Abyssal community analysis from replicate box cores in the central North Pacific

A 0.25 m**2 United States Naval Electronics Laboratory box corer was used to take replicate samples from an oligotrophic bottom under the North Pacific Central Water Mass (~28°N, 155°W). The bottom is a red clay with manganese nodules at a depth of 5500-5800 m. Macrofaunal density ranges from 84 to 160 individuals per m**2 and is therefore much the same as in Northwest Atlantic Gyre waters. Of the macrofaunal taxa, polychaetes dominate (55 %), followed by tanaids (18 %), bivalves (7 %), and isopods (6 %). Meiofaunal taxa were only partially retained by the 297 µm screen used in washing. Even then, they are 1.5-3.9 times as abundant as the macrofaunal taxa, with nematodes being numerically dominant by far. Foraminifera seem to comprise an important portion of the community, but could not be assessed accurately because of the inability to discriminate living and dead tests. Remains of what are probably xenophyophoridans are also very important, but offer the same problem. Faunal diversity is extremely high, with deposit feeders comprising the overwhelming majority. Most species are rare, being encountered only once. The distributions of only three species show any significant deviation from randomness. The polychaete fauna from box cores collected from 90 miles to the north was not significantly different from that of the principal study locality. Concordance appeared at several taxonomic levels, from species through macrofaunal/meiofaunal relationships. As a result, the variation in total animal abundance shows aggregation among cores. We discuss Sokolova's concept of a deep-sea oligotrophic zone dominated by suspension feeders, and reconcile it with our present findings. The high diversity of the fauna combined with the low food level contradict theories that relate diversity directly with productivity.

Depth profiles of dissolved sulfide and sulfate in the pore water of hydrocarbon seep sediments offshore Pakistan

The interaction between fluid seepage, bottom water redox, and chemosynthetic communities was studied at cold seeps across one of the world's largest oxygen minimum zones (OMZ) located at the Makran convergent continental margin. Push cores were obtained from seeps within and below the core-OMZ with a remotely operated vehicle. Extracted sediment pore water was analyzed for sulfide and sulfate concentrations. Depending on oxygen availability in the bottom water, seeps were either colonized by microbial mats or by mats and macrofauna. The latter, including ampharetid polychaetes and vesicomyid clams, occurred in distinct benthic habitats, which were arranged in a concentric fashion around gas orifices. At most sites colonized by microbial mats, hydrogen sulfide was exported into the bottom water. Where macrofauna was widely abundant, hydrogen sulfide was retained within the sediment. Numerical modeling of pore water profiles was performed in order to assess rates of fluid advection and bioirrigation. While the magnitude of upward fluid flow decreased from 11 cm yr**-1 to <1 cm yr**-1 and the sulfate/methane transition (SMT) deepened with increasing distance from the central gas orifice, the fluxes of sulfate into the SMT did not significantly differ (6.6-9.3 mol m**-2 yr**-1). Depth-integrated rates of bioirrigation increased from 120 cm yr**-1 in the central habitat, characterized by microbial mats and sparse macrofauna, to 297 cm yr**-1 in the habitat of large and few small vesicomyid clams. These results reveal that chemosynthetic macrofauna inhabiting the outer seep habitats below the core-OMZ efficiently bioirrigate and thus transport sulfate down into the upper 10 to 15 cm of the sediment. In this way the animals deal with the lower upward flux of methane in outer habitats by stimulating rates of anaerobic oxidation of methane (AOM) with sulfate high enough to provide hydrogen sulfide for chemosynthesis. Through bioirrigation, macrofauna engineer their geochemical environment and fuel upward sulfide flux via AOM. Furthermore, due to the introduction of oxygenated bottom water into the sediment via bioirrigation, the depth of the sulfide sink gradually deepens towards outer habitats. We therefore suggest that - in addition to the oxygen levels in the water column, which determine whether macrofaunal communities can develop or not - it is the depth of the SMT and thus of sulfide production that determines which chemosynthetic communities are able to exploit the sulfide at depth. We hypothesize that large vesicomyid clams, by efficiently expanding the sulfate zone down into the sediment, could cut off smaller or less mobile organisms, as e.g. small clams and sulfur bacteria, from the sulfide source.

The distribution of macrobenthos in surface sediments at Fladen Ground stations, North Sea

A general study of structure, biomass estimates and dynamics on the macrofauna was carried out in August 1975 and March 1976 during PREFLEX (1975) and FLEX (1976), the Fladen Ground Experiment. On the basis of these data an attempt was made to estimate macrobenthic production expressed as minimum production (MP). The macrobenthic production is discussed together with meiobenthic annual production and with indirectly estimated microbenthic production in relation to an energy input from the water column of about 25 g C m**-2 year**-1. From the production estimates of the three benthic components a rough energy budget is proposed. Sampling was performed at five stations for endofauna twice during the time of investigation and for epifauna once. At each station two replicate box core samples (30 X 20 cm) were taken for endofauna. Epifauna was sampled with an Agassiz trawl once at each station. The total numbers of endofauna increased from station 1 to 5. This was valid as well for August 1975 (4,233-12,166 individuals per m**2 and 10 cm sediment depth) as for March 1976 (1,008-2,925 individuals). The polychaetes were the dominant organisms with a share of 33 to 62 %. The densities for the endofauna decreased from August 1975 to March 1976 by a mean factor of 2.8. Abundances of epifauna amounted to values between 11 and 102 individuals per 1000 m**2. The biomass dry weights (DWT) for macrobenthic endofauna varied between 0.97 g DWT m**-2 and 6.42 g DWT m**-2 in August 1975 and between 0.27 g DWT m**-2 and 2.64 g DWT m**-2 in March 1976. The mean amounted to 1.74 g DWT m**-2. Dry weights of epifauna biomass gave values between 4.9 and 83.1 g DWT * 1000 m**-2. The minimum production for the total macro-endofauna at Fladen Ground amounted to 1.43 g DWT m**-2 yr**-1 or 0.82 g C m**-2 yr**-1. This resulted in a minimum turnover rate (P/B) of 0.8. The share produced by the polychaetes amounted to 1.06g DWT m**-2 yr**-1 or 74 %.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Table 1: Station list with macrobenthos biomass results from the ocean floor

Among the Siberian shelf seas the Kara Sea is most strongly influenced by riverine runoff with nearly 1500 km fresh water discharge per year. This fresh water, discharged mainly by Ob and Yenisei, contains about 3.1 * 106 and 4.6 * 106 tons of total organic carbon per year, respectively (Gordeev et al. 1996). Little is known about the relevance of this organic material for biological communities, neither for the Kara Sea nor for the adjacent deep basins of the central Arctic Ocean. Aiming at elucidating the fate of fluvial matter transported from the rivers via estuaries into the central Arctic Ocean and the relative importance of marine organic matter being produced such information is crucial. Here we present calculations on the organic carbon demand of the Kara Sea macrozoobenthos based on measured biomass (total wet weight [ww] per 0.25 m ) from quantitative box corer samples and empirical relationships between biomass, annual production, annual respiration, and carbon remineralisation. This bottom-up approach may serve as a first estimate of the carbon remineralization potential of a given zoobenthos community (or area) as long as no data on in situ respiration rates are available. Our data basis comprises 54 stations sampled in summer seasons 1997, 1999 and 2000 in the Kara Sea at water depths between 10 and 68 m. The geographical area represented by stations analysed covers roughly 178 000 km**2, which is about one fifth of the total Kara Sea area. In this area, 290 species of invertebrate macrozoobenthos were identified with polychaeta, Crustacea, mollusca and echinodermata being the most abundant. For all stations analysed, mean biomass values ranged between 4.3 and 778.1 g ww/m**2 with organic carbon demands between 3.5 and 43.2 mg C/m**2/d. For the area of 178 000 km2 a preliminary total consumption of 1.4 * 10**6t Corg/y (equivalent to 21.5 mg C/m**2/d) was calculated for the macrozoobenthos. An extrapolation of our data would lead to an annual carbon demand of about 5-7 * 106 t for the whole Kara Sea macrozoobenthos (or 15.5-21.7 mg C/m2/d).

Benthos activity, abundance and biomass in surface sediments off Morocco, Northwest Africa

Macro- and meiobenthic abundance and biomass as well as metabolic activity (respiration, ETS activity) have been studied along a transect ranging from 130 to 3000 m water depth off northern Morocco (35° N) during "Meteor" cruise No. 53 (1980). The distribution of chloroplastic pigment concentration (chlorophyll a, pheophytins) in the sediment has been investigated as a measure of sedimented primary organic matter. High chloroplastic pigment concentrations were found on the shelf and around the shelf break, but values declined rapidly between 200 and 600 m depth. Below 1200 m pigment concentrations remained at a relatively uniform low level. Macrobenthic abundance and biomass (wet weight) decreased with increasing water depth and with distance from the shore. Significant changes occurred between the shelf and upper slope and below 2000 m depth. Meiobenthic abundance and biomass (ash free dry weight) followed the same general pattern, but changes were found below 400 and 800 m depth. In the depth range of 1200 to 3000 m values differ only slightly. Meiofauna abundance and biomass show a good correlation with the sedimentary chloroplastic pigment concentrations. Respiratory activity of sediment cores, measured by a shipboard technique at ambient temperatures, decreased with water depth and shows a good correlation with the pigment concentrations. ETS activity was highest on the shelf and decreased with water depth, with significant changes between 200 and 400 m, and below 1200 m depth, respectively. Activity was generally highest in the top 5 cm of the sediment and was measurable, at all stations, down to 15 cm sediment depth. Shelf and upper slope stations exhibited a vertical distribution pattern of ETS activity in the sediment column, different from that of deeper stations. The importance of biological activity measurements as an estimate of productivity is discussed. To prove the thesis that differences in benthic abundance, biomass and activity reflect differences in pelagic surface primary production, in the case of the NW-African coast caused by different upwelling intensities, the values from 35° N were compared with data from 21° N (permanent upwelling activity) and 17° N (ca. 9 months upwelling per year). On the shelf and upper slope (< 500 m) hydrographical conditions (currents, internal waves) influence the deposition of organic matter and cause a biomass minimum between 200 and 400 m depth in some regions. But, in general, macrobenthic abundance and biomass increases with enhanced upwelling activity and reaches a maximum in the area off Cape Blanc (21° N). On the shelf and in the shelf break region meiofauna densities are higher at 35° N in comparison to 21° N; but in contrast to the decreasing meiofauna abundance with increasing water depth at 35° N, an abundance maximum between 400 and 1200 m depth is formed in the Cape Blanc region; this maximum coincides with the maximum of sedimentary chloroplastic pigment equivalents. The comparison of ETS activities between 35° N and 21° N shows on the shelf activity at 21° N is up to 14 times higher and on the slope 4-9 times higher, which demonstrates that benthic activity responds to the surface productivity regime.

Meiofauna and the structure of the benthic community in the Iberian deep sea basin

1. On the cruises 3 and 15 of R.V. "Meteor" 6 grab samples, and 6 hauls with the 6 m Agassiztrawl were taken and at 2 stations the deep sea camera was lowered. This material gave quantitative results on the meiofauna and minimum counts of the macrofauna. 2. The nematodes constitute nearly 95% of the meiofauna, the copepoda only 2%. With increasing sediment depth the density of animals decrease gradually. In the uppermost centimeter of sediment 42.6% of the meiofauna are found while only 3.7% live in layer 6-7 cm. Meiofauna weight ranges from 0.6-5.7 mg/25 m**2 surface i.e. 0.24-2.8 g/m**2. 3. Mean numbers of individuals and weights show standard errors of 20-30 %. As an approximate average values for further considerations the weight of the meiofauna in the area was taken as 1 g/m**2 4. Quantitative information on the macrofauna is derived from the trawls and the photographs for the actinia Chitonanthus abyssorum only, which is found in the rate of 1 individual/36-72 m**2, but seems to be less abundant generally. 5. Animal density does not decrease steadily from nearshore to offshore biocoenoses, i.e. generally with increasing depth. The decrease is more pronounced for macro- than for meiofauna. For the deep sea the weight proportion of macrofauna : meiofauna is of the order of 1 : 1. 6. With the assumption, that adaptation of metabolism to deep sea conditions is similar in macro- and meiofauna total metabolism of invertebrates is ascribed to meiofauna to more than 80%. 7. The structure of the biocoenosis of the deep sea floor is characterized by the meiofauna living on and in the sediment and by the dominance of sediment feeders in the macrofauna. 8. Considering the large numbets and high partition rates of bacteria a comparative large part of the metabolism in the deep sea sediment must be ascribed to bacteria. This favours the hypothesis, that with increasing depth and decreasing addition of organic material to the sediment, the importance of meiofauna and microorganisms for total metabolism increases. 9. Considering the different modes of food transport to the deep sea environment, i.e. sinking of dead particles, transport by vertical migration of organisms, aggregation of organic particles, adsorption of dissoloved organic substance to inorganic particles, and heterotrophy, the sediment may be assumed to contain more food for invertebrates than the water above the bottom. 10. Suspensions feeders of macrofauna are fixed to hard substrates in the sediment surface. Some of them are shown to bend themselves down to the bottom in underwater photographs. This suggests the idea that some deep sea suspension feeders partly depend on food from the sediment surface, on which they feed directly.

Carbon analysis of surface sediments from the Laptev Se

During the past six years organic geochemical, micropaleontological, and sedimentological investigations were carried out within the framework of the multidisciplinary bilateral German-Russian research project ''System Laptev Sea'' and detailed biological investigations within the project ''German-Russian Investigations of the Marginal Seas of the Eurasian Arctic'', In order to understand the Laptev Sea ecosystem and to obtain information about sources and fate of organic carbon, the distribution of phyto- and zooplankton, diatoms, chlorophyll a benthic macrofauna, palynomorphs, grain size, total organic carbon, d13Corg and biomarkers (n-alkanes, fatty acids) were determined. In general, the influence of the major rivers draining into the Laptev Sea, is reflected in the water column as well as in the surface sediments. In both habitats three ecological provinces can be distinguished, i.e., the southeastern Laptev Sea, the central Laptev Sea, and the northern Laptev Sea. Additionally, clear differences between the western and the eastern Laptev Sea occur. The comparison of the different data sets of the water column and the surface sediments provide information about organic carbon sources and pathways in the Laptev Sea shelf and continental slope area.

Pigments of sediment cores of the Northwest European Continental Margin

In the context of the European OMEX Programme this investigation focused on gradients in the biomass and activity of the small benthic size spectrum along a transect across the Goban Spur from the outer Celtic Sea into Porcupine Abyssal Plain. The effects of food pulses (seasonal, episodic) on this part of the benthic size spectrum were investigated. Sediments sampled during eight expeditions at different seasons covering a range from 200 m to 4800 m water depth were assayed with biochemical bulk measurements: determinations of chloroplastic pigment equivalents (CPE), the sum of chlorophyll a and its breakdown products, provide information concerning the input of phytodetrital matter to the seafloor; phospholipids were analyzed to estimate the total biomass of small benthic organisms (including bacteria, fungi, flagellata, protozoa and small metazoan meiofauna). A new term 'small size class biomass' (SSCB) is introduced for the biomass of the smallest size classes of sediment-inhabiting organisms; the reduction of fluorescein-di-acetate (FDA) was determined to evaluate the potential activity of ester-cleaving bacterial exoenzymes in the sediment samples. At all stations benthic biomass was predominantly composed of the small size spectrum (90% on the shelf; 97-98% in the bathyal and abyssal parts of the transect). Small size class biomass (integrated over a 10 cm sediment column) ranged from 8 g C/m**2 on the shelf to 2.1 g C/m**2 on the adjacent Porcupine Abyssal Plain, exponentially decreasing with increasing water depth. However, a correlation between water depth and SSCB, macrofauna biomass as well as metazoan meiofauna biomass exhibited a significantly flatter slope for the small size classes in comparison to the larger organisms. CPE values indicated a pronounced seasonal cycle on the shelf and upper slope with twin peaks of phytodetrital deposition in mid spring and late summer. The deeper stations seem to receive a single annual flux maximum in late summer. SSCB and heterotrophic activity are significantly correlated to the amount of sediment-bound pigments. Seasonality in pigment concentrations is clearly followed by SSCB and activity. In contrast to macro- and megafauna which integrate over larger periods (months/years), the small benthic size classes, namely bacteria and foraminifera, proved to be the most reactive potential of the benthic communities to any perturbations on short time scales (days/weeks). The small size classes, therefore, occupy a key role in early diagenetic processes.