Processing of allochthonous macrophyte subsidies by sandy beach consumers: estimates of feeding rates and impacts on food resources

Allochthonous subsidies of organic material can profoundly influence population and community structure; however, the role of consumers in the processing of these inputs is less understood but may be closely linked to community and ecosystem function. Inputs of drift macrophytes subsidize sandy beach communities and food webs in many regions. We estimated feeding rates of dominant sandy beach consumers, the talitrid amphipods (Megalorchestia corniculata, in southern California, USA, and Talitrus saltator, in southern Galicia, Spain), and their impacts on drift macrophyte subsidies in field and laboratory experiments. Feeding rate varied with macrophyte type and, for T. saltator, air temperature. Size-specific feeding rates of talitrid amphipods were greatest on brown macroalgae (Macrocystis, Egregia, Saccorhiza and Fucus). Rates for large individuals of both species ranged from ∼40 mg wet wt individual⁻¹12 h⁻¹ on brown macroalgae to negligible feeding by M. corniculata on a vascular plant (surfgrass). Amphipod growth rates were also greatest on Macrocystis and lowest on surfgrass, Phyllospadix. For a Californian beach with substantial inputs of macrophyte wrack (>70 kg wet wt m⁻¹ month⁻¹ in summer), we estimated that the population of talitrid amphipods could process an average of 55% of the palatable Macrocystis input. Our results indicate that talitrid amphipod populations can have a significant impact on drift macrophyte processing and fate and that the quantity and composition of drift macrophytes could, in turn, limit populations of beach consumers.     

Dietary contribution of the microphytobenthos to infaunal deposit feeders in an estuarine mudflat in Japan

The food sources of benthic deposit feeders were investigated at three stations in an estuarine mudflat (Idoura Lagoon, Sendai Bay, Japan) during July and August 2005, using δ¹³C and δ¹⁵N ratios. Sediment at the stations was characterized by low chlorophyll (chl) a content (0–1 cm depth, <4 μg cm⁻²) and the dominance of riverine–terrestrial materials (RTM) in the sediment organic matter (SOM) pool. Surface-deposit feeders (Macoma contabulata, Macrophthalmus japonicus, and Cyathura muromiensis) exhibited much higher δ¹³C values (−18.4 to −12.4‰) than did the SOM pool (<−25‰). A δ¹³C-based isotopic mixing model estimated that benthic diatoms comprised 45–100% (on average) of their assimilated diet, whereas RTM comprised a lesser fraction (29% maximum). The major diet of the deep-deposit feeding polychaetes Notomastus sp. and Heteromastus sp. was benthic diatoms and/or marine particulate organic matter (POM), with little RTM assimilated (39% maximum). The consumers appeared to lack specific digestive enzymes and to use detritus-derived carbon only after its transfer to the microbial biomass. The isotopic mixing model also showed that the dietary contribution of RTM increased slightly (15% maximum) in the vicinity of freshwater input, suggesting that spatial changes in RTM supply affect the dietary composition of deposit feeders. These results clearly demonstrate that deposit feeders selectively ingest and/or assimilate the more nutritious microalgal fractions in the SOM pool. Such adaptations may allow enhanced energy gain in estuarine mudflats that are rich in vascular plant detritus with low nutritive value.     

Microbial activity and carbon, nitrogen, and phosphorus content in a subtropical seagrass estuary (Florida Bay): evidence for limited bacterial use of seagrass production

Bacterial abundance, production, and extracellular enzyme activity were determined in the shallow water column, in the epiphytic community of Thalassia testudinum, and at the sediment surface along with total carbon, nitrogen, and phosphorus in Florida Bay, a subtropical seagrass estuary. Data were statistically reduced by principle components analysis (PCA) and multidimensional scaling and related to T. testudinum leaf total phosphorus content and phytoplankton biomass. Each zone (i.e., pelagic, epiphytic, and surface sediment community) was significantly dissimilar to each other (Global R = 0.65). Pelagic aminopeptidase and sum of carbon hydrolytic enzyme (esterase, peptidase, and α- and β-glucosidase) activities ranged from 8 to 284 mg N m⁻² day⁻¹ and 113–1,671 mg C m⁻² day⁻¹, respectively, and were 1–3 orders of magnitude higher than epiphytic and sediment surface activities. Due to the phosphorus-limited nature of Florida Bay, alkaline phosphatase activity was similar between pelagic (51–710 mg P m⁻² day⁻¹) and sediment (77–224 mg P m⁻² day⁻¹) zones but lower in the epiphytes (1.1–5.2 mg P m⁻² day⁻¹). Total (and/or organic) C (111–311 g C m⁻²), N (9.4–27.2 g N m⁻²), and P (212–1,623 mg P m⁻²) content were the highest in the sediment surface and typically the lowest in the seagrass epiphytes, ranging from 0.6 to 8.7 g C m⁻², 0.02–0.99 g N m⁻², and 0.5–43.5 mg P m⁻². Unlike nutrient content and enzyme activities, bacterial production was highest in the epiphytes (8.0–235.1 mg C m⁻² day⁻¹) and sediment surface (11.5–233.2 mg C m⁻² day⁻¹) and low in the water column (1.6–85.6 mg C m⁻² day⁻¹). At an assumed 50% bacterial growth efficiency, for example, extracellular enzyme hydrolysis could supply 1.8 and 69% of epiphytic and sediment bacteria carbon demand, respectively, while pelagic bacteria could fulfill their carbon demand completely by enzyme-hydrolyzable organic matter. Similarly, previously measured T. testudinum extracellular photosynthetic carbon exudation rates could not satisfy epiphytic and sediment surface bacterial carbon demand, suggesting that epiphytic algae and microphytobenthos might provide usable substrates to support high benthic bacterial production rates. PCA revealed that T. testudinum nutrient content was related positively to epiphytic nutrient content and carbon hydrolase activity in the sediment, but unrelated to pelagic variables. Phytoplankton biomass correlated positively with all pelagic components and sediment aminopeptidase activity but negatively with epiphytic alkaline phosphatase activity. In conclusion, seagrass production and nutrient content was unrelated to pelagic bacteria activity, but did influence extracellular enzyme hydrolysis at the sediment surface and in the epiphytes. This study suggests that seagrass-derived organic matter is of secondary importance in Florida Bay and that bacteria rely primarily on algal/cyanobacteria production. Pelagic bacteria seem coupled to phytoplankton, while the benthic community appears supported by epiphytic and/or microphytobenthos production.     

Productivity of bacteria and microalgae and the effect of grazing by holothurians in sediments on a coral reef flat

Bacterial productivity in sandy sediments on reef flats at Lizard Island, Great Barrier Reef was determined from the rate of incorporation of tritiated thymidine into DNA. The study was conducted during January 1982 and July 1983. A small diurnal increase occurred in sediments having a dense population of microalgae. Bacterial production was 120 to 370 mg C m^-2 d^-1 in summer on reef flats, which was equivalent to 30–40% of primary production by benthic microalgae. In winter, rates of primary production by benthic microalgae and secondary production by bacteria were about one-half to one-fifth of those in summer. There was much variation in production, due to patchiness in the distribution of benthic microbes, especially microalgae. Doubling times for the bacteria in surface sediment were 1 to 2 d in summer and 4 to 16 d in winter on the reef flats. These high productivity values for bacteria indicated that a net input of organic matter to the sediment was needed to support the growth of bacteria. Sediment bacteria thus have a very important role in transforming organic matter on the reef flats. Grazing by Holothuria atra depressed both primary production and bacterial production. It was estimated that these holothurians ate about 10 to 40% of bacterial carbon produced each day in summer, and thus have an important role in the carbon cycle. Harpacticoid copepods were numerically important components of the benthic meiofaunal community and probably had a significant impact on bacterial density as grazers.     

Carbon and nitrogen exchange between sandy beach clams (Donax serra) and kelp beds in the Benguela coastal upwelling region

Carbon consumption and nitrogen requirements were estimated for populations of the sandy beach bivalve Donax serra on nine beaches of the west coast of South Africa. Subtidal populations composed mainly of adult clams were responsible for the bulk of standing stock (3538 g C m⁻¹), annual carbon consumption (13 444 g C m⁻¹ yr⁻¹), faeces production (6478 g C m⁻¹ yr⁻¹ ) and nitrogen regeneration (2525 g N m⁻¹ yr⁻¹). Kelp detritus, bacteria and kelp consumers' faeces available in the water column surpass several times the carbon and nitrogen requirements of intertidal and subtidal clam populations. Individual Donax serra pop ulations, in turn, may regenerate up to 3.2% of the total nitrogen requirements of all primary producers from kelp beds and 14% of the requirements of phytoplankton. These high standing stocks of clams are presumably supported mainly by organic matter originating from kelp which, in contrast to phytoplankton, is in constant supply and comprises the largest proportion of the annual production of particulate organic matter on this coast. Wide and shallow continental shelves with gentle slopes probably limit the penetration of upwelled waters to the nearshore waters, decreasing the influence of external inputs and increasing the importance of internal flows of nutrients and carbon within the nearshore zone. In this context, sandy beaches, rocky shores and kelp beds may be more closely interlinked compartments of a larger ecosystem encompassing the whole nearshore than traditionally thought. Received: 28 August 1996 / Accepted: 7 October 1996     

In situ net primary productivity and photosynthesis of Antarctic sea ice algal, phytoplankton and benthic algal communities

Primary production at Antarctic coastal sites is contributed from sea ice algae, phytoplankton and benthic algae. Oxygen microelectrodes were used to estimate sea ice and benthic primary production at several sites around Casey, a coastal area in eastern Antarctica. Maximum oxygen export from sea ice was 0.95 mmol O₂ m⁻² h⁻¹ (~11.7 mg C m⁻² h⁻¹) while from the sediment it was 6.08 mmol O₂ m⁻² h⁻¹ (~70.8 mg C m⁻² h⁻¹). When the ice was present O₂ export from the benthos was either low or negative. Sea ice algae assimilation rates were up to 3.77 mg C (mg Chl-a)⁻¹ h⁻¹ while those from the benthos were up to 1.53 mg C (mg Chl-a)⁻¹ h⁻¹. The contribution of the major components of primary productivity was assessed using fluorometric techniques. When the ice was present approximately 55–65% of total daily primary production occurred in the sea ice with the remainder unequally partitioned between the sediment and the water column. When the ice was absent, the benthos contributed nearly 90% of the primary production.     

Macroinfauna of a Southern New England salt marsh: seasonal dynamics and production

The animal-habitat relationships and seasonal dynamics of the benthic macroinfauna were investigated from November 1986 to October 1988 in the Great Sippe-wissett salt marsh (Massachusetts, USA). Total macrofaunal abundance varied seasonally, displaying a peak in late spring and early summer, then declining sharply during late summer and recovering briefly in fall before collapsing in winter. Three macroinfaunal assemblages were found in the marsh, distributed along gradients of environmental factors. These included a sandy non-organic sediment assemblage, a sandy organic sediment assemblage and a muddy sediment assemblage. The species groups characteristic of unstable sandy non-organic sediments included the polychaetes Leitoscoloplos fragilis, Aricidea jefreyssi, Magelona rosea and Streptosyllis verrilli, the oligochaete Paranais litoralis, and the crustacean Acanthohaustorius millsi. Sandy organic sediments were characterized by the polychaetes Marenzelleria viridis, Capitella capitata, Neanthes succinea, N. arenaceodonta, Polydora ligni and Heteromastus filiformis, the oligochaete Lumbricillus sp., and the mollusc Gemma gemma. In muddy sites, the polychaete Streblospio benedicti and the oligochaetes Paranais litoralis and Monopylephorus evertus were the dominant species. Secondary production of benthic macroinfauna in each of these habitats was estimated. The highest values of biomass and production were recorded in the sandy organic sediments. Secondary production was estimated to be 1850 kJ m^-2 yr^-1 in sandy organic areas, but only 281 kJ m^-2 yr^-1 in sandy non-organic areas and 113 kJ m^-2 yr^-1 in muddy areas. This results in an area-weighted average production of 505 kJ m^-2 yr^-1 for the unvegetated areas of the marsh. The Great Sippewissett salt marsh has an area of 483800 m², the total secondary production of the macroinfauna for the whole unvegetated area of the marsh was estimated as 4651 kg dry wt yr^-1, expressed as somatic growth. This production value seems consistent with production data obtained for other intertidal North Atlantic environments.     

Quantitative Untersuchungen zur Ökologie der Gastrotrichen in Gezeiten-Sandstrand der Insel Sylt

Quantitative investigations were made on the distribution of 21 species of Gastrotricha Macrodasyoidea and 2 species of Chaetonotoidea in 10 different tidal beach areas of the island Sylt (North Sea). Most species live in the upper few centimeters of sandy flats near the low tide level. We counted up to 989 Gastrotricha in 50 cm³ of sediment. This group comprises up to 24% of the total interstitial fauna in about 25 samples. The beach slopes are inhabited by different species which are much less abundant (up to 165 animals per 50 cm³ sediment, and up to 7% of the total interstitial fauna). Each species inhabits a distinct zone of the sand flat or the beach slope. These zones run parallel to the water line and follow one another in a characteristic succession from the sublittoral to the supralittoral. It is assumed that several factors, controlled by exposure — e.g. variations of temperature and salinity as well as oxygen avilability — are responsible for the distribution patterns observed.     

Meiofauna response to a dynamic river plume front

 The benthic response to a plume front was studied in two areas of the northern Adriatic (Mediterranean Sea) differently influenced by the Po River freshwater input. Sediment samples were collected in June 1996 and February 1997 from 12 stations. The adopted sampling strategy was able to identify the front line in real time by satellite images and to locate sampling stations along an inner–outer plume gradient in order to cover the benthic area beneath the river plume, where enhanced biological production was expected, and open-sea sediments not directly influenced by freshwater inputs. Meiofaunal parameters were compared to the physical conditions and to phytodetritus inputs, organic matter accumulation and bacterial secondary production. The sediments of the Adriatic Sea were characterised by high concentrations of phytopigments (0.6 to 13.9 μg g⁻¹ for chlorophyll a and 1.2 to 17.7 μg g⁻¹ for phaeopigments) and biopolymeric organic carbon (0.15 to 3.02 mg g⁻¹). The plume system extended for a large sector of the northern Adriatic. In the northern area, a large and highly dynamic plume area was coupled with a sediment organic matter concentration significantly higher than in open-sea sediments. In the southern sector, where the plume area and the front line did not change markedly during the year, plume–benthic coupling was evident only in the sediments beneath the front, and corresponded to phaeopigment accumulation. Bacterial parameters and secondary production were high and significantly higher in the frontal area than at open-sea stations. Meiofauna density (1342 to 8541 ind. 10 cm⁻²) did not change either by season or between areas and was significantly correlated with phaeopigments and bacterial secondary production. Meiofauna displayed different responses to plume inputs in the two sampling areas. In the northern sector, meiofauna density was coupled with organic matter distribution and displayed highest values beneath the plume. In the southern sector, the densities of copepods, turbellarians and kinorhynchs displayed highest values under the front in summer, and the same applied to total meiofauna density in winter. Juvenile decapods and copepod nauplii significantly increased their densities in sediments beneath the front. Data presented in the present study suggest that plume inputs and frontal systems, enhancing phytodetritus accumulation and benthic bacterial response, might influence density, composition and distribution of meiofaunal assemblages. As river plumes are highly variable systems affecting the trophic characteristics of the sediments underneath, their dynamics should be considered when analysing mesoscale spatial changes of meiofaunal assemblages. Received: 30 November 1999 / Accepted: 24 May 2000     

Bacterivory of a mudflat nematode community under different environmental conditions

The fate of the benthic bacterial biomass is a topic of major importance in understanding how soft-bottom environments function. Because of their high abundance, production and nutritional value, benthic bacteria may constitute an important food resource for benthic fauna. The trophic role of bacteria for a nematode community on the Brouage mudflat (Marennes-Oléron-France), dominated by three species: Chromadora macrolaima (64% of the abundance), Daptonema oxycerca (15%) and Ptycholaimellus jacobi (8%), was determined in grazing experiments using ¹⁵N pre-enriched bacteria. On intertidal flats, seasonal, tidal and circadian cycles induce strong variations in environmental conditions. Grazing experiments were performed in order to measure the effects of abiotic (temperature, salinity and luminosity) and biotic (bacterial and algal abundances) factors on assimilation rates of bacteria by nematodes. In order to assess simultaneously bacteria and algal assimilation rates, algal abundances were modified adding ¹³C pre-enriched Navicula phyllepta. Assimilation rate was significantly lower at 5°C; moreover, general trend shows a prominent temperature effect with an optimum around 30°C. Assimilation at salinity 18 was not significantly different from the assimilation at salinity 31. Assimilation was higher under light conditions than in the dark. Above 10⁹ bacteria ml⁻¹, assimilation of bacteria remained unaffected by bacterial abundance. However, assimilation of algae increased with the algal concentration. Nematode kept feeding under conditions of stress, which are typical of the surficial sediment habitat and they appeared to be principally dependent on the algal resource.     

The amphipod Corophium multisetosum (Corophiidae) in Ria de Aveiro (NW Portugal). II. Abundance, biomass and production

 The abundance and biomass of Corophium multisetosum Stock, 1952 were determined from benthic corer samples collected monthly over 1 yr in the upper reaches of Canal de Mira (Ria de Aveiro, Portugal). Both density and biomass over the sampling period were negatively correlated with water temperature and positively correlated with chlorophyll a concentration in the sediment. C. multisetosum density was significantly negatively correlated with plant biomass and positively correlated with salinity. The nature of the sediment, favourable environmental conditions, high availability of food and low interspecific competition allowed the population to reach a maximal density of 200 × 10³ individuals m⁻² and a maximal biomass (ash-free dry wt, AFDW) of 62 g_AFDW m⁻². The population was highly productive, especially during the autumn/winter period. Production, estimated by two different methods (Hynes method: 251 g_AFDW m⁻² yr⁻¹; Morin–Bourassa method: 308 g_DW m⁻² yr⁻¹), was much higher than the values reported for other Corophium species. The annual P:Bˉ ratio (10) was high, but similar to values reported for Swedish populations of C. volutator and lower than the values estimated from Mediterranean populations of C. insidiosum. Received: 8 October 1999 / Accepted: 22 June 2000     

Effects of river sediment addition to the beach microbial communities

During the processes of claiming land from the sea, river sediments are used to fill and transform the sea area along the margin of islands and lands into new lands. These activities would probably affect microbial ecosystems of the beach sediment. However, little is known about these effects. In this study, a simulation test was conducted to evaluate these effects. Pyrosequencing technique was employed to assess the effects of river sediment addition to the beach bacterial communities. The used river sediments were collected from different rivers. The results indicated that river sediment addition greatly impacted microbial ecosystems of the beach and caused a clear shift in the beach bacterial community composition. These processes remarkably increased toxic metals and decreased the bacterial diversity in the beach sediment, mainly including the phyla of Bacteroidetes, Cyanobacteria, Proteobacteria, and Spirochaetes. River sediment addition caused an increase in potentially pathogenic bacterial genera of the beach sediment. Bacterial phylotype richness in the beach without river sediment addition was higher than that in the beach with river sediment input. There were significant differences in bacterial communities between beach sediments with and without river sediment addition, and the most dominant classes were Gammaproteobacteria and Flavobacteria.     

Population dynamics and growth of <Emphasis Type="Italic">Nassarius reticulatus </Emphasis>(Gastropoda: Nassariidae) in Rhosneigr (Anglesey,UK)

Seasonal changes in catch rate, growth and mortality of Nassarius reticulatus from an intertidal lagoon and a wave-exposed beach at Rhosneigr (Anglesey, North Wales, UK) are described. The number of N. reticulatus caught in baited traps from the lagoon was significantly higher (>125 individuals trap⁻¹) during the summer (>18°C), than at <12°C (<65 individuals trap⁻¹), and the numbers caught in the lagoon were an order of magnitude greater than on the beach, >13 individuals trap⁻¹ in July (>16°C), and <5 individuals trap⁻¹ between December and April (<9.5°C). Predictions of shell growth attained by N. reticulatus annually in the lagoon using graphical modal progression analysis (MPA) of length frequency data, were similar to the growth of marked and recaptured lagoon N. reticulatus. Predictions of shell growth using computerised length frequency distribution analysis (LFDA), however, did not reflect the growth as accurately as MPA. Modal progression analysis demonstrated that N. reticulatus from the lagoon achieved a higher asymptotic maximum shell length (L _∞) and a lower growth constant (K) than animals from the beach. Shell growth was seasonal with growth of the lagoon individuals slowing down towards the end of September and resuming in early April, about a month later than the beach individuals. Mortality of N. reticulatus was greater during the summer, and survival was lower in the lagoon than on the beach. Recruitment patterns were similar in the lagoon and on the beach, and MPA and LFDA predicted that larval N. reticulatus settled between late summer and early autumn, with juveniles (7–8.9 mm) appearing in the population the following year, between February and April. Growth of male and female N. reticulatus in the laboratory was similar and was temperature and size dependent. The different growth patterns between N. reticulatus from the two habitats, predicted using MPA, were maintained when individuals were reared under laboratory conditions for ∼6 months; N. reticulatus <21 mm from the beach grew faster than individuals from the lagoon, although N. reticulatus >21 mm from the lagoon grew faster and attained a larger length (26 mm) than individuals from the beach (24 mm). Low food availability did not affect N. reticulatus survival in the laboratory but significantly suppressed shell growth.     

Hydrodynamic and chemical conditions during onset of a red-tide assemblage in an estuarine upwelling ecosystem

The hydrodynamics and nitrogen/silicon biogeochemistry accompanying the development of a red-tide assemblage were examined in the Ría de Vigo (northwest Spain), a coastal embayment affected by upwelling, during an in situ diel experiment in September 1991. Despite a low N:Si molar ratio (0.5) of nutrients entering the surface layer, which was favourable for diatom growth, the diatom population began to decline. Limited N-nutrient input, arising from moderate coastal upwelling in a stratified water column, restricted net community production (NCP = 630 mg C m⁻² d⁻¹). In addition, light-limitation of gross primary production (GPP = 1525 mg C m⁻² d⁻¹) was observed. The relatively high f-ratio (= NCP:GPP) recorded (0.41, characteristic of intense upwelling conditions) would have been as low as 0.15 had not GPP been limited by light intensity. Temporal separation of carbohydrate synthesis during the photoperiod from protein synthesis in the dark could be inferred from the time-course of the C:N ratio of particulate organic matter. Severe light-limitation would lead to diatom collapse were the diatoms not able to meet all their energy requirements during the hours of darkness. Under the hydrodynamic, nutrient and light conditions of the experiment, an assemblage of red-tide-forming species began to develop, aided by their ability to migrate vertically and to synthesize carbohydrates during the light in surface waters and protein during the dark at the 4 m-deep pycnocline. Thermal stratification, reduced turbulence, intense nutrient mineralization, and the limited nitrogen input through moderate upwelling were all favourable to the onset of a red-tide assemblage. Received: 15 February 1997 / Accepted: 26 September 1997     

Community metabolism of intertidal flats in the Ems-Dollard estuary

To obtain an insight into the flux of carbon through intertidal sediments of the Ems-Dollard estuary, the annual cycles of gross benthic primary production and community respiration were measured at six stations, together with a set of environmental parameters. In a stepwise multiple regression analysis it was shown that temperature alone and temperature plus viable bacteria explained 50 and 70% respectively of the observed variation in community respiration. Other variables, including the rate of primary production and amount of organic carbon in the sediment were less important. The rate of primary production could not be fitted adequately into a multiple regression equation. The annual values of community respiration (177–794 gO₂·m^-2·yr^-1) and primary production (82–628 gO₂·m^-2·yr^-1) were within the range of published values. except for one station in the vicinity of a wastewater outfall, which had an extreme production (average 984 gO₂·m^-2·yr^-1). At four stations, annual community respiration exceeded primary production by 40%. It is concluded that the main carbon flux within the sediment, from CO₂ to benthic primary producers, to benthic consumers and from there to CO₂ again,was completed within a month or so, leaving untouched the large bulk of organic matter within the sediment. Possible effects of wastewater discharges on community metabolism are discussed.Publication No. 43 of the project Biological Research in the Ems-Dollard Estuary     

Biomass partitioning of benthic microbes in a Baltic inlet: relationships between bacteria, algae, heterotrophic flagellates and ciliates

 The structure of a benthic microbial food web and its seasonal changes were studied in the shallow brackish waters of the island of Hiddensee, northeastern Germany, at two sites in close proximity by monthly or bimonthly sampling from July 1995 to June 1996. Abundance and biomass of phototrophic and non-phototrophic bacteria, heterotrophic flagellates (HF) and ciliates as well as the biomass of microphytobenthos were determined in the upper 0.3 cm sediment layer. Abundance of organisms showed strong positive correlation with water temperature, with the exception of the bacteria. Non-phototrophic bacterial numbers ranged from 7 × 10⁸ to 6.7 × 10⁹ cells cm⁻³ and phototrophic bacterial abundance from 4 × 10⁷ to 2.7 × 10⁸. Heterotrophic protist abundance ranged from 8 × 10³ to 104 × 10³ ind cm⁻³ for HF and from 39 to 747 ind cm⁻³ for ciliates. The biomass partitioning demonstrated the primary importance of non-phototrophic bacteria (min. 0.83, max. 84.87 μg C cm⁻³), followed by the microphytobenthos (min. 1.32, max. 50.93 μg C cm⁻³). The heterotrophic protists contributed roughly the same fraction to the total microbial biomass, with the biomass of the HF being slightly higher (HF 0.23 to 1.76 μg C cm⁻³, ciliates 0.04 to 1.17 μg C cm⁻³). Taxonomic classification of the benthic HF revealed the euglenids to be the most important group in terms of abundance and biomass, followed by thaumatomastigids and kinetoplastids. Other important groups were apusomonads, cercomonads, pedinellids and cryptomonads. The structure of the HF assemblage showed strong seasonal changes with euglenids being the most abundant taxa in summer, while apusomonads and thaumatomastigids were predominant in winter. Similar to the pelagic microbial food web, benthic picophototrophic bacteria were occasionally abundant, and the feeding modes of heterotrophic protists exhibited a great variety (predominantly omnivores, bacterivores, herbivores or predators). Filter-feeding HF were of little importance. Contrary to the pelagic environment, a top-down control on total benthic bacterial numbers by HF seemed unlikely at the studied stations which were characterised by muddy sand. Received: 6 January 1999 / Accepted: 21 October 1999     

Food sources used by sediment meiofauna in an intertidal Zostera noltii seagrass bed: a seasonal stable isotope study

In an intertidal Zostera noltii Hornem seagrass bed, food sources used by sediment meiofauna were determined seasonally by comparing stable isotope signatures (δ¹³C, δ¹⁵N) of sources with those of nematodes and copepods. Proportions of different carbon sources used by consumers were estimated using the SIAR mixing model on δ¹³C values. Contrary to δ¹⁵N values, food source mean δ¹³C values encompassed a large range, from −22.1 ‰ (suspended particulate organic matter) to −10.0 ‰ (Z. noltii roots). δ¹³C values of copepods (from −22.3 to −12.3 ‰) showed that they use many food sources (benthic and phytoplanktonic microalgae, Z. noltii matter). Nematode δ¹³C values ranged from −14.6 to −11.4 ‰, indicating a strong role of microphytobenthos and/or Z. noltii matter as carbon sources. The difference of food source uses between copepods and nematodes is discussed in light of source accessibility and availability.     

Benthic community respiration in the N.W. Atlantic Ocean: in situ measurements from 40 to 5200 m

Benthic community respiration was measured in situ at 9 stations along the Gay Head-Bermuda transect from depths of 40 to 5200 m. Three methods were used; bell jar respirometers, grab respirometers, and free vehicle respirometers. Benthic community respiration rates spanned three orders of magnitude, decreasing from 21.5 ml O₂ m^-2 h^-1 at 40 m in November to 0.02 ml O₂ m^-2 h^-1 at 5200 m. Rates decreased two orders of magnitude between 40 and 1800 m and then significantly declined again between the continental rise (3650 m) and the abyssal plain stations. Predictive equations for benthic community respiration along the transect reflect a strong correlation with depth of water. Of lesser significance are the correlations with water temperature, dissolved oxygen, benthic animal biomass, surface primary productivity and sediment organic matter. Calculations show that annual benthic respiration can utilize 1 to 2% of the surface primary productivity. Of the 2 to 7% organic carbon fixed at the surface which supposedly reaches the bottom, only 15 to 29% is utilized by the benthic community at 2200, 3000, and 3650 m. The energy requirements of other biological components of deep-sea benthic communities, such as benthopelagic and macro-epibenthic animals, not included in these measurements, must also be considered in calculating a balance of carbon.Contribution from Scripps Institution of Oceanography.     

Strong variability in bacterioplankton abundance and production in central and western Bay of Bengal

With large influx of freshwater that decreases sea-surface salinities, weak wind forcing of <10 m s⁻¹ and almost always warm (>28°C) sea-surface temperature that stratifies and shallows the mixed layer leading to low or no nutrient injections into the surface, primary production in Bay of Bengal is reportedly low. As a consequence, the Bay of Bengal is considered as a region of low biological productivity. Along with many biological parameters, bacterioplankton abundance and production were measured in the Bay of Bengal during post monsoon (September–October 2002) along an open ocean transect, in the central Bay (CB, 88°E) and the other transect in the western Bay (WB). The latter representing the coastal influenced shelf/slope waters. Bacterioplankton abundances (<2 × 10⁹cells l⁻¹) were similar to those reported from the HNLC equatorial Pacific and the highly productive northern Arabian Sea. Yet, the thymidine uptake rates along CB (average of 1.46 pM h⁻¹) and WB (average of 1.40 pM h⁻¹) were less than those from the northwestern Indian Ocean. These abundances and uptake rates were higher than those in the oligotrophic northwestern Sargasso Sea (<7 × 10⁸ cells l⁻¹; av 1.0 pM h⁻¹). Concentrations of chlorophyll a (chl a), primary production rates and total organic carbon (TOC) were also measured for a comparison of heterotrophic and autotrophic production. In the WB, bacterioplankton carbon biomass equaled ∼ 95% of chl a carbon than just 31% in the CB. Average bacterial:primary production (BP:PP) ratios accounted for 29% in the CB and 31% in the WB. This is mainly due to lower primary productivity (PP) in the WB (281 mg C m⁻² d⁻¹) than in the CB (306 mg C m⁻² day⁻¹). This study indicates that bacteria–phytoplankton relationship differs in the open (CB) and coastal waters (WB). Higher abundance and contrastingly low bacterial production (BP) in WB may be because of the riverine bacteria, brought in through discharges, becoming dormant and unable to reproduce in salinities of 28 or more psu. Heterotrophic bacteria appear to utilize in situ DOC rather rapidly and their carbon demand is ∼50% of daily primary production. It is also apparent that allochthonous organic matter, in particular in the western Bay, is important for meeting their carbon demand.     

Nutrient regeneration in shallow-water sediments of the estuarine plume region of the nearshore Georgia Bight,USA

Benthic community respiration and the cycling of N and P were seasonally investigated in the unprotected, sandy sediments (Z5m) of the nearshore zone of the Georgia Bight, USA in 1981 and 1982. Nutrient exchange across the sediment-water interface was calculated from a diffusive model, measured by in-situ enclosure experiments and estimated from whole core incubations. Seasonally changing pore water profiles indicated that the sediments were not in steady-state with respect to N and P and showed the characteristics of enhanced interstitial water movement by benthic animals. Over an annual period the total flux of nitrogen measured in situ averaged 1812 mol m^-2 d^-1 from the sediments. NH ₄ ⁺ flux accounted for the vast majority of the total directly measured N flux (77%), followed by nitrate + nitrite (14%), and dissolved organic nitrogen (9%). Phosphorus flux averaged 537 mol m^-2 d^-1. A large ratio of in-situ fluxes to calculated diffusive fluxes (5.2:1) indicated flux enhancement due to benthic animal activity. ammonium fluxes measured in situ did not agree well with the rate of NH ₄ ⁺ produced in incubated whole cores (11.7 mmol m^-2 d^-1). Relative rates of C, N and P release throughout the year fluctuated considerably. Generally, nutrient fluxes were not simply related to respiration or temperature. As respiration was highly correlated with temperature, however, this suggested that respiration-regeneration was temporarily decoupled from exchange across the sediment-water interface. The annual C-N-P flux stoichiometry was 130:3.1:1. Using the rate at which NH ₄ ⁺ was produced in incubated cores the stoichiometry was 120:21:1. The anomalously low N flux measured in situ was attributed to a combination of denitrification and wave-and current-induced sediment nutrient flushing. The potential for sediment flushing is high as experiments showed that sediments were fluidized or resuspended down to 25 cm during large storms. Benthic nutrient flux contributed 40% to the annual P but only 11% to the annual N requirements of the pelagic primary producers.This is Contribution No. 558 from the University of Georgia Marine Institute. This work was supported by the Georgia Sea Grant College Program maintained by the National Oceanic and Atmospheric Administration, US Department of Commerce, under Grant No. NA80AA-D-00091