Distribution patterns of benthic microalgal standing stock at McMurdo Sound,Antarctica

Summary During the austral summer of 1975–76 and winter of 1977 benthic and water column chlorophyll a and phaeopigments were measured at several sites along the east and west sides of McMurdo Sound, Antarctica. Estimates of in situ primary productivity were made at some McMurdo Sound locations. Additionally, water column samples were collected at 5 stations in the Ross Sea during January, 1976. Standing stock data are analyzed to identify seasonal and spatial patterns. Variability in algal standing stock was related to ambient light levels and appeared to be mediated by ice and snow cover whereby the highest algal standing stock was present under high light conditions (low ice and snow cover, shallow water, summer). Differences in published benthic invertebrate densities appear to be closely allied to differences in benthic primary production, and less so to in situ planktonic ice microalgal production.     

A multiple biomarker approach to tracking the fate of an ice algal bloom to the sea floor

In ice-covered Arctic seas, the ice algal production can be the main input of organic matter to the ecosystem. Pelagic–benthic coupling is thought to be particularly tight in those areas. The increase in ice algal production in Franklin Bay from January/February to April/May 2004 paralleled an increase in benthic oxygen demand. However, sedimentary chlorophyll a, which is usually an indicator of “fresh” organic matter inputs to the sea floor, did not increase. Consequently, it was asked what was the fate of the ice algal phytodetritus arriving at the sea floor? To answer this question, photosynthetic pigments from the sea ice, water column particulate organic matter, and sediment, as well as diatom frustules in the sediment, were studied from January to May 2004. The number of ice diatom cells in the sediment showed an increase in April/May, confirming higher inputs of fresh ice algae to the sediment. Changes in sedimentary pigment profiles in the first 10 cm suggested an increase in bioturbation due to enhanced benthic activities. Finally, the decrease in the ratio of chlorophyll a to phaeophorbide a implied an increase in macrobenthic activity. Benthic macrofauna consumed some of the deposited material and mixed some within the top five cm of sediment. The response of sedimentary pigments to an ice algal input can be studied at different levels and it is only the combination of these studies that will allow an understanding of the overall fate of phytodetritus in the benthic compartment.     

Limited use of sea ice by the Ross seal (<Emphasis Type="Italic">Ommatophoca rossii</Emphasis>), in Amundsen Sea,Antarctica, using telemetry and remote sensing data

To understand the use and importance of the Antarctic sea ice to the Ross seal (Ommatophoca rossii), four adult females were tagged with Argos satellite transmitters in the Amundsen Sea, Antarctica. The Ross seal is the least studied of the Antarctic seal species and nothing was previously known about their behaviour in the Amundsen Sea. During almost 1 year, their movements, haul out behaviour and time spent at different temperatures were logged. By comparing their movements with daily ice maps, distances to the ice edge were calculated, and seals dependence on sea ice for resting, breeding and moulting was analysed. The tagged seals spent on average 70.8 % (range 66.8–77.8 %) of their time in the water and hauled out mainly during the moult in December–January, and in late October–mid-November during breeding. During the pelagic period, they were on average 837.5 km (range 587–1,282 km) from the ice edge indicating a fully pelagic life during several months. Their pelagic behaviour suggests that Ross seals, although being an ice obligate species, may adapt comparatively easy to climate change involving ice melting and recession and thereby potentially being less sensitive to the reduction of sea ice than other Antarctic seal species. Although nothing is known about their mating behaviour, they appear to be relatively stationary during moulting and breeding, hence requiring a small ice surface. Although previous studies in other parts of Antarctica have found similar results, still many questions remain about this peculiar species.     

Measurements of Seasonal Rates and Annual Budgets of Organic Carbon Fluxes in an Antarctic Coastal Environment at Signy Island, South Orkney Islands, Suggest a Broad Balance between Production and Decomposition

We report here the first comprehensive seasonal study of benthic microbial activity in an Antarctic coastal environment. Measurements were made from December 1990 to February 1992 of oxygen uptake and sulfate reduction by inshore coastal sediments at Signy Island, South Orkney Islands, Antarctica. From these measurements the rate of benthic mineralization of organic matter was calculated. In addition, both the deposition rate of organic matter to the bottom sediment and the organic carbon content of the bottom sediment were measured during the same period. Organic matter input to the sediment was small under winter ice cover, and the benthic respiratory activity and the organic content of the surface sediment declined during this period as available organic matter was depleted. On an annual basis, about 32% of benthic organic matter mineralization was anoxic, but the proportion of anoxic compared with oxic mineralization increased during the winter as organic matter was increasingly buried by the amphipod infauna. Fresh organic input occurred as the sea ice melted and ice algae biomass sedimented onto the bottom, and input was sustained during the spring after ice breakup by continued primary production in the water column. The benthic respiratory rate and benthic organic matter content correspondingly increased towards the end of winter with the input of this fresh organic matter. The rates of oxygen uptake during the southern summer (80 to 90 mmol of O₂ m^-2 day^-1) were as high as those reported for other sediments at much higher environmental temperatures, and the annual mineralization of organic matter was equally high (12 mol of C m^-2 year^-1). Seasonal variations of benthic activity in this antarctic coastal sediment were regulated by the input and availability of organic matter and not by seasonal water temperature, which was relatively constant at between -1.8 and 0.5°C. We conclude that despite the low environmental temperature, organic matter degradation broadly balanced organic matter production, although there may be significant interrannual variations in the sources of the organic matter inputs.     

Contribution of sea ice microbial production to Antarctic benthic communities is driven by sea ice dynamics and composition of functional guilds

Organic matter produced by the sea ice microbial community (SIMCo) is an important link between sea ice dynamics and secondary production in near‐shore food webs of Antarctica. Sea ice conditions in McMurdo Sound were quantified from time series of MODIS satellite images for Sept. 1 through Feb. 28 of 2007–2015. A predictable sea ice persistence gradient along the length of the Sound and evidence for a distinct change in sea ice dynamics in 2011 were observed. We used stable isotope analysis (δ¹³C and δ¹⁵N) of SIMCo, suspended particulate organic matter (SPOM) and shallow water (10–20 m) macroinvertebrates to reveal patterns in trophic structure of, and incorporation of organic matter from SIMCo into, benthic communities at eight sites distributed along the sea ice persistence gradient. Mass‐balance analysis revealed distinct trophic architecture among communities and large fluxes of SIMCo into the near‐shore food web, with the estimates ranging from 2 to 84% of organic matter derived from SIMCo for individual species. Analysis of patterns in density, and biomass of macroinvertebrate communities among sites allowed us to model net incorporation of organic matter from SIMCo, in terms of biomass per unit area (g/m²), into benthic communities. Here, organic matter derived from SIMCo supported 39 to 71 per cent of total biomass. Furthermore, for six species, we observed declines in contribution of SIMCo between years with persistent sea ice (2008–2009) and years with extensive sea ice breakout (2012–2015). Our data demonstrate the vital role of SIMCo in ecosystem function in Antarctica and strong linkages between sea ice dynamics and near‐shore secondary productivity. These results have important implications for our understanding of how benthic communities will respond to changes in sea ice dynamics associated with climate change and highlight the important role of shallow water macroinvertebrate communities as sentinels of change for the Antarctic marine ecosystem.     

Meiofauna distribution and mesoscale variability in two sites of the Ross Sea (Antarctica) with contrasting food supply

Meiofauna abundance, biomass and community structure were investigated in two comparable deep sites of the Ross Sea (Antarctica) characterized by different trophic and sediment characteristics. Site B (567 m depth, dominated by muddy sediments) and site C (439 m depth, characterized by the presence of calcareous debris and coarse sand) were located at increasing distance from the polynyas and were subject to different inputs of organic material to the seabed. Total meiofauna abundance ranged from 192.0 to 1191.2 ind. 10 cm⁻², and total biomass varied between 9.5 and 50.3 μgC 10 cm⁻². Meiofauna densities from the Ross Sea are, on average, 2–7 times lower than those reported from other similar deep polar regions and displayed significant differences between the sites. Nematodes dominated the samples at both sites, but their relative significance changed between the sites (80% at site B and 56% at site C), followed by copepods (1.6% and 35% at sites B and C, respectively). Meiofauna composition at site B appeared similar to that reported for deep-sea antarctic or temperate sediments, whereas the composition at site C was similar to that of coastal areas. On a macroscale, the different inputs of utilizable organic material at the two sites were reflected in meiofaunal distribution patterns, indicating that meiofaunal communities from the Ross Sea are dependent on particulate organic matter fluxes from the photic layer and are coupled to pelagic phenomena. Very low microscale variations (i.e. between replicates) in meiofauna density contrasted with large mesoscale variability, which was related to the concentration of the main food indicators (phytopigments, proteins, carbohydrates and lipids). Accepted: 18 February 1999     

Unusual trophic strategies of Hydractinia angusta (Cnidaria, Hydrozoa) from Terra Nova Bay, Antarctica

Hydractinia angusta Hartlaub, 1904 has been recorded at Terra Nova Bay (Ross Sea, Antarctica) as epizoic on shells of the Antarctic scallop Adamussium colbecki. The species can exploit different trophic resources: first, polyps are able to detach and ingest tube feet and pedicellariae from the sea urchins Sterechinus neumayeri, grazing on the scallop shell, and second, they also eat masses of benthic diatoms settled among the hydrorhiza of the colony. The particular relationship observed between the hydroid and one of the most common Antarctic sea urchins may prevent or reduce the damage to A. colbecki shells, otherwise caused by the grazing of sea urchins on the algal film of the upper valve of the scallops. H. angusta is the first known species of hydroid that exploits prey several times its own size and the second that does not ingest entire prey but portions of them. The use of benthic diatoms as a food resource has previously been documented for the sub-Antarctic marine hydroid Silicularia rosea. Accepted: 20 December 1999     

The nutrient status in sea ice of the Weddell Sea during winter: effects of sea ice texture and algae

Summary Cores and brine samples from sea ice of the Weddell Sea were analyzed for nutrients (phosphate, nitrate and silicate), salinity and chlorophyll a during winter. Stratigraphic analyses of the cores were also carried out. Bulk nutrient concentrations in the sea ice fluctuated widely and did not correlate with salinity. Nutrient concentrations in cores were normalized to sea-water salinity to facilitate comparison. They varied between zero and two or three times those measured in the water column. Differentiation into young and old sea ice, however, revealed that nutrient concentrations in the young ice in many cases corresponded to those in surface seawater. In older ice, nutrients showed signs of increase as well as depletion or exhaustion relative to the water column. Differentiation of core sections according to ice textural classes and analyses of brine samples clarified some relationships between nutrients, salinity and algal biomass. Most of the changes in the nutrient concentrations are attributed to an increase in biological activity as the seasons progress. Silicate is expected to become the first nutrient likely to limit growth of diatoms in the ice which is ascribed to slower regeneration or dissolution of this nutrient relative to phosphate and nitrate. A consequence of silicate exhaustion may be the succession of different algal assemblages, from a diatom dominated community to one in which autotrophic flagellates form the largest component.     

Composition, abundance and stratification of soft-bottom macrobenthos from selected areas of the Ross Sea shelf (Antarctica)

Composition, abundance and stratification of soft-bottom macrobenthos were studied at three selected sites on the Ross Sea shelf (Antarctica) with different geomorphology and sedimentation regimes. Sites A (southwest Ross Sea, 810 m depth) and B (Joides basin, 580 m depth) were characterized by biogenic mud and clay sediments, whereas site C (Mawson bank, 450 m depth) featured sandy sediments mixed with a conspicuous biogenic component characterized by shells and tests of calcareous invertebrates (mainly barnacles of the genus Bathylasma). The macrofauna of sites A and B was mainly composed of infaunal polychaetes and bivalves. The assemblages comprised both surface and sub-surface deposit feeders, including some conveyor-belt polychaetes (Maldanidae and Capitellidae) that are responsible for high sediment mixing and bioturbation. The macrobenthos of site C was dominated by crustaceans, polychaetes and echinoderms (ophiuroids), and mainly by filter feeders and epifaunal or interstitial forms. Abundances were higher (up to 1040 ind. m⁻²) at site B than at sites A and C (430 and 516 ind. m⁻², respectively). At sites A and B the benthos was mainly concentrated in the upper 5 cm of the sediment, and abundances declined sharply in the deeper sediment layers. These results indicate a high degree of consistency between sediment features and benthic community structure, which are both strongly related to local hydrography and bottom dynamics. Sites A and B represent areas where the organic input to the seafloor by vertical sedimentation from the upper water column is high. Site C, however, is characterized by high sediment instability and food particles advecting mainly horizontally. The community is more physically controlled by unpredictable, and probably frequent, disturbance events (e.g., bottom turbid currents, sediment reworking and displacement). Individuals were relatively small, indicating that probably they are not able to grow up to the adult size and reproduce. The community may be represented by “pseudopopulations” depending on the settlement of larvae invading from neighbouring areas. Accepted: 23 October 1998     

Photoacclimation in Antarctic bottom ice algae: an experimental approach

The aim of the study was to investigate the capacity of microalgae from the extremely low light habitat of bottom ice to acclimate to different light conditions. During austral spring 1997 the bottom layer of land-fast ice in Terra Nova Bay displayed high values of microalgal biomass up to 2,400 μg Chla L⁻¹ concentrated in a few centimetres ice layer. The algal assemblage was dominated by benthic pennate diatoms. Photoacclimation of the microalgae was addressed in terms of pigment spectra and photosynthetic parameters. Immediate and long term (minutes to days) changes in the photoprotective pigments (DD-cycle) were analysed. Severe photodamage occurred in microalgal assemblages exposed to high light. However, part of the bottom ice algal community showed a notable ability to acclimate to high irradiance levels. Changes in photosynthetic parameters preceded the sudden abrupt changes in pigment synthesis and the rapid increase in biomass and growth rates. This article belongs to a special topic: Five articles on Sea-ice communities in Terra Nova Bay (Ross Sea), coordinated by L. Guglielmo and V. Saggiomo, appear in this issue of Polar Biology. The studies were conducted in the frame of the National Program of Research in Antarctica (PNRA) of Italy.     

Organic matter composition in coastal sediments at Terra Nova Bay (Ross Sea) during summer 1995

We illustrate the spatial and vertical distribution of sediment phytopigments and organic matter biochemical composition at Terra Nova Bay (Ross Sea) during summer 1995. Coastal sediments displayed high phytopigments concentrations associated with huge amounts of labile organic matter largely dominated by proteins. This result was opposite to previous observations in the same area. Such comparison suggested that whilst organic matter quantity in the sediments depended upon the vertical input from the water column, temporal changes in its biochemical composition were related to benthic processes. As considerably high concentrations of biopolymeric organic carbon were found even at 6-cm depth and according to the “loss type” functioning of the coastal waters of the Ross Sea, we stress the summer time occurrence in coastal sediments of an important organic matter burial. Accepted: 24 October 1999     

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.     

The inshore marine ecosystem off the Vestfold Hills,Antarctica

The planktonic, ice/water interface, and benthic communities at three sites off the coast of the Vestfold Hills, Antarctica, were examined over a complete year.The planktonic flora and fauna were composed predominantly of oceanic species with diatoms and copepods the numerically dominant groups. Primary production was largely restricted to the summer months except for epontic algae which developed in spring. The zooplankton exhibited a similar seasonal cycle but lagged some months behind that of the phytoplankton.The ice/water interface (epontic) fauna consisted of species from the plankton and benthos. Copepods were major contributors; however, two amphipod species dominated. Seasonality of the fauna in this habitat was determined by ice formation and breakout, and development of ice algae.Each of the benthic substrates supported a characteristic macrofaunal assemblage, although infaunal amphipods and tanaids were similar at each site. Infauna exhibited a distinct seasonal cycle related to that of the primary producers whereas macrofauna showed no seasonal changes in abundance.Species composition of each community in this coastal antarctic region was comparable with that of similar habitats in other antarctic coastal areas, supporting the circumpolarity of antarctic marine communities.     

Comparison of bottom sea-ice algal characteristics from coastal and offshore regions in the Arctic Ocean

Ice-core observations were conducted at three arctic sites in early April 2003: one clean sediment-free site (Chukchi), one sediment-rich site (Beaufort) on landfast sea ice offshore Barrow, and one oceanic Ice Exercise (ICEX) site. Concentrations of inorganic nutrients at coastal sites were similar to but higher than those at the oceanic site. Chlorophyll-a (Chl-a) concentration was much higher in the Chukchi bottom ice than at the other two sites. However, large size Chl-a (>20 μm) dominated (>70%) the bottom-ice algal Chl-a at both the Chukchi and Beaufort sites, whereas the oceanic ICEX site was evenly occupied by large (>20 μm), medium (5–20 μm), and small Chl-a (0.7–5 μm). These in situ data were incorporated into an ocean-ecosystem model. The model results revealed that: (1) strong light attenuation by trapped sediments controlled ice-algal production at the Beaufort site; (2) the peak in ice algae occurred later at the oceanic site than at the Chukchi site because of thicker ice and the consequently reduced amount of light reaching the ice algae at the ice bottom; and (3) maximum production at the oceanic site reached only 10% the level at the Chukchi site because of nutrient limitation.     

Rapid Response of a Marine Mammal Species to Holocene Climate and Habitat Change

Environmental change drives demographic and evolutionary processes that determine diversity within and among species. Tracking these processes during periods of change reveals mechanisms for the establishment of populations and provides predictive data on response to potential future impacts, including those caused by anthropogenic climate change. Here we show how a highly mobile marine species responded to the gain and loss of new breeding habitat. Southern elephant seal, Mirounga leonina, remains were found along the Victoria Land Coast (VLC) in the Ross Sea, Antarctica, 2,500 km from the nearest extant breeding site on Macquarie Island (MQ). This habitat was released after retreat of the grounded ice sheet in the Ross Sea Embayment 7,500–8,000 cal YBP, and is within the range of modern foraging excursions from the MQ colony. Using ancient mtDNA and coalescent models, we tracked the population dynamics of the now extinct VLC colony and the connectivity between this and extant breeding sites. We found a clear expansion signal in the VLC population ~8,000 YBP, followed by directional migration away from VLC and the loss of diversity at ~1,000 YBP, when sea ice is thought to have expanded. Our data suggest that VLC seals came initially from MQ and that some returned there once the VLC habitat was lost, ~7,000 years later. We track the founder-extinction dynamics of a population from inception to extinction in the context of Holocene climate change and present evidence that an unexpectedly diverse, differentiated breeding population was founded from a distant source population soon after habitat became available.     

Too much of a good thing: sea ice extent may have forced emperor penguins into refugia during the last glacial maximum

The relationship between population structure and demographic history is critical to understanding microevolution and for predicting the resilience of species to environmental change. Using mitochondrial DNA from extant colonies and radiocarbon‐dated subfossils, we present the first microevolutionary analysis of emperor penguins (Aptenodytes forsteri) and show their population trends throughout the last glacial maximum (LGM, 19.5–16 kya) and during the subsequent period of warming and sea ice retreat. We found evidence for three mitochondrial clades within emperor penguins, suggesting that they were isolated within three glacial refugia during the LGM. One of these clades has remained largely isolated within the Ross Sea, while the two other clades have intermixed around the coast of Antarctica from Adélie Land to the Weddell Sea. The differentiation of the Ross Sea population has been preserved despite rapid population growth and opportunities for migration. Low effective population sizes during the LGM, followed by a rapid expansion around the beginning of the Holocene, suggest that an optimum set of sea ice conditions exist for emperor penguins, corresponding to available foraging area.     

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.     

Nonlinear effects of winter sea ice on the survival probabilities of Adélie penguins

The population dynamics of Antarctic seabirds are influenced by variations in winter sea ice extent and persistence; however, the type of relationship differs according to the region and the demographic parameter considered. We used annual presence/absence data obtained from 1,138 individually marked birds to study the influence of environmental and individual characteristics on the survival of Adélie penguins Pygoscelis adeliae at Edmonson Point (Ross Sea, Antarctica) between 1994 and 2005. About 25% of 600 birds marked as chicks were reobserved at the natal colony. The capture and survival rates of Adélie penguins at this colony increased with the age of individuals, and five age classes were identified for both parameters. Mean adult survival was 0.85 (SE = 0.01), and no effect of sex on survival was evident. Breeding propensity, as measured by adult capture rates, was close to one, indicating a constant breeding effort through time. Temporal variations in survival were best explained by a quadratic relationship with winter sea ice extent anomalies in the Ross Sea, suggesting that for this region optimal conditions are intermediate between too much and too little winter sea ice. This is likely the result of a balance between suitable wintering habitat and food availability. Survival rates were not correlated with the Southern Oscillation Index. Low adult survival after a season characterized by severe environmental conditions at breeding but favorable conditions during winter suggested an additional mortality mediated by the reproductive effort. Adélie penguins are sensitive indicators of environmental changes in the Antarctic, and the results from this study provide insights into regional responses of this species to variability in winter sea ice habitat.     

Response to nutrient enrichment by the plankton of Antarctic coastal lakes and the inshore Ross Sea

Summary Water samples from coastal lakes and ponds on Ross Island (77°S) and from the adjacent Ross Sea were analyzed to determine nutrient supply relative to the population size of planktonic algae and bacteria. An enrichment test was applied to each water to measure N and P accumulation by the seston as a guide to nutrient demand and deficiency. In all waters, including the sea, the dissolved inorganic N-to-P ratios were low, but inorganic nitrogen was generally present at concentrations that satisfied or exceeded the current growth requirements of the algae. None of the samples responded to added P, and only 3 of the 13 plankton assemblages tested responded positively to nitrogen enrichment. These waters are therefore characterized by an abundance of resources relative to demand. This apparently r-selecting coastal environment contrasts markedly in many ecosystem properties with the inland meromictic lakes of Antarctica.     

SHADE ADAPTED BENTHIC DIATOMS BENEATH ANTARCTIC SEA ICE1

A dense community of shade adapted microalgae dominated by the diatom Trachyneis aspera is associated with a siliceous sponge spicule mat in McMurdo Sound, Antarctica. Diatoms at a depth of 20 to 30 m were found attached to spicule surfaces and in the interstitial water between spicules. Ambient irradiance was less than 0.6 μE · m^?2· s^?1 due to light attenuation by surface snow, sea ice, ice algae, and the water column. Photosynthesis-irradiance relationships determined by the uptake of NaH¹⁴CO₃ revealed that benthic diatoms beneath annual sea ice were light-saturated at only 11 μE·m^?2·s^?1, putting them among the most shade adapted microalgae reported. Unlike most shade adapted microalgae, however, they were not photoinhibited even at irradiances of 300 μE·m^?2·s^?1. Although in situ primary production by benthic diatoms was low, it may provide a source of fixed carbon to the abundant benthic invertebrates when phytoplankton or ice algal carbon is unavailable.