John R. Squires

Wolverine gene flow across a narrow climatic niche

Wolverines (Gulo gulo) are one of the rarest carnivores in the contiguous United States. Effective population sizes in Montana, Idaho, and Wyoming, where most of the wolverines in the contiguous United States exist, were calculated to be 35 (credible limits, 28 52) suggesting low abundance. Landscape features that influence wolverine population substructure and gene flow are largely unknown. Recent work has identified strong associations between areas with persistent spring snow and wolverine presence and range. We tested whether a dispersal model in which wolverines prefer to disperse through areas characterized by persistent spring snow cover produced least-cost paths among all individuals that correlated with genetic distance among individuals. Models simulating large preferences for dispersing within areas characterized by persistent spring snow explained the data better than a model based on Euclidean distance. Partial Mantel tests separating Euclidean distance from spring snow-cover-based effects indicated that Euclidean distance was not significant in describing patterns of genetic distance. Because these models indicated that successful dispersal paths followed areas characterized by spring snow cover, we used these understandings to derive empirically based least-cost corridor maps in the U.S. Rocky Mountains. These corridor maps largely explain previously published population subdivision patterns based on mitochondrial DNA and indicate that natural colonization of the southern Rocky Mountains by wolverines will be difficult but not impossible.

The bioclimatic envelope of the wolverine (Gulo gulo): do climatic constraints limit its geographic distribution?

We propose a fundamental geographic distribution for the wolverine (Gulo gulo (L., 1758)) based on the hypothesis that the occurrence of wolverines is constrained by their obligate association with persistent spring snow cover for successful reproductive denning and by an upper limit of thermoneutrality. To investigate this hypothesis, we developed a composite of MODIS classified satellite images representing persistent snow cover from 24 April to 15 May, which encompasses the end of the wolverine’s reproductive denning period. To investigate the wolverine’s spatial relationship with average maximum August temperatures, we used interpolated temperature maps. We then compared and correlated these climatic factors with spatially referenced data on wolverine den sites and telemetry locations from North America and Fennoscandia, and our contemporary understanding of the wolverine’s circumboreal range. All 562 reproductive dens from Fennoscandia and North America occurred at sites with persistent spring snow c...

Effect of forest canopy on GPS‐based movement data

Abstract The advancing role of Global Positioning System (GPS) technology in ecology has made studies of animal movement possible for larger and more vagile species. A simple field test revealed that lengths of GPS-based movement data were strongly biased (P<0.001) by effects of forest canopy. Global Positioning System error added an average of 27.5% additional length to tracks recorded under high canopy, while adding only 8.5% to open-canopy tracks, thus biasing comparisons of track length or tortuosity among habitat types. Other studies may incur different levels of bias depending on GPS sampling rates. Ninety-nine percent of track errors under high canopy were ≤7.98 m of the true path; this value can be used to set the scale-threshold at which movements are attributed to error and not biologically interpreted. This bias should be considered before interpreting GPS-based animal movement data.

Climate change predicted to shift wolverine distributions, connectivity, and dispersal corridors

Boreal species sensitive to the timing and duration of snow cover are particularly vulnerable to global climate change. Recent work has shown a link between wolverine (Gulo gulo) habitat and persistent spring snow cover through 15 May, the approximate end of the wolverines reproductive denning period. We modeled the distribution of snow cover within the Columbia, Upper Missouri, and Upper Colorado River Basins using a downscaled ensemble climate model. The ensemble model was based on the arithmetic mean of 10 global climate models (GCMs) that best fit historical climate trends and patterns within these three basins. Snow cover was estimated from resulting downscaled temperature and precipitation patterns using a hydrologic model. We bracketed our ensemble model predictions by analyzing warm (miroc 3.2) and cool (pcm1) downscaled GCMs. Because Moderate-Resolution Imaging Spectroradiometer (MODIS)-based snow cover relationships were analyzed at much finer grain than downscaled GCM output, we conducted a second analysis based on MODIS-based snow cover that persisted through 29 May, simulating the onset of spring two weeks earlier in the year. Based on the downscaled ensemble model, 67% of predicted spring snow cover will persist within the study area through 2030-2059, and 37% through 2070-2099. Estimated snow cover for the ensemble model during the period 2070- 2099 was similar to persistent MODIS snow cover through 29 May. Losses in snow cover were greatest at the southern periphery of the study area (Oregon, Utah, and New Mexico, USA) and least in British Columbia, Canada. Contiguous areas of spring snow cover become smaller and more isolated over time, but large (.1000 km 2 ) contiguous areas of wolverine habitat are predicted to persist within the study area throughout the 21st century for all projections. Areas that retain snow cover throughout the 21st century are British Columbia, north-central Washington, northwestern Montana, and the Greater Yellowstone Area. By the late 21st century, dispersal modeling indicates that habitat isolation at or above levels associated with genetic isolation of wolverine populations becomes widespread. Overall, we expect wolverine habitat to persist throughout the species range at least for the first half of the 21st century, but populations will likely become smaller and more isolated.

DNA Analysis of Hair and Scat Collected Along Snow Tracks to Document the Presence of Canada Lynx

Abstract Snow tracking is often used to inventory carnivore communities, but species identification using this method can produce ambiguous and misleading results. DNA can be extracted from hair and scat samples collected from tracks made in snow. Using DNA analysis could allow positive track identification across a broad range of snow conditions, thus increasing survey accuracy and efficiency. We investigated the efficacy of DNA identification using hairs and scats collected during the winter along putative Canada lynx (Lynx canadensis) snow tracks and compared our findings to those obtained using hair-snaring techniques during the summer. We were able to positively identify 81% and 98% of the hair and scat samples, respectively, that were collected in or near snow tracks. Samples containing amplifiable lynx DNA were collected at rates of 1.2–1.3 per km of lynx tracks followed. These amplification rates and encounter frequencies validate the collection and use of DNA samples from snow tracks as a feasible technique for identifying Canada lynx and possibly other rare carnivores. We recommend that biologists include the collection of hairs and scats for DNA analysis as part of snow-tracking surveys whenever species identification is a high priority.

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.

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.

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 24°. 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.

Nest-site preference of northern goshawks in southcentral Wyoming

In 1992, we studied the nest-site preference of goshawks (Accipiter gentilis atricapillus) nesting in lodgepole pine (Pinus contorta) forests of the Medicine Bow National Forest, southcentral Wyoming. For 39 active pairs of goshawks, we described nesting habitat at 3 spatial scales: nest tree, nest-tree area (0.04 ha circle centered at nest tree), and nest stand (homogeneous forest stand surrounding nest). Nest stands ranged from 0.4 ha to 13.0 ha (x = 2.7 ha, SE = 0.4). We compared habitat characteristics at nest-sites to those measured at random sites. The mean diameter at breast height (dbh) of nest trees was larger (P < 0.001) than the mean dbh of trees in either the nest-tree area or the nest stand. Nest trees also were taller (P < 0.001) and had greater dbh (P < 0.001) than trees in random stands. Slopes at goshawk nests were more (P = 0.04) moderate (x = 11%, SE 1.1, range 1 to 34%) compared to those available. Aspects at goshawk nests were similar (P = 0.61) to those available. The tree density in goshawk nest stands was lower (P = 0.045) than random stands. However, nest stands had a higher (P < 0.001) density of large trees compared to random stands. Trees in nest stands also were taller (P < 0.001) with greater (P = 0.006) heights to live canopy compared to trees in random stands. The mean density of small trees at nest stands was less than (P = 0.001) one-half those present in random stands. Nest stands were not old-growth in the classic sense of being multi-storied stands with large diameter trees, high canopy closure, and abundant woody debris. Rather, nest stands were in even-aged, single-storied, mature forest stands with high canopy closure (x = 65%, SE 1.4) and clear forest floors. We recommend changes in procedures for identifying mature and old-growth lodgepole pine forests and describe silvicultural methods for creating goshawk nest stands.

Sources and Patterns of Wolverine Mortality in Western Montana

Abstract We instrumented 36 wolverines (Gulo gulo) on 2 study areas in western Montana and one study area on the Idaho–Montana (USA) border: 14 (9 M, 5 F) on the Pioneer study area, 19 (11 M, 8 F) on the Glacier study area, and 3 (2 M, 1 F) on the Clearwater study area. During 2002–2005, harvest from licensed trapping accounted for 9 (6 M, 3 F) of 14 mortalities, including individuals from all 3 study areas. Based on Akaikes Information Criterion adjusted for small sample sizes (AICc) rankings of 8 a priori models, a trapping model and a trapping-by-sex interaction model were equally supported (ΔAICc < 2) in explaining wolverine survival. Estimated annual survival was 0.80 when we did not consider harvest, whereas additive mortality from harvest reduced annual survival to 0.57. Glacier National Park in the Glacier study area provided some refuge as evidenced by an annual survival rate of 0.77 compared to 0.51 for the Pioneer Mountain study area. We incorporated these survival rates into a simple Lefkovitch stage-based model to examine rates of population change. The finite rate of population change (λ) for the Glacier study area was 1.1, indicating a stable to slightly increasing population, whereas λ for the Pioneer study area was 0.7, indicating a 30% annual population decrease during our study. Changes in λ for both study areas were most sensitive to adult survival. In 2004, we used a Lincoln Index to estimate that 12.8 ± 2.9 (95% CI) wolverines resided in the 4 mountain ranges comprising the Pioneer study area, suggesting that small, island ranges in western Montana supported few individuals. Our results suggest that if wolverines are harvested, they should be managed within individual mountain ranges or small groupings of mountain ranges to limit mortality to within biologically defined limits in recognition of the increased vulnerabilities owing to low fecundity and low population numbers in small mountain ranges. We found that refugia, such as Glacier National Park, were important by reducing trap mortality and providing immigrants to the surrounding population, but even large parks were inadequate to provide complete protection to wolverines from trapping as they ranged outside park borders.