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

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. <br><br> 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, 27 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 <i>inter alia</i> hydrothermal vents or cold seeps, where 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 deep-sea communities, which are adapted to low energy availability. In most of the heterotrophic deep-sea settings, 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) and chemoautotrophy through symbiotic relationships is dominant in reducing habitats. <br>...

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.

Effective monitoring of populations and communities is a prerequisite for ecosystembased management of marine areas. However, monitoring programs often neglect important sources of error and thus can lead to biased estimates, spurious conclusions and false management actions. One such source of error is 'imperfect detectability', i.e. the inability of investigators to detect all individuals or all species in a surveyed area. Although there has been great effort to develop monitoring methods that account for imperfect detectability, the application of such methods in the marine environment is not as apparent as in other systems. Plot sampling is by far the most commonly applied method for biological monitoring in the marine environment, yet it largely ignores detectability issues. However, distance sampling, mark-recapture methods, repeated presence-absence surveys for occupancy estimation, and removal methods do estimate detection probabilities and provide unbiased estimates of state variables. We review these methods and the relevant tools for their application in studies on marine populations and communities, with the aim of assisting marine biologists and managers to understand the limitations and pitfalls associated with some approaches and to select the best available methods for their monitoring needs.

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