Winter Habitat Selection by Canada Lynx in Maine: Prey Abundance or Accessibility?

Abstract: We related winter habitat selection by Canada lynx (Lynx canadensis), relative abundance of snowshoe hares (Lepus americanus), and understory stem densities to evaluate whether lynx select stands with the greatest snowshoe hare densities or the greatest prey accessibility. Lynx (3 F, 3 M) selected tall (4.4-7.3 m) regenerating clear-cuts (11-26 yr postharvest) and established partially harvested stands (11-21 yr postharvest) and selected against short (3.4-4.3 m) regenerating clear-cuts, recent partially harvested stands (1-10 yr), mature second-growth stands (>40 yr), and roads and their edges (30 m on either side of roads). Lynx selected stands that provided intermediate to high hare density and intermediate cover for hares (i.e., prey access) but exhibited lower relative preference for stand types with highest hare densities where coniferous saplings exceeded 14,000 stems/ha.     

Canada lynx occurrence and forest management in the Acadian Forest

We evaluated patterns of occurrence and non-occurrence for Canada lynx (Lynx canadensis) across a 16,530-km² study area in Maine to provide a better understanding of lynx habitat selection and habitat ecology on commercially managed forestlands in the Acadian Forest. Because of the influence of forest structure on lynx habitat selection and abundance of their primary prey, the snowshoe hare (Lepus americanus), and to improve our ability to build robust models, we used habitat information derived from a time series of Landsat satellite imagery spanning the period 1973–2004. We defined and mapped 10 forest types based on forest harvest history, time since harvest, and current forest condition. We compared a suite of models to evaluate relative influences of forest composition, habitat patch configuration, and hare density on habitat selection by lynx at the landscape scale. Occupied areas had greater average hare densities and percentage of mature conifer. Average hare density in occupied areas (0.74 hares/ha) was greater than in unoccupied areas (0.62 hares/ha), but was less than previous research has suggested may be necessary to support lynx populations in the southern portion of the species' range. No occupied areas occurred where average hare density was <0.5 hares/ha. Average hare density at the landscape-scale was strongly influenced by amount of high-quality hare habitat (i.e., conifer or mixedwood regenerating forest, 15–35 yr post-harvest). Edge density between mature conifer and high-quality hare habitat was substantially greater in occupied areas compared to unoccupied areas. Juxtaposition of those 2 forest types may provide edge habitat where lynx experience easier travel and improved access to prey in landscapes with extensive areas of high-quality hare habitat where travel and access may be somewhat limited by high understory stem density. Probability of occurrence declined nonlinearly with changes in hare density and percent mature conifer forest in the landscape; thus, suitability of currently occupied landscapes could change markedly with future changes in landscape-level hare densities and changing habitat associated with forest management. Where lynx conservation is a priority, we recommend that managers focus on creating and maintaining a minimum of 27% high-quality hare habitat within 100-km² areas to promote landscape-scale hare densities >0.5 hares/ha. © The Wildlife Society, 2013     

Habitat Conditions Associated With Lynx Hunting Behavior During Winter in Northern Washington

Abstract Effectively managing habitat for threatened populations of Canada lynx (Lynx canadensis) requires knowledge of habitat conditions that provide for the ecological needs of lynx. We snow-tracked lynx to identify habitat conditions associated with hunting behavior and predation during winters of 2002–2003 and 2003–2004 in the northern Cascade Range in Washington state, USA. We recorded number and success of predation attempts, prey species killed, and trail sinuosity on 149 km of lynx trails. Lynx killed snowshoe hares (Lepus americanus), red squirrels (Tamiasciurus hudsonicus), and cricetids more than expected in Englemann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) forests, where snowshoe hare densities were highest. Lynx killed prey less than expected in Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) forests and forest openings. We used the sinuosity of lynx trails as an index of quality of habitat hunted. Lynx trails that included predation attempts were more sinuous than trail segments without predation attempts. Lynx trails had greater sinuosity in forest stands with high hare densities dominated by Engelmann spruce and subalpine fir than in stands with low hare densities dominated by Douglas-fir and ponderosa pine or in forest openings. We encourage forest managers to maintain or create sufficient understory cover to support high densities of snowshoe hares as foraging habitat for lynx.     

Within-Stand Selection of Canada Lynx Natal Dens in Northwest Maine,USA

Abstract Canada lynx (Lynx canadensis) were listed as threatened in the contiguous United States under the Endangered Species Act in March 2000. Little information on lynx ecology at the southern extent of their range was available at the time of listing, and no ecological studies had been conducted in the eastern USA. Between 1999 and 2004, we investigated habitat selection at natal dens in northern Maine to address questions on the importance of forest conditions to denning requirements. We compared within-stand characteristics of 26 den sites to general characteristics of the stands containing dens. We used logistic regression to identify components within stands that distinguished natal dens from the residual stand and used the information-theoretic approach to select models that best explained lynx den-site selection. The top-ranked model had 2 variables: tip-up mounds of blown-down trees and visual obscurity at 5 m from the den (w_i = 0.92). Within-stand structure was useful for predicting lynx den-site selection in managed forests in Maine and suitable denning habitat did not appear to be limiting.     

Winter Prey Selection of Canada Lynx in Northwestern Montana

ABSTRACT The roles that diet and prey abundance play in habitat selection of Canada lynx (Lynx canadensis) in the contiguous United States is poorly understood. From 1998–2002, we back-tracked radiocollared lynx (6 F, 9 M) for a distance of 582 km and we located 86 kills in northwestern Montana, USA. Lynx preyed on 7 species that included blue grouse (Dendragapus obscurus), spruce grouse (Canachites canadensis), northern flying squirrel (Glaucomys sabrinus), red squirrel (Tamiasciurus hudsonicus), snowshoe hare (Lepus americanus), least weasel (Mustela nivalis), and white-tailed deer (Odocoileus virginianus). Snowshoe hares (69 kills) accounted for 96% (4-yr average, range = 94–99%) of prey biomass during the sample period. Red squirrels were the second-most-common prey (11 kills), but they only provided 2% biomass of the winter diet. Red squirrels contributed little to the lynx diet despite low hare densities. A logistic regression model of snowshoe hare, red squirrel, and grouse abundance, as indexed by the number of track crossings of use and available lynx back-tracks, was a significant (Wald statistic = 19.03, df = 3, P < 0.001) predictor of habitat use. As we expected, lynx (P < 0.001) selected use-areas with higher snowshoe hare abundance compared to random expectation. However, the red squirrel index had a weak (P = 0.087) negative relationship to lynx use, and grouse was nonsignificant (P = 0.432). Our results indicate that lynx in western Montana prey almost exclusively on snowshoe hares during the winter with little use of alternative prey. Thus, reductions in horizontal cover for hares would degrade lynx habitat.     

Habitat Fragmentation and the Persistence of Lynx Populations in Washington State

Abstract Lynx (Lynx canadensis) occur in the northern counties of Washington state, USA; however, current distribution and status of lynx in Washington is poorly understood. During winters 2002–2004 we snow-tracked lynx for 155 km within a 211-km² area in northern Washington, to develop a model of lynx-habitat relationships that we could use to assess their potential distribution and status in the state. We recorded movements and behaviors of lynx with a Global Positioning System and overlaid digitized lynx trails on various habitat layers using a Geographic Information System. Based on univariate analyses, lynx preferred Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) forests, with moderate canopy and understory cover, and elevations ranging from 1,525 m to 1,829 m but avoided Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) forests, openings, recent burns, open canopy and understory cover, and steep slopes. A map of suitable lynx habitat based on a logistic regression model built using these candidate variables revealed that habitats at elevations >1,400 m where lynx historically occurred in Washington are intersected and fragmented by landscape features and forest conditions that are generally avoided by lynx. Our habitat suitability map predicts 3,800 km² of lynx habitat in Washington that could support 87 lynx, far fewer than previous estimates. Since 1985, natural fires have burned >1,000 km² of forested habitat in Okanogan County, the only region in Washington where lynx occurrence has been documented during that period. Loss of suitable habitat from natural and human-caused disturbances, and the lack of verifiable evidence of lynx occurrence in historic lynx range, suggests that fragmented landscape conditions may have impeded recolonization of these areas by lynx. Consequently, translocations may be necessary to ensure lynx persistence in Washington. We suggest that managers assess the potential for translocation by first identifying the scale and distribution of potential foraging habitats for lynx based on our or similar habitat models, survey various habitat conditions to obtain reliable estimates of snowshoe hare densities, and identify a genetically compatible source population of lynx. If habitat and source populations are adequate, reintroducing lynx to areas of their historic range may be an appropriate conservation strategy.     

Circadian Activity Patterns of Canada Lynx in Western Montana

Abstract: We recorded activity data for 6 male and 5 female lynx in winter and 3 male and 6 female lynx during summer in western Montana, USA, using motion-sensitive radiocollars. Lynx diel activity appeared to vary by sex, season, and reproductive status. During summer, male lynx exhibited a crepuscular activity pattern, whereas females with kittens remained active throughout the photoperiod. During winter, lynx of both sexes were most active during the afternoon and early evening. Although we observed no evidence that predation risk drove lynx activity patterns, such patterns may be a function of kitten-rearing activity, thermoregulatory strategies, and foraging behavior.     

Spatial Ecology of a Canada Lynx Population in Northern Maine

Abstract Canada lynx (Lynx canadensis) were listed as a federally threatened species in 14 states at the southern extent of their geographic range in March 2000, with Maine being the only state in the northeastern United States known to support a resident population. Relatively little information is known about the ecology of lynx living at the southern edge of their range, including range requirements, movements, and spatial organization. Basic knowledge of lynx ecology is needed for federal recovery planning efforts. Between 1999 and 2004, we trapped and radiocollared 43 lynx (21 M, 22 F) in northern Maine in an intensively managed and predominantly early successional forested landscape. We estimated diurnal annual and seasonal home-range size for male and female lynx using the 85% fixed-kernel home-range estimator. Annual home ranges of adult male lynx (x̄ = 53.6 km²) were more than twice the size of adult female home ranges (x̄ = 25.7 km²). Home ranges of adult females during snow periods (x̄ = 38.3 km²) were nearly 3 times larger than their snow-free-period ranges (x̄ = 14.3 km²), whereas, snow-free ranges of adult males (x̄ = 58.8 km²) were slightly larger than their snow-period ranges (x̄ = 45.2 km²). We observed a limited amount of home-range overlap among lynx of the same sex (F: x̄ = 17.2%; M: x̄ = 11.8%). Lynx of opposite sex showed more extensive overlap (x̄ = 24.3%). Most home-range shifts of resident lynx were typically not extensive. Based on territory mapping, we estimated a minimum lynx density of 9.2–13.0 lynx/100 km². We observed lynx spatial ecology and densities that were more similar to northern lynx populations when hares were abundant than to other southern lynx populations, suggesting that region-specific studies under varying habitat conditions and hare densities are needed to ensure realistic recovery goals and effective management of lynx at the southern extent of their range.     

Seasonal Resource Selection of Canada Lynx in Managed Forests of the Northern Rocky Mountains

ABSTRACT We investigated seasonal patterns in resource selection of Canada lynx (Lynx canadensis) in the northern Rockies (western MT, USA) from 1998 to 2002 based on backtracking in winter (577 km; 10 M, 7 F) and radiotelemetry (630 locations; 16 M, 11 F) in summer. During winter, lynx preferentially foraged in mature, multilayer forests with Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) in the overstory and midstory. Forests used during winter were composed of larger diameter trees with higher horizontal cover, more abundant snowshoe hares (Lepus americanus), and deeper snow compared to random availability; multilayer, spruce-fir forests provided high horizontal cover with tree branching that touched the snow surface. During winter, lynx killed prey at sites with higher horizontal cover than that along foraging paths. Lynx were insensitive to snow depth or penetrability in determining where they killed prey. During summer, lynx broadened their resource use to select younger forests with high horizontal cover, abundant total shrubs, abundant small-diameter trees, and dense saplings, especially spruce-fir saplings. Based on multivariate logistic-regression models, resource selection occurred primarily at a fine spatial scale as was consistent with a sight-hunting predator in dense forests. However, univariate comparisons of patch-level metrics indicated that lynx selected homogenous spruce-fir patches, and avoided recent clear-cuts or other open patches. Given that lynx in Montana exhibit seasonal differences in resource selection, we encourage managers to maintain habitat mosaics. Because winter habitat may be most limiting for lynx, these mosaics should include abundant multistory, mature spruce-fir forests with high horizontal cover that are spatially well-distributed.     

Effects of Precommercial Thinning on Snowshoe Hares in Maine

Abstract: Snowshoe hares (Lepus americanus) are an important prey species and a dominant herbivore across much of their North American range, and researchers have questioned the influences of forestry practices that alter habitat for hares and the potential community-level effects on carnivores. We examined the effects of precommercial thinning (PCT) from 1 to 11 years posttreatment on snowshoe hares. In the commercial forests of northern Maine, USA, we counted and cleared hare pellets twice a year during 2001 and 2002 on >46 km of pellet transects across 30 regenerating conifer stands (17 treated with PCT) previously treated with an aerial application of herbicide. We compared densities of snowshoe hare pellets among 3 development classes with (1 yr after thinning, 6 yr after thinning, and 11 yr after thinning) and without thinning (stands with a similar history of clearcut and herbicide treatment but no thinning). During both years, densities of hares were lower in stands treated with PCT than in similar unthinned stands across the 3 development classes and during both leaf-off and leaf-on seasons (P < 0.001). Within both thinned and unthinned stands, hare density was greatest in stands in the 1-year development class when compared to the 6-year and 11-year development classes, but a statistical difference (P = 0.048) among classes was evident only during leaf-off seasons. Precommercial thinning was associated with densities of snowshoe hares that were approximately half of those in similar unthinned stands up to at least 11 years posttreatment; however, thinned stands may retain densities of hares greater than stands managed using other forest harvesting regimes. Our results apply to core portions of stands with crop trees spaced at 1.8–2.4-m intervals following complete overstory removal and herbicide treatment. We advocate caution when applying our results to other thinning regimes or across broader spatial scales.     

Hierarchical Den Selection of Canada Lynx in Western Montana

Abstract We studied den selection of Canada lynx (Lynx canadensis; hereafter lynx) at multiple ecological scales based on 57 dens from 19 females located in western Montana, USA, between 1999 and 2006. We considered 3 spatial scales in this analysis, including den site (11-m-radius circle surrounding dens), den area (100-m-radius circle), and den environ (1-km radius surrounding dens). Lynx denned in preexisting sheltered spaces created by downed logs (62%), root-wads from wind-thrown trees (19%), boulder fields (10%), slash piles (6%), and live trees (4%). Lynx preferentially selected den sites with northeasterly aspects that averaged 248. Average distance between dens of 13 females monitored in consecutive years was 2,248 m, indicating low den site fidelity. Lynx exhibited habitat selection at all 3 spatial scales. Based on logistic regression, den sites differed from the surrounding den areas in having higher horizontal cover and log volume. Abundant woody debris from piled logs was the dominant habitat feature at den sites. Lynx generally denned in mature spruce-fir (Picea-Abies) forests with high horizontal cover and abundant coarse woody debris. Eighty percent of dens were in mature forest stands and 13% in mid-seral regenerating stands; young regenerating (5%) and thinned (either naturally sparse or mechanically thinned) stands with discontinuous canopies (2%) were seldom used. Female lynx selected den areas with greater spruce-fir tree basal area, higher horizontal cover, and larger-diameter trees compared to random locations within their home range. Lynx selected den environs in topographically concave or drainage-like areas, and farther from forest edges than random expectation. Maintaining mature and mid-seral regenerating spruce-fir forests with high horizontal cover and abundant woody debris would be most valuable for denning when located in drainages or in concave, drainage-like basins. Management actions that alter spruce-fir forests to a condition that is sparsely stocked (e.g., mechanically thinned) and with low canopy closure (<50%) would create forest conditions that are poorly suitable for lynx denning.     

Assessment of Canada Lynx Research and Conservation Needs in the Southern Range: Another Kick at the Cat

Abstract The ecology of Canada lynx (Lynx canadensis) and their main prey, snowshoe hares (Lepus americanus), is poorly understood in southern Canada and the contiguous United States compared to the boreal forest of Canada and Alaska, USA, where both species are well studied. However, given recent listing of lynx under the Endangered Species Act, accurate understanding of lynx and snowshoe hare ecology and conservation requirements in the United States is a high priority. We critically examined unchallenged perceptions and important research needs related to lynx and hare ecology and conservation at the southern extent of their range. Contrary to popular dogma, lynx do not require old-growth forest for denning, but further research on lynx and hare use of fragmented landscapes at lower latitudes is required. The contention that southern lynx are subject to higher interference or exploitative competition compared to their northern counterparts remains without strong empirical support. Lynx rely more on red squirrels (Tamiasciurus hudsonicus) and possibly other alternate prey at lower latitudes, but hares are the predominant food type for lynx across their range. Southern lynx and hare populations do not exhibit periodic cyclicity, but harvest statistics suggest that lynx abundance in the southern range is highly variable, implying that numerical fluctuations likely are fueled by immigration from Canada. Southern lynx population viability in the absence of ingress is suspect and thus maintaining connectivity with northern areas of occupancy should be a priority. Successful conservation of lynx populations in the contiguous United States will require 1) improved understanding of lynx population and habitat ecology at lower latitudes, 2) protection and management of large tracts of lynx and snowshoe hare habitat, and 3) ensured connectivity between lynx populations at the core and periphery of the species' range. However, in light of the numerous challenges facing conservation of populations of many species at their southern distributional limit, the long-term prognosis for lynx in the southern range currently is uncertain.     

Movement and Habitat Use of Canada Lynx During Denning in Minnesota

Abstract Establishing whether conditions are suitable for reproduction would help determine if immigration is necessary for Canada lynx (Lynx canadensis) to persist at the southern edge of the species range. We located den sites and monitored reproduction of radiocollared lynx in Minnesota from 2004 to 2006. Movement rates of denning females measured with Global Positioning System collars were similar to movement rates of lynx elsewhere. Female lynx with kittens used different habitat types in predenning, denning, and postdenning periods. Landscape composition at the scale of the foraging radius around a den site contained more lowland conifer, upland conifer, and regenerating forest than did home ranges or the area used by radiocollared lynx in Minnesota, USA. We used the spatial distribution of cover-type composition around known den sites to predict and map potential denning habitat in northeastern Minnesota. Techniques for identifying the spatial distribution of suitable denning habitat provide a biological basis for management actions that could enhance recovery of Canada lynx populations in the southern part of the species range.     

Special Section on Canada Lynx-Habitat Relations in the Contiguous United States

Abstract The southern extent of the range of Canada lynx (Lynx canadensis) extends into the contiguous United States in locations from Washington State in the West to northern Maine in the East. Lynx persist in various habitats across this range from high-elevation wilderness to intensively managed industrial forests. Lynx habitat use at the species' southern range boundary was poorly understood before the species was listed as Threatened under the Endangered Species Act in 2000, because most research had been conducted in boreal forest. The papers in this special section outline a variety of questions regarding lynx populations at the southern extent of their range, address topics on lynx space use and denning habitat, and provide generally consistent results in terms of features important to lynx habitat use despite the wide variation in specific habitats among the study areas.