Estimating the average distribution of Antarctic krill Euphausia superba at the northern Antarctic Peninsula during austral summer and winter.

This study was performed to aid the management of the fishery for Antarctic krill Euphausia superba. Krill are an important component of the Antarctic marine ecosystem, providing a key food source for many marine predators. Additionally, krill are the target of the largest commercial fishery in the Southern Ocean, for which annual catches have been increasing and concentrating in recent years. The krill fishery is managed by the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR), which has endorsed a new management framework that requires information about the spatial distribution and biomass of krill. Here, we use krill density estimates from acoustic surveys and a GAMM framework to model habitat properties associated with high krill biomass during summer and winter in the northern Antarctic Peninsula region, an area important to the commercial fishery. Our models show elevated krill density associated with the shelf break, increased sea surface temperature, moderate chlorophyll-a concentration and increased salinity. During winter, our models show associations with shallow waters (< 1500 m) with low sea-ice concentration, medium sea-level anomaly and medium current speed. Our models predict temporal averages of the distribution and density of krill, which can be used to aid CCAMLR's revised ecosystem approach to fisheries management. Our models have the potential to help in the spatial and temporal design of future acoustic surveys that would preclude the need for modelled extrapolations. We highlight that the ecosystem approach to fisheries management of krill critically depends upon such field observations at relevant spatial and temporal scales. Supplementary Information: The online version contains supplementary material available at 10.1007/s00300-022-03039-y.

Positive selection over the mitochondrial genome and its role in the diversification of gentoo penguins in response to adaptation in isolation.

Although mitochondrial DNA has been widely used in phylogeography, evidence has emerged that factors such as climate, food availability, and environmental pressures that produce high levels of stress can exert a strong influence on mitochondrial genomes, to the point of promoting the persistence of certain genotypes in order to compensate for the metabolic requirements of the local environment. As recently discovered, the gentoo penguins (Pygoscelis papua) comprise four highly divergent lineages across their distribution spanning the Antarctic and sub-Antarctic regions. Gentoo penguins therefore represent a suitable animal model to study adaptive processes across divergent environments. Based on 62 mitogenomes that we obtained from nine locations spanning all four gentoo penguin lineages, we demonstrated lineage-specific nucleotide substitutions for various genes, but only lineage-specific amino acid replacements for the ND1 and ND5 protein-coding genes. Purifying selection (dN/dS < 1) is the main driving force in the protein-coding genes that shape the diversity of mitogenomes in gentoo penguins. Positive selection (dN/dS > 1) was mostly present in codons of the Complex I (NADH genes), supported by two different codon-based methods at the ND1 and ND4 in the most divergent lineages, the eastern gentoo penguin from Crozet and Marion Islands and the southern gentoo penguin from Antarctica respectively. Additionally, ND5 and ATP6 were under selection in the branches of the phylogeny involving all gentoo penguins except the eastern lineage. Our study suggests that local adaptation of gentoo penguins has emerged as a response to environmental variability promoting the fixation of mitochondrial haplotypes in a non-random manner. Mitogenome adaptation is thus likely to have been associated with gentoo penguin diversification across the Southern Ocean and to have promoted their survival in extreme environments such as Antarctica. Such selective processes on the mitochondrial genome may also be responsible for the discordance detected between nuclear- and mitochondrial-based phylogenies of gentoo penguin lineages.

Hemispheric asymmetry in ocean change and the productivity of ecosystem sentinels.

Climate change and other human activities are causing profound effects on marine ecosystem productivity. We show that the breeding success of seabirds is tracking hemispheric differences in ocean warming and human impacts, with the strongest effects on fish-eating, surface-foraging species in the north. Hemispheric asymmetry suggests the need for ocean management at hemispheric scales. For the north, tactical, climate-based recovery plans for forage fish resources are needed to recover seabird breeding productivity. In the south, lower-magnitude change in seabird productivity presents opportunities for strategic management approaches such as large marine protected areas to sustain food webs and maintain predator productivity. Global monitoring of seabird productivity enables the detection of ecosystem change in remote regions and contributes to our understanding of marine climate impacts on ecosystems.

A comparison of baleen whale density estimates derived from overlapping satellite imagery and a shipborne survey.

As whales recover from commercial exploitation, they are increasing in abundance in habitats that they have been absent from for decades. However, studying the recovery and habitat use patterns of whales, particularly in remote and inaccessible regions, frequently poses logistical and economic challenges. Here we trial a new approach for measuring whale density in a remote area, using Very-High-Resolution WorldView-3 satellite imagery. This approach has capacity to provide sightings data to complement and assist traditional sightings surveys. We compare at-sea whale density estimates to estimates derived from satellite imagery collected at a similar time, and use suction-cup archival logger data to make an adjustment for surface availability. We demonstrate that satellite imagery can provide useful data on whale occurrence and density. Densities, when unadjusted for surface availability are shown to be considerably lower than those estimated by the ship survey. However, adjusted for surface availability and weather conditions (0.13 whales per km2, CV = 0.38), they fall within an order of magnitude of those derived by traditional line-transect estimates (0.33 whales per km2, CV = 0.09). Satellite surveys represent an exciting development for high-resolution image-based cetacean observation at sea, particularly in inaccessible regions, presenting opportunities for ongoing and future research.

Applying science to pressing conservation needs for penguins.

More than half of the world's 18 penguin species are declining. We, the Steering Committee of the International Union for Conservation of Nature Species Survival Commission Penguin Specialist Group, determined that the penguin species in most critical need of conservation action are African penguin (Spheniscus demersus), Galápagos penguin (Spheniscus mendiculus), and Yellow-eyed penguin (Megadyptes antipodes). Due to small or rapidly declining populations, these species require immediate scientific collaboration and policy intervention. We also used a pairwise-ranking approach to prioritize research and conservation needs for all penguins. Among the 12 cross-taxa research areas we identified, we ranked quantifying population trends, estimating demographic rates, forecasting environmental patterns of change, and improving the knowledge of fisheries interactions as the highest priorities. The highest ranked conservation needs were to enhance marine spatial planning, improve stakeholder engagement, and develop disaster-management and species-specific action plans. We concurred that, to improve the translation of science into effective conservation for penguins, the scientific community and funding bodies must recognize the importance of and support long-term research; research on and conservation of penguins must expand its focus to include the nonbreeding season and juvenile stage; marine reserves must be designed at ecologically appropriate spatial and temporal scales; and communication between scientists and decision makers must be improved with the help of individual scientists and interdisciplinary working groups.

Aplicación de Ciencia en las Necesidades de Conservación Urgentes para los Pingüinos. Resumen Más de la mitad de las 18 especies de pingüinos del mundo están disminuyendo. Nosotros, el Comité Directivo de la Unión Internacional para la Conservación de la Naturaleza, Grupo de Especialistas en Pingüinos, determinamos que las especies de pingüinos con necesidades críticas de conservación son el pingüino africano (Spheniscus demersus), el pingüino de las Galápagos (Spheniscus mendiculus) y el pingüino de ojos amarillos (Megadyptes antipodes). Debido a que sus poblaciones son pequeñas o están declinando rápidamente, estos pingüinos requieren colaboración científica e intervención política inmediatas. También utilizamos un método de clasificación por pares para priorizar las necesidades de investigación y conservación para todas las especies de pingüinos. Entre las 12 áreas de investigación que identificamos, las más prioritarias fueron: cuantificación de las tendencias poblacionales, estimación de las tasas demográficas, predicción de las patrones de cambio ambiental y mejora del conocimiento de las interacciones con pesquerías. Las mayores necesidades de conservación fueron: optimizar la planificación marina espacial, mejorar la colaboración de las partes interesadas y desarrollar planes de manejo de desastres y de acción para cada especie. Coincidimos en que, para mejorar la traducción de la ciencia en la conservación efectiva de los pingüinos, la comunidad científica y los organismos financiadores deben reconocer la importancia de la investigación a largo plazo y apoyarla; la investigación sobre pingüinos y su conservación debe expandir su enfoque para incluir la época no reproductiva y la etapa juvenil; las reservas marinas deben ser diseñadas a escalas espaciotemporales ecológicamente apropiadas; y la comunicación entre científicos y tomadores de decisiones debe mejorar con la ayuda de científicos individuales y grupos de trabajo interdisciplinario.

Unravelling the relative roles of top-down and bottom-up forces driving population change in an oceanic predator.

In the open ocean ecosystem, climate and anthropogenic changes have driven biological change at both ends of the food chain. Understanding how the population dynamics of pelagic predators are simultaneously influenced by nutrient-driven processes acting from the "bottom-up" and predator-driven processes acting from the "top-down" is therefore considered an urgent task. Using a state-space demographic model, we evaluated the population trajectory of an oceanic predator, the Macaroni Penguin (Eudyptes chrysolophus), and numerically assessed the relative importance of bottom-up and top-down drivers acting through different demographic rates. The population trajectory was considerably more sensitive to changes in top-down control of survival compared to bottom-up control of survival or productivity. This study integrates a unique set of demographic and covariate data and highlights the benefits of using a single estimation framework to examine the links between covariates, demographic rates and population dynamics.

A draft fur seal genome provides insights into factors affecting SNP validation and how to mitigate them.

Custom genotyping arrays provide a flexible and accurate means of genotyping single nucleotide polymorphisms (SNPs) in a large number of individuals of essentially any organism. However, validation rates, defined as the proportion of putative SNPs that are verified to be polymorphic in a population, are often very low. A number of potential causes of assay failure have been identified, but none have been explored systematically. In particular, as SNPs are often developed from transcriptomes, parameters relating to the genomic context are rarely taken into account. Here, we assembled a draft Antarctic fur seal (Arctocephalus gazella) genome (assembly size: 2.41 Gb; scaffold/contig N50 : 3.1 Mb/27.5 kb). We then used this resource to map the probe sequences of 144 putative SNPs genotyped in 480 individuals. The number of probe-to-genome mappings and alignment length together explained almost a third of the variation in validation success, indicating that sequence uniqueness and proximity to intron-exon boundaries play an important role. The same pattern was found after mapping the probe sequences to the Walrus and Weddell seal genomes, suggesting that the genomes of species divergent by as much as 23 million years can hold information relevant to SNP validation outcomes. Additionally, reanalysis of genotyping data from seven previous studies found the same two variables to be significantly associated with SNP validation success across a variety of taxa. Finally, our study reveals considerable scope for validation rates to be improved, either by simply filtering for SNPs whose flanking sequences align uniquely and completely to a reference genome, or through predictive modelling.

The influence of preceding dive cycles on the foraging decisions of Antarctic fur seals.

The foraging strategy of many animals is thought to be determined by their past experiences. However, few empirical studies have investigated whether this is true in diving animals. We recorded three-dimensional movements and mouth-opening events from three Antarctic fur seals during their foraging trips to examine how they adapt their behaviour based on past experience--continuing to search for prey in the same area or moving to search in a different place. Each dive cycle was divided into a transit phase and a feeding phase. The linear horizontal distance travelled after feeding phases in each dive was affected by the mouth-opening rate during the previous 244 s, which typically covered two to three dive cycles. The linear distance travelled tended to be shorter when the mouth-opening rate in the previous 244 s was higher, i.e. seals tended to stay in the same areas with high prey-encounter rates. These results indicate that Antarctic fur seals follow decision-making strategies based on the past foraging experience over time periods longer than the immediately preceding dive.

Where do penguins go during the inter-breeding period? Using geolocation to track the winter dispersion of the macaroni penguin.

Although penguins are key marine predators from the Southern Ocean, their migratory behaviour during the inter-nesting period remains widely unknown. Here, we report for the first time, to our knowledge, the winter foraging movements and feeding habits of a penguin species by using geolocation sensors fitted on penguins with a new attachment method. We focused on the macaroni penguin Eudyptes chrysolophus at Kerguelen, the single largest consumer of marine prey among all seabirds. Overall, macaroni penguins performed very long winter trips, remaining at sea during approximately six months within the limits of the Southern Ocean. They departed from Kerguelen in an eastward direction and distributed widely, over more than 3.10(6) km(2). The penguins spent most of their time in a previously unrecognized foraging area, i.e. a narrow latitudinal band (47-49 degrees S) within the central Indian Ocean (70-110 degrees E), corresponding oceanographically to the Polar Frontal Zone. There, their blood isotopic niche indicated that macaroni penguins preyed mainly upon crustaceans, but not on Antarctic krill Euphausia superba, which does not occur at these northern latitudes. Such winter information is a crucial step for a better integrative approach for the conservation of this species whose world population is known to be declining.

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.

Spatial and temporal operation of the Scotia Sea ecosystem: a review of large-scale links in a krill centred food web.

The Scotia Sea ecosystem is a major component of the circumpolar Southern Ocean system, where productivity and predator demand for prey are high. The eastward-flowing Antarctic Circumpolar Current (ACC) and waters from the Weddell-Scotia Confluence dominate the physics of the Scotia Sea, leading to a strong advective flow, intense eddy activity and mixing. There is also strong seasonality, manifest by the changing irradiance and sea ice cover, which leads to shorter summers in the south. Summer phytoplankton blooms, which at times can cover an area of more than 0.5 million km2, probably result from the mixing of micronutrients into surface waters through the flow of the ACC over the Scotia Arc. This production is consumed by a range of species including Antarctic krill, which are the major prey item of large seabird and marine mammal populations. The flow of the ACC is steered north by the Scotia Arc, pushing polar water to lower latitudes, carrying with it krill during spring and summer, which subsidize food webs around South Georgia and the northern Scotia Arc. There is also marked interannual variability in winter sea ice distribution and sea surface temperatures that is linked to southern hemisphere-scale climate processes such as the El Niño-Southern Oscillation. This variation affects regional primary and secondary production and influences biogeochemical cycles. It also affects krill population dynamics and dispersal, which in turn impacts higher trophic level predator foraging, breeding performance and population dynamics. The ecosystem has also been highly perturbed as a result of harvesting over the last two centuries and significant ecological changes have also occurred in response to rapid regional warming during the second half of the twentieth century. This combination of historical perturbation and rapid regional change highlights that the Scotia Sea ecosystem is likely to show significant change over the next two to three decades, which may result in major ecological shifts.

Female fur seals show active choice for males that are heterozygous and unrelated.

Much debate surrounds the exact rules that influence mating behaviour, and in particular the selective forces that explain the evolution of female preferences. A key example is the lek paradox, in which female choice is expected rapidly to become ineffective owing to loss of additive genetic variability for the preferred traits. Here we exploit a remarkable system in which female fur seals exert choice by moving across a crowded breeding colony to visit largely static males. We show that females move further to maximize the balance between male high multilocus heterozygosity and low relatedness. Such a system shows that female choice can be important even in a strongly polygynous species, and at the same time may help to resolve the lek paradox because heterozygosity has low heritability and inbreeding avoidance means there is no single 'best' male for all females.

Genetic tagging reveals extreme site fidelity in territorial male Antarctic fur seals Arctocephalus gazella.

Genetic tagging, the identification of individuals using their genotypes, provides a powerful tool for studying animals that are difficult to observe or identify using conventional techniques. However, despite being widely adopted by conservation biologists, the full potential of this approach has yet to be realized. Here we used genetic recapture data to quantify male site fidelity at a colony of Antarctic fur seals where an aerial walkway provides unprecedented access and individual positions are determined daily to 1 m accuracy. Because males are too large and aggressive to be captured and fitted with conventional tags, we remotely collected 770 tissue samples over eight consecutive seasons and used nine-locus microsatellite genotypes to reveal 306 genetic recaptures among 464 unique individuals. Within seasons, males are highly site-faithful, with any movements that occur tending to take place before the period when females come into oestrus. Of those males that return to breed over successive seasons, almost half return to within a body length of where they were before. The discovery of such extreme site faithfulness has implications for the population structure and mating system of fur seals and potentially other colonially breeding species.

Modelling Southern Ocean ecosystems: krill, the food-web, and the impacts of harvesting.

The ecosystem approach to fisheries recognises the interdependence between harvested species and other ecosystem components. It aims to account for the propagation of the effects of harvesting through the food-web. The formulation and evaluation of ecosystem-based management strategies requires reliable models of ecosystem dynamics to predict these effects. The krill-based system in the Southern Ocean was the focus of some of the earliest models exploring such effects. It is also a suitable example for the development of models to support the ecosystem approach to fisheries because it has a relatively simple food-web structure and progress has been made in developing models of the key species and interactions, some of which has been motivated by the need to develop ecosystem-based management. Antarctic krill, Euphausia superba, is the main target species for the fishery and the main prey of many top predators. It is therefore critical to capture the processes affecting the dynamics and distribution of krill in ecosystem dynamics models. These processes include environmental influences on recruitment and the spatially variable influence of advection. Models must also capture the interactions between krill and its consumers, which are mediated by the spatial structure of the environment. Various models have explored predator-prey population dynamics with simplistic representations of these interactions, while others have focused on specific details of the interactions. There is now a pressing need to develop plausible and practical models of ecosystem dynamics that link processes occurring at these different scales. Many studies have highlighted uncertainties in our understanding of the system, which indicates future priorities in terms of both data collection and developing methods to evaluate the effects of these uncertainties on model predictions. We propose a modelling approach that focuses on harvested species and their monitored consumers and that evaluates model uncertainty by using alternative structures and functional forms in a Monte Carlo framework.

Penguin-mounted cameras glimpse underwater group behaviour.

Marine birds and mammals spend most of their lives in the open ocean far from human observation, which makes obtaining information about their foraging behaviour difficult. Here, we show, by use of a miniaturized digital camera system, the first direct evidence (to our knowledge) of underwater group behaviour in free-ranging penguins. Penguins swim closely accompanied by other bird(s) during 24% of their possible foraging dives. This finding confirms that such miniaturized camera technology has broad applicability for advancing our knowledge about the previously unknown social interactions of marine animals at depth.

Environmental change and Antarctic seabird populations.

Recent changes in Antarctic seabird populations may reflect direct and indirect responses to regional climate change. The best long-term data for high-latitude Antarctic seabirds (Adélie and Emperor penguins and snow petrels) indicate that winter sea-ice has a profound influence. However, some effects are inconsistent between species and areas, some in opposite directions at different stages of breeding and life cycles, and others remain paradoxical. The combination of recent harvest driven changes and those caused by global warming may produce rapid shifts rather than gradual changes.