# Home range characteristics of a subordinate predator: selection for refugia or hunt opportunity?

> Where does a cougar choose to live when wolves are moving back in? Following 28 collared cougars over ten years near Yellowstone, researchers found that cougars picked home ranges with good hunting and kept their distance from wolf packs, but did not seek out extra forest or rugged ground. Females chose the best hunting areas, which may bring them closer to wolves after the same prey.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/2014-lendrum-elbroch-home-range-characteristics-subordinate/

## Citation and access

- **Authors:** P. E. Lendrum, L. M. Elbroch, H. Quigley, D. J. Thompson, M. Jimenez and D. Craighead
- **Year published:** 2014
- **Journal or source:** Journal of Zoology
- **Full citation:** Lendrum, P. E., Elbroch, L. M., Quigley, H., Thompson, D. J., Jimenez, M. and Craighead, D. 2014. Home range characteristics of a subordinate predator: selection for refugia or hunt opportunity? Journal of Zoology 294: 59–67.
- **DOI:** 10.1111/jzo.12153
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.1111/jzo.12153

## Tags

- **Species:** Mammals > Mountain lion
- **Region:** North America > United States
- **Behaviour and ecology:** Behaviour; Carnivore coexistence; Habitat selection; Home range; Hunting; Interspecific competition; Intraguild interactions; Predation risk
- **Methods and technology:** Analysis of variance (ANOVA); Geographic Information System (GIS); Global Positioning System (GPS) collars; Radio telemetry; Resource selection function; Utilization distribution; Very high frequency (VHF) radio collars
- **Data type:** New field data
- **Observer qualification:** Not needed
- **Discipline:** Behaviour and ecology
- **Year published:** 2014

## Detailed keywords

- **Species:** Studied: cougar (Puma concolor), 28 adult resident individuals. Dominant competitor whose pack locations were used: wolf (Canis lupus); wolves were not themselves studied, only the centres of known pack territories supplied by the US Fish and Wildlife Service and Wyoming Game and Fish Department. In the study area but not studied: brown bear (Ursus arctos), American black bear (U. americana), coyote (C. latrans), red fox (Vulpes vulpes); ungulate prey species elk (Cervus elaphus), mule deer (Odocoileus hemionus), white-tailed deer (O. virginianus), Shiras moose (Alces alces shirasi), bighorn sheep (Ovis canadensis) and North American pronghorn (Antilocapra americana). Examples from the literature: cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus), African lion (Panthera leo) and spotted hyena (Crocuta crocuta).
- **Location and habitat:** Southern Yellowstone Ecosystem (SYE), southern Teton County, northwestern Wyoming, United States. Elevations from 1800 m in the valleys to over 3600 m; short cool summers and long cold winters, snow up to over 245 cm deep. Foothill grasslands, big sagebrush (Artemisia tridentata, printed as “tridentate”) shrub-steppe, Douglas-fir and aspen forests, higher conifer forests of lodgepole pine, subalpine fir, Engelmann spruce and whitebark pine, and cottonwood and willow riparian corridors. The study area is described as about 2300 km² in the methods, but the minimum convex polygon used as the study area in the results is 7012 km².
- **Topics:** home range size and placement; core areas (50%) and 95% home ranges; male versus female home ranges; Very High Frequency (VHF) versus Global Positioning System (GPS) collar data; refugia (forest cover, terrain ruggedness, distance from roads and wolves); “hunt opportunity” as a stand-in for prey availability, mapped from cougar kill sites; subordinate predators; competition with recolonizing wolves; human hunting of cougars; roads; land cover classes; capture with trailing hounds.
- **Statistics or models:** Fixed-kernel density estimator (KDE) home ranges (least squares cross-validation smoothing for VHF data, plug-in method for GPS data) in the Geospatial Modeling Environment; minimum convex polygon (MCP) for the study area; random points (200 per core area, 400 per 95% home range, 2000 in the study area) in ArcGIS 10; vector ruggedness measure (VRM); a kill-site resource selection function from an earlier study (Elbroch et al. 2013) turned into a “hunt opportunity” surface; correlation matrix for collinearity; generalized linear models with likelihood ratio (chi-square) tests (JMP 10); analyses of variance (ANOVAs) with Tukey’s honestly significant difference (HSD) tests; root and log10 transformations. Inconsistencies: (1) the text says VHF home ranges were 1.4 to 1.9 times larger than GPS home ranges, but the means in Table 1 give ratios of about 1.8 (male 95%), 2.2 (male core), 2.6 (female 95%) and 3.4 (female core); (2) the text says 16 females wore VHF collars, but the 3000 and 6000 VHF female points (200 and 400 per cougar) imply 15; (3) the text says “all cougar home ranges” were further from wolf packs, but Table 1 shows wolf distances only for GPS home ranges (VHF rows are n/a); (4) the discussion says cougars selected home ranges with higher hunt opportunity than the study area, but Table 1 shows GPS male 95% home ranges were statistically the same as the study area; (5) study area given as about 2300 km² in the methods and 7012 km² in the results; (6) in Table 1 the GPS female core area maximum distance to wolves (92,945 m) is far above the study area maximum (20,483 m).
- **Data type:** New field data: cougars captured each year November to March; VHF collars 2001 to 2005 and GPS collars 2006 to 2011; 67 annual home ranges from 28 cougars. Existing data: wolf pack territories from agency partners and a kill-site analysis from an earlier study.

## Synopsis

### 1. What they did

Cougars are often called “habitat generalists”, yet little is known about what shapes where they set up their home ranges. In the Southern Yellowstone Ecosystem of Wyoming, cougars share the land with wolves and bears, which dominate them, and with human hunters. The authors asked whether cougars choose home ranges for safety (refugia such as forest, rugged terrain, and distance from roads and wolves), for good hunting, or for both.

- Capture and collaring. Each winter, from November to March, they used trailing hounds to tree or bay cougars, then immobilized them and fitted Very High Frequency (VHF) radio collars (2001 to 2005) or Global Positioning System (GPS) collars (2006 to 2011). GPS collars took four to eight locations a day.

- Home ranges. They mapped annual 95% home ranges and 50% “core areas” for adult resident cougars, and tested whether sex, collar type, the amount of forest or “hunt opportunity” explained home range size.

- Hunt opportunity. Counting prey across such large areas is hard, so they used a map of how likely a cougar was to make a kill at each spot, built from cougar kills found in the field in an earlier study.

- What is inside each home range. They placed random points inside each home range and across the whole study area, and compared hunt opportunity, terrain ruggedness, distance to roads, distance to the centre of known wolf pack territories, and land cover.

### 2. What they found

The cougars

- 67 annual home ranges from 28 cougars: 9 females and 5 males with GPS collars, and 16 females and 5 males with VHF collars. Seven cougars wore both collar types in different years.

Home range size

- Sex and collar type explained home range size at both scales. Male home ranges were 1.9 to 3.3 times larger than female ones (for example, GPS male 95% home ranges averaged 567 km² and GPS female ones 210 km²).

- Home ranges from VHF data were larger than those from GPS data (the text gives 1.4 to 1.9 times; see the keywords for the table values).

- The amount of forest partly explained the size of core areas. Hunt opportunity did not explain home range size.

What home ranges contained

- Hunt opportunity: highest in female core areas and lowest in GPS male 95% home ranges. Female home ranges had more hunt opportunity than the study area as a whole; GPS male 95% home ranges did not differ from it.

- Wolves: home ranges (GPS data) were further from the centres of known wolf pack territories than expected, and males were further away than females.

- Roads: all home ranges were closer to roads than expected. Male 95% home ranges were further from roads than female ones.

- Ruggedness: terrain ruggedness hardly differed, being only slightly higher than the study area in some cases.

- Land cover: home ranges mostly mirrored the study area. All had less meadow than expected; most had more shrub; female VHF core areas had more forest; females had less barren ground than males.

- Apart from size, home range features from VHF and GPS data were remarkably similar.

### 3. What they concluded

- Cougars in the Southern Yellowstone Ecosystem choose home ranges that give good hunting and keep them away from wolves, rather than ones with more refuge such as forest or rugged ground.

- Distance from wolves was the only safety feature they selected. Their home ranges were closer to roads than expected, where the authors assumed they were more exposed to human hunters, likely because roads follow the low valleys where prey gather in winter.

- Females chose better hunting areas than males, supporting the long-held idea that females pick home ranges with enough prey for themselves and their young. Because wolves also focus on prey-rich areas, this may bring females closer to wolves.

- Avoiding re-established wolves likely limits good habitat for cougars, with possible effects on young cougars’ survival and dispersal and on the population.

- VHF data give larger home ranges than GPS data, so sampling methods need to be standardized. Because home range features were so similar, earlier VHF-based cougar studies still hold value.

### 4. How tracking was or was not used

The paper does not report people tracking cougars. Cougars were found for capture with trailing hounds, which chased them up a tree or onto a rocky outcrop. The “hunt opportunity” map was built from cougar kills located and verified in the field in an earlier study (Elbroch et al. 2013), not in this one.

### 6. Other technologies, skills or innovations

- VHF collars (Telonics) located by triangulation from the ground or from fixed-wing aircraft (Program LOAS), and GPS collars (Telonics, Televilt or Vectronics); all collars had mortality sensors.

- A “hunt opportunity” map built from a kill-site resource selection function, used as a stand-in for prey availability.

- Vector ruggedness measure from a digital elevation model, and Gap Analysis Program land cover at 30 m resolution reclassified into five classes.

- Seven cougars collared with both collar types in alternating years, allowing a direct comparison of VHF and GPS home ranges.

### 7. Implications for conservation (from the paper)

- Competition with recolonizing wolves likely limits the availability of good cougar habitat, with implications for young cougars’ survival, their success in dispersing and cougar population dynamics.

- Cougars are a trophy species hunted in Wyoming, and their home ranges lie closer to roads than expected.

- Home range sizes vary widely with sampling method, so methods need to be standardized.

## Cite this summary

Lawrence, K. 2026. Summary of: Home range characteristics of a subordinate predator: selection for refugia or hunt opportunity? (Lendrum and Elbroch et al., 2014). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/2014-lendrum-elbroch-home-range-characteristics-subordinate/