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.     

Detecting glacial refugia in the Southern Ocean

Throughout the Quaternary, the continental-based Antarctic ice sheets expanded and contracted repeatedly. Evidence suggests that during glacial maxima, grounded ice eliminated most benthic (bottom-dwelling) fauna across the Antarctic continental shelf. However, paleontological and molecular evidence indicates most extant Antarctica benthic taxa have persisted in situ throughout the Quaternary. Where and how the Antarctic benthic fauna survived throughout repeated glacial maxima remain mostly hypothesised. If understood, this would provide valuable insights into the ecology and evolution of Southern Ocean biota over geological timescales. Here we synthesised and appraised recent studies and presented an approach to demonstrate how genetic data can be effective in identifying where and how Antarctic benthic fauna survived glacial periods. We first examined the geological and ecological evidence for how glacial periods influenced past species demography in order to provide testable frameworks for future studies. We outlined past ice-free areas from Antarctic ice sheet reconstructions that could serve as glacial refugia and discussed how benthic fauna with pelagic or non-pelagic dispersal strategies moved into and out of glacial refugia. We also reviewed current molecular studies and collated proposed locations of Southern Ocean glacial refugia on the continental shelf around Antarctica, in the deep sea, and around sub-Antarctic islands. Interestingly, the proposed glacial refugia based on molecular data generally do not correspond to the ice-free areas identified by Antarctic ice sheet reconstructions. The potential biases in sampling and in the choice of molecular markers in current literature are discussed, along with the future directions for employing testable frameworks and genomic methods in Southern Ocean molecular studies. Continued data syntheses will elucidate greater understanding of where and how Southern Ocean benthic fauna persisted throughout glacial periods and provide insights into their resilience against climate changes in the future.     

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.     

Projected Polar Bear Sea Ice Habitat in the Canadian Arctic Archipelago

Background

Sea ice across the Arctic is declining and altering physical characteristics of marine ecosystems. Polar bears (Ursus maritimus) have been identified as vulnerable to changes in sea ice conditions. We use sea ice projections for the Canadian Arctic Archipelago from 2006 – 2100 to gain insight into the conservation challenges for polar bears with respect to habitat loss using metrics developed from polar bear energetics modeling.

Principal Findings

Shifts away from multiyear ice to annual ice cover throughout the region, as well as lengthening ice-free periods, may become critical for polar bears before the end of the 21^st century with projected warming. Each polar bear population in the Archipelago may undergo 2–5 months of ice-free conditions, where no such conditions exist presently. We identify spatially and temporally explicit ice-free periods that extend beyond what polar bears require for nutritional and reproductive demands.

Conclusions/Significance

Under business-as-usual climate projections, polar bears may face starvation and reproductive failure across the entire Archipelago by the year 2100.     

Adaptation of Arctic and Antarctic ice metazoa to their habitat

Sea ice is a unique habitat in polar seas. A diverse assemblage of plants and animals lives in its interior parts and at the ice-water interface. Their distribution is to a large extent controlled by abiotic parameters such as light, salinity and space, as well as food availability. In both the Arctic and Antarctic, the highest metazoan concentrations occur mostly in the bottom centimetres of the sea ice. Dominant metazoans are nematodes, turbellarians, rotifers and crustaceans. The ice-water interface itself houses in addition to endemic amphipods migrants from both the ice and the pelagic realm. To survive with the environmental conditions of the sea ice habitat, the ice biota is adapted, specifically to seasonal salinity variations from below 5 to above 60 PSU. Sea ice metazoans feed mainly on the algae growing within the sea ice. The loss of habitat during ice melt periods can lead to substantial sedimentation of ice fauna to the sea floor, where it might act as food source for the benthos.     

Evidence for Permo-Triassic colonization of the deep sea by isopods

The deep sea is one of the largest ecosystems on Earth and is home to a highly diverse fauna, with polychaetes, molluscs and peracarid crustaceans as dominant groups. A number of studies have proposed that this fauna did not survive the anoxic events that occurred during the Mesozoic Era. Accordingly, the modern fauna is thought to be relatively young, perhaps having colonized the deep sea after the Eocene/Oligocene boundary. To test this hypothesis, we performed phylogenetic analyses of nuclear ribosomal 18S and 28S and mitochondrial cytochrome oxidase I and 16S sequences from isopod crustaceans. Using a molecular clock calibrated with multiple isopod fossils, we estimated the timing of deep-sea colonization events by isopods. Our results show that some groups have an ancient origin in the deep sea, with the earliest estimated dates spanning 232–314 Myr ago. Therefore, anoxic events at the Permian–Triassic boundary and during the Mesozoic did not cause the extinction of all the deep-sea fauna; some species may have gone extinct while others survived and proliferated. The monophyly of the ‘munnopsid radiation’ within the isopods suggests that the ancestors of this group evolved in the deep sea and did not move to shallow-water refugia during anoxic events.     

Climate change impacts on wildlife in a High Arctic archipelago – Svalbard,Norway

The Arctic is warming more rapidly than other region on the planet, and the northern Barents Sea, including the Svalbard Archipelago, is experiencing the fastest temperature increases within the circumpolar Arctic, along with the highest rate of sea ice loss. These physical changes are affecting a broad array of resident Arctic organisms as well as some migrants that occupy the region seasonally. Herein, evidence of climate change impacts on terrestrial and marine wildlife in Svalbard is reviewed, with a focus on bird and mammal species. In the terrestrial ecosystem, increased winter air temperatures and concomitant increases in the frequency of ‘rain‐on‐snow’ events are one of the most important facets of climate change with respect to impacts on flora and fauna. Winter rain creates ice that blocks access to food for herbivores and synchronizes the population dynamics of the herbivore–predator guild. In the marine ecosystem, increases in sea temperature and reductions in sea ice are influencing the entire food web. These changes are affecting the foraging and breeding ecology of most marine birds and mammals and are associated with an increase in abundance of several temperate fish, seabird and marine mammal species. Our review indicates that even though a few species are benefiting from a warming climate, most Arctic endemic species in Svalbard are experiencing negative consequences induced by the warming environment. Our review emphasizes the tight relationships between the marine and terrestrial ecosystems in this High Arctic archipelago. Detecting changes in trophic relationships within and between these ecosystems requires long‐term (multidecadal) demographic, population‐ and ecosystem‐based monitoring, the results of which are necessary to set appropriate conservation priorities in relation to climate warming.     

Patterns and controlling factors of species diversity in the Arctic Ocean

Aim The Arctic Ocean is one of the last near‐pristine regions on Earth, and, although human activities are expected to impact on Arctic ecosystems, we know very little about baseline patterns of Arctic Ocean biodiversity. This paper aims to describe Arctic Ocean‐wide patterns of benthic biodiversity and to explore factors related to the large‐scale species diversity patterns. Location Arctic Ocean. Methods We used large ostracode and foraminiferal datasets to describe the biodiversity patterns and applied comprehensive ecological modelling to test the degree to which these patterns are potentially governed by environmental factors, such as temperature, productivity, seasonality, ice cover and others. To test environmental control of the observed diversity patterns, subsets of samples for which all environmental parameters were available were analysed with multiple regression and model averaging. Results Well‐known negative latitudinal species diversity gradients (LSDGs) were found in metazoan Ostracoda, but the LSDGs were unimodal with an intermediate maximum with respect to latitude in protozoan foraminifera. Depth species diversity gradients were unimodal, with peaks in diversity shallower than those in other oceans. Our modelling results showed that several factors are significant predictors of diversity, but the significant predictors were different among shallow marine ostracodes, deep‐sea ostracodes and deep‐sea foraminifera. Main conclusions On the basis of these Arctic Ocean‐wide comprehensive datasets, we document large‐scale diversity patterns with respect to latitude and depth. Our modelling results suggest that the underlying mechanisms causing these species diversity patterns are unexpectedly complex. The environmental parameters of temperature, surface productivity, seasonality of productivity, salinity and ice cover can all play a role in shaping large‐scale diversity patterns, but their relative importance may depend on the ecological preferences of taxa and the oceanographic context of regions. These results suggest that a multiplicity of variables appear to be related to community structure in this system.     

Effects of Late-Cenozoic Glaciation on Habitat Availability in Antarctic Benthic Shrimps (Crustacea: Decapoda: Caridea)

Marine invertebrates inhabiting the high Antarctic continental shelves are challenged by disturbance of the seafloor by grounded ice, low but stable water temperatures and variable food availability in response to seasonal sea-ice cover. Though a high diversity of life has successfully adapted to such conditions, it is generally agreed that during the Last Glacial Maximum (LGM) the large-scale cover of the Southern Ocean by multi-annual sea ice and the advance of the continental ice sheets across the shelf faced life with conditions, exceeding those seen today by an order of magnitude. Conditions prevailing at the LGM may have therefore acted as a bottleneck event to both the ecology as well as genetic diversity of today''s fauna. Here, we use for the first time specific Species Distribution Models (SDMs) for marine arthropods of the Southern Ocean to assess effects of habitat contraction during the LGM on the three most common benthic caridean shrimp species that exhibit a strong depth zonation on the Antarctic continental shelf. While the shallow-water species Chorismus antarcticus and Notocrangon antarcticus were limited to a drastically reduced habitat during the LGM, the deep-water shrimp Nematocarcinus lanceopes found refuge in the Southern Ocean deep sea. The modeling results are in accordance with genetic diversity patterns available for C. antarcticus and N. lanceopes and support the hypothesis that habitat contraction at the LGM resulted in a loss of genetic diversity in shallow water benthos.     

Melting barriers to faunal exchange across ocean basins

Accelerated loss of sea ice in the Arctic is opening routes connecting the Atlantic and Pacific Oceans for longer periods each year. These changes may increase the ease and frequency with which marine birds and mammals move between the Pacific and Atlantic Ocean basins. Indeed, recent observations of birds and mammals suggest these movements have intensified in recent decades. Reconnection of the Pacific and Atlantic Ocean basins will present both challenges to marine ecosystem conservation and an unprecedented opportunity to examine the ecological and evolutionary consequences of interoceanic faunal exchange in real time. To understand these changes and implement effective conservation of marine ecosystems, we need to further develop modeling efforts to predict the rate of dispersal and consequences of faunal exchange. These predictions can be tested by closely monitoring wildlife dispersal through the Arctic Ocean and using modern methods to explore the ecological and evolutionary consequences of these movements.     

Integrated abundance and biomass of sympagic meiofauna in Arctic and Antarctic pack ice

The abundance and biomass of sympagic meiofauna were studied during three cruises to the Antarctic and one summer expedition to the central Arctic Ocean. Ice samples were collected by ice coring and algal pigment concentrations and meiofauna abundances were determined for entire cores. Median meiofauna abundances for the expeditions ranged from 4.4 to 139.5 × 10³ organisms m⁻² in Antarctic sea ice and accounted for 40.6 × 10³ organisms m⁻² in Arctic multi-year sea ice. While most taxa (ciliates, foraminifers, turbellarians, crustaceans) were common in both Arctic and Antarctic sea ice, nematodes and rotifers occurred only in the Arctic. Based on the calculated biomass, the potential meiofauna ingestion rates were determined by applying an allometric model. For both hemispheres, daily and yearly potential ingestion rates were below the production values of the ice algal communities, pointing towards non-limited feeding conditions for ice meiofauna year-round. Accepted: 29 March 1999     

Paleoceanography of the mid-Pleistocene

The fossil record of the Pleistocene calcareous nannoplankton indicates that during the mid-Pleistocene (0.93–1.25 my) occurred an episode of overwhelming dominance of smallGephyrocapsa. During this episode normally abundant, large size specimens of this genus (mainlyGephyrocapsa oceanica) were virtually excluded from the phytoplankton of tropical and subtropical oceans. The best modern analog of this dominantly smallGephyrocapsa assemblage is the subpolarEmiliania huxleyi assemblage, which implies that nutrient content was significantly greater and water temperature was lower in the photic water column of the tropical oceans than they are today. Increased equatorial upwelling in the oceans, on a scale much greater than today, may explain the above pattern.To achieve such broad equatorial upwelling there must be a source and a drive for cold, dense water. The Arctic Ocean, which was probably seasonally free of ice during this interval of the mid-Pleistocene, is capable of providing the requisite source as well as a drive for the inferred equatorial upwelling. The energy balance of a predominantly ice-free Arctic Ocean requires an approximately three to seven fold increase of hydrospheric heat transport from the North Atlantic to the Arctic Ocean, which dictates a corresponding or even greater increase in the volume of warm water entering the Arctic Ocean at the surface and cold dense water exiting at depth to the North Atlantic. Such enhanced dense water formation in the Arctic Ocean could drive the intensified equatorial upwelling implied by the smallGephyrocapsa dominance interval.If the above scenario is correct then the climate of the earth's northern hemisphere during the mid-Pleistocene may have been very different from the younger Pleistocene climate. One manifestation of this difference may be the mid-Pleistocene shift in climatic cycle periodicity from 40 ky to 100 ky. Another important aspect is that the enhanced greenhouse effect expected during the next century because of an increase of atmospheric CO₂ is thought to lead directly to melting of the Arctic Ocean ice cover and of the Greenland ice sheet. Thus, the “greenhouse” Arctic Ocean and its attendant ocean circulation would resemble the inferred mid-Pleistocene conditions.     

Vocalizations of bearded seals (Erignathus barbatus) and their influence on the soundscape of the western Canadian Arctic

The soundscape is a crucial habitat feature for marine mammals. This study investigates the contribution of bearded seal vocalizations to the soundscape in the western Canadian Arctic, and also the vocal characteristics of bearded seals relative to sea ice conditions. Passive acoustic data were recorded near Sachs Harbour between August 2015 and July 2016. Sound pressure levels (SPL) in the 50–1,000 Hz and 1–10 kHz bands increased as the total duration of all bearded seal vocalizations increased, and this relationship was moderated by sea ice concentration. Bearded seals in this region had an overlapping vocal repertoire with bearded seals in other areas of the Arctic, and had seven additional vocalizations that have not been previously documented for this region. This study is the first detailed assessment of the influence of bearded seal calls on SPL, which shows the high potential of bearded seals to influence underwater sound levels during the mating season. Bearded seals live in a changing Arctic seascape, and their influence on the soundscape may shift as sea ice continues to diminish. It is imperative that acoustic monitoring continues within the Arctic, and this study provides a baseline for future monitoring as the Arctic continues to change.     

北冰洋海域微食物环研究进展

海洋微食物环在海洋生态系统中起着重要作用.北冰洋因常年为海冰所覆盖,对微食物环的研究较为有限.现有研究表明,微食物环在北冰洋生态系统中的作用与海域和季节相关.近年来环境的快速变化、特别是夏季海冰覆盖面积的迅速减少,会对微食物环的结构和功能产生重大影响,已有研究显示其生态作用有望进一步提高.综合近年来已有的研究成果,对北冰洋微食物环的主要类群:原核生物、真核浮游植物、原生动物和浮游病毒等的基本生态特征进行了概述,讨论了各类群间的相互关系,并对未来的研究重点进行了展望.     

Transgressions: rethinking Beringian glaciation

The purpose of this investigation is to encourage a fresh look at Pleistocene Beringia. Heretofore, flooding of Bering Strait has been cited as the only barrier to migration, with marine sea transgressions being a “sea gate” that closed off migration during glacial interstadials and interglaciations. However, the possibility exists that glacial advances were also barriers, with marine ice transgressions being an “ice gate” that closed off migration during glacial stadials and glacial maxima. This possibility proceeds from the Marine Ice Transgression Hypothesis (MITH), which states that marine ice sheets form on the broad Arctic continental shelf of Northern Hemisphere continents when sea ice thickens, grounds and domes in shallow water, and then transgresses landward as continental ice sheets and seaward as floating ice shelves (Hughes, 1987). Landward transgression is onto coastal lowlands. During Pleistocene glaciations, a marine ice sheeet extending from Spitsbergen to Greenland may have transgressed the circumpolar continental landmass at its lowest and narrowest gap, central Beringia, and calved into the Pacific Ocean.

Four models of Beringian glaciation are presented, based on the distinction between marine glaciation and highland glaciation. Central Beringia was glaciated only in highlands in the traditional model (Hopkins et al., 1982), was also glaciated by a self-sustaining ice shelf floating over the deep ocean basins of the Bering Sea in the model by Grosswald and Vozovik (1984), was glaciated by a marine ice sheet that covered highlands, the continental shelf, and supplied the ice shelf in a model for maximum Pleistocene glaciation, and was glaciated by a marine ice sheet in the Chukchi Sea that merged with highland glaciers, transgressed the continental shelf of the western Bering Sea, and calved into the southern Bering Sea along the edge of the continental shelf in a model for the last glaciation. Field tests are suggested to assess the viability of these four models. The first model is already established for highland glaciation in Alaska, but less established in Siberia. The last model should be the easiest to evaluate for marine glaciation. The last model limits human migration across the Beringian land bridge to brief intervals between stadials and interstadials of the last glaciation cycle, when both the ice gate and the sea gate were opened to human migration. This model can influence the sea change now underway among Quaternary scientists studying peopling of the Americas, based on the archaeological, linguistic and ethnic diversity among native American populations.     

Timing of blooms,algal food quality and Calanus glacialis reproduction and growth in a changing Arctic

The Arctic bloom consists of two distinct categories of primary producers, ice algae growing within and on the underside of the sea ice, and phytoplankton growing in open waters. Long chain omega‐3 fatty acids, a subgroup of polyunsaturated fatty acids (PUFAs) produced exclusively by these algae, are essential to all marine organisms for successful reproduction, growth, and development. During an extensive field study in the Arctic shelf seas, we followed the seasonal biomass development of ice algae and phytoplankton and their food quality in terms of their relative PUFA content. The first PUFA‐peak occurred in late April during solid ice cover at the onset of the ice algal bloom, and the second PUFA‐peak occurred in early July just after the ice break‐up at the onset of the phytoplankton bloom. The reproduction and growth of the key Arctic grazer Calanus glacialis perfectly coincided with these two bloom events. Females of C. glacialis utilized the high‐quality ice algal bloom to fuel early maturation and reproduction, whereas the resulting offspring had access to ample high‐quality food during the phytoplankton bloom 2 months later. Reduction in sea ice thickness and coverage area will alter the current primary production regime due to earlier ice break‐up and onset of the phytoplankton bloom. A potential mismatch between the two primary production peaks of high‐quality food and the reproductive cycle of key Arctic grazers may have negative consequences for the entire lipid‐driven Arctic marine ecosystem.     

Atlantic walrus signal latitudinal differences in the long-term decline of sea ice-derived carbon to benthic fauna in the Canadian Arctic

Climate change is altering the biogeochemical and physical characteristics of the Arctic marine environment, which impacts sea ice algal and phytoplankton bloom dynamics and the vertical transport of these carbon sources to benthic communities. Little is known about whether the contribution of sea ice-derived carbon to benthic fauna and nitrogen cycling has changed over multiple decades in concert with receding sea ice. We combined compound-specific stable isotope analysis of amino acids with highly branched isoprenoid diatom lipid biomarkers using archived (1982–2016) tissue of benthivorous Atlantic walrus to examine temporal trends of sea ice-derived carbon, nitrogen isotope baseline and trophic position of Atlantic walrus at high- and mid-latitudes in the Canadian Arctic. Associated with an 18% sea ice decline in the mid-Arctic, sea ice-derived carbon contribution to Atlantic walrus decreased by 75% suggesting a strong decoupling of sea ice-benthic habitats. By contrast, a nearly exclusive amount of sea ice-derived carbon was maintained in high-Arctic Atlantic walrus (98% in 1996 and 89% in 2006) despite a similar percentage in sea ice reduction. Nitrogen isotope baseline or the trophic position of Atlantic walrus did not change over time at either location. These findings indicate latitudinal differences in the restructuring of carbon energy sources used by Atlantic walrus and their benthic prey, and in turn a change in Arctic marine ecosystem functioning between sea ice–pelagic–benthic habitats.     

Late Quaternary environmental and climate history of Rauer Group, East Antarctica

The Rauer Group is an archipelago in Prydz Bay, East Antarctica. The ice-free islands and the surrounding shallow marine areas provide valuable archives for the reconstruction of the late Pleistocene and Holocene environmental and climatic history of the region. Two sediment records from two marine inlets of Rauer Group have been studied for their sedimentological, geochemical, and biological characteristics. Radiocarbon ages from one of the inlets indicate ice-free conditions within the last glacial cycle, probably during the second half of Marine Isotope Stage 3. Subsequent ice sheet coverage of Rauer Group during the Last Glacial Maxiumum (LGM) can be inferred from a till layer recovered in one of the basins. The inlets became ice-free prior to 11,200 cal yr BP, when biogenic sedimentation started. Deglacial processes in the catchments, however, influenced the inlets until ~ 9200 cal yr BP as evidenced by the input of minerogenic material. Marine productivity under relatively open water conditions indicates an early Holocene climate optimum until 8200 cal yr BP, which is followed by a cooler period with increased sea ice. Warmer conditions are inferred for the mid Holocene, when both basins experienced an input of freshwater between ~ 5700-3500 cal yr BP, probably due to ice-sheet melting and increased precipitation on the islands. Neoglacial cooling in the late Holocene since c. 3500 cal yr BP is reflected by an increase in sea ice in both inlets.     

An avian terrestrial predator of the Arctic relies on the marine ecosystem during winter

Top predators of the arctic tundra are facing a long period of very low prey availability during winter and subsidies from other ecosystems such as the marine environment may help to support their populations. Satellite tracking of snowy owls, a top predator of the tundra, revealed that most adult females breeding in the Canadian Arctic overwinter at high latitudes in the eastern Arctic and spend several weeks (up to 101 d) on the sea‐ice between December and April. Analysis of high‐resolution satellite images of sea‐ice indicated that owls were primarily gathering around open water patches in the ice, which are commonly used by wintering seabirds, a potential prey. Such extensive use of sea‐ice by a tundra predator considered a small mammal specialist was unexpected, and suggests that marine resources subsidize snowy owl populations in winter. As sea‐ice regimes in winter are expected to change over the next decades due to climate warming, this may affect the wintering strategy of this top predator and ultimately the functioning of the tundra ecosystem.     

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.