# A Track Count for Estimating Mountain Lion Felis concolor californica Population Trend

> Can you follow the fortunes of a whole state’s mountain lions by counting their tracks on dusty back roads? In California, a single surveyor on a motorcycle counted lion track sets across randomly chosen survey areas, and found it cheap and practical. The tracks also showed that lions favour roads along streams and mountain slopes, not the ridge tops where many surveyors looked for them.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/1995-smallwood-fitzhugh-track-count-estimating-mountain/

## Citation and access

- **Authors:** K. Shawn Smallwood and E. Lee Fitzhugh
- **Year published:** 1995
- **Journal or source:** Biological Conservation
- **Full citation:** Smallwood, K. S. and Fitzhugh, E. L. 1995. A track count for estimating mountain lion Felis concolor californica population trend. Biological Conservation 71(3): 251-259.
- **DOI:** 10.1016/0006-3207(94)00034-N
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.1016/0006-3207(94)00034-N

## Tags

- **Species:** Mammals > American black bear; Mammals > Bobcat; Mammals > Coyote; Mammals > Domestic dog; Mammals > Grey fox; Mammals > Mountain lion; Mammals > Red fox
- **Region:** North America > United States
- **Tracking:** Sign surveys; Track counts; Track measurements; Tracking; Tracks; Tracks and signs identification
- **Behaviour and ecology:** Behaviour; Habitat selection; Movement
- **Methods and technology:** Chi-square test; Observer reliability; Population trends; Road transects; Survey costs
- **Data type:** New field data
- **Observer qualification:** Reported but not validated
- **Discipline:** Behaviour and ecology; Population trends; Wildlife tracking
- **Year published:** 1995

## Detailed keywords

- **Species:** Studied: mountain lion (Felis concolor californica). Also counted on the 1992 survey and analysed for habitat, topography and travel: black bear (Ursus americanus), bobcat (Felis rufus), coyote (Canis latrans), foxes (red fox Vulpes vulpes and grey fox Urocyon cinereoargenteus, not told apart) and domestic dog (Canis familiaris). Mountain lion and black bear tracks were split into adults and “families” (tracks including juveniles). Human tracks are mentioned as often occurring with dog tracks in chaparral.
- **Location and habitat:** California, United States: 0.5 degree latitude and longitude quadrats statewide, excluding mostly urban or agricultural areas, quadrats inaccessible by road, and coast and border quadrats that were mostly ocean or in another state. Dusty dirt roads in mountain lion breeding range; Coast Range, Sierra Nevada and Southern Mountains. Topography: ridge tops, ridge slopes, streams (1st- and 2nd-order), mountain slopes, knolls and peaks, basins and plateaus. Habitats: chaparral, riparian, mature conifer forest, conifer forest with understory, montane hardwood-conifer forest (with and without understory), oak woodland, sage with pine or juniper, sage, and clearcuts. Desert quadrats were included in 1986 but the authors recommend excluding them.
- **Topics:** population index; population trend monitoring; track counts on road transects; track sets; presence/absence versus counts; volunteer local biologists versus a single surveyor; track identification errors; survey costs; efficient sample size; associations of tracks with topography and habitat; side of the road used (uphill, middle, downhill); direction of travel; co-occurrence of carnivore tracks; age and sex classes from rear heel pad width; transient animals; calibrating track counts against direct counts; survey design recommendations.
- **Statistics or models:** Observed/expected association ratio (M) on 0.16 km road segments with chi-square tests; negative binomial distribution fitted to track counts (chi-square = 3.85, 5 degrees of freedom, p = 0.84); Elliott’s (1971) sample size formula for contagious distributions, as simplified by Eberhardt (1978); binomial equation for the first sample size. Recommended for future surveys (not used here): Wilcoxon matched-pairs signed-ranks test, Friedman two-way analysis of variance, Cochran’s Q test, analysis of covariance.
- **Data type:** New field data. Local teams set up transects in 1985 and surveyed them in the summers of 1985 and 1986. Smallwood surveyed the transects in 1986 and 1992; the other carnivore species were recorded in 1992 only. Inconsistencies in the paper: the abstract gives 33.8 km of transect per quadrat, the text 33.9 km (three transects of 11.3 km); quadrats are described as “50 km²” although a 0.5 degree grid gives far larger quadrats (perhaps 50 km sides were meant); the abstract says 48 quadrats, while the methods give a planned 50, with 36 and 38 completely surveyed by the two survey types.

## Synopsis

### 1. What they did

Large carnivores are hard and costly to count over big areas, so their population trends are often unknown. In California, mountain lion numbers had been estimated in different ways by different investigators. The authors set out to build a practical, low-cost index of the mountain lion population trend based on counting sets of tracks on dirt roads.

- Survey layout. California was divided into quadrats along 0.5 degree lines of latitude and longitude, and quadrats were picked at random. In each, local wildlife biologists chose up to three dirt-road transects of 11.3 km, at least 16 km apart, using the authors’ written guidelines. East-west roads were requested, because low sun makes tracks easier to see.

- Two ways of surveying (1986). Local teams (a driver and a professional wildlife biologist) drove the transects at 5 to 8 km per hour and recorded only whether mountain lion tracks were present. They sketched one track from each set and measured heel pad and toe width, so the authors could check the identification. Separately, Smallwood rode the same transects on a motorcycle at 10 to 16 km per hour and counted every track set (a continuous series of footprints, presumably from one animal). He traced tracks onto acetate and noted direction of travel and which side of the road the animal used.

- Other carnivores and habitat (1992). Smallwood repeated the survey in 1992 and also recorded tracks of black bear, coyote, dog, bobcat and fox. He mapped habitat and topography every 0.8 km, and the authors tested where tracks were found more or less often than expected by chance.

- Costs and sample size. They compared the costs, ease and reliability of the surveys, and estimated how many quadrats a future survey needs.

### 2. What they found

The two surveys

- Local biologists found mountain lion tracks on 20 of 36 completely surveyed quadrats (55.6%); Smallwood found them on 21 of 38 (55.3%).

- On the 29 quadrats surveyed by both, tracks were found on 72% (84% of the 25 non-desert quadrats).

- Local biologists misidentified 9.5% (three) of the tracks they sketched, and five teams (14.3%) did not send sketches.

- The cheapest option was one person surveying all quadrats. All three options costed were inexpensive: about 4,600 to 11,300 US dollars per statewide survey, based on 1986 costs.

Sample size

- Track counts were clumped (they fitted a negative binomial distribution), with a mean of 1.26 track sets per quadrat.

- About 44 quadrats would give a mean within 40% of the true mean at 95% confidence; 52 quadrats would give 30% at 90% confidence.

- This estimate was conservative: the 15 most productive quadrats were not fully surveyed, and they held about six times the track density of the rest. With them completed, 37 quadrats might have been enough.

Where the tracks were

- Carnivore tracks were absent on most road segments and turned up together mostly where adult mountain lion tracks were.

- Mountain lion tracks were found 2.3 times more often than expected along streams and 1.9 times along mountain slopes, and less often on ridge tops. They were found more often in oak woodland and montane hardwood-conifer forest.

- Tracks of mountain lion families and black bear families occurred together 26 times more often than expected by chance, but they avoided segments with tracks of most other carnivores.

- Coyote and dog tracks were found more often on basins and plateaus, fox tracks on knolls and peaks.

- Species used different sides of the road. Adult mountain lions and coyotes preferred the uphill side along streams; adult black bears the middle; mountain lion and black bear families the downhill side.

- Direction of travel differed by region and species. For example, adult mountain lion tracks showed east-west travel on 67% of segments in the Coast Range, southward on 70% in the Sierra Nevada, and eastward on 76% in the Southern Mountains.

Age and sex from tracks

Using rear heel pad width (over 52 mm adult male, 43 to 52 mm mostly adult female, under 43 mm juvenile), the ratio of adult males to adult females to juveniles was 1.0 : 1.0 : 0.1 (45 track sets). Far fewer juveniles showed up than in a radio-telemetry study in Idaho. The authors suggest juveniles do not travel on roads in proportion to their numbers.

### 3. What they concluded

- The track survey was inexpensive, easy to run and able to cope with changes in staff and road conditions. Dividing each quadrat into three transects spread the effort and let one person or team finish a quadrat in a day. A statewide survey could be done in 30 days.

- Using the same survey, Smallwood (1994) had found a population decline and a change in age and sex make-up between 1986 and 1992, both linked to timber loss.

- Many local teams placed transects on ridge tops hoping to find more tracks, but mountain lions avoided ridge tops. Choosing transects along streams, on mountain slopes and in the habitats lions favour would cut the sample size, cost and effort needed.

- Track counts would be more useful if calibrated against direct counts of animals of known age and sex in smaller intensive study areas, because transient animals and different age groups are detected at different rates.

- Recommendations: keep methods consistent between years rather than trying to survey every quadrat on the same day; use about 44 quadrats; exclude desert quadrats, where coarse dry sand hides track detail; use two trained people on motorcycles or vehicles with the best view; fewer trackers using standard methods would make results more consistent between years. Non-parametric tests that account for differences between quadrats are suggested for detecting trends.

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

Tracking was the core of the method. Surveyors identified mountain lion footprints on dusty dirt roads and counted track sets. Local teams sketched and measured tracks, and these sketches were used to check identifications: a sketch counted as an error when it showed claw marks, fewer than three heel lobes or the wrong size. Smallwood traced tracks onto acetate and read direction of travel and which side of the road each animal used. In 1992 he also identified tracks of black bear, bobcat, coyote, fox and dog. Rear heel pad width was used to sort mountain lion tracks into age and sex classes. No trailing was reported. The authors mention their earlier method for identifying individual mountain lions from tracks (Smallwood and Fitzhugh 1993) but do not apply it here.

### 5. Tracker or observer certification

Reported but not validated. The local teams were professional wildlife biologists given written instructions and tracings of tracks of mountain lion, bobcat, dog, coyote and black bear; their identifications were checked from sketches (9.5% wrong). Smallwood, who did the counts, is referred to as an “expert tracker” in the costing table, but his skill was not tested or certified in this study.

### 6. Other technologies, skills or innovations

- Motorcycle surveys, for a wider field of view and faster travel.

- Track sketches on pre-printed grids and tracings on acetate.

- Dust ratings at the start, middle and end of each transect and at tracks (no link was found between dust ratings and success in finding tracks, Smallwood 1994).

- An observed/expected association measure for linking tracks with topography, habitat and other species on 0.16 km road segments.

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

- Knowing the status and trend of large carnivores is critical for their conservation, and standard monitoring exists for only a few carnivore populations worldwide. A periodic road track survey can detect a changing mountain lion population cheaply over a whole state.

- The same survey can collect information on other carnivores, their habitats and their associations, which helps explain trends.

- The method can be adapted for some other large carnivores.

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

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

- Where the surveyors placed transects mattered: local teams chose ridge tops, where they expected lions, but lions mostly used roads along streams and mountain slopes. Knowledge of how a species moves through the landscape can make track surveys more efficient.

- Different age groups and species used different sides of the road and travelled in different directions, which affects what a surveyor will find on a given stretch.

## Cite this summary

Lawrence, K. 2026. Summary of: A Track Count for Estimating Mountain Lion Felis concolor californica Population Trend (Smallwood and Fitzhugh, 1995). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/1995-smallwood-fitzhugh-track-count-estimating-mountain/