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Antarctic krill (Euphausia superba) and Ice krill (Euphausia crystallorophias) are key species within Southern Ocean marine ecosystems. Given their importance in regional food webs, coupled with the uncertain impacts of climate change, the on-going recovery of krill-eating marine mammals, and the expanding commercial fishery for Antarctic krill, there is an increasing need to improve current estimates of their circumpolar habitat distribution. Here, we provide an estimate of the austral summer circumpolar habitat distribution of both species using an ensemble of habitat models and updated environmental covariates. Our models were able to resolve the segregated habitats of both species. We find that extensive potential habitat for Antarctic krill is mainly situated in the open ocean and concentrated in the Atlantic sector of the Southern Ocean, while Ice krill habitat was concentrated more evenly around the continent, largely over the continental shelf. Ice krill habitat was mainly predicted by surface oxygen concentration and water column temperature, while Antarctic krill was additionally characterized by mixed layer depth, distance to the continental shelf edge, and surface salinity. Our results further improve understanding about these key species, helping inform sustainable circumpolar management practices.

The Southern Ocean is a major sink of anthropogenic CO2 and an important foraging area for top trophic level consumers. However, iron limitation sets an upper limit to primary productivity. Here we report on a considerably dense late summer phytoplankton bloom spanning 9000 km2 in the open ocean of the eastern Weddell Gyre. Over its 2.5 months duration, the bloom accumulated up to 20 g C m−2 of organic matter, which is unusually high for Southern Ocean open waters. We show that, over 1997–2019, this open ocean bloom was likely driven by anomalies in easterly winds that push sea ice southwards and favor the upwelling of Warm Deep Water enriched in hydrothermal iron and, possibly, other iron sources. This recurring open ocean bloom likely facilitates enhanced carbon export and sustains high standing stocks of Antarctic krill, supporting feeding hot spots for marine birds and baleen whales.

Numerous novel adaptations characterise the radiation of notothenioids, the dominant fish group in the freezing seas of the Southern Ocean. To improve understanding of the evolution of this iconic fish group, here we generate and analyse new genome assemblies for 24 species covering all major subgroups of the radiation, including five long-read assemblies. We present a new estimate for the onset of the radiation at 10.7 million years ago, based on a time-calibrated phylogeny derived from genome-wide sequence data. We identify a two-fold variation in genome size, driven by expansion of multiple transposable element families, and use the long-read data to reconstruct two evolutionarily important, highly repetitive gene family loci. First, we present the most complete reconstruction to date of the antifreeze glycoprotein gene family, whose emergence enabled survival in sub-zero temperatures, showing the expansion of the antifreeze gene locus from the ancestral to the derived state. Second, we trace the loss of haemoglobin genes in icefishes, the only vertebrates lacking functional haemoglobins, through complete reconstruction of the two haemoglobin gene clusters across notothenioid families. Both the haemoglobin and antifreeze genomic loci are characterised by multiple transposon expansions that may have driven the evolutionary history of these genes.

Many populations of southern hemisphere baleen whales are recovering and are again becoming dominant consumers in the Southern Ocean. Key to understanding the present and future role of baleen whales in Southern Ocean ecosystems is determining their abundance on foraging grounds. Distance sampling is the standard method for estimating baleen whale abundance but requires specific logistic requirements which are rarely achieved in the remote Southern Ocean. We explore the potential use of tourist vessel-based sampling as a cost-effective solution for conducting distance sampling surveys for baleen whales in the Southern Ocean. We used a dataset of tourist vessel locations from the southwest Atlantic sector of the Southern Ocean and published knowledge from Southern Ocean sighting surveys to determine the number of tourist vessel voyages required for robust abundance estimates. Second, we simulated the abundance and distributions of four baleen whale species for the study area and sampled them with both standardized line transect surveys and non-standardized tourist vessel-based surveys, then compared modeled abundance and distributions from each survey to the original simulation. For the southwest Atlantic, we show that 12-22 tourist vessel voyages are likely required to estimate abundance for humpback and fin whales, with relative estimates for blue, sei, Antarctic minke, and southern right whales. Second, we show tourist vessel-based surveys outperformed standardized line transect surveys at reproducing simulated baleen whale abundances and distribution. These analyses suggest tourist vessel-based surveys are a viable method for estimating baleen whale abundance in remote regions. For the southwest Atlantic, the relatively cost-effective nature of tourist vessel-based survey and regularity of tourist vessel voyages could allow for annual and intra-annual estimates of abundance, a fundamental improvement on current methods, which may capture spatiotemporal trends in baleen whale movements on forging grounds. Comparative modeling of sampling methods provided insights into the behavior of general additive model-based abundance modeling, contributing to the development of detailed guidelines of best practices for these approaches. Through successful engagement with tourist company partners, this method has the potential to characterize abundance across a variety of marine species and spaces globally, and deliver high-quality scientific outcomes relevant to management organizations.

Introduction: The Scotia Sea and Antarctic Peninsula are warming rapidly and changes in species distribution are expected. In predicting habitat shifts and considering appropriate management strategies for marine predators, a community-level understanding of how these predators are distributed is desirable. Acquiring such data, particularly in remote areas, is often problematic given the cost associated with the operation of research vessels. Here we use cruise vessels as sampling platforms to explore seabird distribution relative to habitat characteristics. Methods: Data on seabird at-sea distribution were collected using strip-transect counts throughout the Antarctic Peninsula and Scotia Sea in the austral summer of 2019-2020. Constrained correspondence analysis (CCA) and generalized additive models (GAM) were used to relate seabird community composition, density, and species richness to environmental covariates. Results: Species assemblages differed between oceanographic areas, with sea surface temperature and distance to coast being the most important predictors of seabird distribution. Our results further revealed a geographic separation of distinct communities rather than hotspot regions in the study area in summer. Discussion: These findings highlight the importance of large-scale environmental characteristics in shaping seabird community structure, presumably through underlying prey distribution and interspecific interactions. The present study contributes to the knowledge of seabird distribution and habitat use as well as the baseline for assessing the response of Antarctic seabird communities to climate warming. We argue that cruise vessels, when combined with structured research surveys, can provide a cost-effective additional tool for the monitoring of community and ecosystem level changes.

Song is produced by a variety of terrestrial and marine animals and is particularly common among baleen whales. Fin whale (Balaenoptera physalus) song is comprised of relatively simple 20 Hz pulses produced at regular intervals. The timing of these intervals, in addition to the presence and frequency of overtones, appears to be unique to each population. The purpose of this study was to characterize Western Antarctic Peninsula fin whale song and describe temporal pattern variations in song type and occurrence. Recordings were collected in the area from 2001–2004 and again 2014–2016. One song type was identified with a primary inter-pulse interval (IPI) of approximately 14 s and secondary IPI of 12.5 s. This song occurred in three pattern variants: singlet, doublet, and long triplet. The interval between pulses increased by 1.5 s between recording periods while the frequency of the overtones decreased from 89 Hz to 86 Hz. Song was never recorded in August and while it was recorded at other times in some years, it was consistently present in recordings from April through June across all years. While multiple pattern variants were present each year, singlets were generally the most prevalent variant. Doublets and triplets occurred from February through June, with highest levels of variants in February. In later years the triplet variant presence increased and in 2016 it comprised 53% of recorded song bouts. Further research is needed to understand the reasons why song changes over time and to examine the feasibility of using song to delineate and identify populations.

Global warming is affecting the population dynamics and trophic interactions across a wide range of ecosystems and habitats. Translating these real-time effects into their long-term consequences remains a challenge. The rapid and extreme warming period that occurred after the Last Glacial Maximum (LGM) during the Pleistocene–Holocene transition (7–12 thousand years ago) provides an opportunity to gain insights into the long-term responses of natural populations to periods with global warming. The effects of this post-LGM warming period have been assessed in many terrestrial taxa, whereas insights into the impacts of rapid global warming on marine taxa remain limited, especially for megafauna. In order to understand how large-scale climate fluctuations during the post-LGM affected baleen whales and their prey, we conducted an extensive, large-scale analysis of the long-term effects of the post-LGM warming on abundance and inter-ocean connectivity in eight baleen whale and seven prey (fish and invertebrates) species across the Southern and the North Atlantic Ocean; two ocean basins that differ in key oceanographic features. The analysis was based upon 7032 mitochondrial DNA sequences as well as genome-wide DNA sequence variation in 100 individuals. The estimated temporal changes in genetic diversity during the last 30,000 years indicated that most baleen whale populations underwent post-LGM expansions in both ocean basins. The increase in baleen whale abundance during the Holocene was associated with simultaneous changes in their prey and climate. Highly correlated, synchronized and exponential increases in abundance in both baleen whales and their prey in the Southern Ocean were indicative of a dramatic increase in ocean productivity. In contrast, the demographic fluctuations observed in baleen whales and their prey in the North Atlantic Ocean were subtle, varying across taxa and time. Perhaps most important was the observation that the ocean-wide expansions and decreases in abundance that were initiated by the post-LGM global warming, continued for millennia after global temperatures stabilized, reflecting persistent, long-lasting impacts of global warming on marine fauna.

Mesoscale eddies shape the foraging ecology of predators such as marine mammals and seabirds. A growing number of animal tracking studies show that predators alter their swimming, diving, and foraging behavior within mesoscale eddies. However, little is known about how Southern Ocean eddies influence the distribution of mesopelagic micronekton (fish, squid, and crustaceans), which are major prey items of megafauna. Studies in other oceanic regions have found that eddies can influence the abundance and community composition of micronekton. Here, we analyze acoustic observations from a 14-day survey of a cyclonic mesoscale eddy, its surrounding waters, and the Polar Frontal Zone (PFZ) waters where the eddy formed. We report and interpret spatial patterns of acoustic backscatter at 18 and 75 kHz, proxies indicating combined changes in species, size, and abundance of micronekton. We find that the vertical distribution of acoustic backscatter matched the underwater light conditions characteristic of the eddy core, periphery, and surrounding waters, at scales smaller than 10 km. The median water-column integrated acoustic backscatter values in the eddy core were only half of those measured in the Sub-Antarctic Zone waters surrounding the eddy, but similar to those measured in the PFZ, where the eddy originated 27 days prior. These results suggest that, as for physical and chemical tracers, the eddy maintained its biological characteristics from its source waters creating a unique habitat compared to its surroundings.

Fine-scale knowledge of spatiotemporal dynamics in cetacean distribution and abundance throughout the Western Antarctic Peninsula (WAP) is sparse yet essential for effective ecosystem-based management (EBM). Cruise vessels were used as platforms of opportunity to collect data on the distribution and abundance of humpback whales (Megaptera novaeangliae) during the austral summer of 2019/2020 in a region that is also important for the Antarctic krill (Euphausia superba) fishery, to assess potential spatiotemporal interactions for future use in EBM. Data were analyzed using traditional design-based line transect methodology and spatial density surface hurdle models fitted using a set of physical environmental covariates to estimate the abundance and distribution of whales in the area, and to describe their temporal dynamics. Our results indicate a rapid increase in humpback whale abundance in the Bransfield and Gerlache Straits through December, reaching a stable abundance by mid-January. The distribution of humpback whales appeared to change from a patchier distribution in the northern Gerlache Strait to a significantly concentrated presence in the central Gerlache and southern Bransfield Straits, followed by a subsequent dispersion throughout the area. Abundance estimates agreed well with previous literature, increasing from approximately 7000 individuals in 2000 to a peak of 19,107 in 2020. Based on these estimates, we project a total krill consumption of between 1.4 and 3.7 million tons based on traditional and contemporary literature on per capita krill consumption of whales, respectively. When taken in the context of krill fishery catch data in the study area, we conclude that there is minimal spatiotemporal overlap between humpback whales and fishery activity during our study period of November–January. However, there is potential for significant interaction between the two later in the feeding season, but cetacean survey efforts need to be extended into late season in order to fully characterize this potential overlap.

Species with similar ecological requirements that overlap in range tend to segregate their niches to minimize competition for resources. However, the niche segregation possibilities for centrally foraging predators that breed on isolated Subantarctic islands may be reduced by spatial constraints and limitations in the availability of alternative prey. In this study we examined spatial and trophic aspects of the foraging niches of two sympatrically breeding penguin species, macaroni (Eudyptes chrysolophus; MAC) and chinstrap (Pygoscelis antarcticus; CHIN) penguins, at Bouvetøya over two breeding seasons. To measure at-sea movements and diving behaviour, 90 MACs and 49 CHINs were equipped with GPS loggers and dive recorders during two austral summer breeding seasons (2014/15 and 2017/18). In addition, blood samples from tracked birds were analysed for stable isotopes to obtain dietary information. CHINs displayed marked interannual variation in foraging behaviour, diving deeper, utilizing a larger foraging area and displaying enriched values of δ15N in 2014/15 compared to the 2017/18 breeding season. In contrast, MACs dove to similar depths and showed similar δ15N values, while consistently utilizing larger foraging areas compared to CHINs. We suggest that low krill abundances in the waters around Bouvetøya during the 2014/15 season resulted in CHINs shifting toward a diet that increased their niche overlap with MACs. Our findings may partly explain the decreasing number of breeding CHINs at the world’s most remote island, Bouvetøya, while also highlighting the importance of characterizing niche overlap of species using multi-season data sets.

Polynyas represent regions of enhanced primary production because of the low, or absent, sea-ice cover coupled with the proximity of nutrient sources. However, studies throughout the Southern Ocean suggest elevated primary production does not necessarily result in increased carbon export. Three coastal polynyas in East Antarctica and an off-shelf region were visited during the austral summer from December 2016 to January 2017 to examine the vertical distribution and concentration of particulate organic carbon (POC). Carbon export was also examined using thorium-234 (234Th) as a proxy at two of the polynyas. Our results show that concentrations and integrated POC stocks were higher within the polynyas compared to the off-shelf sites. Within the polynyas, vertical POC concentrations were higher in the Mertz and Ninnis polynyas compared to the Dalton polynya. Similarly, higher carbon export was measured in the diatom-dominated Mertz polynya, where large particles (>53 μm) represented a significant fraction of the particulate 234Th and POC (average 50% and 39%, respectively), compared to the small flagellate-dominated Dalton polynya, where almost all the particulate 234Th and POC were found in the smaller size fraction (1–53 μm). The POC to Chlorophyll-a ratios suggest that organic matter below the mixed layer in the polynyas consisted largely of fresh phytoplankton at this time of the year. In combination with a parallel study on phytoplankton production at these sites, we find that increased primary production at these polynyas does lead to greater concentrations and export of POC and a higher POC export efficiency.

Allometric relationships between body properties of animals are useful for a wide variety of purposes, such as estimation of biomass, growth, population structure, bioenergetic modelling and carbon flux studies. This study summarizes allometric relationships of zooplankton and nekton species that play major roles in polar marine food webs. Measurements were performed on 639 individuals of 15 species sampled during three expeditions in the Southern Ocean (winter and summer) and 2374 individuals of 14 species sampled during three expeditions in the Arctic Ocean (spring and summer). The information provided by this study fills current knowledge gaps on relationships between length and wet/dry mass of understudied animals, such as various gelatinous zooplankton, and of animals from understudied seasons and maturity stages, for example, for the krill Thysanoessa macrura and larval Euphausia superba caught in winter. Comparisons show that there is intra-specific variation in length–mass relationships of several species depending on season, e.g. for the amphipod Themisto libellula. To investigate the potential use of generalized regression models, comparisons between sexes, maturity stages or age classes were performed and are discussed, such as for the several krill species and T. libellula. Regression model comparisons on age classes of the fish E. antarctica were inconclusive about their general use. Other allometric measurements performed on carapaces, eyes, heads, telsons, tails and otoliths provided models that proved to be useful for estimating length or mass in, e.g. diet studies. In some cases, the suitability of these models may depend on species or developmental stages.

Survival of larval Antarctic krill (Euphausia superba) during winter is largely dependent upon the presence of sea ice as it provides an important source of food and shelter. We hypothesized that sea ice provides additional benefits because it hosts fewer competitors and provides reduced predation risk for krill larvae than the water column. To test our hypothesis, zooplankton were sampled in the Weddell-Scotia Confluence Zone at the ice-water interface (0–2 m) and in the water column (0–500 m) during August–October 2013. Grazing by mesozooplankton, expressed as a percentage of the phytoplankton standing stock, was higher in the water column (1.97 ± 1.84%) than at the ice-water interface (0.08 ± 0.09%), due to a high abundance of pelagic copepods. Predation risk by carnivorous macrozooplankton, expressed as a percentage of the mesozooplankton standing stock, was significantly lower at the ice-water interface (0.83 ± 0.57%; main predators amphipods, siphonophores and ctenophores) than in the water column (4.72 ± 5.85%; main predators chaetognaths and medusae). These results emphasize the important role of sea ice as a suitable winter habitat for larval krill with fewer competitors and lower predation risk. These benefits should be taken into account when considering the response of Antarctic krill to projected declines in sea ice. Whether reduced sea-ice algal production may be compensated for by increased water column production remains unclear, but the shelter provided by sea ice would be significantly reduced or disappear, thus increasing the predation risk on krill larvae.

Global targets for area-based conservation and management must move beyond threshold-based targets alone and must account for the quality of such areas. In the Southern Ocean around Antarctica, a region where key biodiversity faces unprecedented risks from climate change and where there is a growing demand to extract resources, a number of marine areas have been afforded enhanced conservation or management measures through two adopted marine protected areas (MPAs). However, evidence suggests that additional high quality areas could benefit from a proposed network of MPAs. Penguins offer a particular opportunity to identify high quality areas because these birds, as highly visible central-place foragers, are considered indicator species whose populations reflect the state of the surrounding marine environment. We compiled a comprehensive dataset of the location of penguin colonies and their associated abundance estimates in Antarctica. We then estimated the at-sea distribution of birds based on information derived from tracking data and through the application of a modified foraging radius approach with a density decay function to identify some of the most important marine areas for chick-rearing adult penguins throughout waters surrounding Antarctica following the Important Bird and Biodiversity Area (IBA) framework. Additionally, we assessed how marine IBAs overlapped with the currently adopted and proposed network of key management areas (primarily MPAs), and how the krill fishery likely overlapped with marine IBAs over the past five decades. We identified 63 marine IBAs throughout Antarctic waters and found that were the proposed MPAs to be adopted, the permanent conservation of high quality areas for penguin species would increase by between 49 and 100% depending on the species. Furthermore, our data show that, despite a generally contracting range of operation by the krill fishery in Antarctica over the past five decades, a consistently disproportionate amount of krill is being harvested within marine IBAs compared to the total area in which the fishery operates. Our results support the designation of the proposed MPA network and offer additional guidance as to where decision-makers should act before further perturbation occurs in the Antarctic marine ecosystem.

Knowing the magnitude and timing of pelagic primary production is important for ecosystem and carbon sequestration studies, in addition to providing basic understanding of phytoplankton functioning. In this study we use data from an ecosystem cruise to Kong Håkon VII Hav, in the Atlantic sector of the Southern Ocean, in March 2019 and more than two decades of satellite-derived ocean color to study phytoplankton bloom phenology. During the cruise we observed phytoplankton blooms in different bloom phases. By correlating bloom phenology indices (i.e., bloom initiation and end) based on satellite remote sensing to the timing of changes in environmental conditions (i.e., sea ice, light, and mixed layer depth) we studied the environmental factors that seemingly drive phytoplankton blooms in the area. Our results show that blooms mainly take place in January and February, consistent with previous studies that include the area. Sea ice retreat controls the bloom initiation in particular along the coast and the western part of the study area, whereas bloom end is not primarily connected to sea ice advance. Light availability in general is not appearing to control the bloom termination, neither is nutrient availability based on the autumn cruise where we observed non-depleted macronutrient reservoirs in the surface. Instead, we surmise that zooplankton grazing plays a potentially large role to end the bloom, and thus controls its duration. The spatial correlation of the highest bloom magnitude with marked topographic features indicate that the interaction of ocean currents with sea floor topography enhances primary productivity in this area, probably by natural fertilization. Based on the bloom timing and magnitude patterns, we identified five different bloom regimes in the area. A more detailed understanding of the region will help to highlight areas with the highest relevance for the carbon cycle, the marine ecosystem and spatial management. With this gained understanding of bloom phenology, it will also be possible to study potential shifts in bloom timing and associated trophic mismatch caused by environmental changes.

Estimates of the distribution and density of Antarctic krill (Euphausia superba Dana, 1850) were derived from a large-scale survey conducted during the austral summer in the Southwest Atlantic sector of the Southern Ocean and across the Scotia Sea in 2018–19, the ‘2018–19 Area 48 Survey’. Survey vessels were provided by Norway, the Association of Responsible Krill harvesting companies and Aker BioMarine AS, the United Kingdom, Ukraine, Republic of Korea, and China. Survey design followed the transects of the Commission for the Conservation of Antarctic Marine Living Resources synoptic survey, carried out in 2000 and from regular national surveys performed in the South Atlantic sector by the U.S., China, Republic of Korea, Norway, and the U.K. The 2018–19 Area 48 Survey represents only the second large-scale survey performed in the area and this joint effort resulted in the largest ever total transect line (19,500 km) coverage carried out as one single exercise in the Southern Ocean. We delineated and integrated acoustic backscatter arising from krill swarms to produce distribution maps of krill areal biomass density and standing stock (biomass) estimates. Krill standing stock for the Area 48 was estimated to be 62.6 megatonnes (mean density of 30 g m–2 over 2 million km2) with a sampling coefficient variation of 13%. The highest mean krill densities were found in the South Orkney Islands stratum (93.2 g m–2) and the lowest in the South Georgia Island stratum (6.4 g m–2). The krill densities across the strata compared to those found during the previous survey indicate some regional differences in distribution and biomass. It is currently not possible to assign any such differences or lack of differences between the two survey datasets to longer term trends in the environment, krill stocks or fishing pressure.