Influence of the mode of macrofauna-mediated bioturbation on the vertical distribution of living benthic foraminifera: First insight from axial tomodensitometry

We investigated the influence of bioturbation by macrofauna on the vertical distribution of living (stained) benthic foraminifera in marine intertidal sediments. We investigated the links between macrofaunal bioturbation and foraminiferal distribution, by sampling from stations situated on a gradient of perturbation by oyster-farming, which has a major effect on benthic faunal assemblages. Sediment cores were collected on the French Atlantic coast, from three intertidal stations: an oyster farm, an area without oysters but affected by oyster biodeposits, and a control station. Axial tomodensitometry (CT-scan) was used for three-dimensional visualization and two-dimensional analysis of the cores. Biogenic structure volumes were quantified and compared between cores. We collected the macrofauna, living foraminifera, shells and gravel from the cores after scanning, to validate image analysis. We did not investigate differences in the biogenic structure volume between cores. However, biogenic structure volume is not necessarily proportional to the extent of bioturbation in a core, given that many biodiffusive activities cannot be detected on CT-scans. Biodiffusors and larger gallery-diffusors were abundant in macrofaunal assemblage at the control station. By contrast, macrofaunal assemblages consisted principally of downward-conveyors at the two stations affected by oyster farming. At the control station, the vertical distribution of biogenic structures mainly built by the biodiffusor Scorbicularia plana and the large gallery-diffusor Hediste diversicolor was significantly correlated with the vertical profiles of living foraminifera in the sediment, whereas vertical distributions of foraminifera and downward-conveyors were not correlated at the station affected by oyster farming. This relationship was probably responsible for the collection of foraminifera in deep sediment layers (> 6 cm below the sediment surface) at the control station. As previously suggested for other species, oxygen diffusion may occur via the burrows built by S. plana and H. diversicolor, potentially increasing oxygen penetration and providing a favorable microhabitat for foraminifera in terms of oxygen levels. By contrast, the absence of living foraminifera below 6 cm at the stations affected by oyster farming was probably associated with a lack of biodiffusor and large gallery-diffusor bioturbation. Our findings suggest that the effect of macrofaunal bioturbation on the vertical distribution of foraminiferal assemblages in sediments depends on the effects of the macrofauna on bioirrigation and sediment oxidation, as deduced by Eh values, rather than on the biogenic structure volume produced by macrofauna. The loss of bioturbator functional diversity due to oyster farming may thus indirectly affect infaunal communities by suppressing favorable microhabitats produced by bioturbation.     

Breaching biogeographic barriers: the invasion of <Emphasis Type="Italic">Haynesina germanica</Emphasis> (Foraminifera,Protista) in the Bahía Blanca estuary,Argentina

Large populations of the living benthic foraminifera Haynesina germanica (Ehrenberg 1840) are reported for the first time from marsh and intertidal mud flat sediments of the Bahía Blanca estuary (Argentina). Maximum abundance of living specimens was recorded in shallow intertidal environments. The species was previously recorded from many European and North American shallow-water coastal settings, but has not been documented from Argentina. Comparative faunal assemblage analysis from dated core sample material from within the Bahía Blanca estuary shows that the species has not been present for at least the last 8,200 years. This supports the hypothesis that the species has been accidentally introduced outside its natural range as a probable result of ballast water and/or shipping activities. This study is the first report of a successful invasion of non-indigenous benthic foraminifera to the South Atlantic coast of Argentina. The introduction of nonnative foraminiferal species provides new evidence that human-mediated breaching of biogeographic barriers, will ultimately result in the biotic homogenization of foraminiferal intertidal faunal assemblages.     

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.     

Benthic foraminiferal population fluctuations related to anoxia: Santa Barbara Basin

The pore-water geochemistry and benthic foraminiferal assemblages of sediments from two slope sites and within the central portion of the Santa Barbara Basin were characterized between February 1988 and July 1989. The highest foraminiferal numerical densities (1197 cm^–3 as determined by an ATP assay) occurred at a slope site in June 1988 (550 m) in partially laminated sediments. In continuously laminated sediments from the central basin, foraminifera were found living (as determined by ATP assay) in October 1988 to depths of 4 cm, and specimens prepared for transmission electron microscopy were found with intact organelles to 3 cm, indicating their inhabitation of anoxic pore waters. Ultrastructural data from Nonionella stella is consistent with the hypothesis that this species can survive by anaerobic respiration. However, the benthic foraminifera appear unable to survive prolonged anoxia. The benthic foraminiferal population was completely dead in July 1989 when bottom water O₂ was undetectable.     

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.     

Rates and controls of anaerobic microbial respiration across spatial and temporal gradients in saltmarsh sediments

This study was undertaken to determine the rates and controls ofanaerobic respiration reactions coupled to organic matter mineralization as afunction of space and time along a transect from a bioturbated creekbank to themidmarsh in Georgia saltmarsh sediments. Sulfate reduction rates (SRR) weremeasured at 3 sites during 5 sampling periods throughout the growth season. Thesites differed according to hydrologic regime and the abundance of dominantplants and macrofauna. SRR and pore water / solid phase geochemistry showedevidence of enhanced sediment oxidation at sites exposed to intensebioturbation. Iron(III) reduction rates (FeRR) were directly determined insaltmarsh sediments for the first time, and in agreement with measured SRR,higher rates were observed at the bioturbated, unvegetated creekbank (BUC) andbioturbated, vegetated levee (BVL) sites in comparison to a vegetated mid-marsh(MM) site. An unexpected result was the fact that SRR varied nearly as muchbetween sites (2–3 x) as it did with temperature or season (3–4 x).The BVL site, vegetated by the tall form of Spartinaalterniflora, always exhibited the highest SRR and carbon oxidationrates (> 4000 nmol cm^–3 d^–1) with high activity levels extending deep ( 50 cm)into the sediment, while the MM site, dominated by the short form ofSpartina, always exhibited the lowest SRR which werelocalized to the top 15 cm of sediment. SRR and FeRR at BUC wereintermediate between those measured at the BVL and MM. Acetate was the mostabundant microbial fermentation product (concentrations up to > 1mM) in marsh porewaters, and its distribution reflectedrespirationactivity. Chemical exchange, caused by bioturbation, appeared to be the primarycontrol explaining trends in rates of sulfate and Fe(III) reduction withmacrophytes and carbon source acting as secondary controls.     

Benthic foraminiferal distribution and abundance changes in Skagerrak surface sediments: 1937 (Höglund) and 1992/1993 data compared

Both living (stained) and dead (unstained) foraminiferal assemblages from surface sediments (0–2 cm) in the northwestern part of the Skagerrak have been studied in order to (1) define and characterize the distribution of various modern benthic environments and (2) by comparing these findings with surface samples collected 40–60 years ago, to document possible faunal changes that might have occurred. The investigated area is physiographically divided into the Norwegian slope, the Skagerrak Basin, and the Danish slope. The latter is under the influence of the Jutland Current, while the basin and the investigated parts of the Norwegian slope are bathed in Atlantic water. All areas have bottom waters with a high oxygen concentration. Three living (stained) and three dead (unstained) assemblages occupy the three physiographic areas. Only one assemblage (on the Norwegian slope) is common to both the living and dead assemblages but the boundaries between them lie at comparable depths. The higher standing crops are found on the fertile Danish slope while the lower ones are in the deep basin where the diversity is at a maximum. In the dead assemblages, the relative abundance of agglutinated tests increases with depth. Comparison with data collected 40 to 60 years ago shows increases in absolute numbers of tests, especially in the deep basin. There are changes in assemblage compositions in all areas. The dominant species found in 1937 are different from those of 1992/1993. There is a major change in the basin where one agglutinated species has changed its depth distribution downslope and two present day abundant species are new arrivals. These faunal events are probably linked to environmental changes.     

Response of deep-sea benthic foraminifera to Miocene paleoclimatic events, DSDP Site 289

Changes in the Miocene deep-sea benthic foraminifera at DSDP Site 289 closely correlate to the climatically induced variations in deep and bottom waters in the Pacific Ocean. In early Miocene time, oxygen and carbon isotopes indicate that bottom waters were relatively warm and poorly oxygenated. Benthic foraminiferal assemblages are characterized by various species inherited from the Oligocene. Expansion of the Antarctic icecap in the early middle Miocene, 14–16 m.y. ago, increased oxygen isotope values, produced cold, more oxygenated bottom waters and lead to a turnover in the benthic foraminifera. An Oligocene—early Miocene assemblage was replaced by a cibicidoid-dominated assemblage. Some species became extinct and benthic faunas became more bathymetrically restricted with the increased stratification of deep waters in the ocean. In mid-Miocene time, Epistominella exigua and E. umbonifera, indicative of young, oxygenated bottom waters, are relatively common at DSDP Site 289. Further glacial expansion 5–9 m.y. ago lowered sealevel, increased oceanic upwelling and associated biological productivity and intensified the oxygen minima. Abundant hispid and costate uvigerines become a dominant faunal element at shallow depths above 2500 m as E. umbonifera becomes common to abundant below 2500 m. By late Miocene time, benthic faunas similar in species composition and proportion to modern faunas on the Ontong-Java plateau, had become established.     

The roles of elevation and salinity as primary controls on living foraminiferal distributions: Cowpen Marsh, Tees Estuary, UK

Benthic foraminiferal assemblages in subrecent deposits are commonly used to reconstruct past sea level. Interpretations are generally made by comparison with either modern dead or total (live plus dead) assemblages. In both cases there will have been post-mortem changes that have differentially affected preservation. It is therefore important to establish the primary ecological controls by analysis of the living assemblages. We have determined the spatial and temporal variability of intertidal benthic foraminifera in the surface (0–1 cm) sediments from a time series survey of 31 sampling stations at Cowpen Marsh, for a period of 12 months. We counted 112,067 live foraminifera assigned to 28 species. The fauna was dominated by two agglutinated species (Jadammina macrescens and Trochammina inflata) on the high and middle marshes, and three calcareous species (Elphidium williamsoni, Haynesina germanica and Quinqueloculina spp.) on the low marsh and tidal flat.The standing crop of the whole intertidal zone, including the high, middle, low marsh and tidal flat habitats, and the individual species varied both temporally and spatially. The standing crop of the intertidal zone as a whole was greatest in the summer months and showed a positive correlation with elevation. The standing crops of the high and middle marshes showed similar temporal variation with peaks in summer and autumn and a trough in winter. The low marsh showed numerous peaks and troughs of standing crop during the year, whereas the tidal flat showed a single peak in summer. The standing crops of Jadammina macrescens and Trochammina inflata on the high and middle marshes peaked from April to May and August to October with troughs in winter. These agglutinated species showed a strong correlation with elevation. Haynesina germanica peaked in May to August and November to January on the low marsh, whereas on the tidal flat there was a single peak in July. The standing crops of E. williamsoni on the low marsh and tidal flat were relatively high in June and May, and July, respectively. Quinqueloculina spp. peaked in May to July on the low marsh and July on the tidal flat. The species was also found in the middle marsh from July to May and high marsh from September to November. Haynesina germanica showed a strong negative correlation with elevation, whereas the other two dominant calcareous species demonstrated weak negative correlations with both elevation and salinity.Reconstructing former sea level depends primarily on the recognition of high and middle marsh assemblages and in this study these are shown to be strongly controlled by elevation rather than salinity. Caution may be needed in interpreting low marsh and tidal flat data as salinity plays a more important role here.     

The effect of bioturbation by polychaetes (Opheliidae) on benthic foraminiferal assemblages and test preservation

Biological activity such as burrowing can alter benthic foraminiferal shell preservation and may also modify benthic foraminiferal assemblages by vertical mixing, inducing sediment homogenization. Here, we analyse benthic foraminiferal assemblages and taphonomy of upper Miocene marine deposits from Conil de la Frontera (Cádiz, south‐western Spain). The deposits consist of marls displaying a pervasive alternation of intensively bioturbated beds dominated by Macaronichnus segregatis traces (ichnofabric index 4–5) and non‐bioturbated beds. Benthic foraminiferal assemblages are dominated by Cibicidoides mundulus and Cibicides refulgens, indicating that the marls were deposited on an oligotrophic, well‐oxygenated upper slope. The impact of burrowing on the preservation of benthic foraminiferal tests was tested using Q‐mode cluster analysis, which found two well‐differentiated groups of samples, one including the non‐bioturbated beds and the other encompassing the bioturbated ones. Fragmentation and recrystallization account for the differentiation of these groups, both being higher in the bioturbated sediments. Aggressive chemical digestion by the Macaronichnus trace‐makers, assumed to be a polychaete worm of the family Opheliidae, etched the microfossil shells, making them more vulnerable to fragmentation. Intense bioturbation favoured the circulation of pore fluids, encouraging recrystallization. Pervasive burrowing resulted in significant vertical reworking of microfossils. As a consequence, benthic foraminiferal assemblages in the bioturbated beds were homogenized in the mixed layer; that is, the uppermost layer of the substrate totally burrowed. The alternation of bioturbated and non‐bioturbated beds reflects episodic transfer of food particles down slope from shallower parts of the shelf as well as from the continent due to storms under otherwise homogeneous oligotrophic marine conditions.     

Distribution of recent benthic foraminifera on the Sunda Shelf (South China Sea)

In this study we investigate the species composition and spatial distribution patterns of Rose Bengal stained and unstained benthic foraminifera from the central part of the Sunda Shelf in the south-western South China Sea in relation to environmental factors. The uppermost centimetre of the surface sediments (> 150 μm) from 45 sites from inner (60 m) to outer shelf (226 m) water depths revealed 584 species including 443 stained species.The univariate analyses of individual species abundances and community parameters and next canonical correspondence analysis were used to relate the faunal data to a set of measured environmental parameters. Four biofacies recognised on the Sunda Shelf are most strongly correlated to water depth, primary production and sediment type of the habitat. The inner shelf biofacies (CCA cluster A), defined by Ammomassilina alveoliniformis and Asterorotalia pulchella, occurs in fine grained sediments classified as modern terrigenous mud in the region with the highest primary production values. The high-energy inner shelf biofacies (CCA cluster B), defined by Heterolepa dutemplei and Textularia lythostrota, occurs in modern terrigenous sand and silt dominated sediments, northeast from the Natuna Island. The high-energy outer shelf biofacies (CCA cluster C), defined by Cibicidoides pachyderma and Textularia bocki, is sandwiched between assemblages of biofacies D. It occurs in the region characterised by neritic relict sand. In the shallow-waters on the Sunda Shelf the relationship of benthic foraminiferal faunal composition to grain size of sediments indirectly signals the prevailing bottom hydrodynamic conditions. The dominance of the epibenthic foraminifera attached to bigger particles (e.g. Cibicides lobatulus, Planulina arimiensis) and much higher abundances of empty tests suggest greater current velocities northeast of Natuna Island. The outer shelf biofacies (CCA cluster D) is defined by Facetocochlea pulchra and Bulimina marginata. It occurs in an area covered with modern terrigenous silt and mud and is characterised by lower annual primary production, but seasonally influenced by weak upwelling.     

The effects of bioturbation on particle redistribution in Mediterranean coastal sediment. Preliminary results

In order to quantify bioturbation processes in a coastal Mediterranean ecosystem, experiments were performed to determine sediment mixing rates resulting from macrobenthos activity. Particle flux was measured in situ for 22 days using luminophores, which are colored sediment particles with sizes ranging from 10 to 200 µm.In sediment depths from 0–5 cm, particle mixing was intensive due to high macrobenthos abundance. A small quantity of luminophores was transported down to a depth of 14 ± 2 cm, where the macrofauna was represented principally by Polychetes. In a control experimental structure — without benthic fauna — no transfer of luminophores into the sediment was recorded.Sediment particle mixing measured in the ecosystem studied is intensive, and is the result of high macrobenthos activity. Different mixing modes occur with scales and rates depending on the organisms present. The luminophore profile resulting from bioturbation processes is explained by an intensive bioadvection sediment mixing added to a biodiffusive mixing with an order of magnitude of 10^–6 cm² s^–1. Tracer accumulations between 1 and 2 ± 1 cm and between 4 and 5 ± 1 cm are attributed to bioadvection activity of two or more distinct populations. Studies over a larger time scale have been undertaken to monitor developments in the observed subsurface maxima.     

The distribution of Recent benthic foraminifera in the Polar Front region, southwest Atlantic

Twenty-five core-top samples from the Maurice Ewing Bank (MEB) and Islas Orcadas Rise (IOR) were examined to determine the distribution of benthic foraminifera in the vicinity of the Polar Front in the southwest Atlantic Ocean. The Polar Front has a subsurface expression that effects the areal and depth distribution of benthic foraminifera in this region.Three faunal assemblages were identified by Q-mode factor analysis. The shallowest assemblage, dominated by Bulimina aculeata, is present from 1500 to 2600 m on the Maurice Ewing Bank and is associated with potential temperatures of 1.71-0.50°C, salinities of 34.74-34.70‰ and potential density values of 45.84–46.04 sigma-4. A second assemblage, dominated by Uvigerina peregrina, occurs in water depths from 2600 to 3100 m and is associated with potential temperatures of 0.40-0.26°C, salinities of 34.70-34.69‰ and potential density values of 46.05–46.07 sigma-4. The third assemblage is dominated by Nuttallides umbonifera, Ehrenbergina trigona and secondarily by Oridorsalis umbonatus and Pullenia bulloides (the N. umbonifera-E. trigona assemblage) is present form 2770 to 3120 m on the Islas Orcadas Rise. This assemblage is associated with potential temperatures of 0.36-0.14°C, salinities of 34.69-34.68‰, and potential density values of 46.06–46.09 sigma-4. Although the U. peregrina assemblage and the N. umbonifera-E. trigona assemblage overlap bathymetrically, they are present in waters of slightly different properties. The Bulimina aculeata assemblage is within the core of the Lower Circumpolar Deep Water (LCDW), while the other two assemblages occur within transition zones between the LCDW and Weddell Sea Deep Water (WSDW).The difference in the benthic foraminiferal assemblages at similar depths on the Islas Orcadas Rise and the Maurice Ewing Bank is the result of different water-mass regimes separated by the Polar Front.     

Stable oxygen and carbon isotope information on the establishment of a new, opportunistic foraminiferal fauna in a Swedish Skagerrak fjord basin, in 1979/1980

Significant faunal changes reported from recent, coastal environments, which are not directly influenced by urban and industrial impact, are rarely seen. In Gullmar Fjord on the Swedish west coast, a significant foraminiferal fauna change occurred in connection with severe low-oxygen conditions that evolved in the winter of 1979/1980. A foraminiferal fauna marked by common Skagerrak–Kattegat species, which had previously characterised the deep fjord basin, was replaced by the opportunistic, low-oxygen tolerant species Stainforthia fusiformis (Williamsson).To study this phenomenon further we performed stable oxygen and carbon isotope analyses on the indicator species itself, S. fusiformis, both on specimens from sediment cores representing approximately the last 85 years and on living (stained) individuals taken from a transect across the deep fjord basin. Our purpose was to detail how and why S. fusiformis, came to dominate the fauna.The oxygen isotope results suggest that salinities and temperatures in the deep basin have been relatively constant over the last c. 85 years, while the carbon isotopes show a significant change towards more negative values in association with the faunal shift of 1979/1980. The combined results from both the cores and the surface sediments suggest that S. fusiformis did not inhabit the deep basin until 1980. Before then, almost all specimens of S. fusiformis were small sized and their carbon isotope values suggest they were re-deposited shallow-water specimens that had been transported down to the central, deep basin as part of a suspension load. After a major faunal extinction in 1979–1980, S. fusiformis of all sizes suddenly appeared in large numbers and their carbon isotopic values were similar to the signal from registered in the recent, living fauna within the deep basin. This suggests that the opportunistic S. fusiformis established itself in the deep basin as a consequence of the severe low-oxygen event and the faunal crash of the previously dominating Skagerrak–Kattegat fauna.     

Benthic foraminiferal turnover across the Cretaceous/Paleogene boundary at Agost (southeastern Spain): paleoenvironmental inferences

We studied Upper Cretaceous and Lower Paleogene benthic foraminifera from the Agost section (southeastern Spain) to infer paleobathymetrical changes and paleoenvironmental turnover across the Cretaceous/Paleogene (K/P) transition. Benthic foraminifera indicate uppermost bathyal depths at Agost during the Abathomphalus mayaroensis Biochron (from about 400 kyr before the K/P boundary) through the early Plummerita hantkeninoides Biochron (about 120–150 kyr before that boundary). The depth increased to middle bathyal for the remainder of the Cretaceous, and remained so for the Danian part of the studied section (Parasubbotina pseudobulloides Biochron, at least 200 kyr after the K/P boundary). There were no perceivable bathymetrical changes at the K/P boundary, where 5% of the species became extinct, and the species composition of the benthic foraminiferal fauna changed considerably. Below the boundary, infaunal morphogroups constitute up to 65–73% of the faunas. Directly above the boundary, in the black clays of the lower Guembelitria cretacea Biozone, benthic foraminifera are rare. Several opportunistic taxa (e.g. the agglutinant Haplophragmoides sp.) have short peaks in relative abundance, possibly reflecting low-oxygen conditions as well as environmental instability, with benthos receiving food from short-lived, local blooms of primary producers. Above the clays through the end of the studied interval, epifaunal morphogroups dominate (up to 70% of the assemblages) or there is an even mixture or epifaunal and infaunal morphogroups. Infaunal groups do not recover to pre-extinction relative abundances, indicating that the food supply to the benthos did not recover fully over the studied interval (about 200 kyr after the K/P boundary). The benthic foraminiferal faunal changes are compatible with the direct and indirect effects of an asteroid impact, which severely destabilized primary producers and the oceanic food web that was dependent upon them.     

Age/size relations and food of two snailfishes,Liparis gibbus and Careproctus reinhardii (Teleostei,Liparididae) from Spitsbergen coastal waters

Summary The snailfish family Liparididae was well represented in bottom trawl hauls (205–370 m) from the Spitsbergen area, with Liparis gibbus being the dominant species. Careproctus reinhardti was less numerous. The present paper is based on 539 specimens of C. gibbus (size range 6–25cm) and 120 specimens of L. reinhardti (size range 5–26 cm). Both species showed a rather similar size and age distribution. The stomach contents reveal that both snailfish species feed both benthically and pelagically. Crustaceans, especially amphipods and decapods, were the most common prey items. With increasing size of the fish, the decapod Pandalus borealis becomes more important, particularly for L. gibbus. Juvenile Liparis (1.3–4.3 cm in length) were found pelagically (35–200 m), and their diet mainly consisted of copepods and young hyperiid amphipods.     

Spatial distribution of live benthic foraminifera in the Rhône prodelta: Faunal response to a continental–marine organic matter gradient

Benthic foraminifera were collected in the Rhône prodelta (Gulf of Lions, Mediterranean Sea), an enriched zone with high organic matter content. In June 2005, sediment cores were sampled at depths ranging from 20 to 100 m. Four distinct foraminiferal assemblages were determined in the study area, reflecting the geographical distribution of the impact of river supply. The living foraminiferal faunas present a typical picture, with strongly impoverished faunas composed exclusively of stress-tolerant taxa (Fursenkoina fusiformis, Bulimina aculeata, Leptohyalis scottii, and Adelosina longirostra) in the immediate vicinity of the river mouth. This assemblage is well adapted to a high input of continental organic matter and a minimum oxygen penetration depth into the sediment. To the southwest, under the main corridor followed by the river plume, high organic input with a dominantly terrestrial signature (more refractory) may be stressful for many taxa which need organic matter of a more labile quality. In this area, Nonion scaphum, Nonionella turgida and Rectuvigerina phlegeri are present in low densities. On the edge of this area, these taxa show much higher densities. A greater proportion of marine organic carbon could explain their increasing abundances in this area. Towards the east and towards the deepest stations, in the outer part of the enriched zone, biodiversity increases. Faunas at these stations have intermediate densities and contain a number of taxa (Cassidulina carinata, Epistominella vitrea, Valvulineria bradyana, Nonionella iridea/bradyi) at the deepest stations; Bolivina dilatata/spathulata and Textularia porrecta at the eastern stations) that seem to benefit from more marine organic matter. The comparison of geochemical measurements and foraminiferal data strongly suggests that the spatial distribution of foraminifera in the Rhône prodelta is mainly governed by the quality and the quantity of organic matter reaching the sediment–water interface. Since bottom waters are well oxygenated (215–260 µmol/L), and oxygen penetration into the sediment is less than 1 cm at all stations, benthic ecosystem oxygenation appears to have only a minor impact on regional differences in faunal distribution.     

Benthic foraminifera off West Africa (1°N to 32°S): Do live assemblages from the topmost sediment reliably record environmental variability?

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.     

Hydrologic,vegetation, and substrate characteristics of floating marshes in sediment-rich wetlands of the Mississippi river delta plain,Louisiana, USA

Floating marshes occur over 70% of the western Terrebonne Basin, Louisiana, USA, freshwater coastal wetlands. They are of several types: A free-floating thick-mat (45–60 cm) marsh dominated by Panicum hemitomon and Sagittaria lancifolia; a thick mat marsh dominated by Panicum hemitomon and Sagittaria lancifolia that floats part of the year, but whose vertical floating range is damped compared to adjacent water; and an irregularly-floating thin mat (< 30 cm) dominated by Eleocharis spp. in the spring and Ludwigia leptocarpa and Bidens laevis in the summer and fall. Floating mats must be almost entirely organic in order to be buoyant enough to float. The western Terrebonne wetlands receive large winter/spring supplies of suspended sediments from the Atchafalaya River. Even though sediment concentrations in the adjacent bayou are as high as 100 mg l^–1, the Panicum hemitomon/Sagittaria lancifolia free-floating marsh probably receives no over-surface sediments since it floats continuously. The bulk density data of the damped-floating marsh, however, suggest some mineral sediment input, probably during winter when this marsh is submerged. These two types of floating marsh could not have developed in the present sediment regime of the Atchafalaya River, but as long as they remain floating can continue to exist. Thin floating mats are found in areas receiving the least sediment (<20 mg 1^–1 suspended sediment concentration in adjacent bayous). This low sediment environment probably made possible their formation within the past 20 years. They may represent a transitional stage in mat succession from (1) existing thick-mat floating marsh to a degrading floating marsh, or (2) a floating marsh developing in shallow open water.Corresponding editor: D. Whigham     

Relationship between current velocity,depth and the invertebrate community in a stable river system

The relationship between invertebrate densities, current velocity and water depth was studied in the Dan River, northern Israel. Maximum current preferences ranged from 5–120 cm sec^–1, and depth preferences ranged from 5–60 cm. Thirty-five taxa of invertebrates were collected by means of colonization cages. Larval and adult stages of 3 Elmidae (Coleoptera) species were treated separately: Limnius letourneuxi, Grouvellinus caucasicus and Elmis rioloides. Differences in current preference were observed between larval and adult stages of the same species of Elmidae. Taxa were also grouped according to preference for turbulence. Wide ranges of depth and current velocity preferences were observed. Most of the taxa were found at between 80–100 cm sec^–1 and at depths of less than 30 cm. A correlation between species diversity and current velocity was established. Velocities of 60–80 cm sec^–1 contained the greatest overlap of faunal preference. The sensitivity of selected species to stream flow reduction is discussed.IES Laboratory, Department of Zoology, Hebrew University; and the Nature Reserves Authority