Nord‐ und Südpolarmeer im Klimawandel. Ein biologischer Vergleich

Impacts of climate change on polar seas The polar seas in the Arctic and Antarctic are characterized by extreme cold and the prevalence of sea ice, which provides a unique polar habitat but also strongly affects the pelagic and benthic biota beneath. Life conditions for the marine fauna and flora differ considerably between the Arctic and Southern Oceans, as a result of contrasts in geography, geological history, as well as seasonal dynamics in light regime, sea ice cover and, hence, biological production. Climate change is particularly obvious in the Arctic Ocean and off the Antarctic Peninsula where warming results in a rapid shrinkage of the summer sea ice cover. Such decline threatens the sea‐ice communities and their associated fauna and will also have far reaching effects for the plankton and benthos of the polar seas.     

Environmental constraints on life histories in Antarctic ecosystems: tempos, timings and predictability

Knowledge of Antarctic biotas and environments has increased dramatically in recent years. There has also been a rapid increase in the use of novel technologies. Despite this, some fundamental aspects of environmental control that structure physiological, ecological and life-history traits in Antarctic organisms have received little attention. Possibly the most important of these is the timing and availability of resources, and the way in which this dictates the tempo or pace of life. The clearest view of this effect comes from comparisons of species living in different habitats. Here, we (i) show that the timing and extent of resource availability, from nutrients to colonisable space, differ across Antarctic marine, intertidal and terrestrial habitats, and (ii) illustrate that these differences affect the rate at which organisms function. Consequently, there are many dramatic biological differences between organisms that live as little as 10 m apart, but have gaping voids between them ecologically.Identifying the effects of environmental timing and predictability requires detailed analysis in a wide context, where Antarctic terrestrial and marine ecosystems are at one extreme of the continuum of available environments for many characteristics including temperature, ice cover and seasonality. Anthropocentrically, Antarctica is harsh and as might be expected terrestrial animal and plant diversity and biomass are restricted. By contrast, Antarctic marine biotas are rich and diverse, and several phyla are represented at levels greater than global averages. There has been much debate on the relative importance of various physical factors that structure the characteristics of Antarctic biotas. This is especially so for temperature and seasonality, and their effects on physiology, life history and biodiversity. More recently, habitat age and persistence through previous ice maxima have been identified as key factors dictating biodiversity and endemism. Modern molecular methods have also recently been incorporated into many traditional areas of polar biology. Environmental predictability dictates many of the biological characters seen in all of these areas of Antarctic research.     

Diel vertical migration of Arctic zooplankton during the polar night

High-latitude environments show extreme seasonal variation in physical and biological variables. The classic paradigm of Arctic marine ecosystems holds that most biological processes slow down or cease during the polar night. One key process that is generally assumed to cease during winter is diel vertical migration (DVM) of zooplankton. DVM constitutes the largest synchronized movement of biomass on the planet, and is of paramount importance for marine ecosystem function and carbon cycling. Here we present acoustic data that demonstrate a synchronized DVM behaviour of zooplankton that continues throughout the Arctic winter, in both open and ice-covered waters. We argue that even during the polar night, DVM is regulated by diel variations in solar and lunar illumination, which are at intensities far below the threshold of human perception. We also demonstrate that winter DVM is stronger in open waters compared with ice-covered waters. This suggests that the biologically mediated vertical flux of carbon will increase if there is a continued retreat of the Arctic winter sea ice cover.     

On the direct impact of ice on marine benthic communities, a review

Ice has a significant impact on the polar and sub-polar benthos, but relationships between corresponding physical and biological processes are not yet sufficiently understood. Sea ice contributes to a vertical zonation in shallow waters, which also experience other important disturbances. Due to the length of the non-glaciated coastline, sea ice is of greater relevance in the north than in the south. Scouring by icebergs and ridged sea ice causes an increased diversity when different recolonisation stages coexist. Frequently scoured areas do not recover, especially in the Antarctic, due to slow growth rates of the fauna. Iceberg grounding in the Arctic is mainly restricted to the western Eurasian and northeastern American shelf, including Greenland. Around Antarctica, scouring is more evenly distributed. Glacier termini prevent sessile animals from settling in their proximity where only few motile species occur.     

Marine Ecology of Seabirds in Polar Oceans

Patterns of seabird species' distributions differ between theAntarctic and the Arctic. In the Antarctic, distributions areannular or latitudinal, with strong similarities in speciescomposition of seabird communities in all ocean basins at agiven latitude. In the Arctic, communities are arranged meridionally,and show strong differences between ocean basins and, at a givenlatitude, between sides of ocean basins. These differences betweenthe seabird communi ies in the Northern Hemisphere and the SouthernHemisphere reflect differences in the patterns of flow of majorocean current systems. At smaller spatial scales, in both hemispheresthe species composition of seabird communities is sensitiveto changes in watermass characteristics. The distribution of avian biomass is affected by both physicaland biological features of the ocean. In the Antarctic, muchseabird foraging is over deep water, and withinseason, small-scalepatchiness in prey abundance and availability in ice-free watersis likely to be controlledprimarily by the behavior of the prey,rather than by physical features. Thus, prey availability maybe unpredictable in time and space. In contrast, in the NorthernHemisphere, most seabirdforaging is concentrated over shallowcontinental shelves, where currents interact with bathymetryto produce predictable physical features capable of concentratingprey or making prey more easily harvested by seabirds. Ice cover appears to be the most important physical featurein the Antarctic. An entire community of birds is specializedto use prey taken near the ice edge. These prey consist of avariety of species, some of which are normally found much deeperin the water than the birds takingthem can dive. The open-waterportion of the marginal ice zone is also an important foraginghabitat for Antarctic marine birds. In the Arctic, a food webbased on underice algae is used by marine birds, but few ifany data exist on avian use of the open water segment of themarginal ice zone. Recent simultaneous surveys of birds and their prey indicatethat only rarely does the small-scale abundance of birds matchthat of their prey; correlations between predators and preyaregenerally stronger at larger scales. Evidence is accumulatingin the Antarctic that the largestaggregations of krill may bedisproportionately important to foraging seabirds.     

北极浮冰生态学研究进展

近年来随着北极地区的开放和全球变化对北极地区生态环境和海冰现存量的影响日益显现,北极浮冰生态学研究得到了广泛的重视和实质性的进展.最新研究结果显示,浮冰本身包含了一个复杂的生物群落,高纬度浮冰生物群落的初级产量远高于原先的估算,浮冰生物群落在北极海洋生态系统中的作用被进一步确认.但由于对浮冰生物群落的研究受后勤保障条件的制约,目前尚有大量科学问题有待今后进一步深入研究,预期我国科学家将在其中做出贡献.     

Animal survival strategies in Neoproterozoic ice worlds

The timing of the first appearance of animals is of crucial importance for understanding the evolution of life on Earth. Although the fossil record places the earliest metazoans at 572–602 Ma, molecular clock studies suggest a far earlier origination, as far back as ~850 Ma. The difference in these dates would place the rise of animal life into a time period punctuated by multiple colossal, potentially global, glacial events. Although the two schools of thought debate the limitations of each other's methods, little time has been dedicated to how animal life might have survived if it did arise before or during these global glacial periods. The history of recent polar biota shows that organisms have found ways of persisting on and around the ice of the Antarctic continent throughout the Last Glacial Maximum (33–14 Ka), with some endemic species present before the breakup of Gondwana (180–23 Ma). Here we discuss the survival strategies and habitats of modern polar marine organisms in environments analogous to those that could have existed during Neoproterozoic glaciations. We discuss how, despite the apparent harshness of many ice covered, sub-zero, Antarctic marine habitats, animal life thrives on, in and under the ice. Ice dominated systems and processes make some local environments more habitable through water circulation, oxygenation, terrigenous nutrient input and novel habitats. We consider how the physical conditions of Neoproterozoic glaciations would likely have dramatically impacted conditions for potential life in the shallows and erased any possible fossil evidence from the continental shelves. The recent glacial cycle has driven the evolution of Antarctica's unique fauna by acting as a “diversity pump,” and the same could be true for the late Proterozoic and the evolution of animal life on Earth, and the existence of life elsewhere in the universe on icy worlds or moons.     

Metagenomic analysis of stress genes in microbial mat communities from Antarctica and the High Arctic

Polar and alpine microbial communities experience a variety of environmental stresses, including perennial cold and freezing; however, knowledge of genomic responses to such conditions is still rudimentary. We analyzed the metagenomes of cyanobacterial mats from Arctic and Antarctic ice shelves, using high-throughput pyrosequencing to test the hypotheses that consortia from these extreme polar habitats were similar in terms of major phyla and subphyla and consequently in their potential responses to environmental stresses. Statistical comparisons of the protein-coding genes showed similarities between the mats from the two poles, with the majority of genes derived from Proteobacteria and Cyanobacteria; however, the relative proportions differed, with cyanobacterial genes more prevalent in the Antarctic mat metagenome. Other differences included a higher representation of Actinobacteria and Alphaproteobacteria in the Arctic metagenomes, which may reflect the greater access to diasporas from both adjacent ice-free lands and the open ocean. Genes coding for functional responses to environmental stress (exopolysaccharides, cold shock proteins, and membrane modifications) were found in all of the metagenomes. However, in keeping with the greater exposure of the Arctic to long-range pollutants, sequences assigned to copper homeostasis genes were statistically (30%) more abundant in the Arctic samples. In contrast, more reads matching the sigma B genes were identified in the Antarctic mat, likely reflecting the more severe osmotic stress during freeze-up of the Antarctic ponds. This study underscores the presence of diverse mechanisms of adaptation to cold and other stresses in polar mats, consistent with the proportional representation of major bacterial groups.     

Retention of ice-associated amphipods: possible consequences for an ice-free Arctic Ocean

Recent studies predict that the Arctic Ocean will have ice-free summers within the next 30 years. This poses a significant challenge for the marine organisms associated with the Arctic sea ice, such as marine mammals and, not least, the ice-associated crustaceans generally considered to spend their entire life on the underside of the Arctic sea ice. Based upon unique samples collected within the Arctic Ocean during the polar night, we provide a new conceptual understanding of an intimate connection between these under-ice crustaceans and the deep Arctic Ocean currents. We suggest that downwards vertical migrations, followed by polewards transport in deep ocean currents, are an adaptive trait of ice fauna that both increases survival during ice-free periods of the year and enables re-colonization of sea ice when they ascend within the Arctic Ocean. From an evolutionary perspective, this may have been an adaptation allowing success in a seasonally ice-covered Arctic. Our findings may ultimately change the perception of ice fauna as a biota imminently threatened by the predicted disappearance of perennial sea ice.     

Psychrophilic versus psychrotolerant bacteria--occurrence and significance in polar and temperate marine habitats.

The numerical dominance and ecological role of psychrophilic bacteria in bottom sediments, sea ice, surface water and melt pools of the polar oceans were investigated using isolates, colony forming units (CFU) and metabolic activities. All sediment samples of the Southern Ocean studied showed a clear numerical dominance of cold-loving bacteria. In Arctic sediments underlying the influence of cold polar water bodies psychrophiles prevailed also but they were less dominant in sediments influenced by the warm Atlantic Water. A predominance of psychrophiles was further found in consolidated Antarctic sea ice as well as in multiyear Arctic sea ice and in melt pools on top of Arctic ice floes. A less uniform adaptation response was, however, met in polar surface waters. In the very northern part of the Fram Strait (Arctic Ocean) we found bacterial counts and activities at 1 degree C exceeding those at 22 degrees C. In surface water of the Weddell Sea (Southern Ocean) psychrophiles also dominated numerically in early autumn but the dominance declined obviously with the onset of winter-water and a decrease of chlorphyll a. Otherwise in surface water of the Southern Ocean CFUs were higher at 22 degrees C than at 1 degree C while activities were vice versa indicating at least a functional dominance of psychrophiles. Even in the temperate sediments of the German Bight true psychrophiles were present and a clear shift towards cold adapted communities in winter observed. Among the polar bacteria a more pronounced cold adaptation of Antarctic in comparison with Arctic isolates was obtained. The results and literature data indicate that stenothermic cold adapted bacteria play a significant role in the global marine environment. On the basis of the temperature response of our isolates from different habitats it is suggested to expand the definition of Morita in order to meet the cold adaptation strategies of the bacteria in the various cold habitats.     

United States legislation and the polar oceans

The United States has important national interests vested in both the Arctic and Antarctic Oceans. Thus, in recent decades the United States has progressively codified its national commitment to conserve and manage both these marine regions and their resources and to protect them from activities that might produce adverse impacts. The U.S. legislation passed since the 1960s selectively affects both polar regions and supports major American policy objectives there: to maintain the Arctic and Antarctic as areas of international cooperation for peaceful purposes; to satisfy economic needs, especially hydrocarbon, mineral, and living resources; to protect opportunities for scientific research; to protect the marine environment; and to conserve living resources in the circumpolar seas. Future trends suggest a shift in U.S. policy attention to the Arctic. Accordingly, new U.S. legislation will be needed to regulate increased activities in the polar north.     

Likely responses of the Antarctic benthos to climate-related changes in physical disturbance during the 21st century, based primarily on evidence from the West Antarctic Peninsula region

Disturbance has always shaped the evolution and ecology of organisms and nowhere is this more apparent that on the iceberg gouged continental shelves of the Antarctic Peninsula (AP). The vast majority of currently described polar biodiversity occurs on the Southern Ocean shelf but current and projected climate change is rapidly altering disturbance intensities in some regions. The AP is now amongst the fastest warming and changing regions on earth. Seasonal sea ice has decreased in time and extent, most glaciers in the region have retreated, a number of ice shelves have collapsed, and the surface waters of the seas west of the AP have warmed. Here, we review the influences of disturbance from ice, sedimentation, freshening events, wave action and humans on shallow water benthic assemblages, and suggest how disturbance pressures will change during the 21st century in the West Antarctic Peninsula (WAP) and Scotia Arc region. We suggest that the intensity of ice scouring will increase in the region over the next few decades as a result of decreased winter sea ice periods and increased ice loading into coastal waters. Thus, the most frequently disturbed environment on earth will become more so, which will lead to considerable changes in community structure and species distributions. However, as ice fronts retreat past their respective grounding lines, sedimentation and freshening events will become relatively more important. Human presence in the region is increasing, through research, tourism, and resource exploitation, which represents a considerable threat to polar biodiversity over the next century. Adapting to or tolerating multiple, changing environmental stressors will be difficult for a fauna with typically slow generation turnovers that has evolved largely in isolation. We suggest that intensifying acute and chronic disturbances are likely to cause significant changes in ecosystem structure, and probably a considerable loss of polar marine biodiversity, over relatively short timescales.     

The oxygen transport system in three species of the boreal fish family Gadidae. Molecular phylogeny of hemoglobin

The Arctic and Antarctic marine faunas differ by age and isolation. Fishes of the two polar regions have undergone different regional histories that have driven the physiological diversities. Antarctic fish are highly stenothermal, in keeping with stable water temperatures, whereas Arctic fish, being exposed to seasonal temperature variations, exhibit higher physiological plasticity. This study reports the characterization of the oxygen transport system of three Arctic species of the family Gadidae, namely the Arctic cod Arctogadus glacialis, the polar cod Boreogadus saida, and the Atlantic cod Gadus morhua. Unlike Antarctic notothenioids, the blood displays high multiplicity, i.e. it has three hemoglobins, similar to many other acanthomorph teleosts. In the most abundant hemoglobin, oxygen binding is modulated by heterotropic effectors, with marked Bohr and Root effects. Remarkably, in two species (A. glacialis and B. saida), the Hill coefficient is very close to one in the whole pH range, indicating the apparent absence of cooperativity. The amino acid sequences have been used to gain insight into the evolution history of globins of polar fish. The results indicate that Arctic and Antarctic globins have different phylogenies and lead us to suggest that the selective pressure of environment stability allows the phylogenetic signal to be maintained in the Antarctic sequences, whereas environmental variability would tend to disrupt this signal in the Gadidae sequences.     

Life strategy, ecophysiology and ecology of seaweeds in polar waters

Polar seaweeds are strongly adapted to the low temperatures of their environment, Antarctic species more strongly than Arctic species due to the longer cold water history of the Antarctic region. By reason of the strong isolation of the Southern Ocean the Antarctic marine flora is characterized by a high degree of endemism, whereas in the Arctic only few endemic species have been found so far. All polar species are strongly shade adapted and their phenology is finely tuned to the strong seasonal changes of the light conditions. The paper summarises the present knowledge of seaweeds from both polar regions with regard to the following topics: the history of seaweed research in polar regions; the environment of seaweeds in polar waters; biodiversity, biogeographical relationships and vertical distribution of Arctic and Antarctic seaweeds; life histories and physiological thallus anatomy; temperature demands and geographical distribution; light demands and depth zonation; the effect of salinity, temperature and desiccation on supra-and eulittoral seaweeds; seasonality of reproduction and the physiological characteristics of microscopic developmental stages; seasonal growth and photosynthesis; elemental and nutritional contents and chemical and physical defences against herbivory. We present evidence to show that specific characteristics and adaptations in polar seaweeds help to explain their ecological success under environmentally extreme conditions. In conclusion, as a perspective and guide for future research we draw attention to many remaining gaps in knowledge. Dedicated to Prof. Dr. Gunter O. Kirst and to Prof. Dr. Klaus Lüning on occasion of their retirement 28. Februar 2006 and 31. March 2006, respectively.     

Sea Ice-Upper Ocean Ecosystems and Global Changes in the Arctic

In the Arctic, a clear temperature trend toward warming has been observed during the last two decades. The warming has led to a reduction in the area and thickness of the Arctic sea ice cover because of increased melting. Melt water is accumulated within the 0- to 30-m surface layer, markedly changing its hydrological and hydrochemical characteristics: this water is warmer and fresher, with lower nutrient concentrations. A stable thermocline at depths of 30–35 m weakens vertical mixing of water and hampers active exchange between nutrient-enriched water lying below the thermocline and nutrient-poor water under ice. This affects the qualitative and quantitative composition of upper-ocean phyto- and zooplankton. The changes in the climate and the ice environment cause marked alterations in the composition and structure of the biological communities of sea ice and under-ice surface water. Comparative analysis of materials collected in the anticyclonic Arctic Gyre at the ice stations Severnyi Polyus-22 (1975– 1981) and SHEBA (1997–1998) shows that over the last two decades the number of species of ice diatoms markedly decreased and the role of freshwater algae increased. The number of nematodes, copepods, amphipods, and turbellarians, which were the dominant groups in perennial ice, also declined sharply. The results suggest that the sea ice-upper ocean ecosystem is changing from typically marine to brackish-water. The main reason for these changes is probably global warming in the Arctic.     

Arctic marine fishes and their fisheries in light of global change

In light of ocean warming and loss of Arctic sea ice, harvested marine fishes of boreal origin (and their fisheries) move poleward into yet unexploited parts of the Arctic seas. Industrial fisheries, already in place on many Arctic shelves, will radically affect the local fish species as they turn up as unprecedented bycatch. Arctic marine fishes are indispensable to ecosystem structuring and functioning, but they are still beyond credible assessment due to lack of basic biological data. The time for conservation actions is now, and precautionary management practices by the Arctic coastal states are needed to mitigate the impact of industrial fisheries in Arctic waters. We outline four possible conservation actions: scientific credibility, ‘green technology’, legitimate management and overarching coordination.     

Gene flow on ice: the role of sea ice and whaling in shaping Holarctic genetic diversity and population differentiation in bowhead whales (Balaena mysticetus)

Sea ice is believed to be a major factor shaping gene flow for polar marine organisms, but it remains unclear to what extent it represents a true barrier to dispersal for arctic cetaceans. Bowhead whales are highly adapted to polar sea ice and were targeted by commercial whalers throughout Arctic and subarctic seas for at least four centuries, resulting in severe reductions in most areas. Both changing ice conditions and reductions due to whaling may have affected geographic distribution and genetic diversity throughout their range, but little is known about range‐wide genetic structure or whether it differed in the past. This study represents the first examination of genetic diversity and differentiation across all five putative stocks, including Baffin Bay‐Davis Strait, Hudson Bay‐Foxe Basin, Bering‐Beaufort‐Chukchi, Okhotsk, and Spitsbergen. We also utilized ancient specimens from Prince Regent Inlet (PRI) in the Canadian Arctic and compared them with modern stocks. Results from analysis of molecular variance and demographic simulations are consistent with recent and high gene flow between Atlantic and Pacific stocks in the recent past. Significant genetic differences between ancient and modern populations suggest PRI harbored unique maternal lineages in the past that have been recently lost, possibly due to loss of habitat during the Little Ice Age and/or whaling. Unexpectedly, samples from this location show a closer genetic relationship with modern Pacific stocks than Atlantic, supporting high gene flow between the central Canadian Arctic and Beaufort Sea over the past millennium despite extremely heavy ice cover over much of this period.     

Faunal evidence for a late quaternary trans‐Antarctic seaway

Collapse of the West Antarctic Ice Sheet (WAIS) would raise global sea level by ～3.3–5 m. Ice‐sheet models and geological data suggest at least one collapse has happened during the last 1.1 Ma, and some scenarios of future climate change predict a collapse within the next two centuries. A complete WAIS collapse would open shallow seaways across West Antarctica, potentially enabling exchange of animals between West Antarctic seas. We investigated biological evidence for past connectivity between different regions of Antarctica by comparing the composition of modern bryozoan assemblages from the continental margin around Antarctica. Surprisingly, we found most similarity between two areas which are not currently connected – the shelves of the Weddell Sea (WS) and Ross Sea (RS). We evaluated three hypotheses to explain this and conclude that bryozoans most likely dispersed through a trans‐Antarctic seaway that opened in response to a WAIS collapse and connected the WS and RS shelves. These bryozoans must have survived glaciations(s) during subsequent ice ages in refuges, whereas they were wiped out in most other regions of the Antarctic shelf. After the last glacial period, bryozoan assemblages could freely disperse between many of the regions we examined (e.g. Antarctic Peninsula and South Shetland Islands), which has allowed recolonization of areas in which bryozoans had been eradicated during the last ice age. For the bryozoans on the WS and RS shelves to be more similar than those which are in close proximity means the trans‐Antarctic seaway may have been as late as the last few interglacials. Current rates of warming are exceptional compared with the near past glacial cycles so our study, the strongest faunal evidence of WAIS collapse during the recent geological past, thus supports predictions of a near future WAIS collapse (with considerable global sea level implications) and resultant future major faunal exchanges.     

Molecular evolution of haemoglobins of polar fishes

The Arctic and the Antarctic differ by age and isolation of the respective marine faunas. Antarctic fish are highly stenothermal, in response to stable water temperatures, whereas the Arctic ones are exposed to seasonal and latitudinal temperature variations. The knowledge of the mechanisms of phenotypic response to cold exposure in species of both polar habitats offers fundamental insights into the nature of environmental adaptation. In the process of cold adaptation, the evolutionary trend of Antarctic fish has led to unique specialisations, including modification of haematological characteristics, e.g. decreased amounts and multiplicity of haemoglobins.Unlike Antarctic Notothenioidei, Arctic teleosts have high haemoglobin multiplicity. Although the presence of functionally and structurally distinct haemoglobins is a plesiomorphic condition for many perciform-like fishes, it seems that the oxygen-transport system of teleost fish in the Arctic region has been adjusted to temperature differences and fluctuations of Arctic waters, much larger than in the Antarctic. The amino-acid sequences used to gain insight into the evolution history of α and β globins of polar fish have clearly shown that Antarctic and Arctic globins have different phylogenies, leading to the hypothesis that the selective pressure of environment stability allows the phylogenetic signal to be maintained in the Antarctic sequences, whereas environmental variability would tend to disrupt this signal in Arctic sequences.     

Is the poleward expansion by Atlantic cod and haddock threatening native polar cod, <Emphasis Type="Italic">Boreogadus saida</Emphasis>?

During a recent period of increased influx of warm Atlantic water to the western coast of Svalbard, we have observed a northward expansion of boreal Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) into areas dominated by the native polar cod (Boreogadus saida). To determine the potential impact of new ecological interactions, we studied the diet of co-occurring juvenile gadoids in fjords, open water, and sea ice around Svalbard. We also reviewed the available literature on polar cod feeding in different habitats across the Arctic to determine whether region, habitat, or fish size may influence diet. Feeding by polar cod in the pelagic zone was size dependent, with small fish primarily consuming Calanus spp. and smaller copepods, with an increasing ration of Themisto spp. at larger sizes. In benthic habitats, diets were more varied and included considerably more unidentified material and sediment. Less than 40% dietary overlap was detected among the three species when they were found together. Stable isotope analyses indicated these patterns were representative of longer-term assimilation. The low interspecific dietary overlap suggests little direct competition. Future increases in abundance and the high predation potential of the boreal taxa, however, may impact the persistence of polar cod on some Arctic shelves.