Surface Primary Production Research Articles

Deep-sea faunal provinces and their inferred environments in the Indian Ocean based on distribution of Recent benthic foraminifera

Multivariate analysis was performed on percentages of 46 species of unstained deep-sea benthic foraminifera from 131 core-top to near-core-top samples (322–5013 m) from across the Indian Ocean. Faunal data are combined with GEOSECS geochemical data to investigate any relationship between benthic foraminifera (assemblages and species) and deep-sea properties. In general, benthic foraminifera show a good correlation to surface productivity, organic carbon flux to the sea floor, deep-sea oxygenation and, to a lesser extent, to bottom temperature, without correlation with the water depths. The foraminiferal census data combined with geochemical data has enabled the division of the Indian Ocean into two faunal provinces. Province A occupies the northwestern Indian Ocean (Arabian Sea region) where surface primary production has a major maximum during the summer monsoon season and a secondary maximum during winter monsoon season that leads to high organic flux to the seafloor, making the deep-sea one of the most oxygen-deficient regions in the world ocean, with a pronounced oxygen minimum zone (OMZ). This province is dominated by benthic foraminifera characteristic of low oxygen and high organic food flux including Uvigerina peregrina, Robulus nicobarensis, Bolivinita pseudopunctata, Bolivinita sp., Bulimina aculeata, Bulimina alazanensis, Ehrenbergina carinata and Cassidulina carinata. Province B covers southern, southeastern and eastern parts of the Indian Ocean and is dominated by Nuttallides umbonifera, Epistominella exigua, Globocassidulina subglobosa, Uvigerina proboscidea, Cibicides wuellerstorfi, Cassidulina laevigata, Pullenia bulloides, Pullenia osloensis, Pyrgo murrhina, Oridorsalis umbonatus, Gyroidinoides (= Gyroidina) soldanii and Gyroidinoides cf. gemma suggesting well-oxygenated, cold deep water with low (oligotrophic) and pulsed food supply.

Composition of macrobenthos from the Central Indian Ocean Basin

The deep sea is well known for its high faunal diversity. But the current interest in its abundant polymetallic nodules, poses a threat to the little known benthic organisms surviving in this unique environment. The present study is the first attempt to document the Indian Ocean abyssal benthic diversity of macroinvertebrates and to investigate its relation to the surface primary production (chl-a), sediment labile organic matter, organic carbon and texture. The present study is based on 87 individuals. Altogether 39 macroinvertebrate genera were obtained from water depths of 4500–5500 m from 23 box cores. Reduction in macrobenthic density was seen towards the southern latitudes. The area was dominated by deposit feeding macrobenthos. Vertically, the fauna was distributed down to 30 cm depth, with the highest faunal density in the top 2–5 cm sediment section. The values for population density were strongly correlated with surface water chl-a and sediment protein, indicating supply of fresh organic matter as a critical factor for maintaining the deep sea benthic diversity and abundance.

Effects of biological and physical factors on seasonal oxygen dynamics in a stratified, eutrophic coastal ecosystem

Using a dual‐budget approach based on oxygen concentrations and stable isotopes, we examined primary production, respiration, and production to respiration ratios (P : R) across the Louisiana continental shelf to determine the relative roles of biological and physical factors in controlling seasonal oxygen dynamics. Regression models for δ18O and P :R indicated that the concentration of algal biomass explained most of the variability of the seasonal oxygen dynamics in surface waters. Physical factors, such as salinity, temperature, water‐column stability, and station depth were only of minor importance. Seasonal oxygen dynamics were more pronounced in bottom waters than in surface waters as hypoxia was common in bottom waters during summer months (May‐October). During cooler winter months (November‐April), oxygen concentrations throughout the water column were close to equilibrium with the atmosphere. The relative importance of benthic respiration to water‐column respiration in bottom waters was more pronounced during summer months when the water column was stratified. A strong physical disturbance caused by Hurricane Claudette during the summer of 2003 resulted in lower surface productivity relative to the calm 2001 and 2002 summertime conditions. Even though estimates of surface primary production and P :R were lower in July 2003 compared to July 2002, concentration of algal biomass (particulate organic carbon, C: N) remained the most important factor for d18O and P :R dynamics. The physical disturbance due to the hurricane, however, diminished the relative importance of benthic respiration, so that the system rather resembled wintertime conditions.

Impact of the subtropical mode water biogeochemical properties on primary production in the North Atlantic: New insights from an idealized model study

An idealized biophysical model of the North Atlantic was designed to investigate the setting and variability of the subtropical mode water (STMW) biogeochemical properties and its impact on surface primary production in the North Atlantic. The model solution first emphasizes that the exact timing of STMW formation versus the timing of the spring bloom is of primary importance for setting the STMW biogeochemical properties. The surface primary production reaches its maximum in March in the STMW formation region just before its subduction. Thus the spring bloom consumes nitrate at the surface before STMW subducts, and STMW leaves the upper layers depleted in nutrients and fueled in organic matter. This spring consumption explains the low nutrient content of STMW observed near its source region by J. B. Palter et al. (2005). Furthermore, the model suggests that STMW plays a key role in exporting dissolved organic matter (DOM) at subsurface. The spring bloom produces a significant amount of DOM sequestrated in the mode waters after its subduction. This large pool of DOM is then remineralized with time along the transit of STMW through the subtropical gyre. Consequently, the nutrient content of STMW increases as it moves away from its source region. Finally, the model shows also that STMW is very important in controlling primary production in the western boundary current (WBC) region. Indeed, STMW remains isolated from the surface along its trajectory within the subtropical gyre. It joins the mixed layer by obduction in the WBC region only. This nutrient‐rich old STMW irrigates and fertilizes the euphotic zone primarily in the WBC and then spreads along the boundary between the two gyres by advection.

Food-web structure and trophodynamics of mesopelagic–suprabenthic bathyal macrofauna of the Algerian Basin based on stable isotopes of carbon and nitrogen

The trophodynamics of mesopelagic (macrozooplankton/micronekton) and benthic boundary layer (suprabenthos=hyperbenthos) faunas from the Algerian Basin were characterized on a seasonal scale through stable carbon and nitrogen isotopic analyses of a total of 34 species and two broad taxa (Copepoda and Cumacea). This is the first study simultaneously focused on trophodynamics of deep-sea zooplankton and suprabenthos. Samples were collected southeast of Mallorca (Algerian Basin, Western Mediterranean), on the continental slope close to Cabrera Archipelago, at 650–780 m depths, ca. bi-monthly between August 2003 and June 2004. Mean δ 13C values of suprabenthos ranged from −21.1‰ ( Munnopsurus atlanticus) to −16.7‰ ( Cyclaspis longicaudata). Values of δ 15N ranged from 2.8‰ ( Lepechinella manco) to 9.9‰ (larvae of Gnathia sp.). The stable isotope ratios of suprabenthic fauna displayed a continuum of values, confirming a wide spectrum of feeding guilds (from filter feeders/surface deposit feeders to predators). According to the available information on diets for suprabenthic species, the highest annual mean δ 15N values were found for the hematophagous isopod Gnathia sp. parasite on fish (represented by Praniza larvae) and carnivorous amphipods (e.g. Rhachotropis spp., Nicippe tumida) consuming copepods, and the lowest δ 15N values were found for two cumaceans ( Cyclaspis longicaudata and Platysympus typicus) feeding on detritus. Assuming a 15N-enrichment factor of 2.5‰ and deposit feeders as baseline, we found three trophic levels in suprabenthic food webs. δ 13C ranges were particularly wide among deposit feeders (ranging from −21.8% to −17.3‰) and omnivores (from −20.5% to −18.8‰), suggesting exploitation of particulate organic matter (POM) of different characteristics. Our isotopic analyses revealed lower ranges of δ 13C and δ 15N for macrozooplankton/micronekton, compared with suprabenthos. δ 13C values of zooplankton taxa ranged from −21.1‰ (the hyperiid Phrosina semilunata) to −19.9‰ (the decapod Pasiphaea multidentata), while δ 15N values ranged from 3.9‰ ( P. semilunata) to 7.5‰ ( P. multidentata). Among zooplankton, more enriched δ 15N values were found among carnivores (e.g. the fish Cyclothone spp. and Pasiphaea multidentata) preying on copepods, hyperiids, euphausiids and small fish. The lowest δ 15N values were found for hyperiids that feed on the mucus nets of salps (e.g. Vibilia armata). After contrasting isotope analysis with dietary data, we conclude there were two trophic levels among zooplankton/micronekton. Strong correlation between the mean annual δ 15N and δ 13C values was found for zooplankton ( R 2=0.7), but not for suprabenthos, which suggests a single source of carbon for plankton. We found a general seasonal trend for δ 13C enrichment from late autumn (November) to late winter–spring (February–April) for both suprabenthos and zooplankton. The δ 13C enrichment in February–April was correlated in zooplankton with higher surface chlorophyll a concentration 1 month before sampling. As evidenced by δ 13C–δ 15N correlations, the response of zooplankton to the peak of surface primary production was almost immediate (an increase of δ 13C–δ 15N correlations in February), and stronger than for suprabenthos. The response among suprabenthos was weak, with slight increase in δ 13C–δ 15N relationships in April–June.

Regional patterns of nematode assemblages in the Arctic deep seas

Surface primary productivity and carbon flux in the Arctic Seas are higher along the warm Spitsbergen Water Current than along the ice-infested East Greenland Current. These contrasting oceanographic conditions are reflected in the deep-sea environment and may shape nematode assemblages. However, the paucity of samples in the Arctic deep seas precludes any regional scale assessment. In the present study, nematode assemblages were investigated in relation to a range of environmental variables along the 2,000 m isobath between latitudes 72°N and 79°N for both East and Western margins of the northern North Atlantic. Results showed that both margins had distinct environmental characteristics, with respect to chloroplastic pigments, sediment water content, sediment-bound organic matter, phospholipids and particulate proteins. Nematode assemblages varied according to these environmental changes. Along the more oligotrophic western margin, chloroplastic pigments increased towards the North, while the other environmental variables, nematode abundances and species richness decreased. In contrast, along the eastern margin, we observed higher quantities of organic matter and particulate protein, which supported higher abundance and species richness. Nematode assemblages along both margins varied according to food availability with species composition more variable in areas with lower amounts. Seventy percent of the species occurred in both margins indicating a low turnover of species. The present results support the hypothesis of a positive latitudinal gradient across the North Atlantic and further suggest that contemporary climate and recent ecological processes may predict nematode diversity patterns at larger scales.

Seasonal and spatial variability of coccolithophore export production at the South-Western margin of Crete (Eastern Mediterranean)

Six moorings were deployed at different locations in the deep submarine canyons along the south–west margin of Crete, providing a total of eight sediment-trap time series from June 2005 to May 2006. Within this dataset, we analyzed the record from intact coccospheres, which represent the signal of export production from the coccolithophore community. The most abundant species at all stations during the whole investigated period were E. huxleyi and A. robusta, followed by S. pulchra HET, G. flabellatus, H. carteri, F. profunda, S. pulchra HOL oblonga, while the rest of the species represented ≤ 1% of the assemblage. Overall the assemblage composition was comparable at all stations, with slight variations mostly related to the different preservation of coccosphere integrity at the different collection depths. The consistent pattern of seasonal variation in species distribution and total coccolithophore export allowed us to define the occurrence of three main periods: a) March to June, with high overall coccosphere flux (up to 4.3 × 10 5–3.4 × 10 6 coccospheres m − 2 day − 1 ), increased abundance of E. huxleyi and subordinate H. carteri s.s., Umbilicosphaera spp. and S. pulchra; b) June to November, with high but gradually decreasing total coccosphere flux (up to 7 × 10 5–1.4 × 10 6 coccospheres m − 2 day − 1 ) and high relative abundance of the deep photic zone species A. robusta, F. profunda, G. flabellatus as well as S. pulchra and Coronosphaera spp., R. clavigera, U. tenuis, D. tubifera and holococcolithophores; c) November to February, with low overall export fluxes (3.5–9 × 10 4 coccospheres m − 2 day − 1 ) and high relative abundance of A. robusta, S. pulchra and Syracosphaera spp. These three periods correspond to the seasonal changes in sea surface temperature, surface mixed layer depth and rainfall and are associated with varying total surface primary production, as detected through remote sensing in the surface waters.

13C‐isotope analyses reveal that chemolithoautotrophicGamma‐ andEpsilonproteobacteriafeed a microbial food web in a pelagic redoxcline of the central Baltic Sea

Marine pelagic redoxclines are zones of high dark CO(2) fixation rates, which can correspond up to 30% of the surface primary production. However, despite this significant contribution to the pelagic carbon cycle, the identity of most chemolithoautotrophic organisms is still unknown. Therefore, the aim of this study was to directly link the dark CO(2) fixation capacity of a pelagic redoxcline in the central Baltic Sea (Landsort Deep) with the identity of the main chemolithoautotrophs involved. Our approach was based on the analysis of natural carbon isotope signatures in fatty acid methyl esters (FAMEs) and on measurements of CO(2) incorporation in (13)C-bicarbonate pulse experiments. The incorporation of (13)C into chemolithoautotrophic cells was investigated by rRNA-based stable isotope probing (RNA-SIP) and FAME analysis after incubation for 24 and 72 h under in situ conditions. Our results demonstrated that fatty acids indicative of Proteobacteria were significantly enriched in (13)C slightly below the chemocline. RNA-SIP analyses revealed that two different Gammaproteobacteria and three closely related Epsilonproteobacteria of the Sulfurimonas cluster were active dark CO(2)-fixing microorganisms, with a time-dependent community shift between these groups. Labelling of Archaea was not detectable, but after 72 h of incubation the (13)C-label had been transferred to a potentially bacterivorous ciliate related to Euplotes sp. Thus, RNA-SIP provided direct evidence for the contribution of chemolithoautotrophic production to the microbial food web in this marine pelagic redoxcline, emphasizing the importance of dark CO(2)-fixing Proteobacteria within this habitat.

Parameterization of surface irradiance and primary production in Århus Bay, SW Kattegat, Baltic Sea

The aims of the present study were to develop a parameterization of a one-year-long observed PAR time-series, apply the PAR parameterization in a primary production relation, and compare calculated and observed time-series of primary production. The PAR parameterization was applied in the generally used relation for the primary production (P d): P d = a(BI 0 Z 0) + b with observed photic depth (Z 0) and Chl-a concentrations (B). It was tested whether the PAR parameterization in combination with this simple relation for primary production was able to describe the actual measured primary production. The study is based on a one year long time-series of PAR, CTD-casts (n = 45), and primary production measurements (n = 24) from Arhus Bay (56°09′ N; 10°20′ E), south west Kattegat. Results showed a high and positive correlation between observed and calculated primary production in the bay, as based on the present PAR parameterization combined with the simple primary production relation. The developed PAR parameterization, which calculates total daily surface irradiance per day (M photons m−2 d−1), can be applied in any ecological application taking into account that it was developed for the latitude of 56° N.

The feeding and diet of the deep-sea shrimp Aristeus antennatus off the Balearic Islands (Western Mediterranean): Influence of environmental factors and relationship with the biological cycle

Spatio-temporal variation of feeding intensity and diet in the red shrimp Aristeus antennatus was studied at two locations around the island of Mallorca (Balearic Islands, Western Mediterraean) in August, September, and November 2003, and in February, April and June 2004 at depths between 550 and 750 m. The two areas, with different oceanographic conditions, were respectively located in the northwest (Sóller) and the south (Cabrera) of Mallorca. Off Sóller, feeding intensity of A. antennatus showed a significant increase from February to April and June 2004 in all the three size-classes studied (small shrimps: CL < 30 mm; medium: CL between 30 and 40 mm; large: CL ⩾ 40 mm). Off Cabrera, the highest fullness was recorded in November 2003 among small and medium shrimp, while only large specimens showed patterns similar to that found off Sóller. Off Sóller, the diet of both small (CL < 34 mm) and large (CL ⩾ 34 mm) A. antennatus was mainly influenced by season, with three dietary groups corresponding to August–September 2003, to November 2003/February 2004, and to hauls from April to June 2004. Off Cabrera, hauls (representing diets) were grouped by depth, never by season. The most remarkable seasonal shift in the diet of A. antennatus off Sóller was the increase of mesopelagic prey in April–June relative to other months. In all size categories there was an increase off Sóller in the energy intake of prey ingested from February to June 2004, an increase not found off Cabrera. Degree of digestion of mesopelagic prey indicated nocturnal feeding on mesopelagic fauna. These prey probably have a shallower depth distribution at night than found in our daylight sampling, and possible migratory movements among prey and A. antennatus at night would explain the lack of correlation between prey abundance in guts and in the environment found during daylight periods for most micronekton mesopelagic prey (euphausiids, myctophids and sergestids). Off Sóller, fullness and diet were significantly linked to temporal changes in water column productivity (e.g., Chl a readings, fluorescence) and to changes in the shrimp biology (lipid content of hepatopancreas, Gonado-somatic Index, GSI). Off Cabrera, we found a higher dependence of fullness and diet with T and S, both variables in turn related to depth. The increase of stomach fullness and dietary energy intake in pre-reproductive females from February to April–June 2004 found off Sóller, coupled with the consumption of mesopelagic prey, was parallel to a significant increase of the gonad weight (GSI, fecundity) in June. Most individuals attain gonad development in the period May–June, after two months of the peak of primary production at the surface. The strong link found between pelagic resources and reproductive processes in a deep-sea species such as the shrimp Aristeus antennatus, situated near the top of the trophic web, suggests a rapid energy flow via mesopelagic fauna between surface primary production and bathyal megabenthic communities at oligotrophic insular areas. In contrast to mainland areas off the Catalan coasts submitted to the influence of submarine canyons, around the island of Mallorca the empoverishment of benthos biomass may enhance consumption of micronektonic prey and a possible accumulation of pre-reproductive females of A. antennatus in areas (e.g., steep slopes and persistent frontal systems found off Sóller) with high zooplankton aggregations.

Testing the FOODBANCS hypothesis: Seasonal variations in near-bottom particle flux, bioturbation intensity, and deposit feeding based on 234Th measurements

Abstract Naturally occurring 234Th (24-d half-life) was used on the West Antarctic continental shelf to evaluate temporal variations in the flux of particulate material reaching the seabed, bioturbation intensity, the seasonal continuity of feeding by benthic fauna, and trends in particle selection during ingestion for six common detritivores (four surface deposit feeders and two subsurface deposit feeders). These measurements were made at three stations during the five FOODBANCS cruises (December 1999, March, June, and October 2000, and March 2001) to assess the nature of pelagic–benthic coupling on the shelf and to evaluate the seabed as a potential food bank for deposit feeders when surface primary production is minimal. Two summer regimes were sampled (March 2000 and March 2001) with the latter exhibiting a distinct 1–2-cm-thick phytodetritus layer in nearly all sediment core samples. At site B, the 234Th fluxes into the near-bottom (150/170 mab) sediment traps were indistinguishable for the December–March 2000, March–June 2000, and June–October 2000 sampling intervals (fluxes ranging from 170 to 280 dpm m−2 d−1). However, the sediment-trap 234Th flux measured for the October 2000–March 2001 interval (1000 dpm m−2 d−1) was ∼5-fold greater than during the other three sampling periods, consistent with the deposition of a phytodetritus layer. The steady-state 234Th fluxes derived from seabed inventories at site B were 2.4–2.7 times greater than the sediment-trap 234Th fluxes, indicating substantial scavenging of this particle-reactive radiotracer in the bottom 150 m of the water column and/or lateral transport near the seabed. The seabed 234Th inventories at the three stations showed no variation during the first four cruises, but were significantly greater during cruise FB-V (March 2001), when the phytodetritus layer occurred. Based on 234Th distributions in the seabed, bioturbation intensities (quantified using the diffusive mixing coefficient, Db) varied from 0.5 to 97 cm2 yr−1. Mixing coefficients were significantly lower during FB-II than during FB-III, but no other time comparisons were statistically significant (p>0.05). 234Th Dbs showed no distinct seasonality, and no correlation with either organic carbon flux into the sediment traps or 234Th inventory in the seabed. The four surface deposit feeders examined, an echiuran worm and three holothurians (Protelpidia murrayi, Bathyplotes fuscivinculum, and Peniagone vignoni), exhibited greater particle selection for recently deposited sediment during ingestion than the two subsurface deposit feeders studied (a head-down deposit feeding holothurian, Molpadia musculus, and an irregular urchin, Amphipneustes lorioli). All six deposit feeders contained excess 234Th activity in gut sediments during all five cruises, indicating sediment ingestion year round, even during the austral winter. The lack of seasonal variation in bioturbation intensity and the demonstration of year-round feeding in deposit feeders are consistent with the hypothesis that the seafloor sediments accumulate labile organic matter produced during periods of high primary production and that deposit feeders utilize this food source as a food bank on a year-round basis.

Phytoplankton production characteristics in the Eastern Atlantic and Atlantic sector of the Southern Ocean in October–November 2004

Production parameters of surface phytoplankton were measured along three transects: La Manche-Cape Town (I); Cape Town-54°S (II); 0°-49°W (along 54°S) (III). The Canary upwelling waters were most productive along transect I, where the surface chlorophyll a (Chl 0) and the surface primary production (PP 0) were as high as 4.3 mg/m3 and 173 mg C/m3 per day, respectively. Mosaic patterns in the distribution of these parameters were recorded in the northeastern regions of the South Subtropical Anticyclonic Gyre (Chl 0 = 0.03–0.35 mg/m3; PP 0 = 1.6–12.6 mg C/m3 per day). Along transect II, the average twofold southward increase in Chl 0 (from 0.2 to 0.4 mg/m3) and the concurrent decline of the phytoplankton assimilation activity ( AN 0) resulted in deviations from typical latitudinal changes inPP 0. At most sites, PP 0 values varied between 6 and 15 mg C/m3 per day. Negligible changes in Chl 0 (0.36–0.85 mg/m3), PP 0 (8–19 mg C/m3 per day), and AN 0 (0.7–1.6 mg C/mg chl a per hour) were registered for the oceanic waters along transect III. Along all the transects, PP 0 depended on Chl 0 to a greater extent than AN 0. The values of the latter parameter were largely determined by the water temperature and showed a slight correlation with the insolation. Along transect II, the integrated primary production (PP int) and the layer-integrated chlorophyll a in the upper 200 m (Chl 0–200) generally varied from 180 to 360 mg C/m2 per day and from 30 to 70 mg/m2, respectively. In the Polar Front region, an increase in Chl 0–200, PP int, Chl 0, and PP 0 up to respective values of 190 mg/m2, 520 mg C/m2 per day, 1.2 mg/m3, and 32 mg C/m3 per day was observed. A comparison of the water column (0–100 m) stability with the vertical distribution of the primary production and chlorophyll content along transect II implies that the thick (>100 m) upper mixed layer (UML) formed in response to the strong water cooling and wind forcing was largely responsible for the limited primary production in the Subantarctic and Antarctic regions. The large UML thickness resulted in an intense removal of plant cells from the photosynthetic layer and light starvation of a significant (up to 60%) part of the phytoplankton community.

Macrofaunal diversity in the Central Indian Ocean Basin

The deep-sea is characterized by four stable physical variables: hydrostatic pressure, temperature, salinity and oxygen. This harsh environment is comprised of diverse benthic faunal groups. Presently these organisms are at threat due to the increasing interest of mankind in the non-living resources and destructive deep-sea fishing practices present in these areas. The polymetallic nodule is one such resource, looked upon as an alternative to land-based minerals. The Central Indian Ocean Basin (CIOB) abyss is densely covered with these nodules. Commercial mining can cause danger to the species present, some of which may be as yet unknown to science. The deep-sea fauna in the CIOB is the least studied among the three oceans, hence, the aim of the current work is to investigate the composition and feeding patterns of the macrofauna in the CIOB. A total of 27 species were obtained in the current study with a dominance of polychaetes. The CIOB was composed of deposit feeding macrofauna, this is supported by the positive correlation between fauna and sediment organic carbon (p<0.05). Compared to the other oceans, the density of macrofauna is low in CIOB, since it is sustained by low surface primary productivity.

Primary production and chlorophyll distributions in the subtropical and tropical waters of the Atlantic Ocean in the autumn of 2002

In October and November 2002, high and relatively high values of the chlorophyll a concentration at the sea surface (Cchl) were observed in the English Channel (0.47 mg/m3), in the waters of the North Atlantic Current (0.25 mg/m3), in the tropical and subtropical anticyclonic gyres (0.07–0.42 mg/m3), and also in the southwestern region of the southern subtropical anticyclonic gyre (usually 0.11–0.23 mg/m3). The central regions of the southern subtropical anticyclonic gyre (SATG) and the North Atlantic tropical gyre (NATR) were characterized by lower values of Cchl (0.02–0.08 mg/m3 for the SATG and 0.07–0.14 mg/m3 for the NATR). At most of the SATG stations, the values of the surface primary production (Cphs) varied from 2.5 to 5.5 mg C/m3 per day and were mainly defined by the fluctuations of Cchl (r = +0.78) rather than by those of the assimilation number (r = +0.54). The low assimilation activity of phytoplankton in these waters (1.3–4.6 mg chl a per hour) pointed to a lack of nutrients. An analysis of the variability of their concentration and the composition of photosynthetic pigments showed that, in the waters north of 30° N, the growth of phytoplankton was mostly restricted by the deficiency of nitrogen, while, in more southern areas, at the majority of stations (about 60%), the phosphorus concentrations were the minimum. At the low concentrations of nitrates and nitrites, ammonium represented itself as a buffer that prevented planktonic algae from extreme degrees of nitric starvation. In the tropical waters and in the waters of the SATG, the primary production throughout the water column varied from 240 to 380 mg C/m2 30° per day. This level of productivity at stations with low values of Cchl (<0.08 mg/m3) was provided by a well-developed deep chlorophyll maximum and a high transparency of the water. The light curves of photosynthesis based on in situ measurements point to the high efficiency of utilizing the penetrating solar radiation by phytoplankton on cloudy days.

Estimates of atmospheric dry deposition and associated input of nutrients to Gulf of Aqaba seawater

Dry deposition rates and associated inputs of soluble inorganic nitrogen (N) phosphorus (P) and iron (Fe) were calculated for the north coast of the Gulf of Aqaba in Eilat, Israel, between August 2003 and September 2005. The main inorganic N compounds in the water soluble fraction of aerosol particles were nitrate (60%) and ammonium (38%), with mean concentrations of 35 and 22 nmol m−3 of air, respectively. Soluble phosphate ranged between 0.09 and 2 nmol m−3 of air with a mean value of 0.4 nmol m−3 of air. The soluble inorganic nitrogen and soluble phosphate account for approximately 86% and 69% of total soluble nitrogen and total soluble phosphorus, respectively; the difference is assigned to organic N and P compounds. The mean concentration of soluble aerosol Fe was 0.3 nmol m−3 of air. Our measurements of the soluble nutrient concentrations are comparable to data previously reported for the eastern Mediterranean area. Dry deposition fluxes of nutrients were estimated for each sampling date using a size‐dependent deposition model. The estimated fluxes were highly variable over the sampling period with the mean fluxes of 38, 0.2 and 0.02 μmol m−2 d−1 for seawater soluble inorganic N, P and Fe, respectively. The soluble phosphate flux shows a seasonal pattern with higher input during the winter (September to December) than in other seasons. The inorganic N/P molar ratios in the seawater‐soluble fraction of the dry deposition (ranging from 32 to 541) were well above the Redfield ratio (N/P = 16), suggesting that atmospheric inputs of nutrients increase the likelihood for P limitation in the Gulf. Atmospheric deposition could contribute a substantial fraction (35%) of dissolved inorganic N to the euphotic zone during the stratification period (April to October), and the N flux from this source could support over 10% of surface primary production and possibly all of the new production during the summer; however, these estimates have a relatively large uncertainty due to error associated with deposition flux calculation and the temporal variability in dust flux. Atmospheric input of seawater‐soluble Fe is in large excess compared to that required for the phytoplankton growth driven by the N deposition.

Influence of basin-scale and mesoscale physical processes on biological productivity in the Bay of Bengal during the summer monsoon

Physical forcing plays a major role in determining biological processes in the ocean across the full spectrum of spatial and temporal scales. Variability of biological production in the Bay of Bengal (BoB) based on basin-scale and mesoscale physical processes is presented using hydrographic data collected during the peak summer monsoon in July–August, 2003. Three different and spatially varying physical processes were identified in the upper 300 m: (I) anticyclonic warm gyre offshore in the southern Bay; (II) a cyclonic eddy in the northern Bay; and (III) an upwelling region adjacent to the southern coast. In the warm gyre (>28.8 °C), the low salinity (33.5) surface waters contained low concentrations of nutrients. These warm surface waters extended below the euphotic zone, which resulted in an oligotrophic environment with low surface chlorophyll a (0.12 mg m −3), low surface primary production (2.55 mg C m −3 day −1) and low zooplankton biovolume (0.14 ml m −3). In the cyclonic eddy, the elevated isopycnals raised the nutricline upto the surface (NO 3–N > 8.2 μM, PO 4–P > 0.8 μM, SiO 4–Si > 3.5 μM). Despite the system being highly eutrophic, response in the biological activity was low. In the upwelling zone, although the nutrient concentrations were lower compared to the cyclonic eddy, the surface phytoplankton biomass and production were high (Chl a – 0.25 mg m −3, PP – 9.23 mg C m −3 day −1), and mesozooplankton biovolume (1.12 ml m −3) was rich. Normally in oligotrophic, open ocean ecosystems, primary production is based on ‘regenerated’ nutrients, but during episodic events like eddies the ‘production’ switches over to ‘new production’. The switching over from ‘regenerated production’ to ‘new production’ in the open ocean (cyclonic eddy) and establishment of a new phytoplankton community will take longer than in the coastal system (upwelling). Despite the functioning of a cyclonic eddy and upwelling being divergent (transporting of nutrients from deeper waters to surface), the utilization of nutrients leading to enhanced biological production and its transfer to upper trophic levels in the upwelling region imply that the energy transfer from primary production to secondary production (mesozooplankton) is more efficient than in the cyclonic eddy of the open ocean. The results suggest that basin-scale and mesoscale processes influence the abundance and spatial heterogeneity of plankton populations across a wide spatial scale in the BoB. The multifaceted effects of these physical processes on primary productivity thus play a prominent role in structuring of zooplankton communities and could consecutively affect the recruitment of pelagic fisheries.

Bacterial abundance and production in the western Black Sea

Abstract The unusual stability and vertical extent of the oxic–anoxic interface of the central Black Sea water column facilitate studying microbially mediated redox processes. Bacterial abundance, cell volume, 3 H-thymidine and -leucine incorporation were measured in the Black Sea during May–June 2001. Sampling was conducted along a shelf–gyre transect, and was focused at the suboxic–anoxic interface at the deep stations. Bacterial variables (abundance, incorporation rates, biomass, production, and specific growth rates) were significantly higher on the northwest shelf compared to the Southwestern Gyre, and were generally intermediate at shelf-break stations. Mean bacterial abundances ranged from 1.4 to 2.9×10 9 cells l −1 in the mixed layer along the shelf–gyre transect, while isotope incorporation rates ranged from 7 to 74 pM-TdR h −1 and 60 to 209 pM-Leu h −1 . Bacterial biomass ranged from 6.7 to 13.5 μg C l −1 , bacterial production rates ranged from 2.3 to 7.7 μg C l −1 d −1 , and specific growth rates ranged from 0.2 to 1.5 d −1 . In the Southwestern Gyre, surface bacterial production was equivalent to about 50% of 14 C-primary production. In the oxycline layer, mean bacterial abundance (0.38×10 9 cells l −1 ), leucine incorporation (4.1 pM-Leu h −1 ), and bacterial production (0.15 μg C l −1 d −1 ) were significantly greater compared to the suboxic and anoxic layers, while cell volume and biomass increased with depth. Large filamentous bacteria (>10 μm in length) were present below the suboxic boundary, and comprised up to 10% of the total bacterial abundance and up to 53% of the total bacterial biomass. Approximately 60% of the surface primary production was consumed above the suboxic–anoxic interface.

Prokaryotic respiration and production in the meso- and bathypelagic realm of the eastern and western North Atlantic basin

We measured prokaryotic production and respiration in the major water masses of the North Atlantic down to a depth of ,4,000 m by following the progression of the two branches of North Atlantic Deep Water (NADW) in the oceanic conveyor belt. Prokaryotic abundance decreased exponentially with depth from 3 to 0.4 3 105 cells mL21 in the eastern basin and from 3.6 to 0.3 3 105 cells mL21 in the western basin. Prokaryotic production measured via 3H-leucine incorporation showed a similar pattern to that of prokaryotic abundance and decreased with depth from 9.2 to 1.1 mmol C m23 d21 in the eastern and from 20.6 to 1.2 mmol C m23 d21 in the western basin. Prokaryotic respiration, measured via oxygen consumption, ranged from about 300 to 60 mmol C m23 d21 from ,100 m depth to the NADW. Prokaryotic growth efficiencies of ,2% in the deep waters (depth range 1,200–4,000 m) indicate that the prokaryotic carbon demand exceeds dissolved organic matter input and surface primary production by 2 orders of magnitude. Cell-specific prokaryotic production was rather constant throughout the water column, ranging from 15 to 32 3 1023 fmol C cell21 d21 in the eastern and from 35 to 58 3 1023 fmol C cell21 d21 in the western basin. Along with increasing cell-specific respiration towards the deep water masses and the relatively short turnover time of the prokaryotic community in the dark ocean (34–54 d), prokaryotic activity in the meso- and bathypelagic North Atlantic might be higher than previously assumed.

Spatial variability of meiofaunal communities at areas of contrasting depth and productivity in the Aegean Sea (NE Mediterranean)

The abundance and community structure of metazoan meiofauna were studied in deep-sea sediments from the north and south Aegean Sea (Eastern Mediterranean) in summer 1997 and spring 1998. The two areas varied in their surface primary productivity with the northern area being more productive. Meiofaunal densities displayed strong spatial variability while no temporal changes were observed. Total metazoan density ranged from 128 to 1251 ind./10 cm 2, with significantly higher values in the north. Meiofaunal biomass ranged from 27 to 391 μgC/10 cm 2 with higher values also in the north. At all stations nematodes dominated the community, comprising on average more than 91% of the total abundance. On a broad scale, meiofaunal densities displayed a positive correlation with food availability (sediment–bound chloroplastic pigments, carbohydrates and lipids); carbon mineralization (an indicator of organic matter turnover) was significantly higher in the northern Aegean, providing evidence of high organic–matter input and intense benthic–pelagic coupling. The spatial structure of the nematode community indicated that the two areas were similar in terms of their dominant genera ( Halalaimus, Acantholaimus, and Thalassomonhystera).

Rivers as groundwater-dependent ecosystems: a review of degrees of dependency, riverine processes and management implications

Many rivers are classified as groundwater-dependent ecosystems (GDEs), owing to the contribution of groundwater to their base flow. However, there has been little explicit recognition of the way groundwater influences riverine biota or processes, how degrees of ecological dependency may vary, and the management implications of this dependency. The permeable beds and banks of these GDEs where surface water and groundwater exchange are termed ‘hyporheic zones’. They are often inhabited by invertebrates, with varying reliance on groundwater, although the ecological roles of these invertebrates are little known. Upwelling hyporheic water can promote surface primary productivity, influence sediment microbial activity, and affect organic matter decomposition. In many intermittent streams, variable groundwater inputs alter the duration of flow or water permanence, and the duration and timing of these largely govern the biota and rates of many ecosystem processes (e.g. leaf decomposition). Not only is the physical presence of water important, thermal and chemical conditions arising from groundwater inputs also have direct and indirect effects on riverine biota and rates or types of in-stream processes. Differing degrees of dependency of rivers on groundwater mediate all these influences, and may change over time and in response to human activities. Alteration of groundwater inputs through extraction from riparian wells or changes in local water table have an impact on these GDEs, and some current management plans aim to restrict groundwater extraction from near permeable river channels. However, these are often ‘blanket’ restrictions and the mechanisms of GDE dependency or timing of groundwater requirements are poorly understood, hampering refinement of this management approach. More effective management of these GDEs into the future can result only from a better understanding of the mechanisms of the dependency, how these vary among river types and what in-stream changes might be predicted from alteration of groundwater inputs.