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.

Summary1. Time and energy are key currencies in animal ecology, and judicious management of these is a primary focus for natural selection. At present, however, there are only two main methods for estimation of rate of energy expenditure in the field, heart rate and doubly labelled water, both of which have been used with success; but both also have their limitations. 2. The deployment of data loggers that measure acceleration is emerging as a powerful tool for quantifying the behaviour of free-living animals. Given that animal movement requires the use of energy, the accelerometry technique potentially has application in the quantification of rate of energy expenditure during activity. 3. In the present study, we test the hypothesis that acceleration can serve as a proxy for rate of energy expenditure in free-living animals. We measured rate of energy expenditure as rates of O 2 consumption ( ) and CO 2 production ( ) in great cormorants ( Phalacrocorax carbo ) at rest and during pedestrian exercise. and were then related to overall dynamic body acceleration (ODBA) measured with an externally attached three-axis accelerometer. 4. Both and were significantly positively associated with ODBA in great cormorants. This suggests that accelerometric measurements of ODBA can be used to estimate and and, with some additional assumptions regarding metabolic substrate use and the energy equivalence of O 2 and CO 2 , that ODBA can be used to estimate the activity specific rate of energy expenditure of free-living cormorants. 5. To verify that the approach identifies expected trends in from situations with variable power requirements, we measured ODBA in free-living imperial cormorants ( Phalacrocorax atriceps ) during foraging trips. We compared ODBA during return and outward foraging flights, when birds are expected to be laden and not laden with captured fish, respectively. We also examined changes in ODBA during the descent phase of diving, when power requirements are predicted to decrease with depth due to changes in buoyancy associated with compression of plumage and respiratory air. 6. In free-living imperial cormorants, ODBA, and hence estimated , was higher during the return flight of a foraging bout, and decreased with depth during the descent phase of a dive, supporting the use of accelerometry for the determination of activityspecific rate of energy expenditure.

Behavior is an important mechanism of evolution and it is paid for through energy expenditure. Nevertheless, field biologists can rarely observe animals for more than a fraction of their daily activities and attempts to quantify behavior for modeling ecological processes often exclude cryptic yet important behavioral events. Over the past few years, an explosion of research on remote monitoring of animal behavior using acceleration sensors has smashed the decades-old limits of observational studies. Animal-attached accelerometers measure the change in velocity of the body over time and can quantify fine-scale movements and body postures unlimited by visibility, observer bias, or the scale of space use. Pioneered more than a decade ago, application of accelerometers as a remote monitoring tool has recently surged thanks to the development of more accessible hardware and software. It has been applied to more than 120 species of animals to date. Accelerometer measurements are typically collected in three dimensions of movement at very high resolution (>10 Hz), and have so far been applied towards two main objectives. First, the patterns of accelerometer waveforms can be used to deduce specific behaviors through animal movement and body posture. Second, the variation in accelerometer waveform measurements has been shown to correlate with energy expenditure, opening up a suite of scientific questions in species notoriously difficult to observe in the wild. To date, studies of wild aquatic species outnumber wild terrestrial species and analyses of social behaviors are particularly few in number. Researchers of domestic and captive species also tend to report methodology more thoroughly than those studying species in the wild. There are substantial challenges to getting the most out of accelerometers, including validation, calibration, and the management and analysis of large quantities of data. In this review, we illustrate how accelerometers work, provide an overview of the ecological questions that have employed accelerometry, and highlight the emerging best practices for data acquisition and analysis. This tool offers a level of detail in behavioral studies of free-ranging wild animals that has previously been impossible to achieve and, across scientific disciplines, it improves understanding of the role of behavioral mechanisms in ecological and evolutionary processes. AbstractResumen: El comportamiento es un mecanismo importante de la evolución y que se paga a través del gasto de energía. Sin embargo, los biólogos de campo raramente observan los animales durante más de una fracción de sus actividades y los intentos de cuantificar el comportamiento para el modelado de los procesos ecológicos a menudo excluyen eventos crípticos pero importantes. En los últimos años se produjeron avances importantes en el monitoreo remoto del comportamiento de los animales, utilizando sensores de telemétro de aceleración (acelerómetros) que empujan los límites tradicionales de los estudios observacionales. Acelerómetros uni...

The advantages of transmission telemetry or logging systems for studying free-living animals are multiple and have driven designs for many and varied devices, each with its own particular usage sector. However, lack of fundamental data on species with conservation issues shows that there is an urgent need for a single generic system to document the major elements of animal biology. Such a tag could provide a broad picture of wild animal biology and specifically allow previously unidentified factors that might be important in an animal's conservation to be determined. This work describes the major features and operating mode of a single device, the 'daily diary', an animal equivalent of the aeroplane 'black box flight recorder' which is designed to be used on a wide variety of species and which has already been tested on animals including albatrosses, badgers, cheetahs, cormorants, domestic dogs, horses, penguins, sharks, sunfish and turtles. The unit is designed to record animal (1) movement (2) behaviour (3) energy expenditure and (4) the physical characteristics of the animal's environment by logging 14 parameters at infra-second frequencies.