Many free-ranging predators have to make foraging decisions with little, if any, knowledge of present resource distribution and availability. The optimal search strategy they should use to maximize encounter rates with prey in heterogeneous natural environments remains a largely unresolved issue in ecology. Lévy walks are specialized random walks giving rise to fractal movement trajectories that may represent an optimal solution for searching complex landscapes. However, the adaptive significance of this putative strategy in response to natural prey distributions remains untested. Here we analyse over a million movement displacements recorded from animal-attached electronic tags to show that diverse marine predators-sharks, bony fishes, sea turtles and penguins-exhibit Lévy-walk-like behaviour close to a theoretical optimum. Prey density distributions also display Lévy-like fractal patterns, suggesting response movements by predators to prey distributions. Simulations show that predators have higher encounter rates when adopting Lévy-type foraging in natural-like prey fields compared with purely random landscapes. This is consistent with the hypothesis that observed search patterns are adapted to observed statistical patterns of the landscape. This may explain why Lévy-like behaviour seems to be widespread among diverse organisms, from microbes to humans, as a 'rule' that evolved in response to patchy resource distributions.

An optimal search theory, the so-called Lévy-flight foraging hypothesis 1 , predicts that predators should adopt search strategies known as Lévy flights where prey is sparse and distributed unpredictably, but that Brownian movement is sufficiently efficient for locating abundant prey [2][3][4] . Empirical studies have generated controversy because the accuracy of statistical methods that have been used to identify Lévy behaviour has recently been questioned 5,6 . Consequently, whether foragers exhibit Lévy flights in the wild remains unclear. Crucially, moreover, it has not been tested whether observed movement patterns across natural landscapes having different expected resource distributions conform to the theory's central predictions. Here we use maximum-likelihood methods to test for Lévy patterns in relation to environmental gradients in the largest animal movement data set assembled for this purpose. Strong support was found for Lévy search patterns across 14 species of open-ocean predatory fish (sharks, tuna, billfish and ocean sunfish), with some individuals switching between Lévy and Brownian movement as they traversed different habitat types. We tested the spatial occurrence of these two principal patterns and found Lévy behaviour to be associated with less productive waters (sparser prey) and Brownian movements to be associated with productive shelf or convergence-front habitats (abundant prey). These results are consistent with the Lévy-flight foraging hypothesis 1,7 , supporting the contention 8,9 that organism search strategies naturally evolved in such a way that they exploit optimal Lévy patterns.Lévy flights are a special class of random walk with movement displacements (steps) drawn from a probability distribution with a power-law tail (the so-called Pareto-Lévy distribution) 1,10 , and give rise to stochastic processes closely linked to fractal geometry and anomalous diffusion phenomena 7,11 . Lévy flights describe a movement pattern characterized by many small steps connected by longer relocations, with this pattern having scale invariance under projection, such that the probability density function, P(l j ), has a power-law tail in the long-distance regime: P(l j ) < l j 2m , where l j is the flight length (step length of move j), and m, 1 , m # 3, is the power-law exponent. Lévy flights comprise instantaneous steps and hence involve infinite velocities, whereas a Lévy walk 10 refers to a finitevelocity walk such that displacement is determined after a time t, reflecting a dynamical process such as movement 1,10,11 . Lévy flights and walks are theorized to be the most efficient movement pattern for locating patchy prey in low concentrations on spatial scales beyond a searcher's sensory range, with an optimal search having a power-law exponent of m < 2 (refs 4, 13). It is proposed that organisms have therefore naturally evolved search patterns that can be modelled as optimal Lévy flights 1,7,13 .However, burgeoning empirical support for this hypothesis recently foundered following studies sug...

Summary1. Diel vertical migration (DVM) is a widespread phenomenon among marine and freshwater organisms and many studies with various taxa have sought to understand its adaptive significance. Among crustacean zooplankton and juveniles of some fish species DVM is accepted widely as an antipredator behaviour, but little is known about its adaptive value for relatively large-bodied, adult predatory fish such as sharks. Moreover, the majority of studies have focused on pelagic forms, which raises the question of whether DVM occurs in bottom-living predators. 2. To investigate DVM in benthic predatory fish in the marine environment and to determine why it might occur we tracked movements of adult male dogfish ( Scyliorhinus canicula ) by short-and long-term acoustic and archival telemetry. Movement studies were complemented with measurements of prey abundance and availability and thermal habitat within home ranges. A thermal choice experiment and energy budget modelling was used to investigate trade-offs between foraging and thermal habitat selection. 3. Male dogfish undertook normal DVM (nocturnal ascent) within relatively small home ranges ( ∼ 100 × 100 m) comprising along-bottom movements up submarine slopes from deeper, colder waters occupied during the day into warmer, shallow prey-rich areas above the thermocline at night. Few daytime vertical movements occurred. Levels of activity were higher during the night above the thermocline compared to below it during the day indicating they foraged in warm water and rested in colder depths. 4. A thermal choice experiment using environmentally realistic temperatures supported the field observation that dogfish positively avoided warmer water even when it was associated with greater food availability. Males in laboratory aquaria moved into warm water from a cooler refuge only to obtain food, and after food consumption they preferred to rest and digest in cooler water. 5. Modelling of energy budgets under different realistic thermal-choice scenarios indicated dogfish adopting a 'hunt warm − rest cool' strategy could lower daily energy costs by just over 4%. Our results provide the first clear evidence that are consistent with the hypothesis that a benthic marine-fish predator utilizes DVM as an energy conservation strategy that increases bioenergetic efficiency.

Climatic change has been implicated as the cause of abundance fluctuations in marine fish populations worldwide, but the effects on whole communities are poorly understood. We examined the effects of regional climatic change on two fish assemblages using independent datasets from inshore marine (English Channel, 1913-2002) and estuarine environments (Bristol Channel, 1981-2001). Our results show that climatic change has had dramatic effects on community composition. Each assemblage contained a subset of dominant species whose abundances were strongly linked to annual mean sea-surface temperature. Species' latitudinal ranges were not good predictors of species-level responses, however, and the same species did not show congruent trends between sites. This suggests that within a region, populations of the same species may respond differently to climatic change, possibly owing to additional local environmental determinants, interspecific ecological interactions and dispersal capacity. This will make species-level responses difficult to predict within geographically differentiated communities.

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