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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 research team is responsible for the discovery of new information needed to manage and conserve threatened, endangered, and sensitive species throughout the Rocky Mountain Region.  We consider the scientific and practical challenges of managing TES wildlife within a multi-use land-management context based on empirical field data.  Current studies include:

  • Determine how the demography of Canada lynx is affected by landscape mosaics in upper montane forests;
  • Study lynx movements relative to landscape pattern;
  • Determine resource selection of Canada lynx in the Northern Rockies at multiple scales;
  • Document resource selection and spatial-use patterns of Canada lynx in spruce beetle-impacted forests of southern Colorado;
  • Investigate behavior and resource-use patterns of lynx and wolverines to motorized and non-motorized recreation during winter;
  • Assess the population viability of lynx in the Northern Rockies and relate to prey abundance;
  • Document fine-scale genetic sub-structure of lynx in the Northern Rockies;
  • Delineate the distribution of lynx and other carnivores in the Greater Yellowstone Area;
  • Determine how lynx and their primary prey, snowshoe hares, respond to increase fire disturbance in the Northern Rockies and associated post-fire treatments.

I also lead an integrated research program that investigates how ferruginous hawks and other prairie raptors respond to energy development in sage-steppe ecosystems. Research topics include:

  • Occupancy modeling of ferruginous hawks;
  • Density and detection of ferruginous hawks and golden eagles;
  • Landscape genetics of ferruginous hawks in North America;
  • Movement and spatial use of ferruginous hawks relative to energy infrastructure based on GPS telemetry;
  • Resource selection of ferruginous hawks and golden eagles in sage steppe;
  • Raptor population monitoring protocols given field-estimated detection probabilities and occupancy.

Research Interests

My current research interest concerns the management and conservation of Canada lynx, wolverines, and other sensitive species. Lynx and wolverine are highly mobile and depend on broad landscapes to meet their resource needs. Thus, my interest includes multi-scale evaluations of resource selection, forest carnivore movements and connectivity, factors affecting population viability, the effects of forest management, recreation and other human-induced impacts on species persistence, developing detection and monitoring methods for forest carnivores, and determining the effects of climate change on lynx and wolverine. I also have had a life-long interest in raptors and their management, and I am interested in helping agencies and industry develop energy reserves in ways that also facilitate raptor conservation.  

My primary research themes include:

  • Ecology of Threatened, Endangered, and Sensitive species
  • Climate change and disturbance ecology
  • Forest silviculture with repect to Canada lynx conservation and mangement
  • Human-wildlife interactions
  • Carnivore and raptor monitoring
  • Highways and wildlife
  • Population and landscape genetic connectivity

Past Research

Throughout my career I have been interested in the conservation and management of threatened/endangered/sensitive species.  My past research focused on habitat-use relationships of avian and mammalian species in sage steppe and forested ecosystems.  I approach wildlife research from the perspective of collecting new empirical information from field studies that provide direct measures of animal movement and resource-use.  In the late 1980’s, I investigated trumpeter swan behavior and resource-use in the Greater Yellowstone Areas, especially in and adjacent to Teton National Park.  In the early 1990’s, I investigated nest selection and winter movements of northern goshawks.  In the late 1990’s, I moved to Missoula, MT to initiate lynx studies in western Montana.  In early 2000’s, I expanded my research to investigate wolverines in southcentral Montana.  My studies focused on: the management and conservation needs of northern goshawks; patterns of wolverine mortality; seasonal evaluations of resource-use, denning, movements, and population connectivity of Canada lynx; and the development of detection and monitoring methods for mid-sized carnivores. 

Why This Research is Important

The conservation and management needs of mid-sized forest carnivores, like lynx and wolverine, were poorly understood for populations in the continental United States when I initiated my research in 1997.  Land managers were in the difficult position of prescribing management actions without knowing impacts to these species. Forest carnivores are elusive and difficult to study so there still remains much to learn today. However, results from our studies informed land management across millions of acres of federal land in ways that balanced multi-use land management with forest carnivore conservation.  We continue to provide new empirical information that is highly relevant to forest silviculture, fire management, recreation, species monitoring, and energy development.  Winter recreation is a major industry that influences local economies across the western United States.  The information from my studies directly address how lynx and wolverines respond to winter recreation so that recreation planners can promote winter outdoor sports in ways that minimize potential impacts to these sensitive carnivores.

The severity and frequency of high-elevation wildfires in lynx habitat increased sharply in the last decade across the western United States. Fire disturbance is expected to increase with climate change in much of the West, is now a major issue affecting the extent and distribution of lynx habitat in the Northern Rocky Mountians.  Given the scale of fire and insect outbreaks in lynx habitat, our results are central to determining the ways that forest silviculture and management can provide tools that speed habitat recovery following these disturbances.

Although the public supports increased energy independence, the scale of energy development potentially impacts many sensitive species that depend on sage steppe ecosystems from direct disturbance or habitat loss.  Our research is important because we provide empirical data regarding how ferruginous hawks and golden eagles respond to increased energy development.  This information is central to identifying appropriate mitigation that reduces energy-related impacts to raptors, as we further develop domestic energy reserves. 


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


    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; 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
    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?
    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)
    Buskirk, Steven W.; Ruggiero, Leonard F.; Aubry, Keith B.; Pearson, Dean E.; Squires, John R.; McKelvey, Kevin S., 2000. 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., 2000. Conservation of lynx in the United States: A systematic approach to closing critical knowledge gaps [Chapter 17]
    Aubry, Keith B.; Koehler, Gary M.; Squires, John R., 2000. Ecology of Canada lynx in southern boreal forests [Chapter 13]
    McDaniel, Gregory W.; McKelvey, Kevin S.; Squires, John R.; Ruggiero, Leonard F., 2000. Efficacy of lures and hair snares to detect lynx
    Ruggiero, Leonard F.; Aubry, Keith B.; Buskirk, Steven W.; Koehler, Gary M.; Krebs, Charles J.; McKelvey, Kevin S.; Squires, John R., 2000. 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., 2000. The scientific basis for lynx conservation: Qualified insights [Chapter 16]
    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
    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 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.   
    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.