Ratkova.indd 95Ratkova & Wassmann 2005: Polar Research 24(1–2), 95–110 Sea ice algae in the White and Barents seas: composition and origin Tatjana N. Ratkova & Paul Wassmann To examine algae populations, three expeditions (in March 2001, April 2002 and February 2003) were conducted in the Guba Chupa (Chupa Estuary; north-western White Sea), and one cruise was carried out in the open part of the White Sea in April 2003 and in the northern part of the Barents Sea in July 2001. Sea ice algae and phytoplankton composition and abundance and the content of sediment traps under the land-fast ice in the White Sea and annual and multi-year pack ice in the Barents Sea were investigated. The community in land-fast sea ice was dominated by pen- nate diatoms and its composition was more closely related to that of the underlying sediments than was the community of the pack ice, which was dominated by fl agellates, dinofl agellates and centric diatoms. Algae were far more abundant in land-fast ice: motile benthic and ice-benthic spe- cies found favourable conditions in the ice. The pack ice community was more closely related to that of the surrounding water. It originated from plankton incorporation during sea ice formation and during seawater fl ood events. An additional source for ice colonization may be multi-year ice. Algae may be released from the ice during brine drainage or sea ice melting. Many sea ice algae developed spores before the ice melt. These algae were observed in the above-bottom sediment traps all year around. Three possible fates of ice algae can be distinguished: 1) suspension in the water column, 2) sinking to the bottom and 3) ingestion by herbivores in the ice, at the ice–water interface or in the water column. T. N. Ratkova, Shirshov Institute of Oceanology, Academy of Sciences of Russia, Nakhimovsky Ave. 36, 117997, Moscow, Russia, trat@orc.ru; P. Wassmann, Norwegian College of Fishery Science, University of Tromsø, NO-9037 Tromsø, Norway. High concentrations of organisms in polar sea ice have been well documented (e.g. Horner 1985; Lizotte 2003). Sea ice is inhabited by diverse communities, including bacteria, algae, protists and metazoans (e.g. Melnikov 1997). Often the fl ora and fauna are not evenly distributed in the ice, but occur in specifi c microhabitats in differ- ent parts of the ice fl oes (Horner 1985; Syvertsen 1991; Gradinger 1999). Such communities have been studied in detail, but the current knowl- edge of the biological processes involved during sea ice formation and melting are still inadequate (Gradinger & Ikävalko 1998; Weissenberger & Grossmann 1998). There are several hypotheses regarding the origin and fate of sea ice commu- nities (Horner 1985). Sieving of water by densely packed ice crystals has been observed repeated- ly and appears to constitute an important mech- anism for accumulation of algae cells on the ice under-surface (Syvertsen 1991; Melnikov 1997). The composition of sea ice algae off the shelves suggests that this population derives from pelagic algae inhabiting underlying water masses or from sea ice algae at the ice edge or polynyas (Syvert- sen 1991; Druzhkov et al. 2001). In shallow water areas benthic algae may be included in the suc- 96 Sea ice algae in the White and Barents seas cession cycle (Poulin 1990; Gogorev 1998). Through various research programmes we studied and compared the ice fl ora of the White and Barents seas, two regions inadequately stud- ied with regard to ice biota. We investigated in particular the peculiarities of the vertical distri- bution of the sea ice algae under different con- ditions and the succession of this distribution during the late winter/early spring in the land-fast ice in the White Sea. The differences in the algae species composition in different types of ice and the relationships between taxa within the ice and water column were also quantifi ed. Materials and methods Three cruises were conducted to the Guba Chupa (Chupa Estuary; north-western White Sea) in February 2003, March 2001 and April 2002. One cruise was carried out in the open part of the White Sea (April 2003), where the stations were located in the Marginal Ice Zone (MIZ) in the southern part of Gorlo (north-eastern White Sea). One cruise was carried out in the MIZ of the northern part of the Barents Sea in July 2001 (Table 1). Ice cores from the White Sea were obtained with a ring corer (diameter 14.5 cm) and divided into three or four sections: upper 10 cm, lower 3 - 5 cm, and a middle part, which was cut into two parts if the total ice thickness exceeded 40 cm. The sections were melted at low temper- atures (< 2 °C) without addition of fi ltered water. The salinity of the White Sea ice is low (Krell et al. 2003), and we deem that the algae were not being exposed to osmotic stress during melt- ing. In the Barents Sea, ice cores were collected with an ice auger (diameter 9 cm). The lowermost 20 - 30 cm of each ice core was melted in double its volume of fi ltered seawater at low tempera- tures. Samples from the water column below the ice were also obtained in both regions (Appen- dix). The volume of the sea ice and water sam- ples ranged from 100 to 4500 ml, according to the abundance of the algae. To investigate the pos- sible importance of sea ice algae for the vertical fl ux of biogenic matter, cylindrical sediment traps were deployed below the ice: in the White Sea (Guba Chupa) the traps were preserved with for- maldehyde and exposed for a week at 20 m depth. Traps were also deployed above the bottom in the Kandalaksha Bay (depth 280 m) and nine sam- ples (each representing one month) were taken from August 2002 to May 2003. In the Barents Sea, traps were deployed without preservation for one day at 30 m depth . The height/diameter ratio of the traps was 3.11 for the White Sea and 3 for the Barents Sea. Temperature and salinity were measured with standard CTD profi lers. All samples were preserved with a glutardial- dehyde–lugol–ethanol solution and developed according to Ratkova & Wassmann (2002). Small fl agellates were counted in a Fuchs-Rosenthal chamber at 600× magnifi cation prior to any con- centration of samples. After they had settled, larger algae were counted in chambers with rul- Table 1. Biomass (mg C m–3) of the sea-ice algae (“algae”) in the lowermost layer of the ice in the Barents (5 - 34 cm of 100 - 135 cm cores) and in the White Sea (5 - 10 cm of 37 - 56 cm cores) and of phytoplankton (“phyto”) below the ice. Barents Sea White Sea (pack ice) White Sea (land-fast ice) 77° 50' N, 29° 45' E 78° 20' N, 27° 20' E 82° 00' N, 26° 00' E 65° 40' - 66° 10' N, 40° 10' - 40° 50' E 66° 20’N, 33° 40' E 3 July 2001 6 July 2001 9 July 2001 19–20 April 2003 8 February 2003 18–26 March 2001 6–8 April 2002 algae phyto algae phyto algae phyto algae phyto algae phyto algae phyto algae phyto Diatoms 7.58 5 - 44 49.2 1 - 7 254.6 1 - 7 44 - 478 11 - 12 1.49 0. 07 - 0.31 94 - 598 1 - 64 1136 - 35188 1 - 94 Dino- fl agellates 0.49 34 - 80 14.0 25 - 33 11.0 1 - 2 1.3 - 21.7 0.01 - 1.06 0.49 0.03 - 0.18 3.1 - 37.5 0.02 - 1 24.5 - 178.6 1 - 28 Other fl agellates 17.0 82 - 299 86.8 82 - 173 76.5 32 - 74 36 - 49 27 - 94 4.4a 0.5 - 2.2a 8.6 - 35.7 11 - 359 3.8 - 61.7b 1 - 525b Total 28.3 154 - 407 150 117 - 218 343.8 35 - 83 87 - 565 39 - 109 6.49 0.12 - 2.69 115.9 - 674.1 15 - 444 1164 - 35428 3 - 647 a According Sazhin et al. in press. b According Sazhin et al. 2004. 97Ratkova & Wassmann 2005: Polar Research 24(1–2), 95–110 ings, at 300× magnifi cation in a 0.06 ml chamber and at 150× magnifi cation in a 1.0 ml chamber. The carbon content of the algae was calculated accord- ing to Strathmann (1967) for diatoms with a cell volume > 3000 µm3, and according to Menden- Deuer & Lessard (2000) for all other protists. Results Physical environment The temperature of the water in the White Sea seldom reached the freezing point, and the ice developed mainly from the surface (Krell et al. 2003; Pantyulin 2003). The ice was thin during all of the expeditions (thickness 34 - 57 cm) and often fl ooded with seawater. Granular snow ice dominated the entire ice column at all times in the White Sea. Only in March 2001 and in February 2003 was columnar ice observed in the middle part of the ice cores. The sea ice structure was determined by visual observation of the ice core, but was supported by investigation of the crystal structure of thin ice core sections (February and April 2002) (Krell et al. 2003). The winter of 2001 was unusually mild and ice formation did not take place until February. The ice was rather porous, granular and white in the upper and in the lowermost parts and columnar and grey in the middle part of the ice cores. The ice was 44 - 57 cm thick and was covered with a layer of snow which increased in thickness from ca. 10 cm on the fi rst day of the expedition to ca. 16 cm a day later, after a snowfall. Below the ice there was a < 1 m thick brackish layer (psu < 15 ‰) with a temperature of –1.2 °C—well above the freezing point. In April 2002 a < 1 m thick brackish layer (psu < 6 ‰, temperature: –1.2 °C) was encountered below the ice; the sea ice below a thin snow cover was granular, white-grey and semi-transparent, and wide channels were observed in the lower 10 cm of ice cores (thickness 34 - 54 cm). A brownish discolouration was observed at the low- ermost surface. In February 2003 the white, dull granular ice was covered with a thick layer of slush (up to 15 cm) and snow (ca. 10 cm). In the middle part, columnar semi-transparent ice was observed. In the lowermost few centimetres, the ice was opaque, white and granular. The water salinity was high (psu 26.1 ‰) and the temperature was –1.2 °C directly beneath the ice (thickness 34 - 52 cm); both parameters increased with the depth. In April 2003 the depth of the snow cover ranged from 10 to 20 cm on different ice fl oes. The ice fl oes, which remained intact after a storm event, showed no signs of surface melting. However, considerable ice melting occurred from the sides and from below, due to relatively warm water. The lower part of the ice cores (thickness 45 cm) was semi-transparent, granular and had large caverns. The middle part was greyish, opaque and dense. The upper part of the ice layer was not much dif- ferent from the middle part, though it was softer. In the water column, salinity and temperature varied little from the surface to the bottom. From the north-west part of the Gorlo Strait (66° 10' N; 40° 10' E), where most of the water is from the Barents Sea, to the south-east part of the strait (65°40' N; 40°50' E), where White Sea water dom- inates, the temperature increased from –1.52 to –1.08 °C and the salinity decreased from 29.7 ‰ to 27.2 ‰ psu (Kosobokova et al. 2004). A peculiar feature of the physical oceanogra- phy in the MIZ of the northern Barents Sea in July 2001 was a 20 m thick layer of meltwater (–1 to 0.4 °C; psu 31 ‰) above the Arctic Water (< –1 °C; psu 34 ‰). The ice conditions in the investigated fl oes varied from annual pack ice fl oes (thickness about 1 m) to multi-year pack ice (thickness > 1.35 m). Differentiation between the multi-year and annual ice was based mainly on ice thickness and the density of the Melosira arc- tica mats observed by the divers. The snow cover was rather thin (< 10 cm). Phytoplankton and sea ice algae: composition A total of 306 algae species (Appendix) were identifi ed in the ice, in the water and in the sedi- ment traps. In White Sea land-fast ice, 275 species were encountered. The White Sea pack ice was inhabited by fewer species (106), and only four of them were not also observed in the land-fast ice. In the Barents Sea, 201 species were found. In general, the algae composition in the two regions was rather similar: 156 species, including 73 dia- toms, 39 dinofl agellates, all silicofl agellates and coccolithophorides and 31 other fl agellates were observed in both regions. The main differences were encountered among the pennate diatoms. In the White Sea land-fast ice, 110 pennate species were found, while only 49 and 52 species were found in the White Sea and the Barents Sea pack 98 Sea ice algae in the White and Barents seas ice, respectively. Most of the species were found both in the water column and the ice, but 50 species were not observed in the ice. Most of these were ben- thic ones which only occasionally occurred in the water, but there were also some common plank- tonic species which were not found in the ice, for example, the diatoms Chaetoceros borealis, C. brevis, C. decipiens, Corethron criophylum, Coscinodiscus centralis, C. radiatus, Probos- cia eumorpha, Rhizosolenia hebetata f. semispi- na; the dinofl agellates Dinophysis contracta, D. arctica, Heterocapsa triquetra, Protoperidinium pallidum, P. pellucidum, P. pyriforme, Warnowia reticulata and a few other fl agellates. Some species were observed only in the ice: 37 diatoms (e.g. Navicula algida, N. glacialis, N. gelida, Nitzshia hybridae, N. polaris, 22 other pennate and 10 centric species) and 7 dinofl ag- ellates (e.g. Gymnodinium wulfi i, Karenia brevis, Protoperidinium granii). Diatoms and euglenophytes were represented by more species in the ice (178 and 5, respective- ly) than in the water (154 and 4, respectively). All other groups were represented by more species in the water; except for chlorophytes, which were represented by the same number of species in the ice and in the water. The snow covering the ice demonstrated poor species diversity (44 species in the snow in com- parison to 228 species in the ice). All the spe- cies observed in the snow were also found in the underlying ice. The abundance of species also differed between sea ice and the water column: sea ice algae species were less abundant in the water, where plankton species had the higher abundance (Appendix). On the basis of morphological and ecological distinctions, some assemblages of the sea ice spe- cies were selected for comparison: A) pennate ice plankton species that develop ribbon-shaped colo- nies (Fossula arctica, Fragilaria striatula, Fragi- lariopsis spp., Pauliella taeniata and some species of Navicula and Nitzschia—hereafter “ribbon dia- toms”); B) single-celled ice-benthic pennate spe- cies that sometimes develop barrel-shaped colonies in ice (Enthomoneis spp., Undatella cf. quadrata, Navicula lineola—“barrel diatoms”); C) epiphytic species that are associated with Melosira arctica and Nitzschia frigida (Synedropsis spp., Gompho- nemopsis cf. exigua, Gomphonema septentriona- lis, Pseudogomphonema kamchaticum and Atteya septentrionalis—“associated diatoms”). Phytoplankton and sea ice algae: abundance In the White Sea land-fast ice, fl agellates domi- nated the phytoplankton carbon in all samples, but diatoms dominated in the sea ice (Table 1). Flagel- lates comprised < 25 % in the ice and 73 - 80 % in the water (3.8 - 62 and 0.5 - 525 mg C m–3, respec- tively). In February 2003 the species composition and abundance of diatoms and dinofl agellates in the White Sea land-fast ice were rather similar to those in the annual ice of the Barents Sea in July 2001, i.e. centric diatoms dominated in a sparse sea ice algal assemblage (Fig. 1). In March 2001 the species composition in the White Sea land-fast ice was quite different: barrel and other benthic and ice-benthic diatoms domi- nated the algae carbon in the ice. Benthic single- celled pennate diatoms were most abundant in the lowermost few centimetres of the ice core on 18 March before a snowfall (> 1 × 105 cells l–1) and in the upper 10 cm of the ice (Fig. 2) after the snowfall on 26 March (2.5 × 105 cells l–1). These species were rare in the water, where centric dia- toms dominated (Fig. 1). In April 2002, Nitzschia frigida dominated the total carbon of diatoms and dinofl agellates in the sea ice and water column (Fig. 1). Two types of White Sea pack ice were studied: pack ice fl oes in the southern part of the Gorlo Strait, probably formed far from the coast (“clean ice”), and fragmented ice in the northern part of the Gorlo Strait with characteristic brown miner- al insertions (“dirty” ice). Algae biomass in the “dirty” ice (407 mg С m–3) was close to the high- est values in the “clean” ice (87 - 565 mg С m–3). The composition of the algal populations differed Fig 1 (opposite page). Composition of the diatoms and dino- fl agellates abundance in the lowermost part of the ice, in the upper part of the water column and in sediment traps in the northern Barents Sea in July 2001 and in the White Sea in February 2003, March 2001, April 2002, and in the lower- most part of the ice and in the upper part of the water column in April 2003. “Ribbon” denotes plankton diatoms forming ribbon-shaped colonies; “barrel” is benthic single-celled dia- toms forming barrel-shaped colonies in the ice; and “Associ- ated” stands for species found in association with Melosira arctica and Nitzschia frigida. Each bar represents the value of the single sample in the Barents Sea and in March 2001 and April 2003 in the White Sea. The axes showing biomass are different. Each bar is the mean value from the 4 - 5 samples collected in April 2002 and in February 2003. Low biomasses of algae in the White Sea in the water in April 2003 and in the traps in February 2003 are indicated by numbers instead of bars. 99Ratkova & Wassmann 2005: Polar Research 24(1–2), 95–110 100 Sea ice algae in the White and Barents seas 101Ratkova & Wassmann 2005: Polar Research 24(1–2), 95–110 considerably: pennate ice-planktonic species were dominant in the “clean ice”, while plankton- ic centric algae were dominant in “dirty” ice (Fig. 1). Compared to the White Sea land-fast ice, pack ice in the White Sea had higher concentrations of fl agellates (Table 1). Sedimentation of algae was low in Guba Chupa (0.28 - 2.2 mg C m–2 day–1) and was dominated by plankton centric diatoms in February and March. In April, Nitzschia frigida was more abundant than centric diatoms. Planktonic centric diatoms dominated the vertical fl ux above the bottom in the deepest part of Kandalaksha Bay, but the per- manent occurrence of sea ice algae, even during the summer months, was remarkable. In contrast to the rather low abundance of phy- toplankton below the ice in the White Sea, in the Barents Sea, the biomass of phytoplankton was two orders of magnitude higher than the biomass of sea ice algae (Fig. 1). Flagellates dominated the sea ice algae and phytoplankton carbon in all samples from the Barents Sea (Table 1). They comprised 60 - 90 % of the total (1.1 - 3.9 mg C m–3 in the ice and 30 - 300 mg C m–3 in the water). In the lowermost part of the annual ice, centric diatoms were abundant (> 50 % of total carbon of diatoms and dinofl agellates). Plankton algae dominated the sea ice assem- blage and the vertical fl ux in the Barents Sea. In the water and sediment traps dinofl agellates were the second most abundant group. Only one sea ice species (Nitzschia frigida) was numerous in the traps. The composition of the sea ice algae pop- ulation in the multi-year ice was rather similar to that of the sea ice algae in the White Sea in April 2002: N. frigida and ribbon diatoms com- prised the bulk of the total diatom and dinofl ag- ellate carbon. However, in the water column of the Barents Sea, total carbon of diatoms was dominated by centric species, whereas in the White Sea it was dominated by pennates, N. frigida and ribbon species. Total algae concentration was higher in the water than in the ice in the Barents Sea, but the abundance of Melosira arctica and N. frigida was higher in the ice. The sediment trap content under the multi-year ice in the Barents Sea was rather similar to the algae composition in the ice, in contrast to the traps under the annual ice, where the species M. arctica and N. frigida were sparse. Discussion Sea ice properties The temperature of the water in the White Sea seldom reached the freezing point, and the ice developed mainly from the surface (Krell et al. 2003; Pantyulin 2003). The ice was thin during all expeditions (range: 34 - 57 cm) and was often fl ooded by seawater. In the White Sea, ice orig- inated mainly from snow: δ18O varied between –1.51 and –8.72 (Krell et al. 2003). The granu- lar snow ice dominated the entire ice column at all times in the White Sea. Only during the cold- est part of the year (March 2001, February 2002 and February 2003) was columnar ice observed in the lowermost layer of the ice. Congelation ice may only be found during the fi rst stages of ice development in the White Sea (Melnikov et al. in press). The ice melted from below because the under-ice water temperature never dropped below –1.5 °C in the course of our investiga- tions. The snow removed the bulk salinity from the brine channels and reduced the total sea ice salinity to psu 0 - 4 ‰. Thick snow cover insulat- ed the ice from the cold air, and the temperature in the ice was rather high (> –2 °C). Low salini- ty and the high temperature of the ice led to wide brine channels and low salinity in the brine water (Melnikov et al. in press). There are no data regarding the sea ice struc- ture in the investigated region of the Barents Sea. However, it is well known that congelation ice dominates the total ice column in the Barents Sea, as in other Arctic areas. Only the uppermost 10 cm may be represented by snow ice. Salinity of the brine water may be as high as psu 70 - 144 ‰ (Gradinger et al. 1999). In summer, the total salin- ity and temperature of the lowermost part of the ice (psu 3 - 5 ‰ and –1.2 oC) appear to be compa- rable with the White Sea ice. Fig 2 (opposite page). Vertical distribution of the sea ice dia- toms in the White Sea in March 2001, April 2002 and Feb- ruary 2003. “Barrel” refers to benthic single-celled diatoms forming barrel-shaped colonies in the ice; “Ribbon” is plank- ton diatoms forming ribbon-shaped colonies. The axes are the same within each column, but the left and right columns have different axes. Each bar represents a single sample. 102 Sea ice algae in the White and Barents seas Composition and abundance Most of the species that were observed in the sea ice were similar to those recorded from other parts in the Arctic. In the high latitude Arctic this sim- ilarity may be attributed to the long-range trans- port of ice (von Quillfeldt et al. 2003), but this explanation does not hold for the White Sea ice. The similarity between sea ice assemblages in the White Sea and in other ice-covered regions may be explained by a similar origin of the ice fl ora in all Arctic regions (Sazhin et al. 2004). Three types of sea ice organisms can be distinguished: ice specialists, ice-benthic and ice-plankton spe- cies. Additionally, plankton and benthic species may also be observed in sea ice. The sea ice spe- cialists were neither observed in bottom sedi- ments nor in plankton in summer, but they were found in the bottom water sediment trap, prob- ably due to resuspension and stirring up of the sediment. As soon as the ice melted, sea ice spe- cialists decayed or developed resting spores (e.g. Undatella cf. quadrata, Melosira arctica). Closely related to these species were ice-ben- thic species (e.g. Diploneis litoralis, Enthomon- eis spp., Navicula lineola), but these species were also observed in bottom sediments as active cells (Gogorev 1998). Benthic species may be includ- ed into the sea ice during its formation in shallow estuaries or bays. They may survive in the ice, but in contrast to the ice-benthic species, their abundance is not high (e.g. Mastogloia spp., Pin- nularia spp., Triblionella litoralis). Most of these ice-benthic and benthic species were observed in the White Sea bottom sediments in March 2001 (F. Sapozhnikov, pers. comm.) and in the plank- ton in the Pechora Sea during the ice formation in November 2003 (T. Ratkova, unpubl. data). These species were observed in the bottom water trap in the White Sea deep throughout the year (T. Ratk- ova, unpubl. data). Ice-benthic species were not abundant in pack ice, probably because they cannot recolonize ice from the bottom in deeper waters. The sea ice algae that inhabit the pack ice survive the summer in the water column (ice-plankton algae) or in the multi-year ice (sea ice specialists). The ice-plankton species were frequent in land-fast and pack ice (e.g. ribbon diatoms and Nitzschia frigida, accompanied with Atteya sep- tentrionalis, Synedropsis hyperborea and Gom- phonema-like species). The plankton centric diatom species may be incorporated into the ice during ice formation and during fl ood events (Buck et al. 1998). However, they were not abun- dant in the interior of the ice, but only in the upper and lower layers, where the ice is in contact with the underlying water or fl oodwater. Centric dia- toms are regarded as allochthonous for the ice realm. Thus, pennate diatoms dominate among the sea ice algae from the Chukchi, East Siberian and Laptev seas (Okolodkov 1992). The lower part of the annual sea ice in the northern Barents Sea was dominated by ice-plankton and plankton dia- toms. The sub-ice assemblage was dominated by ice-plankton (ribbon diatoms and Nitzschia frig- ida). The ice specialist Melosira arctica, the ice- benthic species (Enthomoneis spp., Diploneis lito- ralis) and associated species were also observed. Planktonic algae dominated the water and sed- iment traps. Only one sea ice species (N. frigi- da) was numerous in the traps. The sea ice com- munity of the annual ice in the northern Barents Sea included more planktonic algae than in the White Sea, but the same benthic species occurred in both areas. The diatom population in the lower part of the multi-year ice was dominated by ribbon diatoms and by N. frigida. M. arctica developed into thick mats below the ice. The plankton diatom Chae- toceros socialis dominated the water below the ice, but in the upper few metres M. arctica was also abundant. M. arctica was abundant in the sediment traps below the multi-year ice, in con- trast to those below annual ice, indicating that this species is primarily introduced to the Barents Sea from the Arctic Ocean. Many resting spores of M. arctica were observed in the water column and in sediment traps. Evidently, part of the M. arctica mats were detached from the lower sur- face of the ice. Coastal sea ice of the White Sea forms in pro- ductive systems where a continuous nutrient supply from the water column (rivers, sediment, mixing by tidal currents) sustains high algal bio- mass, with diatoms as the dominating taxon (Krell et al. 2003). Oceanic ice habitats of the Barents Sea, however, are characterized by more regener- ative food webs, with lower biomass and a higher contribution of fl agellates (Gradinger 1999). This illustrates the main difference between the White Sea land-fast ice and White and Barents seas pack ice. These two different scenarios con- tribute to the observed differences in composition and abundance of the ice algae in these two types of sea ice. 103Ratkova & Wassmann 2005: Polar Research 24(1–2), 95–110 The difference between the abundance of sea ice algae in the White Sea and in the Barents Sea may be attributed in part to the different timing of growth. In the northern Barents Sea the produc- tivity of sea ice algae is highest in July (Kuznetsov & Schoschina 2003), but in the White Sea it is in April (Ilyash et al. 2003). Our observations in the Barents Sea were conducted at the end of sea ice algae vegetation and before and during the max- imum of sea ice algae development in the White Sea. The species composition in the ice changes little with the age of the ice, whereas the relative abundance of each species may change marked- ly: plankton species, which are more abundant and dominate total algae numbers and carbon in young ice, may be replaced by ice species, which are more abundant in older ice. The composition of the sea ice algae results primarily from the com- position of algae in the underlying water masses in regions that are not ice-covered during a part of the year, or from the composition of the sea ice algae community in the ice-edge or in polynyas (Syvertsen 1991; Weissenberger 1998; Druzhkov et al. 2001). In shallow water areas benthic algae may also be included in the succession cycles (Poulin 1990; Gogorev 1998). The stickiness of pennate diatoms (Riebesell et al. 1991) may par- tially explain their high enrichment rates. Usual- ly the planktonic and benthic algae entrapped by the ice crystals in autumn (Syvertsen 1991; Mel- ni kov 1997; von Quillfeldt 1997) will be sparse during the entire winter. They start their develop- ment in spring (Zhitina & Mikhailovsky 1990). However, if sea ice forms earlier (in October in the northern Barents Sea), the entrapped algae may develop immediately (Horner 1990; Druzhk- ov et al. 2001), overwintering as a well-organized community. The timing of the development of the sea ice algae community can be explained mainly by the light conditions and the taxonomic com- position of the algae: benthic algae may be better adapted to lower irradiance and lower tempera- ture than planktonic ones (Gogorev 1998). The vertical distribution of sea ice algae The benthic algae entrapped by the ice crystals during ice formation in February 2001 may have developed immediately in the lower part of the thin granular ice because the light conditions in newly formed ice are favourable for algae growth. When the irradiance decreased after a heavy snowfall, they may have actively moved through the brine channels to the upper part of the ice. Consequently, on 26 March 2001 the highest bio- mass of these species was observed in the upper 10 cm of the ice. The same vertical distribution was also observed in February 2003, when the snow cover was exceptionally thick. The ice-plankton- ic and ice-benthic species may demonstrate new morphological peculiarities when they live in ice. Some of the species became longer (Fragilariop- sis cylindrus, F. oceanica, Nitzschia longissima); others (Enthomoneis spp, Undatella cf. quadrata and Navicula lineola) developed large barrel-like colonies. These morphological changes may indi- cate an adaptation to new conditions. In March 2001 we also observed an upward movement of the ice-benthic diatom Diploneis litoralis. It had its highest abundance in the lower 5 cm of the ice core on 18 March and was dis- placed to the upper 10 cm by 22–26 March. We speculate that the vertical species succession, which usually occurs over a time span of months (Syvertsen 1991), took place in a week in 2001 because of the late development of the sea ice. Conclusion A total of 306 algae species were identifi ed in ice, water and sediment traps in the Barents and White seas. In the White Sea land-fast ice, 275 species were recorded. In the White Sea pack ice, species were less numerous (106 species), and only four of these species were not observed in the land-fast ice. In the Barents Sea, 201 species were found. In general, the algae composition in the two regions was rather similar. The main differences were encountered among the pennate diatoms: in the White Sea land-fast ice, 110 pennate species were found, while only 49 and 52 species were found in the White Sea and Barents Sea pack ice, respectively. Most of the species were found both in the water column and in the ice, but a few spe- cies were not observed in the ice. The algae com- munities of the ice and ice-covered waters in the White and Barents seas are highly variable in space and time (Sazhin et al. 2004; von Quillfeldt et al. 2003). To understand the dynamics of these algae assemblages, one must take into account the formation history of the ice. The algae communities of the land-fast ice, dominated by pennate diatoms, are more closely related to the bottom sediments than the commu- 104 Sea ice algae in the White and Barents seas nity of the pack ice, which is dominated by centric diatoms. The algae in the land-fast ice communi- ty are more numerous and variable than those of the pack ice. Motile benthic and ice-benthic spe- cies fi nd favourable conditions in the ice because of the close proximity and similarity between the bottom sediments and the sea ice interior (Gogor- ev 1998). The plankton and ice-plankton species fi nd stable illumination and refuge from graz- ers in the ice. However, conditions here are less favourable for plankton algae, which are well- adapted to life in the water column, where algae circulate passively with the turbulent fl ow and where conditions for nutrient uptake are better. Algae living in ice, in contrast, must contend with the laminar fl ow in the narrow channels of the ice, where the nutrient supply in the upper layer may be very low (Gradinger & Ikävalko 1998). Benth- ic and ice-benthic ice algae are better adapted to such conditions, attaching themselves to the sub- strates or actively moving along the walls of the channels. The pack ice community is related to that in the surrounding water. It originates from it and is released into it after the ice melt. An addition- al source for ice algae colonization may be the multi-year ice. Algae may be released from the ice during warming events, because of the brine drainage, or during the sea ice melting. Some of the sea ice algae develop spores before the ice melt. These spores may be transported to other ice fl oes, or sink down to the bottom of shelf regions until the next season. Grazers may quickly ingest the released algae: in the White Sea almost none of the released diatoms reach the bottom. The sedimentation rate of the algae is therefore extremely low and a high rate of faecal pellet export was observed (Kosobokova et al. 2003). Acknowledgements.—We are grateful to E. Arashkevich for general discussions and to two anonymous referees for reading and commenting on the manuscript. The technical assistance of E. Halttunen, K. Riser, J. Søreide, A. Belov and A. Novi- gatsky is gratefully acknowledged. 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Moscow: Institute of Oceanology of the USSR Academy of Sciences. 106 Sea ice algae in the White and Barents seas Appendix The following table indicates the relative abun- dance, geographic ranges and ecology of the spe- cies recorded from ice cores and sub-ice water in the White Sea in February, March and April 2001– 03 and in the Barents Sea in July 2001. (Identifi - cations are based on Krishtofovich & Proshkina- Lavrenko [1949, 1950], Hendey [1964], Semina [1974], Semina & Sergeeva [1983], Medlin & Prid- dle [1991], Thomsen [1992], Snoeijs [1993], Snoe- ijs & Vilbaste [1994], Snoeijs & Potapova [1995], Snoeijs & Kasperovičienė [1996], von Quillfeldt [1996, 2000], Konovalova [1997], Tomas [1997], Okolodkov [1998], Snoeijs & Balashova [1998], Tuschling [2000], Ulanova [2003], Throndsen et al. [2003] and von Quillfeldt et al. [2003].) Geo- graphic ranges (in the column headed “Range”) are: cosmopolitan (c), Arctic–boreal (Ab), bipo- lar (b), tropical–Arctic–boreal (tAb), tropical– boreal (tb) and tropical (t). Ecological categories (“Ecology”) are: freshwater (F), brackish water (B), marine (M), eurihaline (E), neritic (N), ice– neritic (In), marine–neritic (Mn), epiphytic (Ep) and benthic (Be). Relative abundance (mean num- ber of cells for all available samples) categories are: 1 (< 1 104), 2 (1 104 - 1 105), 3 (1 105 - 1 106) and 4 (> 1 106 cells l–1. Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Division Chromophyta Class Bacillariophyceae Achnanthes brevipes Agardh 4000 c B, Be 1 - 1 - - A. fl exella (Kütz.) Brun 9000 - F, Be - - 1 - - Achnantidium minutissima (Kütz.) Czarm. 250 - F, Be 1 - 1 - - Actinocyclus curvatulus Ehrenberg 67 500 c M 1 - 1 - - A. octonarius Ehrenberg 48 000 c M 1 - - - - Amphora spp. 1 - 1 - 1 Asterionella formosa Hassal 800 c F - - 1 - - Attheya septentrionalis (Øestrup) Crawford 150 Ab M, Ep, In 3 2 1 3 1 Bacterosira bathyomphala (Gran) Syvertsen & Hasle 2200 Ab M, N 1 - 1 - 1 Caloneis sp. 1 - - - - Campylodiscus fastuosus Ehrenberg 42 000 Ab? M, Be - - 1 - - Chaetoceros affi nis Lauder 4600 Ab? M, N 1 - 1 - - C. borealis Bailey 11 250 tAb M - - 1 - - C. brevis Schütt 1500 Ab M, N - - - - 1 C. ceratosporus Ostenfeld 50 Ab M, N 2 - 1 - 2 C. compressus Lauder 600 tAb M 1 - 1 - 1 C. concavicornis Mangin 2200 Ab M 1 - 1 - - C. danicus Cleve 7500 tAb B, N 1 - 1 - - C. debilis Cleve 600 tAb M, N 1 - 1 - 1 C. decipiens Cleve 13 500 c M - - 1 - 1 C. diadema (Ehrenberg) Gran 3100 Ab M, N 1 - - 1 1 C. diversum Cleve 720 tb M, N - - 1 - - C. fallax Proshkina-Lavrenko 1100 - - 1 - - - - C. furcellatus Bailey 200 Ab M, N 1 - 1 - 1 C. gracilis Schütt 110 Ab M, N 1 - 1 1 1 C. holsaticus Schütt 720 Ab B, N 1 - - - - C. invisibilis Gogorev 30 - M, In 2 - 1 3 - C. laciniosus Schütt 400 tAb M, N 1 1 - - 1 C. perpusillus Cleve 300 tAb M, N - - - - 1 C. similis Cleve 300 Ab M, N 1 - - - - C. simplex Ostenfeld 350 tAb E, N 1 3 1 - 1 C. socialis Lauder 100 c E, N 2 3 3 3 3 C. wighamii Brightwell 600 Ab E, N 1 - - - - Cocconeis costata Gregory 4600 Ab? M, N 1 - 1 - - C. pediculus Ehrenberg 1440 - B, Ep - - 1 - - C. scutellum Ehrenberg 9000 c? M, N 1 - 1 - - C. stauroneiformis (Van Heurck) Okuno 4050 - B, Ep - - 1 - 1 Corethron criophyllum Cas- tracane 25 200 c M - - 1 - - Coscinodiscus asterom- phalus Ehrenberg 1012 500 c M - - 1 1 - C. centralis Ehrenberg 844 000 c M - - 1 - - C. concinnus W.Smith 1350 000 c M 1 - 1 - - C. radiatus Ehrenberg 42 300 c M - - 1 - - Ctenophora pulchella (Ralfs & Kützing) Williams & Round 6370 Ab E, Be 1 - 1 1 - Cyclotella choctawhatchea- na (Proshkina-Lavrenko) Prasad 1700 Ab E, N 1 - 1 1 - C. litoralis Lange & Syvert- sen 16 000 b E, N 1 - 1 - - C. striata (Kützing) Grunow 3000 Ab? E, N 1 - 1 - - Cylindropyxis tremulans Hendey 140 - - 1 - 1 - - Cylindrotheca closterium (Ehrenberg) Lewin & Rei- mann 450 c E, N 1 1 1 1 1 Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Table continued from previous column. 107Ratkova & Wassmann 2005: Polar Research 24(1–2), 95–110 Dactyliosolen fragilissimus (Bergon) Hasle 5100 c? - 1 - - - - Detonula confervacea (Cleve) Gran 270 Ab M, N - - 1 - 1 Diatoma tenuis Agardh 800 - F 1 - 1 - - Didymosphaenia geminata (Lyngb.) M.Schmidt 56 900 - F, Be - - 1 - - Diploneis didyma (Ehren- berg) Ehrenberg 10 000 Ab? B, N 1 - 1 - - D. litoralis var. litoralis (Øestrup) Cleve 2300 Ab M, N 2 - 1 1 1 D. litoralis var. arctica (Øestrup) Cleve 2700 Ab M, In 1 - - - - D. litoralis var. clathrata (Øestrup) Cleve 5300 Ab M, In 1 - 1 - - Ditylum brightwellii (West) Grunow 11 800 tb M 1 - 1 1 - Entomoneis alata (Ehrenberg) Poulin & Cardinal 3000 - - 1 - 1 1 1 E. kjelmanii (Cleve in Cleve & Grunow) Poulin & Car- dinal 78 000 Ab M, In 1 1 1 - - E. kjelmanii var. subtilis (Grunow) Poulin & Cardinal 16 000 Ab M, In 1 - 1 - - E. paludosa (W. Smith) Poulin & Cardinal 28 800 Ab B, In 1 - 1 1 - E. paludosa var. hyperbo- rea (Grunow in Cleve & Grunow) Poulin & Cardinal 29 400 Ab M, In 1 - 1 - - E. pseudopulex Osada & Kobayasi 23 600 - - 1 - - - - E. punctulata (Grunow) Osada & Kobayasi 18 700 Ab B, In 1 - 1 1 - Entomoneis sp. 25 200 1 - 1 - - Eucampia groenlandica Cleve 1500 Ab M, N 1 - 1 - 1 Fallacia forcipata (Greville) Stickle & Mann 23 520 c M 1 - 1 - - Fossula arctica Hasle, Syvert- sen & von Quillfeldt 1400 Ab M, In 2 1 1 3 1 Fragilaria striatula Lyngbye 2560 Ab M, In 2 1 1 1 - F. ulna (Nitzsch) Lange-Ber- talot 7300 - F 1 - 1 - - Fragilariopsis cylindrus (Grunow) Krieger 100 b M, In 2 1 1 2 1 F. oceanica (Cleve) Hasle 400 Ab M, In 2 1 1 1 1 Gomphonemopsis cf. exigua (Simonsen) Medlin 1920 Ab? M, Ep, In 2 1 1 1 1 Grammatophora arctica Cleve 12000 Ab? M, N 1 - 1 - - G. oceanica (Ehrenberg) Grunow 9000 Ab? M, N - - 1 - - Guinardia delicatula (Cleve) Hasle 7000 c M, N 1 - 1 - - Gyrosigma compactum Gre- ville 4300 - - 1 - 1 1 - G. fasciola var. tenuirostris (Grunow) Cleve 7000 Ab? M, N 1 - - - - G . hudsonii Poulin & Car- dinal 18 000 Ab M, In 1 1 - - - G. tenuissimum var. hyperbo- rea (Grunow) Cleve 3400 - M, N 1 - 1 1 - Hannaea arcus (Ehrenberg) Patrick 3200 - F - - 1 - - Hantzschia sp. - - 1 1 - Haslea sp. 1 - - - 1 Lauderia annulata Cleve 22 800 tb M 1 - - 1 1 Leptocylindrus danicus Cleve 1200 tAb? M, N 1 - 1 - 1 L. minimus Gran 600 tAb M 1 - 1 - - Licmophora communis (Heib- erg) Grunow 4100 Ab? M, N 1 - 1 - - L. gracilis (Ehrenberg) Grunow 18 200 - M, N 1 - 1 - - Mastogloia sp. 1 - 1 1 - Melosira arctica Dickie 1500 Ab M, In 1 - 1 2 1 M. moniliformis (O.F.Müller) Agardh 25 200 B, N 1 - - - - M. nummuloides Agardh 43 500 c M, N 1 - 1 - - Melosira sp. - - 1 - - Meridion circularis (Grev.) Agardh 400 - F - - 1 - - Meuniera membranacea (Cleve) P.C. Silva 15 000 Ab - 1 - - - - Navicula algida Grunow 120 000 Ab M, In 1 - - - N. directa (W.Smith) Ralfs 9000 Ab - 1 - 1 - - N. distans (W.Smith) Ralfs 56 300 Ab? Be 1 - 1 - - N. gelida Grunow 15 000 Ab M, In 1 - - - - N. glacialis Cleve 7300 Ab M, In 1 - - - - N. granii (Jørgensen) Gran 2700 Ab M, In 2 - 1 1 - N. cf. lineola Grunow 9050 Ab M, In 1 - 1 1 - N. menisculus var. meniscus (Schum.) Hust. 2200 - F 1 - - - - N. microcephala Grunow 540 - F 1 - - - - N. monilifera Cleve 10 000 Ab? M 1 - - - - N. pelagica Cleve 1100 Ab E, N 2 - 1 2 1 N. pellucidula Hustedt 7200 Ab M, In 1 - - - - N. septentrionalis (Grunow) Gran 800 Ab M, N 1 - 1 - - N. superba Cleve 9600 Ab M, In 1 - - - - N. transitans var. derasa (Grunow) Cleve 34 900 Ab - 1 - 1 - - N. vanhoeffenii Gran 2200 Ab E, N 1 - 1 - - Nitzschia angularis W.Smith 3200 - - - - 1 - - N. dissipata (Kützing) Grunow 2250 - F 1 - - - - Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Table continued from previous column. Table continued from previous column. 108 Sea ice algae in the White and Barents seas N. frigida Grunow 4500 Ab M, In 4 1 1 3 1 N. hybrida Grunow 2400 Ab B, In 1 - - - - N. longissima (Brebisson) Ralfs 830 b M, N 1 1 1 1 - N. neofrigida Medlin 19 000 Ab M, In 1 1 1 - - N. pellucida Grunow 1100 Ab? B 1 - - - - N. polaris Grunow 6500 Ab M, In 1 - - - - N. promare Medlin 560 Ab M, In 2 1 1 1 - N. recta Hantzsch. 1400 - F 1 - - - - N. scrabra Cleve 12 000 Ab M 1 1 1 - - N. sigmoidea (Ehrenberg) Wm. Smith 9000 - B 1 - 1 1 - Odontella aurita Agardh 10 500 c? B, N 1 - 1 - 1 Paralia sulcata (Ehrenberg) Cleve 3300 c? M, Be 1 - 1 1 - Pauliella taeniata (Grunow) Round et Basson 1000 Ab E, Mn 2 1 1 3 1 Pinnularia guadrataera var. baltica Grun 15 000 tAb M, N 1 - 1 - - P. semiinfl ata (Øestrup) Gran 5400 Ab M, In 1 - - - Plagiotropis lepidoptera (Gregory) Poulin & Cardinal 60 000 tAb M, N 1 - 1 - - P. scaligera Grunow in Cleve & Grunow 10 000 - - 1 - 1 1 - Planotidium delicatulum (Kützing) Round & Bukht. 200 Ab? E, N 1 - - - - Pleurosigma angulatum (Quekett) Wm.Smith 6000 tAb? B, N 1 1 1 1 - P. clevei Grunow in Cleve & Grunow 35 000 Ab M, In 1 - 1 1 - P. fi nmarchicum Cleve & Grunow 67 500 Ab? Mn 1 - 1 1 - P. formosum Wm.Smith 18 700 - - 1 - 1 - - P. normanii Ralfs 4500 c? - 1 1 1 1 - Porosira glacialis (Grunow) Jørgensen 38 800 b N 1 - 1 1 1 Proboscia eumorpha (Cast- racane) Takahashi, Jordan & Priddle 3200 c M, Mn - - - - 1 Pseudogomphonema kamt- chaticum (Grunow) Medlin 7200 - Ep 1 - 1 2 - P. septentrionale (Øestrup) Medlin 3200 Ab? M, Ep, In 1 - - - - Pseudo-nitzschia australis (Cleve) Heiden 3100 tb M, Mn 1 - 1 - 1 P. calliantha (Hasle) Lund- holm, Moestrup & Hasle 100 Ab M, Mn 1 - 1 1 - P. delicatissima (Cleve) Heiden 200 Ab M, Mn 1 - 1 - - P. granii (Hasle) Hasle 30 Ab? - 1 - 1 - 1 P. pungens (Grunow & Cleve) Hasle 1400 c M, Mn 1 - - - - P. seriata (Hasle) Hasle 3200 Ab M, Mn - - 1 - - P. seriata f. obtusa (Hasle) Hasle 2200 Ab - 1 - 1 - - Rhabdonema arcuatum (Lyng- bye) Kützing 49 500 Ab Ep 1 - 1 - - Rhizosolenia hebetata f. semi- spina (Hensen) Gran 37 500 Ab M, Mn - - 1 Rhizosolenia setigera Bright- well 18 000 c M, Mn 1 - - - - Rhoicosphenia curvata (Kütz- ing) Grunow 4000 - B, N 1 - 1 - - Skeletonema costatum (Gre- ville) Cleve 200 c M, Mn 2 1 1 2 1 Stauroneis amphioxys Gre- gory 24 000 Ab? N 1 - - - - Stenoneis sp. 1 - - - - Synedropsis hyperborea (Grunow) Hasle, Medlin & Syvertsen 270 Ab M, In 2 - 1 2 1 Synedropsis hypeboreoides Hasle, Medlin & Syvertsen 250 Ab M, In 1 - 1 1 - Tabellaria binalis (Ehren- berg) Grunow 4800 - F 1 - 1 - - T. fenestrata (Lyngbye) Kütz- ing 30 000 - F 1 - 1 1 - T. fl occulosa (Roth.) Kützing 18 700 - F - - 1 1 - Thalassionema nitzschioides (Grunow) Grunow & Hustedt 400 tAb M, Mn 1 1 1 - - Thalassiosira angulata (Gre- gory) Hasle 4300 Ab M, Mn 1 1 1 1 1 T. anguste-lineata (A.Schmidt) Fryxell & Hasle 12 000 tAb M, N 1 - 1 - 1 T. antarctica var. borealis Fryxell, Douc. & Hubb. 14 580 Ab M, Mn 1 1 1 1 1 T. bioculata (Grunow) Osten- feld 18 000 Ab? - 1 - - - - T. bulbosa Syvertsen 400 Ab - 1 - 1 - 1 T. conferta Hasle 2000 tb - 2 1 1 - - T. eccentrica (Ehrenberg) Cleve 23 400 tAb E, Mn - - 1 - - T. gravida Cleve 9000 b M, Mn 1 - 1 - - T. hyalina (Grunow) Gran 6500 Ab M, Mn 2 1 1 1 - T. hyperborea (Grunow) Hasle & Lange 25 000 Ab B, In 1 1 1 1 - T. nordenskioeldii Cleve 2900 Ab E, N 1 1 1 1 1 Trachyneis aspera (Ehren- berg) Cleve 160 000 Ab M, N 1 - 1 - - Tryblionella litoralis (Grunow) D.G.Mann 18 000 - M, Be 1 - 1 1 - Undatella cf. quadrata (Bréb. in Kütz.) Padd. et Sims 24 300 tAb? M, Be 1 - 1 1 - Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Table continued from previous column. Table continued from previous column. 109Ratkova & Wassmann 2005: Polar Research 24(1–2), 95–110 Class Dinophyceae Alexandrium insuetum Balech 1500 tAb? N 1 - 1 1 2 A. ostenfeldii (Paulsen) Balech & Tangen 147 000 tAb? N 1 - 1 - - A. tamarense (Lebour) Balech 19 000 tAb? N 1 - 1 1 - Amylax triacantha (Jør- gensen) Sournia 5000 Ab N - - 1 - - Amphidinium crassum Loh- mann 2300 Ab? N 1 - 1 1 1 A. fusiformis Martin 1050 Ab? N 1 - 1 1 1 A. larvale Lindemann 190 - E, N 1 - 1 - 1 A. latum Lohmann 1350 Ab? N - - - - 1 A. longum Lohmann 1300 c N 1 - 1 1 1 A. sphenoides Wulff 1100 Ab M 1 - 1 - 1 Ceratium fusus (Ehrenberg) Dujardin 91 800 c M 1 - 1 - - C. lineatum (Ehrenberg) Cleve 46 900 tb M - - 1 - - Cochlodinium archimedes (Pouchet) Lemmermann 1400 Ab M 1 - 1 1 1 Cochlodinium brandtii Wulff 8000 Ab Mn - - 1 1 - Cochlodinium schuetti (Kofoid & Swezy) Shiller 6000 - - 1 - 1 1 1 Dinophysis acuminata Cla- parède & Lachmann 19 000 c N - - 1 - - D. acuta Ehrenberg 18 000 b M 1 - - - - D. arctica Mereschkowsky 9000 b N - - 1 - - D. contracta (Kofoid & Skogsberg) Balech 6000 tAb M - - 1 - - D. islandica Paulsen 36 000 Ab N - - 1 - - D. norvegica Claparède & Lachmann 35 000 Ab N 1 - 1 - - Diplopelta parva (Abé) Mat- suoka 5900 Ab N 1 - 1 1 - Enthomosigma peridinioides Shiller 1200 tAb? N 1 - 1 1 - Glenodinium gymnodinium Penard 50 000 - B, N 1 - - - - Gonyaulax digitalis (Pouchet) Kofoid 40 000 b Mn - - 1 - - G. grindleyi Reinecke 18 000 Ab N 1 - 1 1 - G. spinifera (Claparède & Lachmann) Diesing 9900 c Mn 1 - 1 - - Gymnodinium albulum Lin- dermann 150 - B, N 1 1 1 1 2 G. arcticum Wulff 4000 Ab Mn 1 - 1 - 1 G. blax Harris 260 - F 1 - 1 - - G. frigidum Balech 13 500 b N 1 - - - 1 G. japonica Hada 400 - N 1 - 1 1 1 G. heterostriatum Kofoid & Swezy 4700 tAb? N - - 1 - - G. lebourii Pavillard 108 000 - - 1 - 1 - - G. simplex (Lohmann) Kofoid & Swezy 190 tAb N 1 - 1 - 1 G. wulfi i Schiller 4000 tAb N 1 - - - - Gyrodinium cf. aureolum Hulburt 24 000 Ab - 1 - - 1 1 G. cohnii (Seligo) Schiller 4500 Ab? - 1 - - - - G. esturiale Hulburt 1700 c N - - 1 - - G. fusiforme Kofoid & Swezy 1620 tAb Mn 1 - 1 1 1 G. lachryma (Meunier) Kofoid & Swezy 36 700 Ab Mn 1 - 1 - 1 G. prunus (Wulff) Lebour 47 000 c - 1 - - - 1 G. spirale (Berg) Kofoid & Swezy 17 000 c - 1 - 1 1 1 Heterocapsa rotundatum (Lohmann) Loeblich 400 c N 1 - 1 - 2 H. triquedrum (Lohmann) Hansen 6900 c B, N - - 1 - - Karenia brevis (Davis) G.Hansen & Moestrup 4000 Ab N 1 - - - - Karlodinium micrum (Lead- beater & Dodge) J.Larsen 1700 - - 1 1 1 - - K. venefi cum (Ballantine) J.Larsen 900 - - 1 1 1 1 1 K. vitiligo (Ballantine) J.Larsen 1700 - - 1 1 1 1 1 Katodinium glaucum (Lebour) Loeblich 2100 tAb N 1 - 1 1 - Micracanthodinium claytonii (Holmes) Dodge 3800 - - - - - - 1 Oblea baculifera Balech 1700 b - 1 - 1 - - Oxyrrhis marina Dujardin 6500 tAb? E, N 1 - - - - Oxytoxum belgicum Meunier 5250 - - - - 1 - - Peridinella catenata (Lev.) Balech 4700 Ab N 1 - 1 1 - Preperidinium meunieri (Pavillard) Elbrächter 10 000 t M 1 - 1 - 1 Pronoctiluca acuta (Lohm- ann) Schiller 1600 tAb Mn - - 1 - - P. pelagica Fabre-Domergue 9400 c M - - - 1 1 Prorocentrum balticum (Lohmann) Loeblich 400 c M 1 - 1 1 1 P. cordatum (Ostefeld) Dodge 1600 tAb N 1 - 1 - 1 P. micans Ehrenberg 6000 tAb N 1 - 1 - - P. minimum (Pavillard) Schiller 720 c N 1 - 1 - - Protoperidinium bipes (Paulsen) Balech 2000 c N 1 - 1 - 1 P. brevipes (Paulsen) Balech 10 100 Ab N 1 - 1 - 1 P. depressum (Bailey) Balech 130 000 c M - - 1 - - P. granii (Ostefeld) Balech 66 000 c N 1 - - 1 - P. islandicum (Paulsen) Balech 36 000 Ab N - - - - 1 Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Table continued from previous column. Table continued from previous column. 110 Sea ice algae in the White and Barents seas P. nudum (Meunier) Balech 9400 Ab Mn 1 - 1 - - P. pallidum (Ostenfeld) Balech 13 500 b M, N - - 1 - - P. pellucidum Bergh 51 800 c N - - 1 - 1 P. pyriforme subsp. pyriforme (Paulsen) Balech 24 000 c - - - 1 - - P. pyriforme subsp. breve (Paulsen) Balech 23 300 Ab? - - - - - 1 P. subinerme (Paulsen) Loe- blich III 62 500 c - 1 - - - 1 Scrippsiella trochoidea (Stein) Loeblich III 5900 c N 1 1 1 1 1 Torodinium robustum Kofoid & Swezy 4500 - - 1 - 1 - 1 Warnowia maculata (Kofoid & Swezy) Lindemann 15 750 - - - - - - 1 Woloszynkia reticulata Thompson 1700 - B, N - - - 1 1 Class Prymnesiophyceae Corimbellus aureus Green 260 - Mn 1 - 1 - - Emiliania huxleyi (Lohmann) Hay & Mohler 400 c - 1 2 1 2 1 Zigosphaera massilii (Borset- ti & Cati) Heimdal 700 - - 1 - 1 - 1 Phaeocystis pouchetii (Hariot) Lagerheim 35 b M 3 4 3 3 4 Primnesium sp. 1 - - - 2 Class Cryptophyceae Chroomonas marina (Büt- tner) Butcher 1100 - N 1 - 1 - 1 Hilea fusiformis (Schiller) Schiller 64 tAb? N 1 - 2 1 2 H. marina Butcher 18 tAb? N 2 2 3 1 1 Leucocryptos marina (Braarud) Butcher 480 Ab Mn - - 1 - 1 Plagioselmis prolonga Butch- er 750 - - 1 1 1 - 1 Teleaulax acuta (Butcher) Hill 340 - Mn 1 1 1 - 2 Class Chrysophyceae Calicomonas gracilis 20 - - 2 2 2 - - Calicomonas sp. - - - - 3 Dinobryon balticum (Schütt) Lemmermann 800 Ab M 1 - 2 2 3 D. belgica Meunier 600 Ab M 1 - 1 - 1 D. faculiferum (Willén) Willén 260 Ab Mn 1 - 1 - 1 Ochromonas crenata Klebs 300 - B, N 1 1 1 3 3 O. marina Lackey 480 Ab N - - 1 - 1 Class Dictyochophyceae Dictyocha fi bula Ehrenberg 5500 tAb? M 1 - 1 - - D. speculum Ehrenberg 6900 c E, N - - 1 1 1 Division Chlorophyta Class Chlorophyceae Carteria sp. 1 - 1 - 2 Chlamidomonas sp. 1 2 1 - - Tetraselmis sp. 1 - - - - Class Euglenophyceae 1 Eutreptiella braarudii Throndsen 7500 Ab N 1 - 1 1 1 E. eupharyngea Moestrup & Norris 600 Ab N 1 1 1 1 1 E. gymnastica Throndsen 2000 - N 1 - 1 2 1 E. hirudoidea Butcher 560 - N 1 - - - - Eutreptia sp. - - 1 1 - Euglena acus 12000 - - 1 - 1 1 - Class Prasinophyceae Halosphaera viridis Schmitz 171 500 c - 1 - 1 - 1 Micromonas pusilla (Butcher) Manton & Parke 4 tAb? Mn - - - - 2 Pachysphaera marshalia Parke 100 tAb? M 1 - - - 1 Pterosperma vanhoffenii (Jør- gensen) Ostenfeld 32 000 - Mn 1 - 1 - - Pyramimonas grossii Parke 50 c? - 2 1 1 2 2 P. orientalis McFadden, Hill & Wetherbee 30 tAb? N 1 - 2 - 1 Resultor micron (Throndsen) Moestrup 4 - N - - - 1 - Cyanobacteria Anabaenopsis sp. - - 1 - - Synechococcus sp. - - 1 - 2 Phylum Zoomastigophora Class Kinetoplastida Telonema subtilis Griessmann 300 - N 1 - 1 1 1 Class Choanofl agellidea Calliacantha natans (Grøntved) Leadbeater 32 Ab Mn 2 2 2 - - Parvicorbicula socialis ? (Meunier) Defl andre 20 Ab N 3 4 3 2 3 Monosiga marina Grøntved 100 tAb? M 3 3 2 - 2 Pleurasiga reinoldsii Thrond- sen 6900 tAb? N - - 1 - - Class Raphidophyceae Heterosigma akashiwo (Hada) Hada 768 tAb? b, N 1 - 1 3 1 Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Abundance White Sea Bar. Sea C el l v ol . (m µ3 ) R an ge E co lo gy Ic e Sn ow W at er Ic e W at er Table continued from previous column. 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