A trait-based approach to assess niche overlap and functional distinctiveness between non-indigenous and native species.

Our understanding of the community assembly processes acting on non-indigenous species (NIS), as well as the relationship with native species is limited, especially in marine ecosystems. To overcome this knowledge gap we here develop a trait-based approach based on the functional distinctiveness metric to assess niche overlap between NIS and native species, using high-resolution data on benthic invertebrate communities in the Baltic Sea. Our results show that NIS retain a certain degree of similarity with native species, but display one or a few singular unique traits (e.g., bioturbation ability). Furthermore, we demonstrate that community assembly processes, including both environmental filtering and limiting similarity affect NIS establishment, but that their effects may be highly context dependent, as illustrated by pronounced spatial patterns in distinctiveness. Finally, our trait-based approach provides a generic framework applicable to other areas and organisms, to better understand and address biological invasions.

Four decades of functional community change reveals gradual trends and low interlinkage across trophic groups in a large marine ecosystem.

The rate at which biological diversity is altered on both land and in the sea, makes temporal community development a critical and fundamental part of understanding global change. With advancements in trait-based approaches, the focus on the impact of temporal change has shifted towards its potential effects on the functioning of the ecosystems. Our mechanistic understanding of and ability to predict community change is still impeded by the lack of knowledge in long-term functional dynamics that span several trophic levels. To address this, we assessed species richness and multiple dimensions of functional diversity and dynamics of two interacting key organism groups in the marine food web: fish and zoobenthos. We utilized unique time series-data spanning four decades, from three environmentally distinct coastal areas in the Baltic Sea, and assembled trait information on six traits per organism group covering aspects of feeding, living habit, reproduction and life history. We identified gradual long-term trends, rather than abrupt changes in functional diversity (trait richness, evenness, dispersion) trait turnover, and overall multi-trait community composition. The linkage between fish and zoobenthic functional community change, in terms of correlation in long-term trends, was weak, with timing of changes being area and trophic group specific. Developments of fish and zoobenthos traits, particularly size (increase in small size for both groups) and feeding habits (e.g. increase in generalist feeding for fish and scavenging or predation for zoobenthos), suggest changes in trophic pathways. We summarize our findings by highlighting three key aspects for understanding functional change across trophic groups: (a) decoupling of species from trait richness, (b) decoupling of richness from density and (c) determining of turnover and multi-trait dynamics. We therefore argue for quantifying change in multiple functional measures to help assessments of biodiversity change move beyond taxonomy and single trophic groups.

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.

Cover of coastal vegetation as an indicator of eutrophication along environmental gradients.

Coastal vegetation communities are important for primary production, biodiversity, coastal protection, carbon and nutrient cycling which, in combination with their sensitivity to eutrophication, render them potential indicators of environmental status for environmental policies like the EU Water and Marine Strategy Framework Directives. We evaluated one potential indicator for coastal vegetation, the cumulative cover at depths where the vegetation is light limited, by investigating its response to eutrophication along gradients in natural conditions. We used a large data set covering the Swedish coastline, spanning broad gradients in nutrient level, water clarity, seabed substrate, physical exposure and climate in addition to a salinity gradient from 0.5 to 30.5. Macroalgal cover increased significantly along gradients of declining nutrient concentration and increasing water clarity when we had accounted for diver effects, spatio-temporal sampling variability, salinity gradients, wave exposure and latitude. The developed empirical model explained 79% of the variation in algal cover across 130 areas. Based on this, we identified macroalgal cover as a promising indicator across the Baltic Sea, Kattegat and Skagerrak. A parallel analysis of soft-substrate macrophytes similarly identified significant increases in cover with decreasing concentrations of total nitrogen and increasing salinity, but the resulting empirical model explained only 52% of the variation in cover, probably due to the spatially more variable nature of soft-substrate vegetation. The identified general responses of vegetation cover to gradients of eutrophication across wide ranges in environmental settings may be useful for monitoring and management of marine vegetation in areas with strong environmental gradients.

Reducing spatial variation in environmental assessment of marine benthic fauna.

The Benthic Quality Index, BQI, is widely used for benthic quality assessment. Here, we investigated if spatial variation in the BQI can be reduced by accounting for the environmental factors instead of having different boundaries for different salinity regimes between status classes in the EU Water Framework Directive and Marine Strategy Framework Directive. For this purpose we tested salinity, sediment structure, and depth in a regression model to test their contribution to variations in BQI. The spatial variation in BQI was better explained by depth than by salinity or sediment structure. The proposed assessment method uses the residuals from the regression model between BQI and depth. With this method the variance in BQI between samples was reduced by 50% to 75% in the majority of situations. A method to establish the boundary between good and moderate status and how to derive EQR-values according to the WFD is presented.

Calculation of species sensitivity values and their precision in marine benthic faunal quality indices.

A challenging aspect of benthic quality indices used for assessing the marine environment has been to compile reliable measures of the species' sensitivity to disturbances. Sensitivity values and their uncertainties can be calculated, but a problem to cope with is that the results may depend on the actual proportion of samples from disturbed and undisturbed environments. Here we calculated sensitivity values for each species along an artificial disturbance gradient created by bootstrapping varying numbers of samples from disturbed and undisturbed environments. The values were increasing, decreasing, or more or less constant along this gradient. The lowest value with the lowest uncertainty was adopted as the species sensitivity value. Analyses of the uncertainties indicated that the accuracy rather than the precision might be a concern. We suggest a method to exclude species for which the uncertainty is outside predefined limits as a precaution to reduce bias in the environmental status classification.

Theoretical and practical aspects on benthic quality assessment according to the EU-Water Framework Directive--examples from Swedish waters.

A previously presented objective method to calculate each species sensitivity to disturbance is here slightly modified and implemented in the Benthic Quality Index (BQI) for marine benthic invertebrates. A framework for assessment of water bodies based on multi-site BQI-values is also presented, where a certain variation of BQI-values is allowed to cover the heterogeneity within each water body. The 20th percentile, using bootstrapping, from the available sites' BQI-values is compared with the status boundaries for quality assessment. The reliability of the assessment depends on the background information available for the boundary setting as well as the number of sampling sites included in the assessment. Agreement between time series of quality assessments in areas with known changes in anthropogenic disturbances is encouraging. Problems associated with water body assessment based on few or no samples, as well as multiple sampling occasions during the 6-yr WFD cycle are discussed.

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.

Marine quality assessment by use of benthic species-abundance distributions: a proposed new protocol within the European Union Water Framework Directive.

The aim of this study is to develop a new method for classification of marine benthic quality according to the European Union Water Framework Directive. Tolerance values to environmental disturbance were determined in an objective analysis for benthic species along the Swedish west coast by using 4676 samples from 257 stations. Based on a combination of the species tolerance values, abundance and diversity, a benthic quality index (BQI) was calculated for the assessment of environmental status at a particular station. The qualification of BQI was evaluated in relation to known spatial and temporal gradients of disturbance.