Eutrophication and trophic structure in response to the presence of the eelgrass <Emphasis Type="Italic">Zostera noltii</Emphasis>

In estuaries, eelgrass meadows contribute to fundamental ecosystem functions of estuaries, providing food to several predators and buffering the negative effects of eutrophication. We asked whether the presence of the eelgrass Zostera noltii decreased the nitrogen concentration in the overlying water, affected the sources of nitrogen sequestrated by primary producers and changed the benthic and pelagic food web structures. We also studied the importance of these food webs in providing food to fish. We compared bare sediment to sediment covered by a Z. noltii meadow, and examined nutrient concentrations in the water column and δ¹⁵N in primary producers as indicators of anthropogenic inputs of nutrients. We then measured both δ¹³C and δ¹⁵N in the tissues of plants and consumers to establish food web structures. There were no differences in the concentrations and sources of nitrogen between sites. Rather, δ¹⁵N values indicated anthropogenic inputs of N (e.g. sewage discharges, agriculture) in both sites. There were no major differences in the structure of the planktonic food web, which was in part sustained by particulate organic matter and supported most predator fish, and in the structure of the benthic food web. Nonetheless, there were differences in the sources of food for omnivore consumers and for the detritivore Scrobicularia plana. Overall, the benthic food web did not use food derived from the eelgrass or macroalgae deposited on the substratum. Suspension feeders used particulate and sediment organic matter, whereas the δ¹³C and δ¹⁵N values of the other consumers indicated a likely contribution of benthic microalgae. Furthermore, in both habitats we found large variability in the isotope signatures of benthic macrofauna consumers, which did not allow distinguishing clearly different trophic groups and indicated a high level of omnivory and a mixed diet opportunistically making use of the availability of food in the surroundings.     

Seasonal changes of benthic bacteria in a seagrass bed (Posidonia oceanica) of the Ligurian Sea in relation to origin,composition and fate of the sediment organic matter

Variations in number and biomass of benthic bacteria were examined in the surface sediments of a Mediterranean seagrass bed [Posidonia oceanica (L.) Delile] in the Gulf of Marconi (northwestern Mediterranean Sea) from 1990 to 1991. The annual dynamics of benthic bacterial density and biomass were compared to changes in elemental (organic C and total N) and biochemical (lipids, proteins, carbohydrates) composition of sediment organic matter, as well as to microphytobenthic biomass, dissolved inorganic nutrients and ATP. Bacterial densities exhibited marked seasonal variations (5.12 to 322.7x10⁸ cells g^-1 sediment dry wt) with highest values in late spring. Bacterial standing stocks (15.8 to 882.33 g C g^-1 of sediment dry wt) were high. Bacterial biomass did not correlate with organic C, total N or to specific biochemical components, but correlated significantly with chlorophyll a, ATP and porewater phosphate concentrations. There is evidence that benthic bacteria were responding to variations of algal biomass. Bacterial biomass accounted, on average, for 30% of total living carbon (calculated on the basis of the ATP concentrations) and 8.4% of total organic carbon.     

Effect of food type and ration on growth of juvenile Capitella sp. I (Annelida: Polychaeta): macro- and micronutrients

Growth rates of juvenile Capitella sp. I were determined on different rations of six food types: Gerber's mixed cereal, TetraMin fish flakes, benthic diatoms, Ulva sp., spring detritus, and summer detritus. A simple growth model estimated maximum growth rate and maintenance ration for each food. There were differences in the growth response among foods relative to nitrogen content. As juveniles increased in size, differences in growth between foods became more pronounced. For all juveniles, growth rates were correlated with levels of the amino acids histidine, phenylalanine, threonine, and valine, and the polyunsaturated fatty acid 20:5w3; correlations with histidine and phenylalanine levels were the most significant. Regressions of growth rates as a function of these two amino acids suggest a daily maintenance ration of 300 pg histidine and phenylalanine mg^-1 nitrogen biomass. Juvenile worms grew on spring detritus but not on summer detritus, indicating the probable importance of micronutrients (polyunsaturated fatty acids, amino acids) for growth of juvenile Capitella sp. I in the field.     

Benthic response to sedimentation of a spring phytoplankton bloom: Process and budget

The response of benthos to sedimentation of the spring phytoplankton bloom in the Kiel Bight (Western Baltic Sea) is described in terms of biomass (ATP) and activity (heat production and ETS-activity). Input of the bloom (11.5 g C m^-2) over a period from March 25 to April 19, 1980 to the sediment surface was in the form of cells and fresh phytodetritus as indicated by low C/N ratios (7) and high energy charge values (0.78). Benthic microbial activity was immediately stimulated by this input as heat production doubled and the activity of ETS tripled over winter values within 12 d in the absence of a significant increase in ambient temperature. A comparison of the two activity parameters suggests that anaerobic metabolism is more important during the winter (February and March) than after input of the bloom. Meiofauna was not able to take part in the first activity outburst. Benthic ATP-biomass (excluding macrofauna) doubled in late April due to microbial production, and doubled again in early May when meiofauna started reproductive activity. For macrofauna a general statement was not possible, although the sediment surface feeder Macoma baltica commenced a build up of glycogen and lipid resources immediately following bloom input whereas Nephtys ciliata, feeding on sediment and small macrofauna, showed a less pronounced and delayed effect from this input. An energy budget based on heat production measurements was calculated. A daily heat loss of the benthic community of 21.7 KJ m^-2 d^-1 (35.5 KJ m^-2 d^-1) was found, when a depth of 3 cm sediment (5 cm) was assumed. Heat production of macrofauna contributed less than 5% of this activity. The input of the bloom was burned within 21 (13) d. Preliminary estimations for an annual budget suggest that the vertical transport of particulate organic matter via sedimentation can only explain 25% (15%) of the benthic activity in the shallow water ecosystem of the Kiel Bight. This indicates the presence of other sources of organic carbon such as benthic primary production or other transport processes providing carbon to the sediments.Publication No. 384 of the Joint Research Program of Kiel University (Sonderforschungsbereich 95)     

Interlinked temporal changes in environmental conditions, chemical characteristics of sediments and macrofaunal assemblages in an estuarine intertidal sandflat (Seto Inland Sea, Japan)

Five field surveys were conducted in an estuarine intertidal sandflat of the Seto Inland Sea (Japan) between April 1994 and April 1995. Chlorophyll a, pheopigments, total organic carbon and acid-volatile sulphides (AVS) of surface and subsurface sediments, and macrofaunal assemblages were investigated in parallel at 15 stations. Monthly hydrological data of low-tide creek water adjacent to the flat were used as a complementary environmental characterisation of the study area. Strong temporal changes were found among sampling dates, most remarkably in autumn with a major increase of algal detritus and AVS, a sharp reduction in macrofaunal abundances and species richness, and a massive mortality of the clam Ruditapes philippinarum. This dystrophic event was preceded by a photoautotrophic and hypertrophic spring–summer characterized by abundant fresh (i.e., living) algal material, including microphytobenthos and macroalgae (Ulva sp.). In summer, abundant macrofaunal assemblages reached the highest biomass values (455 g wet weight m⁻² or 60.6 g ash free dry weight m⁻²), with a major contribution of filter-feeding bivalves Musculista senhousia and R. philippinarum. These are among the highest values reported in the literature for sedimentary shores. From autumn, there was a progressive recolonisation of macrofauna, initiated by few opportunistic polychaetes (e.g., Cirriformia tentaculata and Polydora sp.), apparently promoting a fast sediment recovery in winter, and followed by new bivalve recruits in the next spring. This study provides the first evidence of significant and interlinked within-year changes in chemical characteristics of sediments and macrofaunal assemblages in an estuarine intertidal flat at a small spatial scale (i.e., tens of meters). This demonstrates the high temporal variability of species–environment relations in these systems and a close relationship in seasonally driven trophodynamic processes among primary producers and benthic consumers. We conclude that a thorough parallel evaluation of the temporal changes in chemical characteristics of sediments should be taken into account in assessing the year-round distribution and changes of intertidal macrofauna, particularly in eutrophic, estuarine intertidal flats.An erratum to this article can be found at     

Recycle of buried macroalgal detritus in sediments: use of dual-labelling experiments in the field

In intertidal sediments, burial and decomposition of macroalgal detritus can fuel the sediment of carbon (C) and nitrogen (N), which can be either promptly mineralised or assimilated to enter the food web. This study investigates the transfer of algal-derived C and N to the sediment and to the infauna feeding primarily on benthic diatoms. Thalli of Ulva spp. were ¹³C- and ¹⁵N-labelled in the mesocosm and frozen to create detritus. Thawed macroalgae were, then, buried in the sediment of an intertidal sand-flat forming a mosaic of small patches (50 × 50 cm²) enriched with the macroalgae interspersed with bare sediment. The area was dominated by Corophium volutator and Hydrobia ulvae. The uptake of ¹³C and ¹⁵N was measured in the residual macroalgae, in the sediment and in those animals. Decomposition of detritus was rapid and after 4 weeks the residual biomass was 3% the amount added. Algal-derived ¹³C and ¹⁵N were moved to the sediment. The total amount of ¹³C and ¹⁵N retained in the sediment after completing the decomposition was 3.4 ± 0.5% ¹³C and 2.7 ± 0.6% ¹⁵N the amount decomposed. During the first 2 weeks more N than C was assimilated (1.7% ¹³C and 13.5% ¹⁵N). During the remaining two weeks, N was released from the sediment, while there was little accumulation of C (+6.4 ± 2.0 % ¹³C and −7.7 ± 3.8% ¹⁵N). At the end of the decomposition, animals were ¹⁵N- and ¹³C-labelled. Considering the total accumulation of label in the sediment, they accounted for 3.5 ± 1.8% ¹³C and 25.8 ± 12.9 % ¹⁵N. Similarly, considering the mass of the heavy isotopes gained (¹³C) or lost (¹⁵N) during the remaining 2 weeks, the animals accounted for 4.7 ± 2.1% of the ¹³C in excess and for 18.6 ± 9.1% of the ¹⁵N loss. The transfer of C and N to the sediment and to the surface deposit-feeders can be a relevant mechanism to remove the excess of detritus from the sediment.     

The capacity of the filter-feeding bivalve Mya arenaria L. to affect water transport in sandy beds

Benthic filter feeding macrofauna organisms may be an important factor linking sediment and water column. Transport of water and concomitantly of suspended matter is directly related to the size of the benthic filter feeding population. This paper aims to determine the potential for water transport by the bivalve Mya arenaria along a coastal stretch of roughly 100 km length in the southern Baltic Sea. Quantification of population filtration rates specific to the area is based on distribution, abundance and biomass of M. arenaria and calculated according to previously published filtration rate–biomass relations. Calculated rates range up to >8 m³ m^–2 day^–1 (at 5–20 m water depth in sandy sediment) with the potential to locally process a volume of water equivalent to the water column within <1 day. Data from 1991–2002 at one site suggest that the area-specific potential population filtration rate remains remarkably constant in time despite changes in population structure of M. arenaria. The related impact on pore water exchange within the permeable sediment associated with the leakage of water from the gape of M. arenaria valves is discussed.Communicated by L. Hagerman, Helsingør     

Role of cold resistance and burial for winter survival and spring initiation of an Ulva spp. (Chlorophyta) bloom in a eutrophic lagoon (Veerse Meer lagoon, The Netherlands)

In the eutrophic Veerse Meer lagoon (The Netherlands) large amounts of free-floating thalli from Ulva spp. are present from May to October. In winter however, no algae seem to occur in the lagoon. Sexual reproduction appears to be negligible, as spore formation and germling growth are observed only sporadically. Results of a field survey showed that in winter, viable Ulva biomass is present buried in the sediment of the shallow parts of the lagoon. Freezing experiments demonstrated that the algae are able to survive temperatures of −5 °C for 2 weeks when kept in darkness. In spring, the buried Ulva thalli are liberated out of the sediment to initiate a bloom. A field experiment indicates that bioturbation by the lugworm Arenicola marina does not stimulate the release of the thalli. Burial and winter survival can explain the rapid increase in Ulva biomass in spring and suggest that the initial spring biomass is one of the major factors determining the maximal biomass in summer. Received: 28 October 1997 / Accepted: 20 January 1998     

The influence of macrofauna on estuarine benthic community metabolism: a microcosm study

Effects of benthic macrofauna (Corophium volutator, Hydrobia sp., Nereis virens) on benthic community metabolism were studied over a 65-d period in microcosms kept in either light/dark cycle (L/D-system) or in continuous darkness (D-system). Sediment and animals were collected in January 1986 in the shallow mesohaline estuary, Norsminde Fjord, Denmark. The primary production in the L/D-system after 10 d acted as a stabilizing agent on the O₂ and CO₂ flux rates, whereas the D-system showed decreasing O₂ and CO₂ flux throughout the period. Mean O₂ uptake over the experimental period ranged from 0.38 to 1.24 mmol m^–2 h^–1 and CO₂ release varied from 0.80 to 1.63 mmol m^–2 h^–1 in both systems. The presence of macrofauna stimulated community respiration rates measured in darknes, 1.4 to 3.0 and 0.9 to 2.0 times for O₂ and CO₂, respectively. In contrast, macrofauna lowered primary production. Gross primary production varied from 1.06 to 2.26 mmol O₂ m^–2 h^–1 and from 1.26 to 2.62 mmol CO₂ m^–2 h^–1. The community respiratory quotient (CRQ, CO₂/O₂) was generally higher in the begining of the experiment (0–20 d, mean 1.89) than in the period from Days 20 to 65 (mean 1.38). The L/D-system exhibited lower CRQ (ca. 1) than the D-system. The community photosynthetic quotient varied for both net and gross primary production from 0.64 to 1.03, mean 0.81. The heterotrophic D-system revealed a sharp decrease in the sediment content of chlorophyll a as compared to the initial content. In the autotrophic L/D-system, a significant increase in chlorophyll a concentration was observed in cores lacking animals and cores with C. volutator (The latter species died during the experiment). Due to grazing and other macrofauna activities other cores of the L/D-system exhibited no significant change in chlorophyll a concentration. Community primary production was linearly correlated to the chlorophyll a content in the 0 to 0.5 cm layer. Fluxes of DIN (NH₄ ⁺+NO₂ ^–+NO₃ ^–) did not reveal significant temporal changes during the experiment. Highest rates were found for the cores containing animals, mainly because of an increased NH₄ ⁺ flux. The release of DIN decreased significantly due to uptake by benthic microalgae in the L/D-system. No effects of the added macrofauna were found on particulate organic carbon (POC), particulate organic nitrogen (PON), total carbon dioxide (TCO₂) and NH₄ ⁺ in the sediment. The ratio between POC and PON was nearly constant (9.69) in all sediment dephts. The relationship between TCO₂ and NH₄ ⁺ was more complex, with ratios below 2 cm depth similar to those for POC/PON, but with low ratios (3.46) at the sediment surface.     

Important organic matter sources and trophic pathways for the nutrition of Hilsa kelee (Cuvier, 1829) and Valamugil buchanani (Bleeker, 1853) in Pangani macro-tidal estuary,Tanzania

The study investigated the spatial variation in the main sources of organic matter (OM) and trophic pathways for zooplanktivorous Hilsa kelee and phytodetritivorous Valamugil buchanani in fresh-water-influenced zone versus sea-water-dominated zone of Pangani estuary. The findings indicated significant inter-specific variations in δ¹³C and δ¹⁵N values (ANOVA, F?≥?84.3, p?F?≥?9.4, p?=?0.001) in both estuarine zones. Results also showed significant zonal-intraspecific variations in stable isotopes (δ¹³C and δ¹⁵N), FA profile and marginal differences in diet for the V. buchanani while no considerable differences were observed for H. kelee from two estuarine zones. The isotope mixing models and FA biomarkers revealed that the most important carbon sources to the nutrition of H. kelee were derived from microphytobenthos, macro-algae and sea grasses transferred through phytoplankton and detrital trophic pathways. In contrast, C₃ terrestrial plants and microphytobenthos were the main carbon source to the diet of V. buchanani; and were transferred via the benthic and detrital trophic pathways. Therefore, both terrestrial and in-situ OM sources were the main trophic resources base fuelling the planktonic and benthic food webs in Pangani estuary.     

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.     

Worldwide effects of non-native species on species–area relationships

Non-native species have invaded most parts of the world, and the invasion process is expected to continue and accelerate. Because many invading non-native species are likely to become permanent inhabitants, future consideration of species-area relationships (SARs) should account for non-native species, either separately or jointly with native species. If non-native species occupy unused niches and space in invaded areas and extinction rate of native species remains low (especially for plants), the resultant SARs (with both native and non-native species) will likely be stronger. We used published and newly compiled data (35 data sets worldwide) to examine how species invasions affect SARs across selected taxonomic groups and diverse ecosystems around the world. We first examined the SARs for native, non-native, and all species. We then investigated with linear regression analyses and paired or unpaired t tests how degree of invasion (proportion of non-native species) affected postinvasion SARs. Postinvasion SARs for all species (native plus non-native) became significantly stronger as degree of invasion increased (r² = 0.31, p = 0.0006), thus, reshaping SARs worldwide. Overall, native species still showed stronger and less variable SARs. Also, slopes for native species were steeper than for non-native species (0.298 vs. 0.153). There were some differences among non-native taxonomic groups in filling new niches (especially for birds) and between islands and mainland ecosystems. We also found evidence that invasions may increase equilibrial diversity. Study of such changing species–area curves may help determine the probability of future invasions and have practical implications for conservation.     

The role of the amphipod <Emphasis Type="Italic">Gammarus locusta</Emphasis> as a grazer on macroalgae in Swedish seagrass meadows

Mesograzers are thought to play a critical role in seagrass beds by preventing overgrowth of ephemeral algae. On the Swedish west coast, eelgrass Zostera marina has decreased in recent decades as a result of eutrophication and increased growth of macroalgal mats (mainly filamentous Ulva spp. and Ectocarpales), with no indication of grazer control of the algae. The aim of this study was to investigate the ability of the amphipod Gammarus locusta to control algal blooms during nutrient-enriched and ambient conditions, using a combination of laboratory, field and model studies. Laboratory experiments demonstrated that juvenile and adult G. locusta could consume both Ulva spp. and Ectocarpales, but that consumption of Ulva spp. was significantly higher. Cannibalism was common in individual treatments involving multiple size-classes of G. locusta, but only large, male gammarids consumed smaller juveniles in the presence of Ulva spp. as an alternative food source. However, no negative effects of cannibalism were found on total grazing impact. A model using size-specific grazing rates and growth rates of G. locusta and of Ulva spp. suggests that approximately 62 young juvenile, or 27 adult G. locusta are needed per gram DW of Ulva spp. to control the algal growth during ambient nutrient conditions, and approximately 2.6 times as many gammarids during enhanced nutrient conditions. On the Swedish west coast, densities and mean sizes of G. locusta in eelgrass beds are below these critical values, suggesting that the gammarids will not be able to control the growth of the filamentous macroalgae. However, in the field cage experiment, immigration of juveniles and reproduction of encaged adult G. locusta resulted in unexpectedly high densities of G. locusta (>4,000 individual m⁻²), and very low biomass of Ulva spp. in both ambient and nutrient-enriched treatments. Although the high numbers of juveniles in all cages precluded any significant treatment effects, this suggests that in the absent of predators, the population of G. locusta can grow significantly and control the biomass of Ulva spp. Furthermore, low grazing of Ectocarpales in the laboratory and high biomass of these filamentous brown algae in the field indicate a preference for the more palatable green algae Ulva spp. This study indicates that the high grazing capacity of G. locusta, in combination with high reproduction and growth rates, would allow the amphipod to play a key role in Z. marina ecosystems by controlling destructive blooms of filamentous green algae. However, high predation pressure appears to prevent large populations of G. locusta in eelgrass beds on the Swedish west coast today.     

Is benthic trophic structure affected by food quality? The Dogger Bank example

The aim of this study was to evaluate the relationship between food quality and trophic structure of macrobenthic communities of the Dogger Bank. Therefore, 21 stations along different depth gradients (a transect across the central Dogger Bank and at a station cross in the north-eastern part) were sampled and analysed for macrofauna trophic structure and food supply, indicated by phytopigment composition and organic matter contents of the sediment. The study revealed differences in the trophic structure of the benthic community in areas with contrasting food supply. The trophic structure reflected differences in relative quality of organic matter. The sandy sediments on top of the bank receiving intermediate quantities of high food quality were inhabited by high abundances of interface feeders and sand-licking amphipods of the genus Bathyporeia. The southern stations characterised by a comparatively high amount of intermediate food quality were dominated by interface feeders such as Amphiura filiformis. The northern and north-eastern stations showing the highest diversity of feeding types were characterised by a comparatively low, but highly variable input of intermediate-quality organic matter consisting of both fresh and degraded organic matter. Results are discussed with a special emphasis on specific Dogger Bank features such as benthic primary production and frontal system characteristics.Communicated by O. Kinne, Oldendorf/Luhe     

Utilization of a fumigated sediment by two benthic deposit-feeders:Abra alba (Mollusca: Bivalvia) andEupolymnia nebulosa (Annelida: Polychaeta)

The present study tested the utilization of dead microbial biomass by two benthic deposit-feeders:Abra alba (Wood) (Mollusca: Bivalvia) andEupolymnia nebulosa (Montagu) (Annelida: Polychaeta). Clams were collected in the Canet lagoon during spring 1989. Worms were collected in the Port-Vendres harbour during spring 1989. The¹⁴C-labelled (glutamic acid, 24 h) sediment used during the study was sterilized with 1% chloroform, washed with sterile seawater, and dried (60°C; 48 h). This sterilisation procedure, called fumigation is the least harmful to the sediment (Novitsky 1986). Both clams and worms were incubated in the presence of the fumigated sediment for 5, 10, 20, and 50 h. At the end of each experiment we recorded the radioactivity in four compartments: (1) sediment, (2) dissolved organic matter (DOM), (3) CO₂, and (4) animals. The radioactivity of the sediment was subdivided into five fractions: (i) soluble in 2N HCl, (ii) soluble in hot 5% trichloroacetic acid (TCA), (iii) soluble in 1N NaOH, (iv) soluble in hot 6N HCl, (v) residual (after combustion in a Leco carbon analyser). In the first set of experiments, after 20 h of incubation, 5.4 and 4.7% of the total radioactivity was taken up by clams and worms, respectively. However, a model revealed that this uptake could have been correlated with the release of radiolabelled DOM (33% of total radioactivity during the first 5 h). In order to test this assumption, we used the same protocol with three additional washes of the fumigated sediment. This resulted in a significantly lower uptake by the clams (1.9% of the total radioactivity byt = 50 h), whereas the worms exhibited an uptake similar to that in the initial experiment (5.1% of total radioactivity byt = 50 h). These results underline the importance of considering interactions with DOM when applying radiotracer techniques to the study of benthic food chains. The average ingestion rates of fumigated sediment byA. alba andE. nebulosa were 5.2 10^–2 mg sediment dry wt mg^–1 clam h^–1 and 3.5 10^–2 mg sediment dry wt mg^–1 worm h^–1, respectively, which is comparable to previous data reported for other deposit-feeding bivalves and polychaetes feeding on natural sediment or detritus. The low radioactivity recorded for CO₂ together with the similarity of the changes in the partitioning of the radioactivity within the sediment between control experiments and experiments carried out in the presence of clams or worms suggest low assimilation efficiencies. Therefore, the present study supports the fact that dead microbial biomass does not constitute an important food source for benthic deposit-feeders.     

Large-scale estimate of somatic growth in Amphiura filiformis (Echinodermata: Ophiuroidea)

Quantitative data on size structure of the ophiuroid Amphiura filiformis (O.F. Müller) from 35 benthic stations in the Kattegat sampled twice with a 143 d interval, June and October 1991, were used to estimate somatic growth. The material was objectively divided into cohorts, and cohorts, from the two occasions were paired to give estimates of growth. The growth constant, K, in the Von Bertalanffy equation, was estimated from a Ford-Walford plot to 0.54 yr^-1. Results were in agreement with previous estimates from a few single sites, and suggest that the main part of dise growth occurs within the first 5 to 7 yr of living. Size specific growth in oral width was density independent despite high densities of A. filiformis (>3000 ind m^-2) and high total benthic biomass (up to 1000 g wet wt m^-2) in some areas. Growth was uncorrelated with plant pigment concentrations in the sediment and showed weak positive correlation with sediment carbon and nitrogen, as well as water depth. Growth was higher in fine sediments. This is the first attempt to estimate growth in this important species over a large area and to relate growth in the field to environmental factors.     

Parasitism as a determinant of community structure on intertidal flats

The burrowing and movement ability of the New Zealand cockle Austrovenus stutchburyi is reduced when infected by echinostome trematodes. Previous experimental evidence from a single site suggests that this parasite-induced behavioural change of a key bivalve can affect the structure of the surrounding benthic community. By using multiple regression analyses on data collected from 17 intertidal flats, we here show that cockle parasitism is associated with macrozoobenthic community structure on a larger spatial scale. Regressions were performed for animal abundance, biomass, species diversity and species richness separately, entering cockle parasitism (infection intensity), presence/absence of ghost shrimps (Callianassa filholi), cockle density, primary producer abundance and organic content, particle size, sorting coefficient and gravel content of the substrate as predictors. Next to ghost shrimps, cockle parasitism was the best predictor of animal abundance by affecting (mainly positively) 8 of the 49 most widespread species significantly. Cockle parasitism was also associated with the biomass of anthozoans (positively), nemerteans (negatively) and bivalves (positively), whereas overall animal biomass was positively related to the sorting coefficient of the substrate. Species diversity was positively associated with cockle parasitism and gravel content of the substrate. Species richness was significantly associated with cockle parasitism (positively), ghost shrimps (negatively) and abundance of primary producers (positively) in combination. The impact of cockle parasitism on benthic community structure is believed governed directly or indirectly by (1) reduced sediment disturbance, (2) increased surface structural complexity and (3) availability of larval trematodes as an additional food source.     

Ecological growth strategies in the seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae): Soluble nitrogen and reserve carbohydrates

The seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae) were grown in an outdoor continuous-flow system at both ambient incident light (I₀) and 0.13 I₀. During the winter, both species accumulated substantial soluble nitrogen reserves (up to 1020 g-at N·g dry wt^-1 in G. foliifera and 630 g-at N·g dry wt^-1 in Ulva sp.). The rate at which these N reserves were depleted was proportional to the growth rate. Seaweeds grown at 0.13 I₀ had lower growth rates and higher levels of soluble tissue N than plants grown at I₀. During the spring-summer growing season, peaks in tissue N followed nutrient peaks in the ambient seawater. Ulva sp. had higher nutrient uptake and growth rates than G. foliifera and showed greater fluctuations in soluble tissue N. This may characterize opportunistic seaweed species with high biomass turnover rates. At I₀, the levels of starch (up to 340 mg·g dry wt^-1 in G. foliifera and 170 mg·g dry wt^-1 in Ulva sp.) were highest during the spring and summer. During this period, fluctuations in starch content were inversely related to growth rate and soluble tissue N. Seaweeds grown at 0.13 I₀ did not accumulate starch. Neither species was found to overwinter with starch reserves.     

Effects of mussel aquaculture on the nitrogen cycle and benthic communities in Kenepuru Sound,Marlborough Sounds,New Zealand

Nitrogen pools and transformations and benthic communities at a Perna canaliculus farm and a nearby reference site without direct influence of marine farming in Kenepuru Sound, New Zealand, were compared on four dates between September 1982 and May 1983. The organic nitrogen pool in the top 12 cm sediment was 7.4 to 10.8 mol m^-2 at the mussel farm and 6.1 to 8.9 mol m^-2 at the reference site. The nitrate and nitrite pools were similar in both sediments, but the ammonium pool in the mussel farm sediment was about twice as high as in the reference sediment. In January, the sediment ammonium concentrations ranged from 418 nmol cm^-3 (surface) to 149 nmol cm^-3 (12 cm depth) at the mussel farm and from 86 to 112 nmol cm^-3 at the reference site. The molar C:N ratio of the sediment organic matter was 6.2 to 7.2 at the mussel farm and 7.9 to 10.0 at the reference site. The molar N:P ratio of the sediment organic matter was 4.3 to 7.2 and 3.3 to 6.1 at mussel farm and reference site, respectively. The total nitrogen mineralisation rate in the top 12 cm sediment ranged from 21.7 to 37.1 mmol m^-2 d^-1 at the mussel farm and from 8.5 to 25.0 mmol m^-2 d^-1 at the reference site. Ammonium excretion by mussels was about 4.7% (January) and 7.4% (May) of the combined nitrogen mineralisation by mussels and sediment. The sediment-denitrification rate was 0.7 to 6.1 mmol m^-2 d^-1 at the mussel farm and 0.1 to 0.9 mmol m^-2 d^-1 at the reference site. In January, 76 and 93% of the nitrate reduced in the sediments were denitrified at the mussel farm and reference site, respectively. The denitrification rate on the mussel lines (determined on detritus-covered mussels) was twice the mussel farm sediment-denitrification rate and 10 times the reference sediment-denitrification rate. Total denitrification at the mussel farm was 21% higher than at the reference site. The loss of nitrogen through mussel harvest and denitrification was 68% higher at the mussel farm. The surface layers of both sediments contained about 75 mg m^-2 chlorophyll a. Sediment phaeophytin levels were 52 mg m^-2 at the reference site and 137 mg m^-2 at the mussel farm. While the benthic infauna of the mussel-farm sediment consisted only of polychaete worms, the reference sediment contained also bivalve molluscs, brittle stars and crustaceans.     

Energy flow model of an oxygen-deficient estuary on the Swedish west coast

The estuary Byfjord (Sweden) is characterized by high primary production, a well developed meiofauna compared to the macrofauna, high epifaunal biomass, a low number of herbivorous copepods and a small fish stock. A simplified energy flow model of the ecosystem of the fjord is given. The energy transfer is approximated to 15%. About one-fourth-300 (metric) tons of carbon — of the annual primary production is suggested to be directly consumed and to produce 5 tons of zooplankton carbon and 40 tons of epifaunal (mainly Mytilus edulis) carbon. About 500 tons of carbon from the detritus pool are probably utilized in animal production. This amount will produce 5 tons of zooplankton carbon, 6 tons of meiofaunal carbon, and 3 tons of carbon from the benthic macrofauna. Production of fish is estimated at 0.3 ton carbon per year. M. edulis seems to be the only food resource in the fjord worth harvesting by man.