Foraminiferal research lies at the border between geology and biology. Benthic foraminifera are a major component of marine communities, highly sensitive to environmental influences, and the most abundant benthic organisms preserved in the deep-sea fossil record. These characteristics make them important tools for reconstructing ancient oceans. Much of the recent work concerns the search for palaeoceanographic proxies, particularly for the key parameters of surface primary productivity and bottom-water oxygenation. At small spatial scales, organic flux and pore-water oxygen profiles are believed to control the depths at which species live within the sediment (their 'microhabitats'). Epifaunal/shallow infaunal species require oxygen and labile food and prefer relatively oligotrophic settings. Some deep infaunal species can tolerate anoxia and are closely linked to redox fronts within the sediment; they consume more refractory organic matter, and flourish in relatively eutrophic environments. Food and oxygen availability are also key factors at large (i.e. regional) spatial scales. Organic flux to the sea floor, and its seasonality, strongly influences faunal densities, species compositions and diversity parameters. Species tend to be associated with higher or lower flux rates and the annual flux range of 2-3 g Corg m-2 appears to mark an important faunal boundary. The oxygen requirements of benthic foraminifera are not well understood. It has been proposed that species distributions reflect oxygen concentrations up to fairly high values (3 ml l-1 or more). Other evidence suggests that oxygen only begins to affect community parameters at concentrations < 0.5 ml l-1. Different species clearly have different thresholds, however, creating species successions along oxygen gradients. Other factors such as sediment type, hydrostatic pressure and attributes of bottom-water masses (particularly carbonate undersaturation and current flow) influence foraminiferal distributions, particularly on continental margins where strong seafloor environmental gradients exist. Epifaunal species living on elevated substrata are directly exposed to bottom-water masses and flourish where suspended food particles are advected by strong currents. Biological interactions, e.g. predation and competition, must also play a role, although this is poorly understood and difficult to quantify. Despite often clear qualitative links between environmental and faunal parameters, the development of quantitative foraminiferal proxies remains problematic. Many of these difficulties arise because species can tolerate a wide range of non-optimal conditions and do not exhibit simple relationships with particular parameters. Some progress has been made, however, in formulating proxies for organic fluxes and bottom-water oxygenation. Flux proxies are based on the Benthic Foraminiferal Accumulation Rate and multivariate analyses of species data. Oxygen proxies utilise the relative proportions of epifaunal (oxyphilic) and deep infaunal (low-oxygen tolerant) species. Yet many problems remain, particularly those concerning the calibration of proxies, the closely interwoven effects of oxygen and food availability, and the relationship between living assemblages and those preserved in the permanent sediment record.

Benthic foraminifera (Protista) as tools in deep-water palaeoceanography: environmental influences on faunal characteristics.

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Cold‐Water Corals: The Biology and Geology of Deep‐Sea Coral Habitats

The fully natatory families of the janiroidean Asellota, the munnopsids sensu lato, include a group of genera that blurs the distinction between the Ilyarachnidae and the Eurycopidae. This work determines the interrelationships of the ilyarachnid-like eurycopids, and shows that they are a monophyletic group. In so doing, the family-level systematics of the munnopsids is revised. The ilyarachnid-like eurycopids are assigned to a newly constituted subfamily, the Lipomerinae, and five genera are described, four of which are new. A diagnosis of the subfamily Lipomerinae with a key to the superspecific taxa is included in the taxonomic part. One species in each superspecific taxon is fully described. Coperonus n. gen. is a primarily Southern Hemisphere group with several species in the south Atlantic and around the Antarctic continent. The most ilyarachnid-like genus is Hapsidohedra n. gen., which may have a cosmopolitan distribution. Lionectes n. gen. is found in Antarctic waters. The pan-Atlantic genus Lipomera Tattersall is further divided into three new subgenera. One of these subgenera, L. (Tetracope) n. subgen., has a coiled gut, a rare occurrence among Crustacea. The cosmopolitan Mimocopelates n. gen.is represented by a North Atlantic species group based on the species M. longipes n. sp.,and an equatorial species M. anchibraziliensis n. sp. Character analyses of most munnopsid genera present the characters that reveal relationships between taxa. The character states were assigned evolutionary polarities by comparison with characters in a presumed munnopsid sister group, the Acanthaspidiidae, and other janiroidean families. Computerized phylogenetic analyses produced cladograms that were not fully resolved, but had significantly lower homoplasy values than a tree based on previous classifications. The Lipomerinae were the most significantly monophyletic group of the munnopsid taxa, and a consensus tree of all possible cladograms confirmed the monophyly of the Lipomerinae. Because the previous classifications of the families were not consistent with the most parsimonious cladograms, the following proposals are made: all munnopsid families should be placed into one large family, the Munnopsidae; the Ilyarachnidae and the Munnopsidae sensu stricto should be demoted to subfamilial status; and the current subfamilies of the Eurycopidae should be retained. This new classification recognizes the monophyly of the Lipomerinae within the Munnopsidae. Alternatives to this classification are discussed. Supporting the text are appendices illustrating and defining morphological terms.

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A Systematic Revision of the Deep-Sea Subfamily Lipomerinae of the Isopod Crustacean Family Munnopsidae

The deep sea plays a critical role in global climate regulation through uptake and storage of heat and carbon dioxide. However, this regulating service causes warming, acidification and deoxygenation of deep waters, leading to decreased food availability at the seafloor. These changes and their projections are likely to affect productivity, biodiversity and distributions of deep-sea fauna, thereby compromising key ecosystem services. Understanding how climate change can lead to shifts in deep-sea species distributions is critically important in developing management measures. We used environmental niche modelling along with the best available species occurrence data and environmental parameters to model habitat suitability for key cold-water coral and commercially important deep-sea fish species under present-day (1951-2000) environmental conditions and to project changes under severe, high emissions future (2081-2100) climate projections (RCP8.5 scenario) for the North Atlantic Ocean. Our models projected a decrease of 28%-100% in suitable habitat for cold-water corals and a shift in suitable habitat for deep-sea fishes of 2.0°-9.9° towards higher latitudes. The largest reductions in suitable habitat were projected for the scleractinian coral Lophelia pertusa and the octocoral Paragorgia arborea, with declines of at least 79% and 99% respectively. We projected the expansion of suitable habitat by 2100 only for the fishes Helicolenus dactylopterus and Sebastes mentella (20%-30%), mostly through northern latitudinal range expansion. Our results projected limited climate refugia locations in the North Atlantic by 2100 for scleractinian corals (30%-42% of present-day suitable habitat), even smaller refugia locations for the octocorals Acanella arbuscula and Acanthogorgia armata (6%-14%), and almost no refugia for P. arborea. Our results emphasize the need to understand how anticipated climate change will affect the distribution of deep-sea species including commercially important fishes and foundation species, and highlight the importance of identifying and preserving climate refugia for a range of area-based planning and management tools.

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Climate‐induced changes in the suitable habitat of cold‐water corals and commercially important deep‐sea fishes in the North Atlantic

The distribution of deep-water corals of the Flemish Cap, Flemish Pass, and the Grand Banks of Newfoundland is described based on bycatch from Spanish/EU bottom trawl groundfish surveys between 40 and 1500 m depth. In all, 37 taxa of deep-water corals were identified in the study area: 21 alcyonaceans (including the gorgonians), 11 pennatulaceans, 2 solitary scleractinians, and 3 antipatharians. The greatest diversity of coral species was on the Flemish Cap. Corals were most abundant along the continental slope, between 600 and 1300 m depth. Soft corals (alcyonaceans), sea fans (gorgonians), and black corals (antipatharians) were most common on bedrock or gravel, whereas sea pens (pennatulaceans) and cup corals (solitary scleractinians) were found primarily on mud. The biomass of deep-water corals in the bycatches was highest in previously lightly trawled or untrawled areas, and generally low in the regularly fished grounds. The information derived from bottom-trawl bycatch records is not sufficient to map vulnerable marine ecosystems (VMEs) accurately, but pending more detailed habitat mapping, it provides a valuable indication of the presence/absence of VMEs that can be used to propose the candidate areas for bottom fishery closures or other conservation measures.

Distribution of deep-water corals of the Flemish Cap, Flemish Pass and the Grand Banks of Newfoundland (Northwest Atlantic Ocean): interaction with fishing activities

The distribution of nine cold-water corals in the cold temperate North Atlantic is described based on existing records. The species studied are Lophelia pertusa, Madrepora oculata, Paragorgia arborea, Primnoa resedaeformis, Paramuricea placomus, Acanella arbuscula, Isidella lofotensis, Radicipes gracilis and Acanthogorgia armata. The compiled number of records were 5,854, of which 4,875 were obtained from databases and 979 from publications. Distribution maps were produced and temperature, broad-scale topography and current patterns were considered to understand the distribution patterns and conditions at which the species thrive. Currents connecting shelves and slopes above 500 m can explain the wide spatial distribution of L. pertusa, P. arborea and P. resedaeformis. However, L. pertusa is scarce on the western side of the North Atlantic, P. arborea has only few records off Iceland and A. arbuscula and A. armata are not found on the Norwegian shelf. The differences in distribution patterns between species indicate that they are differently affected by the topographic barrier between the North Atlantic and the Nordic Seas. Present knowledge of dispersal ability of cold-water corals does not allow a firm causal explanation to the observed distribution patterns. These, however, are indicative of biogeographic provinces relevant to cold-water corals and their habitat requirements.

Distribution of nine cold-water coral species (Scleractinia and Gorgonacea) in the cold temperate North Atlantic: effects of bathymetry and hydrography

In the deep waters of the Nordic Seas and adjacent areas, several benthic habitats such as cold-water coral reefs, coral gardens, and deep-sea sponge aggregations have been classified as vulnerable marine ecosystems (VMEs), due to their uniqueness, limited spatial extent, physical fragility, and slow recovery rate. In the last decade observations carried out by habitat mapping programmes in Norway, Iceland and more recently in the Faroe Islands have substantially increased knowledge on the distribution of VMEs in the Nordic Seas. Nevertheless, large areas have not been explored due to the cost and logistics of obtaining observations in the deep-sea. Species distribution models can be used to predict the distribution of VMEs and their indicator species. Here we present the predicted distribution of 44 VME indicator taxa including 20 sponges, 17 cold-water corals and 7 seapens in the Nordic Seas based on data compiled and models developed by the NovasArc project (2016-2018). Models for 44 VME indicator species were developed using a maximum entropy approach (MaxEnt) using an extensive database compiled from habitat mapping surveys, by-catch data from bottom fish surveys, and records from reports and peer reviewed publications. Modeled distributions showed good agreement with observations. Niche overlap measures were used to identify seven groups and four subgroups of VME indicator taxa that co-occur. These were consistent with the species composition of known biotopes in the study area. A VME Index that combine the predictions for all VME indicator species was computed to identify particularly valuable and vulnerable ecosystems that should be targets of further exploration and conservation efforts. Such areas were identified at shelf break and slope off Iceland, the Faroe Islands, and central Norway, and the continental shelf off southern Greenland. The predicted distribution of VMEs in arctic and sub-arctic waters allows for the evaluation of interactions with fisheries and other anthropogenic activities and provides an important input for managers.

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Predicting the Distribution of Indicator Taxa of Vulnerable Marine Ecosystems in the Arctic and Sub-arctic Waters of the Nordic Seas

Examination of 32 species from seven families of marine asellote isopods (Crustacea, Malacostraca) showed that 15 of these species were infested with ciliate epibionts. The diversity of the ciliate epibiosis was fairly high (mean = 3.8 epibiont ciliate species per host species; range 1 to 10 species). Ciliates were absent from species of the family Janiridae and were rare on desmosomatids. The prevalence was high (≥ 37%) on two ischnomesid species (Ischnomesus norvegicus and Heteromesus frigidus), and on three munnopsid species (genus Ilyarachna). Mean intensity was fairly low (1.3–10.5) overall, but high on I. norvegicus (29.2). Some ciliate species showed host preferences. Predatory ciliates (subclass Suctoria) were most common on host species of the families Munnidae and the Nannoniscidae and some of the Ischnomesidae species, whereas suspension-feeding ciliates (subclass Peritrichia) were found on six host species only, belonging to the family Ischnomesidae or the subfamily Ilyarachninae (Mun...

Ciliate (protozoa) epibionts of deep-water asellote isopods (crustacea): pattern and diversity

This report presents results from the NovasArc project that has collated data on the distribution of vulnerable marine ecosystems (VMEs) in Arctic and sub-Arctic waters. Eleven VMEs were identified ...

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Vulnerable marine ecosystems (VMEs) : Coral and sponge VMEs in Arctic and sub-Arctic waters – Distribution and threats

Stauroteuthis syrtensis and Cirroteuthis muelleri were studied based on representative material collected in Greenland, Iceland, and the Barents Sea and adjacent deep-sea areas (96 stations and 165 individuals for S. syrtensis, and 82 stations and 215 individuals for C. muelleri). The species occur on the border of the Arctic and the northern North Atlantic; S. syrtensis is a southern species and C. muelleri is a northern species, in the area from the Denmark and Davis Straits and associated underwater ridges. These species were demonstrated to be ecological analogues since they: a) occupy the same depths; b) reach the same abundance density; c) reach virtually the same biomass density when corrected for size differences; and d) their food spectra largely coincide. Temperature, not depth, is a principal factor that prevents these species from crossing the border between the Arctic and the northern North Atlantic, with mean habitat temperatures of 3.5 ± 0.06 °C for S. syrtensis and 0.3 ± 0.06 °C for C. muelleri. This study provides unprecedented density estimates for the North Atlantic and Arctic Cirrata: the highest in the world ocean in terms of biomass and second highest in terms of numerical abundance. Food spectra of the studied species consist of Calanoida (including eurybathic Paraeucheta spp. and Calanus spp., and deep-sea Bradyidius similis), Mysidacea and Isopoda (including deep-sea Eurycopinae) (Crustacea), and in C. muelleri also Cumacea, Amphipoda (Crustacea) and Polychaeta (Polynoidae). Additionally, this study: a) provides specific growth and morphometric equations; b) provides equations to estimate mantle length (ML) and body mass (BM) from beak measurements, and BM of fresh individuals (for modelling purposes or predators diet analyses) from BM of fixed individuals; c) reports maximum sizes for C. muelleri (ML 170 mm, total length 400 mm, BM 781.0 g); and d) reports findings of rarely encountered juvenile cirrates. 

Deep and cold: are Boreal and Arctic finned octopods, Stauroteuthis syrtensis and Cirroteuthis muelleri (Cephalopoda, Octopoda, Cirrata), ecological analogues?

Steinunn H. Ólafsdóttir

Papers

Distribution of nine cold-water coral species (Scleractinia and Gorgonacea) in the cold temperate North Atlantic: effects of bathymetry and hydrography

Predicting the Distribution of Indicator Taxa of Vulnerable Marine Ecosystems in the Arctic and Sub-arctic Waters of the Nordic Seas

Ciliate (protozoa) epibionts of deep-water asellote isopods (crustacea): pattern and diversity

Vulnerable marine ecosystems (VMEs) : Coral and sponge VMEs in Arctic and sub-Arctic waters – Distribution and threats

Deep and cold: are Boreal and Arctic finned octopods, Stauroteuthis syrtensis and Cirroteuthis muelleri (Cephalopoda, Octopoda, Cirrata), ecological analogues?