.     

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.     

Lower Silurian animal communities — three case histories

A temporal succession of three marine animal communities of early Silurian age is described from the Midland Valley of Scotland. These formed part of an offshore onshore series of communities developed on the platform of a prograding delta under regressive tectonic conditions. All three communities were trophically structured in the same way as their modern analogues. In the earliest (isorthid-Glassia) community filterers and collectors were dominant; the absence of swallowers is attributable to the nature of the substrate. Soft-bodied swallowers, revealed only by their network of fine burrows, dominated the succeeding ( Skenidioides-Dicaelosia ) community. Filterers, which so dominated the hard-bodied fauna, were of only secondary importance: they lived clustered together in small aggregations, the sediments around which were reworked by the swallowers. A major fall of volcanic ash followed by an increase in the variability of the physical environment resulted in the abrupt replacement of this community by the Eoplectodonta-Visbyella community. Filterers were again dominant, with collectors subordinate; swallowers were comparatively rare.     

Large extinctions of articulate brachiopods in the paleozoic and their ecological and evolutionary consequences

The largest Paleozoic extinctions of articulate brachiopods occurred at the Frasnian—Famennian boundary in the Late Devonian and at the Permian—Triassic boundary. Both extinctions affected taxa of all levels, including orders, but differed in scale, course, and ecological and evolutionary consequences. The Frasnian—Famennian extinction event was selective and evolutionary activity after the crisis varied in different orders. However, in the Early Carboniferous, the brachiopod diversity was mostly restored in comparison with the Devonian maximum. In particular groups, preadaptation played a role in changes in diversity and reconstruction of communities. The brachiopod composition at this boundary changed sharply. The extinction event at the end of Permian was global and accompanied by changes in the biota. Later, in the Meso-Cenozoic, the brachiopod diversity was not restored, and bivalves acquired primary importance in various bottom communities of different sea zones where Paleozoic brachiopods previously dominated. Extinction of brachiopods at this boundary was long and gradual. The symptoms of the ecological crisis in the development of Permian brachiopods are recognized beginning from the Roadian Age, which was probably the onset of this crisis.     

Paleoecology of some Carboniferous Pectinacea

Pectinacean bivalves in the Carboniferous of northern California, USA, occur in all observed biotic communities except bioherms of corals and productoid brachiopods. They show equal abundance in offshore, brachiopod-dominated habitats and nearshore, bivalve-dominated habitats, and are among the only common invertebrates in a 'black shale', restricted marine community. Different morphologic forms of the superfamily show little ecologic division. Pectinacea are least abundant in limestone, and most abundant in terrigenous beds, and bottom type was more important in controlling their distribution than water depth, current activity, or salinity.     

Thick brachiopod shell concentrations from prodelta and siliciclastic ramp in a Tortonian Atlantic–Mediterranean strait (Miocene, Guadix Basin, southern Spain)

Carbonate production by brachiopods in shallow-water habitats is generally expected to be not sufficiently high and temporally persistent to allow them to form very thick and densely packed shell concentrations. The formation of thick brachiopod concentrations requires long-term persistence of populations with high density of individuals, and such circumstances are assumed to be rare especially during the Cenozoic. However, here we show that the large-sized brachiopod Terebratula terebratula, the most common species in benthic assemblages with epifaunal bivalves and irregular echinoids, formed several decameter- to meter-thick, densely packed concentrations in shallow siliciclastic, high-energy environments, in a seaway connecting the Atlantic Ocean with the Mediterranean Sea during the Latest Tortonian (Late Miocene, Guadix Basin, southern Spain). This brachiopod formed (1) meter-scale, thick, parautochthonous concentrations in a prodelta setting and (2) thin, mainly allochthonous, tide- and storm-reworked concentrations in megaripples and dunes. The abundance of brachiopods at the spatial scale of the Guadix Basin seems to be mainly related to intermediate levels of sedimentation rate and current velocity because abundance and thickness of shell concentrations decline both (1) in onshore direction towards delta foresets with high sedimentation rate generated by debris flows and (2) in offshore direction with increasing levels of tide- and storm-induced substrate instability. Although brachiopods in dune and megaripple deposits are more fragmented, disarticulated, and sorted, and have a higher pedicle/brachial valve ratio than in prodelta deposits, taphonomic damage is still relatively high in prodelta deposits. Terebratula terebratula thus formed thick concentrations in spite of that disintegration processes were relatively intense along the whole depositional gradient. Therefore, population dynamic of this species was probably characterized by production maxima that were comparable to some Cenozoic molluscs in terms of their productivity potential to form thick shell concentrations in shallow subtidal environments. We suggest that temporal changes in brachiopod carbonate production have a significant spatial and phylogenetic component because multiple large-sized species of the family Terebratulidae, which underwent radiation during the Cenozoic, attained high abundances and formed shell concentrations in temperate regions.     

Invasive bivalves increase benthic communities complexity in neotropical reservoirs

Invasive bivalves often act as ecosystem engineers, generally causing physical alterations in the ecosystems in which they establish themselves. However, the effects of these physical alterations over benthic macroinvertebrate communities’ structure are less clear. The objective of this study was to characterize the ecological effects of the invasive bivalves Corbicula fluminea and Limnoperna fortunei on the structure of benthic macroinvertebrate communities in neo-tropical reservoirs. Three hypotheses were tested: (1) invasive bivalves act as facilitator species to other benthic macroinvertebrates, resulting in communities with higher number of species, abundance and diversity; (2) invasive bivalves change the taxonomic composition of benthic macroinvertebrate communities; (3) invasive bivalves increase the complexity of benthic macroinvertebrate communities. For that it was used data from 160 sampling sites from four reservoirs. We sampled sites once in each area, during the dry season from 2009 to 2012. The first hypothesis was rejected, as the presence of invasive bivalves significantly decreased the host benthic communities’ number of species and abundance. The second hypothesis was corroborated, as the composition of other benthic macroinvertebrates was shown to be significantly different between sites with and without invasive bivalves. We observed a shift from communities dominated by common soft substrate taxa, such as Chironomidae and Oligochaeta, to communities dominated by the invasive Gastropoda Melanoides tuberculata. The biomass data corroborated that, showing significantly higher biomass of M. tuberculata in sites with invasive bivalves, but significantly lower biomass of native species. Benthic macroinvertebrate communities presenting invasive bivalves showed significantly higher eco-exergy and specific eco-exergy, which corroborate the third hypothesis. These results suggest that while the presence of invasive bivalves limits the abundance of soft bottom taxa such as Chironomidae and Oligochaeta, it enhances benthic communities’ complexity and provide new energetic pathways to benthic communities in reservoirs. This study also suggests a scenario of invasion meltdown, as M. tuberculata was facilitated by the invasive bivalves.     

Developmental stages of Permian bivalves of northeast Asia

In the development of Permian bivalves of northeastern Asia, the following five large stages have been recognized: Asselian-Middle Artinskian, Late Artinskian-Kungurian, Roadian-Wordian, Capitanian-Early Wuchiapingian, and Late Wuchiapingian-Changhsingian. The boundaries between stages correspond to great biotic events and frequently display a sharp change in species diversity of bivalves. During the Permian, brachiopods were gradually replaced by bivalves; this was promoted by repeated sharp changes in the environment.     

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.     

Molluscan assemblages on coral reefs and associated hard substrata in the northern Red Sea

Information on spatial variability and distribution patterns of organisms in coral reef environments is necessary to evaluate the increasing anthropogenic disturbance of marine environments (Richmond 1993; Wilkinson 1993; Dayton 1994). Therefore different types of subtidal, reef-associated hard substrata (reef flats, reef slopes, coral carpets, coral patches, rock grounds), each with different coral associations, were investigated to determine the distribution pattern of molluscs and their life habits (feeding strategies and substrate relations). The molluscs were strongly dominated by taxa with distinct relations to corals, and five assemblages were differentiated. The Dendropoma maxima assemblage on reef flats is a discrete entity, strongly dominated by this encrusting and suspension-feeding gastropod. All other assemblages are arranged along a substrate gradient of changing coral associations and potential molluscan habitats. The Coralliophila neritoidea-Barbatia foliata assemblage depends on the presence of Porites and shows a dominance of gastropods feeding on corals and of bivalves associated with living corals. The Chamoidea-Cerithium spp. assemblage on rock grounds is strongly dominated by encrusting bivalves. The Drupella cornus-Pteriidae assemblage occurs on Millepora-Acropora reef slopes and is strongly dominated by bivalves associated with living corals. The Barbatia setigera-Ctenoides annulata assemblage includes a broad variety of taxa, molluscan life habits and bottom types, but occurs mainly on faviid carpets and is transitional among the other three assemblages. A predicted degradation of coral coverage to rock bottoms due to increasing eutrophication and physical damage in the study area (Riegl and Piller 2000) will result in a loss of coral-associated molluscs in favor of bivalve crevice dwellers in dead coral heads and of encrusters on dead hard substrata.     

Faunal response to sea‐level and climate change in a short‐lived seaway: Jurassic of the Western Interior,USA

Understanding how regional ecosystems respond to sea‐level and environmental perturbations is a main challenge in palaeoecology. Here we use quantitative abundance estimates, integrated within a sequence stratigraphic and environmental framework, to reconstruct benthic community changes through the 13 myr history of the Jurassic Sundance Seaway in the western United States. Sundance Seaway communities are notable for their low richness and high dominance relative to most areas globally in the Jurassic, and this probably reflects steep temperature and salinity gradients along the 2000 km length of the Seaway that hindered colonization of species from the open ocean. Ordination of samples shows a main turnover event at the Middle–Upper Jurassic transition, which coincided with a shift from carbonate to siliciclastic depositional systems in the Seaway, probably initiated by northward drift from subtropical latitudes to more humid temperate latitudes, and possibly global cooling. Turnover was not uniform across the onshore–offshore gradient, but was higher in offshore environments. The higher resilience of onshore communities to third‐order sea‐level fluctuations and to the change from a carbonate to a siliciclastic system was driven by a few abundant eurytopic species that persisted from the opening to the closing of the Seaway. Lower stability in offshore facies was instead controlled by the presence of more volatile stenotopic species. Such increased onshore stability in community composition contrasts with the well‐documented onshore increase in taxonomic turnover rates, and this study underscores how ecological analyses of relative abundance may contrast with taxonomically based analyses. We also demonstrate the importance of a stratigraphic palaeobiological approach to reconstructing the links between environmental and faunal gradients, and how their evolution through time produces local stratigraphic changes in community composition.     

Facies distribution patterns of some marine benthic faunas in early paleozoic platform environments

Worldwide Late Cambrian—Silurian lithofacies patterns indicate that the platforms of that time were sites of accumulation of two essentially different rocks suites: the platform carbonate rocks and the platform terrigenous rocks. Most of the platform rocks accumulated as sediments in shallow marine environments similar to those of the present but far more widely spread.Present-day marine benthic faunas are distributed in depth zones which are primarily controlled by temperature. Faunas tend to occur in substrate-related discrete clusters (communities) within each life zone; similar substrates in different depth zones commonly have different faunal associations. Individual phyletic stocks may encounter environmental optimum or near-optimum conditions in certain areas, that commonly are revealed by an abundance of species and individuals within species in each stock. Environmental optimum conditions depend upon availability of food that may be utilized, modes of feeding of the animals present, water motion, and substrate, among other factors. Organisms in past seas were distributed in patterns similar to those of the present.Carbonate platforms were particularly widespread during the latest Cambrian—Early Ordovician. Intertidal environments spread widely across those platforms during that time and characteristic faunal associations developed in them. Saukiid and related tribolites dominated latest Cambrian carbonate platform intertidal faunas. The Early Ordovician carbonate platform intertidal was dominated by archeogastropod-nautiloid cephalopod faunas. These animals were joined by tabulate corals and certain brachiopods during the latter part of the Ordovician and Silurian as prominent faunal elements in the carbonate platform intertidal—shallow subtidal. Cruziana and related trace fossils, bivalves, and certain tribolites (notably homalonotids and dalmanitids) dominated most terrigenous platform intertidal—shallow subtidal faunas of the Ordovician and Silurian.Articulate brachiopods (primarily orthoids, strophomenoids, and rhynchonelloids) appear to have been relatively prominent during the Early Ordovician in shallow subtidal environments on both carbonate and terrigenous platforms and to have spread down the bathymetric gradient into increasingly deeper subtidal areas of both platforms during the latter part of the Ordovician. Tribolites dominated faunas in relatively moderate to deep subtidal environments on both platforms during the early part of the Ordovician. They were gradually replaced by brachiopods in first the shallower, and later the deeper subtidal as dominant members of the faunas. Brachiopods (primarily pentameroids and spiriferoids) dominated nearly all Silurian warm-water subtidal environments from the shallow subtidal to the edges of the platforms.Platform uplifts in the Middle Ordovician and glacio-eustatic sea-level fluctuations in the Late Ordovician caused environmental changes across the platforms that were accompanied by marked replacements among marine benthic faunas in all environments. The distribution of Ordovician carbonate platforms and glacial deposits suggests that an Ordovician polar region may have been close to present-day equatorial Africa and that Ordovician warm temperate-tropical regions lay close to the present-day North Pole.     

Effect of a transient perturbation on marine bacterial communities with contrasting history

Aims: To evaluate the importance of the bacterial composition on the resilience of the organic matter assimilation in the sea. Methods and Results: Chemostats were inoculated with coastal and offshore bacterial communities. Bacterial density and protein synthesis increased before stabilizing, and this response to confinement was more marked in the offshore chemostats. Before the toluene perturbation the community structure in the coastal chemostats remained complex whereas the offshore chemostats became dominated by Alteromonas sp. After the perturbation, bacterial protein synthesis was inhibited before peaking briefly at a level fivefold to that observed before the perturbation and then stabilizing at a level comparable to that before the perturbation. Alteromonas dominated both the coastal and the offshore communities immediately after the perturbation and the coastal communities did not recover their initial complexity. Conclusions: Cell lysis induced by the toluene perturbation favoured the growth of Alteromonas which could initiate growth rapidly in response to the nutrient pulse. Despite their different community structure in situ, the resilience of protein synthesis of coastal and offshore bacterial communities was dependent on Alteromonas, which dominated in the chemostats. Significance and Impact of the Study: Here we show that although Alteromonas sp. dominated in artificial offshore and coastal communities in chemostats, their response time to the shock was different. This suggests that future perturbation studies on resilience in the marine environment should take account of ecosystem history.     

The Paleoecology,Habitats, and Stratigraphic Range of the Enigmatic Cretaceous Brachiopod Peregrinella

Modern and Cenozoic deep-sea hydrothermal-vent and methane-seep communities are dominated by large tubeworms, bivalves and gastropods. In contrast, many Early Cretaceous seep communities were dominated by the largest Mesozoic rhynchonellid brachiopod, the dimerelloid Peregrinella, the paleoecologic and evolutionary traits of which are still poorly understood. We investigated the nature of Peregrinella based on 11 occurrences world wide and a literature survey. All in situ occurrences of Peregrinella were confirmed as methane-seep deposits, supporting the view that Peregrinella lived exclusively at methane seeps. Strontium isotope stratigraphy indicates that Peregrinella originated in the late Berriasian and disappeared after the early Hauterivian, giving it a geologic range of ca. 9.0 (+1.45/–0.85) million years. This range is similar to that of rhynchonellid brachiopod genera in general, and in this respect Peregrinella differs from seep-inhabiting mollusks, which have, on average, longer geologic ranges than marine mollusks in general. Furthermore, we found that (1) Peregrinella grew to larger sizes at passive continental margins than at active margins; (2) it grew to larger sizes at sites with diffusive seepage than at sites with advective fluid flow; (3) despite its commonly huge numerical abundance, its presence had no discernible impact on the diversity of other taxa at seep sites, including infaunal chemosymbiotic bivalves; and (4) neither its appearance nor its extinction coincides with those of other seep-restricted taxa or with global extinction events during the late Mesozoic. A preference of Peregrinella for diffusive seepage is inferred from the larger average sizes of Peregrinella at sites with more microcrystalline carbonate (micrite) and less seep cements. Because other seep-inhabiting brachiopods occur at sites where such cements are very abundant, we speculate that the various vent- and seep-inhabiting dimerelloid brachiopods since Devonian time may have adapted to these environments in more than one way.     

Microfacies and depositional environment of an Upper Triassic intra-platform carbonate basin: the Fatric Unit of the West Carpathians (Slovakia)

Facies associations of the Rhaetian Fatra Formation from the Veká Fatra Mts. (West Carpathians) were deposited in a storm-dominated, shallow, intra-platform basin with dominant carbonate deposition and variable onshore peritidal and subtidal deposits, with 21 microfacies types supported by a cluster analysis. The deposits are formed by bivalves, gastropods, brachiopods, echinoderms, corals, foraminifers and red algae, ooids, intraclasts and peloids. A typical feature is the considerable variation in horizontal direction. The relative abundance and state of preservation of components as well as the fabric and geometric criteria of deposits can be correlated with depth/water energy-related environmental gradients. Four facies associations corresponding to four types of depositional settings were distinguished: a) peritidal, b) shoreface, above fair-weather wave base (FWWB), c) shallow subtidal, above normal storm wave base and d) above maximum storm wave base. The depositional environment can be characterized as a mosaic of low-relief peritidal flats and islands, shoreface banks and bars, and shallow subtidal depressions. The distribution and preservation of components were mainly controlled by the position of base level (FWWB), storm activity and differences in carbonate production between settings. Poorly or moderately diverse level-bottom macrobenthic assemblages are dominated by molluscs and brachiopods. The main site of patch-reef/biostrome carbonate production was located below the fair-weather wave base. Patch-reef/biostrome assemblages are poorly diverse and dominated by the branched scleractinian coral Retiophyllia, forming locally dm-scale autochthonous aggregations or more commonly parautochthonous assemblages with evidence of storm-reworking and substantial bioerosion by microborings and boring bivalves.Facies types and assemblages are comparable in some aspects to those known from the Upper Triassic of the Eastern and Southern Alps (Hochalm member of the Kössen Formation or Calcare di Zu Formation), pointing to similar intra-platform depositional conditions. The absence of large-scale patch-reefs and poor diversity of level-bottom and patch-reef/biostrome assemblages with abundance of eurytopic taxa indicate high-stress/unstable ecological conditions and more restricted position of the Fatric intra-platform setting from the open ocean than the intra-platform habitats in the Eastern or Southern Alps.     

A model of onshore-offshore change in faunal diversity

Onshore-offshore patterns of faunal change occurred at many taxonomic scales during the Paleozoic Era, ranging from replacement of the Cambrian evolutionary fauna by the Paleozoic fauna to the environmental expansion of many orders and classes. A simple mathematical model is constructed to investigate such change. The environmental gradient across the marine shelf-slope is treated as a linear array of discrete habitats, each of which holds a set number of species, as observed in the fossil record. During any interval of time, some portion of the species in each habitat becomes extinct by background processes, with rates of extinction varying among both clades and habitats, as also observed in the record. After extinction, species are replaced from within the habitat and from immediately adjacent habitats, with proportions dependent on surviving species. This model leads to the prediction that extinction-resistant clades will always diversify at the expense of extinction-prone clades. But if extinction intensity is highest in nearshore habitats, extinction-resistant clades will expand preferentially in the onshore direction, build up diversity there, and then diversify outward toward the offshore. Thus, onshore-offshore patterns of diversification may be the expectation for faunal change quite independently of whether or not clades originate onshore. When the model is parameterized for Paleozoic trilobites and brachiopods, numerical solutions exhibit both a pattern of faunal change and a time span for diversification similar to that seen in the fossil record. They also generate structure similar to that seen in global diversification, including logistic patterns of growth, declining origination but constant extinction within clades through time, and declining overall extinction across clades through time.     

Book review of Littler DM. Littler MM (2000) Caribbean Reef Plants An Identification Guide to the Reef Plants of the Caribbean,Bahamas, Florida and Gulf of Mexico

Information on spatial variability and distribution patterns of organisms in coral reef environments is necessary to evaluate the increasing anthropogenic disturbance of marine environments (Richmond 1993; Wilkinson 1993; Dayton 1994). Therefore different types of subtidal, reef-associated hard substrata (reef flats, reef slopes, coral carpets, coral patches, rock grounds), each with different coral associations, were investigated to determine the distribution pattern of molluscs and their life habits (feeding strategies and substrate relations). The molluscs were strongly dominated by taxa with distinct relations to corals, and five assemblages were differentiated. The Dendropoma maxima assemblage on reef flats is a discrete entity, strongly dominated by this encrusting and suspension-feeding gastropod. All other assemblages are arranged along a substrate gradient of changing coral associations and potential molluscan habitats. The Coralliophila neritoidea–Barbatia foliata assemblage depends on the presence of Porites and shows a dominance of gastropods feeding on corals and of bivalves associated with living corals. The Chamoidea–Cerithium spp. assemblage on rock grounds is strongly dominated by encrusting bivalves. The Drupella cornus–Pteriidae assemblage occurs on Millepora–Acropora reef slopes and is strongly dominated by bivalves associated with living corals. The Barbatia setigera–Ctenoides annulata assemblage includes a broad variety of taxa, molluscan life habits and bottom types, but occurs mainly on faviid carpets and is transitional among the other three assemblages. A predicted degradation of coral coverage to rock bottoms due to increasing eutrophication and physical damage in the study area (Riegl and Piller 2000) will result in a loss of coral-associated molluscs in favor of bivalve crevice dwellers in dead coral heads and of encrusters on dead hard substrata.