Decades of dietary data demonstrate regional food web structures in the Southern Ocean

Understanding regional‐scale food web structure in the Southern Ocean is critical to informing fisheries management and assessments of climate change impacts on Southern Ocean ecosystems and ecosystem services. Historically, a large component of Southern Ocean ecosystem research has focused on Antarctic krill, which provide a short, highly efficient food chain, linking primary producers to higher trophic levels. Over the last 15 years, the presence of alternative energy pathways has been identified and hypotheses on their relative importance in different regions raised. Using the largest circumpolar dietary database ever compiled, we tested these hypotheses using an empirical circumpolar comparison of food webs across the four major regions/sectors of the Southern Ocean (defined as south of 40°S) within the austral summer period. We used network analyses and generalizations of taxonomic food web structure to confirm that while Antarctic krill are dominant as the mid‐trophic level for the Atlantic and East Pacific food webs (including the Scotia Arc and Western Antarctic Peninsula), mesopelagic fish and other krill species are dominant contributors to predator diets in the Indian and West Pacific regions (East Antarctica and the Ross Sea). We also highlight how tracking data and habitat modeling for mobile top predators in the Southern Ocean show that these species integrate food webs over large regional scales. Our study provides a quantitative assessment, based on field observations, of the degree of regional differentiation in Southern Ocean food webs and the relative importance of alternative energy pathways between regions.     

Sea Surface Temperature Influence on Terrestrial Gross Primary Production along the Southern California Current

Some land and ocean processes are related through connections (and synoptic-scale teleconnections) to the atmosphere. Synoptic-scale atmospheric (El Niño/Southern Oscillation [ENSO], Pacific Decadal Oscillation [PDO], and North Atlantic Oscillation [NAO]) decadal cycles are known to influence the global terrestrial carbon cycle. Potentially, smaller scale land-ocean connections influenced by coastal upwelling (changes in sea surface temperature) may be important for local-to-regional water-limited ecosystems where plants may benefit from air moisture transported from the ocean to terrestrial ecosystems. Here we use satellite-derived observations to test potential connections between changes in sea surface temperature (SST) in regions with strong coastal upwelling and terrestrial gross primary production (GPP) across the Baja California Peninsula. This region is characterized by an arid/semiarid climate along the southern California Current. We found that SST was correlated with the fraction of photosynthetic active radiation (fPAR; as a proxy for GPP) with lags ranging from 0 to 5 months. In contrast ENSO was not as strongly related with fPAR as SST in these coastal ecosystems. Our results show the importance of local-scale changes in SST during upwelling events, to explain the variability in GPP in coastal, water-limited ecosystems. The response of GPP to SST was spatially-dependent: colder SST in the northern areas increased GPP (likely by influencing fog formation), while warmer SST at the southern areas was associated to higher GPP (as SST is in phase with precipitation patterns). Interannual trends in fPAR are also spatially variable along the Baja California Peninsula with increasing secular trends in subtropical regions, decreasing trends in the most arid region, and no trend in the semi-arid regions. These findings suggest that studies and ecosystem process based models should consider the lateral influence of local-scale ocean processes that could influence coastal ecosystem productivity.     

Environmental forcing and Southern Ocean marine predator populations: effects of climate change and variability

The Southern Ocean is a major component within the global ocean and climate system and potentially the location where the most rapid climate change is most likely to happen, particularly in the high-latitude polar regions. In these regions, even small temperature changes can potentially lead to major environmental perturbations. Climate change is likely to be regional and may be expressed in various ways, including alterations to climate and weather patterns across a variety of time-scales that include changes to the long interdecadal background signals such as the development of the El Niño–Southern Oscillation (ENSO). Oscillating climate signals such as ENSO potentially provide a unique opportunity to explore how biological communities respond to change. This approach is based on the premise that biological responses to shorter-term sub-decadal climate variability signals are potentially the best predictor of biological responses over longer time-scales. Around the Southern Ocean, marine predator populations show periodicity in breeding performance and productivity, with relationships with the environment driven by physical forcing from the ENSO region in the Pacific. Wherever examined, these relationships are congruent with mid-trophic-level processes that are also correlated with environmental variability. The short-term changes to ecosystem structure and function observed during ENSO events herald potential long-term changes that may ensue following regional climate change. For example, in the South Atlantic, failure of Antarctic krill recruitment will inevitably foreshadow recruitment failures in a range of higher trophic-level marine predators. Where predator species are not able to accommodate by switching to other prey species, population-level changes will follow. The Southern Ocean, though oceanographically interconnected, is not a single ecosystem and different areas are dominated by different food webs. Where species occupy different positions in different regional food webs, there is the potential to make predictions about future change scenarios.     

Length–weight relationships of five fish species associated with krill fishery in the Atlantic sector of the Southern Ocean

Length–weight relationships (LWRs) of five fish species commonly associated with Antarctic krill fishery, were determined. Samples were collected with Chinese krill trawler (codend mesh size 2 cm) in the Atlantic sector of the Southern Ocean from January to August 2016. Data on LWRs and the relationships between weight and standard length as well as weight and total length of those species were updated for the database of FishBase. Those data on fish species derived from Antarctic krill fishery will be very helpful in understanding the interaction between krill fishery and the associated fish community.     

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.     

Genetic variation and population structure of krill (Euphausia superba dana) from the Prydz Bay region of Antarctic waters

Summary Samples of Antarctic krill E. superba from six locations near Prydz Bay were analysed electrophoretically to detect genetically-based protein variation. Analyses of allele distributions at four polymorphic loci indicate no evidence of significant heterogeneity, a result consistent with the hypothesis that all samples were derived from a single breeding population of krill. The results of this study agree closely with genetic data from other studies on krill in the Atlantic sector of the Southern Ocean, suggesting that krill stocks over at least 6000 km of Antarctic waters are derived from a single interbreeding population.     

Genetic homogeneity of krill (Euphausia superba Dana) in the Southern Ocean

Summary Development of a comprehensive picture of the genetic population structure of the Antarctic krill (Euphausia superba) has been hampered by a lack of genetic data from two major areas of the species' distribution, the Bellingshausen Sea and the Ross Sea. Evidence from earlier studies of a discrete Bellingshausen Sea population was based on anomalous allele frequencies in two sample sets that were collected near the west coast of the Antarctic Peninsula rather than in the Bellingshausen Sea proper. In this paper we describe the first biochemical genetic data obtained on krill from the central Bellingshausen Sea and from the Ross Sea. Analyses of eight polymorphic loci in samples from these two areas have failed to provide any evidence of population structuring within the Pacific sector of the Southern Ocean, and have indicated that Pacific sector krill cannot be genetically discriminated from Atlantic sector krill or Indian Ocean sector krill. These findings further support the hypothesis of a single circumpolar breeding population of Antarctic krill.     

Isotopic and genetic evidence for culturally inherited site fidelity to feeding grounds in southern right whales (Eubalaena australis)

Ocean warming will undoubtedly affect the migratory patterns of many marine species, but specific changes can be predicted only where behavioural mechanisms guiding migration are understood. Southern right whales show maternally inherited site fidelity to near-shore winter nursery grounds, but exactly where they feed in summer (collectively and individually) remains mysterious. They consume huge quantities of copepods and krill, and their reproductive rates respond to fluctuations in krill abundance linked to El Niño Southern Oscillation (ENSO). Here we show that genetic and isotopic signatures, analysed together, indicate maternally directed site fidelity to diverse summer feeding grounds for female right whales calving at Península Valdés, Argentina. Isotopic values from 131 skin samples span a broad range (–23.1 to –17.2‰δ¹³C, 6.0 to 13.8‰δ¹⁵N) and are more similar than expected among individuals sharing the same mitochondrial haplotype. This pattern indicates that calves learn summer feeding locations from their mothers, and that the timescale of culturally inherited site fidelity to feeding grounds is at least several generations. Such conservatism would be expected to limit the exploration of new feeding opportunities, and may explain why this population shows increased rates of reproductive failure in years following elevated sea-surface temperature anomalies off South Georgia, the richest known feeding ground for baleen whales in the South Atlantic.     

Evidence of resource partitioning between humpback and minke whales around the western Antarctic Peninsula

For closely related sympatric species to coexist, they must differ to some degree in their ecological requirements or niches ( e.g. , diets) to avoid interspecific competition. Baleen whales in the Antarctic feed primarily on krill, and the large sympatric prewhaling community suggests resource partitioning among these species or a nonlimiting prey resource. In order to examine ecological differences between sympatric humpback and minke whales around the Western Antarctic Peninsula, we made measurements of the physical environment, observations of whale distribution, and concurrent acoustic measurements of krill aggregations. Mantel's tests and classification and regression tree models indicate both similarities and differences in the spatial associations between humpback and minke whales, environmental features, and prey. The data suggest (1) similarities (proximity to shore) and differences (prey abundance versus deep water temperatures) in horizontal spatial distribution patterns, (2) unambiguous vertical resource partitioning with minke whales associating with deeper krill aggregations across a range of spatial scales, and (3) that interference competition between these two species is unlikely. These results add to the paucity of ecological knowledge relating baleen whales and their prey in the Antarctic and should be considered in conservation and management efforts for Southern Ocean cetaceans and ecosystems.     

Not just about sunburn – the ozone hole's profound effect on climate has significant implications for Southern Hemisphere ecosystems

Climate scientists have concluded that stratospheric ozone depletion has been a major driver of Southern Hemisphere climate processes since about 1980. The implications of these observed and modelled changes in climate are likely to be far more pervasive for both terrestrial and marine ecosystems than the increase in ultraviolet‐B radiation due to ozone depletion; however, they have been largely overlooked in the biological literature. Here, we synthesize the current understanding of how ozone depletion has impacted Southern Hemisphere climate and highlight the relatively few documented impacts on terrestrial and marine ecosystems. Reviewing the climate literature, we present examples of how ozone depletion changes atmospheric and oceanic circulation, with an emphasis on how these alterations in the physical climate system affect Southern Hemisphere weather, especially over the summer season (December–February). These potentially include increased incidence of extreme events, resulting in costly floods, drought, wildfires and serious environmental damage. The ecosystem impacts documented so far include changes to growth rates of South American and New Zealand trees, decreased growth of Antarctic mosses and changing biodiversity in Antarctic lakes. The objective of this synthesis was to stimulate the ecological community to look beyond ultraviolet‐B radiation when considering the impacts of ozone depletion. Such widespread changes in Southern Hemisphere climate are likely to have had as much or more impact on natural ecosystems and food production over the past few decades, than the increased ultraviolet radiation due to ozone depletion.     

Important marine habitat off east Antarctica revealed by two decades of multi‐species predator tracking

Satellite telemetry data are a key source of animal distribution information for marine ecosystem management and conservation activities. We used two decades of telemetry data from the East Antarctic sector of the Southern Ocean. Habitat utilization models for the spring/summer period were developed for six highly abundant, wide‐ranging meso‐ and top‐predator species: Adélie Pygoscelis adeliae and emperor Aptenodytes forsteri penguins, light‐mantled albatross Phoebetria palpebrata, Antarctic fur seals Arctocephalus gazella, southern elephant seals Mirounga leonina, and Weddell seals Leptonychotes weddellii. The regional predictions from these models were combined to identify areas utilized by multiple species, and therefore likely to be of particular ecological significance. These areas were distributed across the longitudinal breadth of the East Antarctic sector, and were characterized by proximity to breeding colonies, both on the Antarctic continent and on subantarctic islands to the north, and by sea‐ice dynamics, particularly locations of winter polynyas. These areas of important habitat were also congruent with many of the areas reported to be showing the strongest regional trends in sea ice seasonality. The results emphasize the importance of on‐shore and sea‐ice processes to Antarctic marine ecosystems. Our study provides ocean‐basin‐scale predictions of predator habitat utilization, an assessment of contemporary habitat use against which future changes can be assessed, and is of direct relevance to current conservation planning and spatial management efforts.     

Predicting krill swarm characteristics important for marine predators foraging off East Antarctica

Open ocean predator‐prey interactions are often difficult to interpret because of a lack of information on prey fields at scales relevant to predator behaviour. Hence, there is strong interest in identifying the biological and physical factors influencing the distribution and abundance of prey species, which may be of broad predictive use for conservation planning and evaluating effects of environmental change. This study focuses on a key Southern Ocean prey species, Antarctic krill Euphausia superba, using acoustic observations of individual swarms (aggregations) from a large‐scale survey off East Antarctica. We developed two sets of statistical models describing swarm characteristics, one set using underway survey data for the explanatory variables, and the other using their satellite remotely sensed analogues. While survey data are in situ and contemporaneous with the swarm data, remotely sensed data are all that is available for prediction and inference about prey distribution in other areas or at other times. The fitted models showed that the primary biophysical influences on krill swarm characteristics included daylight (solar elevation/radiation) and proximity to the Antarctic continental slope, but there were also complex relationships with current velocities and gradients. Overall model performance was similar regardless of whether underway or remotely sensed predictors were used. We applied the latter models to generate regional‐scale spatial predictions using a 10‐yr remotely‐sensed time series. This retrospective modelling identified areas off east Antarctica where relatively dense krill swarms were consistently predicted during austral mid‐summers, which may underpin key foraging areas for marine predators. Spatiotemporal predictions along Antarctic predator satellite tracks, from independent studies, illustrate the potential for uptake into further quantitative modelling of predator movements and foraging. The approach is widely applicable to other krill‐dependent ecosystems, and our findings are relevant to similar efforts examining biophysical linkages elsewhere in the Southern Ocean and beyond.     

Relative changes in krill abundance inferred from Antarctic fur seal

Antarctic krill Euphausia superba is a predominant species in the Southern Ocean, it is very sensitive to climate change, and it supports large stocks of fishes, seabirds, seals and whales in Antarctic marine ecosystems. Modern krill stocks have been estimated directly by net hauls and acoustic surveys; the historical krill density especially the long-term one in the Southern Ocean, however, is unknown. Here we inferred the relative krill population changes along the West Antarctic Peninsula (WAP) over the 20th century from the trophic level change of Antarctic fur seal Arctocephalus gazella using stable carbon (δ(13)C) and nitrogen (δ(15)N) isotopes of archival seal hairs. Since Antarctic fur seals feed preferentially on krill, the variation of δ(15)N in seal hair indicates a change in the proportion of krill in the seal's diets and thus the krill availability in local seawater. For the past century, enriching fur seal δ(15)N values indicated decreasing krill availability. This is agreement with direct observation for the past ～30 years and suggests that the recently documented decline in krill populations began in the early parts of the 20th century. This novel method makes it possible to infer past krill population changes from ancient tissues of krill predators.     

Deep-sea pelagic ichthyonekton diversity in the Atlantic Ocean and the adjacent sector of the Southern Ocean

Aim Deep‐sea pelagic diversity is poorly understood. Local (SL) and regional (SR) ichthyonekton species richness are presented and analysed with respect to local and regional environmental factors, and biogeographical processes. Location Sixty‐six stations from the Atlantic Ocean and adjacent sector of the Southern Ocean, 65° N to 57° S. Methods Estimation of SL by means of rarefaction. Stepwise evaluation of SL and SR relationships by means of the second‐order corrected Akaike information criterion (AICc) after locally weighted scatterplot smoothing (LOESS) and linear fitting, analysis of saturation effects by means of slopes of species accumulation curves (log–log plots). Results Latitudinal gradients were present for SL and SR, and were asymmetric between the Northern and Southern hemispheres. Relatively low species richness was encountered for the Southern Ocean. Asymmetry at the regional level by means of higher SR was attributed to area effects in the South Atlantic. Log–log plots indicated saturation of local assemblages and dependence on environmental factors. SL was related to productivity; this relationship was hump‐shaped. SR was positively related to area size and negatively to seasonality of production. Biogeographical effects were indicated in that SR peaks coincided with overlap zones of boreal and tropical faunas as a consequence of historical faunal exchange processes. Main conclusions The stepwise approach allowed for distinction between effects of area size, productivity and biogeographical processes on diversity at local and regional scales. Productivity in particular is important in two ways. At the local scale, the link of productivity to SL is explained by a successional‐functional hypothesis of resource utilization, whereas the seasonality effect for SR reinforces the hypothesis of dependence of deep‐sea fishes on seasonality of production through changes of life‐history traits. The causes of low Antarctic faunal diversity remained unresolved.     

Global climate drives southern right whale (Eubalaena australis) population dynamics

Sea surface temperature (SST) time-series from the southwest Atlantic and the El Ni?o 4 region in the western Pacific were compared to an index of annual calving success of the southern right whale (Eubalaena australis) breeding in Argentina. There was a strong relationship between right whale calving output and SST anomalies at South Georgia in the autumn of the previous year and also with mean El Ni?o 4 SST anomalies delayed by 6 years. These results extend similar observations from other krill predators and show clear linkages between global climate signals and the biological processes affecting whale population dynamics.     

Spatial Distribution, Structure, Biomass, and Physiology of Microbial Assemblages across the Southern Ocean Frontal Zones during the Late Austral Winter

We examined the spatial distributions of picoplankton, nanoplankton, and microplankton biomass and physiological state relative to the hydrography of the Southern Ocean along 90 degrees W longitude and across the Drake Passage in the late austral winter. The eastern South Pacific Ocean showed some large-scale biogeographical differences and size class variability. Microbial ATP biomass was greatest in euphotic surface waters. The horizontal distributions of microbial biomass and physiological state (adenylate energy charge ratio) coincided with internal currents (fronts) of the Antarctic Circumpolar Current. In the Drake Passage, the biological scales in the euphotic and aphotic zones were complex, and ATP, total adenylate, and adenylate energy charge ratio isopleths were compressed due to the extension of the sea ice from Antarctica and constriction of the Circumpolar Current through the narrow passage. The physiological state of microbial assemblages and biomass were much higher in the Drake Passage than in the eastern South Pacific Ocean. The temperature of Antarctic waters, not dissolved organic carbon, was the major variable controlling picoplankton growth. Estimates of picoplankton production based on ATP increments with time suggest that production under reduced predation pressure was 1 to 10 mug of carbon per liter per day. Our results demonstrate the influence of large-scale hydrographic processes on the distribution and structure of microplankton, nanoplankton, and picoplankton across the Southern Ocean.     

Signals from the south; humpback whales carry messages of Antarctic sea‐ice ecosystem variability

Southern hemisphere humpback whales (Megaptera novaeangliae) rely on summer prey abundance of Antarctic krill (Euphausia superba) to fuel one of the longest‐known mammalian migrations on the planet. It is hypothesized that this species, already adapted to endure metabolic extremes, will be one of the first Antarctic consumers to show measurable physiological change in response to fluctuating prey availability in a changing climate; and as such, a powerful sentinel candidate for the Antarctic sea‐ice ecosystem. Here, we targeted the sentinel parameters of humpback whale adiposity and diet, using novel, as well as established, chemical and biochemical markers, and assembled a time trend spanning 8 years. We show the synchronous, inter‐annual oscillation of two measures of humpback whale adiposity with Southern Ocean environmental variables and climate indices. Furthermore, bulk stable isotope signatures provide clear indication of dietary compensation strategies, or a lower trophic level isotopic change, following years indicated as leaner years for the whales. The observed synchronicity of humpback whale adiposity and dietary markers, with climate patterns in the Southern Ocean, lends strength to the role of humpback whales as powerful Antarctic sea‐ice ecosystem sentinels. The work carries significant potential to reform current ecosystem surveillance in the Antarctic region.     

The association of Antarctic krill Euphausia superba with the under-ice habitat

The association of Antarctic krill Euphausia superba with the under-ice habitat was investigated in the Lazarev Sea (Southern Ocean) during austral summer, autumn and winter. Data were obtained using novel Surface and Under Ice Trawls (SUIT), which sampled the 0-2 m surface layer both under sea ice and in open water. Average surface layer densities ranged between 0.8 individuals m(-2) in summer and autumn, and 2.7 individuals m(-2) in winter. In summer, under-ice densities of Antarctic krill were significantly higher than in open waters. In autumn, the opposite pattern was observed. Under winter sea ice, densities were often low, but repeatedly far exceeded summer and autumn maxima. Statistical models showed that during summer high densities of Antarctic krill in the 0-2 m layer were associated with high ice coverage and shallow mixed layer depths, among other factors. In autumn and winter, density was related to hydrographical parameters. Average under-ice densities from the 0-2 m layer were higher than corresponding values from the 0-200 m layer collected with Rectangular Midwater Trawls (RMT) in summer. In winter, under-ice densities far surpassed maximum 0-200 m densities on several occasions. This indicates that the importance of the ice-water interface layer may be under-estimated by the pelagic nets and sonars commonly used to estimate the population size of Antarctic krill for management purposes, due to their limited ability to sample this habitat. Our results provide evidence for an almost year-round association of Antarctic krill with the under-ice habitat, hundreds of kilometres into the ice-covered area of the Lazarev Sea. Local concentrations of postlarval Antarctic krill under winter sea ice suggest that sea ice biota are important for their winter survival. These findings emphasise the susceptibility of an ecological key species to changing sea ice habitats, suggesting potential ramifications on Antarctic ecosystems induced by climate change.     

Climate influences on female survival in a declining population of southern elephant seals (Mirounga leonina)

The Southern Ocean has been disproportionately affected by climate change and is therefore an ideal place to study the influence of changing environmental conditions on ecosystems. Changes in the demography of predator populations are indicators of broader shifts in food web structure, but long‐term data are required to study these effects. Southern elephant seals (Mirounga leonina) from Macquarie Island have consistently decreased in population size while all other major populations across the Southern Ocean have recently stabilized or are increasing. Two long‐term mark‐recapture studies (1956–1967 and 1993–2009) have monitored this population, which provides an opportunity to investigate demographic performance over a range of climatic conditions. Using a 9‐state matrix population model, we estimated climate influences on female survival by incorporating two major climatic indices into our model: The Southern Annular Mode (SAM) and the Southern Oscillation Index (SOI). Our best model included a 1 year lagged effect of SAM and an unlagged SOI as covariates. A positive relationship with SAM1 (lagged) related the previous year''s SAM with juvenile survival, potentially due to changes in local prey availability surrounding Macquarie Island. The unlagged SOI had a negative effect on both juvenile and adult seals, indicating that sea ice dynamics and access to foraging grounds on the East Antarctic continental shelf could explain the different contributions of ENSO events on the survival of females in this population.