Ecology of bathyal polychaete fauna at an Arctic-Atlantic boundary (Faroe-Shetland Channel, North-east Atlantic)

By reference to a series of 15 sampling stations spanning the West Shetland Slope (150-1000 m; Faroe-Shetland Channel, North-east Atlantic) we examined the potential environmental controls on the standing stock, diversity and composition of the polychaete fauna. In contrast to the majority of studied bathyal environments, the Faroe-Shetland Channel has a highly complex and dynamic hydrographic regime, particularly notable for extreme thermal variability at mid-slope depths (i.e. 7°C range at ca. 500 m). Contrary to general expectation, polychaete biomass increased (rather than decreased) with depth. Species diversity exhibited a parabolic pattern with depth, maximum diversity occurring at depths of 350-550 m, rather shallower than observed in other bathyal studies, and possibly linked with a maximum in habitat temperature range. Multivariate analyses of faunal composition suggested a separation of the sampling stations into a shallower and a deeper group, with temperature exerting a major control on polychaete species distributions. The decline in diversity below 600 m (i.e. the descending limb of the parabolic relationship) may be a result of historically limited immigration/recolonization of the thermally isolated Arctic deep-water basins that feed the cold-water flow through the Faroe-Shetland Channel. The bathymetric distribution of polychaetes and other benthos in this region appears to be intimately linked with the thermal regime, having a long-term impact (geological timescales) on the deep-water species pool and leading to local enhancement of diversity where cold- and warm-water masses meet and mix.     

Biodiversity and historical biogeography of stalked crinoids (Echinodermata) in the deep sea

About 95 species of stalked crinoids are now described from 60m to hadal depths, but our knowledge remains far from complete. Depending on which species concept is used, estimates of species richness can be dramatically different. It is necessary to have a homogeneous concept for taxonomic units. The abundance of the crinoid fossil record allows a discussion of the ancestry of deep sea crinoid fauna. Stalked crinoids have a horizontal diversity pattern with three regional centres of high diversity (i.e. western tropical Pacific, western tropical Atlantic and north-eastern Atlantic). Vertical patterns show two faunal strata which vary in importance among provinces. The epibathyal stratum has apparently remained relatively similar in intertropical areas since the Mesozoic. Despite environmental changes related to glaciation since the Middle Miocene, the deepest crinoid fauna (i.e. the deep sea fauna sensu stricto at depths more than 1000 ± 200 m) have a very ancient origin with a dispersion closely related to plate tectonics. The bathyal fauna on hard substrates includes a few living fossils and has a high historical interest.     

Leptostraca,Mysidacea, Isopoda,and Decapoda (Anomura) (Crustacea,Malacostraca) of the Chukchi Sea and Adjacent Waters: Hydrological Regime and Species Composition

The hydrological regime of the region is briefly outlined. The distribution of waters of Arctic and Pacific origin is discussed. Based on the original and literature data, it has been found that 73 species of the investigated orders occur in the northern part of the Bering Sea, in the Chukchi and East Siberian seas, and in the adjoining areas of the continental slope of the Arctic basin at depths down to 500 m. These are 1 leptostracan species, 11 anomurans, 22 mysids, and 39 free-living isopods. The distribution of these species by depth and in relation to the thermal and salinity characteristics of waters is analyzed.     

Cumaceans as Indicators of Atlantic Waters over the Continental Slope of the Arctic Ocean

The material on cumacean fauna of the continental slope of the Arctic Ocean (more than 80 samples) was collected during expeditions with the ships Persei in 1925, 1928, and 1936; Sadko in 1936; and Polarstern in 1990, 1993, and 1995. Examination of the samples (150–900 m depth) revealed 19 species of Cumacea in this region. Three Arcto-Atlantic bathyal species (Hemilamprops uniplicatus, Diastylis echinata, Campylaspis globosa) are markers of the places over the continental slope where Atlantic intermediate waters penetrate into the Arctic Ocean and reach to the bottom. These species are widespread in the North Atlantic (100 to 2882 m depth); on the continental slope of the Arctic Ocean, they occur in a narrow depth range (200 to 900 m) in areas where positive temperatures from 0.09 to 1.78°C and high salinity of 34.75 to 34.87 prevail. The locations of these species is delineated on maps as an interrupted belt extending from northeastern Greenland to the East Siberian Sea and further north up to 79°30 N and 148° E, which corresponds to the route of penetration of Atlantic intermediate waters into the Arctic Ocean.     

Structure and distribution of pelagic ostracods (Ostracoda: Myodocopa) in the Arctic Ocean

The study of the pelagic ostracod fauna of the Arctic Ocean based on materials collected by numerous Russian expeditions (1929–1993) and data from the literature showed the extreme poorness of the Arctic pelagic ostracod fauna, its mainly North Atlantic genesis and complete isolation from the Pacific fauna. Maximum ostracod abundance was observed in the epipelagic zone, and the greatest species diversity occurred in the relatively warm deep Atlantic layer throughout the year. To the north, east, and west of Franz Josef Land and Spitsbergen, the number of species and abundance indices of pelagic ostracods were decreased. In superficial water layers of the Central Arctic, maximum ostracod density and biomass were recorded in June and September. The best bioindicator of warm Atlantic water in the Arctic basin is Obtusoecia obtusata; and of cold polar water in the North Atlantic, Boroecia maxima.     

Demersal fish assemblages and spatial diversity patterns in the Arctic-Atlantic transition zone in the Barents Sea

Direct and indirect effects of global warming are expected to be pronounced and fast in the Arctic, impacting terrestrial, freshwater and marine ecosystems. The Barents Sea is a high latitude shelf Sea and a boundary area between arctic and boreal faunas. These faunas are likely to respond differently to changes in climate. In addition, the Barents Sea is highly impacted by fisheries and other human activities. This strong human presence places great demands on scientific investigation and advisory capacity. In order to identify basic community structures against which future climate related or other human induced changes could be evaluated, we analyzed species composition and diversity of demersal fish in the Barents Sea. We found six main assemblages that were separated along depth and temperature gradients. There are indications that climate driven changes have already taken place, since boreal species were found in large parts of the Barents Sea shelf, including also the northern Arctic area. When modelling diversity as a function of depth and temperature, we found that two of the assemblages in the eastern Barents Sea showed lower diversity than expected from their depth and temperature. This is probably caused by low habitat complexity and the distance to the pool of boreal species in the western Barents Sea. In contrast coastal assemblages in south western Barents Sea and along Novaya Zemlya archipelago in the Eastern Barents Sea can be described as diversity "hotspots"; the South-western area had high density of species, abundance and biomass, and here some species have their northern distribution limit, whereas the Novaya Zemlya area has unique fauna of Arctic, coastal demersal fish. (see Information S1 for abstract in Russian).     

Explaining bathymetric diversity patterns in marine benthic invertebrates and demersal fishes: physiological contributions to adaptation of life at depth

Bathymetric biodiversity patterns of marine benthic invertebrates and demersal fishes have been identified in the extant fauna of the deep continental margins. Depth zonation is widespread and evident through a transition between shelf and slope fauna from the shelf break to 1000 m, and a transition between slope and abyssal fauna from 2000 to 3000 m; these transitions are characterised by high species turnover. A unimodal pattern of diversity with depth peaks between 1000 and 3000 m, despite the relatively low area represented by these depths. Zonation is thought to result from the colonisation of the deep sea by shallow‐water organisms following multiple mass extinction events throughout the Phanerozoic. The effects of low temperature and high pressure act across hierarchical levels of biological organisation and appear sufficient to limit the distributions of such shallow‐water species. Hydrostatic pressures of bathyal depths have consistently been identified experimentally as the maximum tolerated by shallow‐water and upper bathyal benthic invertebrates at in situ temperatures, and adaptation appears required for passage to deeper water in both benthic invertebrates and demersal fishes. Together, this suggests that a hyperbaric and thermal physiological bottleneck at bathyal depths contributes to bathymetric zonation. The peak of the unimodal diversity–depth pattern typically occurs at these depths even though the area represented by these depths is relatively low. Although it is recognised that, over long evolutionary time scales, shallow‐water diversity patterns are driven by speciation, little consideration has been given to the potential implications for species distribution patterns with depth. Molecular and morphological evidence indicates that cool bathyal waters are the primary site of adaptive radiation in the deep sea, and we hypothesise that bathymetric variation in speciation rates could drive the unimodal diversity–depth pattern over time. Thermal effects on metabolic‐rate‐dependent mutation and on generation times have been proposed to drive differences in speciation rates, which result in modern latitudinal biodiversity patterns over time. Clearly, this thermal mechanism alone cannot explain bathymetric patterns since temperature generally decreases with depth. We hypothesise that demonstrated physiological effects of high hydrostatic pressure and low temperature at bathyal depths, acting on shallow‐water taxa invading the deep sea, may invoke a stress–evolution mechanism by increasing mutagenic activity in germ cells, by inactivating canalisation during embryonic or larval development, by releasing hidden variation or mutagenic activity, or by activating or releasing transposable elements in larvae or adults. In this scenario, increased variation at a physiological bottleneck at bathyal depths results in elevated speciation rate. Adaptation that increases tolerance to high hydrostatic pressure and low temperature allows colonisation of abyssal depths and reduces the stress–evolution response, consequently returning speciation of deeper taxa to the background rate. Over time this mechanism could contribute to the unimodal diversity–depth pattern.     

Diversity and biogeographic patterns of the bryozoan fauna of the Faroe Islands

A total of 228 bryozoan species are recorded within the EEZ of the Faroe Islands, 74 of which are new to the area. Analysis of the distribution of the species among six sectors, each characterized by different environmental conditions, showed three faunal assemblages. Variation of the total Faroese bryozoan fauna and of the bryozoan fauna of most sectors, demonstrated significant negative relationships with depth. In general, analysis of the biogeographic composition showed a strong predominance of boreal over arctic species. However, with respect to faunas of each sector, the Norwegian Basin is characterized by a predominance of arctic species and may be regarded as a part of the Arctic Eurasian sub-region of the Arctic biogeographic region. Comparison of the bryozoan species of each sector with the bryozoan faunas of the other 12 areas in the North Atlantic and the neighbouring Arctic regions showed that only the Faroese shelf fauna has significant similarity with part of them, and thus can be regarded as part of the Scandinavian province of the Norwegian high-boreal sub-region of the Atlantic boreal region. Three sectors, the Faroese–Iceland Ridge, the Faroese–Shetland Channel and Norwegian Basin, belong to a transitional zone between the Atlantic Boreal and the Arctic biogeographic regions. The deep south-western sector forms a separate faunal cluster when compared with both the other sectors within the Faroese area and with the faunas of other large geographic areas, and may be regarded as a separate biogeographic zone of the Boreal Atlantic region due to its high proportion of specific species.     

Benthic foraminiferal extinctions linked to late Pliocene–Pleistocene deep-sea circulation changes in the northern Indian Ocean (ODP Sites 722 and 758)

During the late Pliocene–middle Pleistocene, 63 species of elongate, bathyal–upper abyssal benthic foraminifera (Extinction Group = Stilostomellidae, Pleurostomellidae, some Nodosariidae) declined in abundance and finally disappeared in the northern Indian Ocean (ODP Sites 722, 758), as part of the global extinction of at least 88 related species at this time. The detailed record of withdrawal of these species differs by depth and geography in the Indian Ocean. In northwest Indian Ocean Site 722 (2045 m), the Extinction Group of 54 species comprised 2–15% of the benthic foraminiferal fauna in the earliest Pleistocene, but declined dramatically during the onset of the mid-Pleistocene Transition (MPT) at 1.2–1.1 Ma, with all but three species disappearing by the end of the MPT (∼0.6 Ma). In northeast Indian Ocean Site 758 (2925 m), the Extinction Group of 44 species comprised 1–5% of the benthic foraminiferal fauna at ∼3.3–2.6 Ma, but declined in abundance and diversity in three steps, at ∼2.5, 1.7, and 1.2 Ma, with all but one species disappearing by the end of the MPT. At both sites there are strong positive correlations between the accumulation rate of the Extinction Group and proxies indicating low-oxygen conditions with a high organic carbon input. In both sites, there was a pulsed decline in Extinction Group abundance and species richness, especially in glacial periods, with some partial recoveries in interglacials. We infer that the glacial declines at the deeper Site 758 were a result of increased production of colder, well-ventilated Antarctic Bottom Water (AABW), particularly in the late Pliocene and during the MPT. The Extinction Group at shallower water depths (Site 722) were not impacted by the deeper water mass changes until the onset of the MPT, when cold, well-ventilated Glacial North Atlantic Intermediate Water (GNAIW) production increased and may have spread into the Indian Ocean. Increased chemical ventilation at various water depths since late Pliocene, particularly in glacial periods, possibly in association with decreased or more fluctuating organic carbon flux, might be responsible for the pulsed global decline and extinction of this rather specialised group of benthic foraminifera.     

Distribution and seasonal dynamics of <Emphasis Type="Italic">Boroecia maxima</Emphasis> (Ostracoda: Halocypridinae) in the Arctic basin and adjacent Atlantic waters

Based on materials obtained during Soviet and Russian expeditions to the Arctic basin and adjacent Atlantic waters during the period from 1929–1993, as well as data from the literature, we examined the distribution range of the common deep-sea ostracod Boroecia maxima. This species occurs throughout the pelagic water column (down to 3000 m depth and below) and prefers low water temperatures (surface Arctic and intermediate water masses). During the year, at all depths, the ratio between age groups of B. maxima remains virtually unchanged. Breeding apparently continues year-round. During the polar night, B. maxima makes no migration into warm Atlantic water, where wintering predatory zooplankton occur in large numbers, and thus avoids grazing pressure.     

From Europe to America: pliocene to recent trans-atlantic expansion of cold-water north atlantic molluscs

Data on the geographical distribution, phylogeny and fossil record of cool-temperate North Atlantic shell-bearing molluscs that live in waters shallower than 100 m depth belong to two biogeographic provinces, one in eastern North America north of Cape Cod, the other in northern Europe. Amphi-Atlantic species, which are found in both provinces, comprise 30.8% of the 402 species in the northeastern Atlantic and 47.3% of the 262 species in the northwestern Atlantic. Some 54.8% of these amphi-Atlantic species have phylogenetic origins in the North Pacific. Comparisons among fossil Atlantic faunas show that amphi-Atlantic distributions became established in the Middle Pliocene (about 3.5 million years ago), and that all represent westward expansions of European taxa to North America. No American taxa spread eastward to Europe without human assistance. These results are in accord with previous phylogeographic studies among populations within several amphi-Atlantic species. Explanations for the unidirectional expansion of species across the Atlantic remain uncertain, but may include smaller size and greater prior extinction of the North American as compared to the European fauna and biased transport mechanisms. Destruction of the European source fauna may jeopardize faunas on both sides of the Atlantic.     

THE BIOGEOGRAPHIC ORIGIN OF ARCTIC ENDEMIC SEAWEEDS: A THERMOGEOGRAPHIC VIEW1

The Arctic is geologically and biogeographically young, and the origin of its seaweed flora has been widely debated. The Arctic littoral biogeographic region dates from the latest Tertiary and Pleistocene. Following the opening of Bering Strait, about 3.5 mya, the “Great Trans‐Arctic Biotic Interchange” populated the Arctic with a fauna strongly dominated by species of North Pacific origin. The Thermogeographic Model (TM) demonstrates why climate and geography continued to support this pattern in the Pleistocene. Thus, Arctic and Atlantic subarctic species of seaweeds are likely to be evolutionarily “based” in the North Pacific, subarctic species are likely to be widespread in the warmer Arctic, and species of Atlantic Boreal or warmer origin are unlikely in the Arctic and Subarctic. Although Arctic seaweeds have been thought to have a greater affinity with the North Atlantic, we have reanalyzed the Arctic endemic algal flora, using the Thermogeographic Model and evolutionary trees based on molecular data, to demonstrate otherwise. There are 35 congeneric species of the six, abundant Arctic Rhodophyta that we treat in this paper; 32 of these species (91%) occur in the North Pacific, two species (6%) occur in the Boreal or warmer Atlantic Ocean, and a single species is panoceanic, but restricted to the Subarctic. Laminaria solidungula J. Agardh, a kelp Arctic “endemic” species, has 18 sister species. While only eleven (61%) occur in the North Pacific, this rapidly dispersing and evolving genus is a terminal member of a diverse family and order (Laminariales) widely accepted to have evolved in the North Pacific. Thus, both the physical/time‐based TM and the dominant biogeographic pattern of relatives of Arctic macrophytes suggest strong compliance with the evidence of zoology, geology, and paleoclimatology that the Arctic marine flora is largely of Pacific origin.     

Trans‐Pacific and trans‐Arctic pathways of the intertidal macroalga Fucus distichus L. reveal multiple glacial refugia and colonizations from the North Pacific to the North Atlantic

Aim We examined the phylogeography of the cold‐temperate macroalgal species Fucus distichus L., a key foundation species in rocky intertidal shores and the only Fucus species to occur naturally in both the North Pacific and the North Atlantic. Location North Pacific and North Atlantic oceans (42° to 77° N). Methods We genotyped individuals from 23 populations for a mitochondrial DNA (mtDNA) intergenic spacer (IGS) (n = 608) and the cytochrome c oxidase subunit I (COI) region (n = 276), as well as for six nuclear microsatellite loci (n = 592). Phylogeographic structure and connectivity were assessed using population genetic and phylogenetic network analyses. Results IGS mtDNA haplotype diversity was highest in the North Pacific, and divergence between Pacific haplotypes was much older than that of the single cluster of Atlantic haplotypes. Two ancestral Pacific IGS/COI clusters led to a widespread Atlantic cluster. High mtDNA and microsatellite diversities were observed in Prince William Sound, Alaska, 11 years after severe disturbance by the 1989 Exxon Valdez oil spill. Main conclusions At least two colonizations occurred from the older North Pacific populations to the North Atlantic between the opening of the Bering Strait and the onset of the Last Glacial Maximum. One colonization event was from the Japanese Archipelago/eastern Aleutians, and a second was from the Alaskan mainland around the Gulf of Alaska. Japanese populations probably arose from a single recolonization event from the eastern Aleutian Islands before the North Pacific–North Atlantic colonization. In the North Atlantic, the Last Glacial Maximum forced the species into at least two known glacial refugia: the Nova Scotia/Newfoundland (Canada) region and Andøya (northern Norway). The presence of two private haplotypes in the central Atlantic suggests the possibility of colonization from other refugia that are now too warm to support F. distichus. With the continuing decline in Arctic ice cover as a result of global climate change, renewed contact between North Pacific and North Atlantic populations of Fucus species is expected.     

Diversity and species composition of peracarids (Crustacea: Malacostraca) on the South Greenland shelf: spatial and temporal variation

The interannual variability in peracarid (Crustacea: Malacostraca; Amphipoda, Isopoda, Cumacea, Tanaidacea) species composition and diversity on the South Greenland shelf was studied at four stations over a sampling period of 3 years (2001, 2002 and 2004), using a Rauschert sled at depths of about 160 m. The South Greenland peracarids were relatively stable over the 3 years with respect to evenness and diversity. Moderate changes in temperature and salinity had negligible effects on the species composition, while sediment structure was found to be the most important environmental variable shaping the peracarid fauna.     

Benthic infaunal community variability on the northern Svalbard shelf

Marine benthic communities are effective indicators of environmental change. Yet in the Arctic, there are few empirical tests of how sustained climatic change may influence community structure. Northern Svalbard is influenced by both warm Atlantic and cold Arctic water masses, providing an opportunity to assess potential effects of long-term environmental changes by examining spatial variation in community structure. We examined benthic macroinfaunal communities and sediment pigments under Atlantic and Arctic water masses on the northern shelf and fjords of Svalbard. We report on infaunal biomass, abundance, species composition, and diversity at 10 stations spanning 79°–81°N and ranging in depth from 200 to 500?m. Benthic biomass averaged 128?g?WW?m^?2 (48–253?g?WW?m^?2), mean density was 3,635?ind.?m^?2 (780–7,660?ind.?m^?2), and species richness varied from 45 to 136?taxa?stn.^?1. Abundance-based community structure clustered stations in groups related to water mass characteristics, with Atlantic and Arctic shelf stations being well distinguished from each other. Dominant taxa were different in Atlantic- and Arctic-influenced locations. Faunal biomass was highest in the Atlantic-influenced fjords, followed by Arctic fjords and Arctic shelf stations, with Atlantic shelf stations having the lowest biomass. Species richness and diversity were inversely related to biomass. Patterns in faunal biomass were strongly correlated with sedimentary pigments (R ²?=?0.74 for chl a and R ²?=?0.77 for phaeopigments), with large differences in sedimentary pigment concentration among stations. These relationships suggest that benthic fauna on the northern Svalbard shelf are food limited and dependent on predictable, albeit episodic, delivery of organic matter from the water column.     

Patterns in deep-sea macrobenthos at the continental margin: standing crop,diversity and faunal change on the continental slope off Scotland

Depth-related patterns of macrobenthic community structure and composition have been studied from box-core samples from the Scottish continental slope where deep-sea trawling and oil exploration are becoming increasingly important. There is a strong pattern of declining biomass and faunal abundance with increasing depth, but results also indicate reduced biomass and numbers of macrobenthos in the shallowest samples from just below the shelf edge where there are coarse sediments and a regime of strong bottom currents. There is also reduced species diversity at the shallowest stations, probably caused by hydrodynamic disturbance, but no clear mid-slope peak in species diversity as described from the northwest Atlantic. Taxonomic composition of the macrobenthic community shows most change between about 1000 and 1200 m, expressed as a major dichotomy in multivariate analysis by cluster analysis and ordination. It also shows up as a step-like increase in the rate of accumulation of new macrofaunal species. This corresponds to a change in hydrodynamic regime, from a seabed rich in suspension- and interface-feeding epifauna, to one where biogenic traces from large, burrowing deposit feeders are well developed, and visible epifauna rare in seabed photographs. It also corresponds to the depth zone where earlier study of megafaunal echinoderms in trawl and epibenthic sled samples also shows a clear peak in across-slope rate of change in faunal composition.     

A new deep-sea hydroid (Cnidaria: Hydrozoa) from the Bering Sea Basin reveals high genetic relevance to Arctic and adjacent shallow-water species

The marine benthic fauna in Arctic shallow-water is reported to be a relatively young assemblage by species of either Pacific or Atlantic affinity. Whether current deep-sea Pacific species are included in the affinity or not is unknown. Combining morphological comparisons and genetic analyses, a new deep-sea hydroid to science, Sertularia xuelongi sp. nov. (Cnidaria: Hydrozoa: Sertulariidae), is described from the northern margin of the Bering Sea Basin at depths of 800–1570 m collected in 2010. It is characterized by slender and zigzag-shaped hydrocauli, alternately arranged hydrothecae and the absence of distal-lateral horns in fully matured female gonothecae. Its distribution, currently known only from Bering Sea Basin, suggests that it could not be an Arctic species with Pacific affinity. However, phylogenetic analyses based on the mitochondrial 16S rRNA gene show that it is clustered into a distinctive clade with four closely related species recorded from shallow-water of Northwest France, Iceland, Chukchi Sea and/or Bering Sea. In addition, its sequence similarity is highly relevant to these four species: Sertularia argentea (98.6 %), S. cupressina (98.8 %), S. plumosa (98.8 %) and S. robusta (99.4 %). All these provide a new insight into the relevance of North Pacific deep-sea species to the benthic fauna in Arctic and adjacent shallow-water. The taxonomic restriction of the genus Sertularia and the re-validation of the genus Polyserias are discussed. Future researches on more deep-sea species from Pacific and/or Atlantic are required to understand the evolution and speciation pattern involved in polar relevance.     

Global diversity of brittle stars (Echinodermata: Ophiuroidea)

This review presents a comprehensive overview of the current status regarding the global diversity of the echinoderm class Ophiuroidea, focussing on taxonomy and distribution patterns, with brief introduction to their anatomy, biology, phylogeny, and palaeontological history. A glossary of terms is provided. Species names and taxonomic decisions have been extracted from the literature and compiled in The World Ophiuroidea Database, part of the World Register of Marine Species (WoRMS). Ophiuroidea, with 2064 known species, are the largest class of Echinodermata. A table presents 16 families with numbers of genera and species. The largest are Amphiuridae (467), Ophiuridae (344 species) and Ophiacanthidae (319 species). A biogeographic analysis for all world oceans and all accepted species was performed, based on published distribution records. Approximately similar numbers of species were recorded from the shelf (n = 1313) and bathyal depth strata (1297). The Indo-Pacific region had the highest species richness overall (825 species) and at all depths. Adjacent regions were also relatively species rich, including the North Pacific (398), South Pacific (355) and Indian (316) due to the presence of many Indo-Pacific species that partially extended into these regions. A secondary region of enhanced species richness was found in the West Atlantic (335). Regions of relatively low species richness include the Arctic (73 species), East Atlantic (118), South America (124) and Antarctic (126).     

Global diversity of marine isopods (except asellota and crustacean symbionts)

The crustacean order Isopoda (excluding Asellota, crustacean symbionts and freshwater taxa) comprise 3154 described marine species in 379 genera in 37 families according to the WoRMS catalogue. The history of taxonomic discovery over the last two centuries is reviewed. Although a well defined order with the Peracarida, their relationship to other orders is not yet resolved but systematics of the major subordinal taxa is relatively well understood. Isopods range in size from less than 1 mm to Bathynomus giganteus at 365 mm long. They inhabit all marine habitats down to 7280 m depth but with few doubtful exceptions species have restricted biogeographic and bathymetric ranges. Four feeding categories are recognised as much on the basis of anecdotal evidence as hard data: detritus feeders and browsers, carnivores, parasites, and filter feeders. Notable among these are the Cymothooidea that range from predators and scavengers to external blood-sucking micropredators and parasites. Isopods brood 10-1600 eggs depending on individual species. Strong sexual dimorphism is characteristic of several families, notably in Gnathiidae where sessile males live with a harem of females while juvenile praniza stages are ectoparasites of fish. Protandry is known in Cymothoidae and protogyny in Anthuroidea. Some Paranthuridae are neotenous. About half of all coastal, shelf and upper bathyal species have been recorded in the MEOW temperate realms, 40% in tropical regions and the remainder in polar seas. The greatest concentration of temperate species is in Australasia; more have been recorded from temperate North Pacific than the North Atlantic. Of tropical regions, the Central Indo-Pacific is home to more species any other region. Isopods are decidedly asymmetrical latitudinally with 1.35 times as many species in temperate Southern Hemisphere than the temperate North Atlantic and northern Pacific, and almost four times as many Antarctic as Arctic species. More species are known from the bathyal and abyssal Antarctic than Arctic GOODS provinces, and more from the larger Pacific than Atlantic oceans. Two areas with many species known are the New Zealand-Kermadec and the Northern North Pacific provinces. Deep hard substrates such as found on seamounts and the slopes are underrepresented in samples. This, the documented numbers of undescribed species in recent collections and probable cryptic species suggest a large as yet undocumented fauna, potentially an order of magnitude greater than presently known.