Fine-Scale Tracking of Ambient Temperature and Movement Reveals Shuttling Behavior of Elephants to Water

Movement strategies of animals have been well studied as a function of ecological drivers (e.g., forage selection and avoiding predation) rather than physiological requirements (e.g., thermoregulation). Thermal stress is a major concern for large mammals, especially for savanna elephants (Loxodonta africana), which have amongst the greatest challenge for heat dissipation in hot and arid environments. Therefore, elephants must make decisions about where and how fast to move to reduce thermal stress. We tracked 14 herds of elephant in Kruger National Park (KNP), South Africa, for 2 years, using GPS collars with inbuilt temperature sensors to examine the influence of temperature on movement strategies, particularly when accessing water. We first confirmed that collar-mounted temperature loggers captured hourly variation in relative ambient temperatures across the landscape, and thus, could be used to predict elephant movement strategies at fine spatio-temporal scales. We found that elephants moved slower in more densely wooded areas, but unexpectedly, moved faster at higher temperatures, especially in the wet season compared to the dry season. Notably, this speed of movement was highest when elephants were approaching and leaving water sources. Visits to water showed a periodic shuttling pattern, with a peak return rate of 10-30 hours, wherein elephants were closest to water during the hotter times of the day, and spent longer at water sources in the dry season compared to the wet season. When elephants left water, they showed low fidelity to the same water source, and travelled farther in the dry season than in the wet season. In KNP, where water is easily accessible, and the risk of poaching is low, we found that elephants use short, high-speed bursts of movement to get to water at hotter times of day. This strategy not only provides the benefit of predation risk avoidance, but also allows them to use water to thermoregulate. We demonstrate that ambient temperature is an important predictor of movement and water use across the landscape, with elephants responding facultatively to a “landscape of thermal stress”.

[1]  J. Estes,et al.  The functional significance of movements and positions of the pinnae of the African elephant, Loxodonta africana. , 1971, Journal of mammalogy.

[2]  R. Tinnin Interference or Competition? , 1972, The American Naturalist.

[3]  Why do elephants flap their ears , 1984 .

[4]  Do elephants need to sweat , 1984 .

[5]  D. Altman,et al.  STATISTICAL METHODS FOR ASSESSING AGREEMENT BETWEEN TWO METHODS OF CLINICAL MEASUREMENT , 1986, The Lancet.

[6]  R. Owen-Smith,et al.  Megaherbivores: The Influence of Very Large Body Size on Ecology , 1990 .

[7]  T. Williams Heat transfer in elephants: thermal partitioning based on skin temperature profiles , 1990 .

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

[9]  G. Walsberg Small Mammals in Hot Deserts: Some Generalizations Revisited , 2000 .

[10]  Laurance,et al.  Do edge effects occur over large spatial scales? , 2000, Trends in ecology & evolution.

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

[12]  M. V. Rooyen,et al.  Ecological impact of large herbivores on the woody vegetation at selected watering points on the eastern basaltic soils in the Kruger National Park , 2002 .

[13]  Chris J. Johnson,et al.  Movement parameters of ungulates and scale‐specific responses to the environment , 2002 .

[14]  John M. Fryxell,et al.  Foraging ecology of bison at the landscape and plant community levels: the applicability of energy maximization principles , 2002, Oecologia.

[15]  I. J. Goldberg To drink or not to drink? , 2003, The New England journal of medicine.

[16]  Terrie M. Williams,et al.  The cost of foraging by a marine predator, the Weddell seal Leptonychotes weddellii: pricing by the stroke , 2004, Journal of Experimental Biology.

[17]  R. Guralnick The legacy of past climate and landscape change on species’ current experienced climate and elevation ranges across latitude: a multispecies study utilizing mammals in western North America , 2006 .

[18]  Rina C. Grant,et al.  ELEPHANT (LOXODONTA AFRICANA) DIETS IN KRUGER NATIONAL PARK, SOUTH AFRICA: SPATIAL AND LANDSCAPE DIFFERENCES , 2006 .

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

[20]  J. Daudin,et al.  A Segmentation/Clustering Model for the Analysis of Array CGH Data , 2007, Biometrics.

[21]  Alan Y. Chiang,et al.  Generalized Additive Models: An Introduction With R , 2007, Technometrics.

[22]  S. Maloney,et al.  Validation of a biotelemetric technique, using ambulatory miniature black globe thermometers, to quantify thermoregulatory behaviour in ungulates. , 2007, Journal of experimental zoology. Part A, Ecological genetics and physiology.

[23]  S. Pimm,et al.  Ambient temperature as a determinant of landscape use in the savanna elephant, Loxodonta africana , 2007 .

[24]  P. Myles,et al.  Using the Bland-Altman method to measure agreement with repeated measures. , 2007, British journal of anaesthesia.

[25]  P. Bateman,et al.  Body temperature daily rhythm adaptations in African savanna elephants (Loxodonta africana) , 2007, Physiology & Behavior.

[26]  H. Fritz,et al.  Interference competition and temporal niche shifts: elephants and herbivore communities at waterholes , 2007, Oecologia.

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

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

[29]  H. Fritz,et al.  The role of water abundance, thermoregulation, perceived predation risk and interference competition in water access by African herbivores , 2008 .

[30]  N. Owen‐Smith,et al.  Effects of elephants on ecosystems and biodiversity , 2008 .

[31]  R. Slotow,et al.  Modelling the effect of age-specific mortality on elephant Loxodonta africana populations: can natural mortality provide regulation? , 2008, Oryx.

[32]  Sam M. Ferreira,et al.  Elephant spatial use in wet and dry savannas of southern Africa , 2009 .

[33]  D. Macdonald,et al.  Does the risk of encountering lions influence African herbivore behaviour at waterholes? , 2009, Behavioral Ecology and Sociobiology.

[34]  H. Fritz,et al.  Piosphere contribution to landscape heterogeneity: a case study of remote‐sensed woody cover in a high elephant density landscape , 2009 .

[35]  M. Angilletta Thermal Adaptation: A Theoretical and Empirical Synthesis , 2009 .

[36]  Stuart L. Pimm,et al.  Fences and artificial water affect African savannah elephant movement patterns , 2009 .

[37]  David W. Macdonald,et al.  How key habitat features influence large terrestrial carnivore movements: waterholes and African lions in a semi-arid savanna of north-western Zimbabwe , 2010, Landscape Ecology.

[38]  K. Leggett Daily and hourly movement of male desert-dwelling elephants , 2010 .

[39]  M. Thaker,et al.  Do fences create an edge-effect on the movement patterns of a highly mobile mega-herbivore? , 2010 .

[40]  Sassan Saatchi,et al.  Woody Fractional Cover in Kruger National Park, South Africa: Remote Sensing–Based Maps and Ecological Insights , 2010 .

[41]  H. Schwammer,et al.  Thermal windows on the body surface of African elephants (Loxodonta africana) studied by infrared thermography. , 2010 .

[42]  M. Thaker,et al.  Minimizing predation risk in a landscape of multiple predators: effects on the spatial distribution of African ungulates. , 2011, Ecology.

[43]  S. Wieren,et al.  Energy advantages of orientation to solar radiation in three African ruminants , 2011 .

[44]  T. Ruf,et al.  Taking the heat: thermoregulation in Asian elephants under different climatic conditions , 2011, Journal of Comparative Physiology B.

[45]  S. Maloney,et al.  Activity re-assignment and microclimate selection of free-living Arabian oryx: responses that could minimise the effects of climate change on homeostasis? , 2012, Zoology.

[46]  What Is the Use of Elephant Hair? , 2012, PloS one.

[47]  R. Slotow,et al.  Physiological Stress and Refuge Behavior by African Elephants , 2012, PloS one.

[48]  N. Owen‐Smith,et al.  The costs of drinking: comparative water dependency of sable antelope and zebra , 2012 .

[49]  Vito M. R. Muggeo,et al.  Animal Perception of Seasonal Thresholds: Changes in Elephant Movement in Relation to Rainfall Patterns , 2012, PloS one.

[50]  Bram Van Moorter,et al.  Temperature-mediated habitat use and selection by a heat-sensitive northern ungulate , 2012, Animal Behaviour.

[51]  F. Langevelde,et al.  Body temperature variation of South African antelopes in two climatically contrasting environments , 2012 .

[52]  R. Slotow,et al.  Unravelling complex associations between physiological state and movement of African elephants , 2013 .

[53]  G. S. Bakken,et al.  Heat storage in Asian elephants during submaximal exercise: behavioral regulation of thermoregulatory constraints on activity in endothermic gigantotherms , 2013, Journal of Experimental Biology.

[54]  H. Fritz,et al.  African Elephants Adjust Speed in Response to Surface-Water Constraint on Foraging during the Dry-Season , 2013, PloS one.

[55]  A. Mysterud,et al.  Hunting for fear: innovating management of human–wildlife conflicts , 2013 .

[56]  Daniel Fortin,et al.  Moving to stay in place: behavioral mechanisms for coexistence of African large carnivores. , 2013, Ecology.

[57]  Robin C. Dunkin,et al.  Climate influences thermal balance and water use in African and Asian elephants: physiology can predict drivers of elephant distribution , 2013, Journal of Experimental Biology.

[58]  F. Langevelde,et al.  Larger antelopes are sensitive to heat stress throughout all seasons but smaller antelopes only during summer in an African semi-arid environment , 2013, International Journal of Biometeorology.

[59]  Feng Gao,et al.  Landsat Ecosystem Disturbance Adaptive Processing System LEDAPS algorithm description , 2013 .

[60]  K. K. Zander,et al.  Conserving large carnivores: dollars and fence. , 2013, Ecology letters.

[61]  J. D. Stigter,et al.  Optimization of wildlife management in a large game reserve through waterpoints manipulation: a bio-economic analysis. , 2013, Journal of environmental management.

[62]  N. Owen‐Smith,et al.  Coping with savanna seasonality: comparative daily activity patterns of African ungulates as revealed by GPS telemetry , 2014 .

[63]  S. Maloney,et al.  Adaptation to heat and water shortage in large, arid-zone mammals. , 2014, Physiology.

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

[65]  S. Maloney,et al.  Responses of large mammals to climate change , 2014, Temperature.

[66]  D. Bates,et al.  Fitting Linear Mixed-Effects Models Using lme4 , 2014, 1406.5823.

[67]  A. van Niekerk,et al.  The role of elephant Loxodonta africana pathways as a spatial variable in crop-raiding location , 2014, Oryx.

[68]  David M. Scantlebury,et al.  Flexible energetics of cheetah hunting strategies provide resistance against kleptoparasitism , 2014, Science.

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

[70]  S. Harris,et al.  Habitat Selection by African Buffalo (Syncerus caffer) in Response to Landscape-Level Fluctuations in Water Availability on Two Temporal Scales , 2014, PloS one.

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

[72]  A. Bouskila,et al.  Space-Use Patterns of the Asiatic Wild Ass (Equus hemionus): Complementary Insights from Displacement, Recursion Movement and Habitat Selection Analyses , 2015, PloS one.

[73]  H. J. de Knegt,et al.  Modeling elephant-mediated cascading effects of water point closure. , 2015, Ecological applications : a publication of the Ecological Society of America.

[74]  Hugo Valls Fox To drink or not to drink? The influence of resource availability on elephant foraging and habitat selection in a semi-arid savanna , 2015 .

[75]  F. Langevelde,et al.  Prolonged drought results in starvation of African elephant (Loxodonta africana) , 2016 .

[76]  R. van Aarde,et al.  Coping with heat: behavioural and physiological responses of savanna elephants in their natural habitat , 2016, Conservation physiology.

[77]  Andrew K. Skidmore,et al.  Elephant poaching risk assessed using spatial and non-spatial Bayesian models , 2016 .

[78]  Andrew Purdon Environmental determinants of the movement patterns of elephants in the Kruger National Park , 2016 .

[79]  N. Leader‐Williams,et al.  Global priorities for national carnivore conservation under land use change , 2016, Scientific Reports.

[80]  Michael Dixon,et al.  Google Earth Engine: Planetary-scale geospatial analysis for everyone , 2017 .

[81]  Water provisioning in Kruger National Park alters elephant spatial utilisation patterns , 2017 .

[82]  T. Williams,et al.  Energetics and evasion dynamics of large predators and prey: pumas vs. hounds , 2017, PeerJ.

[83]  F. van Langevelde,et al.  Movement Patterns of African Elephants (Loxodonta africana) in a Semi-arid Savanna Suggest That They Have Information on the Location of Dispersed Water Sources , 2018, Front. Ecol. Evol..

[84]  Simon Benhamou,et al.  Identifying stationary phases in multivariate time-series for highlighting behavioural modes and home range settlements , 2018, bioRxiv.

[85]  Jonathan R. Potts,et al.  Spatial scales of habitat selection decisions: implications for telemetry‐based movement modelling , 2018 .

[86]  Edzer Pebesma,et al.  Simple Features for R: Standardized Support for Spatial Vector Data , 2018, R J..

[87]  S. Maloney,et al.  Revisiting concepts of thermal physiology: Predicting responses of mammals to climate change , 2018, The Journal of animal ecology.

[88]  Bart Kranstauber,et al.  move: Visualizing and analyzing animal track data. R package version 3.1.0 , 2018 .