Plant species, biomass and environmental characteristics of relevés along the North America Arctic bioclimate gradient

Question: How do interactions between the physical environment and biotic properties of vegetation influence the formation of small patterned-ground features along the Arctic bioclimate gradient? Location: At 68° to 78°N: six locations along the Dalton Highway in arctic Alaska and three in Canada (Banks Island, Prince Patrick Island and Ellef Ringnes Island). Methods: We analysed floristic and structural vegetation, biomass and abiotic data (soil chemical and physical parameters, the n-factor [a soil thermal index] and spectral information [NDVI, LAI]) on 147 microhabitat releves of zonalpatterned-ground features. Using mapping, table analysis (JUICE) and ordination techniques (NMDS). Results: Table analysis using JUICE and the phi-coefficient to identify diagnostic species revealed clear groups of diagnostic plant taxa in four of the five zonal vegetation complexes. Plant communities and zonal complexes were generally well separated in the NMDS ordination. The Alaska and Canada communities were spatially separated in the ordination because of different glacial histories and location in separate floristic provinces, but there was no single controlling environmental gradient. Vegetation structure, particularly that of bryophytes and total biomass, strongly affected thermal properties of the soils. Patterned-ground complexes with the largest thermal differential between the patterned-ground features and the surrounding vegetation exhibited the clearest patterned-ground morphologies.

Benthic foraminifera of surface samples and sediment cores off Senegal and Gambia, West Africa

Water depth zonation of fifty nine benthonic foraminiferal species in marine sediment surfaces has been described. Some species are combined to groups which mark particular depth zones: an upper and lower shelf-fauna, an upper and lower slope fauna, and a shelf-slope fauna. Dependence on latitude could be ascertained for Textularia panamensis, and submergence effects for Hyalinea balthica.

(Table 2) Species composition of agglutinated foraminifers from the abyssal zone of the Pacific Ocean

The quantitative study of distribution and taxonomic composition of recent living and dead (without plasma) benthic foraminifers revealed three foraminiferal assemblages in bottom sediments of the Pacific Ocean at depths of 3350 to 4981 m. The assemblage dominated by epibenthic Lagenammina difflugiformis, Reophax dentaliniformis, and Saccorhiza ramose occupies slopes of underwater hills. The assemblage with a high share of infaunal Cribrostomoides subglobosum, C. nitidum, and Ammobaculites agglutinans is registered on an abyssal plateau. The assemblage with a significant proportion of large Astrorhiza and Reophax species, which are characterized by active way of life, populates gentle slopes and narrow depressions with potentially strong bottom currents.

Benthic foraminifera in the Arctic Ocean

We examine the quantitative composition of benthic foraminiferal assemblages of Rose Bengal-stained surface samples from 37 stations in the Laptev Sea, and combine this data set with an existing data set along a transect from Spitsbergen to the central Arctic Ocean. Foraminiferal test accumulation rates, diversity, faunal composition and statistically defined foraminiferal associations are analysed for living (Rose Bengal-stained) and dead foraminifers. We compare the results of several benthic foraminiferal diversity indices and statistically defined foraminiferal associations, including Fisher's alpha and Shannon-Wiener diversity indices, Q-mode principal component analysis and correspondence analysis. Diversity and faunal density (standing stock) of living benthic foraminifers are positively correlated to trophic resources. In contrast, the accumulation rate of dead foraminifers (BFAR) shows fluctuating values depending on test disintegration processes. Foraminiferal associations defined by Q-mode principal component analysis and correspondence analysis are comparable. The factor values of the correspondence analysis allow a quantitative correlation between the foraminiferal fauna and the local carbon flux, which may be used as a tool to estimate changes in primary productivity.

Benthic foraminifera off West Africa (1°N to 32°S)

Recent benthic foraminifera (> 125 µm) were investigated from multicorer samples on a latitudinal transect of 20 stations between 1°N and 32°S along the upper slope off West Africa. Samples were selected from a narrow water depth interval, between 1200 and 1500 m, so that changes in water masses are minimized, but changes in surface productivity are important and the only significant environmental variable. Live (Rose Bengal stained) benthic foraminifera were counted from the surface sediment down to a maximum of 12 cm. Dead foraminifera were investigated in the top 5 cm of the sediment only. Five live and five dead benthic foraminiferal assemblages were identified using Q-mode principal component analysis, matching distinct primary productivity provinces, characterized by different systems of seasonal and permanent upwelling. Differences in seasonality, quantity, and quality of food supply are the main controlling parameters on species composition and distribution of the benthic foraminiferal faunas. To test the sensitivity of foraminiferal studies based on the uppermost centimeter of sediment only, a comparative Q-mode principal component analysis was conducted on live and dead foraminiferal data from the top 1 cm of sediment. It has been demonstrated that, on the upper slope off West Africa, most of the environmental signals as recorded by species composition and distribution of the 'total' live and dead assemblages, i.e., including live and dead foraminifera from the surface sediment down to 12 cm and 5 cm, respectively, can be extracted from the assemblages in the top centimeter of sediment only. On the contrary, subsurface abundance maxima of live foraminifera and dissolution of empty tests strongly bias quantitative approaches based on the calculation of standing stocks and foraminiferal numbers in the topmost centimeter.

Communities and microhabitats of living benthic foraminifera from the tropical East Atlantic

Living (Rose Bengal stained) benthic foraminifera were collected with a multicorer from six stations between 2°N and 12°S off West Africa. The foraminiferal communities in the investigated area reflect the direct influence of different productivity regimes, and are characterized by spatially and seasonally varying upwelling activity. At five stations, foraminiferal abundance coincides well with the gradient of surface productivity. However, at one station off the Congo River, the influence of strong fresh water discharge is documented. Although this station lies directly in the center of an upwelling area, foraminiferal standing stocks are surprisingly low. It is suggested that the Congo discharge may induce a fractionation of the organic matter into small and light particles of low nutritional content, by contrast to the relatively fast-sinking aggregates found in the centers of high productivity areas. Quality and quantity of the organic matter seem to influence the distribution of microhabitats as well. The flux of organic carbon to the sea-floor controls the sequence of degradation of organic matter in sediment and the position of different redox fronts. The vertical foraminiferal stratification within sediment closely parallels the distribution of oxygen and nitrate in porewater, and reflects different nutritive strategies and adaptation to different types of organic matter. The epifauna and shallow infauna colonize oxygenated sediments where labile organic matter is available. The intermediate infauna (M. barleeanum) is linked to the zone of nitrate reduction in sediments where epifaunal and shallow infaunal species are not competitive anymore, and must feed on bacterial biomass or on metabolizable nutritious particles produced by bacterial degradation of more refractory organic matter. The deep infauna shows its maximum distribution in anoxic sediments, where no easily metabolizable organic matter is available.

Recent benthic foraminifera of the Scotia Sea and Argentine Basin

We investigated 88 surface sediment samples taken with a multiple corer from the southwestern South Atlantic Ocean for their live (Rose Bengal stained) and dead benthic foraminiferal content. Using Q-Mode Principal Component Analysis six live and six dead associations are differentiated. Live and dead association distributions correspond fairly well; differences are mainly caused by downslope transport and selective test destruction. In addition, four potential fossil associations are calculated from the dead data set after removal of non-fossilizable species. These potential fossil associations are expected to be useful for paleoceanographic reconstructions. Environments are described in detail for the live and potential fossil associations and for selected species. Along the upper Argentine continental slope strong bottom currents control the occurrence of live, dead and potential fossil Angulogerina angulosa associations. Here, particles of a high organic carbon flux rate remain suspended. Below this high energy environment live, dead and potential fossil Uvigerina peregrina dominated associations correlate with enhanced sediment organic carbon content and still high organic carbon flux rates. The live A. angulosa and U. peregrina associations correlate with high standing crops. Furthermore, live and dead Epistominella exigua-Nuttallides umbonifer associations were separated. Dominance of a Nuttallides umbonifer potential fossil association relates to coverage by Antarctic Bottom Water (AABW) and Lower Circumpolar Deep Water (LCDW), above the Calcite Compensation Depth (CCD). Three associations of mainly agglutinated foraminifera occur in sediments bathed mainly by AABW or CDW. A Reophax difflugiformis association was found in mud-rich and diatomaceous sediments. Below the CCD, a Psammosphaera fusca association occurs in coarse sediments poor in organic carbon while a Cribrostomoides subglobosus-Ammobaculites agglutinans association covers a more variable environmental range with mud contents exceeding 30%. One single Eggerella bradyi-Martinottiella communis association poor in both species and individuals remains from the agglutinated associations below the CCD if only preservable species are considered for calculation.

Living benthic foraminifers from the central Arctic Ocean

Fifty short sediment cores collected with a multiple corer and five box cores from the central Arctic Ocean were analysed to study the ecology and distribution of benthic foraminifers. To work out living faunal associations, standing stock and diversity, separate analyses of living (Rose Bengal stained) and dead foraminifers were carried out for the sediment surface. The size fractions between 63 and 125 µm and >125 µm were counted separately to allow comparison with former Arctic studies and with studies from the adjacent Norwegian-Greenland Sea, Barents Sea and the North Atlantic Ocean. Benthic foraminiferal associations are mainly controlled by the availability of food, and competition for food, while water mass characteristics, bottom current activity, substrate composition, and water depth are of minor importance. Off Spitsbergen in seasonally ice-free areas, high primary production rates are reflected by high standing stocks, high diversities, and foraminiferal associations (>125 µm) that are similar to those of the Norwegian-Greenland Sea. Generally, in seasonally ice-free areas standing stock and diversity increase with increasing food supply. In the central Arctic Ocean, the oligotrophic permanently ice-covered areas are dominated by epibenthic species. The limited food availability is reflected by very low standing stocks and low diversities. Most of these foraminiferal associations do not correspond to those of the Norwegian-Greenland Sea. The dominant associations include simple agglutinated species such as Sorosphaerae, Placopsilinellae, Komokiacea and Aschemonellae, as well as small calcareous species such as Stetsonia horvathi and Epistominella arctica. Those of the foraminiferal species that usually thrive under seasonally ice-free conditions in middle bathyal to lower bathyal water depth are found under permanently ice-covered conditions in water depths about 1000 m shallower, if present at all.