Philip N. Trathan

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.

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.

Climatically driven fluctuations in Southern Ocean ecosystems

Determining how climate fluctuations affect ocean ecosystems requires an understanding of how biological and physical processes interact across a wide range of scales. Here we examine the role of physical and biological processes in generating fluctuations in the ecosystem around South Georgia in the South Atlantic sector of the Southern Ocean. Anomalies in sea surface temperature (SST) in the South Pacific sector of the Southern Ocean have previously been shown to be generated through atmospheric teleconnections with El Niño Southern Oscillation (ENSO)-related processes. These SST anomalies are propagated via the Antarctic Circumpolar Current into the South Atlantic (on time scales of more than 1 year), where ENSO and Southern Annular Mode-related atmospheric processes have a direct influence on short (less than six months) time scales. We find that across the South Atlantic sector, these changes in SST, and related fluctuations in winter sea ice extent, affect the recruitment and dispersal of Antarctic krill. This oceanographically driven variation in krill population dynamics and abundance in turn affects the breeding success of seabird and marine mammal predators that depend on krill as food. Such propagating anomalies, mediated through physical and trophic interactions, are likely to be an important component of variation in ocean ecosystems and affect responses to longer term change. Population models derived on the basis of these oceanic fluctuations indicate that plausible rates of regional warming of 1oC over the next 100 years could lead to more than a 95% reduction in the biomass and abundance of krill across the Scotia Sea by the end of the century.

THE EFFECTS OF GLOBAL CLIMATE VARIABILITY IN PUP PRODUCTION OF ANTARCTIC FUR SEALS

Climate variability has strong effects on marine ecosystems, with repercussions that range in scale from those that impact individuals to those that impact the entire food web. Climate-induced changes in the abundance of species in lower trophic levels can cascade up to apex predators by depressing vital rates. However, the characteristics and predictability of predator demographic responses remain largely unexplored. We investigated the detectability, limits, and nonlinearity of changes in Antarctic fur seal pup production at South Georgia over a 20-year period in response to environmental autocorrelation created by global climate perturbations; these were identified in time series of monthly averaged sea surface temperature (SST). Environmental autocorrelation at South Georgia was evident with frequent SST anomalies between 1990 and 1999, during a decade of warm background (time-averaged) conditions. SST anomalies were preceded by, and cross-correlated with, frequent El Nino-La Nina events between 1987 and 1998, which was also a decade of warm background conditions in the tropical Pacific Ocean. Nonlinear mixed-effects models indicated that positive anomalies at South Georgia explained extreme reductions in Antarctic fur seal pup production over 20 years of study. Simulated environmental time series suggested that the effect of anomalies on Antarctic fur seals was only detectable within a narrow range of positive SST, regardless of the distribution, variance, and autocorrelation structure in SST; this explained the observed nonlinearity in responses in pup production, which were observed only under persistent high SST levels. Such anomalies at South Georgia were likely associated with low availability of prey, largely krill, which affected Antarctic fur seal females over time scales longer than their breeding cycle. Reductions in Antarctic fur seal pup production could thus be predicted in advance by the detection of large-scale anomalies, which appeared to be driven by trends in global climate perturbation.

Key Questions in Marine Megafauna Movement Ecology

It is a golden age for animal movement studies and so an opportune time to assess priorities for future work. We assembled 40 experts to identify key questions in this field, focussing on marine megafauna, which include a broad range of birds, mammals, reptiles, and fish. Research on these taxa has both underpinned many of the recent technical developments and led to fundamental discoveries in the field. We show that the questions have broad applicability to other taxa, including terrestrial animals, flying insects, and swimming invertebrates, and, as such, this exercise provides a useful roadmap for targeted deployments and data syntheses that should advance the field of movement ecology.

Dynamics of antarctic penguin populations in relation to inter-annual variability in sea ice distribution

To investigate the role of sea ice cover on penguin populations we used principal component analysis to compare population variables of Adélie (Pygoscelis adeliae) and chinstrap (Pygoscelis antarctica) penguins breeding on Signy Island, South Orkney Islands with local (from direct observations) and regional (from remote sensing data) sea ice variables. Throughout the study period, the Adélie penguin population size remained stable, whereas that of chinstrap penguins decreased slightly. For neither species were there significant relationships between population size and breeding success, except for an apparent inverse density-dependent relationship between the number of Adélie breeding pairs and the number of eggs hatching. For both species, no general relationship was found between either population size or breeding success and the local sea ice conditions. However, the regional sea ice extent at a particular time prior to the start of the breeding season was related to the number of birds that arrived to breed. For both species, this period occurred before the sea ice reached its maximum extent and was slightly earlier for Adélie than for chinstrap penguins. These results suggest that sea ice conditions outside the breeding season may play an important role in penguin population processes.

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.

Remote sensing of the global light-fishing fleet: an analysis of interactions with oceanography, other fisheries and predators

Publisher Summary The use of United States Defense Meteorological Satellite Program Operational Linescan System (DMSP-OLS) data has enabled the precise location of the distribution of the global light-fishing fleet over a six month period in relation to the large and general mesoscale oceanography of the ecological provinces where they occur. The squid catch in these light fisheries can be identified to genera or species in seven ecological provinces where DMSP-OLS imagery reveals light-fishing activities. The DMSP-OLS data provide the information needed to review the relationship between the squid fisheries using lights and other fisheries for finfish with better spatial resolution than has been previously possible using data for FAO statistical areas alone. This chapter demonstrates that 62–70%, and possibly up to something

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.