Seasonal shifts in ice edge phytoplankton blooms in the Barents Sea related to the water column stability

The development of the phytoplankton bloom and its relation to water column stabilisation during the transition from early to high summer (of 1991) in the seasonally ice-covered zone of the Barents Sea were studied from a meridional transect of repeated hydrographic/biological stations. The water column stabilisation is described in detail with the aid of vertical profiles of the Brunt-Väisälä frequency squared (N²). The contributions of seasonal warming and ice melting to stabilisation are elucidated by determining the effects of temperature and salinity on N². The spring bloom in 1991 migrated poleward from June to July by about 400 km, associated with the retreat of the ice edge. The spring bloom culminated with maximum chlorophyll concentrations in the mixed layer about 100–300 km north of the centre of the meltwater lens, at its northern edge, where the ice cover was still substantial. From the distribution of N² it becomes obvious that the bloom starts at the very beginning of stabilisation, which results solely from the release of meltwater. The increase in temperature due to the seasonal warming does not contribute to the onset of vernal blooming; temperature starts to contribute to the stratification later, when the spring bloom has ceased due to the exhaustion of nutrients in the mixed layer. By that time a deep chlorophyll maximum has formed in the seasonal pycnocline, 20–30 m below the base of the mixed layer. The effect of the seasonal ice cover on the mean areal new primary production is discussed.     

Temporal and spatial variation of the phytoplankton assemblage in the eastern Indian sector of the Southern Ocean in summer 2001/2002

The abundance and species composition of phytoplankton were investigated at stations in a permanently ice-free (61°S) and seasonally ice-covered area (64°S and 66°30′S) in the eastern Indian sector of the Southern Ocean between November 2001 and March 2002. Although a phytoplankton bloom occurred just after retreat of the sea ice at both stations in the seasonally ice-covered area, vertical stability of the water column during the bloom was weak at the most southerly station. This shows that a bloom can form even under weak vertical stability. In the bloom, diatoms dominated under weak vertical stability and Phaeocystis under strong vertical stability. In the latter case, ice algae largely contributed to development of the bloom. In the later observation period, a subsurface chlorophyll maximum (SCM) was observed at 61°S and 64°S. Species composition was different between the mixed layer and SCM at 64°S, but was uniform with depth at 61°S, indicating that the SCM is formed by different mechanisms.     

Increased sea ice melt as a driver of enhanced Arctic phytoplankton blooming

Phytoplankton primary production in the Arctic Ocean has been increasing over the last two decades. In 2019, a record spring bloom occurred in Fram Strait, characterized by a peak in chlorophyll that was reached weeks earlier than in other years and was larger than any previously recorded May bloom. Here, we consider the conditions that led to this event and examine drivers of spring phytoplankton blooms in Fram Strait using in situ, remote sensing, and data assimilation methods. From samples collected during the May 2019 bloom, we observe a direct relationship between sea ice meltwater in the upper water column and chlorophyll a pigment concentrations. We place the 2019 spring dynamics in context of the past 20 years, a period marked by rapid change in climatic conditions. Our findings suggest that increased advection of sea ice into the region and warmer surface temperatures led to a rise in meltwater input and stronger near-surface stratification. Over this time period, we identify large-scale spatial correlations in Fram Strait between increased chlorophyll a concentrations and increased freshwater flux from sea ice melt.     

On processes determining the vertical stability of surface waters in the marginal ice zone of the north-western Weddell Sea and their relationship with phytoplankton bloom development

Summary Ice-edge-related phytoplankton blooms following the retreating sea-ice in the marginal ice zone are frequently observed phenomena. Such blooms are generally short-lived and are followed by a strong decrease in the chlorophyll concentration towards the open ocean, generally explained by the degradation of the vertical stability. Solar heating and ice melting, which control the stability of the surface water of the north-western Weddell Sea during sea-ice retreat in spring were analysed in order to relate the spatial features of the phytoplankton ice-edge bloom in 1988 to the density field. Solar heating has little effect on the density of seawater in ice-covered areas because the thermal expansion coefficient is very low close to the freezing temperature. Outside the marginal ice zone, the temperature effect on stratification increases and the combined contribution of melting and heating on buoyancy input to the surface layer is roughly constant. As a consequence, the low phytoplankton stocks in the open ocean adjacent to the marginal ice zone, after an initial bloom peak following the retreating ice in spring, are not necessarily caused by deep turbulent mixing, in contrast to what is often assumed.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Spring bloom dynamics in Kongsfjorden,Svalbard: nutrients,phytoplankton, protozoans and primary production

The marine ecosystem in Kongsfjorden (79°N), a glacial fjord in Svalbard, is to a large extent well known with regard to hydrography, mesozooplankton and higher trophic levels. Research on primary production and lower trophic levels is still scare and especially investigations from winter and spring periods. The spring bloom dynamics in Kongsfjorden were investigated in 2002. The development in nutrient conditions, phytoplankton, protozoans and primary production were followed from 15 April until 22 May. The winter/spring in 2002 was categorized as a cold year with sea ice cover and water masses dominated by local winter-cooled water. The spring bloom started around 18 April and lasted until the middle of May. The bloom probably peaked in late April, but break-up of sea ice made it impossible to sample frequently in this period. Diatoms dominated the phytoplankton assemblage. We estimated the total primary production during the spring bloom in 2002 to range 27–35 g C m⁻². There was a mismatch situation between the mesozooplankton and the phytoplankton spring bloom in 2002.     

Dynamics of ice algae and phytoplankton in Frobisher Bay

Summary Vertical and seasonal variations of ice algae and phytoplankton were studied in relation to their physico-chemical environments in Frobisher Bay from 1979 to 1986. The biomass, estimated by both chlorophyll a concentrations and cell counts, was greater in the ice algae than in the phytoplankton in the underlying sea-water during winter and spring. Algal distribution in the sea ice varied vertically and seasonally, while in the underlying water column the phytoplankton distribution was much less variable. The ice algal bloom occurred at the bottom of the ice, particularly in the lower 5 cm during late spring, while the phytoplankton bloom took place at depths between 1 and 10 m during early summer after the ice bloom was over. The community structure of the ice algae changed from pennate to centric diatoms as the ice melted. The centrics dominated through the fall, and then decreased as the pennates increased in dominance when the ice formed again in winter. Species diversity and number were greater in the sea ice than in the seawater, but they were similar vertically within each habitat. The evenness of the species distribution did not vary with ice thickness or water depth. Species composition, abundance and dominance of ice algae and phytoplankton continually change both vertically and seasonally. The differential abilities of the species to attain maximal growth rates under various environmental conditions may result in species succession. Evidence is given for the major role of environmental factors regulating the dynamics of ice algae and phytoplankton.     

Growth rates of arctic juvenile Scolelepis squamata (Polychaeta: Spionidae) isolated from Chukchi Sea fast ice

In spring, Arctic coastal fast ice is inhabited by high densities of sea ice algae and, among other fauna, juveniles of benthic polychaetes. This paper investigates the hypothesis that growth rates of juveniles of the common sympagic polychaete, Scolelepis squamata (Polychaeta: Spionidae), are significantly faster at sea ice algal bloom concentrations compared to concurrent phytoplankton concentrations. Juvenile S. squamata from fast ice off Barrow, Alaska, were fed with different algal concentrations at 0 and 5?°C, simulating ambient high sea ice algal concentrations, concurrent low phytoplankton concentrations, and an intermediate concentration. Growth rates, calculated using a simple linear regression equation, were significantly higher (up to 115?times) at the highest algal concentration compared to the lowest. At the highest algal concentration, juveniles grew faster at 5?°C compared to those feeding at 0?°C with a Q ₁₀ of 2.0. We conclude that highly concentrated sea ice algae can sustain faster growth rates of polychaete juveniles compared to the less dense spring phytoplankton concentrations. The earlier melt of Arctic sea ice predicted with climate change might cause a mismatch between occurrence of polychaete juveniles and food availability in the near future. Our data indicate that this reduction in food availability might counteract any faster growth of a pelagic juvenile stage based on forecasted increased water temperatures.     

Climate change enhances primary production in the western Antarctic Peninsula

Intense regional warming was observed in the western Antarctic Peninsula (WAP) over the last 50 years. Here, we investigate the impact of climate change on primary production (PP) in this highly productive region. This study is based on temporal data series of ozone thickness (1972–2010), sea ice concentration (1978–2010), sea‐surface temperature (1990–2010), incident irradiance (1988–2010) and satellite‐derived chlorophyll a concentration (Chl‐a, 1997–2010) for the coastal WAP. In addition, we apply a photosynthesis/photoinhibition spectral model to satellite‐derived data (1997–2010) to compute PP and examine the separate impacts of environmental forcings. Since 1978, sea ice retreat has been occurring earlier in the season (in March in 1978 and in late October during the 2000s) while the ozone hole is present in early spring (i.e. August to November) since the early 1990s, increasing the intensity of ultraviolet‐B radiation (UVBR, 280–320 nm). The WAP waters have also warmed over 1990–2010. The modelled PP rates are in the lower range of previously reported PP rates in the WAP. The annual open water PP in the study area increased from 1997 to 2010 (from 0.73 to 1.03 Tg C yr^?1) concomitantly with the increase in the production season length. The coincidence between the earlier sea ice retreat and the presence of the ozone hole increased the exposure to incoming radiation (UVBR, UVAR and PAR) and, thus, increased photoinhibition during austral spring (September to November) in the study area (from 0.014 to 0.025 Tg C yr^?1). This increase in photoinhibition was minor compared to the overall increase in PP, however. Climate change hence had an overall positive impact on PP in the WAP waters.     

Influence of sea ice on the composition of the spring phytoplankton bloom in the northern Baltic Sea

During the late winter and spring of 1994, the influence of sea ice on phytoplankton succession in the water was studied at a coastal station in the northern Baltic Sea. Ice cores were taken together with water samples from the underlying water and analysed for algal composition, chlorophyll a and nutrients. Sediment traps were placed under the ice and near the bottom, and the sedimented material was analysed for algal composition. The highest concentration of ice algae (4.1 mmol C m⁻²) was found shortly before ice break-up in the middle of April, coincidental with the onset of an under-ice phytoplankton bloom. The ice algae were dominated by the diatoms Chaetoceros wighamii Brightwell, Melosira arctica (Ehrenberg) Dickie and Nitzschia frigida Grunow. Under the ice the diatom Achnanthes taeniata Grunow and the dinoflagellate Peridiniella catenata (Levander) Balech were dominant. Calculations of sinking rates and residence times of the dominant ice algal species in the photic water column indicated that only one ice algal species (Chaetoceros wighamii) had a seeding effect on the water column: this diatom dominated the spring phytoplankton bloom in the water together with Achnanthes taeniata and Peridiniella catenata. Received: 9 May 1997 / Accepted: 15 February 1998     

Abundance,population structure and vertical distribution of dominant calanoid copepods on the eastern Weddell Sea shelf during a spring phytoplankton bloom

The main emphasis of this study was to analyse the short-term development of abundance, population structure and vertical distribution of the dominant calanoid copepods during a phytoplankton bloom in the coastal area of the eastern Weddell Sea in December 2003. Microcalanus pygmaeus was by far the most abundant calanoid species. Metridia gerlachei, Ctenocalanus citer, Calanoides acutus, Calanus propinquus and the ice-associated Stephos longipes were also present in considerable proportions. The observed changes in the population characteristics and parameters of these species are described in detail and discussed in the context of the spring phytoplankton bloom. A conspicuous event occurring during the final stage of the study was the development of a strong storm. While the results suggest that this storm did not have any considerable influence on the populations of all other investigated copepod species, it very likely caused pronounced changes in the S. longipes population present in the water column. Before the storm, S. longipes was found primarily in the upper 100 m of the water column, and its population was dominated by adults (mean proportion = 41%) and the copepodite stage I (mean proportion = 30%). After the storm, the abundance increased considerably, and the copepodite stage I contributed by far the largest proportion (53%) of the total population indicating that the early copepodite stages probably had been released from the sea ice into the under ice water layer due to ice break-up and ice melt processes caused by the storm.     

Subsurface phytoplankton blooms fuel pelagic production in the North Sea

The seasonal phytoplankton biomass distribution pattern in stratifiedtemperate marine waters is traditionally depicted as consistingof spring and autumn blooms. The energy source supporting pelagicsummer production is believed to be the spring bloom. However,the spring bloom disappears relatively quickly from the watercolumn and a large proportion of the material sedimenting tothe bottom following the spring bloom is often comprised ofintact phytoplankton cells. Thus, it is easy to argue that thespring bloom is fueling the energy demands of the benthos, butmore difficult to argue convincingly that energy fixed duringthe spring bloom is fueling the pelagic production occurringduring summer months. We argue here that periodic phytoplanktonblooms are occurring during the summer in the North Sea at depthsof >25 m and that the accumulated new production [sensu (Dugdaleand Goering, Limnol. Oceanogr., 12, 196–206, 1967)] occurringin these blooms may be greater than that occurring in the springbloom in the same regions. Thus, such blooms may explain apparentdiscrepancies in production yields between different temperatemarine systems.     

Seasonal and developmental variation in the fatty acid composition of Mysis mixta (Mysidacea) and Acanthostepheia malmgreni (Amphipoda) from the hyperbenthos of a cold-ocean environment (Conception Bay, Newfoundland)

Fatty acid composition in different life-history stages of Mysismixta (Crustacea, Mysidacea) and Acanthostepheia malmgreni (Crustacea,Amphipoda) from Conception Bay, Newfoundland, was examined throughout1999 and 2000. The primary aim was to relate the seasonal fattyacid dynamics to each species’ life cycle and to publishedinformation on the occurrence and quality of the annual phytoplanktonbloom. Divergent patterns in fatty acid composition and specificfatty acid marker ratios reflected different life styles, dietsand critical periods of energy accumulation and utilizationin M. mixta and A. malmgreni. Changes in fatty acids reflectedthe sequence of plankton taxa during and following the springbloom, starting with diatoms and dinoflagellates and endingwith copepods. Immature mysids exhibited a particularly rapidaccumulation of diatom-associated fatty acids at the start ofthe spring bloom, probably owing to the high degree of motilityof M. mixta and the broad range of prey types available to thisspecies. In contrast, immature amphipods did not begin to accumulatesignificant amounts of phytoplankton fatty acids until afterthe spring bloom material had settled to the hyperbenthos inMay. Differences in fatty acid composition indicated that A.malmgreni was restricted to a lower quality diet than was M.mixta and that the trophic connection between A. malmgreni andproduction in the euphotic zone was weaker.     

Year-round acoustic detection of bearded seals (Erignathus barbatus) in the Beaufort Sea relative to changing environmental conditions, 2008–2010

Bearded seals (Erignathus barbatus) are pan-Arctic pinnipeds that are often seen in association with pack ice, and are known for their long, loud trills, produced underwater primarily in the spring. Acoustic recordings were collected from August 2008 to August 2010 at two locations and a single year (2008–2009) at a third location, in the western Beaufort Sea. Three recorders in 2008–2009 had a 30 % duty cycle and a bandwidth of 10–4,096 Hz. One recorder in 2009–2010 had a 45 % duty cycle and a bandwidth of 10–4,096 Hz and the second had a 20 % duty cycle and bandwidth of 10–8,192 Hz. Spectrograms of acoustic data were examined for characteristic patterns of bearded seal vocalizations. For each recorder, the number of hours per day with vocalizations was compared with in situ water temperature and satellite-derived daily sea ice concentrations. At all sites, bearded seals were vocally active year-round. Call activity escalated with the formation of pack ice in the winter and the peak occurred in the spring, coinciding with mating season and preceding breakup of the sea ice. There was a change in the timing of seasonal sea ice formation and retreat between the two consecutive years that was reflected in the timing of peak bearded seal call activity. This study provides new information on fall and winter bearded seal vocal behavior and the relationship between year-round vocal activity and changes in annual sea ice coverage and in situ water temperature.     

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

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

ALGAL ASSEMBLAGES IN ANTARCTIC PACK ICE AND IN ICE-EDGE PLANKTON1

A significant amount of the primary production in the Southern Ocean and other ice-covered oceans takes place in localized ice edge plankton blooms. The dynamics of these blooms appear to be closely related to seasonal melting of sea ice. Algal cells released from the ice are a possible source of ice edge planktonic assemblages, but evidence for this “seeding” has been equivocal. We compared algal assemblages in ice and water in the Weddell Sea during the austral spring of 1983 at a receding ice edge with a well-developed ice edge bloom. The high degree of similarity between ice and water column assemblages, the spatial and temporal patterns in the distribution and abundances of species, and preliminary evidence for the viability and growth of ice-associated species provide evidence for seeding from sea ice of some species in Antarctica.     

Does spring ice cover influence nest initiation date and clutch size in common eiders?

Understanding how environmental factors affect ecological parameters is important to understanding and predicting impacts of environmental change. Given evidence and anticipated impacts of climate variability, this is especially true with respect to sea ice and its role in animal life history in northern regions. We examined relationships between the extent of consolidated spring ice cover (pack and landfast), nest initiation and clutch size in common eiders (Somateria mollissima) in northern Labrador, a sub-Arctic region on the east coast of Canada. Our initial prediction was that eiders would delay nesting and have smaller clutches in years with more extensive spring ice cover. Between 1998 and 2003, we surveyed coastal islands for breeding eiders and collected information on nest age and clutch size. For those years, we estimated ice cover based on Radarsat-1 images supplied by the Canadian Ice Service during the spring period (approximately June 7–12). We found that spring ice cover was a significant positive predictor of nest initiation date, and the regression equation indicated that if the average extent of ice cover around nesting islands increased by 18 ha, average nesting date was delayed by approximately 1 day. Nest initiation date was a significant negative predictor of clutch size, and the regression equation indicated that a 20 day delay in nesting reduced average clutch size by approximately 1 egg. However, ice cover itself was not a significant predictor of clutch size. Our findings suggest that eiders breed when ice is present, but ice extent may negatively influence aspects of their breeding ecology.     

Environmental Variability and Its Impact on the Reproductive Cycle of Antarctic Krill

"Recruitment potential" in Antarctic krill in the Palmer Long-TermEcological Research (LTER) study region west of the AntarcticPeninsula varied significantly over the 7-yr time series betweenJanuary 1993 and January 1999. Timing of ovarian maturation,the percent of the population reproducing, and individual reproductiveoutput (batch volume, embryo diameter) were measured. Indiceshave been developed to quantify the timing and intensity ofreproduction in Antarctic krill. One finding important to estimatesof population fecundity for this long-lived species is thatthe percent of the population reproducing can vary widely, from10 to 98%. Each season was characterized as having delayed,average or advanced ovarian development. In this study we relatethese indices to direct and indirect indicators of spring orannual food availability. The timing of the spring sea ice retreatand the extent of sea ice in the spring (September through November)appear to significantly affect the intensity and timing of reproductionin the population. Intensity of reproduction was highest under"average" conditions, and oöcyte development fastest withconditions of a late retreat and high spring sea ice extent.     

Protist assemblages in winter sea ice: setting the stage for the spring ice algal bloom

This study documents, for the first time, the abundance and species composition of protist assemblages in Arctic sea ice during the dark winter period. Lack of knowledge of sea-ice assemblages during the dark period has left questions about the retention and survival of protist species that initiate the ice algal bloom. Sea-ice and surface water samples were collected between December 27, 2007 and January 31, 2008 within the Cape Bathurst flaw lead, Canadian Beaufort Sea. Samples were analyzed for protist identification and counts, chlorophyll (chl) a, and total particulate carbon and nitrogen concentrations. Sea-ice chl a concentrations (max. 0.27 μg l⁻¹) and total protist abundances (max. 4 × 10³ cells l⁻¹) were very low, indicating minimal retention of protists in the ice during winter. The diversity of winter ice protists (134 taxa) was comparable to spring ice assemblages. Pennate diatoms dominated the winter protist assemblage numerically (averaging 77% of total protist abundances), with Nitzschia frigida being the most abundant species. Only 56 taxa were identified in surface waters, where dinoflagellates were the dominant group. Our results indicate that differences in the timing of ice formation may have a greater impact on the abundance than structure of protist assemblages present in winter sea ice and at the onset of the spring ice algal bloom.     

Temporal and vertical variations of lipid biomarkers during a bottom ice diatom bloom in the Canadian Beaufort Sea: further evidence for the use of the IP25 biomarker as a proxy for spring Arctic sea ice

Variations in the concentrations of the sea ice diatom biomarker, IP₂₅ (Ice Proxy with 25 carbon atoms), were measured in the bottom 10 cm of sea ice collected from the eastern Beaufort Sea and Amundsen Gulf from January to June 2008, as part of the International Polar Year–Circumpolar Flaw Lead system study. Temporal and vertical changes in IP₂₅ concentrations were compared against other biomarkers and indicators of ice algal production. IP₂₅ was not detected in sea ice samples collected from mid-winter to early spring, likely as a result of light-limiting conditions for algal growth and accumulation. From early March to mid-June, IP₂₅ concentrations correlated well with those of fatty acids (r = 0.79; P < 0.001), less so with total sterols (r = 0.63; P < 0.001) and qualitatively with chlorophyll a concentrations and diatom cell abundances from adjacent sea ice cores. Approximately 90% of the total sea ice IP₂₅ accumulation occurred from mid-March to late-May, coincident with the ice algal bloom period. The majority (ca. 87–93%) of IP₂₅ was biosynthesised within the lower 5 cm of the sea ice where brine volume fractions were >5% which is consistent with the hypothesis that brine channel connectivity limits the internal colonisation of sea ice by diatoms. Maximum IP₂₅ concentrations occurred at 1–3 cm from the ice–water interface providing further evidence for a selective sea ice diatom origin for this biomarker. In contrast, vertical concentration profiles for fatty acids and sterols indicated mixed sources for these biomarkers.     

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.