Turn costs change the value of animal search paths.

The tortuosity of the track taken by an animal searching for food profoundly affects search efficiency, which should be optimised to maximise net energy gain. Models examining this generally describe movement as a series of straight steps interspaced by turns, and implicitly assume no turn costs. We used both empirical- and modelling-based approaches to show that the energetic costs for turns in both terrestrial and aerial locomotion are substantial, which calls into question the value of conventional movement models such as correlated random walk or Lévy walk for assessing optimum path types. We show how, because straight-line travel is energetically most efficient, search strategies should favour constrained turn angles, with uninformed foragers continuing in straight lines unless the potential benefits of turning offset the cost.

[1]  M. Kennedy,et al.  Factors Affecting Response of Raccoons to Traps and Population Size Estimation , 1985 .

[2]  Wilson,et al.  UNDERWATER SWIMMING AT LOW ENERGETIC COST BY PYGOSCELID PENGUINS , 1994, The Journal of experimental biology.

[3]  H. Larralde,et al.  Lévy walk patterns in the foraging movements of spider monkeys (Ateles geoffroyi) , 2003, Behavioral Ecology and Sociobiology.

[4]  L. Halsey,et al.  The relationship between oxygen consumption and body acceleration in a range of species. , 2009, Comparative biochemistry and physiology. Part A, Molecular & integrative physiology.

[5]  G. Viswanathan,et al.  Lévy flights and superdiffusion in the context of biological encounters and random searches , 2008 .

[6]  S. Benhamou How to reliably estimate the tortuosity of an animal's path: straightness, sinuosity, or fractal dimension? , 2004, Journal of theoretical biology.

[7]  Joel s. Brown,et al.  Foraging : behavior and ecology , 2007 .

[8]  A. Minetti,et al.  Skyscraper running: physiological and biomechanical profile of a novel sport activity , 2011, Scandinavian journal of medicine & science in sports.

[9]  G. Nevitt,et al.  Sensory ecology on the high seas: the odor world of the procellariiform seabirds , 2008, Journal of Experimental Biology.

[10]  Coen P. H. Elemans,et al.  Walking the line: search behavior and foraging success in ant species , 2011 .

[11]  C. J. Pennycuick,et al.  Modelling the Flying Bird , 2008 .

[12]  Susanne Åkesson,et al.  Island-finding ability of marine turtles , 2003, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[13]  Alan M. Wilson,et al.  Flying in a flock comes at a cost in pigeons , 2011, Nature.

[14]  Edward A. Codling,et al.  Random walk models in biology , 2008, Journal of The Royal Society Interface.

[15]  John David Anderson,et al.  Introduction to Flight , 1985 .

[16]  G. Saunders,et al.  Factors Affecting Bait Uptake and Trapping Success for Feral Pigs (Sus Scrofa) in Kosciusko National Park. , 1993 .

[17]  Charles H. Janson,et al.  Experimental analysis of food detection in capuchin monkeys: effects of distance, travel speed, and resource size , 1997, Behavioral Ecology and Sociobiology.

[18]  Francisco Bozinovic,et al.  The influence of habitat on travel speed, intermittent locomotion, and vigilance in a diurnal rodent , 2002 .

[19]  I. Boyd,et al.  Evaluating the Prudence of Parents: Daily Energy Expenditure Throughout the Annual Cycle of a Free-Ranging Bird, the Macaroni Penguin Eudyptes Chrysolophus , 2009 .

[20]  S. Cooke Biotelemetry and biologging in endangered species research and animal conservation: relevance to regional, national, and IUCN Red List threat assessments , 2008 .

[21]  K Schmidt-Nielsen,et al.  Locomotion: energy cost of swimming, flying, and running. , 1972, Science.

[22]  A. Hofgaard,et al.  Foraging and movement paths of female reindeer: insights from fractal analysis, correlated random walks, and Lévy flights , 2002 .

[23]  Martin Wikelski,et al.  Conservation physiology. , 2020, Trends in ecology & evolution.

[24]  H. Stanley,et al.  Optimizing the success of random searches , 1999, Nature.

[25]  Rory P. Wilson,et al.  Construction of energy landscapes can clarify the movement and distribution of foraging animals , 2012, Proceedings of the Royal Society B: Biological Sciences.

[26]  Nicolas E. Humphries,et al.  Scaling laws of marine predator search behaviour , 2008, Nature.

[27]  G. Pyke Optimal Foraging Theory: A Critical Review , 1984 .

[28]  P. A. Prince,et al.  Lévy flight search patterns of wandering albatrosses , 1996, Nature.

[29]  Simon Benhamou,et al.  Detecting an orientation component in animal paths when the preferred direction is individual-dependent. , 2006, Ecology.

[30]  Emily L. C. Shepard,et al.  Pushed for time or saving on fuel: fine-scale energy budgets shed light on currencies in a diving bird , 2009, Proceedings of the Royal Society B: Biological Sciences.

[31]  Rory P. Wilson,et al.  Prying into the intimate details of animal lives: use of a daily diary on animals , 2008 .

[32]  Sergio A. Lambertucci,et al.  Energy Landscapes Shape Animal Movement Ecology , 2013, The American Naturalist.

[33]  T H Witte,et al.  Accuracy of non-differential GPS for the determination of speed over ground. , 2004, Journal of biomechanics.

[34]  N. Lecomte Foraging: Behaviour and Ecology , 2008 .

[35]  Emily L. C. Shepard,et al.  Energy Beyond Food: Foraging Theory Informs Time Spent in Thermals by a Large Soaring Bird , 2011, PloS one.

[36]  Frederic Bartumeus,et al.  ANIMAL SEARCH STRATEGIES: A QUANTITATIVE RANDOM‐WALK ANALYSIS , 2005 .

[37]  P. Beier,et al.  INFLUENCE OF VEGETATION, TOPOGRAPHY, AND ROADS ON COUGAR MOVEMENT IN SOUTHERN CALIFORNIA , 2005 .

[38]  H. J. Hensbergen,et al.  Climatic factors affecting trapping success of some South African small mammals , 1993 .

[39]  Henri Weimerskirch,et al.  Does Prey Capture Induce Area‐Restricted Search? A Fine‐Scale Study Using GPS in a Marine Predator, the Wandering Albatross , 2007, The American Naturalist.

[40]  Peter I. Corke,et al.  Monitoring Animal Behaviour and Environmental Interactions Using Wireless Sensor Networks, GPS Collars and Satellite Remote Sensing , 2009, Sensors.