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...

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Summary Fundamental ecological research is both intrinsically interesting and provides the basic knowledge required to answer applied questions of importance to the management of the natural world. The 100th anniversary of the British Ecological Society in 2013 is an opportune moment to reflect on the current status of ecology as a science and look forward to high‐light priorities for future work. To do this, we identified 100 important questions of fundamental importance in pure ecology. We elicited questions from ecologists working across a wide range of systems and disciplines. The 754 questions submitted (listed in the online appendix) from 388 participants were narrowed down to the final 100 through a process of discussion, rewording and repeated rounds of voting. This was done during a two‐day workshop and thereafter. The questions reflect many of the important current conceptual and technical pre‐occupations of ecology. For example, many questions concerned the dynamics of environmental change and complex ecosystem interactions, as well as the interaction between ecology and evolution. The questions reveal a dynamic science with novel subfields emerging. For example, a group of questions was dedicated to disease and micro‐organisms and another on human impacts and global change reflecting the emergence of new subdisciplines that would not have been foreseen a few decades ago. The list also contained a number of questions that have perplexed ecologists for decades and are still seen as crucial to answer, such as the link between population dynamics and life‐history evolution. Synthesis. These 100 questions identified reflect the state of ecology today. Using them as an agenda for further research would lead to a substantial enhancement in understanding of the discipline, with practical relevance for the conservation of biodiversity and ecosystem function.

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