Richness, growth, and persistence of life under an Antarctic ice shelf.

Where polar ice sheets meet the coast, they can flow into the sea as floating ice shelves. The seabed underneath is in complete darkness, and may be Earth's least known surface habitat. Few taxa there have been fully identified to named species (see Supplemental information) - remarkable for a habitat spanning nearly 1.6 million km2. Glimpses of life there have come from cameras dropped through 10 boreholes, mainly at the three largest Antarctic ice shelves - the Ross (McMurdo), Filchner-Ronne and Amery. Pioneering studies of life under boreholes found distinct morphotypes of perhaps >50 species. Here, we report remarkable growth and persistence over thousands of years of benthic faunal species collected in 2018 from the seabed under the Ekström Ice Shelf (EIS), Weddell Sea.

Antarctic ecosystems in transition - life between stresses and opportunities.

Important findings from the second decade of the 21st century on the impact of environmental change on biological processes in the Antarctic were synthesised by 26 international experts. Ten key messages emerged that have stakeholder-relevance and/or a high impact for the scientific community. They address (i) altered biogeochemical cycles, (ii) ocean acidification, (iii) climate change hotspots, (iv) unexpected dynamism in seabed-dwelling populations, (v) spatial range shifts, (vi) adaptation and thermal resilience, (vii) sea ice related biological fluctuations, (viii) pollution, (ix) endangered terrestrial endemism and (x) the discovery of unknown habitats. Most Antarctic biotas are exposed to multiple stresses and considered vulnerable to environmental change due to narrow tolerance ranges, rapid change, projected circumpolar impacts, low potential for timely genetic adaptation, and migration barriers. Important ecosystem functions, such as primary production and energy transfer between trophic levels, have already changed, and biodiversity patterns have shifted. A confidence assessment of the degree of 'scientific understanding' revealed an intermediate level for most of the more detailed sub-messages, indicating that process-oriented research has been successful in the past decade. Additional efforts are necessary, however, to achieve the level of robustness in scientific knowledge that is required to inform protection measures of the unique Antarctic terrestrial and marine ecosystems, and their contributions to global biodiversity and ecosystem services.

Do drivers of biodiversity change differ in importance across marine and terrestrial systems - Or is it just different research communities' perspectives?

Cross-system studies on the response of different ecosystems to global change will support our understanding of ecological changes. Synoptic views on the planet's two main realms, the marine and terrestrial, however, are rare, owing to the development of rather disparate research communities. We combined questionnaires and a literature review to investigate how the importance of anthropogenic drivers of biodiversity change differs among marine and terrestrial systems and whether differences perceived by marine vs. terrestrial researchers are reflected by the scientific literature. This included asking marine and terrestrial researchers to rate the relevance of different drivers of global change for either marine or terrestrial biodiversity. Land use and the associated loss of natural habitats were rated as most important in the terrestrial realm, while the exploitation of the sea by fishing was rated as most important in the marine realm. The relevance of chemicals, climate change and the increasing atmospheric concentration of CO2 were rated differently for marine and terrestrial biodiversity respectively. Yet, our literature review provided less evidence for such differences leading to the conclusion that while the history of the use of land and sea differs, impacts of global change are likely to become increasingly similar.

Impacts of Climate Variability and Change on (Marine) Animals: Physiological Underpinnings and Evolutionary Consequences.

Understanding thermal ranges and limits of organisms becomes important in light of climate change and observed effects on ecosystems as reported by the IPCC (2014). Evolutionary adaptation to temperature is presently unable to keep animals and other organisms in place; if they can these rather follow the moving isotherms. These effects of climate change on aquatic and terrestrial ecosystems have brought into focus the mechanisms by which temperature and its oscillations shape the biogeography and survival of species. For animals, the integrative concept of oxygen and capacity limited thermal tolerance (OCLTT) has successfully characterized the sublethal limits to performance and the consequences of such limits for ecosystems. Recent models illustrate how routine energy demand defines the realized niche. Steady state temperature-dependent performance profiles thus trace the thermal window and indicate a key role for aerobic metabolism, and the resulting budget of available energy (power), in defining performance under routine conditions, from growth to exercise and reproduction. Differences in the performance and productivity of marine species across latitudes relate to changes in mitochondrial density, capacity, and other features of cellular design. Comparative studies indicate how and why such mechanisms underpinning OCLTT may have developed on evolutionary timescales in different climatic zones and contributed to shaping the functional characteristics and species richness of the respective fauna. A cause-and-effect understanding emerges from considering the relationships between fluctuations in body temperature, cellular design, and performance. Such principles may also have been involved in shaping the functional characteristics of survivors in mass extinction events during earth's history; furthermore, they may provide access to understanding the evolution of endothermy in mammals and birds. Accordingly, an understanding is emerging how climate changes and variability throughout earth's history have influenced animal evolution and co-defined their success or failure from a bio-energetic point of view. Deepening such understanding may further reduce uncertainty about projected impacts of anthropogenic climate variability and change on the distribution, productivity and last not least, survival of aquatic and terrestrial species.

Community dynamics of nematodes after Larsen ice-shelf collapse in the eastern Antarctic Peninsula.

Free-living marine nematode communities of the Larsen B embayment at the eastern Antarctic Peninsula were investigated to provide insights on their response and colonization rate after large-scale ice-shelf collapse. This study compares published data on the post-collapse situation from 2007 with new material from 2011, focusing on two locations in the embayment that showed highly divergent communities in 2007 and that are characterized by a difference in timing of ice-shelf breakup. Data from 2007 exposed a more diverse community at outer station B.South, dominated by the genus Microlaimus. On the contrary, station B.West in the inner part of Larsen B was poor in both numbers of individuals and genera, with dominance of a single Halomonhystera species. Re-assessment of the situation in 2011 showed that communities at both stations diverged even more, due to a drastic increase in Halomonhystera at B.West compared to relatively little change at B.South. On a broader geographical scale, it seems that B.South gradually starts resembling other Antarctic shelf communities, although the absence of the genus Sabatieria and the high abundance of Microlaimus still set it apart nine years after the main Larsen B collapse. In contrast, thriving of Halomonhystera at B.West further separates its community from other Antarctic shelf areas.

Different Oceanographic Regimes in the Vicinity of the Antarctic Peninsula Reflected in Benthic Nematode Communities.

Marine free-living nematode communities were studied at similar depths (~500 m) at two sides of the Antarctic Peninsula, characterised by different environmental and oceanographic conditions. At the Weddell Sea side, benthic communities are influenced by cold deep-water formation and seasonal sea-ice conditions, whereas the Drake Passage side experiences milder oceanic conditions and strong dynamics of the Antarctic Circumpolar Current. This resulted in different surface primary productivity, which contrasted with observed benthic pigment patterns and varied according to the area studied: chlorophyll a concentrations (as a proxy for primary production) were high in the Weddell Sea sediments, but low in the surface waters above; this pattern was reversed in the Drake Passage. Differences between areas were largely mirrored by the nematode communities: nematode densities peaked in Weddell stations and showed deeper vertical occurrence in the sediment, associated with deeper penetration of chlorophyll a and indicative of a strong bentho-pelagic coupling. Generic composition showed some similarities across both areas, though differences in the relative contribution of certain genera were noted, together with distinct community shifts with depth in the sediment at all locations.

The Southern Ocean ecosystem under multiple climate change stresses--an integrated circumpolar assessment.

A quantitative assessment of observed and projected environmental changes in the Southern Ocean (SO) with a potential impact on the marine ecosystem shows: (i) large proportions of the SO are and will be affected by one or more climate change processes; areas projected to be affected in the future are larger than areas that are already under environmental stress, (ii) areas affected by changes in sea-ice in the past and likely in the future are much larger than areas affected by ocean warming. The smallest areas (<1% area of the SO) are affected by glacier retreat and warming in the deeper euphotic layer. In the future, decrease in the sea-ice is expected to be widespread. Changes in iceberg impact resulting from further collapse of ice-shelves can potentially affect large parts of shelf and ephemerally in the off-shore regions. However, aragonite undersaturation (acidification) might become one of the biggest problems for the Antarctic marine ecosystem by affecting almost the entire SO. Direct and indirect impacts of various environmental changes to the three major habitats, sea-ice, pelagic and benthos and their biota are complex. The areas affected by environmental stressors range from 33% of the SO for a single stressor, 11% for two and 2% for three, to <1% for four and five overlapping factors. In the future, areas expected to be affected by 2 and 3 overlapping factors are equally large, including potential iceberg changes, and together cover almost 86% of the SO ecosystem.

Asteroids, ophiuroids and holothurians from the southeastern Weddell Sea (Southern Ocean).

Until the early 1980s, the composition and distribution of the asteroid (starfish), ophiuroid (brittle star) and holothurian (sea cucumber) bottom fauna of the southeastern Weddell Sea was virtually unknown. This southernmost part of the Atlantic sector of the Southern Ocean is a typical high-latitude Antarctic region located in the circumpolar permanent pack-ice zone. It became accessible for large-scale scientific surveys only through the availability of modern ice-breaking research vessels, such as the German RV "Polarstern". Here, we describe a dataset of the faunal composition and abundance of starfish, brittle star and sea cucumber assemblages in this area, based on collections from trawl catches carried out during three "Polarstern" cruises in 1983, 1984 and 1985. The set comprises a total of 4,509 records of abundances of 35 asteroid species (with a total of 2,089 specimens) and 38 ophiuroid species (with a total of 18,484 specimens) from 34 stations, as well as of 66 holothurian species (with a total of 20,918 specimens) from 59 stations including zero-abundances (absences). A synthesizing zoogeographical community analysis confirms the presence of three distinct assemblages of asteroids, ophiuroids, and holothurians with highest species richness on the eastern shelf. Overall, starfishes, brittle stars and sea cucumbers were present at all sites investigated in the study area but composition and abundance of asterozoan (asteroids and ophiuroids together) and holothurian fauna varied considerably. A synthesizing zoogeographical community analysis confirms the presence of three distinct assemblages of asteroids, ophiuroids, and holothurians with highest species richness on the eastern shelf. In the case of asterozoans, water depth and latitude seemed to be the most important drivers of assemblage distribution and composition. One of the holothurian assemblages was part of the rich macrozoobenthic community dominated by a diverse and abundant epifauna, mainly sponges and gorgonians. Another one was mainly composed of vagrant deposit-feeding species inhabiting a predominantly non-colonised substratum. In addition, a mixed holothurian assemblage was identified.

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota.

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

Understanding and protecting the world's biodiversity: the role and legacy of the SCAR programme "Evolution and Biodiversity in the Antarctic".

Current global changes are prompting scientists and governments to consider the risk of extinction of species inhabiting environments influenced by ice. Concerted, multidisciplinary, international programmes aimed at understanding life processes, evolution and adaptations in the Polar Regions will help to counteract such an event by protecting polar life and ecosystems. There is a long tradition of international scientific cooperation in Antarctica that provides a strong foundation for such approaches. While basic understanding is emerging, we still largely lack predictive biological models, and need to achieve further integration amongst biological and non-biological disciplines. The ongoing SCAR Science Research Programme, "Evolution and Biodiversity in the Antarctic (EBA)" has successfully carried out its crucial role of providing an overarching umbrella for SCAR research in Life Sciences. Now is the time for aiming to progress beyond this important role, and the Antarctic biology community is proposing two programmes, focussed on distinct but complementary aspects of polar biology and working across marine, freshwater and terrestrial environments: "State of the Antarctic Ecosystem (AntEco)", and "Antarctic Thresholds--Ecosystem Resilience and Adaptation (AnT-ERA)". These programmes are the legacy of EBA, and they are key to understanding and protect Antarctic biodiversity.

Semi-automated image analysis for the assessment of megafaunal densities at the Arctic deep-sea observatory HAUSGARTEN.

Megafauna play an important role in benthic ecosystem function and are sensitive indicators of environmental change. Non-invasive monitoring of benthic communities can be accomplished by seafloor imaging. However, manual quantification of megafauna in images is labor-intensive and therefore, this organism size class is often neglected in ecosystem studies. Automated image analysis has been proposed as a possible approach to such analysis, but the heterogeneity of megafaunal communities poses a non-trivial challenge for such automated techniques. Here, the potential of a generalized object detection architecture, referred to as iSIS (intelligent Screening of underwater Image Sequences), for the quantification of a heterogenous group of megafauna taxa is investigated. The iSIS system is tuned for a particular image sequence (i.e. a transect) using a small subset of the images, in which megafauna taxa positions were previously marked by an expert. To investigate the potential of iSIS and compare its results with those obtained from human experts, a group of eight different taxa from one camera transect of seafloor images taken at the Arctic deep-sea observatory HAUSGARTEN is used. The results show that inter- and intra-observer agreements of human experts exhibit considerable variation between the species, with a similar degree of variation apparent in the automatically derived results obtained by iSIS. Whilst some taxa (e. g. Bathycrinus stalks, Kolga hyalina, small white sea anemone) were well detected by iSIS (i. e. overall Sensitivity: 87%, overall Positive Predictive Value: 67%), some taxa such as the small sea cucumber Elpidia heckeri remain challenging, for both human observers and iSIS.

Characterisation of the nematode community of a low-activity cold seep in the recently ice-shelf free Larsen B area, Eastern Antarctic Peninsula.

BACKGROUND: Recent climate-induced ice-shelf disintegration in the Larsen A (1995) and B (2002) areas along the Eastern Antarctic Peninsula formed a unique opportunity to assess sub-ice-shelf benthic community structure and led to the discovery of unexplored habitats, including a low-activity methane seep beneath the former Larsen B ice shelf. Since both limited particle sedimentation under previously permanent ice coverage and reduced cold-seep activity are likely to influence benthic meiofauna communities, we characterised the nematode assemblage of this low-activity cold seep and compared it with other, now seasonally ice-free, Larsen A and B stations and other Antarctic shelf areas (Weddell Sea and Drake Passage), as well as cold-seep ecosystems world-wide. PRINCIPAL FINDINGS: The nematode community at the Larsen B seep site differed significantly from other Antarctic sites in terms of dominant genera, diversity and abundance. Densities in the seep samples were high (>2000 individuals per 10 cm(2)) and showed below-surface maxima at a sediment depth of 2-3 cm in three out of four replicates. All samples were dominated by one species of the family Monhysteridae, which was identified as a Halomonhystera species that comprised between 80 and 86% of the total community. The combination of high densities, deeper density maxima and dominance of one species is shared by many cold-seep ecosystems world-wide and suggested a possible dependence upon a chemosynthetic food source. Yet stable (13)C isotopic signals (ranging between -21.97±0.86‰ and -24.85±1.89‰) were indicative of a phytoplankton-derived food source. CONCLUSION: The recent ice-shelf collapse and enhanced food input from surface phytoplankton blooms were responsible for the shift from oligotrophic pre-collapse conditions to a phytodetritus-based community with high densities and low diversity. The parthenogenetic reproduction of the highly dominant Halomonhystera species is rather unusual for marine nematodes and may be responsible for the successful colonisation by this single species.