# Estimating larger carnivore numbers from track counts and measurements

> How many leopards, caracals and brown hyaenas share a small game ranch in South Africa’s Waterberg? Reading 283 tracks on sand roads over six months, two researchers compared three ways of estimating carnivore numbers from tracks, from a subjective comparison of track measurements to formal statistics, and found that all three gave similar answers.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/2005-gusset-burgener-estimating-larger-carnivore-numbers/

## Citation and access

- **Authors:** Markus Gusset and Nicole Burgener
- **Year published:** 2005
- **Journal or source:** African Journal of Ecology
- **Full citation:** Gusset, M. and Burgener, N. 2005. Estimating larger carnivore numbers from track counts and measurements. African Journal of Ecology 43(4): 320-324.
- **DOI:** 10.1111/j.1365-2028.2005.00581.x
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.1111/j.1365-2028.2005.00581.x

## Tags

- **Species:** Mammals > African wild cat; Mammals > Black-backed jackal; Mammals > Brown hyena; Mammals > Caracal; Mammals > Leopard; Mammals > Serval
- **Region:** Africa > South Africa
- **Tracking:** Individual identification; Track counts; Track measurements; Tracking; Tracks; Tracks and signs identification
- **Methods and technology:** Analysis of variance (ANOVA); Linear regression; Population estimation; Road transects
- **Data type:** New field data
- **Observer qualification:** Not reported
- **Discipline:** Wildlife tracking
- **Year published:** 2005

## Detailed keywords

- **Species:** Studied: leopard (Panthera pardus), caracal (Caracal caracal), serval (Leptailurus serval), African wildcat (Felis silvestris libyca), brown hyaena (Parahyaena brunnea) and black-backed jackal (Canis mesomelas). Discussed from other studies: cougar or mountain lion (Puma concolor), lion (Panthera leo) and African wild dog (Lycaon pictus).
- **Location and habitat:** Shayamanzi Game Ranch (1500 ha), Waterberg region, Limpopo Province, northern South Africa. Mainly private properties in a larger ecosystem that carnivores move through. Sour bushveld, or Waterberg moist mountain bushveld, of the savannah biome. Tracks were read mainly on sand roads.
- **Topics:** indirect census of carnivores; track counts as an index of density; individual identification from track measurements; comparing a subjective (qualitative) and two statistical (quantitative) estimates; track density versus true density (Stander’s regression method) with a correction factor for local leopard density; foot placement of caracal and African wildcat (hind foot over front foot) leaving incomplete track sets; effect of substrate and slope on measurements; survey time (about 10 minutes to measure one track set). Inconsistency: Table 2 is headed as the number of carnivores on the ranch, but the leopard figure from the regression method (3.2) is a density per 100 km², while the ranch is only 15 km² (1500 ha); the paper compares it directly with the counts of individuals from the other two methods without explaining this.
- **Statistics or models:** Simple linear regression of track density on true density from Stander (1998) (slope 1.9, intercept 0), multiplied by a correction factor of 1.3 for the higher leopard density in the Waterberg; repeated measures analysis of variance (ANOVA) with Tukey-Kramer multiple comparisons post-tests (InStat V2.04, GraphPad Software), P < 0.05; means and standard deviations of track measurements.
- **Data type:** New field data: tracks recorded periodically over 6 months in 2001 (months not given).

## Synopsis

### 1. What they did

Knowing how many large carnivores live in an area matters for their conservation, but counting them directly is difficult, expensive and slow. The authors set out to compare three indirect ways of estimating carnivore numbers from tracks, because earlier work had called for several independent surveys to reach a reliable figure.

They worked on Shayamanzi Game Ranch, a 1500 hectare game ranch in the Waterberg region of Limpopo Province, South Africa. Over 6 months in 2001 they recorded all the fresh tracks of cats, hyaenas and members of the dog family that they could find, identifying them with the field guide by Stuart and Stuart (1998). Most tracks were on sand roads, the only substrate where tracks could be identified and measured, and the roads were not prepared beforehand. For each set of tracks, the same person recorded the location and measured the total length, total width and main pad height of the front and hind feet, as well as straddle and stride.

They then estimated numbers in three ways:

- Qualitative estimate. Individuals were told apart subjectively by comparing track measurements, before any statistics were run.

- Regression method (leopards only). They used the straight-line relationship between track density and true leopard density found in Namibia by Stander (1998), and multiplied the result by 1.3 because radio-tracking had shown leopard density in the Waterberg to be higher than in Stander’s study area.

- ANOVA method. A statistical test (repeated measures analysis of variance, ANOVA, with follow-up comparisons) on the foot measurements counted how many track sets differed significantly from each other. Stride and straddle were left out because they can change with slope and with what the animal is doing.

### 2. What they found

- They found and identified 283 tracks of six larger carnivores: leopard, caracal, serval, African wildcat, brown hyaena and black-backed jackal. Of these, 138 could be measured.

- Tracks of every species were found throughout the 6 months and all over the ranch.

- Caracal and African wildcat often placed the hind foot exactly over the front foot, so many of their tracks gave measurements for one foot only.

- Average track sizes matched those given by Stuart and Stuart (1998).

- Leopard track density was 4.7 tracks per 100 km of sand road, from about 300 km of road surveyed.

- The ANOVA found significant differences between track sets for all six species, so individual track sets could be separated.

Estimates (qualitative, regression, ANOVA)

- Leopard: 4; 3.2; 3

- Caracal: 6; not applicable; 5

- Serval: 3; not applicable; 3

- African wildcat: 5; not applicable; 5

- Brown hyaena: 13; not applicable; 12

- Black-backed jackal: 10; not applicable; 7

The regression figure for leopards (3.2 per 100 km²) is higher than the 1.9 per 100 km² found earlier in the Waterberg with radio-tracking. The authors suggest this may be due to less persecution and more prey since game ranching began, or to the difference in methods.

### 3. What they concluded

- All three methods gave similar results that seem reasonable given the animals’ social and spatial organisation, and may reflect the real number of larger carnivores living in or passing through the ranch.

- The estimates could not be checked against a direct count, which the authors call a major drawback. Sample sizes were small, and the ranch is smaller than the home ranges of all these species.

- Stander’s method needs only counting, not measuring, which saves time: measuring one complete track set took about 10 minutes. More studies should test it against direct counts and develop equations for other species and habitats. Its drawback is that individuals cannot be identified, unless experienced local trackers who can tell individual tracks apart without measuring them are available.

- The subjective approach also seemed fairly accurate, although it usually gave higher numbers than the statistical methods. This differs from earlier work on cougars, which found that subjective identification of individuals does not usually work.

- Future studies should record the substrate and slope where each track is measured.

- Overall, tracking data can give a reliable and cost-effective indirect estimate of larger carnivore numbers.

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

The whole study rests on tracks and signs identification and track measurement. Fresh tracks were found mainly on sand roads, identified to species with a field guide, and measured to tell individuals apart. Tracks were counted per 100 km of road for the leopard regression method. No trailing was reported, and tracking conditions were not assessed.

### 5. Tracker or observer certification

Not reported. The paper says only that the measurements were always taken by the same person, and that tracks were identified with a field guide. Nothing is said about that person’s tracking experience, which matters here because the qualitative estimate depends on a subjective judgement of which tracks belong to which individual.

### 6. Other technologies, skills or innovations

- Car odometer readings to estimate the distance of sand road surveyed.

- Statistical software (InStat V2.04, GraphPad Software) for the repeated measures ANOVA and Tukey-Kramer post-tests.

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

- Indirect estimates from tracking data appear reliable and cost-effective, which is promising for research and conservation of larger carnivores where direct counts are not possible for practical or financial reasons.

- All the larger carnivores expected in the area were present, which the authors take as a sign of good habitat for them. They point out the presence of the brown hyaena, whose role as a scavenger makes them an integral part of the ecosystem.

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

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

- The authors name experienced local trackers as the way to identify individuals without measuring tracks. This suggests that tracker skill could combine the speed of simple track counts with individual identification, something this study did not test.

- The foot placement of caracal and African wildcat, with the hind foot landing on the front track, is a practical reminder that what a track shows depends on gait as well as foot size.

## Cite this summary

Lawrence, K. 2026. Summary of: Estimating larger carnivore numbers from track counts and measurements (Gusset and Burgener, 2005). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/2005-gusset-burgener-estimating-larger-carnivore-numbers/