Sea ice seasonality during the Holocene, Adélie Land, East Antarctica

Thin sections of laminated cores from different Antarctic coastal areas have demonstrated the potential of diatom species to document climate change at the seasonal scale. Here we present the relative abundances of four diatom species and species groups (Fragilariopsis curta group as a proxy for yearly sea ice cover, F. kerguelensis as a proxy for summer sea-surface temperature, Chaetoceros Hyalochaete resting spores as a proxy for spring sea ice melting and the Thalassiosira antarctica group as a proxy for autumn sea ice formation) in core MD03-2601 retrieved off Adélie Land on the Antarctic continental shelf. These abundances were compared to surface temperatures and sea ice cover modelled over the last 9000 years. Both the marine records and the simulated climate demonstrated a cooler Early Holocene (9000–7700 years BP), a warmer Mid-Holocene (7700–4000 years BP) and a colder Late Holocene (4000–1000 years BP). Yearly sea ice cover followed an inverse pattern to temperatures with less sea ice during the Mid-Holocene Hypsithermal than during the Late Holocene Neoglacial. However, diatom census counts and model output indicate that sea ice spring melting happened earlier in the season, as expected, but that autumn sea ice formation also occurred earlier in the season during the Hypsithermal than during the colder Neoglacial, thereby following seasonal changes in local insolation.     

Modern distribution of diatoms in sediments from the George V Coast, Antarctica

Currently, little information exists concerning high frequency climate change in the Antarctic, data that are essential to gaining a more complete understanding of global climate change. Sediments from the George V Coast continental margin potentially contain a high resolution record of the late Holocene. The relatively high rate of sedimentation ( 3 mm/year) and the presence of laminated intervals which contain undisturbed sequences make these sediments valuable for future downcore work. In order to utilize diatom assemblages as proxies of paleoclimatic and paleoceanographic parameters, a preliminary study of the surface sediment distribution of diatoms from the George V Coast continental margin was completed. Nitzschia curta (Van Heurck) Hasle and Nitzschia kerguelensis (O'Meara) Hasle present great potential for tracking the Holocene retreat of ice across the continental shelf of this region. Concentrations of N. curta, an “ice-related” form, decreases in an offshore direction, whereas N. kerguelensis, an “oceanic” species associated with circumpolar flow, displays an increase in sediments offshore. The genus Thalassiosira Cleve displays a clear association with areas of open water primary production, as has been noted in others parts of the Antarctic. Downcore variability in relative abundance of this genus will be useful in understanding the history of sea ice versus open water production on the George V Coast shelf. In contrast, the distribution of Chaetoceros Ehrenberg resting spores appears to be more complex, but may be related to the position of the stationary ice edge associated with the annual minimal extent of sea ice. Finally, an understanding of the downcore variability of Thalassiothrix antarctica Schimper ex Karsten, a species which is very abundant in the laminated sections of the sediment cores, will have to rely on studies of its distribution in other areas of the Antarctic, as it is sparsely distributed along the George V Coast today.     

Paleoenvironmental change in the middle Okinawa Trough since the last deglaciation: Evidence from the sedimentation rate and planktonic foraminiferal record

Well-dated, high-resolution records of planktonic foraminifera and oxygen isotopes from two sediment cores, A7 and E017, in the middle Okinawa Trough reveal strong and rapid millennial-scale climate changes since ∼ 18 to 17 thousand years before present (kyr B.P.). Sedimentation rate shows a sudden drop at ∼ 11.2 cal. kyr B.P. due to a rapid rise of sea level after the Younger Dryas (YD) and consequently submergence of the large continental shelf on the East China Sea (ECS) and the retreat of the estuary providing sediment to the basin. During the last deglaciation, the relative abundance of warm and cold species of planktonic foraminifera fluctuates strongly, consistent with the timing of sea surface temperature (SST) variations determined from Mg/Ca measurements of planktonic foraminifera from one of the two cores. These fluctuations are coeval with climate variation recorded in the Greenland ice cores and North Atlantic sediments, namely Heinrich event 1 (H1), Bølling–Allerød (B/A) and YD events. At about 9.4 kyr B.P., a sudden change in the relative abundance of shallow to deep planktonic species probably indicates a sudden strengthening of the Kuroshio Current in the Okinawa Trough, which was synchronous with a rapid sea-level rise at 9.5–9.2 kyr B.P. in the ECS, Yellow Sea (YS) and South China Sea (SCS). The abundance of planktonic foraminiferal species, together with Mg/Ca based SST, exhibits millennial-scale oscillations during the Holocene, with 7 cold events (at about 1.7, 2.3–4.6, 6.2, 7.3, 8.2, 9.6, 10.6 cal. kyr BP) superimposed on a Holocene warming trend. This Holocene trend, together with centennial-scale SST variations superimposed on the last deglacial trend, suggests that both high and low latitude influences affected the climatology of the Okinawa Trough.     

An r-strategist in Antarctic pack ice

Summary The Antarctic copepod Drescheriella glacialis, an inhabitant of sea ice, is the first polar invertebrate metazoan to have been cultured throughout its life cycle. We describe its demographic characteristics on the basis of a laboratory cohort study and correlative field data. When compared to its closest temperate-zone relatives, D. glacialis shows temperature compensation of developmental and reproductive rates. A genuine r-strategist in every respect, it does not fit established trends for Antarctic invertebrates but appears well adapted to the peculiar spatio-temporal variability of the sea ice habitat.     

Early Pliocene paleoenvironment of the Sørsdal Formation, Vestfold Hills, based on diatom data

Comparison of diatom data from modern surface sediments in Prydz Bay and the Kerguelen Plateau with diatom assemblages from the Sørsdal Formation, Vestfold Hills, indicates that the climate was warmer than present during the early Pliocene (4.5–4.1 Ma). Extant, sea-ice associated diatoms are significantly less abundant throughout the Sørsdal Formation than in the modern Antarctic coastal zone. Extant diatoms in the Sørsdal Formation, including Stellarima stellaris, Thalassiosira oliverana, Fragilariopsis sublinearis, Pseudo-nitzschia turgiduloides and Eucampia antarctica var. recta, are consistent with annual sea-surface temperatures (SST) of between −1.8 and 5.0°C. The presence of S. stellaris indicates that the summer SSTs were >3°C during some intervals. The absence of calcareous coccoliths and the silicoflagellate Dictyocha suggests that the upper limit for summer SST was <5°C. These data indicate that early Pliocene summer SST were between 1.6 and 3°C warmer than today. Abundant Chaetoceros cysts infer that stratified, open-water conditions were present during summer/spring. Ice sheet models suggest that warming of the magnitude evident in the Sørsdal Formation (≤3°C) should have resulted initially in increased snow accumulation and ice sheet growth. However, ice sheet growth was probably short-lived, as the long-term response to this warming in the early Pliocene resulted in a significant decrease in ice volume and deposition of the Sørsdal Formation. Other factors, such as increased basal-ice sliding and higher discharge (icebergs and melt-water), probably led to significantly elevated ablation rates from the Pliocene ice sheet, resulting in ice sheet retreat.     

Effects of the ice-edge bloom and season on the metabolism of copepods in the Weddell Sea,Antarctica

The metabolic responses of several species of Antarctic copepods to primary productivity and changes between seasons were investigated. To examine the influence of the spring ice-edge bloom on the metabolism of copepods, oxygen consumption rates were determined on specimens from three zones of widely different ice coverage and chlorophyll biomass: pack ice (pre-bloom), ice edge (bloom) and open water (post-bloom). Summer metabolic rates were compared with published winter rates. Field work was done in the Weddell Sea in the region of 60 °S, 36°W in late November and December 1993. Oxygen consumption rates were determined by placing individuals in syringe respirometers and monitoring the oxygen partial pressure for 10–20 hours. Higher metabolic rates were observed in the primarily herbivorous copepods, Calanoides acutus, Rhincalanus gigas and Calanus propinquus in regions of higher primary production: ice edge and open water. The carnivorous Paraeuchaeta antarctica showed a similar pattern. The omnivorous copepods Metridia gerlachei and Gaetanus tenuispinus showed no changes in metabolism between zones. Data on routine rates of copepods from the winter were available for C. propinquus and P. antarctica. In P. antarctica, rates were higher in the summer. Calanus propinquus showed a higher metabolic rate in the summer than in the winter, but the difference was not significant at the 0.05 level. It was concluded that copepods near the ice zone in the ice zone in the Antarctic rely on the spring ice-edge bloom for growth and completion of their life cycle.     

Copepods in summer platelet ice in the eastern Weddell Sea,Antarctica

Copepods in platelet-ice layers underlying fast ice and in the water column below were studied at Drescher Inlet, eastern Weddell Sea in February 1998. Three copepod species were found: Drescheriella glacialis and Paralabidocera antarctica occurred in platelet-ice layers, while Stephos longipes was only present in the water column. The distribution of all species varied considerably between station and depth. D. glacialis dominated the platelet-ice community and occurred at all five platelet-ice sampling sites, except one, with numbers of up to 26 ind. l^–1. In contrast, P. antarctica was only found in low numbers (up to 2 ind. l^–1) at one site. The total copepod abundance in the platelet ice was not associated with algal biomass, although it was strongly correlated with high ammonium concentrations (up to 9 M) in the interstitial water between the platelets. This is the first indirect evidence to support the hypothesis that zooplankton excretion can partly account for the high ammonium values often found in platelet-ice layers.     

Biogeographic responses of the copepod Calanus glacialis to a changing Arctic marine environment

Dramatic changes have occurred in the Arctic Ocean over the past few decades, especially in terms of sea ice loss and ocean warming. Those environmental changes may modify the planktonic ecosystem with changes from lower to upper trophic levels. This study aimed to understand how the biogeographic distribution of a crucial endemic copepod species, Calanus glacialis, may respond to both abiotic (ocean temperature) and biotic (phytoplankton prey) drivers. A copepod individual‐based model coupled to an ice‐ocean‐biogeochemical model was utilized to simulate temperature‐ and food‐dependent life cycle development of C. glacialis annually from 1980 to 2014. Over the 35‐year study period, the northern boundaries of modeled diapausing C. glacialis expanded poleward and the annual success rates of C. glacialis individuals attaining diapause in a circumpolar transition zone increased substantially. Those patterns could be explained by a lengthening growth season (during which time food is ample) and shortening critical development time (the period from the first feeding stage N3 to the diapausing stage C4). The biogeographic changes were further linked to large‐scale oceanic processes, particularly diminishing sea ice cover, upper ocean warming, and increasing and prolonging food availability, which could have potential consequences to the entire Arctic shelf/slope marine ecosystems.     

Surface-associated diatoms from marine habitats at Cape Evans,Antarctica, including the first record of living <Emphasis Type="Italic">Eunotogramma marginopunctatum</Emphasis>

The diversity of diatom communities found epiphytically on red macroalgae collected under the sea ice at Cape Evans, Antarctica, was studied microscopically. Comparison of communities along a depth profile showed that species diversity decreased as depth below sea ice increased. Dominant taxa also changed with depth, with Cocconeis fasciolata dominant at 10 and 15 m, Porosira glacialis at 20 m and Eunotogramma marginopunctatum at 25 m. Epiphytic communities were also compared to sympagic and sediment-associated communities collected from a single depth. Species richness was greatest for communities associated with surface sediments while the poorest was found in sympagic, or sea-ice, communities. E. marginopunctatum, previously only described from fossil material, was found associated epiphytically on macroalgae from 20 to 25 m below sea ice.     

Zooplankton boom and ice amphipod bust below melting sea ice in the Amundsen Gulf, Arctic Canada

Early summer in the Arctic with extensive ice melt and break-up represents a dramatic change for sympagic–pelagic fauna below seasonal sea ice. As part of the International Polar Year-Circumpolar Flaw Lead system study (IPY-CFL), this investigation quantified zooplankton in the meltwater layer below landfast ice and remaining ice fauna below melting ice during June (2008) in Franklin Bay and Darnley Bay, Amundsen Gulf, Canada. The ice was in a state of advanced melt, with fully developed melt ponds. Intense melting resulted in a 0.3- to 0.5-m-thick meltwater layer below the ice, with a strong halocline to the Arctic water below. Zooplankton under the ice, in and below the meltwater layer, was sampled by SCUBA divers. Dense concentrations (max. 1,400 ind. m⁻³) of Calanus glacialis were associated with the meltwater layer, with dominant copepodid stages CIV and CV and high abundance of nauplii. Less abundant species included Pseudocalanus spp., Oithona similis and C. hyperboreus. The copepods were likely feeding on phytoplankton (0.5–2.3 mg Chl-a m⁻³) in the meltwater layer. Ice amphipods were present at low abundance (<10 ind. m⁻²) and wet biomass (<0.2 g m⁻²). Onisimus glacialis and Apherusa glacialis made up 64 and 51% of the total ice faunal abundance in Darnley Bay and Franklin Bay, respectively. During early summer, the autochthonous ice fauna becomes gradually replaced by allochthonous zooplankton, with an abundance boom near the meltwater layer. The ice amphipod bust occurs during late stages of melting and break-up, when their sympagic habitat is diminished then lost.     

Productivity events of the marine diatom Thalassiosira tumida (Janisch) Hasle recorded in deglacial varves from the East Antarctic Margin

During the last deglaciation calving bay reentrants formed over several deep inner shelf basins around Antarctica as the ice sheets retreated. Marine diatoms flourished in spring and summer within the bays, providing exceptionally high silica flux rates to the basins on an annual basis. As a result, several hundred years of the late deglaciation are archived in many of these deep basins as continuously laminated diatom-rich marine sediments. The laminae are excellently preserved and offer a unique opportunity to study deglacial processes over the Antarctic shelf at very high resolution (subseasonal to seasonal at best). Annual palaeoenvironments from these sediments have been recently reconstructed. However, finer sedimentary detail is also apparent by the preservation of sublaminae of single diatom species, which represent discrete productivity events (blooms). In this paper, we focus on Thalassiosira tumida (Janisch) Hasle sublaminae preserved in the  5 m thick deglacial laminated sequence of NBP01-01 Jumbo Piston Core (JPC) 43B from Iceberg Alley ( 67°S, 63°E) on the Mac.Robertson Shelf, East Antarctic Margin. These T. tumida sublaminae are distinctive, occur several times throughout the deglacial sequence and are always preserved within summer laminae. Importantly they carry ecological and paleoceanographic information for deglacial Iceberg Alley and possibly the entire East Antarctic Margin. We describe and illustrate the T. tumida sublaminae using scanning electron microscope backscattered electron imagery of highly polished thin sections and secondary electron imagery of sublamina fracture surfaces. T. tumida sublaminae range in thickness from 0.57 to 21.07 mm (mean = 5.12; σ = 4.49) and are defined as discrete sedimentary intervals within a summer lamination where T. tumida is the dominant diatom species (abundance > 50%; but frequently > 80%) or is very abundant (abundance 40% to 49%). Based on the stratigraphic position of these sublaminae in the annual/seasonal succession and on published morphological, ecological and distributional data of living specimens in culture and in the field, we surmise an open-water, late summer bloom for T. tumida in deglacial Iceberg Alley, at sea-surface temperatures of 0 to − 1.69 °C, under low or fading light levels just prior to significant sea-ice formation. We suggest that the sublaminae formed by a combination of episodic T. tumida blooms (surface concentration), followed by rapid deposition (high flux) and optimal frustule preservation on the sea floor. Increased sublaminae frequency upcore suggests a lengthening of the summer season, while their complete cessation 43 years prior to the end of lamination deposition in outer Iceberg Alley (JPC43B) may be due to a coastward population shift.     

The Holocene Pulleniatina Minimum Event revisited: Geochemical and faunal evidence from the Okinawa Trough and upper reaches of the Kuroshio current

The Holocene Pulleniatina Minimum Event (PME) is characterized by a very low abundance of the planktonic foraminifer Pulleniatina obliquiloculata between  4.5 and 3 ka. The PME occurs widely in the Okinawa Trough and the South China Sea, and can be correlated throughout this area; it has been related to variability in the Kuroshio current. To further explore the nature of the PME, we studied cores obtained from the southern Okinawa Trough and the upper reaches of the Kuroshio current. Faunal census data indicate that all cores record the PME between  4.5 and  3 ka. The relative abundance of Neogloboquadrina dutertrei is negatively correlated to that of P. obliquiloculata in the southern Okinawa Trough, but not in the sites at the upper reaches. Mg/Ca and δ¹⁸O measurements on Globigerinoides ruber shells from the southern Okinawa Trough indicate that there was no change in sea surface temperature or sea surface salinity during the PME. The vertical structure of the water column as reconstructed by multispecies δ¹⁸O and δ¹³C profiles shows no consistent anomalies in the southern Okinawa Trough and western Philippine Sea during the PME. These observations suggest that: (1) the PME was not restricted to marginal seas, but widespread in the western North Pacific. (2) The high abundance of N. dutertrei during the PME in the Okinawa Trough may be a result of higher food-availability in the absence of P. obliquiloculata. (3) No distinctive, consistent anomalies in the paleoceanographic proxies are associated with the PME, implying there were no changes in hydrography and productivity. The absence of a linkage between faunal variation and paleoceanographic proxies indicates that we do not yet understand what causes changes in planktonic foraminiferal assemblages. This lack of understanding implies that we cannot always trust fauna-based paleothermometry at millennial timescales.     

Winter aggregations of marine mammals and birds in the north-eastern Weddell Sea pack ice

Summary A seabird and mammal census was carried out in the north-eastern Weddell Sea during the austral winter of 1986. The German research icebreaker Polarstern operated in heavy pack ice along the Greenwich Meridian between the northern sea ice boundary and the Antarctic coast. Crabeater seals (Lobodon carcinophagus), minke whales (Balaenoptera acutorostrata), Adélie penguins (Pygoscelis adeliae), Antarctic petrels (Thalassoica antarctica) and snow petrels (Pagodroma nivea) were found to be more abundant in the vicinity of the submarine Maud Rise, about 700 km north of the continental margin, than in other areas of substantial ice cover traversed during that cruise. The aggregations of birds and mammals are expected to reflect aggregations of their principal food, krill (Euphausia superba) wintering underneath the ice cover. The distribution pattern of krill predators coincides with the course of a warm water belt upwelling near Maud Rise. This upwelling could induce local ice melting which in turn may result in an increased release of sea ice algae.     

Acquisition of freeze protection in a sea-ice crustacean through horizontal gene transfer?

Sea ice is permeated by small brine channels, which are characterised by sub-zero temperatures and varying salinities. Despite sometimes extreme conditions a diverse fauna and flora thrives within the brine channels. The dominant calanoid copepods of Antarctic sea ice are Stephos longipes and Paralabidocera antarctica. Here, I report for the first time thermal hysteresis (TH) in the haemolymph of a crustacean, S. longipes, whereas P. antarctica has no such activity. TH, the non-colligative prevention of ice growth, seems to enable S. longipes to exploit all available microhabitats within sea ice, especially the surface layer, in which strong temperature fluctuations can occur. In contrast, P. antarctica only thrives within the lowermost centimetres of sea ice, where temperature fluctuations are moderate. S. longipes possesses two isoforms of a protein with TH activity. A high homology to a group of (putative) antifreeze proteins from diatoms, bacteria and a snow mold and, in contrast, no homologs in any metazoan lineage suggest that this protein was obtained through horizontal gene transfer (HGT). Further analysis of available sequence data from sea-ice organisms indicates that these antifreeze proteins were probably transferred horizontally several times. Temperature and salinity fluctuations within the brine channel system are proposed to provide “natural transformation” conditions enabling HGT and thus making this habitat a potential “hot spot” for HGT.     

Food and feeding ecology of emperor penguins in the eastern Weddell Sea

Summary The diet of the emperor penguin Aptenodytes forsteri in the eastern Weddell Sea, Antarctica was studied during October and November 1986 by stomach content analysis. Emperor penguins fed mainly on Antarctic krill Euphausia superba, Antarctic silverfish Pleuragramma antarcticum and squid Psychroteuthis glacialis. Benthic prey was not found. The prey composition suggests two different feeding strategies, shallow dives exploring the rugged underside of sea ice where krill is taken, and deep dives when mesopelagic fish and squid are consumed. Chicks were fed on average every 1.44 days.     

Post-glacial seasonal diatom record of the Mertz Glacier Polynya,East Antarctica

An ultra-high-resolution post-glacial laminated sediment record from Mertz Ninnis Trough, East Antarctic Margin (EAM), has been analysed using SEM backscattered electron imagery, secondary electron imagery and quantitative diatom abundance. Laminations are classified using visually dominant diatom species and terrigenous content. Four biogenic diatom ooze laminae types, one diatom-bearing terrigenous lamina type and one diatom-bearing terrigenous sub-lamina type have been identified. Diatom ooze lamina types comprise near-monogeneric Hyalochaete Chaetoceros spp. resting spore laminae, laminae characterised by Corethron pennatum, laminae characterised by Rhizosolenia spp. and mixed diatom assemblage laminae. Diatom-bearing terrigenous lamina and sub-lamina types comprise mixed diatom assemblage terrigenous laminae and sub-laminae characterised by Porosira glacialis resting spores. Formation of each of these lamina types is controlled by seasonal changes in nutrients, oceanographic regimes and the Mertz Glacier Polynya dynamics.     

Salinity sensitivity of early embryos of the Antarctic sea urchin, <Emphasis Type="Italic">Sterechinus neumayeri</Emphasis>

Embryos and larvae of the Antarctic sea urchin, Sterechinus neumayeri, have received considerable experimental attention assessing impacts of low temperature on development; however, salinity effects are not well documented because heretofore, the Antarctic coastal marine environment has been remarkably stenohaline. In this study, subtle decreases of 2 and 4 parts per 1,000 in standard salinity were tested to see if the developmental rate of S. neumayeri embryos would be impacted by a potential hyposmotic stress. At 30 psu, significantly fewer embryos (2 individuals out of 198 tested) reached morula stage by 36 h post-fertilization in comparison embryos in control treatments at 34 psu. Antarctic sea urchins are an important component of marine environments due to their grazing activities. Reductions in larval recruitment success due to the influx of freshwater from melting ice shelves resulting from global climate change could have far-reaching impacts on benthic ecosystem structure in Antarctica.     

Summer and spring trophic niche of larval and juvenile <Emphasis Type="Italic">Pleuragramma antarcticum</Emphasis> in the Western Ross Sea,Antarctica

The aim of this paper is to contribute to the knowledge on the feeding habits of larvae and juvenile Pleuragramma antarcticum in the western Ross Sea. In summer, the diet of P. antarcticum postlarvae (8–17 mm) was dominated by calanoid eggs (35.5%), Limacina (32.1%) and tintinnids (17.6%), while the principal food of juveniles consisted mainly of copepods (98.2%), with Oncaea curvata being the most abundant (85.1%) and the most frequently consumed prey. The food composition of P. antarcticum postlarvae (24–29 mm), collected in spring, suggest that they fed actively under the sea ice. Stephos longipes, Harpacticus furcifer and Paralabidocera antarctica sea ice copepods represent, in all their different developmental stages, the most abundant biomass food in Terra Nova Bay in this period. Our results therefore suggest that the diet of younger Pleuragramma specimens shifted in prey composition from the first summer to the following spring. This study draws attention to the key role of the copepod, P. antarctica, in the food web of Terra Nova Bay. This article belongs to a special topic: Five articles coordinated by L. Guglielmo and V. Saggiomo appear in this issue of Polar Biology and are a result of a workshop on Sea-ice communities in Terra Nova Bay (Ross Sea) held in August 2007 in Capo Calavà, Messina, Italy.