Bryozoans of the Weddell Sea continental shelf,slope and abyss: did marine life colonize the Antarctic shelf from deep water,outlying islands or in situ refugia following glaciations?

Aim At the height of glaciations such as the Last Glacial Maximum (LGM), benthic life on polar continental shelves was bulldozed off nearly all of the Antarctic shelf by grounded ice sheets. The origins of the current shelf benthos have become a subject of considerable debate. There are several possible sources for the current Antarctic shelf fauna, the first of which is the continental slope and deep sea of the Southern Ocean. The high levels of reported eurybathy for many Antarctic species are taken as evidence supporting this. A second possible source for colonists is the southern margins of other continents. Finally, shelves could have been recolonized from refugia on the continental shelves or slopes around Antarctica. The current study investigates whether the patchily rich and abundant biota that now occurs on the Antarctic continental shelf recolonized from refugia in situ or elsewhere. Location Weddell Sea, Antarctica. Methods We examined bryozoan samples of the BENDEX, ANDEEP III and SYSTCO expeditions, as well as the literature. Using similarity matrices (Sørensen coefficient), we assessed similarities of benthos sampled from around Antarctica. By assessing numbers of species shared between differing depths and adjacent shelf areas, we evaluated the origins of cheilostome bryozoan communities. Results Bryozoans decreased from 28, 6.5 and 0.3 colonies per trawl, and 0.16, 0.046 and 0.0026 colonies per cm² of hard surface from shelf to slope to abyssal depths. We found little and no support for recolonization of the Weddell Sea shelf by bryozoans from the adjacent slope and abyss, in the scenario of LGM faunal wipe‐out. The Weddell Sea shelf bryozoan fauna was considerably more similar to those on other Antarctic shelves than to that of the adjacent (Weddell Sea) continental slope. The known bryozoan fauna of the Weddell Sea shelf is not a subset of the Weddell Sea slope or abyssal faunas. Main conclusions We consider that the composition of the current Weddell Sea bryozoan fauna is most easily explained by in situ survival. Thus we consider that at least some of the Weddell Sea fauna persisted throughout the LGM, although not necessarily at the same locations throughout, to recolonize the large area currently occupied.     

Detecting glacial refugia in the Southern Ocean

Throughout the Quaternary, the continental-based Antarctic ice sheets expanded and contracted repeatedly. Evidence suggests that during glacial maxima, grounded ice eliminated most benthic (bottom-dwelling) fauna across the Antarctic continental shelf. However, paleontological and molecular evidence indicates most extant Antarctica benthic taxa have persisted in situ throughout the Quaternary. Where and how the Antarctic benthic fauna survived throughout repeated glacial maxima remain mostly hypothesised. If understood, this would provide valuable insights into the ecology and evolution of Southern Ocean biota over geological timescales. Here we synthesised and appraised recent studies and presented an approach to demonstrate how genetic data can be effective in identifying where and how Antarctic benthic fauna survived glacial periods. We first examined the geological and ecological evidence for how glacial periods influenced past species demography in order to provide testable frameworks for future studies. We outlined past ice-free areas from Antarctic ice sheet reconstructions that could serve as glacial refugia and discussed how benthic fauna with pelagic or non-pelagic dispersal strategies moved into and out of glacial refugia. We also reviewed current molecular studies and collated proposed locations of Southern Ocean glacial refugia on the continental shelf around Antarctica, in the deep sea, and around sub-Antarctic islands. Interestingly, the proposed glacial refugia based on molecular data generally do not correspond to the ice-free areas identified by Antarctic ice sheet reconstructions. The potential biases in sampling and in the choice of molecular markers in current literature are discussed, along with the future directions for employing testable frameworks and genomic methods in Southern Ocean molecular studies. Continued data syntheses will elucidate greater understanding of where and how Southern Ocean benthic fauna persisted throughout glacial periods and provide insights into their resilience against climate changes in the future.     

Biodiversity and biogeographic affiliation of Bryozoa from King George Island (Antarctica)

King George Island (KGI), which is located between the Antarctic and South American continents, may play a crucial role in the exchange of Bryozoa amongst the various Antarctic sectors and across the Polar Front. Knowledge regarding the biological diversity of this area could help us understand the evolution of the Antarctic ecosystem and its connectivity to the South American continent as well as the colonization ability of particular species. Here, we investigate the patterns of diversity and biogeographic affiliation of the cheilostome Bryozoa from KGI and the surrounding areas. Of 114 identified taxa from a depth range of 6–492 m, 26 species were reported for the first time in KGI. The most speciose genera were Camptoplites, Osthimosia, Smittina, and Cellarinella. Species richness at KGI consisted of 70% of the total bryozoans at the South Shetland Islands (SSI). Fifty-nine per cent of the bryozoans from KGI are endemic to Antarctica, which closely reflects the previously estimated endemism rate for bryozoans and other Antarctic taxa. Cluster analysis indicated that the strongest faunal links of SSI bryozoans were with Antarctic Peninsula assemblages, corresponding to the physical distance between both locations. The biogeographic similarities between SSI and South America confirm the broad trend of existing Antarctic–South American faunal links previously observed in bryozoans and many other taxa and indicate that SSI might be an important transitional zone between Antarctica and South America.     

Antarctic terrestrial life--challenging the history of the frozen continent?

Antarctica is a continent locked in ice, with almost 99.7% of current terrain covered by permanent ice and snow, and clear evidence that, as recently as the Last Glacial Maximum (LGM), ice sheets were both thicker and much more extensive than they are now. Ice sheet modelling of both the LGM and estimated previous ice maxima across the continent give broad support to the concept that most if not all currently ice-free ground would have been overridden during previous glaciations. This has given rise to a widely held perception that all Mesozoic (pre-glacial) terrestrial life of Antarctica was wiped out by successive and deepening glacial events. The implicit conclusion of such destruction is that most, possibly all, contemporary terrestrial life has colonised the continent during subsequent periods of glacial retreat. However, several recently emerged and complementary strands of biological and geological research cannot be reconciled comfortably with the current reconstruction of Antarctic glacial history, and therefore provide a fundamental challenge to the existing paradigms. Here, we summarise and synthesise evidence across these lines of research. The emerging fundamental insights corroborate substantial elements of the contemporary Antarctic terrestrial biota being continuously isolated in situ on a multi-million year, even pre-Gondwana break-up timescale. This new and complex terrestrial Antarctic biogeography parallels recent work suggesting greater regionalisation and evolutionary isolation than previously suspected in the circum-Antarctic marine fauna. These findings both require the adoption of a new biological paradigm within Antarctica and challenge current understanding of Antarctic glacial history. This has major implications for our understanding of the key role of Antarctica in the Earth System.     

Bacteria beneath the West Antarctic Ice Sheet

Subglacial environments, particularly those that lie beneath polar ice sheets, are beginning to be recognized as an important part of Earth's biosphere. However, except for indirect indications of microbial assemblages in subglacial Lake Vostok, Antarctica, no sub-ice sheet environments have been shown to support microbial ecosystems. Here we report 16S rRNA gene and isolate diversity in sediments collected from beneath the Kamb Ice Stream, West Antarctic Ice Sheet and stored for 15 months at 4°C. This is the first report of microbes in samples from the sediment environment beneath the Antarctic Ice Sheet. The cells were abundant (∼10⁷ cells g⁻¹) but displayed low diversity (only five phylotypes), likely as a result of enrichment during storage. Isolates were cold tolerant and the 16S rRNA gene diversity was a simplified version of that found in subglacial alpine and Arctic sediments and water. Although in situ cell abundance and the extent of wet sediments beneath the Antarctic ice sheet can only be roughly extrapolated on the basis of this sample, it is clear that the subglacial ecosystem contains a significant and previously unrecognized pool of microbial cells and associated organic carbon that could potentially have significant implications for global geochemical processes.     

Biodiversity and biogeography of southern temperate and polar bryozoans

Aim To describe the distribution of biodiversity and endemism of bryozoans in southern temperate and polar waters. We hypothesized that we would find: (1) no strong latitudinal richness gradient; (2) striking contrasts in richness and endemism between clades and between regions; and (3) that faunal similarity of regions would cluster geographically around each southern continent. Location South Atlantic, Indian and Pacific Oceans and the Southern Ocean. Methods We constructed a data base from known literature, regional data bases and recent finds. We regionalized each southern continent, calculated levels of richness and endemism for each region and continent, and used primer 5 to perform multivariate statistical analysis. Results A third (1681) of global bryozoan species described occur south of 30° S, of which c. 87% were cheilostomes. In richness we found no latitudinal cline and change across longitude was stronger. New Zealand was richest and had the most (60%) endemic species, followed by Antarctica at 57%. There were striking contrasts in regional richness and endemism between clades but the highest levels of between‐region similarity were around Antarctica. The timing of past continent connectivity was reflected. Main conclusions Bryozoans show strong hemispherical asymmetry in richness and, like molluscs and corals, decrease away from Australasia rather than with latitude. Species endemism is much lower in Antarctic bryozoans than previously thought, and as this taxon is not particularly dispersive and is now amongst the best studied regionally, maybe Antarctic endemism in general is lower and Antarctica less cut‐off to species dispersal than previously thought. However, Antarctic generic endemism is double the level previously calculated and regional faunal similarities are much higher than around other continents – both reflecting long‐term isolation. Bryozoans, in contrast to the paradigm of Antarctic fauna, may be fairly robust to predicted climate change. Paradoxically, they may also be one of the best taxa to monitor to sensitively detect marine benthic responses.     

Persistent genetic signatures of historic climatic events in an Antarctic octopus

Repeated cycles of glaciation have had major impacts on the distribution of genetic diversity of the Antarctic marine fauna. During glacial periods, ice cover limited the amount of benthic habitat on the continental shelf. Conversely, more habitat and possibly altered seaways were available during interglacials when the ice receded and the sea level was higher. We used microsatellites and partial sequences of the mitochondrial cytochrome oxidase 1 gene to examine genetic structure in the direct‐developing, endemic Southern Ocean octopod Pareledone turqueti sampled from a broad range of areas that circumvent Antarctica. We find that, unusually for a species with poor dispersal potential, P. turqueti has a circumpolar distribution and is also found off the islands of South Georgia and Shag Rocks. The overriding pattern of spatial genetic structure can be explained by hydrographic (with ocean currents both facilitating and hindering gene flow) and bathymetric features. The Antarctic Peninsula region displays a complex population structure, consistent with its varied topographic and oceanographic influences. Genetic similarities between the Ross and Weddell Seas, however, are interpreted as a persistent historic genetic signature of connectivity during the hypothesized Pleistocene West Antarctic Ice Sheet collapses. A calibrated molecular clock indicates two major lineages within P. turqueti, a continental lineage and a sub‐Antarctic lineage, that diverged in the mid‐Pliocene with no subsequent gene flow. Both lineages survived subsequent major glacial cycles. Our data are indicative of potential refugia at Shag Rocks and South Georgia and also around the Antarctic continent within the Ross Sea, Weddell Sea and off Adélie Land. The mean age of mtDNA diversity within these main continental lineages coincides with Pleistocene glacial cycles.     

Faunal migration into the Late Permian Zechstein Basin - Evidence from bryozoan palaeobiogeography

Late Permian bryozoans from the Wegener Halvø, Ravnefjeld and Schuchert Formations in East Greenland have been investigated. 14 genera are recognised.Integration of the new bryozoan data from the Upper Permian of East Greenland with data on the distribution of Permian bryozoans along the northern margin of Pangea is used to test hypotheses concerning Late Palaeozoic evolution of the North Atlantic region. During the Permian, the Atlantic rift system formed a seaway between Norway and Greenland from the boreal Barents Shelf to the warm and arid Zechstein Basin. This seaway is considered to be the only marine connection to the Zechstein Basin and therefore the only possible migration route for bryozoans to enter the basin. The distribution of Permian bryozoans is largely in keeping with such a connection from the cool Barents Shelf past the East Greenland Basin to the warm Zechstein Basin and also corroborates the change in temperature through this connection.     

Benthic palaeoecology of Danian deep-shelf bryozoan mounds in the Danish Basin

Early Danian cool-water bryozoan mounds exposed in the coastal cliff Stevns Klint in Denmark were formed shortly after the Cretaceous–Tertiary mass extinction. They represent a relatively deep-water, highly diverse benthic ecosystem within the epeiric seaway that covered the Danish Basin. The mounds are 50–110 m long and reached a height of about 5–10 m above the seafloor; they are asymmetrical with a steep southern and a gentle northern flank, and were dominated by small suspension feeders. The benthic elements generally occur as fragments set in a carbonate mud matrix. The main skeletal contributors are delicate branching bryozoans with minor contributions of bryozoan sheets, and nodular/arborescent bryozoans. Locally abundant octocorals occur on the mound crests and upper parts of the steep flanks. Echinoids are present in minor amounts, but are locally abundant. Serpulids, crinoids, asteroids, brachiopods, bivalves, massive calcareous sponges, and benthic foraminifers are generally minor contributors to the benthic mound fauna. Influx of planktonic foraminifers, coccoliths and other planktonic organisms was high and was probably a major source of nutrient supply to the mainly suspension-feeding benthic fauna.

The faunal association reflects a relatively low energy environment with a high, possibly seasonal influx of particulate nutrients. The best growth conditions with respect to nutrient influx were on the mound crest and upper steep flank reflected by the diverse and relatively largest benthic faunal elements. Periodic reworking and winnowing occurred across the entire mound structure but most prominent on the gentle northern flanks limiting the benthic growth and notably the colony density and size of delicate branching bryozoans. Vagile benthic faunas were also adapted to different areas on the mound. Irregular echinoids preferred the intermound areas within fine-grained wackestone–packstone facies where they ploughed through the sediment, whereas regular echinoids were epifaunal and preferred the upper parts of the mounds, possibly feeding mainly on bryozoans. Skeletons of both groups became concentrated at the toe of the steep flanks and in the intermound areas by physical reworking during major storms.

Changes in faunal composition on the mound crests occurred rhythmically on both small and large scale during mound growth. Rhythmically recurring faunal assemblages reflect alternating hydrodynamic conditions on the seafloor with respect to nutrient influx and energy, which probably were linked to short-term seasonal and long-term climatic variations; the long-term alternation may be within the Milankovitch frequency band. Blooming events of bryozoan sheets resulted from relatively short periods with large amounts of available food and suitable substrate. Successful colonisation by octocorals on the other hand reflected longer-term favourable conditions on the mounds possibly associated with overall higher energy levels.

A possible Pleistocene analogue to the bryozoan-dominated Danian mounds occurs at the shelf-slope break of the Great Australian Bight. Both of these cool-water mound systems deviate from most other biogenic mounds known from the fossil record in their non-cemented nature, regular geometry and a lack of core and flank facies.     

Extinction and recolonization of maritime Antarctica in the limpet Nacella concinna (Strebel, 1908) during the last glacial cycle: toward a model of Quaternary biogeography in shallow Antarctic invertebrates

Quaternary glaciations in Antarctica drastically modified geographical ranges and population sizes of marine benthic invertebrates and thus affected the amount and distribution of intraspecific genetic variation. Here, we present new genetic information in the Antarctic limpet Nacella concinna, a dominant Antarctic benthic species along shallow ice‐free rocky ecosystems. We examined the patterns of genetic diversity and structure in this broadcast spawner along maritime Antarctica and from the peri‐Antarctic island of South Georgia. Genetic analyses showed that N. concinna represents a single panmictic unit in maritime Antarctic. Low levels of genetic diversity characterized this population; its median‐joining haplotype network revealed a typical star‐like topology with a short genealogy and a dominant haplotype broadly distributed. As previously reported with nuclear markers, we detected significant genetic differentiation between South Georgia Island and maritime Antarctica populations. Higher levels of genetic diversity, a more expanded genealogy and the presence of more private haplotypes support the hypothesis of glacial persistence in this peri‐Antarctic island. Bayesian Skyline plot and mismatch distribution analyses recognized an older demographic history in South Georgia. Approximate Bayesian computations did not support the persistence of N. concinna along maritime Antarctica during the last glacial period, but indicated the resilience of the species in peri‐Antarctic refugia (South Georgia Island). We proposed a model of Quaternary Biogeography for Antarctic marine benthic invertebrates with shallow and narrow bathymetric ranges including (i) extinction of maritime Antarctic populations during glacial periods; (ii) persistence of populations in peri‐Antarctic refugia; and (iii) recolonization of maritime Antarctica following the deglaciation process.     

Correlation of Early Pliocene diatomite to low amplitude Milankovitch cycles in the ANDRILL AND-1B drill core

In the austral summer of 2006/7 the ANDRILL MIS (ANtarctic geological DRILLing-McMurdo Ice Shelf) project recovered a 1285 m sediment core from beneath the Ross Ice Shelf near Ross Island, Antarctica in a flexural moat associated with volcanic loading. The upper ~ 600 m of this core contain sediments recording 38 glacial/interglacial cycles of Early Pliocene to Pleistocene time, including 13 discrete diatomite units (DUs). The longest of these, DU XI, is ~ 76 m-thick, and has been assigned an Early to Mid-Pliocene age (5–3 Ma). A detailed record of the siliceous microfossil assemblages in DU XI is used in conjunction with geochemical and sedimentological data to subdivide DU XI into four discrete subunits of continuous sedimentation. Within each subunit, changes in diatom assemblages have been correlated with the δ¹⁸O record, providing a temporal resolution up to 600 yr, and allowing for the construction of a detailed age model and calculation of associated sediment accumulation rates within DU XI. Results indicate a productivity-dominated sedimentary record with greater proportions of hemipelagic mud accumulating during relatively cool periods. This implies that even during periods of substantial warmth, Milankovitch-paced changes in Antarctic ice volume can be linked to ecological changes recorded in diatom assemblages.     

Incursion and excursion of Antarctic biota: past, present and future

Aim To investigate the major paradigms of intense isolation and little anthropogenic influence around Antarctica and to examine the timings and scales of the modification of the southern polar biota. Location Antarctica and surrounding regions. Methods First, mechanisms of and evidence for long‐term isolation are reviewed. These include continental drift, the development of a surrounding deep‐water channel and the Antarctic Circumpolar Current (ACC). They also include levels of endemism, richness and distinctiveness of assemblages. Secondly, evidence for past and modern opportunities for species transport are investigated. Comparative levels of alien establishments are also examined around the Southern Ocean. Discussion On a Cenozoic time‐scale, it is clear that Gondwana's fragmentation led to increasing geographical isolation of Antarctica and the initiation of the ACC, which restricted biota exchange to low levels while still permitting some movement of biota. On a shorter Quaternary time‐scale, the continental ice‐sheet, influenced by solar (Milankovitch) cycles, has expanded and contracted periodically, covering and exposing terrestrial and continental shelf habitats. There were probably refugia for organisms during each glacial maxima. It is also likely that new taxa were introduced into Antarctica during cycles of ice sheet and oceanic front movement. The current situation (a glacial minimum) is not ‘normal’; full interglacials represent only 10% of the last 430 ka. On short (ecological) time‐scales, many natural dispersal processes (airborne, oceanic eddy, rafting and hitch‐hiking on migrants) enable the passage of biota to and from Antarctica. In recent years, humans have become influential both directly by transporting organisms and indirectly by increasing survival and establishment prospects via climate change. Main conclusions Patterns of endemism and alien establishment are very different across taxa, land and sea, and north vs. south of the Polar Frontal Zone. Establishment conditions, as much as transport, are important in limiting alien establishment. Three time‐scales emerge as important in the modification of Antarctica's biota. The natural ‘interglacial’ process of reinvasion of Antarctica is being influenced strongly by humans.     

Bryozoan colonization of the marine isopod Glyptonotus antarcticus at Signy Island, Antarctica

Sixty specimens of the giant marine isopod Glyptonotus antarcticus Eights, collected from Borge Bay, Signy Island, Antarctica were examined for epizoans. Ten species of cheilostomatid bryozoans were found on the isopods. The purpose of the study was to quantify the prevalence, intensity, abundance, and spatial distribution of the bryozoans on the isopods. The proportion of isopods colonized was 42%. The larger isopods had both significantly more epizoic bryozoan colonies and species. The greatest density of bryozoans was on the fused pleon and telson. There was no significant difference between the dorsal and ventral abundance of bryozoan colonies. The diversity of epizoic bryozoans on the isopods is higher than on other host organisms from more stable environments. This may be because of active selection by settling larvae. The frequency of local substrata being scoured by ice is high around Signy Island, so there may be a selective advantage in colonizing a motile host. Accepted: 20 June 1998     

Climate change,glacier retreat and a new ice‐free island offer new insights on Antarctic benthic responses

The Antarctic Peninsula is among the places on Earth that registered major warming in the last 60 yr. Massive ice losses, represented by glacier retreat, ice‐shelf collapses and sea‐ice reduction are among the main impacts of this regional warming. The loss of sea‐bed ice coverage, on the one hand has been affecting benthic assemblages, but on the other it is opening up new areas for benthic colonisation. Potter Cove (South Shetland Islands) offered the opportunity of assessing both processes. We recently reported a sudden shift of benthic assemblages related to increased sedimentation rates caused by glacier retreat. This glacier retreat also uncovered a new island that presents a natural experiment to study Antarctic benthic colonisation and succession. We sampled the new island by photo‐transects taken up to 30 m depth. Here, we report an unexpected benthic assemblage characterised by high species richness, diversity and structural complexity with a well‐developed three‐dimensional structure and epibiotic relationships. Filter feeders comprised the largest trophic group at all depths, mainly ascidians, sponges and bryozoans. Densities were also surprising, recording only six ascidian species with a mean of ～310 ind. m^–2. These values are at least an order of magnitude higher than previous Antarctic reports on early colonisation. This finding challenges the extended idea of a slow and continuous recruitment in Antarctica. However, it also opens the question of whether these complex assemblages could have been present under the glacier in ice‐free refuges that are now exposed to open sea conditions. Under the current scenario of climate change, these results acquire high relevance as they suggest a two‐fold effect of the Antarctic Peninsula warming: the environmental shifts that threaten coastal ecosystems, and also the opening up of new areas for colonisation that may occur at a previously unimagined speed.     

Why is the South Orkney Island shelf (the world's first high seas marine protected area) a carbon immobilization hotspot?

The Southern Ocean archipelago, the South Orkney Islands (SOI), became the world's first entirely high seas marine protected area (MPA) in 2010. The SOI continental shelf (~44 000 km²), was less than half covered by grounded ice sheet during glaciations, is biologically rich and a key area of both sea surface warming and sea‐ice losses. Little was known of the carbon cycle there, but recent work showed it was a very important site of carbon immobilization (net annual carbon accumulation) by benthos, one of the few demonstrable negative feedbacks to climate change. Carbon immobilization by SOI bryozoans was higher, per species, unit area and ice‐free day, than anywhere‐else polar. Here, we investigate why carbon immobilization has been so high at SOI, and whether this is due to high density, longevity or high annual production in six study species of bryozoans (benthic suspension feeders). We compared benthic carbon immobilization across major regions around West Antarctica with sea‐ice and primary production, from remotely sensed and directly sampled sources. Lowest carbon immobilization was at the northernmost study regions (South Georgia) and southernmost Amundsen Sea. However, data standardized for age and density showed that only SOI was anomalous (high). High immobilization at SOI was due to very high annual production of bryozoans (rather than high densities or longevity), which were 2x, 3x and 5x higher than on the Bellingshausen, South Georgia and Amundsen shelves, respectively. We found that carbon immobilization correlated to the duration (but not peak or integrated biomass) of phytoplankton blooms, both in directly sampled, local scale data and across regions using remote‐sensed data. The long bloom at SOI seems to drive considerable carbon immobilization, but sea‐ice losses across West Antarctica mean that significant carbon sinks and negative feedbacks to climate change could also develop in the Bellingshausen and Amundsen seas.     

Late pleistocene paleoclimates of the South Pacific based on statistical analysis of planktonic foraminifers

Statistical analysis applied to foraminiferal data from 78 South Pacific core tops enables the derivation of a transfer function that relates sea surface temperature to foraminiferal assemblages. Application of this transfer function to eight cores from the southern part of the East Pacific Rise yields estimates of the sea surface temperatures of the last glacial maximum, as well as the paleotemperature record of the past 150,000 years. Comparison of the last glacial temperature estimates with the recent sea surface temperature shows that the greatest change between glacial and present conditions (about 5°C) occurs in a climatically sensitive area near 50°S. Stratigraphic correlation of two cores from this area suggests that the last glacial started in this area with rapid cooling, and that the climate stayed generally cold until the end of the glacial. Similar general shape of the climatic record is found in the high latitudes of the North Atlantic as well as in the ice sheets of Greenland and Antarctica. In contrast to the similarity in the shape of these high-latitude records, they differ distinctly from the foraminiferal oxygen isotope record of several deep-sea cores which indicates a general gradual increase of ice volume from the beginning of the last glacial to the maximum glaciation which occurred about 18,000 years B.P.In the study area the rate of sediment accumulation during the last glacial is about two to three times less than in the last interglacial. There is no indication of increased carbonate solution during the glacial, and it is suggested that the change in the accumulation rate results from a reduction in the supply of calcareous shells to the sediment. It seems that with cooling, the environment becomes less favorable to organisms producing calcium carbonate tests, and therefore carbonate production decreases during the glacial.     

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.     

Have Historical Climate Changes Affected Gentoo Penguin (Pygoscelis papua) Populations in Antarctica?

The West Antarctic Peninsula (WAP) has been suffering an increase in its atmospheric temperature during the last 50 years, mainly associated with global warming. This increment of temperature trend associated with changes in sea-ice dynamics has an impact on organisms, affecting their phenology, physiology and distribution range. For instance, rapid demographic changes in Pygoscelis penguins have been reported over the last 50 years in WAP, resulting in population expansion of sub-Antarctic Gentoo penguin (P. papua) and retreat of Antarctic Adelie penguin (P. adeliae). Current global warming has been mainly associated with human activities; however these climate trends are framed in a historical context of climate changes, particularly during the Pleistocene, characterized by an alternation between glacial and interglacial periods. During the last maximal glacial (LGM∼21,000 BP) the ice sheet cover reached its maximum extension on the West Antarctic Peninsula (WAP), causing local extinction of Antarctic taxa, migration to lower latitudes and/or survival in glacial refugia. We studied the HRVI of mtDNA and the nuclear intron βfibint7 of 150 individuals of the WAP to understand the demographic history and population structure of P. papua. We found high genetic diversity, reduced population genetic structure and a signature of population expansion estimated around 13,000 BP, much before the first paleocolony fossil records (∼1,100 BP). Our results suggest that the species may have survived in peri-Antarctic refugia such as South Georgia and North Sandwich islands and recolonized the Antarctic Peninsula and South Shetland Islands after the ice sheet retreat.     

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.     

Animal survival strategies in Neoproterozoic ice worlds

The timing of the first appearance of animals is of crucial importance for understanding the evolution of life on Earth. Although the fossil record places the earliest metazoans at 572–602 Ma, molecular clock studies suggest a far earlier origination, as far back as ~850 Ma. The difference in these dates would place the rise of animal life into a time period punctuated by multiple colossal, potentially global, glacial events. Although the two schools of thought debate the limitations of each other's methods, little time has been dedicated to how animal life might have survived if it did arise before or during these global glacial periods. The history of recent polar biota shows that organisms have found ways of persisting on and around the ice of the Antarctic continent throughout the Last Glacial Maximum (33–14 Ka), with some endemic species present before the breakup of Gondwana (180–23 Ma). Here we discuss the survival strategies and habitats of modern polar marine organisms in environments analogous to those that could have existed during Neoproterozoic glaciations. We discuss how, despite the apparent harshness of many ice covered, sub-zero, Antarctic marine habitats, animal life thrives on, in and under the ice. Ice dominated systems and processes make some local environments more habitable through water circulation, oxygenation, terrigenous nutrient input and novel habitats. We consider how the physical conditions of Neoproterozoic glaciations would likely have dramatically impacted conditions for potential life in the shallows and erased any possible fossil evidence from the continental shelves. The recent glacial cycle has driven the evolution of Antarctica's unique fauna by acting as a “diversity pump,” and the same could be true for the late Proterozoic and the evolution of animal life on Earth, and the existence of life elsewhere in the universe on icy worlds or moons.