Indirect effects of Atrina zelandica on water column nitrogen and oxygen fluxes: The role of benthic macrofauna and microphytes

The effects of ecosystem engineers can be mediated by direct and indirect interactions. For suspension-feeding bivalves that contribute to benthic-pelagic coupling, indirect effects have been linked to hydrodynamics; however, the influence of these ecosystem engineers may also operate through interactions with surrounding sediments, microphytes and macrofauna that, in turn, affect nutrient and oxygen fluxes. This study investigated the indirect effects of an epibenthic suspension-feeding bivalve (Atrina zelandica) on ammonium and nitrate-nitrite effluxes from the sediment, and oxygen consumption in the overlying waters, under dark conditions, at two sites with different environmental characteristics. Location-dependent effects were observed in the relative strength of the effect of Atrina on microphyte and macrofaunal abundance. The difference between the strength of the effect of Atrina on macrofauna between sites was not driven by a single species or type of species; rather all the species decreasing in abundance away from Atrina were small species that utilised the sediment water interface. Location-dependent effects were also observed in the relative strength of the effect of microphyte and macrofaunal abundance on oxygen and nutrient fluxes. While microphytes were an important consumer of oxygen at one site, at the other site, small infaunal macrofauna were important. Similarly, the strength of the effect of surrounding macrofauna on ammonium efflux varied between sites. These findings demonstrate the importance of natural history and species functions for understanding complex responses. They suggest that indirect effects by key benthic macrofaunal species in marine systems can also be important to benthic-pelagic coupling. In particular, while key species are often large, excretion and respiration of smaller macrofauna can be important to the exchange of nutrients near the seafloor and oxygen consumption in the benthic boundary layer.     

Context-Specific Bioturbation Mediates Changes to Ecosystem Functioning

Species are often grouped according to their biological or functional traits to better understand their contribution to ecosystem functioning. However, it is becoming clear that a single species can perform different roles in different habitats. Austrohelice crassa, a burrow-building mud crab shifts its primary bioturbational role to that of a vertical mixer in non-cohesive sediments as frequent burrow collapse greatly enhances sediment reworking. We conducted in situ crab density manipulations in two sediment environments (a non-cohesive sand and a cohesive muddy-sand) to examine if the context-specific functional roles were linked to changes in solute fluxes across the sediment–water interface. Across both habitats, we show that A. crassa regulated nutrient cycling, creating strong density driven effects on solute exchanges. Increasing crab density increased sediment O₂ demand and the flux of NH₄ ⁺ from the sediment, indicating much of the response was physiologically driven. Clear interactions between A. crassa and microphytobenthos were also detected in both habitats. Despite lowering microphyte standing stock through deposit feeding, A. crassa increased benthic primary production per unit of chlorophyll a. Our experiment also revealed important context-specific differences, most notably for NH₄ ⁺ fluxes, which were higher where burrows and their associated microbial communities were most stable (muddy-sand). This study highlights the need to integrate interactions between organism behavior and habitat type into functional group studies to broaden conceptual frameworks and avoid oversimplification of highly complex organism–sediment interactions.     

Macrofauna communities of the Dogger Bank (central North Sea) in the late 1990s: spatial distribution,species composition and trophic structure

Macrofauna samples were taken in May 1996–1998 at 28 stations on the Dogger Bank (central North Sea) to describe species composition, spatial distribution and trophic structure of the macrofaunal communities. In general, five communities could be distinguished depending on the influence of different water masses, depth, type of sediment and food availability. Except for those species which feed in the sediment, macrofauna communities did not show any correlation to the measured sediment variables. In fact, macrofauna communities of the Dogger Bank were mainly structured by lateral food supply via frontal systems. This was indicated by the dominance of interface feeders foraging in the benthic boundary layer, which are able to switch from surface deposit to suspension feeding depending on flow conditions. The importance of sandlickers in the shallowest parts of the Dogger Bank hints at an important contribution of benthic primary production to the nutritional and energetic needs of the benthic community. Electronic Publication     

Assessment of surface sediment dynamics and response of benthic macrofauna assemblages in Boughrara Lagoon (SW Mediterranean Sea)

A study was conducted in the spring of 2009, the winters of 2010 and 2013, and in the summer of 2012 at 13 stations in Boughrara Lagoon, Tunisia (southern Mediterranean). The country⿿s largest lagoon, it is considered to be an anthropogenically stressed area, though a major tourist centre⿿Djerba Island⿿is located along its northern shores. The lagoon bottoms were studied via analyses of grain size, surface sediment composition, total organic matter (TOM) and through the trophic and functional organisation of benthic macrofauna. The results indicate that the bottoms are composed of fine, medium or coarse sands and that sediment distribution is controlled by water movement. Estimation of TOM content revealed that the studied samples present both normal and imbalanced sediments. The structure and organisation of the lagoon⿿s benthic macrofauna are dominated by select deposit feeders and underwent significant changes during the period 2010⿿2013.Subjected for decades to increased pollution due to growing human activities in the surrounding area, Boughrara Lagoon now appears to be impacted by certain environmental/anthropogenic stressors, as indicated by the presence of pollution-tolerant bio-indicator species in the imbalanced area. The response of the lagoon ecosystem to changes in benthic sediment deposition provides a potential assessment tool for similar habitats elsewhere.     

Large Spatial Scale Variability in Bathyal Macrobenthos Abundance,Biomass, α- and β-Diversity along the Mediterranean Continental Margin

The large-scale deep-sea biodiversity distribution of the benthic fauna was explored in the Mediterranean Sea, which can be seen as a miniature model of the oceans of the world. Within the framework of the BIOFUN project (“Biodiversity and Ecosystem Functioning in Contrasting Southern European Deep-sea Environments: from viruses to megafauna”), we investigated the large spatial scale variability (over >1,000 km) of the bathyal macrofauna communities that inhabit the Mediterranean basin, and their relationships with the environmental variables. The macrofauna abundance, biomass, community structure and functional diversity were analysed and the α-diversity and β-diversity were estimated across six selected slope areas at different longitudes and along three main depths. The macrobenthic standing stock and α-diversity were lower in the deep-sea sediments of the eastern Mediterranean basin, compared to the western and central basins. The macrofaunal standing stock and diversity decreased significantly from the upper bathyal to the lower bathyal slope stations. The major changes in the community composition of the higher taxa and in the trophic (functional) structure occurred at different longitudes, rather than at increasing water depth. For the β-diversity, very high dissimilarities emerged at all levels: (i) between basins; (ii) between slopes within the same basin; and (iii) between stations at different depths; this therefore demonstrates the high macrofaunal diversity of the Mediterranean basins at large spatial scales. Overall, the food sources (i.e., quantity and quality) that characterised the west, central and eastern Mediterranean basins, as well as sediment grain size, appear to influence the macrobenthic standing stock and the biodiversity along the different slope areas.     

Minimum requirements for modelling bivalve carrying capacity

The concept of carrying capacity of an ecosystem fornatural populations is derived from the logisticgrowth curve in population ecology, and defined as themaximum standing stock that can be supported by agiven ecosystem for a given time. This definitionneeds to be modified for the exploitation ofecosystems. Carrying capacity for exploitation isdefined as the standing stock at which the annualproduction of a marketable cohort is maximized. Forbivalve suspension feeders, the dominant factordetermining the exploitation carrying capacity at theecosystem scale is primary production. At a localscale carrying capacity depends on physicalconstraints such as substrate, shelter and food supplyby tidal currents.We evaluate critically some existing models ofexploited ecosystems for shellfish cultivation inorder to formulate the minimum requirements of ageneric carrying capacity model. Generic models canbe developed for both the ecosystem scale and thelocal scale, depending on the aim of the modelling.Transport processes, sediment dynamics and submodelsfor organism and population level processes areminimum requirements for carrying capacity modelling. This revised version was published online in August 2006 with corrections to the Cover Date.     

Food-chain relationships in subtidal silty sand marine sediments and the role of meiofauna in stimulating bacterial productivity

Summary From bibliographic data the biomass correlations (organic dry weight) are constructed for the subsurface layer of a hypothetical 30 m deep silty sand station: 200 g/ml macrofauna (including 120 g/ml subsurface deposit feeders), 50 g/ml meiofauna, 20 g/ml Foraminifera, 1 g/ml Ciliata and Flagellata, and 100 g/ml bacteria. ATP-biomass is discussed.Meiofauna and Foraminifera contribute with 30 and 12% to the living biomass in the sediment, and it is assumed that their contribution to the food of deposit-feeding macrofauna is of a similar percentage. This is corroborated by productivity estimations.Bacteria are the main food of deposit feeding macrofauna, meiofauna, and microfauna. From different calculations it becomes evident that the productivity of bacteria in the sediment is far below figures achieved in experimental cultures: the conclusion is that sediment bacteria, in general, do not live under good environmental conditions.A rather large part of the bacterial population in the sediment seems to be in the stationary phase of life, and only a fraction of the total population exhibits high metabolic rates and rapid duplications. Only these active bacteria are of importance for the breakdown of relatively refractive organic matter in the sediment.In soft bottom marine sediments where the input of organic matter is higher than the remineralization rate, benthic animals stimulate by their activities and by nutrient cycling the decomposition of detritus via bacteria. Though meiofauna, in principle, feeds upon the same food resource as macrofauna, there is no real competition for food, because meiofaunal animals by their activities and by excreting metabolic end products induce a bacterial productivity which would not be there without them, and feed on it. There are a few examples where more specialized interactions exist between benthic animals and bacteria; these interactions have been termed gardening. They could be highly important in the benthic ecosystem.     

Weak and Habitat-Dependent Effects of Nutrient Pollution on Macrofaunal Communities of Southeast Australian Estuaries

Among the impacts of coastal settlements to estuaries, nutrient pollution is often singled out as a leading cause of modification to the ecological communities of soft sediments. Through sampling of 48 sites, distributed among 16 estuaries of New South Wales, Australia, we tested the hypotheses that (1) anthropogenic nutrient loads would be a better predictor of macrofaunal communities than estuarine geomorphology or local sediment characteristics; and (2) local environmental context, as determined largely by sediment characteristics, would modify the relationship between nutrient loading and community composition. Contrary to the hypothesis, multivariate multiple regression analyses revealed that sediment grain size was the best predictor of macrofaunal assemblage composition. When samples were stratified according to median grain size, relationships between faunal communities and nitrogen loading and latitude emerged, but only among estuaries with sandier sediments. In these estuaries, capitellid and nereid polychaetes and chironomid larvae were the taxa that showed the strongest correlations with nutrient loading. Overall, this study failed to provide evidence of a differential relationship between diffuse nutrient enrichment and benthic macrofauna across a gradient of 7° of latitude and 4°C temperature. Nevertheless, as human population growth continues to place increasing pressure on southeast Australian estuaries, manipulative field studies examining when and where nutrient loading will lead to significant changes in estuarine community structure are needed.     

Effects of Deposit-Feeding Macrofauna on Benthic Bacteria,Viruses, and Protozoa in a Silty Freshwater Sediment

In microcosm experiments, we simultaneously tested the effects of increased numbers of deposit-feeding macrofauna (chironomids, oligochaetes and cladocerans) on the standing stock, activities and interactions of heterotrophic bacteria, viruses, and bacterivorous protozoa (heterotrophic nanoflagellates and ciliates) in the aerobic layer of a silty littoral freshwater sediment. On average, bacterial secondary production was stimulated between 11 and 29% by all macrofaunal groups compared to control experiments without macrofauna addition. Bacterial standing stock increased significantly by 8 and 13% in case of chironomids and cladocerans, respectively. Oligochaetes and chironomids produced significant negative effects on viral abundance while the results with cladocerans were inconsistent. The addition of oligochaetes and chironomids resulted in a significant decrease by on average 68 and 32% of viral decay rates, respectively, used as a measure of viral production. The calculated contribution of virus-induced lysis to benthic bacterial mortality was low, with 2.8 to 11.8% of bacterial secondary production, and decreased by 39 to 81% after the addition of macrofauna compared to the control. The abundances of heterotrophic nanoflagellates were significantly reduced by 20% by all tested macrofauna groups, while ciliates showed inconsistent results. The importance of heterotrophic nanoflagellate grazing on benthic bacteria was very low (<1% of bacterial secondary production) and was further reduced by elevated numbers of macrofauna. Thus, the selected deposit feeding macrofauna groups seem to have several direct and indirect and partly antagonistic effects on the benthic bacterial compartment through the enhancement of bacterial production and the reduction of virus-induced cell lysis and protozoan grazing.     

Dynamics of microbial biomass and activity in five habitats of the Okefenokee Swamp ecosystem

A variety of freshwater marsh and swamp habitats are found interspersed in a mosaic pattern throughout the Okefenokee Swamp, Georgia, USA. We examined spatial and temporal patterns in standing stocks and activity in the microbial community of five habitats within this heterogeneous ecosystem. Standing stock dynamics were studied by measuring microbial biomass (ATP) and bacterial numbers (AODC) in both water and sediments over a 14 month period. Abundance varied temporally, being generally lower in winter months than in spring and summer months. However, a large proportion of the measured variability was not correlated with temporal patterns in temperature or with bulk nutrient levels. Spatial variability was characteristic of the Okefenokee at a variety of large and small scales. Habitat-level heterogeneity was evident when microbial standing stocks and activity (measured as [¹⁴C]lignocellulose mineralization) were compared across the five communities, although abundance differences among sites were restricted to nonwinter months when microbial biomass was high. Spatial variation within habitats was also found; patches of surface sediment with differing microbial activity or abundance were measured at scales from 30 cm to 150 m.     

Empirical Evidence Reveals Seasonally Dependent Reduction in Nitrification in Coastal Sediments Subjected to Near Future Ocean Acidification

Research so far has provided little evidence that benthic biogeochemical cycling is affected by ocean acidification under realistic climate change scenarios. We measured nutrient exchange and sediment community oxygen consumption (SCOC) rates to estimate nitrification in natural coastal permeable and fine sandy sediments under pre-phytoplankton bloom and bloom conditions. Ocean acidification, as mimicked in the laboratory by a realistic pH decrease of 0.3, significantly reduced SCOC on average by 60% and benthic nitrification rates on average by 94% in both sediment types in February (pre-bloom period), but not in April (bloom period). No changes in macrofauna functional community (density, structural and functional diversity) were observed between ambient and acidified conditions, suggesting that changes in benthic biogeochemical cycling were predominantly mediated by changes in the activity of the microbial community during the short-term incubations (14 days), rather than by changes in engineering effects of bioturbating and bio-irrigating macrofauna. As benthic nitrification makes up the gross of ocean nitrification, a slowdown of this nitrogen cycling pathway in both permeable and fine sediments in winter, could therefore have global impacts on coupled nitrification-denitrification and hence eventually on pelagic nutrient availability.     

Benthic fauna bio-irrigation effects on nutrient regeneration in fish farm sediments

Impacts of organic enrichment and a modified benthic fauna community (caused by fish farming) on benthic mineralization rates and nutrient cycling were studied in sediments at one Danish and one Cypriote fish farm. Sediment organic matter concentration and macrofauna community composition were manipulated in microcosms and changes in total benthic metabolism (oxygen consumption, TCO₂ production), anaerobic metabolism (sulfate reduction rates), nutrient fluxes and sediment parameters were followed for a period of 3 weeks. Mineralization rates were found to be highly correlated with irrigation velocities and largest fauna effects were found in the Danish sediments with the large and active irrigating climax species (Nereis diversicolor and Macoma balthica). Eastern Mediterranean climax species (Glycera rouxii and Naineris laevigata) also stimulated mineralization rates but to a smaller extent due to lower irrigation, whereas the opportunistic species (Capitella in Danish sediment and Hermodice carunculata in Cypriote sediment) showed less effect on mineralization. Ammonium and phosphate release increased with increasing irrigation velocities, but much less in Cyprus indicating higher nutrient retention at the ultra-oligotrophic location compared to eutrophic Danish site. Irrigation velocities, and thus mineralization rates, increased by organic matter loading, indicating larger fauna-induced oxidation in enriched environments. The result implies that a change in fauna structure in fish farm sediment towards smaller opportunistic polychaete species with lower irrigation will result in slower mineralization rates and potentially increase accumulation of organic waste products.     

Paleoecology of some molluscan assemblages from the badenian (Miocene) marine sandy facies of Poland

Three molluscan assemblages from the Badenian (Miocene) marine sandy facies of Poland are described in terms of their taxonomic composition, diversity, and trophic structure. The structural variation between the molluscan assemblages seems to be largely controlled by a gradient in water energy which in turn results in gradients in substrate mobility and organic content of bottom sediments. Together with a subordinate factor of local sedimentation rate, these gradients control the distribution of lucinoid mucus tube feeders vs. deep burrowing siphonate suspension feeders, of browsers, and of deposit feeders vs. suspension feeders. Some structural features of the molluscan assemblages are regarded as related to biotic factors. The graphs of cumulative species frequencies vs. the logarithm of cumulative individual frequencies are used as a measure of community organization. These graphs indicate distinct differences in ecological maturity between the molluscan assemblages. On this basis, two different stages of ecological succession in an offshore sandy bottom environment are recognized. The evidence from the comparison of the Miocene molluscan assemblages with their recent counterparts may support the view that the evolutionary stability of species populations of a given community, and long-term stability of the community structure are independent. High environmental variability and stress seem to be correlated with evolutionary stability of species populations rather than with long-term permanence of community structure.     

Effect of shade and shading history on species abundances and ecosystem processes in temporary ponds

1. The amount of sunlight that an ecosystem receives is an important determinant of primary productivity, which in turn can influence species diversity and nutrient cycling. Here, we examine the effects of shading and shading history on ecosystem processes, macroinvertebrate diversity and development of dominant ecosystem states in field‐based aquatic mesocosms. 2. We found that there were large effects of the level of shade, but few effects of shading history. Increasing light increased the biomass of filamentous algae (metaphyton) which increased the overall productivity of the ecosystem, and shifted the invertebrate community from one with more mosquitoes (filter feeders) to one with more anuran tadpoles (algal grazers). 3. There was also an effect of shading history, where increased shade led to changes in the macroinvertebrate communities that were maintained after shade was reduced in the later part of the experiment. 4. Finally, our results indicated that correlations between ecosystem processes, specific key macrofauna and the development of pond ecosystem states were greater than correlations of these factors to shading treatments. These results suggest that the history of community assembly can have a greater impact on the development of ecosystem processes than diminishing light by as much as 50%. However, light may have a potentially strong indirect effect and may impact the communities through altered bottom‐up structuring forces.     

Species effects on ecosystem processes are modified by faunal responses to habitat composition

Heterogeneity is a well-recognized feature of natural environments, and the spatial distribution and movement of individual species is primarily driven by resource requirements. In laboratory experiments designed to explore how different species drive ecosystem processes, such as nutrient release, habitat heterogeneity is often seen as something which must be rigorously controlled for. Most small experimental systems are therefore spatially homogeneous, and the link between environmental heterogeneity and its effects on the redistribution of individuals and species, and on ecosystem processes, has not been fully explored. In this paper, we used a mesocosm system to investigate the relationship between habitat composition, species movement and sediment nutrient release for each of four functionally contrasting species of marine benthic invertebrate macrofauna. For each species, various habitat configurations were generated by selectively enriching patches of sediment with macroalgae, a natural source of spatial variability in intertidal mudflats. We found that the direction and extent of faunal movement between patches differs with species identity, density and habitat composition. Combinations of these factors lead to concomitant changes in nutrient release, such that habitat composition effects are modified by species identity (in the case of NH4-N) and by species density (in the case of PO4-P). It is clear that failure to accommodate natural patterns of spatial heterogeneity in such studies may result in an incomplete understanding of system behaviour. This will be particularly important for future experiments designed to explore the effects of species richness on ecosystem processes, where the complex interactions reported here for single species may be compounded when species are brought together in multi-species combinations.     

Interaction effects of fish, nutrients, mixing and sediments on autotrophic picoplankton and algal composition

1. We examined the effects of nutrients, turbulent mixing, mosquitofish, Gambusia affinis Baird and Girard and sediments on algal composition, algal biomass and autotrophic picoplankton (APP) abundance in a 6-week experiment of factorial design in twenty-four 5-m³ outdoor mesocosms during late autumn 1995.
2. Turbulent mixing decreased surface temperature and increased turbidity, which also was increased by the addition of sediments. Total algal biomass was significantly enhanced by nutrients and mixing, and decreased by the sediment treatment. In the mixing × nutrient treatment, algal biomass increased more than expected from the individual effects, while the fish × mixing and mixing × sediment treatments increased algal biomass less than expected.
3.  Cryptomonas (cryptomonad) blooms were observed in the unmixed, high nutrient treatment; Synedra (diatom) blooms were observed in the high nutrient, high sediment treatment; Ulothrix (green algae) blooms were observed in the mixed, high nutrient, low sediment treatment.
4. Eukaryotic APP abundances were increased by sediment addition and by turbulent mixing, and increased synergistically by mixing × sediment and mixing × nutrient interactions. Prokaryotic APP abundances were decreased by nutrient enhancement and by a mixing × nutrient interaction. There were no main effects of fish on APP abundance, but fish were involved in some of the two–way interactions.
5. The large number of significant interaction effects indicates that APP and other phytoplankton are regulated by a complex set of interdependent factors which should be considered simultaneously in studies of phytoplankton population dynamics and community composition.     

Trophic structure of macrobenthos in the Tagus estuary and adjacent coastal shelf

The trophic structure of benthic communities in the Tagus estuary and adjacent coastal shelf was characterized according to a functional guild approach, based on sampling surveys conducted between 1987 and 2000. Macrobenthic organisms were assigned to seven distinct trophic groups (herbivorous, filter feeders, surface deposit feeders, subsurface deposit feeders, carnivores, filter feeders/detritivores, carnivores/detritivores) and the dominance of these groups was related to environmental variables using multivariate ordination techniques. Surface-deposit feeders were numerically dominant in the Tagus estuary, making up 52% of the benthic communities, while in the adjacent coastal shelf the assemblage was dominated by both surface-deposit feeders and filter feeders (37% and 33%, respectively). When biomass was considered, filter feeders and filter feeders/detritivores were the dominant groups in the estuary, while for the adjacent coastal shelf filter feeders represented 83% of the total biomass. Salinity, depth and sediment composition were the main factors structuring spatial distribution. Surface-deposit feeders were the most abundant macrobenthos of the upper estuary. Surface deposit feeders also dominated the middle and the lower estuary but the proportion of filter feeders as well as other trophic groups increased with salinity. Generally, a more even distribution of trophic structure was found at stations with high salinity. In the adjacent coastal shelf, the trophic diversity decreased with depth. The trophic structure revealed that filter feeders dominated in abundance and biomass in shallow sandy sediments (<25 m), while in deeper sandy mud and muddy habitats (>50 m to 260 m), deposit feeders and carnivores were the most important groups in abundance and biomass, respectively.     

Composition, abundance and stratification of soft-bottom macrobenthos from selected areas of the Ross Sea shelf (Antarctica)

Composition, abundance and stratification of soft-bottom macrobenthos were studied at three selected sites on the Ross Sea shelf (Antarctica) with different geomorphology and sedimentation regimes. Sites A (southwest Ross Sea, 810 m depth) and B (Joides basin, 580 m depth) were characterized by biogenic mud and clay sediments, whereas site C (Mawson bank, 450 m depth) featured sandy sediments mixed with a conspicuous biogenic component characterized by shells and tests of calcareous invertebrates (mainly barnacles of the genus Bathylasma). The macrofauna of sites A and B was mainly composed of infaunal polychaetes and bivalves. The assemblages comprised both surface and sub-surface deposit feeders, including some conveyor-belt polychaetes (Maldanidae and Capitellidae) that are responsible for high sediment mixing and bioturbation. The macrobenthos of site C was dominated by crustaceans, polychaetes and echinoderms (ophiuroids), and mainly by filter feeders and epifaunal or interstitial forms. Abundances were higher (up to 1040 ind. m⁻²) at site B than at sites A and C (430 and 516 ind. m⁻², respectively). At sites A and B the benthos was mainly concentrated in the upper 5 cm of the sediment, and abundances declined sharply in the deeper sediment layers. These results indicate a high degree of consistency between sediment features and benthic community structure, which are both strongly related to local hydrography and bottom dynamics. Sites A and B represent areas where the organic input to the seafloor by vertical sedimentation from the upper water column is high. Site C, however, is characterized by high sediment instability and food particles advecting mainly horizontally. The community is more physically controlled by unpredictable, and probably frequent, disturbance events (e.g., bottom turbid currents, sediment reworking and displacement). Individuals were relatively small, indicating that probably they are not able to grow up to the adult size and reproduce. The community may be represented by “pseudopopulations” depending on the settlement of larvae invading from neighbouring areas. Accepted: 23 October 1998     

Population Dynamics of the Polychaetous Annelids of an Intertidal Habitat in Upper Old Tampa Bay,Florida

Following an extensive red-tide induced mass mortality of the benthic macrofauna of a sandy intertidal habitat, the population dynamics of the polychaete species were studied. Detailed studies of the 12 most abundant species are presented. Data concerning total population levels, reproduction, recruitment, distribution within the intertidal zone, and gut content analyses are integrated in order to explain the observed spatial and temporal patterns of distribution and abundance. Potential competive interactions for food are considered to be the most important factor for explaining the observed ecological patterns. The polychaete species studied are divided into three trophic guilds: an omnivorous guild that feeds predaciously and as non-selective deposit feeders, a surface feeding guild consisting of species usually considered to be selective surface deposit feeders, and a subsurface feeding guild usually considered as non-selective infaunal deposit feeders. Within and between guild interactions are discussed.