Register      Login
Wildlife Research Wildlife Research Society
Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

Incorporating movement patterns to discern habitat selection: black bears as a case study

Dana L. Karelus https://orcid.org/0000-0003-0029-2288 A E , J. Walter McCown B , Brian K. Scheick B , Madelon van de Kerk C , Benjamin M. Bolker D and Madan K. Oli A
+ Author Affiliations
- Author Affiliations

A Department of Wildlife Ecology and Conservation, and School of Natural Resources and Environment, 110 Newins-Ziegler Hall, University of Florida, Gainesville, FL 32611, USA.

B Florida Fish and Wildlife Conservation Commission, 1105 SW Williston Road, Gainesville, FL 32601, USA.

C School of Environmental and Forest Sciences, 114 Winkenwerder, University of Washington, Seattle, WA 98195, USA.

D Departments of Mathematics & Statistics and Biology, McMaster University, 314 Hamilton Hall, Hamilton, Ontario L8S 4K1, Canada.

E Corresponding author. Email: dana.karelus@sulross.edu

Wildlife Research 46(1) 76-88 https://doi.org/10.1071/WR17151
Submitted: 30 October 2017  Accepted: 4 November 2018   Published: 15 February 2019

Abstract

Context: Animals’ use of space and habitat selection emerges from their movement patterns, which are, in turn, determined by their behavioural or physiological states and extrinsic factors.

Aim: The aims of the present study were to investigate animal movement and incorporate the movement patterns into habitat selection analyses using Global Positioning System (GPS) location data from 16 black bears (Ursus americanus) in a fragmented area of Florida, USA.

Methods: Hidden Markov models (HMMs) were used to discern the movement patterns of the bears. These results were then used in step-selection functions (SSFs) to evaluate habitat selection patterns and the factors influencing these patterns.

Key results: HMMs revealed that black bear movement patterns are best described by three behavioural states: (1) resting (very short step-lengths and large turning angles); (2) encamped (moderate step-lengths and large turning angles); and (3) exploratory (long step-lengths and small turning angles). Bears selected for forested wetlands and marsh wetlands more than any other land cover type, and generally avoided urban areas in all seasons and when in encamped and exploratory behavioural states. Bears also chose to move to locations farther away from major roads.

Conclusions: Because habitat selection is influenced by how animals move within landscapes, it is essential to consider animals’ movement patterns when making inferences about habitat selection. The present study achieves this goal by using HMMs to first discern black bear movement patterns and associated parameters, and by using these results in SSFs to investigate habitat selection patterns. Thus, the methodological framework developed in this study effectively incorporates state-specific movement patterns while making inferences regarding habitat selection. The unified methodological approach employed here will contribute to an improved understanding of animal ecology as well as informed management decisions.

Implications: Conservation plans focused on preserving forested wetlands would benefit bears by not only providing habitat for resting and foraging, but also by providing connectivity through fragmented landscapes. Additionally, the framework could be applied to species that follow annual cycles and may provide a tool for investigating how animals are using dispersal corridors.

Additional keywords: black bear movement, hidden Markov models, step-selection functions, step-length, Ursus americanus.


References

Abrahms, B., Sawyer, S. C., Jordan, N. R., Mcnutt, J. W., Wilson, A. M., and Brashares, J. S. (2017). Does wildlife resource selection accurately inform corridor conservation? Journal of Applied Ecology 54, 412–422.
Does wildlife resource selection accurately inform corridor conservation?Crossref | GoogleScholarGoogle Scholar |

Alt, G. L., Matula, G. J., Alt, F. W., and Lindzey, J. S. (1980). Dynamics of home range and movements of adult black bears in northeastern Pennsylvania. International Conference on Bear Research and Management 4, 131–136.

Benson, J. F., and Chamberlain, M. J. (2007). Space use and habitat selection by female Louisiana black bears in the Tensas River Basin of Louisiana. The Journal of Wildlife Management 71, 117–126.
Space use and habitat selection by female Louisiana black bears in the Tensas River Basin of Louisiana.Crossref | GoogleScholarGoogle Scholar |

Beyer, H. L., Gurarie, E., Börger, L., Panzacchi, M., Basille, M., Herfindal, I., Van Moorter, B., Lele, S. R., and Matthiapoulos, J. (2016). “You shall not pass!”: quantifying barrier permeability and proximity avoidance by animals. Journal of Animal Ecology 85, 43–53.
“You shall not pass!”: quantifying barrier permeability and proximity avoidance by animals.Crossref | GoogleScholarGoogle Scholar | 25056207PubMed |

Bivand, R., and Rundel, C. (2016). rgeos: interface to Geometry Engine – Open Source (GEOS). R package version 0.3–20. Available at https://CRAN.R-project.org/package=rgeos [Verified December 2018]

Boyce, M. S., Vernier, P. R., Nielsen, S. E., and Schmiegelow, F. K. A. (2002). Evaluating resource selection functions. Ecological Modelling 157, 281–300.
Evaluating resource selection functions.Crossref | GoogleScholarGoogle Scholar |

Boyd, C., Punt, A. E., Weimerskirch, H., and Bertrand, S. (2014). Movement models provide insights into variation in the foraging effort of central place foragers. Ecological Modelling 286, 13–25.
Movement models provide insights into variation in the foraging effort of central place foragers.Crossref | GoogleScholarGoogle Scholar |

Bridges, A. S., Vaughan, M. R., and Klenzendorf, S. (2004). Seasonal variation in American black bear Ursus americanus activity patterns: quantification via remote photography. Wildlife Biology 10, 277–284.

Buchmann, C. M., Schurr, F. M., Nathan, R., and Jeltsch, F. (2012). Movement upscaled – the importance of individual foraging movement for community response to habitat loss. Ecography 35, 436–445.
Movement upscaled – the importance of individual foraging movement for community response to habitat loss.Crossref | GoogleScholarGoogle Scholar |

Burnham, K. P., and Anderson, D. R. (2002). ‘Model Selection and Multimodel Inference: a Practical Information–theoretic Approach.’ 2nd edn. (Springer: New York.)

Carr, M. H., and Zwick, P. D. (2016). Florida 2070: mapping Florida’s future–alternative patterns of development in 2070. Technical Report to Florida Department of Agriculture and Consumer Services & 1000 Friends of Florida. Geoplan Center, University of Florida, Gainesville, Florida, USA.

Clark, J. D., Laufenberg, J. S., Davidson, M., and Murrow, J. L. (2015). Connectivity among subpopulations of Louisiana black bears as estimated by a step selection function. The Journal of Wildlife Management 79, 1347–1360.
Connectivity among subpopulations of Louisiana black bears as estimated by a step selection function.Crossref | GoogleScholarGoogle Scholar |

Costello, C. M., Cain, S. I., Nielson, R. M., Servheen, C., and Schwartz, C. C. (2013). Response of American black bears to the non-motorized expansion of a road corridor in Grand Teton National Park. Ursus 24, 54–69.
Response of American black bears to the non-motorized expansion of a road corridor in Grand Teton National Park.Crossref | GoogleScholarGoogle Scholar |

Coster, S. S., and Kovach, A. I. (2012). Anthropogenic influences on the spatial genetic structure of black bears. Conservation Genetics 13, 1247–1257.
Anthropogenic influences on the spatial genetic structure of black bears.Crossref | GoogleScholarGoogle Scholar |

Cozzi, G., Chynoweth, M., Kusak, J., Coban, E., Coban, A., Ozgul, A., and Sekercioglu, C. (2016). Anthropogenic food resources foster the coexistence of distinct life history strategies: year-round sedentary and migratory brown bears. Journal of Zoology 300, 142–150.
Anthropogenic food resources foster the coexistence of distinct life history strategies: year-round sedentary and migratory brown bears.Crossref | GoogleScholarGoogle Scholar |

Craiu, R. V., and Duchesne, T. (2018). A scalable and efficient covariate selection criterion for mixed effects regression models with unknown random effects structure. Computational Statistics & Data Analysis 117, 154–161.
A scalable and efficient covariate selection criterion for mixed effects regression models with unknown random effects structure.Crossref | GoogleScholarGoogle Scholar |

Craiu, R. V., Duchesne, T., Fortin, D., and Baillargeon, S. (2011). Conditional logistic regression with longitudinal follow-up and individual-level random coefficients: a stable and efficient two-step estimation method. Journal of Computational and Graphical Statistics 20, 767–784.
Conditional logistic regression with longitudinal follow-up and individual-level random coefficients: a stable and efficient two-step estimation method.Crossref | GoogleScholarGoogle Scholar |

Craiu, R. V., Duchesne, T., Fortin, D., and Baillargeon, S. (2016). TwoStepCLogit: conditional logistic regression: a two-step estimation method. R package version 1.2.5. Available at https://CRAN.R-project.org/package=TwoStepCLogit [Verified December 2018]

Ditmer, M. A., Moen, R. A., Windels, S. K., Forester, J. D., Ness, T. E., and Harris, T. R. (2017). Moose at their bioclimatic edge alter their behavior based on weather, landscape, and predators. Current Zoology 64, 419–432.
| 30109872PubMed |

Dobey, S., Masters, D. V., Scheick, B. K., Clark, J. D., Pelton, M. R., and Sunquist, M. E. (2005). Ecology of Florida black bears in the Okefenokee–Osceola ecosystem. Wildlife Monographs 158, 1–41.

Duchesne, T., Fortin, D., and Courbin, N. (2010). Mixed conditional logistic regression for habitat selection studies. Journal of Animal Ecology 79, 548–555.
Mixed conditional logistic regression for habitat selection studies.Crossref | GoogleScholarGoogle Scholar | 20202010PubMed |

Fieberg, J., Matthiopoulos, J., Hebblewhite, M., Boyce, M. S., and Frair, J. L. (2010). Correlation and studies of habitat selection: problem, red herring or opportunity? Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 365, 2233–2244.
Correlation and studies of habitat selection: problem, red herring or opportunity?Crossref | GoogleScholarGoogle Scholar | 20566500PubMed |

Florida Fish and Wildlife Conservation Commission (2012). Florida black bear management plan. Florida Fish and Wildlife Conservation Commission, Tallahassee, FL.

Forester, J. D., Im, H. K., and Rathouz, P. J. (2009). Accounting for animal movement in estimation of resource selection functions: sampling and data analysis. Ecology 90, 3554–3565.
Accounting for animal movement in estimation of resource selection functions: sampling and data analysis.Crossref | GoogleScholarGoogle Scholar | 20120822PubMed |

Forman, R. T. T., and Alexander, L. E. (1998). Roads and their major ecological effects. Annual Review of Ecology and Systematics 29, 207–231.
Roads and their major ecological effects.Crossref | GoogleScholarGoogle Scholar |

Forney, G. D. (1973). The Viterbi algorithm. Proceedings of the IEEE 61, 268–278.
The Viterbi algorithm.Crossref | GoogleScholarGoogle Scholar |

Fortin, D., Morales, J. M., and Boyce, M. S. (2005). Elk winter foraging at fine scale in Yellowstone National Park. Oecologia 145, 335–343.
Elk winter foraging at fine scale in Yellowstone National Park.Crossref | GoogleScholarGoogle Scholar | 15965755PubMed |

Franke, A., Caelli, T., and Hudson, R. J. (2004). Analysis of movements and behavior of caribou (Rangifer tarandus) using hidden Markov models. Ecological Modelling 173, 259–270.
Analysis of movements and behavior of caribou (Rangifer tarandus) using hidden Markov models.Crossref | GoogleScholarGoogle Scholar |

Franke, A., Caelli, T., Kuzyk, G., and Hudson, R. J. (2006). Prediction of wolf (Canis lupus) kill-sites using hidden Markov models. Ecological Modelling 197, 237–246.
Prediction of wolf (Canis lupus) kill-sites using hidden Markov models.Crossref | GoogleScholarGoogle Scholar |

Fryxell, J. M., Hazell, M., Börger, L., Dalziel, B. D., Haydon, D. T., Morales, J. M., McIntosh, T., and Rosatte, R. C. (2008). Multiple movement modes by large herbivores at multiple spatiotemporal scales. Proceedings of the National Academy of Sciences of the United States of America 105, 19114–19119.
Multiple movement modes by large herbivores at multiple spatiotemporal scales.Crossref | GoogleScholarGoogle Scholar | 19060190PubMed |

Garrison, E. P., McCown, J. W., Barrett, M. A., and Oli, M. K. (2012). Denning ecology of Florida black bears in north-central Florida. Southeastern Naturalist 11, 517–528.
Denning ecology of Florida black bears in north-central Florida.Crossref | GoogleScholarGoogle Scholar |

Garshelis, D. L. (1978). Movement ecology and activity behavior of black bears in the Great Smokey Moutains National Park. Masters Thesis, University of Tennessee.

Garshelis, D. L., and Hristienko, H. (2006). State and provincial estimates of American black bear numbers versus assessments of population trend State and provincial estimates of American black bear numbers versus assessments of population trend. Ursus 17, 1–7.
State and provincial estimates of American black bear numbers versus assessments of population trend State and provincial estimates of American black bear numbers versus assessments of population trend.Crossref | GoogleScholarGoogle Scholar |

Garshelis, D. L., and Pelton, M. R. (1980). Activity of black bears in the Great Smoky Mountains National Park. Journal of Mammalogy 61, 8–19.
Activity of black bears in the Great Smoky Mountains National Park.Crossref | GoogleScholarGoogle Scholar |

Garshelis, D. L., and Pelton, M. R. (1981). Movements of black bears in the Great Smoky Mountains. The Journal of Wildlife Management 45, 912–925.
Movements of black bears in the Great Smoky Mountains.Crossref | GoogleScholarGoogle Scholar |

Garshelis, D. L., Quigley, H. B., Villarrubia, C. R., and Pelton, M. R. (1983). Diel movements of black bears in the southern Appalachians. International Conference on Bear Research and Management 5, 11–19.

Gilbert, S. L., Hundertmark, K. J., Person, D. K., Lindberg, M. S., and Boyce, M. S. (2017). Behavioral plasticity in a variable environment: snow depth and habitat interactions drive deer movement in winter. Journal of Mammalogy 98, 246–259.
Behavioral plasticity in a variable environment: snow depth and habitat interactions drive deer movement in winter.Crossref | GoogleScholarGoogle Scholar |

Gurarie, E., Bracis, C., Delgado, M., Meckley, T. D., Kojola, I., and Wagner, C. M. (2015). What is the animal doing? Tools for exploring behavioral structure in animal movements. Journal of Animal Ecology 85, 69–84.
What is the animal doing? Tools for exploring behavioral structure in animal movements.Crossref | GoogleScholarGoogle Scholar | 25907267PubMed |

Hellgren, E. C., and Vaughan, M. R. (1990). Range dynamics of black bears in Great Dismal Swamp, Virginia–North Carolina. Proceedings of the Annual Conference Southeastern Association of Fish and Wildlife Agencies 44, 268–278.

Hellgren, E. C., Vaughan, M. R., and Kirkpatrick, R. L. (1989). Seasonal patterns in physiology and nutrition of black bears in Great Dismal Swamp, Virginia–North Carolina. Canadian Journal of Zoology 67, 1837–1850.
Seasonal patterns in physiology and nutrition of black bears in Great Dismal Swamp, Virginia–North Carolina.Crossref | GoogleScholarGoogle Scholar |

Hellgren, E. C., Vaughan, M. R., and Stauffer, D. F. (1991). Macrohabitat use by black bears in a southeastern wetland. The Journal of Wildlife Management 55, 442–448.
Macrohabitat use by black bears in a southeastern wetland.Crossref | GoogleScholarGoogle Scholar |

Hoctor, T. S., Carr, M. H., and Zwick, P. D. (2000). Identifying a linked reserve system using a regional landscape approach: The Florida Ecological Network. Conservation Biology 14, 984–1000.
Identifying a linked reserve system using a regional landscape approach: The Florida Ecological Network.Crossref | GoogleScholarGoogle Scholar |

Hristienko, H., and McDonald, J. E. (2007). Going into the 21st century: a perspective on trends and controversies in the management of the American black bear. Ursus 18, 72–88.
Going into the 21st century: a perspective on trends and controversies in the management of the American black bear.Crossref | GoogleScholarGoogle Scholar |

Humm, J. M., McCown, J. W., Scheick, B. K., and Clark, J. D. (2017). Spatially explicit population estimates for black bears based on cluster sampling. The Journal of Wildlife Management 81, 1187–1201.
Spatially explicit population estimates for black bears based on cluster sampling.Crossref | GoogleScholarGoogle Scholar |

Johnson, C., and Nielsen, S. (2006). Resource selection functions based on use-availability data: theoretical motivation and evaluation methods. The Journal of Wildlife Management 70, 347–357.
Resource selection functions based on use-availability data: theoretical motivation and evaluation methods.Crossref | GoogleScholarGoogle Scholar |

Jonsen, I. D., Ransom, A. M., and Flemming, J. M. (2003). Meta-analysis of animal movement using state-space models. Ecology 84, 3055–3063.
Meta-analysis of animal movement using state-space models.Crossref | GoogleScholarGoogle Scholar |

Karelus, D. L., McCown, J. W., Scheick, B. K., van de Kerk, M., and Oli, M. K. (2016). Home ranges and habitat selection by black bears in a newly colonized population in Florida. Southeastern Naturalist (Steuben, ME) 15, 346–364.
Home ranges and habitat selection by black bears in a newly colonized population in Florida.Crossref | GoogleScholarGoogle Scholar |

Karelus, D. L., McCown, J. W., Scheick, B. K., van de Kerk, M., Bolker, B. M., and Oli, M. K. (2017). Effects of environmental factors and landscape features on movement patterns of Florida black bears. Journal of Mammalogy 98, 1463–1478.

Langrock, R., King, R., Matthiopoulos, J., Thomas, L., Fortin, D., and Morales, J. M. (2012). Flexible and practical modeling of animal telemetry data: hidden Markov models and extensions. Ecology 93, 2336–2342.
Flexible and practical modeling of animal telemetry data: hidden Markov models and extensions.Crossref | GoogleScholarGoogle Scholar | 23236905PubMed |

Latham, A. D. M., Latham, M. C., Boyce, M. S., and Boutin, S. (2011). Movement responses by wolves to industrial linear features and their effect on woodland caribou in northeastern Alberta. Ecological Applications 21, 2854–2865.
Movement responses by wolves to industrial linear features and their effect on woodland caribou in northeastern Alberta.Crossref | GoogleScholarGoogle Scholar |

Latombe, G., Parrott, L., Basille, M., and Fortin, D. (2014). Uniting statistical and individual-based approaches for animal movement modelling. PLoS One 9, e99938.
Uniting statistical and individual-based approaches for animal movement modelling.Crossref | GoogleScholarGoogle Scholar | 24979047PubMed |

Leos-Barajas, V., Photopoulou, T., Langrock, R., Patterson, T. A., Watanabe, Y. Y., Murgatroyd, M., and Papastamatiou, Y. P. (2017). Analysis of animal accelerometer data using hidden Markov models. Methods in Ecology and Evolution 8, 161–173.
Analysis of animal accelerometer data using hidden Markov models.Crossref | GoogleScholarGoogle Scholar |

Lewis, J. S., and Rachlow, J. L. (2011). Activity patterns of black bears in relation to sex, season, and daily movement rates. Western North American Naturalist 71, 388–395.
Activity patterns of black bears in relation to sex, season, and daily movement rates.Crossref | GoogleScholarGoogle Scholar |

Li, M., and Bolker, B. M. (2017). Incorporating periodic variability in hidden Markov models for animal movement. Movement Ecology 5, 1–12.
Incorporating periodic variability in hidden Markov models for animal movement.Crossref | GoogleScholarGoogle Scholar | 28149522PubMed |

Lund U., and Agostinelli. C. (2012). CircStats: Circular Statistics. R package version 0.2-4. Available at https://cran.r-project.org/package=CircStats [Verified 16 January 2019].

Maehr, D. S., and Brady, J. R. (1984). Food habits of Florida black bears. The Journal of Wildlife Management 48, 230–235.
Food habits of Florida black bears.Crossref | GoogleScholarGoogle Scholar |

Manly, B. F. J., Mcdonald, L. L., Thomas, D. L., McDonald, T. L., and Erickson, W. P. (2002). ‘Resource Selection by Animals: Statistical Design and Analysis for Field Studies.’ 2nd edn. (Kluwer Academic Publishers: Boston, MA.)

Martin, J, van Moorter, B., Revilla, E., Blanchard, P., Dray, S., Quenette, P. Y., Allainé, D., and Swenson, J. E. (2013). Reciprocal modulation of internal and external factors determines individual movements. Journal of Animal Ecology 82, 290–300.
Reciprocal modulation of internal and external factors determines individual movements.Crossref | GoogleScholarGoogle Scholar | 23039315PubMed |

Martin, J., Calenge, C., Quenette, P.-Y., and Allainé, D. (2008). Importance of movement constraints in habitat selection studies. Ecological Modelling 213, 257–262.
Importance of movement constraints in habitat selection studies.Crossref | GoogleScholarGoogle Scholar |

McGreer, M. T., Mallon, E. E., Vander Vennen, L. M., Wiebe, P. A., Baker, J. A., Brown, G. S., Avgar, T., Hagens, J., Kittle, A. M., Mosser, A., Street, G. M., Reid, D. E. B., Rodgers, A. R., Shuter, J., Thompson, I. D., Turetsky, M. J., Newmaster, S. G., Patterson, B. R., and Fryxell, J. M. (2015). Selection for forage and avoidance of risk by woodland caribou (Rangifer tarandus caribou) at coarse and local scales. Ecosphere 6, art288.
Selection for forage and avoidance of risk by woodland caribou (Rangifer tarandus caribou) at coarse and local scales.Crossref | GoogleScholarGoogle Scholar |

McKellar, A. E., Langrock, R., Walters, J. R., and Kesler, D. C. (2015). Using mixed hidden Markov models to examine behavioral states in a cooperatively breeding bird. Behavioral Ecology 26, 148–157.
Using mixed hidden Markov models to examine behavioral states in a cooperatively breeding bird.Crossref | GoogleScholarGoogle Scholar |

Michelot, T., Langrock, R., and Patterson, T. A. (2016). moveHMM: an R package for the statistical modelling of animal movement data using hidden Markov models. Methods in Ecology and Evolution 7, 1308–1315.
moveHMM: an R package for the statistical modelling of animal movement data using hidden Markov models.Crossref | GoogleScholarGoogle Scholar |

Moorcroft, P. R. (2012). Mechanistic approaches to understanding and predicting mammalian space use: recent advances, future directions. Journal of Mammalogy 93, 903–916.
Mechanistic approaches to understanding and predicting mammalian space use: recent advances, future directions.Crossref | GoogleScholarGoogle Scholar |

Moorcroft, P. R., and Barnett, A. (2008). mechanistic home range models and resource selection analysis : a reconciliation and unification. Ecology 89, 1112–1119.
mechanistic home range models and resource selection analysis : a reconciliation and unification.Crossref | GoogleScholarGoogle Scholar | 18481535PubMed |

Morales, J. M., Haydon, D. T., Frair, J., Holsinger, K. E., and Fryxell, J. M. (2004). Extracting more out of relocation data: building movement models as mixtures of random walks. Ecology 85, 2436–2445.
Extracting more out of relocation data: building movement models as mixtures of random walks.Crossref | GoogleScholarGoogle Scholar |

Moyer, M. A., Mccown, J. W., and Oli, M. K. (2007). Factors influencing home range size of female Florida black bears. Journal of Mammalogy 88, 468–476.
Factors influencing home range size of female Florida black bears.Crossref | GoogleScholarGoogle Scholar |

Moyer, M. A., McCown, J. W., and Oli, M. K. (2008). Scale-dependent habitat selection by female Florida black bears in Ocala National Forest, Florida. Southeastern Naturalist (Steuben, ME) 7, 111–124.
Scale-dependent habitat selection by female Florida black bears in Ocala National Forest, Florida.Crossref | GoogleScholarGoogle Scholar |

Nathan, R., Getz, W. M., Revilla, E., Holyoak, M., Kadmon, R., Saltz, D., and Smouse, P. E. (2008). A movement ecology paradigm for unifying organismal movement research. Proceedings of the National Academy of Sciences of the United States of America 105, 19052–19059.
A movement ecology paradigm for unifying organismal movement research.Crossref | GoogleScholarGoogle Scholar | 19060196PubMed |

Noyce, K. V., and Garshelis, D. L. (2011). Seasonal migrations of black bears (Ursus americanus): causes and consequences. Behavioral Ecology and Sociobiology 65, 823–835.
Seasonal migrations of black bears (Ursus americanus): causes and consequences.Crossref | GoogleScholarGoogle Scholar |

Onorato, D. P., Hellgren, E. C., Mitchell, F. S., and Skiles, J. R. (2003). Home range and habitat use of American black bears on a desert montane island in Texas. Ursus 14, 120–129.

Patterson, T. A., Basson, M., Bravington, M. V., and Gunn, J. S. (2009). Classifying movement behaviour in relation to environmental conditions using hidden Markov models. Journal of Animal Ecology 78, 1113–1123.
Classifying movement behaviour in relation to environmental conditions using hidden Markov models.Crossref | GoogleScholarGoogle Scholar | 19563470PubMed |

Patterson, T. A., Parton, A., Langrock, R., Blackwell, P. G., Thomas, L., and King, R. (2017). Statistical modelling of individual animal movement: an overview of key methods and a discussion of practical challenges. AStA. Advances in Statistical Analysis 101, 399–438.
Statistical modelling of individual animal movement: an overview of key methods and a discussion of practical challenges.Crossref | GoogleScholarGoogle Scholar |

Pelton, M. R. (2003). Black bears (Ursus americanus). In ‘Wild Mammals of North America’. 2nd edn. (Eds G. A. Feldhamer, B. C. Thompson and J. A. Chapman.) pp. 547–556. (Johns Hopkins University Press: Baltimore, MD.)

Pohle, J., Langrock, R., van Beest, F. M., and Schmidt, N. M. (2017). Selecting the number of states in hidden Markov models: pragmatic solutions illustrated using animal movement. Journal of Agricultural Biological & Environmental Statistics , .
Selecting the number of states in hidden Markov models: pragmatic solutions illustrated using animal movement.Crossref | GoogleScholarGoogle Scholar |

Powell, R. A., Zimmerman, J. W., and Seaman, D. E. (1997). ‘Ecology and Behaviour of North American Black Bears: Home Ranges, Habitat and Social Organization.’ (Chapman and Hall: London, UK.)

R Core Team (2016). R: a language and environment for statistical computing. (R Foundation for Statistical Computing, Vienna, Austria.) Available at http://www.r-project.org/ [Verified December 2018]

Rayl, N. D., Fuller, T. K., Organ, J. F., McDonald, J. E., Otto, R. D., and Mahoney, S. P. (2014). Den abandonment and transitional day bed use by black bears (Ursus americanus) in Newfoundland. Wildlife Biology 20, 222–228.
Den abandonment and transitional day bed use by black bears (Ursus americanus) in Newfoundland.Crossref | GoogleScholarGoogle Scholar |

Redner, J., and Srinivasan, S. (2014). Florida vegetation and land cover 2014. Final Report. Fish and Wildlife Research Institute grant #6207. (Florida Fish and Wildlife Conservation Commission: Tallahassee, FL.)

Revelt, D., and Train, K. (1998). Mixed logit with repeated choices: households’ choices of appliance efficiency level. The Review of Economics and Statistics 80, 647–657.
Mixed logit with repeated choices: households’ choices of appliance efficiency level.Crossref | GoogleScholarGoogle Scholar |

Reynolds-Hogland, M. J., and Mitchell, M. S. (2007). Effects of roads on habitat quality for bears in the southern Appalachians: a long-term study. Journal of Mammalogy 88, 1050–1061.
Effects of roads on habitat quality for bears in the southern Appalachians: a long-term study.Crossref | GoogleScholarGoogle Scholar |

Scheick, B. K., and McCown, W. (2014). Geographic distribution of American black bears in North America. Ursus 25, 24–33.
Geographic distribution of American black bears in North America.Crossref | GoogleScholarGoogle Scholar |

Scheick, B. K., Cunningham, M. W., McCown, J. W., and Orlando, M. A. (2009). Anchor modification for a foot-hold snare to capture American black bears. Ursus 20, 47–49.
Anchor modification for a foot-hold snare to capture American black bears.Crossref | GoogleScholarGoogle Scholar |

Schick, R. S., Loarie, S. R., Colchero, F., Best, B. D., Boustany, A., Conde, D. A., Halpin, P. N., Joppa, L. N., McClellan, C. M., and Clark, J. S. (2008). Understanding movement data and movement processes: current and emerging directions. Ecology Letters 11, 1338–1350.
Understanding movement data and movement processes: current and emerging directions.Crossref | GoogleScholarGoogle Scholar | 19046362PubMed |

Schliehe-Diecks, S., Kappeler, P. M., and Langrock, R. (2012). On the application of mixed hidden Markov models to multiple behavioural time series. Interface Focus 2, 180–189.
On the application of mixed hidden Markov models to multiple behavioural time series.Crossref | GoogleScholarGoogle Scholar | 23565332PubMed |

Servheen, C., Herrero, S., and Peyton, B. (1999). Bears: status, survey and conservation action plan. IUCN/SSC Action Plans for the Conservation of Biological Diversity. (IUCN/SSC Bear Specialist Group: Gland, Switzerland.)

Squires, J. R., DeCesare, N. J., Olson, L. E., Kolbe, J. A., Hebblewhite, M., and Parks, S. A. (2013). Combining resource selection and movement behavior to predict corridors for Canada lynx at their southern range periphery. Biological Conservation 157, 187–195.
Combining resource selection and movement behavior to predict corridors for Canada lynx at their southern range periphery.Crossref | GoogleScholarGoogle Scholar |

Stratman, M. R., and Pelton, M. R. (1999). Feeding ecology of black bears in northwest Florida. Florida Field Naturalist 27, 95–102.

Stratman, M., Alden, C., Pelton, M., and Sunquist, M. (2001). Habitat use by American black bears in the sandhills of Florida. Ursus 12, 109–114.

Thurfjell, H., Ciuti, S., and Boyce, M. S. (2014). Applications of step-selection functions in ecology and conservation. Movement Ecology 2, 4.
Applications of step-selection functions in ecology and conservation.Crossref | GoogleScholarGoogle Scholar | 25520815PubMed |

Tigas, L. A., Van Vuren, D. H., and Sauvajot, R. M. (2002). Behavioral responses of bobcats and coyotes to habitat fragmentation and corridors in an urban environment. Biological Conservation 108, 299–306.
Behavioral responses of bobcats and coyotes to habitat fragmentation and corridors in an urban environment.Crossref | GoogleScholarGoogle Scholar |

van de Kerk, M., Onorato, D. P., Criffield, M. A., Bolker, B. M., Augustine, B. C., McKinley, S. A., and Oli, M. K. (2014). Hidden semi-Markov models reveal multiphasic movement of the endangered Florida panther. Journal of Animal Ecology 84, 576–585.
Hidden semi-Markov models reveal multiphasic movement of the endangered Florida panther.Crossref | GoogleScholarGoogle Scholar | 25251870PubMed |

van Moorter, B., Rolandsen, C. M., Basille, M., and Gaillard, J.-M. (2016). Movement is the glue connecting home ranges and habitat selection. Journal of Animal Ecology 85, 21–31.
Movement is the glue connecting home ranges and habitat selection.Crossref | GoogleScholarGoogle Scholar | 25980987PubMed |

Visser, I. (2011). Seven things to remember about hidden Markov models: a tutorial on Markovian models for time series. Journal of Mathematical Psychology 55, 403–415.
Seven things to remember about hidden Markov models: a tutorial on Markovian models for time series.Crossref | GoogleScholarGoogle Scholar |

Willey, C. H. (1974). Aging black bears from first premolar tooth sections. The Journal of Wildlife Management 38, 97–100.
Aging black bears from first premolar tooth sections.Crossref | GoogleScholarGoogle Scholar |

Wooding, J. B., and Hardisky, T. S. (1992). Denning by black bears in northcentral Florida. Journal of Mammalogy 73, 895–898.
Denning by black bears in northcentral Florida.Crossref | GoogleScholarGoogle Scholar |

Wooding, J. B., and Hardisky, T. S. (1994). Home range, habitat use, and mortality of black bears in north-central Florida. International Conference on Bear Research and Management 9, 349–356.

Zeller, K. A., Mcgarigal, K., Cushman, S. A., Beier, P., Vickers, T. W., and Boyce, W. M. (2016). Using step and path selection functions for estimating resistance to movement: pumas as a case study. Landscape Ecology 31, 1319–1335.
Using step and path selection functions for estimating resistance to movement: pumas as a case study.Crossref | GoogleScholarGoogle Scholar |

Zucchini, W., and Macdonald, I. L. (2016). ‘Hidden Markov Models for Time Series: an Introduction Using R.’ 2nd edn. (Chapman and Hall/CRC Press: Boca Raton, FL.)