Temperature-mediated habitat use and selection by a heat-sensitive northern ungulate

The behavioural response of animals to unfavourable climatic conditions has received increased attention recently. While many studies have examined the behavioural responses of endotherms to cold temperatures, thermoregulatory behaviour may also occur in response to heat stress. We evaluated whether a heat-sensitive northern ungulate, the moose, Alces alces, showed thermoregulatory behaviour in response to ambient temperature in two populations in southern Norway. We quantified the seasonal habitat use of GPS-collared adult females, as well as fine-scale habitat selection patterns, in relation to time of day and critical temperature thresholds thought to induce heat stress. We also assessed whether temperature-associated changes in spatial behaviour led to a shift in the availability of thermal cover and forage at the chosen location. Frequent exposure to temperatures above critical thresholds occurred in both summer and winter and in both study areas. Moose responded by seeking thermal shelter in mature coniferous forest and avoiding open habitat types, leading to a decrease in local forage availability in summer but not in winter. Differences in habitat choice in response to temperature were most pronounced at twilight. We found that fine-scale habitat selection analyses, using step-selection functions, more effectively revealed thermoregulatory behaviour in both seasons and populations than did habitat use. This is because habitat selection analyses are better able to identify limiting factors operating at different spatiotemporal scales than is habitat use. Our results clearly show that ambient temperature affected fine-scale behavioural decisions of moose with consequences for forage accessibility, especially during summer. As the climate changes, the limiting effect of ambient temperature (cold and high) on animal behaviour is likely to increase, potentially influencing individual fitness and population dynamics. 2012 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

[1]  Marco Apollonio,et al.  Seasonal variations of spatial behaviour in female Alpine ibex (Capra ibex ibex) in relation to climatic conditions and age , 2004 .

[2]  Mark S Boyce,et al.  Lifetime reproductive success and density-dependent, multi-variable resource selection , 2006, Proceedings of the Royal Society B: Biological Sciences.

[3]  V. Savage,et al.  A Framework for Elucidating the Temperature Dependence of Fitness , 2011, The American Naturalist.

[4]  M. Boyce,et al.  WOLVES INFLUENCE ELK MOVEMENTS: BEHAVIOR SHAPES A TROPHIC CASCADE IN YELLOWSTONE NATIONAL PARK , 2005 .

[5]  M. Boyce,et al.  Evaluating resource selection functions , 2002 .

[6]  O. Schmitz.  Thermal constraints and optimization of winter feeding and habitat choice in white-tailed deer , 1991 .

[7]  A. Mysterud,et al.  Icing events trigger range displacement in a high-arctic ungulate. , 2010, Ecology.

[8]  Göran Ericsson,et al.  Risk of capture-related mortality in large free-ranging mammals: experiences from Scandinavia , 2006 .

[9]  F. Bunnell,et al.  Forest cover selection and activity of cow moose in summer , 1995 .

[10]  Atle Mysterud,et al.  How does local weather predict red deer home range size at different temporal scales? , 2010, The Journal of animal ecology.

[11]  J. Gaillard,et al.  Determinants of seasonal variation in activity patterns of mouflon , 2008 .

[12]  E. Solberg,et al.  Differential forage use makes carrying capacity equivocal on ranges of Scandinavian moose (Alces alces) , 2010 .

[13]  O. Liberg,et al.  Activity patterns of predator and prey: a simultaneous study of GPS-collared wolves and moose , 2011, Animal Behaviour.

[14]  Francis K C Hui,et al.  The arcsine is asinine: the analysis of proportions in ecology. , 2011, Ecology.

[15]  A. Mysterud,et al.  Climate change and implications for the future distribution and management of ungulates in Europe. , 2010 .

[16]  S. Côté,et al.  Trade-offs in activity budget in an alpine ungulate: contrasting lactating and nonlactating females , 2008, Animal Behaviour.

[17]  Evelyn H. Merrill,et al.  Thermal constraints on use of cover types and activity time of elk , 1991 .

[18]  S. Wood.  Generalized Additive Models: An Introduction with R , 2006 .

[19]  Wallace P. Erickson,et al.  Resource Selection by Animals , 2002, Springer Netherlands.

[20]  R. Macarthur,et al.  On Optimal Use of a Patchy Environment , 1966, The American Naturalist.

[21]  A. Mysterud,et al.  Activity pattern of arctic reindeer in a predator-free environment: no need to keep a daily rhythm , 2007, Oecologia.

[22]  J. Kie,et al.  THERMAL LANDSCAPES AND RESOURCE SELECTION BY BLACK-TAILED DEER: IMPLICATIONS FOR LARGE HERBIVORES , 2009 .

[23]  Larissa Conradt,et al.  Sex differences in weather sensitivity can cause habitat segregation: red deer as an example , 2000, Animal Behaviour.

[24]  Floris M. van Beest,et al.  Comparative Space Use and Habitat Selection of Moose Around Feeding Stations , 2010 .

[25]  J. Gaillard,et al.  Daily responses of mouflon (Ovis gmelini musimon × Ovis sp.) activity to summer climatic conditions , 2011 .

[26]  O. Hjeljord,et al.  Range-body mass interactions of a northern ungulate – a test of hypothesis , 1999, Oecologia.

[27]  Mark S. Boyce,et al.  Scale for resource selection functions , 2006 .

[28]  P. E. Hertz,et al.  Thermoregulation in a Lacertid Lizard: The Relative Contributions of Distinct Behavioral Mechanisms , 1996 .

[29]  T. Fuller,et al.  Pathogens, Nutritional Deficiency, and Climate Influences on a Declining Moose Population , 2006 .

[30]  M. Lenarz,et al.  Temperature Mediated Moose Survival in Northeastern Minnesota , 2009 .

[31]  R. Hudson,et al.  Behavioral and thermoregulatory responses of moose to high ambient temperatures and insect harassment in aspen-dominated forests , 1990 .

[32]  W. Porter,et al.  Model of Japanese serow (Capricornis crispus) energetics predicts distribution on Honshu, Japan. , 2007, Ecological applications : a publication of the Ecological Society of America.

[33]  A. Houston,et al.  State-dependent life histories , 1996, Nature.

[34]  R. Bowyer,et al.  HABITAT SELECTION BY MULE DEER: FORAGE BENEFITS OR RISK OF PREDATION? , 2004 .

[35]  M. Boyce,et al.  Elk winter foraging at fine scale in Yellowstone National Park , 2005, Oecologia.

[36]  C. Thomas,et al.  The influence of thermal ecology on the distribution of three nymphalid butterflies , 2002 .

[37]  C. Parmesan.  Ecological and Evolutionary Responses to Recent Climate Change , 2006 .

[38]  Floris M. van Beest,et al.  Forage quantity, quality and depletion as scale-dependent mechanisms driving habitat selection of a large browsing herbivore. , 2010, The Journal of animal ecology.

[39]  Ivar Herfindal,et al.  Screening Global Positioning System Location Data for Errors Using Animal Movement Characteristics , 2010 .

[40]  J. Kohler,et al.  Climate, icing, and wild arctic reindeer: past relationships and future prospects. , 2011, Ecology.

[41]  J. Gaillard,et al.  Maternal and individual effects in selection of bed sites and their consequences for fawn survival at different spatial scales , 2009, Oecologia.

[42]  G. Ruxton,et al.  Why do vultures have bald heads? The role of postural adjustment and bare skin areas in thermoregulation , 2008 .

[43]  G. Belovsky.  Optimal activity times and habitat choice of moose , 1981, Oecologia.

[44]  M. Hebblewhite,et al.  Trade-offs between predation risk and forage differ between migrant strategies in a migratory ungulate. , 2009, Ecology.

[45]  Mark Hebblewhite,et al.  A MULTI-SCALE TEST OF THE FORAGE MATURATION HYPOTHESIS IN A PARTIALLY MIGRATORY UNGULATE POPULATION , 2008 .

[46]  M. Rosenzweig,et al.  A Theory of Habitat Selection , 1981 .

[47]  K. Parker,et al.  Estimates of Critical Thermal Environments for Mule Deer , 1990 .

[48]  R. Courtois,et al.  Behavioural responses of moose to thermal conditions in the boreal forest1 , 2004 .

[49]  F. Schwab,et al.  Moose selection of canopy cover types related to operative temperature, forage, and snow depth , 1991 .

[50]  O. Hoegh‐Guldberg,et al.  Ecological responses to recent climate change , 2002, Nature.

[51]  K. Schmidt.  Winter ecology of nonmigratory Alpine red deer , 1993, Oecologia.

[52]  Floris M. van Beest,et al.  What determines variation in home range size across spatiotemporal scales in a large browsing herbivore? , 2011, The Journal of animal ecology.

[53]  Andrew Paul Gutierrez,et al.  Temperature requirements of some aphids and their parasites , 1974 .

[54]  P. Krausman,et al.  Mechanisms of Thermoregulation and Water Balance in Desert Ungulates , 2006 .

[55]  D. Fortin,et al.  An appraisal of the fitness consequences of forest disturbance for wildlife using habitat selection theory , 2010, Oecologia.

[56]  V. V. Ballenberghe,et al.  Activity of Moose during Spring and Summer in Interior Alaska , 1990 .

[57]  J. A. Schaefer,et al.  Lack of behavioral responses of moose (Alces alces) to high ambient temperatures near the southern periphery of their range , 2010 .

[58]  A Sih,et al.  Optimal behavior: can foragers balance two conflicting demands? , 1980, Science.

[59]  Jenny A. Hodgson,et al.  Predicting insect phenology across space and time , 2011 .

[60]  R. Hudson,et al.  Seasonal energy expenditures and thermoregulatory responses of moose , 1986 .

[61]  D. Bates,et al.  Mixed-Effects Models in S and S-PLUS , 2001 .

[62]  O. Hjeljord,et al.  Choice of feeding sites by moose during summer, the influence of forest structure and plant phenology , 1990 .

[63]  Paul J Rathouz,et al.  Accounting for animal movement in estimation of resource selection functions: sampling and data analysis. , 2009, Ecology.

[64]  Bram Van Moorter,et al.  Moose Alces alces habitat use at multiple temporal scales in a human-altered landscape , 2011 .

[65]  B. Bassano,et al.  Temperature constraints on foraging behaviour of male Alpine ibex (Capra ibex) in summer , 2009, Oecologia.

[66]  O. Hjeljord,et al.  Do continental moose ranges improve during cloudy summers , 1991 .

[67]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[68]  J. Arnemo,et al.  CHEMICAL IMMOBILIZATION OF FREE-RANGING MOOSE , 2003 .

[69]  Chris J. Johnson,et al.  Sensitivity of species-distribution models to error, bias, and model design: An application to resource selection functions for woodland caribou , 2008 .

[70]  O Gimenez,et al.  Assessing the impact of climate variation on survival in vertebrate populations , 2008, Biological reviews of the Cambridge Philosophical Society.

[71]  A. Mysterud,et al.  Cover as a habitat element for temperate ungulates: effects on habitat selection and demography , 1999 .

[72]  J. Kie,et al.  Effects of scale on interpreting life‐history characteristics of ungulates and carnivores , 2006 .