Substantial changes in the depth distributions of benthic invertebrates in the eastern Kattegat since the 1880s.

Bottom trawling and eutrophication are well known for their impacts on the marine benthic environment in the last decades. Evaluating the effects of these pressures is often restricted to contemporary benthic data, limiting the potential to observe change from an earlier (preimpact) state. In this study, we compared benthic species records from 1884 to 1886 by CGJ Petersen with recent data to investigate how benthic invertebrate species in the eastern Kattegat have changed since preimpact time. The study shows that species turnover between old and recent times was high, ca. 50%, and the species richness in the investigation area was either unchanged or higher in recent times, suggesting no net loss of species. Elements of metacommunity structure analysis of datasets from the 1880s, 1990s, and 2000s revealed a clear change in the depth distribution structure since the 1880s. The system changed from a Quasi-nested/Random pattern unrelated to depth in the 1880s with many species depth ranges over a major part of the studied depth interval, to a Clementsian pattern in recent times strongly positively correlated with depth. Around 30% of the 117 species recorded both in old and in recent times, including most trawling-sensitive species, that is large, semiemergent species, showed a decrease in maximal depth of occurrence from the deeper zone fished today to the shallower unfished zone, with on average 20 m. Concurrently, the species category remaining in the fished zone was dominated by species less sensitive to bottom trawling like infauna polychaetes and small-sized Peracarida crustaceans, most likely with short longevity. The depth interval and magnitude of the changes in depth distribution and the changes in species composition indicate impacts from bottom trawling rather than eutrophication. Furthermore, the high similarity of results from the recent datasets 10 years apart suggests chronic impact keeping the system in an altered state.

Long-term temporal and spatial trends in eutrophication status of the Baltic Sea.

Much of the Baltic Sea is currently classified as 'affected by eutrophication'. The causes for this are twofold. First, current levels of nutrient inputs (nitrogen and phosphorus) from human activities exceed the natural processing capacity with an accumulation of nutrients in the Baltic Sea over the last 50-100 years. Secondly, the Baltic Sea is naturally susceptible to nutrient enrichment due to a combination of long retention times and stratification restricting ventilation of deep waters. Here, based on a unique data set collated from research activities and long-term monitoring programs, we report on the temporal and spatial trends of eutrophication status for the open Baltic Sea over a 112-year period using the HELCOM Eutrophication Assessment Tool (HEAT 3.0). Further, we analyse variation in the confidence of the eutrophication status assessment based on a systematic quantitative approach using coefficients of variation in the observations. The classifications in our assessment indicate that the first signs of eutrophication emerged in the mid-1950s and the central parts of the Baltic Sea changed from being unaffected by eutrophication to being affected. We document improvements in eutrophication status that are direct consequences of long-term efforts to reduce the inputs of nutrients. The reductions in both nitrogen and phosphorus loads have led to large-scale alleviation of eutrophication and to a healthier Baltic Sea. Reduced confidence in our assessment is seen more recently due to reductions in the scope of monitoring programs. Our study sets a baseline for implementation of the ecosystem-based management strategies and policies currently in place including the EU Marine Strategy Framework Directives and the HELCOM Baltic Sea Action Plan.

Species Sorting of Benthic Invertebrates in a Salinity Gradient - Importance of Dispersal Limitation.

The relative importance of environment and dispersal related processes for community assembly has attracted great interest over recent decades, but few empirical studies from the marine/estuarine realm have examined the possible effects of these two types of factors in the same system. Importance of these processes was investigated in a hypothetical metacommunity of benthic invertebrates in 16 micro-tidal estuaries connected to the same open sea area. The estuaries differed in size and connectivity to the open sea and represented a salinity gradient across the estuaries. The Elements of Metacommunity Structure (EMS) approach on estuary scale was complemented with a mechanistic variance partitioning approach on sample scale to disentangle effects of factors affecting assembly of three trait groups of species with different dispersivity. A quasi-Clementsian pattern was observed for all three traits, a likely response to some latent gradient. The primary axis in the pattern was most strongly related to gradients in estuary salinity and estuary entrance width and correlation with richness indicated nestedness only in the matrix of the most dispersive trait group. In the variance partitioning approach measures of turnover and nestedness between paired samples each from different estuaries were related to environmental distance in different gradients. Distance between estuaries was unimportant suggesting importance of factors characterizing the estuaries. While the high dispersive species mainly were sorted in the salinity gradient, apparently according to their tolerance ranges towards salinity, the two less dispersive traits were additionally affected by estuary entrance width and possibly also area. The results exemplify a mechanism of community assembly in the marine realm where the niche factor salinity in conjunction with differential dispersal structure invertebrates in a metacommunity of connected estuaries, and support the idea that dispersive species are more controlled by the environment than less dispersive species.

The role of recurrent disturbances for ecosystem multifunctionality.

Ecosystem functioning is threatened by an increasing number of anthropogenic stressors, creating a legacy of disturbance that undermines ecosystem resilience. However, few empirical studies have assessed to what extent an ecosystem can tolerate repeated disturbances and sustain its multiple functions. By inducing increasingly recurring hypoxic disturbances to a sedimentary ecosystem, we show that the majority of individual ecosystem functions experience gradual degradation patterns in response to repetitive pulse disturbances. The degradation in overall ecosystem functioning was, however, evident at an earlier stage than for single ecosystem functions and was induced after a short pulse of hypoxia (i.e., three days), which likely reduced ecosystem resistance to further hypoxic perturbations. The increasing number of repeated pulse disturbances gradually moved the system closer to a press response. In addition to the disturbance regime, the changes in benthic trait composition as well as habitat heterogeneity were important for explaining the variability in overall ecosystem functioning. Our results suggest that disturbance-induced responses across multiple ecosystem functions can serve as a warning signal for losses of the adaptive capacity of an ecosystem, and might at an early stage provide information to managers and policy makers when remediation efforts should be initiated.

Assessment of marine benthic quality change in gradients of disturbance: comparison of different Scandinavian multi-metric indices.

Three multi-metric benthic macrofauna indices were used to assess marine benthic ecological quality status (EcoQS) according to the European Water Framework Directive, in seven pollution gradients mainly, western Scandinavia. The impacts included organic load, hypoxia, metals, urban effluents and physical disturbance. The indices responded in a similar threshold fashion, irrespective of impact factor identity. Usually, the border between Good and Moderate EcoQS (G/M), is determined as some deviation from a reference situation. References, however, are difficult to find. An alternative procedure is described to estimate the G/M border, not requiring reference data. Thresholds, where faunal structure deterioration commences, were identified from non-linear regressions between indices and impact factors. Index values from the less impacted side of the thresholds were assumed to come from environments of Good and High EcoQS, and the 5th percentile of these data, was defined as the G/M border. Estimated G/M borders compared well with previous studies.

An approach to the intercalibration of benthic ecological status assessment in the North Atlantic ecoregion, according to the European Water Framework Directive.

The European Water Framework Directive (WFD) establishes a framework for the protection and improvement of transitional and coastal waters; its final objective is to achieve at least 'good water status' for all waters, by 2015. The WFD requires Member States (MSs) to assess the Ecological Status (ES) of water bodies. This assessment will be based upon the status of the biological, hydromorphological and physico-chemical quality elements, by comparing data obtained from monitoring networks to reference (undisturbed) conditions, and then deriving an Ecological Quality Ratio (EQR). One of the biological quality elements to be considered is the benthic invertebrate component and some structural parameters (composition, diversity and disturbance-sensitive taxa) must be included in the ES assessment. Following these criteria, several approaches to benthic invertebrate assessment have been proposed by MSs. The WFD requires that these approaches are intercalibrated. This contribution describes the comparison of the different methodologies proposed by United Kingdom, Spain, Denmark and Norway. Results show a high consistency between the approaches, both with regard to determining the EQR and boundary settings for the ES.