Spatial distribution of digestive enzyme activities of Calanus finmarchicus and C.hyperboreus in Fram Strait/Greenland Sea

Digestive enzymes of copepodite V (CV) Calanus finmarchicusand C.hyperboreus from two different depths were compared duringMIZEX 1984 (June/July) at stations in the ice and open water.CV of both species from 500–200 m showed reduced enzymeactivity, indicating that they were in a resting stage. In moultingexperiments at the end of June using CV from 100–0 m moultingwas delayed and began only after 3 weeks in C.finmarchicus andafter 3 months in C.hyperboreus. These results suggest thatthe deep CV populations are the seed of the new overwinteringstock. In surface CV C.finmarchicus and C. hyperboreus enzymeactivities were generally much higher than in deep CV. In neitherspecies were enzyme activities correlated with chlorophyll concentrations.Activities in C.finmarchicus reflected overall phytoplanktondistribution and were highest in the marginal ice zone, whereasthey decreased under the ice except for polynya stations onthe East Greenland Shelf. In surface C.hyperboreus digestiveenzymes were not correlated with those of C.finmarchicus, pointingto different diets or regulatory mechanisms. Enzyme activitywas lowest in the marginal ice zone and increased under theice. High activities were found at polynya stations and otherclose pack ice. The utilization of ice algae by C.hyperboreuscould explain these discrepancies in digestive enzyme activitiesof both species.     

Ice algal assemblages and vertical export of organic matter from sea ice in the Barents Sea and Nansen Basin (Arctic Ocean)

Algal communities and export of organic matter from sea ice were studied in the offshore marginal ice zone (MIZ) of the northern Barents Sea and Nansen Basin of the Arctic Ocean north of Svalbard by means of ice cores and short-term deployed sediment traps. The observations cover a total of ten stations within the drifting pack ice, visited over a period of 3 years during the period of ice melt in May and July. Maximum flux of particulate organic carbon and chlorophyll a from the ice at 1 m depth (1,537 mg C m⁻² per day and 20 mg Chl a m⁻² per day) exceeded the flux at 30 m by a factor of 2 during spring, a pattern that was reversed later in the season. Although diatoms dominated the ice-associated algal biomass, flagellates at times revealed similarly high biomass and typically dominated the exported algal carbon. Importance of flagellates to the vertical flux increased as melting progressed, whereas diatoms made the highest contribution during the early melting stage. High export of ice-derived organic matter and phytoplankton took place simultaneously in the offshore MIZ, likely as a consequence of ice drift dynamics and the mosaic structure of ice-covered and open water characteristic of this region.     

The North East Water polynya (Greenland Sea)

Physical observations of the North East Water (NEW) polynya, located near the north-eastern corner of Greenland, are presented. Data were collected in June 1991 by RV Polarstern. An idea is put forward to explain how the NEW is generated. A northward coastal current interacts with a persistent shelf ice barrier under which water can flow but that retains ice floes and therefore protects the NEW area from ice advection. Since in summer, the combination of currents, barrier and air-sea heat balance gives rise to a polynya. The distribution of upper water column vertical stability in the NEW is also influenced by its generation process. Surface melt water is retained by the shelf ice barrier, causing neutral vertical stability in its lee. Sea ice melting and land runoff then act as two distinct sources of vertical stability enabling the development of plankton blooms, especially in the northern part of the NEW.     

Spatial distribution and life-cycle timing of zooplankton in the marginal ice zone of the Barents Sea during the summer melt season in 1995

The marginal ice zone (MIZ) in the northern Barents Sea is ecologically important because it represents a highly productive area in Arctic water masses north of the Polar Front. During a multi-disciplinary cruise in 1995, ecological and oceanographic processes were investigated at four stations located in a north-south transect in the MIZ. This study was carried out in Arctic water masses north of the Polar Front where ice conditions varied from dense first-year pack ice to open water. Also, the phytoplankton development varied along the transect from a pre-bloom situation at the northern-most station to a post-bloom situation in the open water. This paper includes a study of the zooplankton community and population structure of some of the dominant copepod species. Numerically, the most important mesozooplankton components were the copepods Calanus glacialis, Pseudocalanus minutus and Oithona similis. Copepods of Atlantic origin, such as Calanus finmarchicus and Oithona atlantica, gave evidence of an advection of Atlantic water masses into the area. It is concluded that the occurrence of new cohorts of Arctic copepods coincides with the onset of the phytoplankton bloom in the MIZ, and, that therefore, the spawning relies on stored energy.      

Pelagic microbial activity in the Northeast Water polynya, summer 1992

 Heterotrophic activity and related measures of pelagic microorganisms in the Northeast Water (NEW) polynya, the largest and northernmost summer polynya in the Arctic, were studied during the Polar Sea cruise of July/August 1992 at stations spanning a range of ice conditions. Utilization (incorporation and respiration) of ¹⁴C-labelled amino acids was measured at in situ temperature as a proxy for microheterotrophic activity in samples from the chlorophyll maxima, intermediate water depths, and the benthic boundary layer. Total activity and bacterial abundance (measured by epifluorescence microscopy) were highest in the sub-zero surface water layer that dominates the northeast Greenland shelf, and particularly in areas most influenced by ice-edge processes and lateral advection. In this cold-water layer activity correlated strongly with particulate organic carbon (POC), to a lesser degree with phytoplankton-derived pigments and bacterial abundance, but not with median bacterial cell size. No significant correlations were detected in the warmer, Atlantic-influenced deep waters of the regional trough system where POC concentrations and other parameters were low. In comparison with temperate environments, pelagic heterotrophic activity in the NEW polynya appears to be low, a finding that bears upon the fate of dissolved organic matter and the efficient supply of particulate food to the benthos in this unique polar setting. Received: 3 December 1995/Accepted:29 May 1996     

Movements of female polar bears (<Emphasis Type="Italic">Ursus maritimus</Emphasis>) in the East Greenland pack ice

The movements of two adult female polar bears ( Ursus maritimus) in East Greenland and the Greenland Sea area were studied by use of satellite telemetry between the fall of 1994 and the summer of 1998. One female was tracked for 621 days, the other for 1,415 days. During this time the females used maternity dens on land. If denning periods on land were excluded, the two females used between 73% and 100% of the tracking time offshore where they were able to navigate in the dynamic pack ice and counteract the fast southward movement of the ice (up to 30 km/h) in the East Greenland Current. Mean monthly movement rates varied between 0.32 and 0.76km/h. Both bears had very large home ranges (242,000 and 468,000 km ²) within the dynamic pack ice of the Greenland Sea. The facts that the bears made extensive use of the offshore sea ice and that there is a marked reduction of the Greenland Sea ice call for a closer monitoring of the effects of this change on the East Greenland polar bear population.     

Satellite tracking of high-arctic northern fulmars

Three northern fulmars (Fulmarus glacialis) were tracked with satellite transmitters while they dispersed from a colony at the North East Water polynya in high-arctic Greenland after breeding failure. The longest cumulative distance recorded was 2043 km in 14 days, giving an average daily movement of 143 km, and the maximum distance covered in 1 day was 369 km. The highest effective flight (ground) speed recorded was 25.8 kmh^–1, and when corrected for non-linear flight path the ground speeds ranged between 27.9 and 38.4 kmh^–1, which is very close to the theoretical most energy efficient airspeed (V_mp) of approximately 36 kmh^–1. Bird flight tracks generally followed ice edges, and on long flights the birds used tail or cross winds. Within areas believed to be foraging areas birds moved cumulative distances of 86–488 km, or 33–147 kmday^–1. While in the high-arctic, the birds appeared to select foraging areas mainly in the marginal ice zones, but all birds left the polynya following breeding failure and moved towards boreal waters. One bird was tracked to the polar front zone near Bear Island where an international fishing fleet operated. The study suggests that when the birds were no longer attached to the North East Water polynya by a breeding attempt, they sought alternative foraging grounds. Satellite transmitters or other devices were tested on a total of eight birds' (including the three tracked individuals), all of which probably failed in their breeding attempt. The birds' reactions to the handling and tagging are presented, and the possible reasons for the breeding failure discussed.     

Phytoplankton in the Marginal Ice Zone of the Greenland Sea during summer, 1984

Summary Phytoplankton biomass and species composition were studied in transects through the ice edge region of the Greeland Sea from 19 July to 8 August 1984. Biomass was estimated by vertical in situ chlorophyll fluorescence and pigment extraction of discrete samples. Preserved material was used for identification of phytoplankton species and calculation of their relative abundances. The results suggest that the various geographical regions of the Greenland Sea differ considerably in their phytoplankton development. Autotrophic biomass and species composition were closely associated with the extent of the annual and seasonal ice cover, hydrographic conditions, nutrient availability and the water masses typical of the different domains. In the NE Greenland polynya a deep mixed layer inhibited the development of a phytoplankton bloom, whereas greatest biomass concentrations were associated with a receding ice edge on the E Greenland Shelf. In the Fram Strait, the position of the relatively stationary ice edge is controlled by frontal dynamics, currents and wind. Due to rapidly changing physical and chemical conditions, phytoplankton biomass showed great variability between stations. High chlorophyll a concentrations may develop locally where melting ice causes stratification or can result from passive accumulation in eddies. In July/August 84 the Fram Strait area was dominated by a typical summer population of flagellates and large diatom species.Contribution 6 of the Alfred-Wegener-Institute for Polar and Marine Research     

EFFECTS OF ENVIRONMENTAL VARIATION ON SINKING RATES OF MARINE PHYTOPLANKTON1

The effects of environmental variables, particularly irradiance, on the sinking rates of phytoplankton were investigated using cultures of Chaetoceros gracilis Schütt and C. flexuosum Mangin in laboratory experiments; these data were compared with results from assemblages in the open ocean and marginal ice zone of the Greenland Sea. In culture experiments both the irradiance under which the diatom was grown and culture growth rate were positively correlated with sinking rates. Sinking rates (ψ) in the Greenland Sea were smallest when determined from chlorophyll (mean ψ_chl= 0.14 m · d^?1) and biogenic silica (ψ_si= 0.14 m · d^?1) and greatest when determined from particulate carbon (ψ_c= 0.55 m · d^?1) and nitrogen (ψ_N= 0.64 m · d^?1). Field measurements indicated that variations in sinking may be associated with changes in irradiance and nitrate concentrations. Because these factors do not directly affect water density, they must be inducing physiological changes in the cell which affect buoyancy. Although a direct response to a single environmental variable was not always evident, sinking rates were positively correlated with growth rates in the marginal ice zone, further indicating a connection to physiological processes. Estimats of carbon flux at stations with vertically mixed euphotic zones indicated that approximately 30% of the daily primary production sank from the euphotic zone in the form of small particulates. Calculated carbon flux tended to increase with primary productivity.     

CETACEAN OCCURRENCE PATTERNS IN THE AMUNDSEN AND SOUTHERN BELLINGSHAUSEN SEA SECTOR, SOUTHERN OCEAN

We conducted 239.5 h and 3,494 km of cetacean surveys in the Amundsen and Bellingshausen seas, from 15 February to 31 March 1994; most of the area, the large portion of which was ice covered, had never before nor has it since been surveyed for cetaceans, even to the date when this paper was prepared (2006). Logistic regression and an information-theoretic approach related the occurrence of Antarctic minke whales Balaenoptera bonaerensis (the most abundant species) to whether we were in open- or pack-ice-covered pelagic or neritic waters, in or out of the marginal ice zone (MIZ), and north or south of the Antarctic Circumpolar Current southern boundary. Other variables included date and distance to the MIZ and shelfbreak front. Statistical analysis showed that the probability of sighting a minke, as well as killer whale—but not the case for an index to whale density—was related to the proximity of coastal polynyas in early autumn, switching offshore to the MIZ once waters within the pack began to freeze persistently later in the season. Probability of detection was higher with distance into the MIZ. Supporting these findings, the density index was strongly related to ice concentration in an inverse relationship. The strong relationship to polynyas and the MIZ indicate that sea-ice divergence altered by decadal or longer-term climate change, as described in the recent literature, could well affect any apparent, long-term trends evident in this species' abundance if surveyed only in open or near-to-ice waters. We speculate on how the minke whale's pagophilic nature (1) could have been encouraged by large-scale industrial whaling and by competition with species more characteristic of open waters and the outer MIZ, and (2) may have protected the population somewhat during industrial whaling resulting in the much greater abundance of this species now compared to other targeted species.     

Hypotheses and tracking results about the longest migration: The case of the arctic tern

The arctic tern Sterna paradisaea completes the longest known annual return migration on Earth, traveling between breeding sites in the northern arctic and temperate regions and survival/molt areas in the Antarctic pack‐ice zone. Salomonsen (1967, Biologiske Meddelelser, Copenhagen Danske Videnskabernes Selskab, 24 , 1) put forward a hypothetical comprehensive interpretation of this global migration pattern, suggesting food distribution, wind patterns, sea ice distribution, and molt habits as key ecological and evolutionary determinants. We used light‐level geolocators to record 12 annual journeys by eight individuals of arctic terns breeding in the Baltic Sea. Migration cycles were evaluated in light of Salomonsen's hypotheses and compared with results from geolocator studies of arctic tern populations from Greenland, Netherlands, and Alaska. The Baltic terns completed a 50,000 km annual migration circuit, exploiting ocean regions of high productivity in the North Atlantic, Benguela Current, and the Indian Ocean between southern Africa and Australia (sometimes including the Tasman Sea). They arrived about 1 November in the Antarctic zone at far easterly longitudes (in one case even at the Ross Sea) subsequently moving westward across 120–220 degrees of longitude toward the Weddell Sea region. They departed from here in mid‐March on a fast spring migration up the Atlantic Ocean. The geolocator data revealed unexpected segregation in time and space between tern populations in the same flyway. Terns from the Baltic and Netherlands traveled earlier and to significantly more easterly longitudes in the Indian Ocean and Antarctic zone than terns from Greenland. We suggest an adaptive explanation for this pattern. The global migration system of the arctic tern offers an extraordinary possibility to understand adaptive values and constraints in complex pelagic life cycles, as determined by environmental conditions (marine productivity, wind patterns, low‐pressure trajectories, pack‐ice distribution), inherent factors (flight performance, molt, flocking), and effects of predation/piracy and competition.     

Biogeographic structure of the microphytoplankton assemblages of the south Atlantic and Southern Ocean during austral summer

Microphytoplankton distribution in the Atlantic sector of theSouthern Ocean was investigated along a transect during theSAAMES II cruise undertaken in late austral summer (January/February) 1993. Samples were collected at 60 km intervals between34 and 70°S for the analysis of mineral nutrients, and theidentification and enumeration of microphytoplankton. Peaksin microphytoplankton abundance were recorded in the neriticwaters of Africa and Antarctica, at all major oceanic fronts,and in the marginal ice zone (MIZ). Partial correlation analysisindicated that 45% of the total variance associated with microphytoplanktonabundance could be explained by silicate and phosphate concentrations,while temperature accounted for 65% (P<0.001). Cluster andordination analyses identified two major groups of stations,one north and one south of the Subantarctic Front (SAF). Thisdivision appears to be related to differences in temperatureand silicate concentrations. Each region comprised distinctmicrophytoplankton subgroups associated with specific watermasses or hydrological features. Indicator species could beidentified for some water masses. In the MIZ, microphytoplanktonspecies composition and succession were strongly affected bysea-ice throughout the summer.     

Small nanoplankton and bacteria in the Western Ross Sea during sea-ice retreat (spring 1994)

Spatial changes of small nanoplankton (2–10 μm) were investigated in relation to sea-ice conditions, hydrography and receding ice processes in the Ross Sea (Antarctica) during spring 1994. Abundance and biomass of heterotrophic and autotrophic nanoplankton, as well as bacterioplankton, were determined along a south-north transect from the open waters polynya towards the pack ice. Autotrophic and heterotrophic nanoplankton biomass ranged from 758 to 4570 mgC m⁻² and from 3 to 387 mgC m⁻², respectively. Heterotrophic nanoplankton accounted, on average, for about 9% of the total (i.e. autotrophic plus heterotrophic) nanoplankton biomass. The size structure of both auto- and heterotrophic nanoplankton in the Ross Sea continental shelf receding ice edge was different from that of nanoplankton associated with the shelf break and open Antarctic ice-edge area. Generally, the highest heterotrophic biomass was found in the pack-ice zone on the continental shelf, while the highest heterotrophic contribution to the total nanoplankton biomass (up to 25%) was encountered at the shelf break where phytoplankton was largely dominated by 2- to 3-μm-size cells. Accepted: 2 May 1999     

Response of nematode communities after large‐scale ice‐shelf collapse events in the Antarctic Larsen area

Owing to large‐scale ice‐shelf disintegration events, the Antarctic Larsen A and B areas recently became ice‐free. During the ANT‐XXIII/8 Polarstern campaign, this region was sampled for the first time. Our study is the first to investigate benthic communities in this area and their response to the collapse of ice shelves in the Antarctic. The nematofauna appears to be strongly influenced by the sudden ice‐cover removal, although its response differs from that of the macro‐ and megabenthos. Our results indicate that precollapse, sub‐ice communities were impoverished and characterized by low densities, low diversity and high dominance of a few taxa. This might still be visible at a station located deep inside the Larsen B embayment, where Halomonhystera was dominant. Post‐collapse recolonization of the ‘inner’ stations, i.e. those located furthermost from the former ice‐shelf edge, is believed to be a long‐time process. At the time of sampling, community structure at the inner stations was not or only slightly influenced by colonization, and might be structured by local environmental conditions. Our results indicate that a locally increased food supply after ice‐cover removal could provoke a faster, local response of the nematode assemblages compared with the response due to recolonization. Thalassomonhystera is recognized as an opportunist, taking advantage of increased food supply at inner stations A_South and B_North. Communities living close to the former ice‐shelf edge are believed to be at an intermediate or late stage of succession, with a dominance of Microlaimus, a common Antarctic genus and quick colonizer. Densities here were comparable with those at other Antarctic stations, whereas they were considerably decreased at the inner stations. In general, the collapse of the Larsen ice shelves initially has a positive effect on the shelf nematode fauna in the area, both in terms of abundance and diversity.     

Export or retention? Copepod abundance,faecal pellet production and vertical flux in the marginal ice zone through snap shots from the northern Barents Sea

The balance between faecal pellet (FP) production and destruction that accelerates or diminishes vertical export has an effect on pelagic-benthic coupling, but is inadequately known. Production, export and retention of copepod FP were investigated in the marginal ice zone (MIZ) of the northern Barents Sea in July 2003. Older stages of Calanus finmarchicus and C. glacialis dominated the copepod biomass and FP production experiments revealed that more than 90% of the FP were produced in the upper 50 m where most of the copepods were located both day and night. Copepod pellets typically made up ∼10% of the vertical particulate organic carbon flux, and significantly less than what was produced by the copepod community. This implies a variable but significant retention of pellets. We suggest that retention of FP is caused partly by the zooplankton themselves and that retention of FP is the rule rather than the exception in the Barents Sea, particularly during non-bloom scenarios.     

Magnitude of mesozooplankton variability: a case study from the Marginal Ice Zone of the Barents Sea in spring

Zooplankton was studied on eight stations in the marginal icezone (MIZ) of the Barents Sea, in May 1999, along two transectsacross the ice edge. On each station, physical background measurementsand zooplankton samples were taken every 6 h over a 24 h periodat five discrete depth intervals. Cluster analysis revealedseparation of open water stations from all ice stations as wellas high similarity level among replicates belonging to particularstation. Based on five replicates per station, analysis of variance(ANOVA) confirmed significant differences (P < 0.05) in abundancesof the main mesozooplankton taxa among stations. Relations betweenthe zooplankton community and environmental parameters wereestablished using redundancy analysis (CANOCO). In total, 55%of mesozooplankton variability within studied area was explainedby eight variables with significant conditional effects: depthstratum, fluorescence, temperature, salinity, bottom depth,latitude, bloom situation, and ice concentration. GLM modelssupported supposition about clear and negative relationshipbetween concentration of Oithona similis, and overall mesozooplanktondiversity. The analyses showed a dynamic relationship betweenmesozooplankton distribution and hydrological conditions onshort-term scale. Furthermore, our study demonstrated that variabilityin the physical environment of dynamic MIZ of the Barents Seahas measurable effect on the Arctic pelagic ecosystem.     

Protists in the marine ice of the Amery Ice Shelf, East Antarctica

Samples of marine ice were collected from the Amery Ice Shelf, a large embayed ice shelf in East Antarctica, during the Austral summer of 2001–2002. The samples came from a site ∼90 km from the iceberg calving front of the shelf, where the ice is 479 m thick and the lower 203 m is composed of accreted marine ice. Protists identified within the marine ice layer of the Amery Ice Shelf include diatoms, chrysophytes, silicoflagellates and dinoflagellates. The numerical dominance of sea ice indicator diatoms such as Fragilariopsis curta, Fragilariopsis cylindrus, Fragilariopsis rhombica and Chaetoceros resting spores, and the presence of cold open water diatoms such as Fragilariopsis kerguelensis and species of Thalassiosira suggest the protist composition of the Amery marine ice is attributable to seeding from melting pack and/or fast ice protist communities in the highly productive waters of Prydz Bay to the north.     

Distribution,abundance and biology of polar cod, <Emphasis Type="Italic">Boreogadus saida</Emphasis>, in Iceland–East Greenland waters

Recent warming has caused a northern extension in the distribution of many southern fish species in Icelandic waters. Polar cod (Boreogadus saida) around Iceland are near the southern limit of their distribution, but are poorly studied in the area. Therefore, material sampled during demersal fish surveys in March 1985–2013 and in pelagic 0-group surveys in Iceland–East Greenland waters in August–September 1974–2003 was used to investigate their distribution, abundance and biology. Demersal polar cod were most often caught on the outer shelf to the north-west and north of Iceland, but during years of widest distribution and highest abundance, they were caught farther to the east on the northern shelf. Pelagic 0-group polar cod were only caught sporadically and mainly confined to the waters off the north-west shelf of Iceland and the East Greenland shelf (southern Denmark Strait). In demersal hauls, the number of stations with polar cod decreased with increasing bottom temperature and polar cod were most widely distributed in the years 1989, 1994 and 1995. Highest numbers of demersal polar cod per haul were caught at temperatures of ?0.5 to 2.5 °C and at 200–450 m depth. The length of demersal polar cod ranged from 5 to 32 cm, while the fish caught in the pelagic trawl ranged from 2.2 to 19 cm. The polar cod in the subarctic waters north of Iceland most likely originate from the waters off East Greenland and further warming and decline in sea ice may eventually lead to the disappearance of polar cod from Icelandic waters.     

Antarctic Phytoplankton Assemblages in the Marginal Ice Zone of the Northwestern Weddell Sea

The waters around the northern tip of the Antarctic Peninsulashow complex patterns of water circulation due to mixing ofdiverse water masses. Physicochemical properties of the differentwater types should affect the distribution, biomass and speciescomposition of the phytoplankton assemblages. We examined thesefeatures in the marginal ice zone (MIZ) of the northwesternWeddell Sea. Areas with the higher biomass were located in theWeddell Sea MIZ where the surface waters were relatively stabledue to the sea-ice melting. In these waters, the colonial stageof Phaeocystis antarctica and micro-sized chain-forming diatomsaccounted for 70% of the total phytoplankton carbon. Watersin the Bransfield Strait region, in contrast, were characterizedby a dominance of nanoflagellates, which accounted for 80% ofthe total phytoplankton carbon. Our observations support thehypothesis that the species composition of phytoplankton communitiesis a function of the different water mass, reflecting the physicalconditions of the upper water column, particularly its stability.     

Phytoplankton uptake of 15N and 14C in the Ross Sea during austral spring 1994

In spring 1994, within the ROSSMIZE research project, combined measurements of nitrogen (¹⁵N) and carbon (¹⁴C) uptake were made in the Ross Sea, passing from the McMurdo polynya to the ice-covered area in the north, in order to study the effect of environmental conditions (light availability, ice cover, vertical stability) on the coupling of N and C cycles. Nitrogen (nitrate and ammonium) and carbon uptakes were measured under simulated in situ conditions. The obtained results revealed, in most situations, much higher C:N uptake ratios than the Redfield ratio for phytoplankton composition; only in the inner part of the pack ice C:N uptake was lower than the balanced composition ratio. The high uptake ratios are ascribed to a larger C requirement during early phases of bloom evolution and to a greater importance of nitrogen sources, such as urea and other dissolved organic compounds, which were not measured in this study. In contrast, the lower C:N ratios in most of the pack-ice environment are ascribed to reduced photosynthesis in comparison to nutrient assimilation at low irradiances and to an increased importance of bacterial processes. Accepted: 3 January 2000