Sam Cushman, Ph.D.

Photo of Sam

Research Landscape Ecologist
Forest and Woodlands Ecosystems Program
Rocky Mountain Research Station
800 E. Beckwith Ave.
Missoula MT 59801
Phone: 406.329.2122
Fax: 406.543.2663
E-mail: scushman [at]
Genetics Lab Website:





B.S. Evergreen State College, Olympia, WA. 1995
M.S. Western Washington University, Bellingham, WA, 1997
Ph.D. University of Massachusetts, Amherst, MA, 2003


  1. McGarigal, K., S.A. Cushman, and S. Stafford. 2000. Multivariate Statistics for Wildlife and Ecology Research. Springer-Verlag, New York.
  2. McGarigal, K., S.A. Cushman, M. Neel and E. Ene. (in prep). Landscape Metrics: A Comprehensive Guide to their Use and Interpretation.
  3. Huettman, F. and S.A. Cushman. (in prep). Spatial Data and Mammal Ecology. Springer-Verlag, Tokyo. The scientist is co-editor of this volume and is contributing four original chapters.
    Refereed Journal Articles
  5. Cushman, S.A., K. McKelvey, C. Flather and K. McGarigal. (submitted). Testing the use of forest communities to evaluate biological diversity. Frontiers in Ecology and the Environment.
  6. Compton, B., K. McGarigal, S.A. Cushman and L. Gamble. 2007. A resistant kernel model of connectivity for vernal pool breeding amphibians. Conservation Biology.[PDF]
  7. Cushman, S.A., M.K. Schwartz, J. Hayden, and K. McKelvey. 2006. Gene flow in complex landscapes: Confronting models with data. American Naturalist 168: 486-499. [PDF]
  8. Schwartz, M.K., S.A. Cushman, J. Hayden, K. McKelvey and C. Engkjer. 2006. Detecting Genotyping Errors and Describing Black Bear Movement in Northern Idaho, USA. Ursus 17:138-148. [PDF]
  9. Cushman, S.A. 2006. Implications of habitat loss and fragmentation for the conservation of pond breeding amphibians: A review and prospectus. Biological Conservation 128: 231-240.
  10. Cushman, S.A., M. Chase, and C. Griffin 2005. Elephants in space and time. Oikos 109: 331-341. [PDF]
  11. Cushman, S.A. and K. McGarigal. 2004. Patterns in the species-environment relationship depend on both scale and choice of response variables. Oikos 105: 117-124.
  12. Neel, M.C., S.A. Cushman, and K. McGarigal. 2004. Behavior and stability of landscape metrics across controlled gradients of landscape structure. Landscape Ecology. 19:534-455.
  13. Cushman, S.A. and K. McGarigal. 2004. Hierarchical analysis of forest bird species-environment relationships in the Oregon Coast Range. Ecological Applications. 14(4): 1090-1105. [HTML]
  14. Grand, J., Buonaccorsi, S.A. Cushman, M. Mello, and C.R. Griffin. 2004. A comparison of predicted and observed bird and moth rarity hotspots in a threatened pitch pine - scrub oak (Pinus rigida Querecus ilicifolia) community. Conservation Biology 18(4): 1063-1077.
  15. Grand, J. and S.A. Cushman. 2003. Multiple-scale analysis of bird-habitat relationships: breeding birds in a pitch pine-scrub oak (Pinus rigida-Quercus ilicifolia) community. Biological Conservation. 12(3):307-317.
  16. Cushman, S.A. and K. McGarigal. 2003. Landscape-level patterns of avian diversity in the Oregon Coast Range. Ecological Monographs 73(2):259-281.
  17. Cushman, S.A. and K. McGarigal. 2002. Hierarchical, multi-scale decomposition of species-environment relationships. Landscape Ecology. 17:637-646.
  18. Cushman, S.A. and D.O. Wallin. 2002. Separating the influences of spatial, environmental, and disturbance factors in forests of the Russian Far East. Forest Ecology and Management. 168 (1-3): 201-215.
  19. McGarigal, K. and S.A. Cushman. 2002. Comparative evaluation of experimental approaches to the study of habitat fragmentation effects. Ecological Applications 12(2): 335-345.
  20. Cushman, S.A. and D.O. Wallin. 2000. Rates and patterns of landscape change in the Sikhote-alin Mountains, Russian Far East. Landscape Ecology 15(7): 643-659.
    Book Chapters
  22. Cushman, S.A. and K. McGarigal. (in press). Landscape metrics and scales of resolution. Chapter 4 in K.V. Gadow, (ed.), Adaptive Design of Forested Landscapes: The Multiple Path Principle. Springer.
  23. McKenzie, D., C.L. Raymond, and S. Cushman. (in press). Modeling understory vegetation and its response to fire. In J.J. Millspaugh and F.R. Thompson, III, (eds.) Models for planning wildlife conservation in large landscapes. Elsevier.
  24. Cushman, S.A., K.S. McKelvey, and K. McGarigal. (solicited). Landscape fire and wildlife habitat. Chapter 5 in D. McKenzie and L.-K. Kellog (eds.) The Landscape Ecology of Fire. Springer.
  25. Cushman, S.A., K. McKelvey, C. Reagan, T. Demeo, and C. Vojta. (accepted). Landscape analysis of wildlife habitat. Chapter 6 in C. Vojta (ed.), USFS Wildlife Habitat Technical Guide.
  26. McKelvey, K.S., S.A. Cushman, M.K. Schwartz, and L. F. Ruggiero. (In Press). Wildlife monitoring across multiple spatial scales using grid-based sampling. Symposium proceedings, 8th Annual Forest Inventory and Analysis Symposium, Monterey, CA, 2006.
  27. Cushman, S.A., K.S. McKelvey, and M.K. Schwartz. (In Press). Case Study I: DNA survey for fisher in Northern Idaho. In R.A. Long, P. MacKay, J.C. Ray, and W.J. Zielinski (eds). Noninvasive Survey Methods for North American Carnivores. Island Press.
  28. Cushman, S.A. and K. McGarigal. 2006. Multi-variate landscape trajectory analysis: An example using simulation modeling of American marten habitat change under three disturbance regimes. in J. A.: Bissonette, J. A., and I. Storch. Temporal Explicitness in Landscape Ecology: Wildlife Responses to Changes in Time. Springer, New York, New York.
  29. McGarigal, K. and S.A. Cushman. 2005. The gradient concept of landscape structure. Pgs. 112-119. in J.Wiens and M. Moss eds., Issues and Perspectives in Landscape Ecology. Cambridge University Press, Cambridge.
  30. Technical papers
  31. Cushman, S.A., D. McKenzie, J. Littell, and D.L. Peterson. (in press). A research prospectus for integrated landscape modelling. USDA For. Serv. Res. Paper RM.
  32. McGarigal, K., S. A. Cushman, and C. Reagan. (in review). Quantifying habitat loss and fragmentation: A protocol. USDA For. Serv. Res. Paper RM.