Running on empty: recharge dynamics from animal movement data.

Vital rates such as survival and recruitment have always been important in the study of population and community ecology. At the individual level, physiological processes such as energetics are critical in understanding biomechanics and movement ecology and also scale up to influence food webs and trophic cascades. Although vital rates and population-level characteristics are tied with individual-level animal movement, most statistical models for telemetry data are not equipped to provide inference about these relationships because they lack the explicit, mechanistic connection to physiological dynamics. We present a framework for modelling telemetry data that explicitly includes an aggregated physiological process associated with decision making and movement in heterogeneous environments. Our framework accommodates a wide range of movement and physiological process specifications. We illustrate a specific model formulation in continuous-time to provide direct inference about gains and losses associated with physiological processes based on movement. Our approach can also be extended to accommodate auxiliary data when available. We demonstrate our model to infer mountain lion (Puma concolor; in Colorado, USA) and African buffalo (Syncerus caffer; in Kruger National Park, South Africa) recharge dynamics.

[1]  J. Fieberg,et al.  Establishing the link between habitat selection and animal population dynamics , 2015 .

[2]  Sonny S. Bleicher,et al.  The landscape of fear conceptual framework: definition and review of current applications and misuses , 2017, PeerJ.

[3]  Jonathan R. Potts,et al.  Energy benefits and emergent space use patterns of an empirically parameterized model of memory‐based patch selection , 2017 .

[4]  E. Revilla,et al.  A movement ecology paradigm for unifying organismal movement research , 2008, Proceedings of the National Academy of Sciences.

[5]  Wayne M Getz,et al.  Methods for assessing movement path recursion with application to African buffalo in South Africa. , 2009, Ecology.

[6]  A. Sinclair,et al.  The African Buffalo: A Study of Resource Limitation of Populations , 1978 .

[7]  H. Prins,et al.  Ecology and Behaviour of the African Buffalo , 1996, Chapman & Hall Wildlife Ecology and Behaviour Series.

[8]  Sadie J. Ryan,et al.  African buffalo Syncerus caffer (Sparrman, 1779) , 2014 .

[9]  Jacob S. Ivan,et al.  Hierarchical animal movement models for population‐level inference , 2016, 1606.09585.

[10]  J. Forester,et al.  Incorporating animal spatial memory in step selection functions. , 2016, The Journal of animal ecology.

[11]  Murray M Humphries,et al.  Heat for nothing or activity for free? Evidence and implications of activity-thermoregulatory heat substitution. , 2011, Integrative and comparative biology.

[12]  Eric M. Gese,et al.  Resource selection by cougars: Influence of behavioral state and season , 2016 .

[13]  Mevin B. Hooten,et al.  Animal movement models for migratory individuals and groups , 2017, 1708.09472.

[14]  Mevin B. Hooten,et al.  A guide to Bayesian model selection for ecologists , 2015 .

[15]  Jan Beran,et al.  Statistics for long-memory processes , 1994 .

[16]  V. Savage,et al.  A quantitative, theoretical framework for understanding mammalian sleep , 2007, Proceedings of the National Academy of Sciences.

[17]  J. D. Toit,et al.  Large predators and their prey in a southern African savanna: a predator's size determines its prey size range , 2004 .

[18]  Romuald N. Lipcius,et al.  Shelter Selection by Spiny Lobster Under Variable Predation Risk, Social Conditions, and Shelter Size , 1992 .

[19]  Jacob S. Ivan,et al.  A functional model for characterizing long‐distance movement behaviour , 2016 .

[20]  E J Milner-Gulland,et al.  The demographic consequences of the cost of reproduction in ungulates. , 2008, Ecology.

[21]  Roland Langrock,et al.  Flexible and practical modeling of animal telemetry data: hidden Markov models and extensions. , 2012, Ecology.

[22]  P. Turchin Quantitative analysis of movement : measuring and modeling population redistribution in animals and plants , 1998 .

[23]  R. Kays,et al.  Terrestrial animal tracking as an eye on life and planet , 2015, Science.

[24]  J. Fryxell,et al.  Are there general mechanisms of animal home range behaviour? A review and prospects for future research. , 2008, Ecology letters.

[25]  Jacob S. Ivan,et al.  Time-varying predatory behavior is primary predictor of fine-scale movement of wildland-urban cougars , 2018, Movement Ecology.

[26]  D. Brillinger Modeling Spatial Trajectories , 2010 .

[27]  Chloe Bracis,et al.  Memory, not just perception, plays an important role in terrestrial mammalian migration , 2017, Proceedings of the Royal Society B: Biological Sciences.

[28]  Jonathan A Green,et al.  The heart rate method for estimating metabolic rate: review and recommendations. , 2011, Comparative biochemistry and physiology. Part A, Molecular & integrative physiology.

[29]  Kenneth H. Pollock,et al.  Modeling capture, recapture, and removal statistics for estimation of demographic parameters for fish and wildlife populations : Past, present, and future , 1991 .

[30]  Christopher C Wilmers,et al.  The golden age of bio-logging: how animal-borne sensors are advancing the frontiers of ecology. , 2015, Ecology.

[31]  F. Cagnacci,et al.  Animal ecology meets GPS-based radiotelemetry: a perfect storm of opportunities and challenges , 2010, Philosophical Transactions of the Royal Society B: Biological Sciences.

[32]  Mevin B. Hooten,et al.  Process convolution approaches for modeling interacting trajectories , 2017, 1703.02112.

[33]  N. T. Hobbs,et al.  Mechanisms of Foraging in Mammalian Herbivores: New Models of Functional Response , 1992, The American Naturalist.

[34]  R J Full,et al.  Effect of variation in form on the cost of terrestrial locomotion. , 1990, The Journal of experimental biology.

[35]  W. Karasov,et al.  Daily Energy Expenditure and the Cost of Activity in Mammals , 1992 .

[36]  Wayne M. Getz,et al.  LoCoH: Nonparameteric Kernel Methods for Constructing Home Ranges and Utilization Distributions , 2007, PloS one.

[37]  Michael Schaub,et al.  Integrated population models: a novel analysis framework for deeper insights into population dynamics , 2011, Journal of Ornithology.

[38]  Mevin B. Hooten,et al.  Bayesian Models: A Statistical Primer for Ecologists , 2015 .

[39]  R. McNeill Alexander,et al.  Principles of Animal Locomotion , 2002 .

[40]  Mevin B. Hooten,et al.  Continuous-time discrete-space models for animal movement , 2012, 1211.1992.

[41]  Juan M. Morales,et al.  EXTRACTING MORE OUT OF RELOCATION DATA: BUILDING MOVEMENT MODELS AS MIXTURES OF RANDOM WALKS , 2004 .

[42]  James E. Dunn,et al.  Analysis of Radio Telemetry Data in Studies of Home Range , 1977 .

[43]  Adrian F. M. Smith,et al.  Sampling-Based Approaches to Calculating Marginal Densities , 1990 .

[44]  Terrie M. Williams,et al.  Energetics‐informed behavioral states reveal the drive to kill in African leopards , 2017 .

[45]  Ran Nathan,et al.  An emerging movement ecology paradigm , 2008, Proceedings of the National Academy of Sciences.

[46]  WAYNE M. GETZ,et al.  Range and Habitat Selection of African Buffalo in South Africa , 2006 .

[47]  Paul Cross,et al.  Habitat quality and heterogeneity influence distribution and behavior in African buffalo (Syncerus caffer). , 2008, Ecology.

[48]  N. Heglund,et al.  Energetics and mechanics of terrestrial locomotion. I. Metabolic energy consumption as a function of speed and body size in birds and mammals. , 1982, The Journal of experimental biology.

[49]  N. Zuntz,et al.  Ueber den Stoffverbrauch des Hundes bei Muskelarbeit , 1897, Archiv für die gesamte Physiologie des Menschen und der Tiere.

[50]  Herbert H. T. Prins,et al.  Ecology and behaviour of the African buffalo : social inequality and decision making , 1996 .

[51]  Mevin B. Hooten,et al.  Dynamic social networks based on movement , 2015, 1512.07607.

[52]  Brett T. McClintock,et al.  A general discrete‐time modeling framework for animal movement using multistate random walks , 2012 .

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

[54]  Mevin B. Hooten,et al.  Agent-Based Inference for Animal Movement and Selection , 2010 .

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

[56]  Jennifer A. Hoeting,et al.  Handbook of Environmental and Ecological Statistics , 2019 .

[57]  Rory P. Wilson,et al.  DETERMINATION OF MOVEMENTS OF AFRICAN PENGUINS SPHENISCUS DEMERSUS USING A COMPASS SYSTEM: DEAD RECKONING MAY BE AN ALTERNATIVE TO TELEMETRY , 1991 .

[59]  Ian D. Jonsen,et al.  ROBUST STATE-SPACE MODELING OF ANIMAL MOVEMENT DATA , 2005 .

[60]  Alan A. Ager,et al.  Analyzing animal movement patterns using potential functions , 2013 .

[61]  S. Foster,et al.  The value of ecosystem services provided by the U.S. National Wildlife Refuge System in the contiguous U.S. , 2008 .

[62]  L. Zanette,et al.  Fear affects parental care, which predicts juvenile survival and exacerbates the total cost of fear on demography. , 2018, Ecology.

[63]  Wayne M. Getz,et al.  SURFACE‐WATER CONSTRAINTS ON HERBIVORE FORAGING IN THE KRUGER NATIONAL PARK, SOUTH AFRICA , 2003 .

[64]  Cang Hui,et al.  Long‐term rainfall regression surfaces for the Kruger National Park, South Africa: a spatio‐temporal review of patterns from 1981 to 2015 , 2018 .

[65]  Chloe Bracis,et al.  Memory Effects on Movement Behavior in Animal Foraging , 2015, PloS one.

[66]  Lucas N Joppa,et al.  Understanding movement data and movement processes: current and emerging directions. , 2008, Ecology letters.

[67]  Y. Ropert‐Coudert,et al.  Accelerometry predicts daily energy expenditure in a bird with high activity levels , 2013, Biology Letters.

[68]  Simon Benhamou,et al.  Spatial memory and animal movement. , 2013, Ecology letters.

[69]  Kevin A. Blecha,et al.  Risk-reward tradeoffs in the foraging strategy of cougar (Puma concolor): Prey distribution, anthropogenic development, and patch selection , 2015 .

[70]  Patrick J Butler,et al.  Biotelemetry: a mechanistic approach to ecology. , 2004, Trends in ecology & evolution.

[71]  P. Colgan,et al.  Animal Motivation , 1989, Chapman and Hall Animal Behaviour Series.

[72]  A. Mysterud,et al.  How many routes lead to migration? Comparison of methods to assess and characterize migratory movements. , 2016, The Journal of animal ecology.

[73]  N. Heglund,et al.  Energetics and mechanics of terrestrial locomotion. , 1982, Annual review of physiology.

[74]  Roland Langrock,et al.  Using mixed hidden Markov models to examine behavioral states in a cooperatively breeding bird , 2015 .

[75]  Mark S. Boyce,et al.  Evaluating Global Positioning System Telemetry Techniques for Estimating Cougar Predation Parameters , 2009 .

[76]  David R. Anderson,et al.  Modeling Survival and Testing Biological Hypotheses Using Marked Animals: A Unified Approach with Case Studies , 1992 .

[77]  Dennis L. Murray,et al.  Assessing differential prey selection patterns between two sympatric large carnivores , 2003 .

[78]  Simon Chamaillé-Jammes,et al.  Partial migration links local surface-water management to large-scale elephant conservation in the world's largest transfrontier conservation area , 2017 .

[79]  A. Houston,et al.  Models of adaptive behaviour , 1999 .

[80]  Mevin B. Hooten,et al.  Basis Function Models for Animal Movement , 2016, 1601.05408.

[81]  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.

[82]  Jacqueline L. Frair,et al.  Building the bridge between animal movement and population dynamics , 2010, Philosophical Transactions of the Royal Society B: Biological Sciences.

[83]  Rob Deardon,et al.  An empirically parameterized individual based model of animal movement, perception, and memory , 2013 .

[84]  J. Laundré,et al.  Wolves, elk, and bison: reestablishing the "landscape of fear" in Yellowstone National Park, U.S.A. , 2001 .

[85]  L. Halsey Terrestrial movement energetics: current knowledge and its application to the optimising animal , 2016, Journal of Experimental Biology.

[86]  James S. Clark,et al.  Using Hierarchical Bayes to Understand Movement, Health, and Survival in the Endangered North Atlantic Right Whale , 2013, PloS one.

[87]  T. Faniran Numerical Solution of Stochastic Differential Equations , 2015 .

[88]  Aristotle Aristotle,et al.  Aristotle's De Motu Animalium: Text with Translation, Commentary, and Interpretive Essays , 1978 .

[89]  Gabriel Hugh Elkaim,et al.  Instantaneous energetics of puma kills reveal advantage of felid sneak attacks , 2014, Science.

[90]  Roland Langrock,et al.  Analysis of animal accelerometer data using hidden Markov models , 2016, 1602.06466.

[91]  Daniel Fortin,et al.  Adaptive models for large herbivore movements in heterogeneous landscapes , 2005, Landscape Ecology.

[92]  Greg Stuart-Hill,et al.  Home on the Range: Factors Explaining Partial Migration of African Buffalo in a Tropical Environment , 2012, PloS one.

[93]  Roland Kays,et al.  Observing the unwatchable through acceleration logging of animal behavior , 2013, Animal Biotelemetry.

[94]  Kim Whoriskey,et al.  A hidden Markov movement model for rapidly identifying behavioral states from animal tracks , 2016, Ecology and evolution.

[95]  Brett T. McClintock,et al.  Animal Movement: Statistical Models for Telemetry Data , 2017 .