# A Comparison of Methods Used to Examine Snowshoe Hare Habitat Use

> Do hare tracks in the snow tell the same story as their droppings, or as live traps? Over two winters in Maine, researchers compared all three and found that each pointed snowshoe hares to dense understory. Snow track counts proved a cheap way to cover large areas, though tracks show where hares travel more than where they linger.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/1985-litvaitis-sherburne-comparison-methods-used-examine/

## Citation and access

- **Authors:** John A. Litvaitis, James A. Sherburne and John A. Bissonette
- **Year published:** 1985
- **Journal or source:** The Journal of Wildlife Management
- **Full citation:** Litvaitis, J. A., Sherburne, J. A. and Bissonette, J. A. 1985. A comparison of methods used to examine snowshoe hare habitat use. The Journal of Wildlife Management 49(3): 693–695.
- **DOI:** 10.2307/3801696
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.2307/3801696

## Tags

- **Species:** Mammals > Snowshoe hare
- **Region:** North America > United States
- **Tracking:** Scat; Sign surveys; Snow tracks; Track counts; Tracking; Tracks
- **Behaviour and ecology:** Behaviour; Habitat selection
- **Methods and technology:** Chi-square test; Plot searches
- **Data type:** New field data
- **Observer qualification:** Not reported
- **Discipline:** Behaviour and ecology; Wildlife tracking
- **Year published:** 1985

## Detailed keywords

- **Species:** Studied: snowshoe hare (Lepus americanus), the only hares or rabbits (leporids) in each study area. Traps were baited with alfalfa (Medicago sativa). Mentioned from another study: mule deer (Odocoileus hemionus), whose defecation rates varied with activity and cover type. Vegetation named: spruce and fir (Picea spp., Abies balsamea), maples (Acer spp.), birches (Betula spp.), American beech (Fagus grandifolia) and blueberry (Vaccinium angustifolium).
- **Location and habitat:** Two study areas in Maine, United States. Pierce Pond (45°15’N, 70°10’W), in the western mountains of Maine: spruce-fir and northern hardwood forest, with dense regenerating stands after logging. Cherryfield (44°35’N, 67°55’W), on the coast about 190 km east: mostly hardwoods, conifers on poorly drained sites, scattered logging, and commercial blueberry barrens kept open by burning every second year. Habitat was classed by understory stem density: open (under 700 stems per hectare), sparse (700 to 7,000), moderate (7,000 to 20,000) and dense (20,000 or more).
- **Topics:** comparing habitat-use indices; fecal pellet counts; snow track counts; live trapping; understory density and cover; habitat use versus availability; assumptions behind indirect indices (defecation rates, distance travelled, bait attraction); choosing a method by study objectives and budget. Possible inconsistency: the text says at least two of the three indices agreed in seven of the eight comparisons, but Table 1, as read here, shows at least two agreeing in all eight.
- **Statistics or models:** Chi-square (χ²) and z statistics (the Neu et al. 1974 method for comparing use with availability), P < 0.05.
- **Data type:** New field data, September 1981 to May 1983. Pellet counts in spring (April to May) and late summer (September); snow track counts January to March 1982 and 1983; live trapping in April to May and October to November, 1981 to 1983. Only data from the leafless season (October to May) were compared.

## Synopsis

### 1. What they did

Researchers study where mammals spend their time either directly (watching, catching or radio-tracking them) or indirectly, from the signs they leave, such as browsing, tracks and droppings. Different methods are often assumed to tell the same story, but few studies had checked. The authors used three methods at the same time on snowshoe hares in Maine to see whether they gave similar answers about habitat use.

They worked in two study areas, Pierce Pond and Cherryfield, from September 1981 to May 1983. Earlier studies had shown that the density of the understory matters most to hares, so each method was compared against understory stem density (open, sparse, moderate and dense).

- Pellet counts. In two 49-hectare sampling areas per study area, they laid out seven 700-metre transects, each with fifteen 1-metre-radius plots 50 metres apart. Fifty more plots were added in each sampling area in the second year. Hare pellets were counted and removed in spring and late summer.

- Snow track counts. Hare tracks were counted on the 50-metre stretches between plots, within 40 hours of a snowstorm and after at least one full night without snow, so tracks could build up. Each stretch was assigned the understory type of the plot before it.

- Live trapping. Box traps baited with alfalfa were set every 100 metres along the transects, for 8 to 16 days in spring and autumn.

Because the true time hares spent in each understory type was unknown, the authors compared the patterns of the three indices, using only data from the leafless months.

### 2. What they found

- In both areas, all three indices were spread unevenly across the understory types, and pellets, tracks and captures all increased with understory density.

- All three methods showed hares using dense understory (20,000 or more stems per hectare) more than expected from how much of it was available.

- For open, sparse and moderate understory, the methods did not always agree on whether use was more, less or about the same as expected. Still, at least two of the three agreed in seven of the eight comparisons, according to the text.

- No method ever showed hares favouring an understory type while another showed them avoiding it.

- In early trials, more hares were caught with alfalfa bait than with apples, vanilla or no bait, but how far hares travelled to reach the bait was unknown.

### 3. What they concluded

- Pellet counts, track counts and live captures give similar information on how snowshoe hares use understory types, so biologists can choose a method to suit their other aims or their budget.

- Each index has weaknesses. Hare defecation rates might vary with activity or cover, as found in mule deer. Track counts measure distance travelled rather than time spent, so they may overestimate time in large, poor patches that hares cross to reach small feeding areas, where they move less. Bait may draw hares from unknown distances.

- Track counts are cheap over large areas but cannot show seasonal habitat use (they need snow).

- Live trapping is costly, labour intensive and usually limited to small areas, but it gives data for mark-recapture estimates of population size.

- Pellet counts are cheap over large areas and may show seasonal changes in habitat use and relative abundance.

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

Two of the three methods relied on tracks and signs: counting hare tracks in fresh snow along transect segments, and counting hare fecal pellets in fixed plots. No trailing was reported. The snow track counts were timed after snowfall, with at least one snow-free night, so that tracks of a known age could accumulate.

### 5. Tracker or observer certification

Not reported. The paper does not say who counted tracks and pellets or how hare tracks and pellets were identified; snowshoe hares were the only hares or rabbits in the study areas.

### 8. Implications for tracking (Original Wisdom's observations)

_These are Original Wisdom's observations, not the authors'._

- Snow track counts gave the same broad picture of habitat use as pellet counts and live trapping, supporting tracks and signs as an inexpensive way to study habitat use over large areas.

- The authors’ point that tracks record distance travelled rather than time spent is a useful reminder when reading track counts: animals passing through a habitat can leave more tracks there than in the places where they actually feed or rest.

## Cite this summary

Lawrence, K. 2026. Summary of: A Comparison of Methods Used to Examine Snowshoe Hare Habitat Use (Litvaitis and Sherburne et al., 1985). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/1985-litvaitis-sherburne-comparison-methods-used-examine/