Abstract. 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 Correspondence to: E. Ramirez-Llodra (ezr@icm.csic.es) 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 Published by Copernicus Publications on behalf of the European Geosciences Union. 2852 E. Ramirez-Llodra et al.: Unique attributes of the world's largest ecosystem 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...

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.

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U-series age patterns obtained on reef framework-forming cold-water corals collected over a nearly 6,000 km long continental margin sector, extending from off Mauritania to the south-western Barents Sea reveal strong climate influences on the geographical distribution and sustained development of these ecosystems. During glacial times densely populated cold-water coral reefs flourished in the temperate east Atlantic, where at present only scarce live coral occurrences exist. In contrast, climate warming induces a rapid northward colonization of cold-water coral reefs with the biogeographic limit advancing from ∼45• N to ∼70• N. Thus, we invoke here that north-south oscillations of the polar front during the past glacial-interglacial cycles and the consequent displacement of cold nutrient-rich intermediate waters and productivity drives the decline and expansion of cold-water coral ecosystems and its biogeographic limits in the northeast Atlantic.