Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years

Animals switch between inactive and active states, simultaneously impacting their energy intake, energy expenditure and predation risk, and collectively defining how they engage with environmental variation and trophic interactions. We assess daily activity responses to long‐term variation in temper...

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Published in:Ecology Letters
Main Authors: Studd, E. K., Menzies, A. K., Siracusa, E. R., Dantzer, B., Lane, J. E., McAdam, A. G., Boutin, S., Humphries, M. M.
Format: Article in Journal/Newspaper
Language:unknown
Published: Wiley Periodicals, Inc. 2020
Subjects:
optimal behaviour
Accelerometer
behaviour
decision‐making
energetic gain
hoarding
metabolic ecology
Tamiasciurus hudsonicus
Ecology and Evolutionary Biology
Science
Arctic
Online Access:https://hdl.handle.net/2027.42/154889
https://doi.org/10.1111/ele.13494
id ftumdeepblue:oai:deepblue.lib.umich.edu:2027.42/154889
record_format openpolar
institution Open Polar
collection University of Michigan: Deep Blue
op_collection_id ftumdeepblue
language unknown
topic optimal behaviour
Accelerometer
behaviour
decision‐making
energetic gain
hoarding
metabolic ecology
Tamiasciurus hudsonicus
Ecology and Evolutionary Biology
Science
spellingShingle optimal behaviour
Accelerometer
behaviour
decision‐making
energetic gain
hoarding
metabolic ecology
Tamiasciurus hudsonicus
Ecology and Evolutionary Biology
Science
Studd, E. K.
Menzies, A. K.
Siracusa, E. R.
Dantzer, B.
Lane, J. E.
McAdam, A. G.
Boutin, S.
Humphries, M. M.
Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years
topic_facet optimal behaviour
Accelerometer
behaviour
decision‐making
energetic gain
hoarding
metabolic ecology
Tamiasciurus hudsonicus
Ecology and Evolutionary Biology
Science
description Animals switch between inactive and active states, simultaneously impacting their energy intake, energy expenditure and predation risk, and collectively defining how they engage with environmental variation and trophic interactions. We assess daily activity responses to long‐term variation in temperature, resources and mating opportunities to examine whether individuals choose to be active or inactive according to an optimisation of the relative energetic and reproductive gains each state offers. We show that this simplified behavioural decision approach predicts most activity variation (R2 = 0.83) expressed by free‐ranging red squirrels over 4 years, as quantified through accelerometer recordings (489 deployments; 5066 squirrel‐days). Recognising activity as a determinant of energetic status, the predictability of activity variation aggregated at a daily scale, and the clear signal that behaviour is environmentally forced through optimisation of gain, provides an integrated approach to examine behavioural variation as an intermediary between environmental variation and energetic, life‐history and ecological outcomes.By assessing daily activity responses to long‐term variation in temperature, resources, and mating opportunities, we examine whether individuals choose to be active or inactive according to an optimization of energetic and reproductive gains. This simplified behavioural decision approach predicts most daily activity variation (R2 = 0.83) expressed by free‐ranging red squirrels over four years, as quantified through accelerometer recordings. Here we provide an integrated approach to examine behavioural variation as an intermediary between environmental variation and energetic, life‐history, and ecological outcomes. Peer Reviewed https://deepblue.lib.umich.edu/bitstream/2027.42/154889/1/ele13494_am.pdf https://deepblue.lib.umich.edu/bitstream/2027.42/154889/2/ele13494.pdf
format Article in Journal/Newspaper
author Studd, E. K.
Menzies, A. K.
Siracusa, E. R.
Dantzer, B.
Lane, J. E.
McAdam, A. G.
Boutin, S.
Humphries, M. M.
author_facet Studd, E. K.
Menzies, A. K.
Siracusa, E. R.
Dantzer, B.
Lane, J. E.
McAdam, A. G.
Boutin, S.
Humphries, M. M.
author_sort Studd, E. K.
title Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years
title_short Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years
title_full Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years
title_fullStr Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years
title_full_unstemmed Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years
title_sort optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years
publisher Wiley Periodicals, Inc.
publishDate 2020
url https://hdl.handle.net/2027.42/154889
https://doi.org/10.1111/ele.13494
genre Arctic
genre_facet Arctic
op_relation Studd, E. K.; Menzies, A. K.; Siracusa, E. R.; Dantzer, B.; Lane, J. E.; McAdam, A. G.; Boutin, S.; Humphries, M. M. (2020). "Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years." Ecology Letters 23(5): 841-850.
1461-023X
1461-0248
https://hdl.handle.net/2027.42/154889
doi:10.1111/ele.13494
Ecology Letters
Mathot, K.J. & Dingemanse, N.J. ( 2015 ). Energetics and behavior: unrequited needs and new directions. Trends in Ecology & Evolution, 30, 199 – 206. https://doi.org/10.1016/J.TREE.2015.01.010.
Krebs, C.J., Boonstra, R., Boutin, S., Sinclair, A.R.E., Smith, J.N.M., Scott Gilbert, B. et al. ( 2014 ). Trophic dynamics of the boreal forests of the kluane region. Arctic, 67, 71 – 81, https://doi.org/10.14430/arctic.2012.12‐109.
LaMontagne, J.M., Peters, S. & Boutin, S. ( 2005 ). A visual index for estimating cone production for individual white spruce trees. Canadian Journal of Forest Research, 35, 3020 – 3026. https://doi.org/10.1139/X05‐210.
Lane, J.E., Boutin, S., Gunn, M.R. & Coltman, D.W. ( 2009 ). Sexually selected behaviour: red squirrel males search for reproductive success. Journal of Animal Ecology, 78, 296 – 304. https://doi.org/10.1111/j.1365‐2656.2008.01502.x.
Lane, J.E., Boutin, S., Speakman, J.R. & Humphries, M.M. ( 2010 ). Energetic costs of male reproduction in a scramble competition mating system. Journal of Animal Ecology, 79, 27 – 34. https://doi.org/10.1111/j.1365‐2656.2009.01592.x.
Lescroël, A., Ballard, G., Toniolo, V., Barton, K.J., Wilson, P.R., Lyver, P. et al. ( 2010 ). Working less to gain more: when breeding quality relates to foraging efficiency. Ecology, 91 ( 7 ), 2044 – 2055.
Levitis, D.A., Lidicker, W.Z. & Freund, G. ( 2009 ). Behavioural biologists do not agree on what constitutes behaviour. Animal Behaviour, 78, 103 – 110. https://doi.org/10.1016/J.ANBEHAV.2009.03.018.
Lichti, N.I., Steele, M.A. & Swihart, R.K. ( 2017 ). Seed fate and decision‐making processes in scatter‐hoarding rodents. Biol. Rev, 92, 474 – 504. https://doi.org/10.1111/brv.12240.
Lima, S. & Dill, L. ( 1990 ). Behavioral decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology, 68, 619 – 640.
Lone, K., Mysterud, A., Gobakken, T., Odden, J., Linnell, J. & Loe, L.E. ( 2016 ). Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators. Oikos, 126, 624 – 632. https://doi.org/10.1111/oik.03486.
Luttbeg, B., Rowe, L. & Mangel, M. ( 2003 ). Prey state and experimental design affect relative size of trait‐and density‐mediated indirect effects. Ecology, 84, 1140 – 1150.
Martin, P. & Bateson, G. ( 1993 ). Measuring Behaviour: An Introductory Guide. Cambridge University Press, Cambridge.
McAdam, A.G., Boutin, S., Sykes, A.K. & Humphries, M.M. ( 2007 ). Life histories of female red squirrels and their contributions to population growth and lifetime fitness. Écoscience, 14, 362 – 369. https://doi.org/10.2307/42902046.
McNamara, J.M. & Houston, A.I. ( 1987 ). Starvation and predation as factors limiting population size. Ecology, 68, 1515 – 1519.
Mueller, P. & Diamond, J. ( 2001 ). Metabolic rate and environmental productivity: Well‐provisioned animals evolved to run and idle fast. PNAS, 23, 12550 – 12554.
Murray, I.W. & Smith, F.A. ( 2012 ). Estimating the influence of the thermal environment on activity patterns of the desert woodrat ( Neotoma lepida ) using temperature chronologies. Canadian Journal of Zoology, 90, 1171 – 1180. https://doi.org/10.1139/z2012‐084.
Pauls, R.W. ( 1977 ). Behavioural strategies relevant to the energy economy of the red squirrel (Tamiasciurus hudsonicus). Canadian Journal of Zoology, 56, 1519 – 1525.
Post, D.M., Conners, M.E. & Goldberg, D.S. ( 2000 ). Prey preference by a top predator and the stability of linked food chains. Ecology, 81, 8 – 14. https://doi.org/10.1890/0012‐9658(2000)081[0008:PPBATP]2.0.CO;2.
Pyke, G.H., Pulliam, H.R. & Charnov, E.L. ( 1977 ). Optimal foraging: a selective review of theory and tests. The Quarterly Review of Biology, 52 ( 2 ), 137 – 154.
Real, L. ( 1990 ). Search theory and mate choice. I. Models of single‐sex discrimination. The American Naturalist, 136, 376 – 405.
Rezende, E., Gomes, F., Chappell, M. & Garland, T. Jr ( 2009 ). Running behavior and its energy cost in mice selectively bred for high voluntary locomotor activity. Physiological and Biochemical Zoology2, 82, 662 – 679.
Scholander, P.F., Hock, R., Walters, V. & Irving, L. ( 1950 ). Adaptation to cold in arctic and tropical mammals and birds in relation to body temperature, insulation, and basal metabolic rate. Biological Bulletin, 99, 259 – 271.
Scholander, P.F., Hock, R., Walters, V. & Johnson, F. ( 1950 ). Heat regulation in some arctic and tropical mammals and birds. Biological Bulletin, 99, 237 – 258.
Studd, E.K., Boutin, S., McAdam, A.G. & Humphries, M.M. ( 2016 ). Nest attendance of lactating red squirrels (Tamiasciurus hudsonicus): influences of biological and environmental correlates. Journal of Mammalogy, 97, 806 – 814. https://doi.org/10.1093/jmammal/gyw010.
Studd, E.K., Boudreau, M.R., Majchrzak, Y.N., Menzies, A.K., Peers, M.J.L., Seguin, J.L., et al. ( 2019 ). Use of acceleration and acoustics to classify behavior, generate time budgets, and evaluate responses to moonlight in free‐ranging snowshoe hares. Frontiers in Ecology and Evolution, 7, 154. https://doi.org/10.3389/fevo.2019.00154.
Studd, E.K., Landry‐Cuerrier, M., Menzies, A.K., Boutin, S., McAdam, A.G., Lane, J.E., et al. ( 2019 ). Behavioral classification of low‐frequency acceleration and temperature data from a free‐ranging small mammal. Ecology and Evolution, 9, 619 – 630. https://doi.org/10.1002/ece3.4786.
Tatler, J., Cassey, P. & Prowse, T.A.A. ( 2018 ). High accuracy at low frequency: detailed behavioural classification from accelerometer data. Journal of Experimental Biology, 221. https://doi.org/10.1242/jeb.184085.
Werner, E.E. & Anholt, B.R. ( 1993 ). Ecological consequences of the trade‐off between growth and mortality rates mediated. The American Naturalist, 142, 242 – 272.
Williams, C.M., Henry, H.A.L. & Sinclair, B.J. ( 2015 ). Cold truths: how winter drives responses of terrestrial organisms to climate change. Biological Reviews, 90, 214 – 235.
Williams, C.T., Barnes, B.M. & Buck, C.L. ( 2016 ). Integrating physiology, behavior, and energetics: Biologging in a free‐living arctic hibernator. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 202, 53 – 62. https://doi.org/10.1016/J.CBPA.2016.04.020.
Yodzis, P. & Innes, S. ( 1992 ). Body size and consumer‐resource dynamics. The American Naturalist, 139, 1151 – 1175.
Levin, S.A. ( 1992 ). The problem of pattern and scale in ecology. Ecology, 73, 1943 – 1967.
Altmann, S.A. & Altmann, J. ( 2006 ). The transformation of behaviour field studies. In: Essays in Animal Behaviour: Celebrating 50 Years of Animal Behaviour (eds Lucas, J.R. & Simmons, L.W. ). Elsevier Academic Press, Burlington, MA, pp. 57 – 80.
Archibald, D.W., Fletcher, Q.E., Boutin, S., McAdam, A.G., Speakman, J.R. & Humphries, M.M. ( 2013 ). Sex‐specific hoarding behavior in North American red squirrels (Tamiasciurus hudsonicus). Journal of Mammalogy, 94, 761 – 770. https://doi.org/10.1644/12‐MAMM‐A‐213.1.
Baum, W.M. ( 2013 ). What counts as behavior? The molar multiscale view. The Behavior Analyst, 36, 283 – 293.
Biro, P.A. & Stamps, J.A. ( 2010 ). Do consistent individual differences in metabolic rate promote consistent individual differences in behavior? Trends in Ecology & Evolution, 25, 653 – 659.
Boggs, C. ( 1992 ). Resource allocation: exploring connections between foraging and life history. Functional Ecology, 6, 508 – 518.
Boonstra, R., Dušek, A., Lane, J. & Boutin, S. ( 2017 ). When the ball is in the female’s court: How the scramble‐competition mating system of the North American red squirrel has shaped male physiology and testosterone dynamics. General and Comparative Endocrinology, 252, 162 – 172.
Brown, J.S. ( 1992 ). Patch use under predation risk: I. Models and predictions. Ann. Zool. Fennici, 29, 301 – 309.
Brown, J.S., Laundre, J. & Gurung, M. ( 1999 ). The ecology of fear: Optimal foraging, game theory, and trophic interactions. Journal of Mammalogy, 80, 385 – 399. https://doi.org/10.2307/1383287.
Brown, J.H., Gillooly, J., Allen, A.P., Savage, V.M. & West, G.B. ( 2004 ). Toward a metabolic theory of ecology. Ecology, 85, 1771 – 1789. https://doi.org/10.1890/03‐9000@10.1002/(ISSN)1939‐9170.MACARTHURAWARD.
Careau, V., Thomas, D., Humphries, M.M. & Réale, D. ( 2008 ). Energy metabolism and animal personality. Oikos, 117, 641 – 653. https://doi.org/10.1111/j.2008.0030‐1299.16513.x.
Careau, V., Thomas, D., Pelletier, F., Turki, L., Landry, F., Garant, D. et al. ( 2011 ). Genetic correlation between resting metabolic rate and exploratory behaviour in deer mice (Peromyscus maniculatus). Journal of Evolutionary Biology, 2 ), 2153 – 2163.
Daly, M. ( 1978 ). The cost of mating. The American Natualist, 112, 771 – 774.
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spelling ftumdeepblue:oai:deepblue.lib.umich.edu:2027.42/154889 2023-08-20T04:03:10+02:00 Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years Studd, E. K. Menzies, A. K. Siracusa, E. R. Dantzer, B. Lane, J. E. McAdam, A. G. Boutin, S. Humphries, M. M. 2020-05 application/pdf https://hdl.handle.net/2027.42/154889 https://doi.org/10.1111/ele.13494 unknown Wiley Periodicals, Inc. Elsevier Academic Press Studd, E. K.; Menzies, A. K.; Siracusa, E. R.; Dantzer, B.; Lane, J. E.; McAdam, A. G.; Boutin, S.; Humphries, M. M. (2020). "Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years." Ecology Letters 23(5): 841-850. 1461-023X 1461-0248 https://hdl.handle.net/2027.42/154889 doi:10.1111/ele.13494 Ecology Letters Mathot, K.J. & Dingemanse, N.J. ( 2015 ). Energetics and behavior: unrequited needs and new directions. Trends in Ecology & Evolution, 30, 199 – 206. https://doi.org/10.1016/J.TREE.2015.01.010. Krebs, C.J., Boonstra, R., Boutin, S., Sinclair, A.R.E., Smith, J.N.M., Scott Gilbert, B. et al. ( 2014 ). Trophic dynamics of the boreal forests of the kluane region. Arctic, 67, 71 – 81, https://doi.org/10.14430/arctic.2012.12‐109. LaMontagne, J.M., Peters, S. & Boutin, S. ( 2005 ). A visual index for estimating cone production for individual white spruce trees. Canadian Journal of Forest Research, 35, 3020 – 3026. https://doi.org/10.1139/X05‐210. Lane, J.E., Boutin, S., Gunn, M.R. & Coltman, D.W. ( 2009 ). Sexually selected behaviour: red squirrel males search for reproductive success. Journal of Animal Ecology, 78, 296 – 304. https://doi.org/10.1111/j.1365‐2656.2008.01502.x. Lane, J.E., Boutin, S., Speakman, J.R. & Humphries, M.M. ( 2010 ). Energetic costs of male reproduction in a scramble competition mating system. Journal of Animal Ecology, 79, 27 – 34. https://doi.org/10.1111/j.1365‐2656.2009.01592.x. Lescroël, A., Ballard, G., Toniolo, V., Barton, K.J., Wilson, P.R., Lyver, P. et al. ( 2010 ). Working less to gain more: when breeding quality relates to foraging efficiency. Ecology, 91 ( 7 ), 2044 – 2055. Levitis, D.A., Lidicker, W.Z. & Freund, G. ( 2009 ). Behavioural biologists do not agree on what constitutes behaviour. Animal Behaviour, 78, 103 – 110. https://doi.org/10.1016/J.ANBEHAV.2009.03.018. Lichti, N.I., Steele, M.A. & Swihart, R.K. ( 2017 ). Seed fate and decision‐making processes in scatter‐hoarding rodents. Biol. Rev, 92, 474 – 504. https://doi.org/10.1111/brv.12240. Lima, S. & Dill, L. ( 1990 ). Behavioral decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology, 68, 619 – 640. Lone, K., Mysterud, A., Gobakken, T., Odden, J., Linnell, J. & Loe, L.E. ( 2016 ). Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators. Oikos, 126, 624 – 632. https://doi.org/10.1111/oik.03486. Luttbeg, B., Rowe, L. & Mangel, M. ( 2003 ). Prey state and experimental design affect relative size of trait‐and density‐mediated indirect effects. Ecology, 84, 1140 – 1150. Martin, P. & Bateson, G. ( 1993 ). Measuring Behaviour: An Introductory Guide. Cambridge University Press, Cambridge. McAdam, A.G., Boutin, S., Sykes, A.K. & Humphries, M.M. ( 2007 ). Life histories of female red squirrels and their contributions to population growth and lifetime fitness. Écoscience, 14, 362 – 369. https://doi.org/10.2307/42902046. McNamara, J.M. & Houston, A.I. ( 1987 ). Starvation and predation as factors limiting population size. Ecology, 68, 1515 – 1519. Mueller, P. & Diamond, J. ( 2001 ). Metabolic rate and environmental productivity: Well‐provisioned animals evolved to run and idle fast. PNAS, 23, 12550 – 12554. Murray, I.W. & Smith, F.A. ( 2012 ). Estimating the influence of the thermal environment on activity patterns of the desert woodrat ( Neotoma lepida ) using temperature chronologies. Canadian Journal of Zoology, 90, 1171 – 1180. https://doi.org/10.1139/z2012‐084. Pauls, R.W. ( 1977 ). Behavioural strategies relevant to the energy economy of the red squirrel (Tamiasciurus hudsonicus). Canadian Journal of Zoology, 56, 1519 – 1525. Post, D.M., Conners, M.E. & Goldberg, D.S. ( 2000 ). Prey preference by a top predator and the stability of linked food chains. Ecology, 81, 8 – 14. https://doi.org/10.1890/0012‐9658(2000)081[0008:PPBATP]2.0.CO;2. Pyke, G.H., Pulliam, H.R. & Charnov, E.L. ( 1977 ). Optimal foraging: a selective review of theory and tests. The Quarterly Review of Biology, 52 ( 2 ), 137 – 154. Real, L. ( 1990 ). Search theory and mate choice. I. Models of single‐sex discrimination. The American Naturalist, 136, 376 – 405. Rezende, E., Gomes, F., Chappell, M. & Garland, T. Jr ( 2009 ). Running behavior and its energy cost in mice selectively bred for high voluntary locomotor activity. Physiological and Biochemical Zoology2, 82, 662 – 679. Scholander, P.F., Hock, R., Walters, V. & Irving, L. ( 1950 ). Adaptation to cold in arctic and tropical mammals and birds in relation to body temperature, insulation, and basal metabolic rate. Biological Bulletin, 99, 259 – 271. Scholander, P.F., Hock, R., Walters, V. & Johnson, F. ( 1950 ). Heat regulation in some arctic and tropical mammals and birds. Biological Bulletin, 99, 237 – 258. Studd, E.K., Boutin, S., McAdam, A.G. & Humphries, M.M. ( 2016 ). Nest attendance of lactating red squirrels (Tamiasciurus hudsonicus): influences of biological and environmental correlates. Journal of Mammalogy, 97, 806 – 814. https://doi.org/10.1093/jmammal/gyw010. Studd, E.K., Boudreau, M.R., Majchrzak, Y.N., Menzies, A.K., Peers, M.J.L., Seguin, J.L., et al. ( 2019 ). Use of acceleration and acoustics to classify behavior, generate time budgets, and evaluate responses to moonlight in free‐ranging snowshoe hares. Frontiers in Ecology and Evolution, 7, 154. https://doi.org/10.3389/fevo.2019.00154. Studd, E.K., Landry‐Cuerrier, M., Menzies, A.K., Boutin, S., McAdam, A.G., Lane, J.E., et al. ( 2019 ). Behavioral classification of low‐frequency acceleration and temperature data from a free‐ranging small mammal. Ecology and Evolution, 9, 619 – 630. https://doi.org/10.1002/ece3.4786. Tatler, J., Cassey, P. & Prowse, T.A.A. ( 2018 ). High accuracy at low frequency: detailed behavioural classification from accelerometer data. Journal of Experimental Biology, 221. https://doi.org/10.1242/jeb.184085. Werner, E.E. & Anholt, B.R. ( 1993 ). Ecological consequences of the trade‐off between growth and mortality rates mediated. The American Naturalist, 142, 242 – 272. Williams, C.M., Henry, H.A.L. & Sinclair, B.J. ( 2015 ). Cold truths: how winter drives responses of terrestrial organisms to climate change. Biological Reviews, 90, 214 – 235. Williams, C.T., Barnes, B.M. & Buck, C.L. ( 2016 ). Integrating physiology, behavior, and energetics: Biologging in a free‐living arctic hibernator. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 202, 53 – 62. https://doi.org/10.1016/J.CBPA.2016.04.020. Yodzis, P. & Innes, S. ( 1992 ). Body size and consumer‐resource dynamics. The American Naturalist, 139, 1151 – 1175. Levin, S.A. ( 1992 ). The problem of pattern and scale in ecology. Ecology, 73, 1943 – 1967. Altmann, S.A. & Altmann, J. ( 2006 ). The transformation of behaviour field studies. In: Essays in Animal Behaviour: Celebrating 50 Years of Animal Behaviour (eds Lucas, J.R. & Simmons, L.W. ). Elsevier Academic Press, Burlington, MA, pp. 57 – 80. Archibald, D.W., Fletcher, Q.E., Boutin, S., McAdam, A.G., Speakman, J.R. & Humphries, M.M. ( 2013 ). Sex‐specific hoarding behavior in North American red squirrels (Tamiasciurus hudsonicus). Journal of Mammalogy, 94, 761 – 770. https://doi.org/10.1644/12‐MAMM‐A‐213.1. Baum, W.M. ( 2013 ). What counts as behavior? The molar multiscale view. The Behavior Analyst, 36, 283 – 293. Biro, P.A. & Stamps, J.A. ( 2010 ). Do consistent individual differences in metabolic rate promote consistent individual differences in behavior? Trends in Ecology & Evolution, 25, 653 – 659. Boggs, C. ( 1992 ). Resource allocation: exploring connections between foraging and life history. Functional Ecology, 6, 508 – 518. Boonstra, R., Dušek, A., Lane, J. & Boutin, S. ( 2017 ). When the ball is in the female’s court: How the scramble‐competition mating system of the North American red squirrel has shaped male physiology and testosterone dynamics. General and Comparative Endocrinology, 252, 162 – 172. Brown, J.S. ( 1992 ). Patch use under predation risk: I. Models and predictions. Ann. Zool. Fennici, 29, 301 – 309. Brown, J.S., Laundre, J. & Gurung, M. ( 1999 ). The ecology of fear: Optimal foraging, game theory, and trophic interactions. Journal of Mammalogy, 80, 385 – 399. https://doi.org/10.2307/1383287. Brown, J.H., Gillooly, J., Allen, A.P., Savage, V.M. & West, G.B. ( 2004 ). Toward a metabolic theory of ecology. Ecology, 85, 1771 – 1789. https://doi.org/10.1890/03‐9000@10.1002/(ISSN)1939‐9170.MACARTHURAWARD. Careau, V., Thomas, D., Humphries, M.M. & Réale, D. ( 2008 ). Energy metabolism and animal personality. Oikos, 117, 641 – 653. https://doi.org/10.1111/j.2008.0030‐1299.16513.x. Careau, V., Thomas, D., Pelletier, F., Turki, L., Landry, F., Garant, D. et al. ( 2011 ). Genetic correlation between resting metabolic rate and exploratory behaviour in deer mice (Peromyscus maniculatus). Journal of Evolutionary Biology, 2 ), 2153 – 2163. Daly, M. ( 1978 ). The cost of mating. The American Natualist, 112, 771 – 774. IndexNoFollow optimal behaviour Accelerometer behaviour decision‐making energetic gain hoarding metabolic ecology Tamiasciurus hudsonicus Ecology and Evolutionary Biology Science Article 2020 ftumdeepblue https://doi.org/10.1111/ele.1349410.14430/arctic.2012.12‐10910.3389/fevo.2019.0015410.1002/ece3.478610.5067/MODIS/MOD13A1.00610.1007/s00265‐003‐0637‐910.1111/j.1095‐8649.2003.00214.x10.1111/j.1461‐0248.2005.00839.x 2023-07-31T20:47:31Z Animals switch between inactive and active states, simultaneously impacting their energy intake, energy expenditure and predation risk, and collectively defining how they engage with environmental variation and trophic interactions. We assess daily activity responses to long‐term variation in temperature, resources and mating opportunities to examine whether individuals choose to be active or inactive according to an optimisation of the relative energetic and reproductive gains each state offers. We show that this simplified behavioural decision approach predicts most activity variation (R2 = 0.83) expressed by free‐ranging red squirrels over 4 years, as quantified through accelerometer recordings (489 deployments; 5066 squirrel‐days). Recognising activity as a determinant of energetic status, the predictability of activity variation aggregated at a daily scale, and the clear signal that behaviour is environmentally forced through optimisation of gain, provides an integrated approach to examine behavioural variation as an intermediary between environmental variation and energetic, life‐history and ecological outcomes.By assessing daily activity responses to long‐term variation in temperature, resources, and mating opportunities, we examine whether individuals choose to be active or inactive according to an optimization of energetic and reproductive gains. This simplified behavioural decision approach predicts most daily activity variation (R2 = 0.83) expressed by free‐ranging red squirrels over four years, as quantified through accelerometer recordings. Here we provide an integrated approach to examine behavioural variation as an intermediary between environmental variation and energetic, life‐history, and ecological outcomes. Peer Reviewed https://deepblue.lib.umich.edu/bitstream/2027.42/154889/1/ele13494_am.pdf https://deepblue.lib.umich.edu/bitstream/2027.42/154889/2/ele13494.pdf Article in Journal/Newspaper Arctic University of Michigan: Deep Blue Ecology Letters 23 5 841 850

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