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.

Biological traits approaches in benthic marine ecology: Dead ends and new paths.

Biological traits analysis (BTA) links community structure to both ecological functions and response to environmental drivers through species' attributes. In consequence, it has become a popular approach in marine benthic studies. However, BTA will reach a dead end if the scientific community does not acknowledge its current shortcomings and limitations: (a) uncertainties related to data origins and a lack of standardized reporting of trait information; (b) knowledge gaps on the role of multiple interacting traits on driving the organisms' responses to environmental variability; (c) knowledge gaps regarding the mechanistic links between traits and functions; (d) a weak focus on the spatial and temporal variability that is inherent to the trait expression of species; and, last but not least, (e) the large reliance on expert knowledge due to an enormous knowledge gap on the basic ecology of many benthic species. BTA will only reach its full potential if the scientific community is able to standardize and unify the reporting and storage of traits data and reconsider the importance of baseline observational and experimental studies to fill knowledge gaps on the mechanistic links between biological traits, functions, and environmental variability. This challenge could be assisted by embracing new technological advances in marine monitoring, such as underwater camera technology and artificial intelligence, and making use of advanced statistical approaches that consider the interactive nature and spatio-temporal variability of biological systems. The scientific community has to abandon some dead ends and explore new paths that will improve our understanding of individual species, traits, and the functioning of benthic ecosystems.

Causal Approach to Determining the Environmental Risks of Seabed Mining.

Mineral deposits containing commercially exploitable metals are of interest for seabed mineral extraction in both the deep sea and shallow sea areas. However, the development of seafloor mining is underpinned by high uncertainties on the implementation of the activities and their consequences for the environment. To avoid unbridled expansion of maritime activities, the environmental risks of new types of activities should be carefully evaluated prior to permitting them, yet observational data on the impacts is mostly missing. Here, we examine the environmental risks of seabed mining using a causal, probabilistic network approach. Drawing on a series of expert interviews, we outline the cause-effect pathways related to seabed mining activities to inform quantitative risk assessments. The approach consists of (1) iterative model building with experts to identify the causal connections between seabed mining activities and the affected ecosystem components and (2) quantitative probabilistic modeling. We demonstrate the approach in the Baltic Sea, where seabed mining been has tested and the ecosystem is well studied. The model is used to provide estimates of mortality of benthic fauna under alternative mining scenarios, offering a quantitative means to highlight the uncertainties around the impacts of mining. We further outline requirements for operationalizing quantitative risk assessments in data-poor cases, highlighting the importance of a predictive approach to risk identification. The model can be used to support permitting processes by providing a more comprehensive description of the potential environmental impacts of seabed resource use, allowing iterative updating of the model as new information becomes available.

Predation risk and competition affect habitat use of adult perch, Perca fluviatilis.

The aim of this study was to examine how the presence of a predator and an interspecific competitor influence the habitat use of adult perch (Perca fluviatilis; size: 15.1 ± 0.5 cm) when given the choice between two adjacent habitats. By conducting aquarium experiments, the habitat occupancy of P. fluviatilis was documented in the presence and absence of a predator (pike Esox lucius; size: 25.4 ± 2.1 cm) and a potential competitor (ruffe Gymnocephalus cernuus; size: 14.1 ± 0.3 cm) fish species. Two P. fluviatilis individuals generally shared the same habitat. In the presence of a conspecific, P. fluviatilis favoured the structurally more-complex, artificial macrophyte habitat over the less-structured rock and sand habitat, which in turn were used equally. In the predator- and competitor treatments, P. fluviatilis seemed to adapt their habitat use to the habitat occupancy of E. lucius and G. cernuus in the Macrophyte vs. Rock and, in the predator treatment, also in the Macrophyte vs. Sand habitat combination, by increasingly occupying a habitat that was used less by the predator or competitor species, respectively. This behaviour suggests that P. fluviatilis tried to avoid the other fish species by choosing a, in some cases less preferred, predator- or competitor-free habitat. This study emphasizes the importance of biological interactions illustrated by the potential of predation risk and competition to structure fish communities by influencing habitat use at small spatial scales.

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.

The importance of benthic-pelagic coupling for marine ecosystem functioning in a changing world.

Benthic-pelagic coupling is manifested as the exchange of energy, mass, or nutrients between benthic and pelagic habitats. It plays a prominent role in aquatic ecosystems, and it is crucial to functions from nutrient cycling to energy transfer in food webs. Coastal and estuarine ecosystem structure and function are strongly affected by anthropogenic pressures; however, there are large gaps in our understanding of the responses of inorganic nutrient and organic matter fluxes between benthic habitats and the water column. We illustrate the varied nature of physical and biological benthic-pelagic coupling processes and their potential sensitivity to three anthropogenic pressures - climate change, nutrient loading, and fishing - using the Baltic Sea as a case study and summarize current knowledge on the exchange of inorganic nutrients and organic material between habitats. Traditionally measured benthic-pelagic coupling processes (e.g., nutrient exchange and sedimentation of organic material) are to some extent quantifiable, but the magnitude and variability of biological processes are rarely assessed, preventing quantitative comparisons. Changing oxygen conditions will continue to have widespread effects on the processes that govern inorganic and organic matter exchange among habitats while climate change and nutrient load reductions may have large effects on organic matter sedimentation. Many biological processes (predation, bioturbation) are expected to be sensitive to anthropogenic drivers, but the outcomes for ecosystem function are largely unknown. We emphasize how improved empirical and experimental understanding of benthic-pelagic coupling processes and their variability are necessary to inform models that can quantify the feedbacks among processes and ecosystem responses to a changing world.

Coastal habitats as surrogates for taxonomic, functional and trophic structures of benthic faunal communities.

Due to human impact, there is extensive degradation and loss of marine habitats, which calls for measures that incorporate taxonomic as well as functional and trophic aspects of biodiversity. Since such data is less easily quantifiable in nature, the use of habitats as surrogates or proxies for biodiversity is on the rise in marine conservation and management. However, there is a critical gap in knowledge of whether pre-defined habitat units adequately represent the functional and trophic structure of communities. We also lack comparisons of different measures of community structure in terms of both between- (ß) and within-habitat (α) variability when accounting for species densities. Thus, we evaluated a priori defined coastal habitats as surrogates for traditional taxonomic, functional and trophic zoobenthic community structure. We focused on four habitats (bare sand, canopy-forming algae, seagrass above- and belowground), all easily delineated in nature and defined through classification systems. We analyzed uni- and multivariate data on species and trait diversity as well as stable isotope ratios of benthic macrofauna. A good fit between habitat types and taxonomic and functional structure was found, although habitats were more similar functionally. This was attributed to within-habitat heterogeneity so when habitat divisions matched the taxonomic structure, only bare sand was functionally distinct. The pre-defined habitats did not meet the variability of trophic structure, which also proved to differentiate on a smaller spatial scale. The quantification of trophic structure using species density only identified an epi- and an infaunal unit. To summarize the results we present a conceptual model illustrating the match between pre-defined habitat types and the taxonomic, functional and trophic community structure. Our results show the importance of including functional and trophic aspects more comprehensively in marine management and spatial planning.

Developing the multitrait concept for functional diversity: lessons from a system rich in functions but poor in species.

Studies focusing on the linkage between numerical and functional trait diversity frequently consider functional diversity indices but rarely evaluate them empirically or evaluate the use of other than continuous traits such as body size. Here, we present an extensive compilation on functional knowledge of benthic macrofauna using the categorical trait approach and scores of both common and rare species for 25 biological traits, including 102 modalities. We empirically quantify functional trait richness, within-trait species richness (redundancy), and trait variability on a large regional scale (> 1000 km), in three environmentally different areas (basins of a sea), over a long time-span (10 years). To develop further the usage of multiple categorical traits as an analysis tool, we examine the effect of sampling effort for the understanding of the functional properties of the benthic meta-assemblages. We also evaluate the relationship between species richness and trait richness in order to understand co-variation between trait modalities and how traits are packaged within species. Results show that the biological diversity in terms of traits could be distinguished between areas of higher and lower salinity, higher and lower anthropogenic stress, and higher and lower species richness. A considerably lower number of samples are needed to portray the functional structure of an area in relation to the taxonomic structure, thereby demonstrating the advantage of using traits when considering management and conservation issues. Using categorical traits empirically requires, as shown within this study, an understanding of the relationship between species richness and expression of traits, covariation of traits at different species richness and composition levels, acknowledgment of differences in trait expressions between common and rare species, and variability in abundance of species. Empirical trait-based analysis can reveal large-scale differences and insights into complexities between assemblage structure and function, and simultaneously be a valid tool for finding generalities.