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The biodiversity of the deep Southern Ocean benthos

The Royal Society
Philosophical Transactions B
Authors:
Angelika Brandt at Senckenberg Research Institute
Angelika Brandt
Claude De Broyer at Royal Belgian Institute of Natural Sciences
Claude De Broyer
Ilse De Mesel at Royal Belgian Institute of Natural Sciences
Ilse De Mesel
Kari E. Ellingsen at Norwegian Institute for Nature Research, Tromsø, Norway
Kari E. Ellingsen
  • Norwegian Institute for Nature Research, Tromsø, Norway
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Abstract and Figures

Our knowledge of the biodiversity of the Southern Ocean (SO) deep benthos is scarce. In this review, we describe the general biodiversity patterns of meio-, macro- and megafaunal taxa, based on historical and recent expeditions, and against the background of the geological events and phylogenetic relationships that have influenced the biodiversity and evolution of the investigated taxa. The relationship of the fauna to environmental parameters, such as water depth, sediment type, food availability and carbonate solubility, as well as species interrelationships, probably have shaped present-day biodiversity patterns as much as evolution. However, different taxa exhibit different large-scale biodiversity and biogeographic patterns. Moreover, there is rarely any clear relationship of biodiversity pattern with depth, latitude or environmental parameters, such as sediment composition or grain size. Similarities and differences between the SO biodiversity and biodiversity of global oceans are outlined. The high percentage (often more than 90%) of new species in almost all taxa, as well as the high degree of endemism of many groups, may reflect undersampling of the area, and it is likely to decrease as more information is gathered about SO deep-sea biodiversity by future expeditions. Indeed, among certain taxa such as the Foraminifera, close links at the species level are already apparent between deep Weddell Sea faunas and those from similar depths in the North Atlantic and Arctic. With regard to the vertical zonation from the shelf edge into deep water, biodiversity patterns among some taxa in the SO might differ from those in other deep-sea areas, due to the deep Antarctic shelf and the evolution of eurybathy in many species, as well as to deep-water production that can fuel the SO deep sea with freshly produced organic matter derived not only from phytoplankton, but also from ice algae.
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... This agrees with the findings of other expeditions e.g., [26,56,117,118]. In general agreement with our results, Crustacea e.g., [56,58,78,[119][120][121], Polychaeta e.g., [99,122,123], and Mollusca e.g., [56,78] have all been reported to be among the dominant taxa. In our study, Polychaeta were the most abundant taxa overall, contributing more than 10% of the overall community composition across all regions and depth zones and over 30% in some regions (Northwest Atlantic). ...
... Ophiuroidea and Gastropoda were present in small amounts throughout the study area with the exception of the Southwest Atlantic shelf, where both taxa dominated the community with 22% abundance each. Ophiuroidea and Gastropoda are found throughout the Atlantic and Southern Ocean e.g., [58,[137][138][139][140][141] but have rarely been found to dominate comparable macrofaunal community compositions e.g., [28,78]. Our results may support the hypothesis that benthic fauna assemblages are distributed patchily e.g., [142,143], with Ophiuroidea and Gastropoda only dominant within a single region and depth group that consists of only one station. ...
Pan-Atlantic Comparison of Deep-Sea Macro-and Megabenthos
Article
Full-text available
  • Jun 2023
  • Diversity
Citation: Kürzel, K.; Brix, S.; Brandt, A.; Brenke, N.; Enderlein, P.; Griffiths, H.J.; Kaiser, S.; Svavarsson, J.; Lörz, A.-N.; Frutos, I.; et al. Pan-Atlantic Comparison of Deep-Sea Macro-and Megabenthos. Diversity 2023, 15, 814. Abstract: Deep-sea benthic fauna is vital for a well-functioning marine ecosystem but is increasingly under threat from a changing environment. To monitor and conserve this fauna, an understanding of their large-scale spatial and bathymetric distribution and their environmental drivers is necessary. In this study, we conduct a multivariate analysis on abundance benthic fauna data collected at the phylum and multitaxon levels using an epibenthic sledge (EBS) across the Atlantic, and identify the environmental factors that affect such data. Our findings show a decrease in abundance with depth in most of the Atlantic but find relatively heterogeneous abundances with depth within the Southern Ocean. Principal component analyses indicate differences in environmental conditions south of the Antarctic Polar Front (~52 • S), outlining contrasts in the quantities of macronutrients and physical factors. Despite this, community composition seemed markedly similar throughout the Atlantic with the Antarctic Circumpolar Current seemingly not affecting benthic community composition for higher taxonomic levels. Those differences that did occur were largely caused by benthic chlorophyll, benthic iron, and surface silicate through a Bio-ENV. Overall, we argue that further large-scale spatial and bathymetric distribution studies are important amid environmental changes that are driving shifts in benthic community abundance and composition.
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... In contrast, other studies found no common effect of depth on species richness. For example, bivalves showed no clear relationship with depth, polychaete richness decreased with depth and isopods had their highest richness in middle abyssal depth (around 3000 m) [22], indicating that depth distribution patterns might be taxon-specific. In addition, earlier studies showed that marine biodiversity typically peaks at continental slope depths around 2000 m [22]. ...
... For example, bivalves showed no clear relationship with depth, polychaete richness decreased with depth and isopods had their highest richness in middle abyssal depth (around 3000 m) [22], indicating that depth distribution patterns might be taxon-specific. In addition, earlier studies showed that marine biodiversity typically peaks at continental slope depths around 2000 m [22]. ...
Distribution and Species Richness of Benthic Polychaeta and Sipuncula in the Northwestern Pacific
Article
Full-text available
  • Apr 2023
  • Diversity
Polychaeta and Sipuncula are abundant inhabitants of benthic marine habitats and have been increasingly sampled in the Northwest Pacific (NWP). However, polychaete and sipunculan species richness, composition, and distribution patterns still require further investigation, despite previous studies due to increasing deep-sea data flow. Using occurrence records for Polychaeta and Sipuncula from the Ocean Biodiversity Information System (OBIS) and the Global Biodiversity Information Facility (GBIF), we analyzed sampling effort (the number of distribution records), alpha (the number of species per 700,000 km 2 hexagon cells) and gamma (the number of species per 5 • latitudinal band) species diversity, and estimated species richness along latitudinal and bathymetric gradients. The species richness estimations were also correlated with multiple environmental and topographic variables, including depth, temperature, dissolved oxygen, chlorophyll, primary production , phytoplankton, current velocity, light, iron, nitrate, phosphate, silicate, and salinity. The dataset included over 30,000 distribution records belonging to polychaete (31,114 records, 98%) and sipunculan (690 records, 2%) species. Half of the distribution records were reported at a species level. The area around the island of Hainan and South Korea showed the highest alpha species richness (the number of species per 700,000 km 2 hexagon cell), yet the estimated species richness (ES50) indicated that there might be many unknown, unsampled, or non-digitized species throughout the whole NWP. Correspondingly, most distribution records (sampling effort) and gamma species richness were found between latitudes 20 and 40 • and decreased towards higher latitudes. Sipuncula were reported relatively more frequently from the deep sea than Polychaeta (62.8% vs. 12%). Overall, the number of species and records decreased with increasing depth, with a peak at about 5000 m. The alpha species richness had the strongest positive correlations with temperature, chlorophyll, primary production, and phytoplankton concentration. Here, we provide an overview of the species richness and distribution of Polychaeta in comparison with Sipuncula in the NWP, in both shallow and deep environments. This study demonstrates where further sampling efforts are needed to fill our knowledge gaps on annelids' distribution and diversity along the NWP. This could improve the analyses of the distribution and diversity of annelids to better understand the current environmental drivers of biodiversity, as well as predicting potential future drivers. The outcome of the environmental correlation provides thus valuable knowledge for predicting the future impacts of global warming on potential distribution shifts of annelids into new environments such as the Arctic Sea, possibly resulting in biological invasions.
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... Like deep-sea areas worldwide, few families and genera dominate Arctic and Antarctic meiofauna (Table 9.3). Species diversity within some nematode genera was studied on Antarctic shelves and slopes Fonseca et al. 2006;Ingels et al. 2006;Vermeeren et al. 2004), reporting high local and regional species diversity (Brandt et al. 2007). The highest number of co-existing species within the same genus was found for Acantholaimus, with 29 species recognized in only one sample (Brandt et al. 2007;De Mesel et al. 2006). ...
... Species diversity within some nematode genera was studied on Antarctic shelves and slopes Fonseca et al. 2006;Ingels et al. 2006;Vermeeren et al. 2004), reporting high local and regional species diversity (Brandt et al. 2007). The highest number of co-existing species within the same genus was found for Acantholaimus, with 29 species recognized in only one sample (Brandt et al. 2007;De Mesel et al. 2006). ...
Polar Meiofauna—Antipoles or Parallels?
Chapter
Full-text available
  • Mar 2023
At opposite ends of our world lie the poles. In the North, the Arctic, an ocean surrounded by coasts; in the South, the Antarctic continent surrounded by an ocean that separates it from the nearest landmasses. At first glance, the poles could not be more dissimilar owing to their contrasting location, geography, and tectonic and evolutionary history. The amplitude and types of ice cover, though differing between the poles, are influenced by the same climatic, atmospheric, and hydrodynamic processes that affect the entire Earth. Freshwater influx into their coastal areas too—beyond the effects of glaciological changes and dynamics such as glacier melt and increasing meltwater discharges—is different: in contrast to the Arctic, the Antarctic continent and sub-Antarctic islands lack major rivers. However, their latitudinal range and low temperatures, ice shelves, icebergs, sea ice, impacts from tidewater and land-based glaciers, significant seasonal variation in light intensity and, hence, primary productivity, offer parallel environments for organisms that have adapted to such conditions. Although we know much about the similarities and differences from an environmental perspective, there are still many unknowns about how benthic communities, especially the meiobenthos, from both regions compare. In this chapter, we provide an overview of the contrasts and parallels between Arctic and Antarctic meiobenthos and place it into context of their extreme habitats. Following a brief account of Arctic and Antarctic evolution and the historical study of their faunas, we (i) compare how extreme polar conditions affect meiofauna across four main habitats: polar coastal areas and fjords, continental shelves and ice shelves, the deep sea, and sea ice, and we (ii) discuss the implications of climate change on meiofauna in these habitats. Reflecting on (i) and (ii) allowed us to identify frontiers for future research of polar meiofauna, which we put forward in the concluding sections of this chapter.
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... Like deep-sea areas worldwide, few families and genera dominate Arctic and Antarctic meiofauna (Table 9.3). Species diversity within some nematode genera was studied on Antarctic shelves and slopes Fonseca et al. 2006;, reporting high local and regional species diversity (Brandt et al. 2007). The highest number of co-existing species within the same genus was found for Acantholaimus, with 29 species recognized in only one sample (Brandt et al. 2007;. ...
... Species diversity within some nematode genera was studied on Antarctic shelves and slopes Fonseca et al. 2006;, reporting high local and regional species diversity (Brandt et al. 2007). The highest number of co-existing species within the same genus was found for Acantholaimus, with 29 species recognized in only one sample (Brandt et al. 2007;. ...
Freshwater Meiofauna—A Biota with Different Rules?
Chapter
  • Mar 2023
Great divergences arise when comparing the ecology of meiofauna in freshwater and marine ecosystems. Emphasizing the main differences between freshwater meiofauna and their marine counterparts, we will go on a stepwise journey through three major frontiers in freshwater research, which in turn are hierarchically interrelated: biodiversity, community organization (e.g. food webs structure), and ecosystem processes (e.g. metabolism and organic carbon breakdown). The starting point of this chapter is one of the utmost frontiers, both in marine and freshwater research: meiofaunal diversity. Especially in freshwater ecosystems diversity becomes evident since, here, habitats extend as highly disconnected biotopes, each characterized by an often fundamentally different biocenosis. From the biodiversity level, we move up the theoretical hierarchy to assess the role of meiofauna as an integral part of benthic food webs. Recent research underlines the role of freshwater meiofauna as highly connected nodes and shows their pivotal role in the transfer of energy and carbon along food chains. Distributed over all trophic levels, this structure contrasts with the prevailing conception of meiofauna in food webs, where meiofauna often are considered rather marginal units. Finally, we apply allometric principles from the metabolic theory of ecology in order to assess the role of freshwater meiofauna in the functioning of the benthic systems. With a novel modelling framework we develop an analytical perspective, showing that secondary production of micro- and meiobenthic communities can predict microbial decomposition rates within the benthic interface. Our results demonstrate that productive micro- and meiobenthos act as catalysers in the system of organic carbon breakdown and recycling. These findings underline the relevance of freshwater meiofauna within the biogeochemical carbon cycle. The mechanistic forces behind the processes involved require future experimental research.
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... Adaptations such as increased mitochondrial concentration and adoption of enzymes more efficiently at low temperatures have been documented (Clarke, 1998). The importance of temperature as a structurer of benthic communities in the deep sea can be seen in the distribution pattern observed in regions such as Antarctica where the bathymetric gradient does not imply a significant temperature gradient and it is very common to record eurybathic species (Brey et al., 1996;Brandt et al., 2007) in such a way that the typical zonation of temperate or tropical regions is not usually recorded. Also, oxygen (Levin and Sibuet, 2012) and the organic matter availability (Cosson et al., 1997;Mamouridis et al., 2011;Bernardino et al., 2016;Brandt et al., 2018;Guggolz et al., 2018) has also been shown to be important factors for the distribution of deep-sea benthic communities. ...
... Brandt et al., 2019). Specially the very diverse and widely deep sea distributed Asellota suborder (Wilson, 2008) which can represent up to 97% of the isopods collected in this environment (Brandt et al., 2007). We recorded a relative isopods abundance lower than tanaidaceans and amphipods (4% in SOGOM 3 and 3% in SOGOM 4). ...
Macrofauna abundance and diversity patterns of deep sea southwestern Gulf of Mexico
Article
Full-text available
  • Jan 2023
Macrofauna abundance and diversity patterns in deep-sea waters of the southwestern Gulf of Mexico
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... The Southern Ocean harbours rich benthic assemblages with high levels of biodiversity and endemism (Clarke and Crame, 1992;Gage, 2004;Brandt et al., 2007). This is due to the high environmental stability, low disturbance and the relative isolation of the deep-sea in Antarctic waters (Grebmeier and Barry, 1991). ...
A new deep-sea species of golden gorgonian (Octocorallia: Scleralcyonacea: Chrysogorgiidae) from Antarctic waters
Article
Full-text available
  • Feb 2024
The Southern Ocean harbours rich deep-sea ecosystems with local hotspots of benthic biodiversity. Still, many species, including deep-water octocorals, remain undescribed despite the fact that the exploration of the deep-sea has improved thanks to recent technological advances. In this context, a new species of golden gorgonian has been collected at 1407–1581 m depth during a recent cruise to Dronning Maud Land in the Eastern Weddell Sea. Currently, after recent changes in Chrysogorgia taxonomy, Chrysogorgia lunae sp. nov. (Octocorallia: Scleralcyonacea) is the only known representative of the genus in Antarctic waters. It corresponds to the "Squamosae typicae" and is characterized by a bushy colony with branching sequence of 2/5L and bell-shaped polyps. A molecular comparison with published sequences of Chrysogorgia species is provided based on a concatenated sequence of two mitochondrial genes, mtMutS and COI, as well as the 28S nuclear ribosomal gene.
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... Most publications on deep-sea biodiversity highlight a limited understanding and the need for further studies, (e.g. Etter et al. 1999;Brandt et al. 2007;Baker et al. 2010;German et al. 2011;Ramirez-Llodra 2020). Furthermore, canyon topography influences current patterns and local upwelling, pumping nutrients into the euphotic zone which stimulates primary productivity (Fernandez-Arcaya et al. 2016 and references therein). ...
Metals and Metalloids
Chapter
Full-text available
  • May 2023
This chapter introduces you to metals and metalloids that are a concern to the health of marine ecosystems. It provides a general chemical understanding of important metals and metalloids, their sources, behaviour, impacts and management. Metals, metalloids and non-metals all make up the periodic table (Appendix II) and are classified into these categories according to their properties. Metals are good conductors of heat and electricity and are malleable and ductile, making them very useful to humans and therefore economically valuable. Metalloids sit on the periodic table in a jagged line at the division between metals and non-metals and have intermediate properties.
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Taxonomy and Biogeography of Bivalves of the Genus Cuspidaria Nardo, 1840, from the Southern Southwestern Atlantic Deep Sea
Article
  • Dec 2022
  • MALACOLOGIA
In this paper, a taxonomic review of the bivalves of the genus Cuspidaria Nardo, 1840 from the southern southwestern Atlantic is conducted. Specimens deposited in malacological collections and samples collected onboard the R/V Puerto Deseado off Mar del Plata (36S) and MPA Namuncur/Burdwood Bank area (54S), between 200 and 3,000 m depth, are the focus of this revision. The specimens were analyzed through conchological and anatomical features. The geographic and bathymetric distributions for each species are provided and possible factors determining biogeographic patterns are discussed. As a result, Cuspidaria infirma n. sp., Cuspidaria cancellata n. sp., Cuspidaria namuncura n. sp., and Cuspidaria cf. kerguelensis (Smith, 1885), are described. In addition, Cuspidaria exigua (Jeffreys, 1876), Cuspidaria bicarinata Jeffreys, 1882, Cuspidaria platensis (Smith, 1885), Cuspidaria tenella Smith, 1907, Cuspidaria infelix Thiele, 1912, and Cuspidaria barnardi Knudsen, 1970 are redescribed after the study of new specimens. Elliptic Fourier analyses were performed for each side of the shell to delimit species objectively using their shape. Results showed a clear differentiation on both valves among species. Cuspidaria bicarinata and Cuspidaria exigua, both North Atlantic species, were recorded for the first time in the southwestern Atlantic, and Cuspidaria infelix and Cuspidaria tenella, both Antarctic/sub-Antarctic species, expanded their distribution northwards. Two cluster analyses, for species and areas respectively, revealed a vertical zonation, separating species into two different groups highly corresponding to deep-sea water mass distributions.
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Meiofauna and Nematode Community Composition in Maxwell Bay, King George Island, Antarctica
Article
  • Nov 2022
The Antarctic Ocean hosts several species adapted to polar environments. However, the Antarctic marine environment is rapidly changing. We studied the meiofauna and nematode communities in Maxwell Bay, King George Island, Antarctica. The meiofaunal densities ranged from 174.3 ± 6.2 to 377.9 ± 7.0 individuals/10 cm2. The nematodes and harpacticoids accounted for 89.7% of the total meiofaunal density. A total of 1448 nematode individuals across 34 genera were recorded. The dominant nematode genus was Molgolaimus. This study offers guidance for future studies on polar benthic organisms and a reference to evaluate anthropogenic effects on polar ecosystems.
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Biodiversity and ecology of benthic foraminiferans from the Antarctic deep sea
Thesis
  • Jan 2005
p>Live foraminferal assemblages (i.e. including soft-walled species) and the metazoan fauna were analysed using replicate multicorer subcores (3.45 cm<sup>2</sup> surface area, 0-1 cm layer; >63 μm fraction for meiofaunal forms) and boxcorer subcores (100 cm<sup>2</sup> surface area, 0-5 cm layer; >300 μm fraction for macrofaunal forms) from stations along a transect (1100 - 4975 m water depth) east of the Antarctic Peninsula and a station in the S. Sandwich Trench (6,300 m water depth). Wet-sorting revealed abundant and diverse meio- and macrofaunal assemblages. Foraminiferans usually accounted for 43-83% of all meiofaunal and 65-96% of macrofaunal organisms. Many faunal trends were related to water depth and distance from land, representing a decrease in organic carbon inputs. Foraminiferan and metazoan densities were broadly coherent across the depth range sampled. Abundance generally decreased with water depth, as did the proportion of calcareous individuals, although there was considerable variability between replicate multicorer subcores at some stations. The location of the Carbonate Compensation Depth at about 3000 m did not seem to limit the bathymetric distribution of live individuals of certain calcareous species. Macrofaunal foraminiferans were concentrated in the surface 1 cm of sediment and sediment penetration increased with water depth. A total of 205 live foraminiferal species were recognized, of which only 93 belonged to described taxa. The species diversity of both meio- and macrofaunal assemblages exhibited a unimodal pattern with depth and was highest on the lower slope. Monothalamous taxa increased and calcareous taxa decreased in relative abundance with increasing depth, especially in the meiofaunal fraction. A large proportion (2.5 - 73.3%) of the meiofaunal assemblages were hidden within phytodetrital aggregates; species such as Epistominella exigua, Alabaminella weddellensis and Tinogullmia riemanni were concentrated within these microhabitats. The phytodetrital assemblages are strikingly similar to those reported from abyssal sites in the North Atlantic.</p
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New species of Tanaella Norman and Stebbing, 1886 (Crustacea: Tanaidacea: Tanaellidae) from the deep-sea off the Antarctic and the Angola Basin, with a key to the genus
Article
  • Mar 2004
Three new deep-sea species in the genus Tanaella are described: two from the Antarctic (T. eltaninae sp. nov., T. kimi sp. nov.) and one from the Angola Basin (T. profunda sp. nov.). This is the first record of Tanaella in the deep-sea of the Antarctic and the southern Atlantic Ocean. A key to the 13 known species of the genus is provided.
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The historical component of marine taxonomic diversity gradients
Chapter
  • Dec 1997
The biodiversity of many ecosystems is under threat and although seas cover the majority of our planet's surface, far less is known about the biodiversity of marine environments than that of terrestrial systems. It is also not clear whether many of the patterns known to occur on land also occur in the sea. Until we have a firmer idea of the diversity of a wide range of marine habitats and what controls it, we have little hope of conserving biodiversity, or determining the impact of human activities such as mariculture, fishing, dumping of waste and pollution. This book brings together key studies from the deep sea and open ocean, to tropical shores and polar regions to consider how comparable the patterns and processes underlying diversity are in these different ecosystems. Marine Biodiversity will be a major resource for all those interested in biodiversity and its conservation.
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Large-scale patterns of species diversity in the deep-sea benthos
Chapter
  • Dec 1997
The biodiversity of many ecosystems is under threat and although seas cover the majority of our planet's surface, far less is known about the biodiversity of marine environments than that of terrestrial systems. It is also not clear whether many of the patterns known to occur on land also occur in the sea. Until we have a firmer idea of the diversity of a wide range of marine habitats and what controls it, we have little hope of conserving biodiversity, or determining the impact of human activities such as mariculture, fishing, dumping of waste and pollution. This book brings together key studies from the deep sea and open ocean, to tropical shores and polar regions to consider how comparable the patterns and processes underlying diversity are in these different ecosystems. Marine Biodiversity will be a major resource for all those interested in biodiversity and its conservation.
View
High benthic species diversity in deep-sea sediments: The importance of hydrodynamics
Chapter
  • Dec 1997
The biodiversity of many ecosystems is under threat and although seas cover the majority of our planet's surface, far less is known about the biodiversity of marine environments than that of terrestrial systems. It is also not clear whether many of the patterns known to occur on land also occur in the sea. Until we have a firmer idea of the diversity of a wide range of marine habitats and what controls it, we have little hope of conserving biodiversity, or determining the impact of human activities such as mariculture, fishing, dumping of waste and pollution. This book brings together key studies from the deep sea and open ocean, to tropical shores and polar regions to consider how comparable the patterns and processes underlying diversity are in these different ecosystems. Marine Biodiversity will be a major resource for all those interested in biodiversity and its conservation.
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Ecological Studies in the Antarctic Sea Ice Zone: Results of EASIZ Midterm Symposium
Book
  • Jan 2002
Until comparatively recently, the remoteness, inaccessibility, and extreme climate have meant that the vast pack-ice zone around the Antarctic continent was one of the least-known marine ecosystems on Earth. Myths and speculations prevailed in the literature, often derived from an anthropocentric way of thinking that considered the sea-ice environment as predominantly hostile to marine life. This picture has changed drastically now as a result of a series of international efforts, the most recent of which has been the highly successful EASIZ (Ecology of the Antarctic Sea Ice Zone) programme of the Scientific Council on Antarctic Research (SCAR). Focusing, in contrast to other international programmes, on life at the seafloor, EASIZ has attempted to link processes in the three major marine subsystems (sea ice, pelagic and benthic) within the pack-ice zone. Work has been carried out from both research ships and shore-based research stations. This work included organisms ranging in size from bacteria to seals and covered topics as diverse as biodiversity, iceberg scour, pelagobenthic coupling, autecology, and ecophysiology. Consequently, we now view the sea-ice zone as a rich system with highly adapted organisms, considerable natural disturbance from ice, low resilience and of great potential importance as an indicator for climate change.
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