Jan Geert Hiddink

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 (P0.9). The pressure indicator measures the proportion of the area where trawling frequency is sufficiently high to prevent reaching predicted B0.9 or P0.9. Time to recovery to B0.9 and P0.9 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 B0.9, and 27.1% was trawled too frequently for production to reach P0.9. As a result of bottom-trawling in 2003, in 56% of the southern North Sea benthic biomass was below B0.9, whereas in 27% of the southern North Sea benthic production was below P0.9. 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.

Predation on 0-group and older year classes of the bivalve Macoma balthica: interaction of size selection and intertidal distribution of epibenthic predators

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea. Shrimps Crangon crangon, adult crabs Carcinus maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinus stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+group Macoma. Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation.

Environmental metabarcoding reveals heterogeneous drivers of microbial eukaryote diversity in contrasting estuarine ecosystems

Assessing how natural environmental drivers affect biodiversity underpins our understanding of the relationships between complex biotic and ecological factors in natural ecosystems. Of all ecosystems, anthropogenically important estuaries represent a ‘melting pot’ of environmental stressors, typified by extreme salinity variations and associated biological complexity. Although existing models attempt to predict macroorganismal diversity over estuarine salinity gradients, attempts to model microbial biodiversity are limited for eukaryotes. Although diatoms commonly feature as bioindicator species, additional microbial eukaryotes represent a huge resource for assessing ecosystem health. Of these, meiofaunal communities may represent the optimal compromise between functional diversity that can be assessed using morphology and phenotype–environment interactions as compared with smaller life fractions. Here, using 454 Roche sequencing of the 18S nSSU barcode we investigate which of the local natural drivers are most strongly associated with microbial metazoan and sampled protist diversity across the full salinity gradient of the estuarine ecosystem. In order to investigate potential variation at the ecosystem scale, we compare two geographically proximate estuaries (Thames and Mersey, UK) with contrasting histories of anthropogenic stress. The data show that although community turnover is likely to be predictable, taxa are likely to respond to different environmental drivers and, in particular, hydrodynamics, salinity range and granulometry, according to varied life-history characteristics. At the ecosystem level, communities exhibited patterns of estuary-specific similarity within different salinity range habitats, highlighting the environmental sequencing biomonitoring potential of meiofauna, dispersal effects or both.

Temperature tracking by North Sea benthic invertebrates in response to climate change

Climate change is a major threat to biodiversity and distributions shifts are one of the most significant threats to global warming, but the extent to which these shifts keep pace with a changing climate is yet uncertain. Understanding the factors governing range shifts is crucial for conservation management to anticipate patterns of biodiversity distribution under future anthropogenic climate change. Soft-sediment invertebrates are a key faunal group because of their role in marine biogeochemistry and as a food source for commercial fish species. However, little information exists on their response to climate change. Here, we evaluate changes in the distribution of 65 North Sea benthic invertebrate species between 1986 and 2000 by examining their geographic, bathymetric and thermal niche shifts and test whether species are tracking their thermal niche as defined by minimum, mean or maximum sea bottom (SBT) and surface (SST) temperatures. Temperatures increased in the whole North Sea with many benthic invertebrates showing north-westerly range shifts (leading/trailing edges as well as distribution centroids) and deepening. Nevertheless, distribution shifts for most species (3.8-7.3 km yr(-1) interquantile range) lagged behind shifts in both SBT and SST (mean 8.1 km yr(-1)), resulting in many species experiencing increasing temperatures. The velocity of climate change (VoCC) of mean SST accurately predicted both the direction and magnitude of distribution centroid shifts, while maximum SST did the same for contraction of the trailing edge. The VoCC of SBT was not a good predictor of range shifts. No good predictor of expansions of the leading edge was found. Our results show that invertebrates need to shift at different rates and directions to track the climate velocities of different temperature measures, and are therefore lagging behind most temperature measures. If these species cannot withstand a change in thermal habitat, this could ultimately lead to a drop in benthic biodiversity.

Context dependence of marine ecosystem engineer invasion impacts on benthic ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.

Predation of intertidal infauna on juveniles of the bivalve Macoma balthica

Juveniles of the bivalve Macoma balthica live on tidal flats in the Wadden Sea. This study examined the interaction of Macoma with the infaunal polychaetes Arenicola marina and Nereis diversicolor and the gastropod Retusa obtusa. The distribution of M. balthica spat on the flats, shortly after settlement in April, showed a positive correlation with the Arenicola distribution and a negative correlation with Nereis distribution. There were no locations where Macoma spat and Retusa occurred together. In August, Macoma spat had grown too large for predation by intertidal infauna. Small individuals of Macoma spat were found in stomachs of Arenicola (0.14 worm 1 ) and Nereis (0.05 worm 1 ). Laboratory experiments showed that Nereis and Retusa could reduce Macoma spat abundance, both in the absence and presence of sediment and alternative prey. Arenicola reduced the abundance of small Macoma (<1 mm) in sediment without, but not with, alternative prey. In field experiments, we manipulated the density of Arenicola in 0.25–1 m 2 plots and of Nereis in 0.03 m 2 cages and examined the effect on Macoma density several weeks later. We found a significant negative relation between densities of polychaetes and Macoma spat for both polychaete species in these experimental plots. Peculiarly, we found a significant positive relation between manipulated Nereis density and adult Macoma density in the cages; we cannot explain this. Consumption rates, calculated both from stomach contents and from field experiments, were 45 to 102 Macoma m 2 d 1 for Arenicola and 5 to 116 Macoma m 2 d 1 for Nereis. These values are higher than recorded consumption rates by epibenthic predators in the same area. Nevertheless, between-year differences in year-class strength could not be explained by differential abundance of these polychaetes. In conclusion, Arenicola and Nereis had a negative effect on the abundance of Macoma <1.5 mm, which was at least partly caused by direct consumption. Retusa obtusa can eat juvenile Macoma, but probably did not so in the study area, because there were no locations where Retusa and Macoma spat occurred together in the period that Macoma was <2 mm. D 2002 Elsevier Science B.V. All rights reserved.

Effects of bottom trawling on fish foraging and feeding

The effects of bottom trawling on benthic invertebrates include reductions of biomass, diversity and body size. These changes may negatively affect prey availability for demersal fishes, potentially leading to reduced food intake, body condition and yield of fishes in chronically trawled areas. Here, the effect of trawling on the prey availability and diet of two commercially important flatfish species, plaice (Pleuronectes platessa) and dab (Limanda limanda), was investigated over a trawling intensity gradient in the Irish Sea. Previous work in this area has shown that trawling negatively affects the condition of plaice but not of dab. This study showed that reductions in local prey availability did not result in reduced feeding of fish. As trawling frequency increased, both fish and prey biomass declined, such that the ratio of fish to prey remained unchanged. Consequently, even at frequently trawled sites with low prey biomass, both plaice and dab maintained constant levels of stomach fullness and gut energy contents. However, dietary shifts in plaice towards energy-poor prey items were evident when prey species were analysed individually. This, together with a potential decrease in foraging efficiency due to low prey densities, was seen as the most plausible cause for the reduced body condition observed. Understanding the relationship between trawling, benthic impacts, fish foraging and resultant body condition is an important step in designing successful mitigation measures for future management strategies in bottom trawl fisheries.

Global analysis of depletion and recovery of seabed biota after bottom trawling disturbance

Significance Bottom trawling is the most widespread source of physical disturbance to the world’s seabed. Predictions of trawling impacts are needed to underpin risk assessment, and they are relevant for the fishing industry, conservation, management, and certification bodies. We estimate depletion and recovery of seabed biota after trawling by fitting models to data from a global data compilation. Trawl gears removed 6–41% of faunal biomass per pass, and recovery times posttrawling were 1.9–6.4 y depending on fisheries and environmental context. These results allow the estimation of trawling impacts on unprecedented spatial scales and for data poor fisheries and enable an objective analysis of tradeoffs between harvesting fish and the wider ecosystem effects of such activities. Bottom trawling is the most widespread human activity affecting seabed habitats. Here, we collate all available data for experimental and comparative studies of trawling impacts on whole communities of seabed macroinvertebrates on sedimentary habitats and develop widely applicable methods to estimate depletion and recovery rates of biota after trawling. Depletion of biota and trawl penetration into the seabed are highly correlated. Otter trawls caused the least depletion, removing 6% of biota per pass and penetrating the seabed on average down to 2.4 cm, whereas hydraulic dredges caused the most depletion, removing 41% of biota and penetrating the seabed on average 16.1 cm. Median recovery times posttrawling (from 50 to 95% of unimpacted biomass) ranged between 1.9 and 6.4 y. By accounting for the effects of penetration depth, environmental variation, and uncertainty, the models explained much of the variability of depletion and recovery estimates from single studies. Coupled with large-scale, high-resolution maps of trawling frequency and habitat, our estimates of depletion and recovery rates enable the assessment of trawling impacts on unprecedented spatial scales.

Dominance, Biomass and Extinction Resistance Determine the Consequences of Biodiversity Loss for Multiple Coastal Ecosystem Processes

Key ecosystem processes such as carbon and nutrient cycling could be deteriorating as a result of biodiversity loss. However, currently we lack the ability to predict the consequences of realistic species loss on ecosystem processes. The aim of this study was to test whether species contributions to community biomass can be used as surrogate measures of their contribution to ecosystem processes. These were gross community productivity in a salt marsh plant assemblage and an intertidal macroalgae assemblage; community clearance of microalgae in sessile suspension feeding invertebrate assemblage; and nutrient uptake in an intertidal macroalgae assemblage. We conducted a series of biodiversity manipulations that represented realistic species extinction sequences in each of the three contrasting assemblages. Species were removed in a subtractive fashion so that biomass was allowed to vary with each species removal, and key ecosystem processes were measured at each stage of community disassembly. The functional contribution of species was directly proportional to their contribution to community biomass in a 1∶1 ratio, a relationship that was consistent across three contrasting marine ecosystems and three ecosystem processes. This suggests that the biomass contributed by a species to an assemblage can be used to approximately predict the proportional decline in an ecosystem process when that species is lost. Such predictions represent “worst case scenarios” because, over time, extinction resilient species can offset the loss of biomass associated with the extinction of competitors. We also modelled a “best case scenario” that accounts for compensatory responses by the extant species with the highest per capita contribution to ecosystem processes. These worst and best case scenarios could be used to predict the minimum and maximum species required to sustain threshold values of ecosystem processes in the future.

An approach for the identification of exemplar sites for scaling up targeted field observations of benthic biogeochemistry in heterogeneous environments

Continental shelf sediments are globally important for biogeochemical activity. Quantification of shelf-scale stocks and fluxes of carbon and nutrients requires the extrapolation of observations made at limited points in space and time. The procedure for selecting exemplar sites to form the basis of this up-scaling is discussed in relation to a UK-funded research programme investigating biogeochemistry in shelf seas. A three-step selection process is proposed in which (1) a target area representative of UK shelf sediment heterogeneity is selected, (2) the target area is assessed for spatial heterogeneity in sediment and habitat type, bed and water column structure and hydrodynamic forcing, and (3) study sites are selected within this target area encompassing the range of spatial heterogeneity required to address key scientific questions regarding shelf scale biogeochemistry, and minimise confounding variables. This led to the selection of four sites within the Celtic Sea that are significantly different in terms of their sediment, bed structure, and macrofaunal, meiofaunal and microbial community structures and diversity, but have minimal variations in water depth, tidal and wave magnitudes and directions, temperature and salinity. They form the basis of a research cruise programme of observation, sampling and experimentation encompassing the spring bloom cycle. Typical variation in key biogeochemical, sediment, biological and hydrodynamic parameters over a pre to post bloom period are presented, with a discussion of anthropogenic influences in the region. This methodology ensures the best likelihood of site-specific work being useful for up-scaling activities, increasing our understanding of benthic biogeochemistry at the UK-shelf scale.