The deep sea, the largest biome on Earth, has a series of characteristics that make this environment both distinct from other marine and land ecosystems and unique for the entire planet. This review describes these patterns and processes, from geological settings to biological processes, biodiversity and biogeographical patterns. It concludes with a brief discussion of current threats from anthropogenic activities to deep-sea habitats and their fauna. Investigations of deep-sea habitats and their fauna began in the late 19th century. In the intervening years, technological developments and stimulating discoveries have promoted deep-sea research and changed our way of understanding life on the planet. Nevertheless, the deep sea is still mostly unknown and current discovery rates of both habitats and species remain high. The geological, physical and geochemical settings of the deep-sea floor and the water column form a series of different habitats with unique characteristics that support specific faunal communities. Since 1840, 28 new habitats/ecosystems have been discovered from the shelf break to the deep trenches and discoveries of new habitats are still happening in the early 21st century. However, for most of these habitats the global area covered is unknown or has been only very roughly estimated; an even smaller – indeed, minimal – proportion has actually been sampled and investigated. We currently perceive most of the deep-sea ecosystems as heterotrophic, depending ultimately on the flux on organic matter produced in the overlying surface ocean through photosynthesis. The resulting strong food limitation thus shapes deep-sea biota and communities, with exceptions only in reducing ecosystems such as inter alia hydrothermal vents or cold seeps. Here, chemoautolithotrophic bacteria play the role of primary producers fuelled by chemical energy sources rather than sunlight. Other ecosystems, such as seamounts, canyons or cold-water corals have an increased productivity through specific physical processes, such as topographic modification of currents and enhanced transport of particles and detrital matter. Because of its unique abiotic attributes, the deep sea hosts a specialized fauna. Although there are no phyla unique to deep waters, at lower taxonomic levels the composition of the fauna is distinct from that found in the upper ocean. Amongst other characteristic patterns, deep-sea species may exhibit either gigantism or dwarfism, related to the decrease in food availability with depth. Food limitation on the seafloor and water column is also reflected in the trophic structure of heterotrophic deep-sea communities, which are adapted to low energy availability. In most of these heterotrophic habitats, the dominant megafauna is composed of detritivores, while filter feeders are abundant in habitats with hard substrata (e.g. mid-ocean ridges, seamounts, canyon walls and coral reefs). Chemoautotrophy through symbiotic relationships is dominant in reducing habitats. Deep-sea biodiversity is among of the highest on the planet, mainly composed of macro and meiofauna, with high evenness. This is true for most of the continental margins and abyssal plains with hot spots of diversity such as seamounts or cold-water corals. However, in some ecosystems with particularly "extreme" physicochemical processes (e.g. hydrothermal vents), biodiversity is low but abundance and biomass are high and the communities are dominated by a few species. Two large-scale diversity patterns have been discussed for deep-sea benthic communities. First, a unimodal relationship between diversity and depth is observed, with a peak at intermediate depths (2000–3000 m), although this is not universal and particular abiotic processes can modify the trend. Secondly, a poleward trend of decreasing diversity has been discussed, but this remains controversial and studies with larger and more robust data sets are needed. Because of the paucity in our knowledge of habitat coverage and species composition, biogeographic studies are mostly based on regional data or on specific taxonomic groups. Recently, global biogeographic provinces for the pelagic and benthic deep ocean have been described, using environmental and, where data were available, taxonomic information. This classification described 30 pelagic provinces and 38 benthic provinces divided into 4 depth ranges, as well as 10 hydrothermal vent provinces. One of the major issues faced by deep-sea biodiversity and biogeographical studies is related to the high number of species new to science that are collected regularly, together with the slow description rates for these new species. Taxonomic coordination at the global scale is particularly difficult, but is essential if we are to analyse large diversity and biogeographic trends.

Mapping trawling pressure on the benthic habitats is needed as background to support an ecosystem approach to fisheries management. The extent and intensity of bottom trawling on the European continental shelf (0–1000 m) was analysed from logbook statistics and vessel monitoring system data for 2010–2012 at a grid cell resolution of 1 × 1 min longitude and latitude. Trawling intensity profiles with seabed impact at the surface and subsurface level are presented for 14 management areas in the North-east Atlantic, Baltic Sea and Mediterranean Sea. The footprint of the management areas ranged between 53–99% and 6–94% for the depth zone from 0 to 200 m (Shallow) and from 201 to 1000 m (Deep), respectively. The footprint was estimated as the total area of all grid cells that were trawled fully or partially. Excluding the untrawled proportions reduced the footprint estimates to 28–85% and 2–77%. Largest footprints per unit landings were observed off Portugal and in the Mediterranean Sea. Mean trawling intensity ranged between 0.5 and 8.5 times per year, but was less in the Deep zone with a maximum intensity of 6.4. Highest intensities were recorded in the Skagerrak-Kattegat, Iberian Portuguese area, Tyrrhenian Sea and Adriatic Sea. Bottom trawling was highly aggregated. For the Shallow zone the seabed area where 90% of the effort occurred comprised between 17% and 63% (median 36%) of the management area. Footprints were high over a broad range of soft sediment habitats. Using the longevity distribution of the untrawled infaunal community, the seabed integrity was estimated as the proportion of the biomass of benthic taxa where the trawling interval at the subsurface level exceeds their life span. Seabed integrity was low (<0.1) in large parts of the European continental shelfs, although smaller pockets of seabed with higher integrity values occur. The methods developed here integrate official fishing effort statistics and industry-based gear information to provide high-resolution pressure maps and indicators, which greatly improve the basis for assessing and managing benthic pressure from bottom trawling. Further they provide quantitative estimates of trawling impact on a continuous scale by which managers can steer.

Bottom trawlers land around 19 million tons of fish and invertebrates annually, almost one-quarter of wild marine landings. The extent of bottom trawling footprint (seabed area trawled at least once in a specified region and time period) is often contested but poorly described. We quantify footprints using high-resolution satellite vessel monitoring system (VMS) and logbook data on 24 continental shelves and slopes to 1,000-m depth over at least 2 years. Trawling footprint varied markedly among regions: from 50% in some European seas. Overall, 14% of the 7.8 million-km² study area was trawled, and 86% was not trawled. Trawling activity was aggregated; the most intensively trawled areas accounting for 90% of activity comprised 77% of footprint on average. Regional swept area ratio (SAR; ratio of total swept area trawled annually to total area of region, a metric of trawling intensity) and footprint area were related, providing an approach to estimate regional trawling footprints when high-resolution spatial data are unavailable. If SAR was ≤0.1, as in 8 of 24 regions, there was >95% probability that >90% of seabed was not trawled. If SAR was 7.9, equal to the highest SAR recorded, there was >95% probability that >70% of seabed was trawled. Footprints were smaller and SAR was ≤0.25 in regions where fishing rates consistently met international sustainability benchmarks for fish stocks, implying collateral environmental benefits from sustainable fishing. Significance We 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.

Megafauna on bioherms (large biological structures) of the scleractinian coral Lophelia pertusa at 240-290 m depth in the Haltenbanken- Fmyabanken area was investigated by video-recording. Sixteen transects from soft bottom with scattered patches of stones below bioherms to top of bioherms were analysed. Fauna patterns were related to the near-bottom currents. The orientation of the gorgonianParamuricea placomus was used as an indicator of the direction of the main currents at the bioherms. The bioherms were 2 to 31 m high, and had a basal area ranging from 1 500 to 50 600 m2. 36 taxa were identified, of which five taxa only occurred on the bioherms, and five only on the soft bottom with scattered stones. The diversity, H', was highest in the zone of dead Lophelia, and lowest on the silty clay. None of the 26 taxa observed on stones were spesific for this habitat, but occurred also on the bioherms or the soft bottom. The area with Lophelia rubble, near the basis of the bioherms, had the lowest number of taxa (15), but the highest average density of individuals (7.92 ind./1Om2). Different sponges, gorgonians (Paragorgia arborea, Paramuricea placomus, Primnoa resedaeformis), squat lobsters (Munida sarsi), redfish (Sebastes spp.) and saithe (Pollachius virens) dominated in terms of individuals per area. Diversity, density of sponges and density of gorgonians were highest on the down-current side of the bioherms. Saithe were observed with highest densities near the basis of the bioherms, on the up-current side, while redfish had highest densities on the parallel-current side of the bioherm top. These results indicate that near bottom currents and turbulence are factors affecting the fauna on Lophelia bioherms