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John R. Squires

Capturing Canada lynx in Montana

Research Wildlife Biologist

800 East Beckwith Avenue
Missoula, MT 59801
Contact John R. Squires

Current Research


My current research focuses on how sensitive wildlife responds to increased natural (i.e., forest insect outbreaks, fire) and human-caused (i.e., recreation, forest fragmentation, energy development) disturbance. Understanding how increased disturbance impacts forests and wildlife is of global concern because these environmental changes are expected to accelerate with continued climate change and with our ever-expanding human footprint in natural landscapes. I am currently studying: 1) how Canada lynx and their prey respond to changes in habitat condition from fire and insect-related disturbance processes; 2) how the demography of Canada lynx is impacted by forest structure and composition; and 3) the response of lynx to forest silviculture and management.

Another important issue facing species conservation at a global scale is understanding how outdoor recreation and nature-based tourism impacts sensitive wildlife. Outdoor recreationists are central in the defense and use of public lands, but they also can impact sensitive species of wildlife through their motorized and non-motorized activities. I just completed studies that investigated how Canada lynx and wolverines respond to motorized (snowmobiles) and non-motorized (backcountry skiing) outdoor recreation in the western United States.

Finally, I have had a life-long interest in the conservation and management of birds of prey and native prairie ecosystems. I am studying how ferruginous hawks and golden eagles respond to energy development in terms of their behavior, movements, migration, resource use, and occupancy.

Research Interests

I am most interested in providing research that discovers better ways to manage and conserve sensitive species and their habitat in natural landscapes. The goal of my research is not only to advance scientific and ecological understanding in general, but to also provide practical solutions to the many management challenges associated with species conservation. My interests include learning how a species’ movements, spatial use, resource selection, behaviors, and viability relate to changes in landscape pattern, including forest silviculture and energy development. I am also very interested in learning how to enhance landscape connectivity for sensitive species, especially forest carnivores, in response to natural and anthropogenic change.

Past Research

Throughout my career, I conducted field studies that provided new scientific information based on direct measures of animal movement, behavior, and resource use; usually this was accomplished by instrumenting animals with GPS-based telemetry. This work included: 1) studying connectivity of wolverines in central Montana; 2) studying with cooperators how snow compaction changed coyote/lynx potential competition; and 3) researching how Canada lynx in Colorado respond to highways and transportation infrastructure. For over 20 years, I continued to lead Canada lynx studies from Montana through southern Colorado. During this time, I studied lynx demography in relation to forest condition, connectivity, seasonal changes in resource selection, denning behavior, movements, diets, and their spatial and temporal responses to forest silviculture. From 2000 to the present, I have continued to develop and evaluate detection and monitoring methods for mid-sized carnivores. During my early years, I studied the ecology of prairie falcons, northern goshawks, and trumpeter swans in Wyoming.  

Why This Research is Important

Wildlife conservation is a core American value. The public’s extensive input regarding conservation plans, regulation, media coverage, wildlife watching, and citizen science is a testimony to their deep concern for wildlife management on federal lands. My research is important because we learn how environmental changes to landscapes may either benefit or impair sensitive species depending on implementation. Results from our studies help inform activities across millions of acres of public land in ways that balance multi-use management with species conservation. Our research continues to provide a bridge of communication between people concerned about species conservation and those individuals and communities that are dependent on Forest Service and other public lands for timber, recreation, and energy.  


  • Colorado State University, B.S., Wildlife Biology, 1979
  • University of Wyoming, M.S., Zoology, 1986
  • University of Wyoming, Ph.D., Zoology, 1991
  • Featured Publications


    Kraberger, Simona; Serieys, Laurel EK.; Richet, Cécile; Fountain-Jones, Nicholas M.; Baele, Guy; Bishop, Jacqueline M.; Nehring, Mary; Ivan, Jacob S.; Newkirk, Eric S.; Squires, John R.; Lund, Michael C.; Riley, Seth PD.; Wilmers, Christopher C.; van Helden, Paul D.; Van Doorslaer, Koenraad; Culver, Melanie; VandeWoude, Sue; Martin, Darren P.; Varsani, Arvind, 2021. Complex evolutionary history of felid anelloviruses
    Bandoo, Rohan Antonio; Bautista, Joshua; Lund, Michael; Newkirk, Eric; Squires, John R.; Varsani, Arvind; Kraberger, Simona, 2021. Identification of novel circovirus and anelloviruses from wolverines using a non-invasive faecal sampling approach
    Olson, Lucretia E.; Bjornlie, Nichole; Hanvey, Gary; Holbrook, Joseph D.; Ivan, Jacob S.; Jackson, Scott; Kertson, Brian; King, Travis; Lucid, Michael; Murray, Dennis; Naney, Robert; Rohrer, John; Scully, Arthur; Thornton, Daniel; Walker, Zachary; Squires, John R., 2021. Improved prediction of Canada lynx distribution through regional model transferability and data efficiency
    Squires, John R.; Holbrook, Joseph D.; Olson, Lucretia E.; Ivan, Jacob S.; Ghormley, Randal W.; Lawrence, Rick L., 2020. A specialized forest carnivore navigates landscape-level disturbance: Canada lynx in spruce-beetle impacted forests
    Balkenhol, Niko; Schwartz, Michael K.; Inman, Robert M.; Copeland, Jeffrey P.; Squires, John R.; Anderson, Neil J.; Waits, Lisette P., 2020. Landscape genetics of wolverines (Gulo gulo): Scale-dependent effects of bioclimatic, topographic, and anthropogenic variables
    Squires, John R.; Olson, Lucretia E.; Wallace, Zachary P.; Oakleaf, Robert J.; Kennedy, Patricia L., 2020. Resource selection of apex raptors: Implications for siting energy development in sagebrush and prairie ecosystems
    Holbrook, Joseph D.; Olson, Lucretia E.; DeCesare, Nicholas J.; Hebblewhite, Mark; Squires, John R.; Steenweg, Robin, 2019. Functional responses in habitat selection: Clarifying hypotheses and interpretations
    Holbrook, Joseph D.; Squires, John R.; Bollenbacher, Barry; Graham, Russ; Olson, Lucretia E.; Hanvey, Gary; Jackson, Scott; Lawrence, Rick L.; Savage, Shannon L., 2019. Management of forests and forest carnivores: Relating landscape mosaics to habitat quality of Canada lynx at their range periphery
    Johnson, Tracey N.; Nasman, Kristen; Wallace, Zachary P.; Olson, Lucretia E.; Squires, John R.; Nielson, Ryan M.; Kennedy, Patricia L., 2019. Survey design for broad-scale, territory-based occupancy monitoring of a raptor: Ferruginous hawk (Buteo regalis) as a case study
    Franklin, Thomas W.; McKelvey, Kevin S.; Golding, Jessie; Mason, Daniel H.; Dysthe, Joseph; Pilgrim, Kristine L.; Squires, John R.; Aubry, Keith B.; Long, Robert A.; Greaves, Samuel; Raley, Catherine M.; Jackson, Scott; MacKay, Paula; Lisbon, Joshua; Sauder, Joel D.; Pruss, Michael T.; Heffington, Don; Schwartz, Michael K., 2019. Using environmental DNA methods to improve winter surveys for rare carnivores: DNA from snow and improved noninvasive techniques
    Miller, Sue; Olson, Lucretia E.; Squires, John R.; Roberts, Elizabeth; Ivan, Jake, 2019. Winter sports and wildlife: Can Canada lynx and winter recreation share the same slope?
    Heinemeyer, Kimberly; Squires, John R.; Hebblewhite, Mark; O’Keefe, Julia J.; Holbrook, Joseph D.; Copeland, Jeffrey, 2019. Wolverines in winter: Indirect habitat loss and functional responses to backcountry recreation
    Golding, Jessie; Schwartz, Michael K.; McKelvey, Kevin S.; Squires, John R.; Jackson, Scott D.; Staab, Cara; Sadak, Rema B., 2018. Multispecies mesocarnivore monitoring: USDA Forest Service multiregional monitoring approach
    Olson, Lucretia E.; Squires, John R.; Roberts, Elizabeth K.; Ivan, Jacob S.; Hebblewhite, Mark, 2018. Sharing the same slope: Behavioral responses of a threatened mesocarnivore to motorized and non-motorized winter recreation
    Holbrook, Joseph D.; Squires, John R.; Bollenbacher, Barry; Graham, Russell T.; Olson, Lucretia E.; Hanvey, Gary; Jackson, Scott; Lawrence, Rick L., 2018. Spatio-temporal responses of Canada lynx (Lynx canadensis) to silvicultural treatments in the Northern Rockies, U.S.
    Miller, Aubrey D.; Vaske, Jerry J.; Squires, John R.; Olson, Lucretia E., 2017. Does zoning winter recreationists reduce recreation conflict?
    Savage, Shannon L.; Lawrence, Rick L.; Squires, John R., 2017. Mapping post-disturbance forest landscape composition with Landsat satellite imagery
    Olson, Lucretia E.; Squires, John R.; Roberts, Elizabeth K.; Miller, Aubrey D.; Ivan, Jacob S.; Hebblewhite, Mark, 2017. Modeling large-scale winter recreation terrain selection with implications for recreation management and wildlife
    Olson, Lucretia E.; Squires, John R.; Oakleaf, Robert J.; Wallace, Zachary P.; Kennedy, Patricia L., 2017. Predicting above-ground density and distribution of small mammal prey species at large spatial scales
    Baigas, Phillip E.; Squires, John R.; Olson, Lucretia E.; Ivan, Jacob S.; Roberts, Elizabeth. K., 2017. Using environmental features to model highway crossing behavior of Canada lynx in the Southern Rocky Mountains
    Holden, Zachary A.; Swanson, Alan; Klene, Anna E.; Abatzoglou, John T.; Dobrowski, Solomon Z.; Cushman, Samuel A.; Squires, John R.; Moisen, Gretchen; Oyler, Jared W., 2016. Development of high-resolution (250 m) historical daily gridded air temperature data using reanalysis and distributed sensor networks for the US northern Rocky Mountains
    Wallace, Zachary P.; Kennedy, Patricia L.; Squires, John R.; Olson, Lucretia E.; Oakleaf, Robert J., 2016. Human-made structures, vegetation, and weather influence ferruginous hawk breeding performance
    Wallace, Zachary P.; Kennedy, Patricia L.; Squires, John R.; Oakleaf, Robert J.; Olson, Lucretia E.; Dugger, Katie M., 2016. Re-occupancy of breeding territories by ferruginous hawks in Wyoming: Relationships to environmental and anthropogenic factors
    Olson, Lucretia E.; Oakleaf, Robert J.; Squires, John R.; Wallace, Zachary P.; Kennedy, Patricia L., 2015. Nesting pair density and abundance of ferruginous hawks (Buteo regalis) and golden eagles (Aquila chrysaetos) from aerial surveys in Wyoming
    McKelvey, Kevin S.; Aubry, Keith B.; Anderson, Neil J.; Clevenger, Anthony P.; Copeland, Jeffrey P.; Heinemeyer, Kimberley S.; Inman, Robert M.; Squires, John R.; Waller, John S.; Pilgrim, Kristine L.; Schwartz, Michael K., 2014. Recovery of wolverines in the western United States: Recent extirpation and recolonization or range retraction and expansion?
    McKelvey, Kevin S.; Aubry, Keith B.; Anderson, Neil J.; Clevenger, Anthony P.; Copeland, Jeffrey P.; Heinemeyer, Kimberley S.; Inman, Robert M.; Squires, John R.; Waller, John S.; Pilgrim, Kristine L.; Schwartz, Michael K., 2014. Recovery of wolverines in the western United States: recent extirpation and recolonization or range retraction and expansion?
    Squires, John R.; Olson, Lucretia E.; Turner, David L.; DeCesare, Nicholas J.; Kolbe, Jay A., 2012. Estimating detection probability for Canada lynx Lynx canadensis using snow-track surveys in the northern Rocky Mountains, Montana, USA
    Berg, Nathan D.; Gese, Eric M.; Squires, John R.; Aubry, Lise M., 2012. Influence of forest structure on the abundance of snowshoe hares in western Wyoming
    Squires, John R.; DeCesare, Nicholas J.; Hebblewhite, Mark; Berger, Joel, 2012. Missing lynx and trophic cascades in food webs: A reply to Ripple et al.
    McKelvey, Kevin S.; Copeland, Jeffrey P.; Schwartz, Michael K.; Littell, Jeremy S.; Aubry, Keith B.; Squires, John R.; Parks, Sean A.; Elsner, Marketa M.; Mauger, Guillaume S., 2011. Climate change predicted to shift wolverine distributions, connectivity, and dispersal corridors
    Olson, Lucretia E.; Squires, John R.; DeCesare, Nicholas J.; Kolbe, Jay A., 2011. Den use and activity patterns in female Canada lynx (Lynx Canadensis) in the northern Rocky Mountains
    Fanson, Kerry V.; Wielebnowski, Nadja C.; Shenk, Tanya M.; Vashon, Jennifer H.; Squires, John R.; Lucas, Jeffrey R., 2010. Patterns of ovarian and luteal activity in captive and wild Canada lynx (Lynx canadensis)
    Fanson, Kerry V.; Wielebnowski, Nadja C.; Shenk, Tanya M.; Jakubas, Walter J.; Squires, John R.; Lucas, Jeffrey R., 2010. Patterns of testicular activity in captive and wild Canada lynx (Lynx canadensis)
    Squires, John R.; DeCesare, Nicholas J.; Kolbe, Jay A.; Ruggiero, Leonard F., 2010. Seasonal resource selection of Canada lynx in managed forests of the northern Rocky Mountains
    Copeland, J. P.; McKelvey, Kevin S.; Aubry, K. B.; Landa, A.; Persson, J.; Inman, R. M.; Krebs, J.; Lofroth, E.; Golden, H.; Squires, John R.; Magoun, A.; Schwartz, Michael K.; Wilmot, J.; Copeland, C. L.; Yates, R. E.; Kojola, I.; May, R., 2010. The bioclimatic envelope of the wolverine (Gulo gulo): do climatic constraints limit its geographic distribution?
    Schwartz, Michael K.; Aubry, Keith B.; McKelvey, Kevin S.; Pilgrim, Kristine L.; Copeland, Jeffrey P.; Squires, John R.; Inman, Robert M.; Wisely, Samantha M.; Ruggiero, Leonard F., 2007. Inferring geographic isolation of wolverines in California using historical DNA
    Kolbe, Jay A.; Squires, John R.; Pletscher, Daniel H.; Ruggiero, Leonard F., 2007. The effect of snowmobile trails on coyote movements within lynx home ranges
    Squires, John R.; Ruggiero, Leonard F., 2007. Winter prey selection of Canada lynx in northwestern Montana
    McKelvey, Kevin S.; von Kienast, Jeffrey; Aubry, Keith B.; Koehler, Gary M.; Maletzke, Bejamin T.; Squires, John R.; Lindquist, Edward L.; Loch, Steve; Schwartz, Michael K., 2006. DNA analysis of hair and scat collected along snow tracks to document the presence of Canada Lynx.
    Ulizio, Todd J.; Squires, John R.; Pletscher, Daniel H.; Schwartz, Michael K.; Claar, James J.; Ruggiero, Leonard F., 2006. The efficacy of obtaining genetic-based identifications from putative wolverine snow tracks
    DeCesare, Nicholas J.; Squires, John R.; Kolbe, Jay A., 2005. Effect of forest canopy on GPS-based movement data
    Boyce, Douglas A.; Kennedy, Patricia L.; Beier, Paul; Ingraldi, Michael F.; MacVean, Susie R.; Siders, Melissa S.; Squires, John R.; Woodbridge, Brian, 2005. When are goshawks not there? Is a single visit enough to infer absence at occupied nest areas? Journal of Raptor Research
    Squires, John R.; McKelvey, Kevin S.; Ruggiero, Leonard F., 2004. A snow-tracking protocol used to delineate local lynx, Lynx canadensis, distributions
    Kolbe, Jay A.; Squires, John R.; Parker, Thomas W., 2003. An effective box trap for capturing lynx
    Biek, Roman; Zarnke, Randall L.; Gillin, Colin; Wild, Margaret; Squires, John R.; Poss, Mary, 2002. Serologic survey for viral and bacterial infections in western populations of Canada Lynx (Lynx canadensis)
    McDaniel, Gregory W.; McKelvey, Kevin S.; Squires, John R.; Ruggiero, Leonard F., 2000. Efficacy of lures and hair snares to detect lynx
    Buskirk, Steven W.; Ruggiero, Leonard F.; Aubry, Keith B.; Pearson, Dean E.; Squires, John R.; McKelvey, Kevin S., 1999. Comparative ecology of lynx in North America [Chapter 14]
    Aubry, Keith B.; Ruggiero, Leonard F.; Squires, John R.; McKelvey, Kevin S.; Koehler, Gary M.; Buskirk, Steven W.; Krebs, Charles J., 1999. Conservation of lynx in the United States: A systematic approach to closing critical knowledge gaps [Chapter 17]
    Ruggiero, Leonard F.; Aubry, Keith B.; Buskirk, Steven W.; Koehler, Gary M.; Krebs, Charles J.; McKelvey, Kevin S.; Squires, John R., 1999. Ecology and conservation of lynx in the United States
    Aubry, Keith B.; Koehler, Gary M.; Squires, John R., 1999. Ecology of Canada lynx in southern boreal forests [Chapter 13]
    Ruggiero, Leonard F.; Aubry, Keith B.; Buskirk, Steven W.; Koehler, Gary M.; Krebs, Charles J.; McKelvey, Kevin S.; Squires, John R., 1999. The scientific basis for lynx conservation: Can we get there from here? [Chapter 18]
    Aubry, Keith B.; Buskirk, Steven W.; Koehler, Gary M.; Krebs, Charles J.; McKelvey, Kevin S.; Squires, John R., 1999. The scientific basis for lynx conservation: Qualified insights [Chapter 16]
    Squires, John R.; Reynolds, Richard T., 1997. Northern goshawk (Accipiter gentilis)
    Squires, John R.; Ruggiero, Leonard F., 1996. Nest-site preference of northern goshawks in southcentral Wyoming
    Squires, John R., 1995. Carrion use by northern goshawks
    Squires, John R.; Anderson, Stanley H., 1995. Trumpeter swan food habitats in the greater Yellowstone ecosystem
    Squires, John R.; Anderson, Stanley H.; Oakleaf, Robert., 1991. Prairie falcons quit nesting in response to spring snowstorm
    Squires, John R.; Anderson, Stanley H.; Oakleaf, Robert, 1989. Food habits of nesting prairie falcons in Campbell County
    A lynx wearing a radio collar walking in snow.
    Canada lynx depend on boreal and subalpine forests that have been structured by natural disturbances for millennia.  The management conundrum is how to salvage beetle-killed trees, while also conserving this iconic species.  We instrumented Canada lynx with GPS collars to learn how they used beetle-impacted forests.  Our research informed how to balance timber salvage with species conservation.
    A suspected lynx has triggered this photo from a game camera in Montana (photo credit: Don Heffington).
    A new project showed that animal footprints in snow contain enough DNA for species identification, even when the snow was many months old. The study extracted DNA from snow samples collected within animal tracks as well as areas where the animal had been photographed months earlier. Newly developed genetic assays were applied and positively detected the DNA of each species, performing nearly flawlessly on samples previously considered too poor to provide usable DNA. This method could revolutionize winter surveys of rare species by greatly reducing or eliminating misidentifications and missed detections.
    A closeup shot of a lynx face
    The management of Canada lynx habitat is an issue that has generated much debate and litigation across the Northern (Montana, Idaho) and Southern (Colorado, Wyoming) Rocky Mountains. This species depends almost exclusively on snowshoe hare for food during winter, and this prey species is sensitive to changes in forest composition and structure. Research conducted by scientists at the Rocky Mountain Research Station, in collaboration with universities and local forest managers, is central in resolving management impasses by learning how changes in forest structure and composition can be implemented in ways that enhance the ability of Canada lynx to produce kittens.  
    Remote camera captures a wolverine as it approaches a researcher's trap.
    Forest Service scientists and their research partners use a novel approach that includes trapping and fitting wolverines with GPS collars that accurately plot their movements in areas of high winter recreation. Thenvolunteer snowmobilers, back-country skiers, and other recreationists carry GPS units in the same areas used by wolverines. Resulting data show how wolverines respond to winter recreation in terms of their movements, behaviors, and resource-use.
    Lynx kitten from a female in spruce-beetle kill
    Spruce-bark beetles impacted about 480,000 acres of spruce-fir forests in southern Colorado and are spreading at the rate of 100,000 acres annually.  A central question is how to salvage for timber production insect-impacted forests in ways consistent with the management and conservation of Canada lynx, a federally-listed species.
    A wolverine
    The U.S. Fish and Wildlife Service is examining the wolverine (Gulo gulo) as a candidate for listing as a threatened or endangered species. RMRS researchers are investigating suitable habitats for wolverine reintroduction efforts, and have found ways to apply models derived from current genetic patterns to future landscapes to inform land management decisions on existing and future corridor locations. While current efforts are focused on wolverines, these newly developed tools can be applied to a variety of organisms to inform their potential future in a changing climate.
    Ferruginous hawk instrumented with a solar GPS transmitter.
    Over the past decade and a half, raptors nesting in prairie ecosystems have been subject to sharp increases in nearby energy development activity. This research documents how nesting ferruginous hawks forage in oil and gas energy fields based on GPS telemetry. The purpose is to help managers and companies reflect conservation needs of this species in the management and arrangement of energy-development infrastructure.   
    Opportunity and Vision
    Land managers require high-quality information on species and habitats at risk to develop effective management strategies. In the absence of information on these species and their habitats, agencies frequently err on the side of the species and make conservative, and often unnecessary, decisions relative to habitat protection. Over 20 years of research by scientists with the Rocky Mountain Research Station are helping address these information needs.
    Canada lynx, and their primary prey snowshoe hares, live in high-elevation spruce-fir forests, which are increasingly modified by spruce-bark beetle outbreaks. The goal of our research is to combine lynx use of insect-impacted forests with measures of forest condition.  Our results will inform forest prescriptions that facilitate timber-salvage and lynx conservation.
    By 2013, a spruce beetle outbreak impacted 85% of the mature spruce-fir forests on the Rio Grande National Forest. These spruce-fir forests provided some of the highest quality lynx habitat in the state. The goal of this project is to research the forest structures and compositions that lynx and snowshoe hare depend within landscapes altered by spruce bark beetle outbreak, in relation to increased post-beetle forest management activities from timber salvage.