Coastal regions of the northern Antarctic Peninsula are key for gentoo populations.

Southern Ocean ecosystems are rapidly changing due to climate variability. An apparent beneficiary of such change in the western Antarctic Peninsula (WAP) is the gentoo penguin Pygoscelis papua, which has increased its population size and expanded its range southward in the last 20 years. To better understand how this species has responded to large-scale changes, we tracked individuals during the non-breeding winter period from five colonies across the latitudinal range of breeding sites in the WAP, including from a recently established colony. Results highlight latitudinal gradients in movement; strong associations with shallow, coastal habitats along the entire Antarctic Peninsula; and movements that are independent of, yet constrained by, sea ice. It is clear that coastal habitats essential to gentoo penguins during the breeding season are similarly critical during winter. Larger movements of birds from northern colonies in the WAP further suggest that leap-frog migration may influence colonization events by facilitating nest-area prospecting and use of new haul-out sites. Our results support efforts to develop a marine protected area around the WAP. Winter habitats used by gentoo penguins outline high priority areas for improving the management of the spatio-temporally concentrated krill (Euphausia superba) fishery that operates in this region during winter.

Acute bottlenecks to the survival of juvenile Pygoscelis penguins occur immediately after fledging.

Estimating when and where survival bottlenecks occur in free-ranging marine predators is critical for effective demographic monitoring and spatial planning. This is particularly relevant to juvenile stages of long-lived species for which direct observations of death are typically not possible. We used satellite telemetry data from fledgling Adélie, chinstrap and gentoo penguins near the Antarctic Peninsula to estimate the spatio-temporal scale of a bottleneck after fledging. Fledglings were tracked up to 106 days over distances of up to 2140 km. Cumulative losses of tags increased to 73% within 16 days of deployment, followed by an order-of-magnitude reduction in loss rates thereafter. The timing and location of tag losses were consistent with at-sea observations of penguin carcasses and bioenergetics simulations of mass loss to thresholds associated with low recruitment probability. A bootstrapping procedure is used to assess tag loss owing to death versus other factors. Results suggest insensitivity in the timing of the bottleneck and quantify plausible ranges of mortality rates within the bottleneck. The weight of evidence indicates that a survival bottleneck for fledgling penguins is acute, attributable to predation and starvation, and may account for at least 33% of juvenile mortality.

Comparing feedback and spatial approaches to advance ecosystem-based fisheries management in a changing Antarctic.

To implement ecosystem-based approaches to fisheries management, decision makers need insight on the potential costs and benefits of the policy options available to them. In the Southern Ocean, two such options for addressing trade-offs between krill-dependent predators and the krill fishery include "feedback management" (FBM) strategies and marine protected areas (MPAs); in theory, the first adjusts to change, while the latter is robust to change. We compared two possible FBM options to a proposed MPA in the Antarctic Peninsula and Scotia Sea given a changing climate. One of our feedback options, based on the density of Antarctic krill (Euphasia superba), projected modest increases in the abundances of some populations of krill predators, whereas outcomes from our second FBM option, based on changes in the abundances of penguins, were more mixed, with some areas projecting predator population declines. The MPA resulted in greater increases in some, but not all, predator populations than either feedback strategy. We conclude that these differing outcomes relate to the ways the options separate fishing and predator foraging, either by continually shifting the spatial distribution of fishing away from potentially vulnerable populations (FBM) or by permanently closing areas to fishing (the MPA). For the krill fishery, we show that total catches could be maintained using an FBM approach or slightly increased with the MPA, but the fishery would be forced to adjust fishing locations and sometimes fish in areas of relatively low krill density-both potentially significant costs. Our work demonstrates the potential to shift, rather than avoid, ecological risks and the likely costs of fishing, indicating trade-offs for decision makers to consider.

What's the catch? Profiling the benefits and costs associated with marine protected areas and displaced fishing in the Scotia Sea.

Both costs and benefits must be considered when implementing marine protected areas (MPAs), particularly those associated with fishing effort displaced by potential closures. The Southern Ocean offers a case study in understanding such tradeoffs, where MPAs are actively being discussed to achieve a range of protection and sustainable use objectives. Here, we evaluated the possible impacts of two MPA scenarios on the Antarctic krill (Euphausia superba) fishery and krill-dependent predators in the Scotia Sea, explicitly addressing the displacement of fishing from closed areas. For both scenarios, we employed a minimally realistic, spatially explicit ecosystem model and considered three alternative redistributions of displaced fishing. We projected both MPAs to provide positive outcomes for many krill-dependent predators, especially when closed areas included at least 50-75% of their foraging distributions. Further, differences between the scenarios suggest ways to improve seal and penguin protection in the Scotia Sea. MPA scenarios also projected increases in total fishery yields, but alongside risks of fishing in areas where relatively low krill densities could cause the fishery to suspend operations. The three alternatives for redistributing displaced fishing had little effect on benefits to predators, but did matter for the fishery, with greater differences in overall catch and risk of fishing in areas of low krill density when displaced fishing was redistributed evenly among the open areas. Collectively, results suggest a well-designed MPA in the Scotia Sea may protect krill-dependent predators, even with displaced fishing, and preclude further spatial management of the krill fishery outside the MPA. More broadly, outcomes denote the importance of delineating fishing and predator habitat, spatial scales, and the critical trade-offs inherent in MPA development.

Long-term observations from Antarctica demonstrate that mismatched scales of fisheries management and predator-prey interaction lead to erroneous conclusions about precaution.

Low catch limits for forage species are often considered to be precautionary measures that can help conserve marine predators. Difficulties measuring the impacts of fisheries removals on dependent predators maintain this perspective, but consideration of the spatio-temporal scales over which forage species, their predators, and fisheries interact can aid assessment of whether low catch limits are as precautionary as presumed. Antarctic krill are targeted by the largest fishery in the Southern Ocean and are key forage for numerous predators. Current krill removals are considered precautionary and have not been previously observed to affect krill-dependent predators, like penguins. Using a hierarchical model and 30+ years of monitoring data, we show that expected penguin performance was reduced when local harvest rates of krill were ≥0.1, and this effect was similar in magnitude to that of poor environmental conditions. With continued climate warming and high local harvest rates, future observations of penguin performance are predicted to be below the long-term mean with a probability of 0.77. Catch limits that are considered precautionary for forage species simply because the limit is a small proportion of the species' standing biomass may not be precautionary for their predators.

Individual variation in migratory movements of chinstrap penguins leads to widespread occupancy of ice-free winter habitats over the continental shelf and deep ocean basins of the Southern Ocean.

A goal of tracking migratory animals is to characterize the habitats they use and to interpret population processes with respect to conditions experienced en route to, and within, overwintering areas. For migratory seabirds with broad breeding ranges, inferring population-level effects of environmental conditions that are experienced during migratory periods would benefit by directly comparing how birds from different breeding aggregations disperse, characterizing the physical conditions of areas they use, and determining whether they occupy shared foraging areas. We therefore tracked 41 adult and juvenile chinstrap penguins (Pygoscelis antarctica) from three breeding locations in the northern Antarctic Peninsula region during the austral winter of 2017. The satellite tracking data revealed overlap of individuals over continental shelf areas during autumn months (Mar-May), shared outbound corridors that track the southern Antarctic circumpolar current front, followed by occupancy of progressively colder, deeper, and ice-free waters that spanned the entire western hemisphere south of the Polar Front. Despite broadly similar physical environments used by individuals from different colonies, the proportion of birds from each colony that remained within 500km of their colony was positively correlated with their local population trends. This suggests that local migration strategies near the Antarctic Peninsula may benefit breeding populations. However, the magnitude of inter-colony and intra-colony overlap was generally low given the broad scale of habitats occupied. High individual variation in winter movements suggests that habitat selection among chinstrap penguins is more opportunistic, without clear colony-specific preference for fine-scale foraging hotspots. Mixing of individuals from multiple colonies across broad regions of the Southern Ocean would expose chinstrap penguins from the Antarctic Peninsula to a shared environmental experience that helps explain the regional decline in their abundance.

Using sea-ice to calibrate a dynamic trophic model for the Western Antarctic Peninsula.

The pelagic ecosystems of the Western Antarctic Peninsula are dynamic and changing rapidly in the face of sustained warming. There is already evidence that warming may be impacting the food web. Antarctic krill, Euphausia superba, is an ice-associated species that is both an important prey item and the target of the only commercial fishery operating in the region. The goal of this study is to develop a dynamic trophic model for the region that includes the impact of the sea-ice regime on krill and krill predators. Such a model may be helpful to fisheries managers as they develop new management strategies in the face of continued sea-ice loss. A mass balanced food-web model (Ecopath) and time dynamic simulations (Ecosim) were created. The Ecopath model includes eight currently monitored species as single species to facilitate its future development into a model that could be used for marine protected area planning in the region. The Ecosim model is calibrated for the years 1996-2012. The successful calibration represents an improvement over existing Ecopath models for the region. Simulations indicate that the role of sea ice is both central and complex. The simulations are only able to recreate observed biomass trends for the monitored species when metrics describing the sea-ice regime are used to force key predator-prey interactions, and to drive the biomasses of Antarctic krill and the fish species Gobionotothen gibberifrons. This model is ready to be used for exploring results from sea-ice scenarios or to be developed into a spatial model that informs discussions regarding the design of marine protected areas in the region.

Parameter estimation using randomized phases in an integrated assessment model for Antarctic krill.

An integrated model assessing the status and productivity of Antarctic krill (Euphausia superba, hereafter krill) was configured to estimate different subsets of 118 potentially estimable parameters in alternative configurations. We fixed the parameters that were not estimated in any given configuration at pre-specified values. The model was fitted to over forty years of fisheries and survey data for krill in Subarea 48.1, a statistical reporting area around the Antarctic Peninsula used by the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR). The number of estimated parameters was gradually increased across model configurations. Configurations that estimated more parameters fitted the data better, but the order in which the parameters were estimated became more important in finding the best fit. Twenty-two configurations estimating from 48 to 107 parameters were able to obtain an invertible Hessian matrix that was subsequently used to estimate parameter uncertainty. Parameter uncertainties calculated using asymptotic approximation around the maximum likelihood estimates were often larger than uncertainties based on Markov chain Monte Carlo sampling for the same parameters. Diagnostics applied to MCMC samples in the best model of each configuration that obtained an invertible Hessian indicated that the most highly parameterized configurations did not reach stationary distributions. A 96-parameter configuration was the best fitting model of those that passed the MCMC diagnostics. The ΔAIC and ΔBIC scores indicated essentially no support relative to the best model for the alternative models that also passed MCMC diagnostics. Simulated data using the configurations as operating models showed that while all configurations passed "self-tests" for spawning biomass and recruitment, there was a small negative bias due to model penalties in the fishing mortality estimates for years with the highest fishing mortalities. "Cross-tests" of configurations that estimated different parameters often differed from the operating model values.

Impacts of rising sea temperature on krill increase risks for predators in the Scotia Sea.

Climate change is a threat to marine ecosystems and the services they provide, and reducing fishing pressure is one option for mitigating the overall consequences for marine biota. We used a minimally realistic ecosystem model to examine how projected effects of ocean warming on the growth of Antarctic krill, Euphausia superba, might affect populations of krill and dependent predators (whales, penguins, seals, and fish) in the Scotia Sea. We also investigated the potential to mitigate depletion risk for predators by curtailing krill fishing at different points in the 21st century. The projected effects of ocean warming on krill biomass were strongest in the northern Scotia Sea, with a ≥40% decline in the mass of individual krill. Projections also suggest a 25% chance that krill biomass will fall below an established depletion threshold (75% of its unimpacted level), with consequent risks for some predator populations, especially penguins. Average penguin abundance declined by up to 30% of its unimpacted level, with up to a 50% chance of falling below the depletion threshold. Simulated krill fishing at currently permitted harvest rates further increased risks for depletion, and stopping fishing offset the increased risks associated with ocean warming in our model to some extent. These results varied by location and species group. Risk reductions at smaller spatial scales also differed from those at the regional level, which suggests that some predator populations may be more vulnerable than others to future changes in krill biomass. However, impacts on predators did not always map directly to those for krill. Our findings indicate the importance of identifying vulnerable marine populations and targeting protection measures at appropriate spatial scales, and the potential for spatially-structured management to avoid aggravating risks associated with rising ocean temperatures. This may help balance tradeoffs among marine ecosystem services in an uncertain future.

Identifying Risk: Concurrent Overlap of the Antarctic Krill Fishery with Krill-Dependent Predators in the Scotia Sea.

Mitigating direct and indirect interactions between marine predators and fisheries is a motivating factor for ecosystem-based fisheries management (EBFM), especially where predators and fisheries compete for a shared resource. One difficulty in advancing EBFM is parameterizing clear functional responses of predators to indices of prey availability. Alternative characterizations of fishery-predator interactions may therefore benefit the implementation of EBFM. Telemetry data identify foraging areas used by predators and, therefore, represent critical information to mitigate potential competition between predators and fisheries. We analyzed six years (2009-2014) of telemetry data collected at Cape Shirreff, Livingston Island and Admiralty Bay, King George Island, Antarctica, on three species of Pygoscelid penguins and female Antarctic fur seals. In this region, all four species are primarily dependent on Antarctic krill. The tracking data demonstrate local movements near breeding colonies during the austral summer and dispersal from breeding colonies during the winter. We then assessed overlap between predators and the Antarctic krill fishery on a suite of spatiotemporal scales to examine how different data aggregations affect the extent and location of overlap. Concurrent overlap was observed on all spatiotemporal scales considered throughout the Antarctic Peninsula and South Orkney Islands region, including near tagging locations and in distant areas where recent fishing activity has concentrated. Overlap occurred at depths where mean krill densities were relatively high. Our results demonstrate that direct overlap of krill-dependent predators with the krill fishery on small spatiotemporal scales is relatively common throughout the Antarctic Peninsula region. As the krill fishery continues to develop and efforts to implement ecosystem-based management mature, indices of overlap may provide a useful metric for indicating where the risks of fishing are highest. A precautionary approach to allocating krill catches in space would be to avoid large increases in catch where overlap on small spatiotemporal scales is common.

Variability in krill biomass links harvesting and climate warming to penguin population changes in Antarctica.

The West Antarctic Peninsula (WAP) and adjacent Scotia Sea support abundant wildlife populations, many of which were nearly extirpated by humans. This region is also among the fastest-warming areas on the planet, with 5-6 °C increases in mean winter air temperatures and associated decreases in winter sea-ice cover. These biological and physical perturbations have affected the ecosystem profoundly. One hypothesis guiding ecological interpretations of changes in top predator populations in this region, the "sea-ice hypothesis," proposes that reductions in winter sea ice have led directly to declines in "ice-loving" species by decreasing their winter habitat, while populations of "ice-avoiding" species have increased. However, 30 y of field studies and recent surveys of penguins throughout the WAP and Scotia Sea demonstrate this mechanism is not controlling penguin populations; populations of both ice-loving Adélie and ice-avoiding chinstrap penguins have declined significantly. We argue in favor of an alternative, more robust hypothesis that attributes both increases and decreases in penguin populations to changes in the abundance of their main prey, Antarctic krill. Unlike many other predators in this region, Adélie and chinstrap penguins were never directly harvested by man; thus, their population trajectories track the impacts of biological and environmental changes in this ecosystem. Linking trends in penguin abundance with trends in krill biomass explains why populations of Adélie and chinstrap penguins increased after competitors (fur seals, baleen whales, and some fishes) were nearly extirpated in the 19th to mid-20th centuries and currently are decreasing in response to climate change.

Divergent responses of Pygoscelis penguins reveal a common environmental driver.

The responses of predators to environmental variability in the Antarctic Peninsula region have exhibited divergent patterns owing to variation in the geographic settings of colonies and predator life-history strategies. Five breeding colonies of Pygoscelis penguins from King George Island and Livingston Island, South Shetland Islands, Antarctica, were examined to (1) compare the responses of sympatric congeners to recent changes in their Antarctic ecosystem and (2) assess underlying causes for such responses. We used linear regression and correlation analyses to compare indices of abundance, recruitment, and summer breeding performance of the Adélie (P. adeliae), gentoo (P. papua), and chinstrap penguins (P. antarctica). Breeding colonies of Adélie and chinstrap penguins have declined by roughly 50% since the mid-1970s, and recruitment indices of Adélie penguins have declined by roughly 80%, but no such patterns are evident for gentoo penguins. Fledging success, however, has remained stable at all breeding colonies. The different trends in abundance and recruitment indices for each species, despite generally similar indices of summer performance, suggest that winter conditions contribute to the divergent responses among the penguins. In particular, strong correlations between indices of penguin and krill recruitment suggest that penguins in the South Shetland Islands may live under an increasingly krill-limited system that has disproportionate effects on the survival of juvenile birds.