Indicators of the Ecological Impact of Bottom-Trawl Disturbance on Seabed Communities

The Ecosystem Approach to Fisheries requires that managers take account of the environmental impacts of fishing. We develop linked state and pressure indicators that show the impact of bottom-trawling on benthic communities. The state indicator measures the proportion of an area where benthic invertebrate biomass (B) or production (P) is more than 90% of pristine benthic biomass (B _0.9) or production (P_0.9). The pressure indicator measures the proportion of the area where trawling frequency is sufficiently high to prevent reaching predicted B_0.9 or P_0.9. Time to recovery to B_0.9 and P_0.9 after trawling, depending on the habitat, was estimated using a validated size-based model of the benthic community. Based on trawling intensity in 2003, 53.5% of the southern North Sea was trawled too frequently for biomass to reach B_0.9, and 27.1% was trawled too frequently for production to reach P_0.9. As a result of bottom-trawling in 2003, in 56% of the southern North Sea benthic biomass was below B_0.9, whereas in 27% of the southern North Sea benthic production was below P_0.9. Modeled recovery times were comparable to literature estimates (2.5 to more than 6 years). The advantages of using the area with an ecological impact of trawling as a pressure indicator are that it is conceptually easy to understand, it responds quickly to changes in management action, it can be implemented at a relevant scale for fisheries management, and the necessary effort distribution data are centrally collected. One of this approach’s greatest utilities, therefore, will be to communicate to policy makers and fishing enterprises the expected medium- to long-term ecological benefits that will accrue if the frequency of trawling in particular parts of fishing grounds is reduced.     

Habitat-Specific Effects of Fishing Disturbance on Benthic Species Richness in Marine Soft Sediments

Around the globe, marine soft sediments on continental shelves are affected by bottom trawl fisheries. In this study, we explore the effect of this widespread anthropogenic disturbance on the species richness of a benthic ecosystem, along a gradient of bottom trawling intensities. We use data from 80 annually sampled benthic stations in the Dutch part of the North Sea, over a period of 6 years. Trawl disturbance intensity at each sampled location was reconstructed from satellite tracking of fishing vessels. Using a structural equation model, we studied how trawl disturbance intensity relates to benthic species richness, and how the relationship is mediated by total benthic biomass, primary productivity, water depth, and median sediment grain size. Our results show a negative relationship between trawling intensity and species richness. Richness is also negatively related to sediment grain size and primary productivity, and positively related to biomass. Further analysis of our data shows that the negative effects of trawling on richness are limited to relatively species-rich, deep areas with fine sediments. We find no effect of bottom trawling on species richness in shallow areas with coarse bottoms. These condition-dependent effects of trawling suggest that protection of benthic richness might best be achieved by reducing trawling intensity in a strategically chosen fraction of space.     

Cost/benefit and the effect of sample preservation procedures on quantitative patterns in benthic ecology

Some benthic assemblages studies have tested the effects of different preservation procedures on biomass, but their influence on quantitative patterns (number of species and abundance) is still unclear. We evaluated the influence of two sample preservation procedures on quantitative patterns in benthic ecology. Ten sampling points were systematically interspersed on two types of sediment (sandy and muddy). At each sediment type, samples from five sampling points were fixed in 10% formalin, and the other five points were preserved in 70% ethanol (without previous fixation). Three replicates were collected at each sampling point, and samples were washed in 0.5 mesh size and sorted in laboratory. A cost/benefit analysis was performed considering the washing time in laboratory and the costs of substances. A total of 1970 individuals were collected (muddy sediment: 132; sandy sediment: 1838), belonging to 121 taxa (muddy: 49; sandy: 83). Assemblages preserved in ethanol were composed of 795 individuals and 80 taxa, while those fixed with formalin had 1173 individuals and 94 taxa. Polychaeta predominated as the most abundant group for both preservation procedures. For the whole benthic community, significant differences occurred only between sediment types. Significant differences in the number of individuals of polychaetes were observed for the different preservation procedures in sandy sediment. Ethanol has the best cost/benefit ratio in both sediment types due to additional costs to attend safety requirements for formalin-fixed samples. Further studies should evaluate how quantitative patterns are affected by exposure time of preservation, anesthesia interaction, and morphological deformations (e.g. impossibility of identification).     

Scale-dependant control of motile epifaunal community structure along a coral reef fishing gradient

Large-scale fishing is mostly conducted using towed gears that reduce the biomass and diversity of benthic invertebrates. However, it is impossible to differentiate between the physical disturbance effect of towed gears from the effect of fish predator removal upon benthic invertebrate communities. Here we explore the impact of fish removal alone on the community structure of small motile coral reef invertebrates (epifauna) along a subsistence fishing intensity gradient in the Lau group, Fiji. We deployed settlement plates at three areas in each of six fishing grounds and examined the density and class richness of the motile epifaunal communities and the associated algal communities in relation to the structure of fish and benthic communities. Motile epifaunal density was unrelated to fishing intensity. However, at smaller inter-area scale (0.5-10 km) motile epifaunal density was negatively related to plate algal biomass, whereas at the larger inter-fishing-ground scale (4-180 km) motile epifaunal density was positively related to the rugosity (substrate complexity) of the surrounding benthos. The class richness and diversity (Margalef's d) of motile epifaunal communities were negatively related to fishing intensity, but unrelated to grazing intensity, rugosity or algal biomass at either scale. Benthic community structure varied significantly with fishing intensity; hard-coral cover was lower and turf-algal cover was higher at high fishing pressure. The variation in benthic community structure was associated with variation in fish community structure, which in turn varied with fishing intensity. Motile epifaunal community structure upon plates was linked to the structure of the surrounding benthic community, but was not directly linked to the plate algal community. We suggest the decline in richness of the motile epifauna community along the fishing gradient is attributable to either to exploiter-mediated coexistence or the reduction in ‘habitat quality’ of the surrounding benthos. At the large spatial scale substrate complexity is the key determinant of motile epifaunal density, suggesting predation by fishes plays an important structuring role at this scale. Assuming that rugosity is inversely related to predation risk then this study represents the first evidence for spatial-dependence on the top-down (predation) vs. bottom-up (algal biomass) control of community structure. We argue fisheries exploitation, in the absence of a physical disturbance can negatively influence motile epifaunal community structure at large spatial scales.     

Assessing functional redundancy in chronically trawled benthic communities

Fishing disturbance on ecosystems leads to changes in community structure and composition, which may have drastic implications for ecosystem functional performance. Functional redundancy, defined as species sharing similar functional roles, is a community property that plays an important role in preventing functional changes in ecosystems under pressure. In this study, we suggest that functional redundancy may be achieved through trait abundance (i.e. large amounts of a trait, hereafter “common traits”), or through trait richness (i.e. large numbers of distinct taxa exhibiting the same trait, hereafter “widespread traits”). We assessed the variability of both measures obtained from epifaunal and infaunal communities in soft-bottom trawling grounds. Sampling sites were located in four Mediterranean areas that were subjected to different levels of trawling effort. Common and widespread traits measures were based on the analysis of biological traits linked to key soft-bottoms functions such as nutrient cycling, bentho-pelagic coupling and habitat provision. The role of rare species in both measures was also assessed and we observed that, in our study sites, rare species generally exhibited the same traits as the most abundant species. Common and widespread traits measures provided complementary information on benthic functional redundancy. Thus, we suggest that a combination of the two measures should be used to appropriately assess benthic functional redundancy in trawling grounds. As redundancy is a component of ecosystem resilience, functional redundancy evaluation is important to assess the overall integrity of ecosystems.     

Response of Antarctic benthic communities to disturbance: first results from the artificial Benthic Disturbance Experiment on the eastern Weddell Sea Shelf, Antarctica

The long-term benthic disturbance experiment (BENDEX) was started on the eastern Weddell Sea shelf off Austasen (Antarctica) during ‘Polarstern’ cruise ANT XXI/2 in December 2003 to simulate the impact of grounding icebergs on the seabed and follow the steps and timescales of recovery of disturbed benthos and demersal fish communities. Here, we report the basic approach and first results for this experimental field study. By means of 11 densely-placed hauls with a modified bottom trawl, a seabed area of approximately 100 × 1000 m was artificially scoured to inflict a similar damage to the benthic habitats as a grounding iceberg. Before the disturbance event and 11 days after it, the seafloor communities were sampled (invertebrate assemblages by multibox corers, the fish fauna by trawl hauls) and comparatively analyzed. Sediment texture and chemistry was not significantly altered by the heavy disturbance inflicted by repeated trawling, whereas the fauna was negatively affected. Invertebrate benthic biomass was drastically reduced by a factor of 10, while mean abundances were only slightly reduced. Demersal fish biomass and abundance were slightly but not significantly smaller after the disturbance. Effects of disturbance became more evident in the composition of the fish fauna, with Trematomus pennelli and T. hansoni being dominant at disturbed sites, whereas Chionodraco myersi was the dominant species in trawl catches from undisturbed stations.     

Development of filamentous green algae in the benthic algal community in a littoral sand-beach zone of Lake Biwa

Temporal changes of biomass and dominant species in benthic algal communities were investigated in a littoral sand-beach zone in the north basin of Lake Biwa from December 1999 to September 2000. Chlorophyll-a amounts of benthic algal communities per unit area of the sandy sediments rapidly increased from late April to June. Increases in biomass of the benthic algal communities are considered to result from the propagation of filamentous green algae Oedogonium sp. and Spirogyra sp. The cell numbers of filamentous green algae and chlorophyll-a amounts of benthic algal communities at depths of 30 and 50cm at a station protected by a breakwater in May were significantly higher than those of a station exposed directly to wave activity. Thus, the biomass accumulation of the benthic algal communities seems to be regulated strongly by wave disturbance. The development of filamentous green algae may contribute to the increase in biomass of the benthic algal community and to the changes in seasonal patterns of biomass in the sand-beach zone of Lake Biwa. We consider that the development of the filamentous green algal community in the littoral zone of Lake Biwa is the result of eutrophication.     

Differences in predator composition alter the direction of structure‐mediated predation risk in macrophyte communities

Structural complexity strongly influences the outcome of predator–prey interactions in benthic marine communities affecting both prey concealment and predator hunting efficacy. How habitat structure interacts with species‐specific differences in predatory style and antipredatory strategies may therefore be critical in determining higher trophic functions. We examined the role of structural complexity in mediating predator–prey interactions across several macrophyte habitats along a gradient of structural complexity in three different bioregions: western Mediterranean Sea (WMS), eastern Indian Ocean (EIO) and northern Gulf of Mexico (NGM). Using sea urchins as model prey, we measured survival rates of small (juveniles) and medium (young adults) size classes in different habitat zones: within the macrophyte habitat, along the edge and in bare sandy spaces. At each site we also measured structural variables and predator abundance. Generalised linear models identified biomass and predatory fish abundance as the main determinants of predation intensity but the efficiency of predation was also influenced by urchin size class. Interestingly though, the direction of structure‐mediated effects on predation risk was markedly different between habitats and bioregions. In WMS and NGM, where predation by roving fish was relatively high, structure served as a critical prey refuge, particularly for juvenile urchins. In contrast, in EIO, where roving fish predation was low, predation was generally higher inside structurally complex environments where sea stars were responsible for much of the predation. Larger prey were generally less affected by predation in all habitats, probably due to the absence of large predators. Overall, our results indicate that, while the structural complexity of habitats is critical in mediating predator–prey interactions, the direction of this mediation is strongly influenced by differences in predator composition. Whether the regional pool of predators is dominated by visual roving species or chemotactic benthic predators may determine if structure dampens or enhances the influence of top–down control in marine macrophyte communities.     

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.     

Impacts of the 2011 Tsunami on Sediment Characteristics and Macrozoobenthic Assemblages in a Shallow Eutrophic Lagoon,Sendai Bay,Japan

A huge tsunami is one of the greatest disturbance events in coastal benthic communities, although the ecological consequences are not fully understood. Here we examined the tsunami-induced changes in the sediment environment and macrozoobenthic assemblage in a eutrophic brackish lagoon in eastern Japan. The 7.2-m-high tsunami completely replaced muddy sediment with drifting sea sand throughout the lagoon, leading to the drastic changes in quantity and quality of sedimental organic matters, sulfide contents, and sediment redox condition. Intensive physical stress devastated the benthic community, but the disappearance of sulfidic muddy bottoms significantly improved the habitat quality for macrozoobenthos. The re-established macrozoobenthic community after 5 months was characterized by (1) a 2-fold higher total density, but sharp declines in species richness, diversity, and evenness; (2) an increased density of opportunistic taxa (e.g., polychaete Pseudopolydora spp. and amphipod Monocorophium uenoi) in newly created sandy bottoms; and (3) disappearance of several dominant taxa including bivalves and chironomid larvae. These findings indicate that the sensitivity and recovery potential of macrozoobenthos were highly taxa-specific, which was closely related to the taxa’s ecological characteristics, including tolerance to physical disturbance, life-history traits, and life form. Our data revealed the rapid recolonization of opportunistic macrozoobenthos after a huge tsunami, which would contribute to the functional recovery of estuarine soft-bottom habitats shortly after a disturbance event.     

A trophic mass balance model of the eastern Chukchi Sea with comparisons to other high-latitude systems

The Chukchi Sea is a seasonally ice-covered, marginal Arctic-shelf sea that possesses both large petroleum reserves and abundant biological communities, including migratory mammals and seabirds. We developed a mass balance food web model for the eastern Chukchi Sea to evaluate the trophic structure of this ecosystem and to compare food web properties of the Chukchi Sea to those of other high-latitude marine ecosystems. We compiled data on biomass levels, diet composition, demographic rates (production, consumption), and fishery removals, and used these data to construct an Ecopath trophic mass balance model. The majority of biomass was concentrated in benthic invertebrates and most of the mass flow above trophic level 2.0 was through these groups. We found that density estimates of most fish groups derived from trawl survey data using area-swept methods were insufficient to match the consumptive demands of predators, and that densities needed to be several-fold greater to meet modeled demand. We also used a set of system metrics derived from a common modeling framework to highlight differences in ecosystem structure between the eastern Chukchi Sea and other high-latitude systems. The extent of benthic dominance observed in the eastern Chukchi Sea was unique among the systems examined, both in terms of food web structure and associated mass flows between benthic and pelagic components. In relation to total biomass density, the eastern Chukchi Sea had low production when compared with the other systems, and this lower turnover rate suggests that recovery from disturbance might be slow.     

Patterns and processes in abundance-body size relationships for marine benthic invertebrates

1. The nature of abundance-body size relationships in animal communities, and especially the drivers behind the observed patterns, have been a focus of persistent debate in animal ecology. In a recent review, Allen et al. (2006) categorized five mechanistic explanations behind the commonly observed polymodality in these relationships: energetic constraints; phylogenetic constraints; biogeographical determinants; habitat structure; and community interactions. Progress in understanding of these patterns and the processes underlying them have been hindered by the use of a range of methods that differ in their validity and robustness. 2. Here, we used data on invertebrate body sizes from a variety of sandy beaches in the UK to test the hypothesis that these communities display modality in their abundance-body size relationships. We quantified modality in the relationships using kernel density estimation and smoothed bootstrap resampling and then evaluated the competing explanations for this modality based on the patterns identified in conjunction with measurements of the physical beach environment. 3. We found bimodal distributions in the body size spectrum for benthic invertebrates at nine of 16 sites. There was a consistent trough in the spectrum at around 0·5-1 mm diameter, which reflected the traditional split between meiofauna and macrofauna. Beaches with finer particle sizes and more heterogeneous macrofauna hosted communities with more than two modes. 4. Our results suggest that modality in sandy beach benthic communities is unlikely to be explained by any single hypothesis. There will be an interplay between physical and biological factors, with different explanations accounting for modality at different scales.     

Drifting algal mats as an alternative habitat for benthic invertebrates: Species specific responses to a transient resource

Patchy occurrences of benthic drift algae (i.e. loose lying macroalgal mats) may increase habitat complexity on normally bare soft bottoms, but at the same time, extensive amounts of drifting algal mats are known to stress the benthic fauna. This paper presents results of the first detailed study of the fauna associated with drift algal mats in the northern Baltic Sea. In order to assess the importance of drifting algae as an alternative habitat for benthic fauna, benthic drift algal mats were sampled on shallow (2-9 m) sandy soft bottoms in the outer archipelago of the ?land Islands (Finland). Species composition, abundance and biomass of the macrofauna associated with algal mats were recorded. The results show that drifting algae at times can harbour very high abundances of invertebrates (up to 1116 individuals/g algal dryweight), surpassing invertebrate densities recorded in seagrass communities. The algal fauna varied between sites and over time, and factors such as ambient benthic fauna, exposure to wind-wave disturbance, depth, and algal coverage and condition influenced the invertebrate community composition of the algal mats. Abundance increased while individual biomass of the animals decreased over time (summer season; July-October). A series of laboratory experiments were conducted in order to test the ability of a few important benthic species to move up into, and survive in a drifting algal mat. Macoma balthica, Hydrobia spp., Nereis diversicolor and Bathyporeia pilosa were used in the experiments, and significant differences in their survival and mobility within drifting algae were recorded. This study shows that benthic species differ significantly in their ability to utilise the algal mats, with mainly opportunistic and mobile taxa such as Hydrobia spp., Chironomidae and Ostracoda benefiting from the algae, whereas infaunal species such as M. balthica and B. pilosa are negatively affected. The occurrence of eutrophication induced drifting macroalgal mats has increased significantly during the last decade in the northern Baltic Sea. Hence, the importance of drifting algae as a stress factor and as an alternative habitat for benthic fauna increases.     

Comparing benthic biogeochemistry at a sandy and a muddy site in the Celtic Sea using a model and observations

Results from a 1D setup of the European Regional Seas Ecosystem Model (ERSEM) biogeochemical model were compared with new observations collected under the UK Shelf Seas Biogeochemistry (SSB) programme to assess model performance and clarify elements of shelf-sea benthic biogeochemistry and carbon cycling. Observations from two contrasting sites (muddy and sandy) in the Celtic Sea in otherwise comparable hydrographic conditions were considered, with the focus on the benthic system. A standard model parameterisation with site-specific light and nutrient adjustments was used, along with modifications to the within-seabed diffusivity to accommodate the modelling of permeable (sandy) sediments. Differences between modelled and observed quantities of organic carbon in the bed were interpreted to suggest that a large part (>90%) of the observed benthic organic carbon is biologically relatively inactive. Evidence on the rate at which this inactive fraction is produced will constitute important information to quantify offshore carbon sequestration. Total oxygen uptake and oxic layer depths were within the range of the measured values. Modelled depth average pore water concentrations of ammonium, phosphate and silicate were typically 5–20% of observed values at the muddy site due to an underestimate of concentrations associated with the deeper sediment layers. Model agreement for these nutrients was better at the sandy site, which had lower pore water concentrations, especially deeper in the sediment. Comparison of pore water nitrate with observations had added uncertainty, as the results from process studies at the sites indicated the dominance of the anammox pathway for nitrogen removal; a pathway that is not included in the model. Macrofaunal biomasses were overestimated, although a model run with increased macrofaunal background mortality rates decreased macrofaunal biomass and improved agreement with observations. The decrease in macrofaunal biomass was compensated by an increase in meiofaunal biomass such that total oxygen demand remained within the observed range. The permeable sediment modification reproduced some of the observed behaviour of oxygen penetration depth at the sandy site. It is suggested that future development in ERSEM benthic modelling should focus on: (1) mixing and degradation rates of benthic organic matter, (2) validation of benthic faunal biomass against large scale spatial datasets, (3) incorporation of anammox in the benthic nitrogen cycle, and (4) further developments to represent permeable sediment processes.     

Consequences of Increasing Hypoxic Disturbance on Benthic Communities and Ecosystem Functioning

Disturbance-mediated species loss has prompted research considering how ecosystem functions are changed when biota is impaired. However, there is still limited empirical evidence from natural environments evaluating the direct and indirect (i.e. via biota) effects of disturbance on ecosystem functioning. Oxygen deficiency is a widespread threat to coastal and estuarine communities. While the negative impacts of hypoxia on benthic communities are well known, few studies have assessed in situ how benthic communities subjected to different degrees of hypoxic stress alter their contribution to ecosystem functioning. We studied changes in sediment ecosystem function (i.e. oxygen and nutrient fluxes across the sediment water-interface) by artificially inducing hypoxia of different durations (0, 3, 7 and 48 days) in a subtidal sandy habitat. Benthic chamber incubations were used for measuring responses in sediment oxygen and nutrient fluxes. Changes in benthic species richness, structure and traits were quantified, while stress-induced behavioral changes were documented by observing bivalve reburial rates. The initial change in faunal behavior was followed by non-linear degradation in benthic parameters (abundance, biomass, bioturbation potential), gradually impairing the structural and functional composition of the benthic community. In terms of ecosystem function, the increasing duration of hypoxia altered sediment oxygen consumption and enhanced sediment effluxes of NH₄ ⁺ and dissolved Si. Although effluxes of PO₄ ³⁻ were not altered significantly, changes were observed in sediment PO₄ ³⁻ sorption capability. The duration of hypoxia (i.e. number of days of stress) explained a minor part of the changes in ecosystem function. Instead, the benthic community and disturbance-driven changes within the benthos explained a larger proportion of the variability in sediment oxygen- and nutrient fluxes. Our results emphasize that the level of stress to the benthic habitat matters, and that the link between biodiversity and ecosystem function is likely to be affected by a range of factors in complex, natural environments.     

Modelling marine ecosystem response to climate change and trawling in the North Sea

The marine ecosystem response to climate change and demersal trawling was investigated using the coupled hydrodynamic-biogeochemical water column model GOTM-ERSEM-BFM for three contrasting sites in the North Sea. Climate change forcing was derived from the HadRM3-PPE-UK regional climate model for the UK for the period 1950–2100 using historical emissions and a medium emissions scenario (SRESA1B). Effects of demersal trawling were implemented as an additional mortality on benthic fauna, and changes in the benthic–pelagic nutrient and carbon fluxes. The main impacts of climate change were (i) a temperature-driven increase in pelagic metabolic rates and nutrient cycling, (ii) an increase in primary production fuelled by recycled nutrients, (iii) a decrease in benthic biomass due to increased benthic metabolic rates and decreased food supply as a result of the increased pelagic cycling, and (iv) a decrease in near-bed oxygen concentrations. The main impacts of trawling were (i) reduced benthic biomass due to the increased mortality, and (ii) the increased benthic–pelagic nutrient fluxes, with these effects counteracting each other, and relatively small changes in other variables. One important consequence was a large decrease in the de-nitrification flux predicted at the two summer-stratified sites because less benthic nitrate was available. The effects of trawling scaled linearly with fishing effort, with greatest sensitivity to fishing in summer compared to fishing in winter. The impacts of climate change and trawling were additive, suggesting little or no non-linear interactions between these disturbances.     

Microbial biomass and activity in the vicinity of a mussel bed built up by the blue mussel Mytilus edulis

Our study focuses on the impact of the biosedimentary system mussel bed (Mytilus edulis) on the spatial variability of benthic microbial biomass and activity in relation to organic deposits. We sampled a transect of six stations from the muddy mussel bed towards a reference station in the sandflat in monthly intervals for 1 year. The sediment grain size fraction <63 μm and the total organic carbon (TOC) concentration decreased significantly towards the sandflat. Bacterial numbers and total microbial biomass (total adenylates) showed a high spatial variability and were not correlated to increasing distance from the mussel bed. However, a significant relationship with the TOC concentration was found. In contrast, the energetic status (adenylate energy charge) of the microbial community in the mussel bed was significantly lower than in the sandflat. A principal component analysis of the substrate-utilization pattern revealed clear differences between the microbial communities in the mussel bed and in the sandflat. Our results indicate that the sandflat may be dominated by a relatively specialized benthic microbial community with an increased efficiency in utilizing organic carbon sources. As a disadvantage, however, such r-strategists are only able to meet environmental changes within a comparatively narrow range. Benthic microbial communities in the vicinity of an M. edulis mussel bed, in contrast, are dominated by relative generalists with a greater physiological capacity to buffer discrete environmental changes. Such K-strategists show a lack of specialization which generally means a reduced efficiency in utilizing a particular resource. Received: 2 May 1999 / Received in revised form: 22 November 1999 / Accepted: 6 December 1999     

Small-spatial scale changes in productivity of suprabenthic and infaunal crustaceans at the continental shelf of Ebro Delta (western Mediterranean)

Production (P) and production/biomass ratio (P/B) were estimated from 21 species of peracarid and eucarid crustaceans belonging to two different compartments (infauna: 10 species and suprabenthos: 11 species) using the Hynes size–frequency method. Calculations were based on 7 cruises performed between December 2002 and November 2003 in the continental shelf of the Ebro river delta (ED, Catalan Sea: northwestern Mediterranean). Within ED, P and P/B were compared between two neighbouring sites, one located on 47 m (S1), the other at 61 m (S2). The possible effects of natural variability on production were considered. Sediments in ED were muddy (%clay between 40 and 65%). No significant trend in production comparing S1 and S2 was detected, with 50% of species showing higher P value in S1, and 50% in S2. A significant trend emerged from P/B estimates, with 18 of 21 species showing higher P/B at S1. Similarly significant trends for infauna and suprabenthos were found after analyzing both compartments separately. It is suggested that the more significant P/B found on S1 both among infauna and suprabenthos could be caused by the higher temperature and organic matter content (%TOC) at S1. In multi-linear regression (MLR) models, differences in P between S1 and S2 were greater with increasing differences in the density of species (RD) between sites. Differences were greater in infauna than in suprabenthos, and greater among rare species (those with low dominance, %D). P/B differences between S1 and S2 were mainly explained by %D and RD, and to a lesser extent by differences between sites in the mean size of species (RW). In other words changes in P and P/B were not explained by removal of large size specimens. Some trawling activity occurred at S2, and it may alter the habitat of small macrofauna. In addition to natural variability, we discussed the possible influence of trawling disturbance on P and P/B, bearing in mind the feeding type, mobility, and dispersal capability of species. We suggest that amphipods of the genus Ampelisca, showing higher P/B on S1, characterized by low mobility (probably tube-dwellers), low trophic level (e.g. filter feeding strategies) and low dispersal capability (not free larvae), could be especially vulnerable to trawling impact.     

Effects of commercial trawling activities in the diet of the flat fish Citharus linguatula (Osteichthyes: Pleuronectiformes) and the starfish Astropecten irregularis (Echinodermata: Asteroidea)

This study assesses the effects of commercial trawling on benthic fauna, by analysing differences in the population structure and diet of benthic organisms inhabiting fishing grounds. The study is focused on two epifaunal species collected from a fishing ground in the north-western Mediterranean, which included a portion of un-trawled seabed used as a reference of undisturbed community. The selected species were the flatfish Citharus linguatula, more abundant at the undisturbed site and previously characterised as a surface predator and vulnerable to trawling, and the starfish Astropecten irregularis, which dominated the epibenthic community and had been characterised as an opportunistic species with low vulnerability to trawling. Both species were selective feeders and ingested similar prey species at each of the two sites, although differences in the relative prey abundance found in the stomachs occurred between sites. A. irregularis at the fished site mainly ingested gastropods, whereas higher proportion of bivalves was ingested at the undisturbed site. C. linguatula diet mainly composed of crustaceans at both sites, but there was an increase in the ingestion of small fish at the fished site. Moreover, prey ingestion of these two species generally increased with fishing activity. Trawling appears to modify the diet of these two species in terms of relative abundance of ingested prey, nevertheless, increased opportunism due to trawling disturbance was not detected and the density of these predators over fishing grounds was more closely related to their vulnerability to trawling.