Deep-sea nematode biodiversity in the Mediterranean basin: testing for longitudinal, bathymetric and energetic gradients

The knowledge of the processes controlling the spatial distribution of species diversity is one of the main challenges of the present ecological research. Spatial patterns of benthic biodiversity in the deep sea are poorly known in comparison with other ecosystems and this limits our understanding of the mechanisms controlling the distribution and maintenance of high biodiversity in the largest ecosystems of our biosphere. Although the Mediterranean basin covers <1% of the world ocean surface, none the less it hosts >7.5% of the global biodiversity. The high biogeographic complexity and the presence of steep ecological gradients contribute in making the Mediterranean a region of very high diversity. Here we report the results of an investigation on the patterns of nematode biodiversity in the deep-Mediterranean Sea, in relation with bathymetric, longitudinal and energetic gradients. Our results indicate that benthic biodiversity in the deep-Mediterranean decreases significantly with increasing depth. Moreover, at equally deep sites, nematode diversity decreased from the western to the eastern basin and longitudinal gradients were evident when comparing sites at 4000-m depth, with 3000-m depth. The analysis of the available energy (measured as labile organic matter content of the sediments) suggests that biodiversity patterns are not controlled by the amounts of food resources, but instead bio-availability is the key factor. A more detailed analysis revealed an extremely high deep-sea beta-diversity (turnover diversity), both among sites at different depths as well as at similar depths of different longitude or within the same basin. This new finding has not only important implications on the estimates of the overall regional diversity (gamma diversity), but also suggests the presence of high biogeographic complexity in the deep benthic domain of the Mediterranean Sea.     

Impact of late Quaternary environmental changes on deep-sea benthic foraminiferal faunas of the Red Sea

Fluctuations in abundance, diversity and species composition of benthic foraminifera from two sites of the northern and southern Red Sea indicate strong variability of deep-sea ecosystems during the last four glacial to interglacial cycles. In total, five and four different benthic foraminiferal assemblages have been identified in the northern core and southern core, respectively. Comparison with recent faunas from the Red Sea and adjacent oceans allowed the reconstruction of temporal changes in deep-water ventilation, salinity and food availability at the seafloor. Generally, the abundance of infaunal and miliolid taxa increase during glacial intervals indicating increased organic matter fluxes, oxygen decrease and salinity increase in deep waters during these times. These fluctuations are attributed to enhanced oxygen consumption rates and temporarily reduced deep-water formation in the northern Red Sea during glacial intervals. The recorded environmental changes are a reflection of both high- and low-latitude climate changes. The northern Red Sea is mainly influenced by glacio-eustatic sea level fluctuations that control deep-water formation rates and by mid-latitude climate changes of the Mediterranean region that control surface productivity. In contrast, deep-sea ecosystem variability of the southern Red Sea is additionally influenced by low-latitude climate changes attributed to the NE monsoon intensity that drives the inflow of nutrient-rich surface waters from the Gulf of Aden. These results demonstrate the high sensitivity of deep-sea ecosystems of the Red Sea to both global and regional climate changes.     

Assemblages of sea stars (Echinodermata: Asteroidea) and brittle stars (Echinodermata: Ophiuroidea) in the Weddell Sea (Antarctica) and off Northeast Greenland (Arctic): a comparison of diversity and abundance

Composition and distribution of asteroid and ophiuroid assemblages were investigated by means of Agassiz trawl catches at 34 stations in 220- to 1,200-m depth in the Weddell Sea and at 17 stations in 90- and 830-m depth off Northeast Greenland. A total of 86 species (48 sea stars, 38 brittle stars) were identified in the Weddell Sea whereas off Northeast Greenland a total of 26 species (16 sea stars, 10 brittle stars) were recorded. In both study areas, brittle stars were numerically more important than sea stars, and abundances generally decreased with water depths. Multivariate analyses revealed a conspicuous depth zonation of sea and brittle stars off Greenland. Very high abundances of Ophiocten sericeum and Ophiura robusta characterized the assemblages on shallow shelf banks whereas in greater depths Ophiopleura borealis, Ophioscolex glacialis and Ophiacantha bidentata became dominant, albeit at significantly lower densities. Mass occurrences of brittle stars, such as those recorded on Greenlandic shelf banks, have not been discovered in the Weddell Sea, where distinct assemblages were discriminated in deep shelf trenches as well as on the eastern and southern shelf. Ophioplocus incipiens, Ophiurolepis martensi and Ophiurolepis brevirima were the most prominent species on the eastern shelf. Ophiacantha antarctica, Ophiurolepis gelida and Ophionotus victoriae on the southern shelf, and Ophiosparte gigas as well as the asteriod Hymenaster sp., in the shelf trenches. Overall, the Weddell Sea housed conspicuously more asterozoan species than the waters off Greenland. Higher species diversity was also evident at both a regional and local scale, especially for the eastern Weddell Sea shelf. However, because many species from the Weddell Sea are closely related, the Weddell Sea assemblages were not significantly different from the Greenland ones in terms of taxonomic diversity and distinctness. Received: 29 April 1996/Accepted: 10 June 1996     

Demersal fish assemblages and spatial diversity patterns in the Arctic-Atlantic transition zone in the Barents Sea

Direct and indirect effects of global warming are expected to be pronounced and fast in the Arctic, impacting terrestrial, freshwater and marine ecosystems. The Barents Sea is a high latitude shelf Sea and a boundary area between arctic and boreal faunas. These faunas are likely to respond differently to changes in climate. In addition, the Barents Sea is highly impacted by fisheries and other human activities. This strong human presence places great demands on scientific investigation and advisory capacity. In order to identify basic community structures against which future climate related or other human induced changes could be evaluated, we analyzed species composition and diversity of demersal fish in the Barents Sea. We found six main assemblages that were separated along depth and temperature gradients. There are indications that climate driven changes have already taken place, since boreal species were found in large parts of the Barents Sea shelf, including also the northern Arctic area. When modelling diversity as a function of depth and temperature, we found that two of the assemblages in the eastern Barents Sea showed lower diversity than expected from their depth and temperature. This is probably caused by low habitat complexity and the distance to the pool of boreal species in the western Barents Sea. In contrast coastal assemblages in south western Barents Sea and along Novaya Zemlya archipelago in the Eastern Barents Sea can be described as diversity "hotspots"; the South-western area had high density of species, abundance and biomass, and here some species have their northern distribution limit, whereas the Novaya Zemlya area has unique fauna of Arctic, coastal demersal fish. (see Information S1 for abstract in Russian).     

Meiobenthic stocks and benthic activity on the NE-Svalbard shelf and in the Nansen Basin

Summary High Arctic meiofaunal distribution, standing stock, sediment chemistry and benthic respiratory activity (determined by sediment oxygen consumption using a shipboard technique) were studied in summer 1980 on the NE Svalbard shelf (northern Barents Sea) and along a transect into the Nansen Basin, over a depth range of 240–3920 m. Particulate sediment proteins, carbohydrates and adenylates were measured as additional measures of benthic biomass. To estimate the sedimentation potential of primary organic matter, sediment bound chloroplastic pigments (chlorophylls, pheopigments) were assayed. Pigment concentrations were found comparable to values in sediments from the boreal and temperate N-Atlantic. Meiofauna, which was abundant on the shelf, decreased in numbers and biomasses with increasing depth, as did sediment proteins, carbohydrates, adenylates and sediment oxygen consumption. Meiofaunal abundances and biomasses within the Nansen Basin were comparable with those observed in abyssal sediments of the North Atlantic. Nematodes clearly dominated in metazoan meiofauna. Protozoans were abundant in shelf sediments. Probably in response to the sedimentation of the plankton bloom, meiofauna abundance and biomass as well as sediment proteins, carbohydrates and adenylates were significantly correlated to the amount of sediment bound chloroplastic pigments, stressing the importance of food quantity to determine benthic stocks. Ninety-four percent of the variance in sediment oxygen consumption were caused by chloroplastic pigments. Benthic respiration, calculated per unit biomass, was 3–10 times lower than in the East Atlantic, suggesting low turnover rates in combination with a high standing stocks for the high Arctic benthos.     

Biostratigraphy and paleoecology of the Oligo-Miocene succession in Fars and Khuzestan areas (Zagros Basin,SW Iran)

Paleontological and biostratigraphical studies on carbonate platform succession from southwest Iran documented a great diversity of shallow-water benthic foraminifera during the Oligocene–Miocene. Larger foraminifera are the main means for the stratigraphic zonation of carbonate sediments. The distributions of larger benthic foraminifera in two outcrop sections (Abolhayat and Lali) in the Zagros Basin, Iran, are used to determine the age of the Asmari Formation. Four assemblage zones have been recognized by distribution of the larger benthic foraminifera in the study areas. Assemblage 3 (Aquitanian age) and 4 (Burdigalian age) have not been recognized in the Abolhayat section (Fars area), due to sea-level fall. The end Chattian sea-level fall restricted marine deposition in the Abolhayat section and Asmari Formation replaced laterally by the Gachsaran Formation. This suggests that the Miocene part of the formation as recognized in the Lali section (Khuzestan area) of the Zagros foreland basin is not present in the Abolhayat outcrop. The distribution of the Oligocene larger benthic foraminifera indicates that shallow marine carbonate sediments of the Asmari Formation at the study areas have been deposited in the photic zone of tropical to subtropical oceans. Based on analysis of larger benthic foraminiferal assemblages and microfacies features, three major depositional environments are identified. These include inner shelf, middle shelf and outer shelf. The inner shelf facies is characterized by wackestone–packstone, dominated by various taxa of imperforate foraminifera. The middle shelf is represented by packstone–grainstone to floatstone with a diverse assemblage of larger foraminifera with perforate wall. Basinwards is dominated by argillaceous wackestone characterized by planktonic foraminifera and large and flat nummulitidae and lepidocyclinidae. Planktonic foraminifera wackestone is the dominant facies in the outer shelf.     

Polychaete assemblages of the northwestern Ross Sea shelf: worming out the environmental drivers of Antarctic macrobenthic assemblage composition

We have limited understanding of which environmental factors structure the distribution patterns and composition of Antarctic macrobenthos assemblages, and the spatial scales on which such factors operate. In 2004, the “BioRoss Survey” was conducted on the northwestern Ross Sea shelf between Cape Adare and Cape Hallett in depths of 50–750 m to describe and quantify the assemblage patterns of benthic macroinvertebrates. In order to determine the influence of primary productivity, disturbance and habitat heterogeneity on the distribution and composition of the macrofaunal assemblages, polychaete data derived from 52 grab samples were analysed. Although differences in the composition of polychaete assemblages among different sampling transects and depth strata were not particularly pronounced (yet statistically significant), the results suggested that large-scale differences in both primary productivity and iceberg disturbance influence distribution patterns. The combination of sediment chl a content, sorting coefficient, sponge spicule content and distance to the nearest iceberg scour best explained polychaete assemblage patterns. This finding supports previous contentions that multiple environmental drivers working at varying scales influence Antarctic shelf assemblages. The results do not supply support for a pronounced decoupling of pelagic and benthic systems, as has been suggested by another study of deeper water benthic assemblages on the Ross Sea shelf.     

Recent benthic foraminiferal assemblages from cold-water coral mounds in the Porcupine Seabight

Cold-water coral ecosystems are characterised by a high diversity and population density. Living and dead foraminiferal assemblages from 20 surface sediment samples from Galway and Propeller Mounds were analysed to describe the distribution patterns of benthic foraminifera on coral mounds in relation to different sedimentary facies. Hard substrates were examined to assess the foraminiferal microhabitats and diversities in the coral framework. We recognised 131 different species, of which 27 prefer an attached lifestyle. Epibenthic species are the main constituents of the living and dead foraminiferal assemblages. The frequent species Discanomalina coronata was associated with coral rubble, Cibicides refulgens showed preference to the off-mound sand veneer, and Uvigerina mediterranea displayed abundance maxima in the main depositional area on the southern flank of Galway Mound, and in the muds around Propeller Mound. The distribution of these species is rather governed by their specific ecological demands and microhabitat availability than by the sedimentary facies. Benthic foraminiferal assemblages from coral mounds fit well into basin-wide-scale distribution patterns of species along the western European continental margin. The diversity of the foraminiferal faunas is not higher on the carbonate mounds as in their vicinity. The living assemblages show a broad mid-slope diversity maximum between 500 and 1,300 m water depth, which is the depth interval of coral mound formation at the Celtic and Amorican Margin. The foraminiferal diversity maximum is about 700 m shallower than comparable maxima of nematodes and bivalves. This suggests that different processes are driving the foraminiferal and metazoan diversity patterns.     

Trace fossil associations related to facies of an upper Ordovician low wave energy shoreface and shelf, Oslo-Asker district, Norway

Three trace fossil associations have been identified from facies interpreted as low wave energy upper, middle and lower shoreface deposits, transitional shoreface/shelf deposits and open epicontinental shelf deposits. These were developed in pre-regressive, regressive and transgressive phases associated with eustatic sea level changes caused by the upper Ordovician glaciation. The three associations are the Skolithos-Diplocraterion association, the Diplichnites-Phycodes association and the Thalassinoides association. The Skolithos-Diplocraterion association inhabited upper, middle and lower shoreface environments, the Diplichnites-Phycodes association a lowermost shoreface environment and the Thalassinoides association inhabited offshore epicontinental shelf environmcnts. The Skolithos-Diplocraferion and Diplichnites-Phycodes associations bear strong affinities with previously described Skolithos ichnofacies assemblages (indicating shoreline environments) and the Cruziana ichnofacies forms (indicating shallow water environments down to wave base) respectively. The Thalassinoides assemblage, however, closely compares with assemblages described from Mesozoic epicontinental shelf sequences. Trace fossil diversity achieves a maximum in the lowermost shoreface environment and reduces in deeper water and shallower water environments. This accords with diversity patterns identified in analogous Jurassic epicontinental palaeonvironments. The stratigraphic distribution of some trace fossil association boundaries accords with those of contemporary body fossil associations. However, several body fossil associations are included within the volume of distribution of a single trace fossil assemblage. □ Trace fossil association, shoreface and shelf, Oslo district, upper Ordovician.     

A nearshore–offshore trend in acritarch distribution from the Early–Middle Ordovician of the Yangtze Platform, South China

The stratigraphical interval of the late Early Ordovician Didymograptus deflexus and the early Middle Ordovician Azygograptus suecicus graptolite Biozones was investigated from seven sections from the upper Yangtze Platform, southern China. These are located on different parts of the platform, between the nearshore environments of the Kunming area, Yunnan Province, and the offshore carbonate shelf of the Yichang area, Hubei Province. The assemblages recovered from the different parts of the platform vary both in terms of diversity and composition. The nearshore environments show low diversity assemblages with about 10 acritarch species, whereas the offshore shelf environments reflect higher diversities with about 40 species. The composition of the assemblages also changes from simple morphologies (micrhystrids, leiosphaerids, fusiform acritarchs) in nearshore environments to specimens with longer and more complexly branched processes on the shelf. The polygonomorph acritarchs are common over all the upper Yantze Platform, while the acanthomorph genera Baltisphaeridium and Peteinosphaeridium are the most abundant taxa on the offshore carbonate shelf area. Phylogenetic or sea-level changes are probably not responsible for the compositional and diversity changes that occurred during the investigated interval. This study confirms previous interpretations that poorly diversified Palaeozoic acritarch assemblages occur in neritic environments and more complex, highly diversified assemblages are found on the shelf.     

Living benthic foraminifera of the Okhotsk Sea: Faunal composition,standing stocks and microhabitats

Live (Rose Bengal stained) benthic foraminifera were investigated in surface sediment samples from the Okhotsk Sea to reveal the relationship between faunal characteristics and environmental parameters. Live benthic foraminifera were quantified in the size fraction > 125 µm in the upper 8 cm of replicate sediment cores, recovered with a multicorer at five stations along the Sakhalin margin, and at three stations on the southwestern Kamchatka slope. The stations are from water depths between 625 to 1752 m, located close or within the present Okhotsk Sea oxygen minimum zone, with oxygen levels between 0.3 and 1.5 ml l^- 1. At the high-productivity and ice-free Kamchatka stations, live benthic foraminifera are characterized by maximal standing stocks (about 1700-3700 individuals per 50 cm²), strong dominance of calcareous species (up to 87-91% of total live faunas), and maximal habitat depths (down to 5.2-6.7 cm depth). Vertical distributions of total faunal abundances exhibit a clear subsurface maximum in sediments. At the Sakhalin stations, which are seasonally ice-covered and less productive, live benthic foraminifera show lower standing stocks (about 200-1100 individuals per 50 cm²), lower abundance of calcareous species (10-64% of total live faunas), and shallower habitat depths (down to 2.5-5.4 cm depth). Faunal vertical distributions are characterized by maximum in the uppermost surface sediments. It is suggested that 1) lower and strongly seasonal organic matter flux, caused by the seasonal sea ice cover and seasonal upwelling, 2) lower bottom water oxygenation (0.3-1.1 ml l^- 1), and 3) more pronounced influence of carbonate undersaturated bottom water along the Sakhalin margin are the main factors responsible for the observed faunal differences. According to species downcore distributions and average living depths, common calcareous species were identified as preferentially shallow, intermediate and deep infaunal. Foraminiferal microhabitat occupation correlates with the organic matter flux and the depth of the oxygenated layer in sediments.     

Living benthic foraminifera of the Okhotsk Sea: Faunal composition, standing stocks and microhabitats

Live (Rose Bengal stained) benthic foraminifera were investigated in surface sediment samples from the Okhotsk Sea to reveal the relationship between faunal characteristics and environmental parameters. Live benthic foraminifera were quantified in the size fraction > 125 µm in the upper 8 cm of replicate sediment cores, recovered with a multicorer at five stations along the Sakhalin margin, and at three stations on the southwestern Kamchatka slope. The stations are from water depths between 625 to 1752 m, located close or within the present Okhotsk Sea oxygen minimum zone, with oxygen levels between 0.3 and 1.5 ml l^- 1. At the high-productivity and ice-free Kamchatka stations, live benthic foraminifera are characterized by maximal standing stocks (about 1700-3700 individuals per 50 cm²), strong dominance of calcareous species (up to 87-91% of total live faunas), and maximal habitat depths (down to 5.2-6.7 cm depth). Vertical distributions of total faunal abundances exhibit a clear subsurface maximum in sediments. At the Sakhalin stations, which are seasonally ice-covered and less productive, live benthic foraminifera show lower standing stocks (about 200-1100 individuals per 50 cm²), lower abundance of calcareous species (10-64% of total live faunas), and shallower habitat depths (down to 2.5-5.4 cm depth). Faunal vertical distributions are characterized by maximum in the uppermost surface sediments. It is suggested that 1) lower and strongly seasonal organic matter flux, caused by the seasonal sea ice cover and seasonal upwelling, 2) lower bottom water oxygenation (0.3-1.1 ml l^- 1), and 3) more pronounced influence of carbonate undersaturated bottom water along the Sakhalin margin are the main factors responsible for the observed faunal differences. According to species downcore distributions and average living depths, common calcareous species were identified as preferentially shallow, intermediate and deep infaunal. Foraminiferal microhabitat occupation correlates with the organic matter flux and the depth of the oxygenated layer in sediments.     

Seasonal rather than spatial variability drives planktonic and benthic bacterial diversity in a microtidal lagoon and the adjacent open sea

Coastal lagoons are highly productive ecosystems, which are experiencing a variety of human disturbances at increasing frequency. Bacteria are key ecological players within lagoons, yet little is known about the magnitude, patterns and drivers of diversity in these transitional environments. We carried out a seasonal study in the Venice Lagoon (Italy) and the adjacent sea, to simultaneously explore diversity patterns in different domains (pelagic, benthic) and their spatio‐temporal variability, and test the role of environmental gradients in structuring assemblages. Community composition differed between lagoon and open sea, and between domains. The dominant phyla varied temporally, with varying trends for the two domains, suggesting different environmental constraints on the assemblages. The percentage of freshwater taxa within the lagoon increased during higher river run‐off, pointing at the lagoon as a dynamic mosaic of microbial taxa that generate the metacommunity across the whole hydrological continuum. Seasonality was more important than spatial variability in shaping assemblages. Network analyses indicated more interactions between several genera and environmental variables in the open sea than the lagoon. Our study provides evidences for a temporally dynamic nature of bacterial assemblages in lagoons and suggests that an interplay of seasonally influenced environmental drivers shape assemblages in these vulnerable ecosystems.     

Fossil plankton depth zones

Varied evidence from the known depth correlated distribution of benthic shelly facies communities in the Ordovician and Silurian clearly shows that graptolite taxonomic diversity increased from the nearshore regions to the shelf margin regions, as well as into the deeper portions of epicontinental basinal areas. We then consider a method for determining the depth distribution of individual graptolite taxa. This method depends on correlating the first appearance of graptolite taxa as one departs from shoreline in terms of the underlying benthos, arrayed in benthic assemblages. The first appearance departing from shoreline corresponds with the upper depth limits of each taxon in terms of the underlying depth indicated by the underlying, associated benthos. The depth of maximum abundance for each taxon should correspond to that point in the increasing relative abundance of that taxon where relative increase in abundance ceases. The lower depth limit of each taxon should correspond to that point on the relative abundance curve where increase in total relative abundance ceases.Using information about the stratigraphic ages indicated by the graptolite faunas, zone-by-zone, combined with that provided by the communities present within the associated and formerly underlying benthic assemblages should provide a more refined stratigraphic zonation than that obtained without combining both.     

Large Spatial Scale Variability in Bathyal Macrobenthos Abundance,Biomass, α- and β-Diversity along the Mediterranean Continental Margin

The large-scale deep-sea biodiversity distribution of the benthic fauna was explored in the Mediterranean Sea, which can be seen as a miniature model of the oceans of the world. Within the framework of the BIOFUN project (“Biodiversity and Ecosystem Functioning in Contrasting Southern European Deep-sea Environments: from viruses to megafauna”), we investigated the large spatial scale variability (over >1,000 km) of the bathyal macrofauna communities that inhabit the Mediterranean basin, and their relationships with the environmental variables. The macrofauna abundance, biomass, community structure and functional diversity were analysed and the α-diversity and β-diversity were estimated across six selected slope areas at different longitudes and along three main depths. The macrobenthic standing stock and α-diversity were lower in the deep-sea sediments of the eastern Mediterranean basin, compared to the western and central basins. The macrofaunal standing stock and diversity decreased significantly from the upper bathyal to the lower bathyal slope stations. The major changes in the community composition of the higher taxa and in the trophic (functional) structure occurred at different longitudes, rather than at increasing water depth. For the β-diversity, very high dissimilarities emerged at all levels: (i) between basins; (ii) between slopes within the same basin; and (iii) between stations at different depths; this therefore demonstrates the high macrofaunal diversity of the Mediterranean basins at large spatial scales. Overall, the food sources (i.e., quantity and quality) that characterised the west, central and eastern Mediterranean basins, as well as sediment grain size, appear to influence the macrobenthic standing stock and the biodiversity along the different slope areas.     

Warming shelf seas drive the subtropicalization of European pelagic fish communities

Pelagic fishes are among the most ecologically and economically important fish species in European seas. In principle, these pelagic fishes have potential to demonstrate rapid abundance and distribution shifts in response to climatic variability due to their high adult motility, planktonic larval stages, and low dependence on benthic habitat for food or shelter during their life histories. Here, we provide evidence of substantial climate‐driven changes to the structure of pelagic fish communities in European shelf seas. We investigated the patterns of species‐level change using catch records from 57 870 fisheries‐independent survey trawls from across European continental shelf region between 1965 and 2012. We analysed changes in the distribution and rate of occurrence of the six most common species, and observed a strong subtropicalization of the North Sea and Baltic Sea assemblages. These areas have shifted away from cold‐water assemblages typically characterized by Atlantic herring and European sprat from the 1960s to 1980s, to warmer‐water assemblages including Atlantic mackerel, Atlantic horse mackerel, European pilchard and European anchovy from the 1990s onwards. We next investigated if warming sea temperatures have forced these changes using temporally comprehensive data from the North Sea region. Our models indicated the primary driver of change in these species has been sea surface temperatures in all cases. Together, these analyses highlight how individual species responses have combined to result in a dramatic subtropicalization of the pelagic fish assemblage of the European continental shelf.     

Polychaete diversity in the Scotia Arc benthic realm: Are polychaetes tracers for faunal exchange?

The Scotia Arc is the only shallow-water and island bridge linking nowadays Patagonia and the Antarctic. The Antarctic Circumpolar Current as an oceanographic peculiarity makes this region an interesting biogeographic transition zone, because this frontal system traditionally is said to isolate the Antarctic fauna from that of the adjacent northern ecosystems. Based on benthos samples from three expeditions onboard R/V Polarstern, we studied distribution patterns of 200 polychaete species and 34 major benthic taxa in order to evaluate the role of polychaetes in the benthic realm of this part of the Southern Ocean. ANOSIM test distinguished three station groups: the central eastern Scotia Sea, the continental shelf off South America and stations at the tip of the Antarctic Peninsula. These station groups differed in organism densities and diversities with stations at the tip of the Antarctic Peninsula hosting the most diverse and dense community. The polychaete diversity patterns in the three assemblages evidenced closer connectivity between the tip of the Antarctic Peninsula and the central eastern Scotia Sea than between the continental shelf off South America with either the stations off the tip of the Peninsula or the central eastern Scotia Sea. This is probably supported by the Polar Front, which divides the island chain into two branches. Species distribution and community patterns of polychaetes appear to be associated with oceanographic and sediment conditions in this region. Most of the shared species showed the capability to tolerate differences in hydrostatic pressure. We suggest that the islands of the Scotia Sea may constitute a bridge for exchange of benthic species, particularly for polychaetes with eurybathic distribution and high dispersal capabilities.     

Coralline red algal limestones of the late Eocene alpine Foreland Basin in Upper Austria: Component analysis, facies and palecology

Summary Late Eocene sediments of the Upper Austrian Alpine Foreland Basin discordantly overlie Mesozoic and crystalline rocks, which are deeply eroded and form a distinct pre-Eocene relief. Late Eocene deposits contain red algal limestones with a remarkable lateral extent and a high diversity of sedimentary facies. Towards the south the algal limestones change into more clastic sediments, which are characterized by larger foraminifera and bryozoans. Main components are coralline algal branches and detritus, coralline crusts, rhodoliths, peyssonneliacean aggregates and crusts, nummulitid and orthophragminid foraminifera, corals, bryozoans, as well as terrigenous components. Rank correlation and factor analysis were calculated in order to obtain informations about relations between components. Hierarchical cluster analysis allowed the designation of 17 facies, most of them are dominated by coralline algae. Actualistic comparisons and correlations obtained from statistical analyses allowed the reconstruction of the depositional environments. Main features of the northern area are huge accumulations of unattached coralline algae (branches, rhodoliths, detritus), which are comparable to the present-day “Maerl”-facies. They formed loose frameworks cut by sand channels. The frequency of coralline detritus decreases upsection. Peyssonneliacean algae in higher parts of the profiles show growth-forms that are comparable to peyssonneliaceans of the Mediterranean circalittoral soft bottoms. This succession can be interpreted by an increasing relative sea level. Besides, crustose coralline algal frameworks were growing on morphological highs which are partially comparable to the present-day “Coralligéne de Plateau” of the Mediterranean Sea. In contrast to the northern area, sedimentation rate of the southern area is too low to keep up with rising sea level. The typical succession from nummulitid- to orthophragminid-and bryozoan-dominated facies can be interpreted by an increasing water depth from shallowest subtidal to the deeper photic zone and finally to the aphotic zone.     

Benthic community structures in the North Sea

Coherent assemblages of marine benthic species have been recognized from the early twentieth century, and the classical papers of Petersen (1914, 1918) were based on studies of limited areas in the North Sea. In 1986, a synoptic survey of the North Sea north to 57°N was undertaken by a group of ten laboratories from seven North Sea countries. The results of this survey have recently been published (Heip et al., 1992a, b; Künitzer et al., 1992; Huys et al., 1992), and some of the results are summarized in this paper. The analysis of the macrofauna is based on slightly more than 700 taxa. In general, the North Sea macrofauna consists of northern species extending south to the northern margins of the Dogger Bank, and southern species extending north to the 100 m depth line. The central North Sea is an area of overlap of southern and northern species, especially around the 70 m depth contour. Consistent groupings of species are recognized that were summarized in seven faunal groupings. Macrofaunal body weight, density and diversity increase linearly towards the north. Macrofaunal biomass for the whole area averages 7 g adwt. m⁻² and decreases from south to north. Distribution patterns and trends within the meiofauna were very different. Nematodes, which are the dominant taxon overall, are least abundant in the sandy sediments of the Southern Bight, then increase to a maximum around 53° 30′ N and slowly decrease again towards the north. Copepod density and diversity are highest in the Southern Bight, due to the presence of many interstitial species. A large number of species new to science were recorded by the North Sea Benthos Survey and about 1500 species are expected to occur. Copepods show very distinct assemblages according to water depth and sediment type. The contrasting patterns in latitudinal gradients of body weight and number of species of macro- and meiofauna can be only partially explained. Latitude and sediment characteristics, such as grain size and content in plant pigments, and water depth, determine part of the variance in species composition, density and biomass of the benthic fauna, but the patterns that are observed are different for different benthic groups, requiring careful consideration as to their use in biological monitoring procedures. Distributions are related to current patterns in the North Sea, annual temperature variations and availability of food. However, large parts of the variance in many parameters remain unexplained.