https://www.cell.com/current-biology/pdf/S0960-9822(12)01138-4.pdf

The magnitude of global marine species diversity

On a role

. 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. Because of their remoteness, anthropogenic impacts on deep-sea ecosystems have not been addressed very thoroughly until recently. The depletion of biological and mineral resources on land and in shallow waters, coupled with technological developments, are promoting the increased interest in services provided by deep-water resources. Although often largely unknown, evidence for the effects of human activities in deep-water ecosystems – such as deep-sea mining, hydrocarbon exploration and exploitation, fishing, dumping and littering – is already accumulating. Because of our limited knowledge of deep-sea biodiversity and ecosystem functioning and because of the specific life-history adaptations of many deep-sea species (e.g. slow growth and delayed maturity), it is essential that the scientific community works closely with industry, conservation organisations and policy makers to develop robust and efficient conservation and management options.

/pdf/deep-diverse-and-definitely-different-unique-attributes-of-30cfoyr2tg.pdf

Deep, diverse and definitely different: unique attributes of the world's largest ecosystem

http://www.vliz.be/imisdocs/publications/130615.pdf

Exponential Decline of Deep-Sea Ecosystem Functioning Linked to Benthic Biodiversity Loss

We present the first global-scale analy- sis of standing stock (abundance and biomass) for 4 major size classes of deep-sea biota: bacteria, meta- zoan meiofauna, macrofauna and megafauna. The community standing stock decreases with depth; this is a universal phenomenon that involves a com- plex transition in the relative importance of the different size groups. Bacterial abundance and biomass show no decline with depth. All 3 animal size groups experience significant exponential de- creases in both abundance and biomass. The abun- dance of larger animals is significantly lower and decreases more rapidly than for smaller groups. The resulting drop in average body size with depth con- firms Thiel's size-structure hypothesis on very large spatial scales. In terms of their proportion of total community biomass, smaller size classes replace larger size classes. The upper continental slope is dominated by macrofaunal biomass, and the abyss by bacterial and meiofaunal biomass. The dramatic decrease in total community standing stock and the ascendancy of smaller organisms with depth has important implications for deep-sea biodiversity. The bathyal zone (200 to 4000 m) affords more eco- logical and evolutionary opportunity in the form of energy availability for larger organisms, and conse- quently supports higher macrofaunal and mega- faunal species diversity than the abyss (> 4000 m).

/pdf/global-bathymetric-patterns-of-standing-stock-and-body-size-1yi0l0isof.pdf

Global bathymetric patterns of standing stock and body size in the deep-sea benthos

The great variety of geological and hydrological conditions in the deep sea generates many different habitats. Some are only recently explored, although their true extent and geographical coverage are still not fully established. Both continental margins and mid-oceanic seafloors are much more complex ecologically, geologically, chemically and hydrodynamically than originally thought. As a result, fundamental patterns of species distribution first observed and explained in the context of relatively monotonous slopes and abyssal plains must now be re-evaluated in the light of this newly recognized habitat heterogeneity. Based on a global database of nematode genus composition, collected as part of the Census of Marine Life, we show that macrohabitat heterogeneity contributes significantly to total deep-sea nematode diversity on a global scale. Different deep-sea settings harbour specific nematode assemblages. Some of them, like coral rubble zones or nodule areas, are very diverse habitats. Factors such as increased substrate complexity in the case of nodules and corals seem to facilitate the co-existence of a large number of genera with different modes of life, ranging from sediment dwelling to epifaunal. Furthermore, strong biochemical gradients in the case of vents or seeps are responsible for the success of particular genera, which are not prominent in more typical soft sediments. Many nematode deep-sea genera are cosmopolitan, inhabiting a variety of deep-sea habitats and oceans, whereas only 21% of all deep-sea genera recorded are restricted to a single habitat. In addition to habitat heterogeneity, regional differences are important in structuring nematode assemblages. For instance, seeps from different regions yield different genera that thrive on the sulphidic sediments. This study also shows that many areas and habitats remain highly under-sampled, affecting our ability to understand fully the contribution of habitat heterogeneity versus regional differences to global nematode diversity.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1439-0485.2009.00352.x

The contribution of deep-sea macrohabitat heterogeneity to global nematode diversity

This chapter throws the attention on the meiobenthos of the deep northeast Atlantic. The main purpose of this chapter is to summarize new results from an area lying between 15°N and 53°N and extending from the continental margin of western Europe and northwest Africa to the Mid-Atlantic Ridge. It considers first the nature and scope of meiofaunal research in the northeast Atlantic and then discuss the environmental parameters, which are believed to influence meiofaunal organisms. This chapter then discusses the various types and scales of pattern observed among meiofaunal populations within the study area, progressing from the large-scale bathymetric and latitudinal trends and then to small-scale horizontal patterns within particular areas. Faunal densities and faunal composition are considered separately and compared with data from other regions. This chapter also deals with the distribution of meiofauna within sediment profiles and the temporal variability of populations. This chapter concludes by discussing the recent review of deep-sea meiofauna, which focused mainly on the abundance and biomass data from different oceans and on the relationship between the biomass of the meiofauna and that of other faunal components

/pdf/meiobenthos-of-the-deep-northeast-atlantic-2ol3itdrzc.pdf

Meiobenthos of the Deep Northeast Atlantic

The organic biogeochemistry of surface sediments from the Madeira Abyssal Plain has been investigated. Total organic carbon contents, organic carbon to nitrogen ratios, concentrations of hydrolysable amino acids, hydrolysable carbohydrates and lipids have been determined. In one case, the detailed lipid compositions were also investigated.



The sedimentary geochemistry appears to have been strongly influenced by the recent (ca. 1000 yrs.) deposition of a turbidite. Normally labile lipids are present in high concentrations through the turbidite and the numbers of the most abundant metazoan meiofauna (nematodes) closely correlates with the concentrations of the former. Concentrations of hydrolysable amino acids and carbohydrates, relative to total organic carbon, also decrease through the turbidite layer, but in the case of the carbohydrates tend to be higher in the underlying pelagic sediments. Deeper sediments are more oxidised than those at the surface. The concentration profile for manganese in one of the examined cores suggests that this anomaly arises through the action of manganese-reducing bacteria that are present in pelagic sediments underlying the turbidite.

Tim Ferrero

Papers

The contribution of deep-sea macrohabitat heterogeneity to global nematode diversity

Meiobenthos of the Deep Northeast Atlantic

The Biogeochemistry of Sediments from the Madeira Abyssal Plain — Preliminary Results