The study of animal foraging behaviour is of practical ecological importance 1 , and exemplifies the wider scientific problem of optimizing search strategies 2 .Lévy flights are random walks whose step lengths come from probability distributions with heavy power-law tails 3, 4 , such that clusters of short steps are connected by rare long steps. flight durations (time intervals between landing on the ocean) were then calculated as consecutive hours for which a bird remained dry, to a resolution of 1 h. It was assumed that birds landed on the water solely to feed, and that flight durations were thus indicative of distances between prey.Time series for 19 separate foraging trips 7 were pooled to give a total of 363 3 flights. The resulting log-log histogram of flight durations gave a straight line with a slope of approximately 2, and is reproduced in Supplementary Fig. 1 from the original raw data. The crux of the conclusion that the albatrosses were performing Lévy flights was that the slope of 2 implied the probability density function (pdf) of flight durations t (in hours), was 7, 10for t ≥ 1 h (leaving out the normalization constant). This is consistent with the Lévy flight definition that the tail of the pdf is of the power-law form t −µ , where 1 < µ ≤ 3 (though technically this is a Lévy walk 4,7,22 We first analyze a newer, larger, and higher resolution data set of albatross flight durations to test for Lévy flights. In 2004, 20 wandering albatrosses on BirdIsland were each fitted with a salt-water logger and a GPS device. The GPS data were too infrequent (at most one location h −1 ) to give distances between landings, but were needed to estimate each bird's departure time from Bird Island, in order to calculate the duration of the initial flight before first landing on the water (we calculated return flights similarly). The resulting data set of flight records was 4 pooled, as in ref. 7, yielding a total of 1416 flights to a resolution of 10 s (Fig. 1).The flights ≥ 1 h are clearly inconsistent with coming from the power law t −2 ascertained 7 for the 1992 data. Furthermore, data from a power law of any exponent (not just 2) would yield a straight line 23 , and this is clearly not the case.In fact, the flight durations t (in h) are consistent with coming from the shifted gamma distribution given by the pdfwhere y = t − 1/120 accounts for the assumed 30 s period before the bird searches for new food sources (see Methods), s = 0.31 is the shape parameter, r = 0.41 h −1 is the rate parameter, and Γ(·) is the gamma function. Equation (2) is valid for flights >30 s; for shorter flights we have f (t) = 0. The exponential term of (2) dominates for large t, implying Poisson behaviour, such that for long enough flights the birds essentially encounter prey randomly with a constant low probability.A Brownian random walker's displacement increases as t H where H = 1/2.If H > 1/2, we have "superdiffusion" as originally inferred in Fig. 2a The gamma distribution (2) has µ = 1 − s = 0.69. This is such a slow powerlaw ...
Determining the form of key predator-prey relationships is critical for understanding marine ecosystem dynamics. Using a comprehensive global database, we quantified the effect of fluctuations in food abundance on seabird breeding success. We identified a threshold in prey (fish and krill, termed "forage fish") abundance below which seabirds experience consistently reduced and more variable productivity. This response was common to all seven ecosystems and 14 bird species examined within the Atlantic, Pacific, and Southern Oceans. The threshold approximated one-third of the maximum prey biomass observed in long-term studies. This provides an indicator of the minimal forage fish biomass needed to sustain seabird productivity over the long term.
Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.
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.
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.
Climate warming and associated sea ice reductions in Antarctica have modified habitat conditions for some species. These include the congeneric Adélie, chinstrap and gentoo penguins, which now demonstrate remarkable population responses to regional warming. However, inconsistencies in the direction of population changes between species at different study sites complicate the understanding of causal processes. Here, we show that at the South Orkney Islands where the three species breed sympatrically, the less iceadapted gentoo penguins increased significantly in numbers over the last 26 years, whereas chinstrap and Adélie penguins both declined. These trends occurred in parallel with regional long-term warming and significant reduction in sea ice extent. Periodical warm events, with teleconnections to the tropical Pacific, caused cycles in sea ice leading to reduced prey biomass, and simultaneous interannual population decreases in the three penguin species. With the loss of sea ice, Adélie penguins were less buffered against the environment, their numbers fluctuated greatly and their population response was strong and linear. Chinstrap penguins, considered to be better adapted to ice-free conditions, were affected by discrete events of locally increased ice cover, but showed less variable, nonlinear responses to sea ice loss. Gentoo penguins were temporarily affected by negative anomalies in regional sea ice, but persistent sea ice reductions were likely to increase their available niche, which is likely to be substantially segregated from that of their more abundant congeners. Thus, the regional consequences of global climate perturbations on the sea ice phenology affect the marine ecosystem, with repercussions for penguin food supply and competition for resources. Ultimately, variability in penguin populations with warming reflects the local balance between penguin adaptation to ice conditions and trophic-mediated changes cascading from global climate forcing.
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