Simeon L. Hill

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.

End-To-End Models for the Analysis of Marine Ecosystems: Challenges, Issues, and Next Steps

Abstract There is growing interest in models of marine ecosystems that deal with the effects of climate change through the higher trophic levels. Such end-to-end models combine physicochemical oceanographic descriptors and organisms ranging from microbes to higher-trophic-level (HTL) organisms, including humans, in a single modeling framework. The demand for such approaches arises from the need for quantitative tools for ecosystem-based management, particularly models that can deal with bottom-up and top-down controls that operate simultaneously and vary in time and space and that are capable of handling the multiple impacts expected under climate change. End-to-end models are now feasible because of improvements in the component submodels and the availability of sufficient computing power. We discuss nine issues related to the development of end-to-end models. These issues relate to formulation of the zooplankton submodel, melding of multiple temporal and spatial scales, acclimation and adaptation, behavioral movement, software and technology, model coupling, skill assessment, and interdisciplinary challenges. We urge restraint in using end-to-end models in a true forecasting mode until we know more about their performance. End-to-end models will challenge the available data and our ability to analyze and interpret complicated models that generate complex behavior. End-to-end modeling is in its early developmental stages and thus presents an opportunity to establish an open-access, community-based approach supported by a suite of true interdisciplinary efforts.

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.

Potential Climate Change Effects on the Habitat of Antarctic Krill in the Weddell Quadrant of the Southern Ocean

Antarctic krill is a cold water species, an increasingly important fishery resource and a major prey item for many fish, birds and mammals in the Southern Ocean. The fishery and the summer foraging sites of many of these predators are concentrated between 0° and 90°W. Parts of this quadrant have experienced recent localised sea surface warming of up to 0.2°C per decade, and projections suggest that further widespread warming of 0.27° to 1.08°C will occur by the late 21st century. We assessed the potential influence of this projected warming on Antarctic krill habitat with a statistical model that links growth to temperature and chlorophyll concentration. The results divide the quadrant into two zones: a band around the Antarctic Circumpolar Current in which habitat quality is particularly vulnerable to warming, and a southern area which is relatively insensitive. Our analysis suggests that the direct effects of warming could reduce the area of growth habitat by up to 20%. The reduction in growth habitat within the range of predators, such as Antarctic fur seals, that forage from breeding sites on South Georgia could be up to 55%, and the habitat’s ability to support Antarctic krill biomass production within this range could be reduced by up to 68%. Sensitivity analysis suggests that the effects of a 50% change in summer chlorophyll concentration could be more significant than the direct effects of warming. A reduction in primary production could lead to further habitat degradation but, even if chlorophyll increased by 50%, projected warming would still cause some degradation of the habitat accessible to predators. While there is considerable uncertainty in these projections, they suggest that future climate change could have a significant negative effect on Antarctic krill growth habitat and, consequently, on Southern Ocean biodiversity and ecosystem services.

Decision‐making for ecosystem‐based management: evaluating options for a krill fishery with an ecosystem dynamics model

Decision-makers charged with implementing ecosystem-based management (EBM) rely on scientists to predict the consequences of decisions relating to multiple, potentially conflicting, objectives. Such predictions are inherently uncertain, and this can be a barrier to decision-making. The Convention on the Conservation of Antarctic Marine Living Resources requires managers of Southern Ocean fisheries to sustain the productivity of target stocks, the health and resilience of the ecosystem, and the performance of the fisheries themselves. The managers of the Antarctic krill fishery in the Scotia Sea and southern Drake Passage have requested advice on candidate management measures consisting of a regional catch limit and options for subdividing this among smaller areas. We developed a spatially resolved model that simulates krill-predator-fishery interactions and reproduces a plausible representation of past dynamics. We worked with experts and stakeholders to identify (1) key uncertainties affecting our ability to predict ecosystem state; (2) illustrative reference points that represent the management objectives; and (3) a clear and simple way of conveying our results to decision-makers. We developed four scenarios that bracket the key uncertainties and evaluated candidate management measures in each of these scenarios using multiple stochastic simulations. The model emphasizes uncertainty and simulates multiple ecosystem components relating to diverse objectives. We summarize the potentially complex results as estimates of the risk that each illustrative objective will not be achieved (i.e., of the state being outside the range specified by the reference point). This approach allows direct comparisons between objectives. It also demonstrates that a candid appraisal of uncertainty, in the form of risk estimates, can be an aid, rather than a barrier, to understanding and using ecosystem model predictions. Management measures that reduce coastal fishing, relative to oceanic fishing, apparently reduce risks to both the fishery and the ecosystem. However, alternative reference points could alter the perceived risks, so further stakeholder involvement is needed to identify risk metrics that appropriately represent their objectives.

Ecosystem services of the Southern Ocean: trade-offs in decision-making

Abstract Ecosystem services are the benefits that mankind obtains from natural ecosystems. Here we identify the key services provided by the Southern Ocean. These include provisioning of fishery products, nutrient cycling, climate regulation and the maintenance of biodiversity, with associated cultural and aesthetic benefits. Potential catch limits for Antarctic krill (Euphausia superba Dana) alone are equivalent to 11% of current global marine fisheries landings. We also examine the extent to which decision-making within the Antarctic Treaty System (ATS) considers trade-offs between ecosystem services, using the management of the Antarctic krill fishery as a case study. Management of this fishery considers a three-way trade-off between fisheries performance, the status of the krill stock and that of predator populations. However, there is a paucity of information on how well these components represent other ecosystem services that might be degraded as a result of fishing. There is also a lack of information on how beneficiaries value these ecosystem services. A formal ecosystem assessment would help to address these knowledge gaps. It could also help to harmonize decision-making across the ATS and promote global recognition of Southern Ocean ecosystem services by providing a standard inventory of the relevant ecosystem services and their value to beneficiaries.

Prospects for a Sustainable Increase in the Availability of Long Chain Omega 3s: Lessons from the Antarctic Krill Fishery

The Global Summit on Nutrition, Health and Human Behaviour (GSNHHB) identified a target intake of long chain omega-3 (LC-omega-3) of around 1 g/day and therefore a need to “increase the availability of LC-omega-3 (especially DHA) for human consumption in a sustainable, environmentally responsible way” (1). Papers elsewhere in this volume make the case for increased consumption of LC-omega-3. The issue of a sustainable increase in availability also merits serious consideration. Marine fish are the main source of the two key LC-omega-3s for human consumption: docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA) (2). Any increase in demand for of LC-omega-3 is likely to increase pressure on marine living resources. Historically, increases in demand for marine living resources have often resulted in the degradation of marine ecosystems’ ability to supply the relevant product. The GSNHHB’s commitment to sustainability and environmental responsibility indicates an intention to avoid exacerbating this situation, but it also presents a considerable challenge in terms of both defining and achieving sustainability.

The Importance of Krill Predation in the Southern Ocean

Antarctic krill is a major prey species for a diverse array of Southern Ocean predators. The amount of krill that predators consume, and how this changes over space and time, is a key issue in understanding both regional and circumpolar aspects of the Southern Ocean food-web. We assess current knowledge of consumption by the various predator groups, and the ecological processes through which krill and its predators influence each other. Knowledge has improved greatly over recent decades and has revealed a high level of complexity in the processes that govern krill consumption. We focus on the Antarctic Peninsula and Scotia Sea region where both krill and its consumers occur in significant concentrations and where an updated estimate of krill consumption by the main vertebrate groups is 55 million tonnes per year. Research has mainly focused on mammalian and avian predators of post-larval krill, particularly penguins. Potentially important consumer groups like fish, cephalopods and carnivorous zooplankton remain poorly understood, as does consumption of the early life stages of krill. As a consequence of these knowledge gaps and the variability that arises from complexity, a reliable seasonally, spatially or taxonomically resolved description of krill consumption remains elusive. One of the key motivations for attempting to estimate krill consumption is to understand how changes in krill availability impact predator populations. Such understanding is an important requirement for ecosystem based management of the Antarctic krill fishery. We therefore propose that integrated study areas in strategic fishing locations should be developed to directly assess the response of krill predators to changes in krill availability.

From strategic ambiguity to technical reference points in the Antarctic krill fishery: the worst journey in the world?

The goals of ecosystem based management (EBM) are strategically ambiguous, meaning that they require interpretation to identify objectives for ecosystem state. Ecosystem states that are useful for achieving such objectives are known as reference points. Soft reference points specify both a state and a probability of the ecosystem being in that state. They are used with simulation models to identify management measures for which the risk of the ecosystem entering an undesirable state is below a specified level. The Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) is responsible for the EBM of Antarctic krill fisheries. CCAMLR used soft reference points for the krill stock in the Scotia Sea and southern Drake Passage to set a regional catch limit. However, this catch limit needs spatial subdivision to protect predators from localized depletion. Model-based evaluations of different options for subdividing the catch limit used illustrative reference points to assess the depletion risk to multiple predators. This study demonstrates that the apparent risk is sensitive to the choice of reference point and method for aggregating modelled predators. EBM practitioners and stakeholders need to be aware that these factors could therefore bias comparisons of management measures. Nonetheless, qualitative distinctions between different spatial subdivision options are relatively consistent except at high levels of aggregation and extreme reference points. This study also demonstrates a lack of generality in the relationship between current and future ecosystem state. Thus, the EBM goal of maintaining ecosystem resilience implies different reference points for the current state of different ecosystem components. Despite early progress in defining soft reference points for the krill stock, CCAMLR has not yet defined reference points for krill predators. Structured dialogue aimed at identifying collective objectives might be necessary to achieve further progress in CCAMLR and other EBM organizations.

Distribution of short-finned squid Illex argentinus (Cephalopoda: Ommastrephidae) inferred from the diets of Southern Ocean albatrosses using stable isotope analyses

The diets of marine predators are a potential source of information about range shifts in their prey. For example, the short-finned squid Illex argentinus, a commercially fished species on the Patagonian Shelf in the South Atlantic, has been reported in the diet of grey-headed, Thalassarche chrysostoma; black-browed, T. melanophris; and wandering, Diomedea exulans, albatrosses breeding at Bird Island, South Georgia (54°S 28°W) in the Southern Ocean. Tracking data suggest that these birds may feed on I. argentinus while foraging in Southern Ocean waters during their breeding season. This led to the hypothesis that I. argentinus may occur south of the Antarctic Polar Front. To test this hypothesis, we used stable isotope analyses to assess the origin of I. argentinus. We compared I. argentinus beaks from the diets of the three albatross species with beaks of cephalopod species endemic to the Patagonian Shelf and others from the Southern Ocean. Our results show that I. argentinus from the diet of albatrosses at Bird Island have ?13C values in the range ?18.77 to ?15.28‰. This is consistent with ?13C values for Octopus tehuelchus, a typical species from the Patagonian Shelf. In contrast, Alluroteuthis antarcticus, a Southern Ocean squid, has typically Antarctic ?13C in the range ?25.46 to ?18.61‰. This suggests that I. argentinus originated from warmer waters of the Patagonian Shelf region. It is more likely that the albatross species obtained I. argentinus by foraging in the Patagonian Shelf region than that I. argentinus naturally occurs south of the Antarctic Polar Front.