Biological structures exert a major influence on species diversity at both local and regional scales on deep continental margins. Some organisms use other species as substrates for attachment, shelter, feeding or parasitism, but there may also be Mutual benefits from the association. Here, we highlight the structural attributes and biotic effects of the habitats that corals, sea pens, sponges and xenophyophores offer other organisms. The environmental setting of the biological structures influences their species composition. The importance of benthic species as substrates seems to increase with depth as the complexity of the surrounding geological substrate and food supply decline. There are marked differences in the degree of mutualistic relationships between habitat-forming taxa. This is especially evident for scleractinian corals, which have high numbers of facultative associates (commensals) and few obligate associates (mutualists), and gorgonians, with their few commensals and many obligate associates. Size, flexibility and architectural complexity of the habitat-forming organism are positively related to species diversity for both sessile and mobile species. This is mainly evident for commensal species sharing a facultative relationship with their host. Habitat complexity is enhanced by the architecture of biological structures, as well as by biological interactions. Colony morphology has a great influence on feeding efficiency for suspension feeders. Suspension feeding, habitat-forming organisms modify the environment to optimize their food uptake. This environmental advantage is also passed on to associated filter-feeding species. These effects are poorly understood but represent key points for understanding ecosystems and biodiversity on continental margins. In this paper we explore the contributions of organisms and the biotic structures they create (rather than physical modifications) to habitat heterogeneity and diversity on the deep continental margins

SignificanceWe conducted a systematic, high-resolution analysis of bottom trawl fishing footprints for 24 regions on continental shelves and slopes of five continents and New Zealand. The proportion of seabed trawled varied >200-fold among regions (from 0.4 to 80.7% of area to a depth of 1,000 m). Within 18 regions, more than two-thirds of seabed area remained untrawled during study periods of 2–6 years. Relationships between metrics of total trawling activity and footprint were strong and positive, providing a method to estimate trawling footprints for regions where high-resolution data are not available. Trawling footprints were generally smaller in regions where fisheries met targets for exploitation rates, implying collateral environmental benefits of effective fisheries management.

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The European Water Framework Directive (WFD; 2000/60/EG) and the European Marine Strategy Framework Directive (MSFD; 2008/56/EC) were umbrella legislations for fresh and marine waters. It is a challenge for the scientific community to translate the principles of these directives into realistic and accurate approaches. Both directives have the same concept, comparing the current state of an area with that which would be expected under minimal or sustainable human use of that area and in case of degradation, intervening to bring it back to the desired good status. However, each directive used specific principles to fill it in. For the WFD, this was executed during the last decade, and many results of it were already published. This process delivered valuable knowledge on which the implementation of the MSFD can be founded.Therefore, the ICES Benthos Ecology Working Group aimed to stress and discuss some issues, with focus on benthic macro-invertebrates, related to the fulfillment of the principles of both directives. This through the description of (1) how the principles are theoretically filled in by both directives

Marine benthic ecosystems are difficult to monitor and assess, which is in contrast to modern ecosystem-based management requiring detailed information at all important ecological and anthropogenic impact levels. Ecosystem management needs to ensure a sustainable exploitation of marine resources as well as the protection of sensitive habitats, taking account of potential multiple-use conflicts and impacts over large spatial scales. The urgent need for large-scale spatial data on benthic species and communities resulted in an increasing application of distribution modelling (DM). The use of DM techniques enables to employ full spatial coverage data of environmental variables to predict benthic spatial distribution patterns. Especially, statistical DMs have opened new possibilities for ecosystem management applications, since they are straightforward and the outputs are easy to interpret and communicate. Mechanistic modelling techniques, targeting the fundamental niche of species, and Bayesian belief networks are the most promising to further improve DM performance in the marine realm. There are many actual and potential management applications of DMs in the marine benthic environment, these are (i) early warning systems for species invasion and pest control, (ii) to assess distribution probabilities of species to be protected, (iii) uses in monitoring design and spatial management frameworks (e.g. MPA designations), and (iv) establishing long-term ecosystem management measures (accounting for future climate-driven changes in the ecosystem). It is important to acknowledge also the limitations associated with DM applications in a marine management context as well as considering new areas for future DM developments. The knowledge of explanatory variables, for example, setting the basis for DM, will continue to be further developed: this includes both the abiotic (natural and anthropogenic) and the more pressing biotic (e.g. species interactions) aspects of the ecosystem. While the response variables on the other hand are often focused on species presence and some work undertaken on species abundances, it is equally important to consider, e.g. biological traits or benthic ecosystem functions in DM applications. Tools such as DMs are suitable to forecast the possible effects of climate change on benthic species distribution patterns and hence could help to steer present-day ecosystem management.

The MAREANO (Marine AREA database for NOrwegian coast and sea areas) mapping programme includes acquisition of multibeam bathymetry and backscatter data together with a comprehensive, integrated biological and geological sampling programme. Equipment used includes underwater video, box corer, grab, epibenthic sled and beam trawl. Habitat maps are produced by combining information on landscapes, landscape elements, sediment types and biological communities. Video observations provide information about the megafauna diversity of large ([1 cm) epifauna and bottom types, whilst bottom samples describe the composition of epifauna, hyperfauna (crustaceans living in the upper part of the sediment and/or swimming just above the substratum) and infauna, and sediment composition. In this study, two biological data sets are used to study fauna response to environmental heterogeneity at two different spatial scales: (1) broad scale, megahabitat (1-10s km), based on information about megafauna taxa observed during video surveys in the Nordland/ Troms area, (2) fine scale, mesohabitat (10s m-1 km), based on information about species composition documented with video records and bottom sampling gear from the bank ''Tromsøflaket''. In general, the highest diversity is found on bottoms with mixed substrates indicating that substratum heterogeneity is very important for the biodiversity at both scales. The number of taxa shows a maximum at depths between 200 and 700 m followed by a gradual decrease down to 2,200 m. At the broad scale, multibeam data provides a variety of terrain variables that indicate environmental variation (e.g. exposure to currents, interpreted substrates). This analysis identifies six fauna groups associated to specific landscape elements. Diversity of megafauna shows a strong correlation with number of bottom types occurring along video transects. It is highest at the shelf break and decreased with depth on the slope in parallel with a decrease in habitat heterogeneity and temperature. At a fine scale, six biotopes are identified based on megafauna composition with habitat characteristics ranging from homogenous muddy bottom, biotope 1, to the most heterogeneous bottom with [20% rocks and several bottom types present in biotope 6. The macrofauna 123Hydrobiologia ( ) 685:191-219 DOI 10.1007 sampled is used for description of the whole benthic community, including diversity, biomass and production, related to these six biotopes. The variation in percentage cover of substrate types and in particular the cover of hard substrates demonstrate to be a good proxy for the benthic community composition (megaand macrofauna) and its diversity.

A framework to assess the impact of mobile fishing gear on the seabed and benthic ecosystem is presented. The framework that can be used at regional and local scales provides indicators for both trawling pressure and ecological impact. It builds on high-resolution maps of trawling intensity and considers the physical effects of trawl gears on the seabed, on marine taxa, and on the functioning of the benthic ecosystem. Within the framework, a reductionist approach is applied that breaks down a fishing gear into its components, and a number of biological traits are chosen to determine either the vulnerability of the benthos to the impact of that gear component, or to provide a proxy for their ecological role. The approach considers gear elements, such as otter boards, twin trawl clump, and groundrope, and sweeps that herd the fish. The physical impact of these elements on the seabed, comprising scraping of the seabed, sediment mobilization, and penetration, is a function of the mass, size, and speed of the individual component. The impact of the elements on the benthic community is quantified using a biological-trait approach that considers the vulnerability of the benthic community to trawl impact (e.g. sediment position, morphology), the recovery rate (e.g. longevity, maturation age, reproductive characteristics, dispersal), and their ecological role. The framework is explored to compare the indicators for pressure and ecological impact of bottom trawling in three main seabed habitat types in the North Sea. Preliminary results show that the Sublittoral mud (EUNIS A5.3) is affected the most due to the combined effect of intensive fishing and large proportions of long-lived taxa.