Trophic plasticity of Antarctic echinoids under contrasted environmental conditions

Echinoids are common members of Antarctic zoobenthos, and different groups can show important trophic diversity. As part of the ANT-XXIX/3 cruise of RV Polarstern, trophic plasticity of sea urchins was studied in three neighbouring regions (Drake Passage, Bransfield Strait and Weddell Sea) featuring several depth-related habitats offering different trophic environments to benthic consumers. Three families with contrasting feeding habits (Cidaridae, Echinidae and Schizasteridae) were studied. Gut content examination and stable isotopes ratios of C and N suggest that each of the studied families showed a different response to variation in environmental and food conditions. Schizasteridae trophic plasticity was low, and these sea urchins were bulk sediment feeders relying on sediment-associated organic matter in all regions and/or depth-related habitats. Cidaridae consumed the most animal-derived material. Their diet varied according to the considered area, as sea urchins from Bransfield Strait relied mostly on living and/or dead animal material, while specimens from Weddell Sea fed on a mixture of dead animal material and other detritus. Echinidae also showed important trophic plasticity. They fed on various detrital items in Bransfield Strait, and selectivity of ingested material varied across depth-related habitats. In Weddell Sea, stable isotopes revealed that they mostly relied on highly ¹³C-enriched food items, presumably microbially reworked benthic detritus. The differences in adaptive strategies could lead to family-specific responses of Antarctic echinoids to environmental and food-related changes.     

Antarctic, Sub-Antarctic and cold temperate echinoid database

This database includes spatial data of Antarctic, Sub-Antarctic and cold temperate echinoid distribution (Echinodermata: Echinoidea) collected during many oceanographic campaigns led in the Southern Hemisphere from 1872 to 2010. The dataset lists occurrence data of echinoid distribution south of 35°S latitude, together with information on taxonomy (from species to genus level), sampling sources (cruise ID, sampling dates, ship names) and sampling sites (geographic coordinates and depth). Echinoid occurrence data were compiled from the Antarctic Echinoid Database (David et al. 2005a), which integrates records from oceanographic cruises led in the Southern Ocean until 2003. This database has been upgraded to take into account data from oceanographic cruises led after 2003. The dataset now reaches a total of 6160 occurrence data that have been checked for systematics reliability and consistency. It constitutes today the most complete database on Antarctic and Sub-Antarctic echinoids.     

Will krill fare well under Southern Ocean acidification?

Antarctic krill embryos and larvae were experimentally exposed to 380 (control), 1000 and 2000 µatm pCO₂ in order to assess the possible impact of ocean acidification on early development of krill. No significant effects were detected on embryonic development or larval behaviour at 1000 µatm pCO₂; however, at 2000 µatm pCO₂ development was disrupted before gastrulation in 90 per cent of embryos, and no larvae hatched successfully. Our model projections demonstrated that Southern Ocean sea water pCO₂ could rise up to 1400 µatm in krill''s depth range under the IPCC IS92a scenario by the year 2100 (atmospheric pCO₂ 788 µatm). These results point out the urgent need for understanding the pCO₂-response relationship for krill developmental and later stages, in order to predict the possible fate of this key species in the Southern Ocean.     

Interannual pteropod variability in sediment traps deployed above and below the aragonite saturation horizon in the Sub-Antarctic Southern Ocean

Anthropogenic inputs of CO₂ are altering ocean chemistry and may alter the role of marine calcifiers in ocean ecosystems. Laboratory research and ocean models suggest calcifiers in polar waters are especially at risk, particularly pteropods: pelagic aragonite-shelled molluscs. However, baseline data for natural populations of pteropods are limited, especially for polar and sub-polar waters. In order to establish baseline data on diversity, preservation state and shell flux of in situ populations of Sub-Antarctic Southern Ocean pteropods, we deployed sediment traps above (1,000 m) and below (2,000 m) the aragonite saturation horizon (ASH) (currently at 1,200 m) from 1997 to 2006 at 47°S, 142°E. We identified seven pteropod taxa. We applied a shell opacity index to each shell collected and found 50% of shells collected above the ASH to be in pristine condition but only 3% of the shells collected below the ASH showed such a high degree of preservation. We estimated pteropod shell mass fluxes for the region (0.17–4.99 mg m⁻² day⁻¹), and we identified significant reductions in shell flux for Limacina helicina antarctica forma rangi and Clio recurva to the trap series above the ASH and for Limacina helicina antarctica forma rangi and Limacina helicina antarctica forma antarctica to the trap series below the ASH over the interval 1997–2006. Our data establish a temporal and vertical snapshot of the current Sub-Antarctic pelagic pteropod community and provide a baseline against which to monitor Southern Ocean pteropods responses, if any, to changing ocean conditions projected for the region in the coming decades.     

Antarctic Marine Biodiversity – What Do We Know About the Distribution of Life in the Southern Ocean?

The remote and hostile Southern Ocean is home to a diverse and rich community of life that thrives in an environment dominated by glaciations and strong currents. Marine biological studies in the region date back to the nineteenth century, but despite this long history of research, relatively little is known about the complex interactions between the highly seasonal physical environment and the species that inhabit the Southern Ocean. Oceanographically, the Southern Ocean is a major driver of global ocean circulation and plays a vital role in interacting with the deep water circulation in each of the Pacific, Atlantic, and Indian oceans. The Census of Antarctic Marine Life and the Scientific Committee on Antarctic Research Marine Biodiversity Information Network (SCAR-MarBIN) have strived to coordinate and unify the available scientific expertise and biodiversity data to improve our understanding of Southern Ocean biodiversity. Taxonomic lists for all marine species have been compiled to form the Register of Antarctic Marine Species, which currently includes over 8,200 species. SCAR-MarBIN has brought together over 1 million distribution records for Southern Ocean species, forming a baseline against which future change can be judged. The sample locations and numbers of known species from different regions were mapped and the depth distributions of benthic samples plotted. Our knowledge of the biodiversity of the Southern Ocean is largely determined by the relative inaccessibility of the region. Benthic sampling is largely restricted to the shelf; little is known about the fauna of the deep sea. The location of scientific bases heavily influences the distribution pattern of sample and observation data, and the logistical supply routes are the focus of much of the at-sea and pelagic work. Taxa such as mollusks and echinoderms are well represented within existing datasets with high numbers of georeferenced records. Other taxa, including the species-rich nematodes, are represented by just a handful of digital records.     

Ocean acidification stimulates particulate organic carbon accumulation in two Antarctic diatom species under moderate and high natural solar radiation

Impacts of rising atmospheric CO₂ concentrations and increased daily irradiances from enhanced surface water stratification on phytoplankton physiology in the coastal Southern Ocean remain still unclear. Therefore, in the two Antarctic diatoms Fragilariopsis curta and Odontella weissflogii, the effects of moderate and high natural solar radiation combined with either ambient or future pCO₂ on cellular particulate organic carbon (POC) contents and photophysiology were investigated. Results showed that increasing CO₂ concentrations had greater impacts on diatom physiology than exposure to increasing solar radiation. Irrespective of the applied solar radiation regime, cellular POC quotas increased with future pCO₂ in both diatoms. Lowered maximum quantum yields of photochemistry in PSII (F_v/F_m) indicated a higher photosensitivity under these conditions, being counteracted by increased cellular concentrations of functional photosynthetic reaction centers. Overall, our results suggest that both bloom‐forming Antarctic coastal diatoms might increase carbon contents under future pCO₂ conditions despite reduced physiological fitness. This indicates a higher potential for primary productivity by the two diatom species with important implications for the CO₂ sequestration potential of diatom communities in the future coastal Southern Ocean.     

Ammonia-oxidizing Archaea in the Arctic Ocean and Antarctic coastal waters

We compared abundance, distributions and phylogenetic composition of Crenarchaeota and ammonia-oxidizing Archaea (AOA) in samples collected from coastal waters west of the Antarctic Peninsula during the summers of 2005 and 2006, with samples from the central Arctic Ocean collected during the summer of 1997. Ammonia-oxidizing Archaea and Crenarchaeota abundances were estimated from quantitative PCR measurements of amoA and 16S rRNA gene abundances. Crenarchaeota and AOA were approximately fivefold more abundant at comparable depths in the Antarctic versus the Arctic Ocean. Crenarchaeota and AOA were essentially absent from the Antarctic Summer Surface Water (SSW) water mass (0–45 m depth). The ratio of Crenarchaeota 16S rRNA to archaeal amoA gene abundance in the Winter Water (WW) water mass (45–105 m depth) of the Southern Ocean was much lower (0.15) than expected and in sharp contrast to the ratio (2.0) in the Circumpolar Deep Water (CDW) water mass (105–3500 m depth) immediately below it. We did not observe comparable segregation of this ratio by depth or water mass in Arctic Ocean samples. A ubiquitous, abundant and polar-specific crenarchaeote was the dominant ribotype in the WW and important in the upper halocline of the Arctic Ocean. Our data suggest that this organism does not contain an ammonia monooxygenase gene. In contrast to other studies where Crenarchaeota populations apparently lacking amoA genes are found in bathypelagic waters, this organism appears to dominate in well-defined, ammonium-rich, near-surface water masses in polar oceans.     

Adult antarctic krill feeding at abyssal depths

Antarctic krill (Euphausia superba) is a large euphausiid, widely distributed within the Southern Ocean [1], and a key species in the Antarctic food web [2]. The Discovery Investigations in the early 20(th) century, coupled with subsequent work with both nets and echosounders, indicated that the bulk of the population of postlarval krill is typically confined to the top 150 m of the water column [1, 3, 4]. Here, we report for the first time the existence of significant numbers of Antarctic krill feeding actively at abyssal depths in the Southern Ocean. Biological observations from the deep-water remotely operated vehicle Isis in the austral summer of 2006/07 have revealed the presence of adult krill (Euphausia superba Dana), including gravid females, at unprecedented depths in Marguerite Bay, western Antarctic Peninsula. Adult krill were found close to the seabed at all depths but were absent from fjords close inshore. At all locations where krill were detected they were seen to be actively feeding, and at many locations there were exuviae (cast molts). These observations revise significantly our understanding of the depth distribution and ecology of Antarctic krill, a central organism in the Southern Ocean ecosystem.     

Pacific-wide contrast highlights resistance of reef calcifiers to ocean acidification

Ocean acidification (OA) and its associated decline in calcium carbonate saturation states is one of the major threats that tropical coral reefs face this century. Previous studies of the effect of OA on coral reef calcifiers have described a wide variety of outcomes for studies using comparable partial pressure of CO₂ (pCO₂) ranges, suggesting that key questions remain unresolved. One unresolved hypothesis posits that heterogeneity in the response of reef calcifiers to high pCO₂ is a result of regional-scale variation in the responses to OA. To test this hypothesis, we incubated two coral taxa (Pocillopora damicornis and massive Porites) and two calcified algae (Porolithon onkodes and Halimeda macroloba) under 400, 700 and 1000 μatm pCO₂ levels in experiments in Moorea (French Polynesia), Hawaii (USA) and Okinawa (Japan), where environmental conditions differ. Both corals and H. macroloba were insensitive to OA at all three locations, while the effects of OA on P. onkodes were location-specific. In Moorea and Hawaii, calcification of P. onkodes was depressed by high pCO₂, but for specimens in Okinawa, there was no effect of OA. Using a study of large geographical scale, we show that resistance to OA of some reef species is a constitutive character expressed across the Pacific.     

Antarctic marine top predators revisited: homeotherms do not leak much CO2 to the air

Gross growth efficiencies observed for populations are applied to reasonably detailed box models of food webs of coastal Antarctic waters and the marginal ice zone. Seabirds, seals and whales apparently exhale 3% of primary net production as CO₂ into the atmosphere. For the Southern Ocean at large, the figure may be <2%.     

Interactive effects of near-future temperature increase and ocean acidification on physiology and gonad development in adult Pacific sea urchin,Echinometra sp. A

Increased atmospheric CO₂ will have a twofold impact on future marine ecosystems, increasing global sea surface temperatures and uptake of CO₂ (Ocean Acidification). Many experiments focus on the investigation of one of these stressors, but under realistic future climate predictions, these stressors may have interactive effects on individuals. Here, we investigate the effect of warming and acidification in combination. We test for interactive effects of potential near-future (2100) temperature (+2 to 3 °C) and pCO₂ (~860–940 μAtm) levels on the physiology of the tropical echinoid Echinometra sp. A. The greatest reduction in growth was under simultaneous temperature and pH/pCO₂ stress (marginally significant temperature × pH/pCO₂ interaction). This was mirrored by the physiological data, with highest metabolic activity (measured as respiration and ammonium excretion) occurring at the increased temperature and pCO₂ treatment, although this was not significant for excretion. The perivisceral coelomic fluid pH was ~7.5–7.6, as typical for echinoids, and showed no significant changes between treatments. Indicative of active calcification, internal magnesium and calcium concentrations were reduced compared to the external medium, but were not different between treatments. Gonad weight was lower at the higher temperature, and this difference was more distinct and statistically significant for males. The condition of the gonads assessed by histology declined in increased temperature and low pH treatments. The Echinometra grew in all treatments indicating active calcification of their magnesium calcite tests even as carbonate mineral saturation decreased. Our results indicate that the interactive temperature and pH effects are more important for adult echinoids than individual stressors. Although adult specimens grow and survive in near-future conditions, higher energy demands may influence gonad development and thus population maintenance.     

In situ observations of a possible skate nursery off the western Antarctic Peninsula

A dense aggregation of skate egg cases was imaged during a photographic survey of the sea floor along the western Antarctic Peninsula in November 2013. Egg cases were noted in a narrow band between 394 and 443 m depth. Although some skate species in other oceans are known to utilize restricted areas to deposit eggs in great numbers, such nurseries have not been described in the Southern Ocean.     

Aerial surveys for Antarctic minke whales (Balaenoptera bonaerensis) reveal sea ice dependent distribution patterns

This study investigates the distribution of Antarctic minke whales (AMW) in relation to sea ice concentration and variations therein. Information on AMW densities in the sea ice‐covered parts of the Southern Ocean is required to contextualize abundance estimates obtained from circumpolar shipboard surveys in open waters, suggesting a 30% decline in AMW abundance. Conventional line‐transect shipboard surveys for density estimation are impossible in ice‐covered regions, therefore we used icebreaker‐supported helicopter surveys to obtain information on AMW densities along gradients of 0%–100% of ice concentration. We conducted five helicopter surveys in the Southern Ocean, between 2006 and 2013. Distance sampling data, satellite‐derived sea‐ice data, and bathymetric parameters were used in generalized additive models (GAMs) to produce predictions on how the density of AMWs varied over space and time, and with environmental covariates. Ice concentration, distance to the ice edge and distance from the shelf break were found to describe the distribution of AMWs. Highest densities were predicted at the ice edge and through to medium ice concentrations. Medium densities were found up to 500 km into the ice edge in all concentrations of ice. Very low numbers of AMWs were found in the ice‐free waters of the West Antarctic Peninsula (WAP). A consistent relationship between AMW distribution and sea ice concentration weakens the support for the hypothesis that varying numbers of AMWs in ice‐covered waters were responsible for observed changes in estimated abundance. The potential decline in AMW abundance stresses the need for conservation measures and further studies into the AMW population status. Very low numbers of AMWs recorded in the ice‐free waters along the WAP support the hypothesis that this species is strongly dependent on sea ice and that forecasted sea ice changes have the potential of heavily impacting AMWs.     

Environmental control on the structure of echinoid assemblages in the Bellingshausen Sea (Antarctica)

The Bellingshausen Sea is one of the most remote and least surveyed seas of the Southern Ocean, so that little was known about benthic communities and those factors that determine community structuring until recently. The present work aims at characterizing the structure and spatial distribution of echinoid assemblages in the Bellingshausen Sea, as well as identifying the environmental factors that determine assemblage structuring. Echinoids were collected at 32 stations using an Agassiz trawl, at depths of 86–3,304?m, during BENTART oceanographic expeditions led in 2003 and 2006. Sediment and bottom water properties were analysed using an USNEL-type box corer and a Neil Brown Instrument System Mark III CTD, respectively. Echinoids were found at all stations, except Peter I Island. Seventeen species were identified, representing 22?% of the echinoid species present in the Southern Ocean and increasing twofold the number of species recorded in the Bellingshausen Sea so far. The echinoid fauna is dominated by the very abundant species Sterechinus antarcticus. Depth is the key factor that determines the nature of echinoid assemblages, which are mainly divided into the continental shelf, the slope and the deep-sea basin. In addition, sediment properties, namely redox values, organic matter and mud content, best match species dispersion on the shelf. Sediment properties affect echinoid distribution depending on species food range and feeding strategy. As it might be expected, sediment properties more strongly influence specialist feeders (Schizasteridae and Cidaridae) than generalists (Echinidae).     

Sea urchins, sea stars and brittle stars from Terra Nova Bay (Ross Sea, Antarctica)

Although echinoderms constitute some of the most conspicuous taxa of the Antarctic benthic communities, the echinoderm fauna of Terra Nova has not been described yet. The present study provides the first species list of echinoids, ophiuroids and asteroids from Terra Nova Bay (30–500 m depth) and describes the depth distribution of these species. Preliminary observations of the summer reproductive condition of some of the species are also included.     

WHY ARE THERE SO MANY SPECIES OF BROODING ANTARCTIC ECHINOIDS?

Marine invertebrates display a great variety of life-history traits and reproductive strategies. In echinoids, four patterns of larval development are generally recognized: planktotrophy, pelagic lecithotrophy, bottom dwelling, and brood protecting. Each broad type of free and protected development is found in all the oceans, but comparisons of the principal reproductive modes between different geographic regions have shown that they are not equally distributed. Frequency of pelagic development (planktotrophic and lecithotrophic) decreases from equator to Antarctic, where brood protecting becomes dominant. Numerous theories have been proposed to explain the richness of nonpelagic development in most marine invertebrates within the Southern Ocean. These theories can be grouped into three categories: (1) larval survival, where selection acts on larval; (2) energy allocation; and (3) dispersal. All of them consider the adaptative significance of brood protecting as the key to the success of this strategy in the Antarctic. However, the adaptative significance of brooding and the evolutionary success of this strategy in the Antarctic must be considered as two separate questions. To consider the problem at an evolutionary level, we have examined the consequences of different reproductive strategies on the genetic structure of species and on the long-term evolution of the clade. We examine this problem in the case of echinoids, a clade particularly well suited to addressing this question. In echinoids, the reduction of larval-stage duration is associated with a decrease in gene flow and consequently in the geographical scale of genetic differentiation. This allows us to reconsider the high-speciation-rate model, which leads to an increase in the number of low-dispersal species (isolation by distance). This model, previously tested by means of fossils is not satisfactory in living echinoids. Thus, the model is rebuilt with the addition of differential extinction rate between planktotrophic and brooding species in relation with the climatic history of the Antarctic.     

Sponge communities of the Antarctic Peninsula: influence of environmental variables on species composition and richness

Sponge communities on the Antarctic continental shelf currently represent one of the most extensive sponge grounds in the world, and all sponge classes are known to occur in the Southern Ocean. Main objectives of this study conducted at the tip of the Antarctic Peninsula were (1) to identify all sampled sponges and (2) to investigate whether the species composition and species richness of Southern Ocean sponge communities in the area of the Antarctic Peninsula are significantly influenced by environmental variables. The studied material originated from 25 AGT catches and was sampled during the expedition ANT-XXIX/3 of RV Polarstern. Samples were collected in three large-scale areas in the vicinity of the Antarctic Peninsula: Bransfield Strait, Drake Passage and Weddell Sea. The following six environmental variables were measured from bottom water samples (except for sea-ice cover): depth (m), light transmission (%), oxygen (µmol/kg), salinity, sea-ice cover (%) and temperature (°C). Two hundred and sixty-three sponge samples were analyzed, and 81 species of 33 genera from all Porifera classes (Calcarea, Demospongiae, Hexactinellida and Homoscleromorpha) were identified. Total numbers of sponge species per sample station ranged from 1 to 29. A detrended correspondence analysis and a backward-stepwise model selection were performed to check whether species composition and richness were significantly influenced by environmental variables. The analyses revealed that none of the measured environmental variables significantly influenced species composition but that species richness was significantly influenced by (1) temperature and (2) the combination of temperature and depth. Results of this study are of crucial importance for development, performance and assessment of future protection strategies in case of ongoing climatic changes at the Antarctic Peninsula.     

Effects of ocean acidification on Antarctic marine organisms: A meta‐analysis

Southern Ocean waters are among the most vulnerable to ocean acidification. The projected increase in the CO₂ level will cause changes in carbonate chemistry that are likely to be damaging to organisms inhabiting these waters. A meta‐analysis was undertaken to examine the vulnerability of Antarctic marine biota occupying waters south of 60°S to ocean acidification. This meta‐analysis showed that ocean acidification negatively affects autotrophic organisms, mainly phytoplankton, at CO₂ levels above 1,000 μatm and invertebrates above 1,500 μatm, but positively affects bacterial abundance. The sensitivity of phytoplankton to ocean acidification was influenced by the experimental procedure used. Natural, mixed communities were more sensitive than single species in culture and showed a decline in chlorophyll a concentration, productivity, and photosynthetic health, as well as a shift in community composition at CO₂ levels above 1,000 μatm. Invertebrates showed reduced fertilization rates and increased occurrence of larval abnormalities, as well as decreased calcification rates and increased shell dissolution with any increase in CO₂ level above 1,500 μatm. Assessment of the vulnerability of fish and macroalgae to ocean acidification was limited by the number of studies available. Overall, this analysis indicates that many marine organisms in the Southern Ocean are likely to be susceptible to ocean acidification and thereby likely to change their contribution to ecosystem services in the future. Further studies are required to address the poor spatial coverage, lack of community or ecosystem‐level studies, and the largely unknown potential for organisms to acclimate and/or adapt to the changing conditions.     

MORPHOLOGICAL AND PHYSIOLOGICAL EFFECTS IN PROBOSCIA ALATA (BACILLARIOPHYCEAE) GROWN UNDER DIFFERENT LIGHT AND CO2 CONDITIONS OF THE MODERN SOUTHERN OCEAN1

The combined effects of different light and aqueous CO₂ conditions were assessed for the Southern Ocean diatom Proboscia alata (Brightwell) Sundström in laboratory experiments. Selected culture conditions (light and CO_2(aq)) were representative for the natural ranges in the modern Southern Ocean. Light conditions were 40 (low) and 240 (high) μmol photons · m^?2 · s^?1. The three CO_2(aq) conditions ranged from 8 to 34 μmol · kg^?1 CO_2(aq) (equivalent to a pCO₂ from 137 to 598 μatm, respectively). Clear morphological changes were induced by these different CO_2(aq) conditions. Cells in low [CO_2(aq)] formed spirals, while many cells in high [CO_2(aq)] disintegrated. Cell size and volume were significantly affected by the different CO_2(aq) concentrations. Increasing CO_2(aq) concentrations led to an increase in particulate organic carbon concentrations per cell in the high light cultures, with exactly the opposite happening in the low light cultures. However, other parameters measured were not influenced by the range of CO_2(aq) treatments. This included growth rates, chlorophyll a concentration and photosynthetic yield (F_V/F_M). Different light treatments had a large effect on nutrient uptake. High light conditions caused an increased nutrient uptake rate compared to cells grown in low light conditions. Light and CO₂ conditions co‐determined in various ways the response of P. alata to changing environmental conditions. Overall P. alata appeared to be well adapted to the natural variability in light availability and CO_2(aq) concentration of the modern Southern Ocean. Nevertheless, our results showed that P. alata is susceptible to future changes in inorganic carbon concentrations in the Southern Ocean.