# Spoor counts as indices of large carnivore populations: the relationship between spoor frequency, sampling effort and true density

> Do more tracks on the road really mean more animals? In northern Namibia, Ju/Hoansi San trackers counted leopard, lion and wild dog spoor while the true number of animals was known from individual recognition. Spoor density rose in a straight line with true density, but lions and wild dogs, which travel farther each day, left more spoor per animal than leopards.

Summary by Kersey Lawrence, Original Wisdom. Updated 5 November 2026. Page: https://www.originalwisdom.com/tracking-literature/1998-stander-spoor-counts-indices-large/

## Citation and access

- **Authors:** P. E. Stander
- **Year published:** 1998
- **Journal or source:** Journal of Applied Ecology
- **Full citation:** Stander, P. E. 1998. Spoor counts as indices of large carnivore populations: the relationship between spoor frequency, sampling effort and true density. Journal of Applied Ecology 35(3): 378–385.
- **DOI:** 10.1046/j.1365-2664.1998.00313.x
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.1046/j.1365-2664.1998.00313.x

## Tags

- **Species:** Mammals > African wild dog; Mammals > Leopard; Mammals > Lion
- **Region:** Africa > Namibia
- **Tracking:** Footprints; Individual identification; Spoor age estimation; Track counts; Tracking; Tracks and signs identification; Trailing
- **Behaviour and ecology:** Behaviour; Movement
- **Methods and technology:** Analysis of covariance (ANCOVA); Bootstrapping; Double sampling; Kolmogorov–Smirnov test; Linear regression; Mann–Whitney U test; Population estimation; Radio telemetry; Road transects
- **Data type:** New field data
- **Observer qualification:** Other experienced trackers
- **Discipline:** Indigenous knowledge; Wildlife tracking
- **Year published:** 1998

## Detailed keywords

- **Species:** Studied: leopard (Panthera pardus), lion (Panthera leo) and African wild dog (Lycaon pictus). Plants named in the study area: Burkea africana, Commiphora africana, Terminalia sericea and Grewia species. Humans: Ju/Hoansi San trackers (hunters) carried out the spoor work.
- **Location and habitat:** Kaudom Game Reserve (3842 km²; Table 1 sample zone 2345 km²) and Tsumkwe District (4869 km²; Table 1 sample zone 2731 km²), northern Namibia. Tsumkwe District had no formal conservation status and was inhabited by Ju/Hoansi San at 0.39 people per km². The experimental zone was a 244 km² zone in the centre of Tsumkwe District, chosen for its uniform habitat and its 20 randomly placed roads (“cutlines” made for geological prospecting). Habitat: forest and shrub savannah woodland; 400–500 mm summer rainfall; warm semi-arid steppe climate with temperatures from 5 to 36 °C. Substrate: arenosol soils with a hard surface over soft sand, where spotting spoor on roads was moderate to easy.
- **Topics:** spoor counts as an index of population density; true density from recognized individuals and groups; spoor frequency (km per spoor) and spoor density (spoor per 100 km); sampling effort and road penetration; sampling intensity and precision; sex ratio from spoor; group size from spoor; individual recognition from spoor; daily travel distance and home range size; differences between species in the spoor-to-density relationship; habitat uniformity and road use.
- **Statistics or models:** Bootstrap analyses (means, coefficient of variance and 95% confidence intervals with increasing sample size); Kolmogorov–Smirnov two-sample test; ordinary least squares linear regression; analysis of covariance (ANCOVA); Mann–Whitney U test; chi-square test; binomial probabilities (tracker accuracy, from Stander et al. 1997).
- **Data type:** New field data. Direct population estimates were made between 1991 and 1995. Spoor counts were made over 6 months (the paper does not give the exact dates).

## Synopsis

### 1. What they did

Counting large carnivores directly, by recognizing individual animals and groups, is expensive and slow. Counting their spoor is cheaper and repeatable, but few studies had compared spoor counts with known true densities. Stander did both in the same area and time frame, in northern Namibia.

True density, measured directly

- Between 1991 and 1995, he measured the populations of lions and wild dogs in Kaudom Game Reserve, and of lions, wild dogs and leopards in Tsumkwe District, by recognizing and monitoring individual animals and groups.

- In total, 86 animals were immobilized, some with a traditional San bow and arrow adapted for darting. Wild dogs (46) were recognized from photographs, and 47 lions were marked with individual brands. Eighteen leopards, 14 lions and five wild dogs were radio-collared.

- By the end of 1992, every leopard in the experimental zone, and at least 80% of the lions and wild dogs, were marked or individually recognizable.

Spoor counts, made separately

- Over 6 months, experienced San hunters scanned the roads in all three zones daily from an open vehicle driven at 20 km per hour, with two to four people on the bumper and sides.

- Only early-morning counts on “fresh” roads (not disturbed by vehicles, rain or wind) were used, with at least one observer on the bumper and one on the side.

- For each fresh spoor, the trackers recorded species, group size, age and sex. Counts were weighted by group size, so each spoor stood for one animal, and each individual was counted only once per day.

- Sampling effort was described by “road penetration”: km² of area per 1 km of road.

Leopards in the experimental zone

- Most of the data came from leopards in the 244 km² experimental zone. All leopards using it were darted by the San and radio-collared. This was only possible because the trackers could recognize individuals from spoor: they followed the spoor of unknown leopards to capture and mark them, and often followed marked leopards, which confirmed their identifications.

- The 20 roads were sampled 821 times (11 to 26 times each), covering 3089 km. Only spoor made the previous night was recorded, and each leopard’s identity was determined from it.

- Bootstrap analyses tested how accuracy and precision changed with more sampling.

- To compare spoor with true density, he split the leopard population into four independent groups of known individuals, using each leopard’s home range overlap with the zone.

### 2. What they found

Leopards

- There was one leopard spoor for every 38.1 km of road (± 3.83 km). Spoor frequency did not differ between roads and did not depend on road length, in this uniform habitat with random roads.

- Road penetration: with few roads, estimates varied widely. From 1 km of road per 6.5 km² of area, the average spoor frequency (41.49 ± 8.42 km) changed little with more roads.

- Sampling intensity: estimates stabilized at about 30 spoor samples, or 1200 km of driving. Precision rose sharply over the first 10 samples (500 km). Going from 30 to 80 samples improved precision by only 4.6%.

- Spoor density and true density had a strongly significant linear relationship (r² = 0.98).

- Sex ratio: the sex ratio from spoor (1 male to 0.75 female) matched the true sex ratio. This estimate needed more sampling to stabilize: 45 to 60 spoor samples, or 1400 km.

Lions and wild dogs

- True densities were 1.45 ± 0.25 and 0.3 ± 0.1 lions per 100 km², and 0.53 ± 0.25 and 1.20 ± 0.23 wild dogs per 100 km², in two zones.

- Despite small samples, the spoor density of both species had a strong linear relationship with true density (combined r² = 0.98). Lions and wild dogs showed the same slope.

Differences between species

- The slope for leopards was significantly different from the slope for lions and wild dogs. In theory, at the same density, lions and wild dogs would leave more spoor than leopards.

- Leopards travelled shorter daily distances (9.7 ± 0.9 km) than lions (19.4 ± 1.4 km) and wild dogs (24.6 ± 1.7 km), and had smaller home ranges than both.

- Stander suggests the extra spoor of lions and wild dogs may come from their longer daily movements and larger ranges.

### 3. What they concluded

- For leopards, lions and wild dogs, spoor density is a function of true density. The relationships are strongly linear, with intercepts at zero, which fits the expectation that no animals means no spoor.

- The slope of the relationship reflects each species’ ecology, such as daily movement, range use, habitat use and behaviour. So it differs between species.

- Spoor counts depend on thorough sampling design: how many roads and how much road length, and how often they are sampled. The most efficient sampling intensity may vary between areas with different carnivore densities.

- In uniform habitat with random roads, leopards used roads at random. In mixed habitats, carnivores favour certain vegetation or geological types, so roads through their preferred habitats show more spoor than expected.

- “Double sampling” (measuring true density and spoor counts in the same place) can calibrate cheaper indirect methods, but it is expensive and has rarely been done.

- Limitations: the sample sizes are statistically extremely small. Stander argues the data still matter ecologically and economically, given the cost and effort involved.

- The paper does not aim to provide a technique for estimating density or monitoring trends. Its aim is to contribute to understanding how spoor counts relate to true density.

- Highly skilled trackers, especially their ability to recognize individuals, improved understanding of how spoor is distributed, but are not essential for future monitoring designs. Observers with moderate tracking skills, such as hunters, could record spoor distribution along roads or transects.

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

Tracking was central to both parts of the study:

- Spoor counts: trackers identified the species, sex, age and group size from fresh spoor on roads, and judged which spoor had been made the previous night.

- Leopard identity: trackers recognized individual leopards from their spoor.

- Trailing: trackers followed the spoor of unknown leopards so they could be captured, darted with a traditional bow and arrow, and radio-collared. They also followed marked leopards often, which confirmed their identifications.

### 5. Tracker or observer certification

- The trackers were experienced Ju/Hoansi San hunters. They were not certified.

- Their reliability had been tested in an earlier study (Stander et al. 1997). The paper reports high probabilities of correct identification from spoor: species (147 cases), sex (69), individually known animals (32), and group size for lions (39) and wild dogs (12).

- The acknowledgements thank Ghau, Tsisaba, ǂOma, ǀUi, Kaqece, Young ǀUi, Nisa, ǀAm and Dam.

### 6. Other technologies, skills or innovations

- Radio telemetry was used to measure leopard home ranges and true density.

- A traditional San bow and arrow was adapted for darting to immobilize the animals.

- Wild dogs were recognized from photographs, and lions were marked with individual brands.

- Bootstrap analyses were used to find the point where more sampling stops improving the estimates.

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

- Why it matters: reliable estimates of population density are needed for effective conservation management. Most large carnivore populations are declining worldwide, so practical, accurate ways to count them and monitor trends are urgently needed.

- What spoor counts offer: they are repeatable and inexpensive. They are also objective and free from personal bias, open to scrutiny, and can be improved as better information becomes available.

- Planning a survey: spoor counts must be designed with enough roads and enough sampling effort. Habitat use should be considered, because spoor on roads in preferred habitats overstates density.

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

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

- The counting rules shaped the results: only fresh spoor from the previous night, each animal counted once per day, and group size weighted. Anyone comparing spoor counts between places or years can check which rules were used.

- Because lions and wild dogs left more spoor per animal than leopards, a raw spoor count cannot be compared directly between species. Each species needs its own relationship between spoor and density.

## Cite this summary

Lawrence, K. 2026. Summary of: Spoor counts as indices of large carnivore populations: the relationship between spoor frequency, sampling effort and true density (Stander, 1998). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/1998-stander-spoor-counts-indices-large/