, Volume 190, Issue 1, pp 11–23 | Cite as

Hourly movement decisions indicate how a large carnivore inhabits developed landscapes

  • Michael J. EvansEmail author
  • Jason E. Hawley
  • Paul W. Rego
  • Tracy A. G. Rittenhouse
Highlighted Student Research


The ecology of wildlife living in proximity to humans often differs from that in more natural places. Animals may perceive anthropogenic features and people as threats, exhibiting avoidance behavior, or may acclimate to human activities. As development expands globally, changes in the ecology of species in response to human phenomena may determine whether animals persist in these changing environments. We hypothesize that American black bears (Ursus americanus) persist within developed areas by effectively avoiding risky landscape features. We test this by quantifying changes in the movements of adult females from a population living within exurban and suburban development. We collected hourly GPS data from 23 individuals from 2012 to 2014 and used step-selection functions to estimate selection for anthropogenic features. Females were more avoidant of roads and highways when with cubs than without and were more responsive to increased traffic volume. As bears occupied greater housing densities, selection for housing increased, while avoidance of roads and responsiveness to traffic increased. Behavioral flexibility allowed bears in highly developed areas to alter selection and avoidance for anthropogenic features seasonally. These findings support the hypothesis that black bears perceive human activity as risky, and effectively avoid these risks while inhabiting developed areas. We document a high amount of individual variation in selection of anthropogenic features within the study population. Our findings suggest that initially, wildlife can successfully inhabit developed landscapes by effectively avoiding human activity. However, variation among individuals provides the capacity for population-level shifts in behavior over time.


Adaptation Functional response Movement behavior Risk avoidance Selection Urban 



We thank L. S. Eggert for help with manuscript preparation. Funding provided by Federal Aid in Wildlife Restoration Act under Project W-49-R “Wildlife Investigations” administered by the Connecticut Department of Energy and Environmental Protection, Wildlife Division.

Author contribution statement

MJE conceived the study; JEH and PWR designed data collection and conducted fieldwork; and MJE designed and conducted analyses and led the writing of the manuscript, with contributions and supervision from TAGR.

Supplementary material

442_2018_4307_MOESM1_ESM.pdf (236 kb)
Changes in model-averaged selection parameter estimates (βω) and standard errors (SEω) of the relationship between anthropogenic variables and log odds of black bear steps estimated by step selection functions using increasing numbers of simulated steps. Different symbols correspond to different individual bears, selected at random from the study population. (PDF 235 kb)


  1. Akaike H (1974) A new look at the statistical model identification. IEEE Trans Autom Control 19:716–723CrossRefGoogle Scholar
  2. Animal Care and Use Committee (1998) Guidelines for the capture, handling, and care of mammals as approved by the American Society of Mammalogists. J Mammal 79:1416–1431CrossRefGoogle Scholar
  3. Anthony LL, Blumstein DT (2000) Integrating behaviour into wildlife conservation: the multiple ways that behaviour can reduce Ne. Biol Conserv 95:303–315CrossRefGoogle Scholar
  4. Baker PJ, Dowding CV, Molony SE, White PC, Harris S (2007) Activity patterns of urban red foxes (vulpes vulpes) reduce the risk of traffic-induced mortality. Behav Ecol 18:716–724CrossRefGoogle Scholar
  5. Barber JR, Crooks KR, Fristrup KM (2010) The costs of chronic noise exposure for terrestrial organisms. Trends Ecol Evol 25:180–189CrossRefGoogle Scholar
  6. Baruch-Mordo S, Wilson KR, Lewis DL, Broderick J, Mao JS, Breck SW (2014) Stochasticity in natural forage production affects use of urban areas by black bears: implications to management of human-bear conflicts. PLoS One 9:e85122CrossRefGoogle Scholar
  7. Bateman P, Fleming P (2012) Big city life: carnivores in urban environments. J Zool 287:1–23CrossRefGoogle Scholar
  8. Bates D, Maechler M, Bolker B, Walker S (2014) lme4: Linear mixed-effects models using Eigen and S4. R Package Version 1:1–12Google Scholar
  9. Beckmann JP, Berger J (2003a) Using black bears to test ideal-free distribution models experimentally. J Mammal 84:594–606CrossRefGoogle Scholar
  10. Beckmann JP, Berger J (2003b) Rapid ecological and behavioral changes in carnivores: the response of black bears to (Ursus americanus) to altered food. J Zool 261:207–212CrossRefGoogle Scholar
  11. Beyer HL (2014) Geospatial modeling environment. Accessed May 2015
  12. Chapron G, Lopez-Bao JV (2014) Conserving carnivores: politics in play. Science 343:1199–1200CrossRefGoogle Scholar
  13. Clark JD, Eastridge R (2006) Growth and sustainability of black bears at white river national wildlife refuge, Arkansas. J Wildl Manag 70:1094–1101CrossRefGoogle Scholar
  14. Coulon A, Morellet N, Goulard M, Cargnelutti B, Angibault J, Hewison AJM (2008) Inferring the effects of landscape structure on roe deer movements using a step selection function. Landsc Ecol 23:603–614CrossRefGoogle Scholar
  15. Dahle B, Swenson JE (2003) Seasonal range size in relation to reproductive strategies in brown bears Ursus arctos. J Anim Ecol 72:660–667CrossRefGoogle Scholar
  16. Dall SR (2004) Behavioural biology: fortune favours bold and shy personalities. Curr Biol 14:470–472CrossRefGoogle Scholar
  17. Darrow PA, Shivik JA (2009) Bold, shy, and persistent: variable coyote response to light and sound stimuli. Appl Anim Behav Sci 116:82–87CrossRefGoogle Scholar
  18. Delibes M, Gaona P, Ferreras P (2001) Effects of an attractive sink leading into maladaptive habitat selection. Am Nat 158:277–285CrossRefGoogle Scholar
  19. DeStefano S, DeGraaf RM (2003) Exploring the ecology of suburban wildlife. Front Ecol Environ 1:95–101CrossRefGoogle Scholar
  20. Ditchkoff SS, Saalfeld ST, Gibson CJ (2006) Animal behavior in urban ecosystems: modifications due to human-induced stress. Urban Ecosyst 9:5–12CrossRefGoogle Scholar
  21. Eiler JH, Wathen WG, Pelton MR (1989) Reproduction in black bears in the southern Appalachian Mountains. J Wildl Manag 53:353–360CrossRefGoogle Scholar
  22. Elfstrom M, Zedrosser A, Stoen O-G, Swenson JE (2012) Ultimate and proximate mechanisms underlying the occurrence of bears close to human settlements: review and management implications. Mamm Rev 44:5–18CrossRefGoogle Scholar
  23. Evans MJ, Hawley JE, Rego PW, Rittenhouse TAG (2017) Black bear recolonization patterns on human-dominated landscapes vary based on housing: new insights from spatially explicit density models. Landsc Urban Plan 162:13–24CrossRefGoogle Scholar
  24. Fortin D, Boyce MS, Merrill EH, Fryxell JM (2004) Foraging costs of vigilance in large mammalian herbivores. Oikos 107:172–180CrossRefGoogle Scholar
  25. Fortin D, Beyer HL, Boyce MS, Smith DW, Duchesne T, Mao JS (2005) Wolves influence elk movements: behavior shapes a trophic cascade in Yellowstone National Park. Ecology 86:1320–1330CrossRefGoogle Scholar
  26. Francis CD, Barber JR (2013) A framework for understanding noise impacts on wildlife: an urgent conservation priority. Front Ecol Environ 11:305–313CrossRefGoogle Scholar
  27. Frid A, Dill LM (2002) Human-caused disturbance stimuli as a form of predation risk. Conserv Ecol 6:11CrossRefGoogle Scholar
  28. Fry JA, Xian G, Jin S, Dewitz JA, Homer CG, Yang L, Barnes CA, Herold ND, Wickham JD (2011) Completion of the 2006 national land cover database for the conterminous united states. Photogramm Eng Remote Sens 77(9):858–864Google Scholar
  29. Garshelis DL, Noyce KV, Ditmer MA (2012) Ecology and population dynamics of black bears in Minnesota. In: Cornicelli L, Carstensen M, Grund MD, Larson MA, Lawrence JS (eds) Summaries of wildlife research findings 2012. Minnesota DNR, St. Paul, pp 13–27Google Scholar
  30. Gehrt SD, Brown JL, Anchor C (2011) Is the urban coyote a misanthropic synanthrope? The case from Chicago. Cities Environ 4:3CrossRefGoogle Scholar
  31. Groffman PM, Baron JS, Blett T, Gold AJ, Goodman I, Gunderson LH, Levinson BM, Palmer MA, Paerl HW, Peterson GD (2006) Ecological thresholds: the key to successful environmental management or an important concept with no practical application? Ecosystems 9:1–13CrossRefGoogle Scholar
  32. Hopkins JB (2013) Use of genetics to investigate socially learned foraging behavior in free-ranging black bears. J Mammal 94:1214–1222CrossRefGoogle Scholar
  33. Hosmer DW, Leeshow S (2000) Applied logistic regression. Wiley Inc., New YorkCrossRefGoogle Scholar
  34. Hristienko H, McDonald JE Jr (2007) Going into the 21st century: a perspective on trends and controversies in the management of the American black bear. Ursus 18:72–88CrossRefGoogle Scholar
  35. Hulbert IA, French J (2001) The accuracy of GPS for wildlife telemetry and habitat mapping. J Appl Ecol 38:869–878CrossRefGoogle Scholar
  36. Johnson DH (1980) The comparison of usage and availability measurements for evaluating resource preference. Ecology 61:65–71CrossRefGoogle Scholar
  37. Johnson HE, Breck SW, Baruch-Mordo S, Lewis DL, Lackey CW, Wilson KR, Broderick J, Mao JS, Beckmann JP (2015) Shifting perceptions of risk and reward: dynamic selection for human development by black bears in the western United States. Biol Conserv 187:164–172CrossRefGoogle Scholar
  38. Kertson BN, Spencer RD, Marzluff JM, Hepinstall-Cymerman J, Grue CE (2011) Cougar space use and movements in the wildland–urban landscape of western Washington. Ecol Appl 21:2866–2881CrossRefGoogle Scholar
  39. Knopff AA, Knopff KH, Boyce MA, St Clair CC (2014) Flexible habitat selection by cougars in response to anthropogenic development. Biol Conserv 178:136–145CrossRefGoogle Scholar
  40. Kohl MT, Stahler DR, Metz MC, Forester JD, Kauffman MJ, Varley N, White PJ, Smith DW, MacNulty DR (2018) Diel predator activity drives a dynamic landscape of fear. Ecol Monogr 88(4):638–652CrossRefGoogle Scholar
  41. Koops MA, Abrahams MV (1998) Life history and the fitness consequences of imperfect information. Evol Ecol 12:601–613CrossRefGoogle Scholar
  42. Lele SR, Merrill EH, Keim J, Boyce MS (2013) Selection, use, choice and occupancy: clarifying concepts in resource selection studies. J Anim Ecol 82:1183–1191CrossRefGoogle Scholar
  43. Lewis D, Breck S, Wilson K, Webb C (2014) Modeling black bear population dynamics in a human-dominated stochastic environment. Ecol Model 294:51–58CrossRefGoogle Scholar
  44. Lima SL, Bednekoff PA (1999) Temporal variation in danger drives antipredator behavior: the predation risk allocation hypothesis. Am Nat 153:649–659CrossRefGoogle Scholar
  45. Linnell JD, Swenson JE, Anderson R (2001) Predators and people: conservation of large carnivores is possible at high human densities if management policy is favourable. Anim Conserv 4:345–349CrossRefGoogle Scholar
  46. Lowry H, Lill A, Wong B (2013) Behavioural responses of wildlife to urban environments. Biol Rev 88:537–549CrossRefGoogle Scholar
  47. Lukacs PM, Burnham KP, Anderson DR (2009) Model selection bias and Freedman’s paradox. Ann Inst Stat Math 62:117–125CrossRefGoogle Scholar
  48. Martin JG, Réale D (2008) Temperament, risk assessment and habituation to novelty in eastern chipmunks, Tamias striatus. Anim Behav 75:309–318CrossRefGoogle Scholar
  49. Matthiopoulos J, Hebblewhite M, Aarts G, Fieberg J (2011) Generalized functional responses for species distributions. Ecology 92:583–589CrossRefGoogle Scholar
  50. Mazur R, Seher V (2008) Socially learned foraging behavior in wild black bears, Ursus americanus. Anim Behav 75:1503–1508CrossRefGoogle Scholar
  51. McKinney ML (2006) Urbanization as a major cause of biotic homogenization. Biol Conserv 127:247–260CrossRefGoogle Scholar
  52. Merkle JA, Robinson HS, Krausman PR, Alaback P (2013) Food availability and foraging near human developments by black bears. J Mammal 94:378–385CrossRefGoogle Scholar
  53. Messmer TA (2009) Human–wildlife conflicts: emerging challenges and opportunities. Hum Wildl Confl 3:10–17Google Scholar
  54. Naves J, Wiegand T, Revilla E, Delibes M (2003) Endangered species constrained by natural and human factors: the case of brown bears in northern Spain. Conserv Biol 17:1276–1289CrossRefGoogle Scholar
  55. Nellemann C, Støen O, Kindberg J, Swenson JE, Vistnes I, Ericsson G, Katajisto J, Kaltenborn BP, Martin J, Ordiz A (2007) Terrain use by an expanding brown bear population in relation to age, recreational resorts and human settlements. Biol Conserv 138:157–165CrossRefGoogle Scholar
  56. Ordiz A, Støen O, Delibes M, Swenson JE (2011) Predators or prey? Spatio-temporal discrimination of human-derived risk by brown bears. Oecologia 166:59–67CrossRefGoogle Scholar
  57. Powell RA, Zimmerman JW, Seaman DE, Gilliam JF (1996) Demographic analyses of a hunted black bear population with access to a refuge. Conserv Biol 10:224–234CrossRefGoogle Scholar
  58. Powell RA, Zimmerman JW, Seaman DE (1997) Ecology and behaviour of North American black bears: home ranges, habitat, and social organization. Springer, New YorkGoogle Scholar
  59. R Core Team (2016) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Accessed Sept 2016
  60. Remeš V (2000) How can maladaptive habitat choice generate source-sink population dynamics? Oikos 91:579–582CrossRefGoogle Scholar
  61. Riley SPD, Sauvajot RM, Fuller TK, York EC, Kamradt DA, Bromley C, Wayne RK (2003) Effects of urbanization and habitat fragmentation on bobcats and coyotes in southern California. Conserv Biol 17:566–576CrossRefGoogle Scholar
  62. Robertson BA, Rehage JS, Sih A (2013) Ecological novelty and the emergence of evolutionary traps. Trends Ecol Evol 28:552–560CrossRefGoogle Scholar
  63. Rode KD, Farley SD, Robbins CT (2006) Sexual dimorphism, reproductive strategy, and human activities determine resource use by brown bears. Ecology 87:2636–2646CrossRefGoogle Scholar
  64. Rodriguez-Prieto I, Fernández-Juricic E, Martín J, Regis Y (2009) Antipredator behavior in blackbirds: habituation complements risk allocation. Behav Ecol 20:371–377CrossRefGoogle Scholar
  65. Roever CL, Boyce MS, Stenhouse GB (2010) Grizzly bear movements relative to roads: application of step selection functions. Ecography 33:1113–1122CrossRefGoogle Scholar
  66. Rogers LL, Allen W (1987) Habitat suitability index models: black bear upper great lakes region. U.S. Fish Wildlife Service Biology Report 82(10.144)Google Scholar
  67. Schlaepfer MA, Runge MC, Sherman PW (2002) Ecological and evolutionary traps. Trends Ecol Evol 17:474–480CrossRefGoogle Scholar
  68. Shochat E, Warren PS, Faeth SH, McIntyre NE, Hope D (2006) From patterns to emerging processes in mechanistic urban ecology. Trends Ecol Evol 21:186–191CrossRefGoogle Scholar
  69. Sih A, Bell A, Johnson JC (2004) Behavioral syndromes: an ecological and evolutionary overview. Trends Ecol Evol 19:372–378CrossRefGoogle Scholar
  70. Slater P (1981) Individual differences in animal behavior. In: Bateson PPG, Klopfer PH (eds) Perspectives in ethology. Springer, New York, pp 35–49CrossRefGoogle Scholar
  71. Spencer RD, Beausoleil RA, Martorello DA (2007) How agencies respond to human-black bear conflicts: a survey of wildlife agencies in North America. Ursus 18:217–229CrossRefGoogle Scholar
  72. Steyaert SMJG, Zedrosser A, Elfstrom M, Ordiz A, Leclerc M, Frank SC, Kindberg J, Stoen O-G, Brunberg S, Swenson JE (2016) Ecological implications from spatial patterns in human-caused brown bear mortality. Wildl Biol 22(4):144–152CrossRefGoogle Scholar
  73. Therneau T (2015) A package for survival analysis in S v 2.38Google Scholar
  74. Thurfjell H, Ciuti S, Boyce M (2014) Applications of step-selection functions in ecology and conservation. Mov Ecol 2:4CrossRefGoogle Scholar
  75. Tuomainen U, Candolin U (2011) Behavioural responses to human-induced environmental change. Biol Rev 86:640–657CrossRefGoogle Scholar
  76. Van Horne B (1983) Density as a misleading indicator of habitat quality. J Wildl Manag 47:893–901CrossRefGoogle Scholar

Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  • Michael J. Evans
    • 1
    Email author
  • Jason E. Hawley
    • 2
  • Paul W. Rego
    • 2
  • Tracy A. G. Rittenhouse
    • 1
  1. 1.Department of Natural Resources and the EnvironmentUniversity of Connecticut, Wildlife and Fisheries Conservation CenterStorrsUSA
  2. 2.Connecticut Department of Energy and Environmental Protection Wildlife DivisionBurlingtonUSA

Personalised recommendations