# The importance of crossroads in faecal marking behaviour of the wolves (Canis lupus)

> Where do wolves leave their scats? In northwest Spain, researchers searched 837 km of forest roads and found that wolves pile their scats at crossroads, especially well-connected junctions linking different parts of their territory, and often on raised objects where they are easy to see and smell. For wolves, a crossroads is a message board.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/2004-barja-de-miguel-importance-crossroads-faecal-marking/

## Citation and access

- **Authors:** Isabel Barja, Francisco Javier de Miguel and Felipe Bárcena
- **Year published:** 2004
- **Journal or source:** Naturwissenschaften
- **Full citation:** Barja, I., de Miguel, F. J. and Bárcena, F. 2004. The importance of crossroads in faecal marking behaviour of the wolves (Canis lupus). Naturwissenschaften 91(10): 489–492.
- **DOI:** 10.1007/s00114-004-0557-1
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.1007/s00114-004-0557-1

## Tags

- **Species:** Mammals > Grey wolf
- **Region:** Europe > Spain
- **Tracking:** Scat; Sign surveys; Tracking; Tracks and signs identification
- **Behaviour and ecology:** Behaviour; Communication
- **Methods and technology:** Chi-square test; Road transects
- **Data type:** New field data
- **Observer qualification:** Not reported
- **Discipline:** Behaviour and ecology; Wildlife tracking
- **Year published:** 2004

## Detailed keywords

- **Species:** Studied: wolf (Canis lupus). Mentioned from other studies: red fox (Vulpes vulpes) and Iberian lynx (Lynx pardinus), which also pile scats at crossroads.
- **Location and habitat:** Montes do Invernadeiro Natural Park (12,000 ha), northwest Spain. A craggy region of low mountains and deep valleys, crossed by a dense network of firebreaks and forestry roads that wolves often use. About 0.9 crossroads per kilometre of road.
- **Topics:** scent marking; faecal (scat) marking; territorial marking; chemical communication; visual and olfactory signals; crossroads (road intersections) as marking sites; conspicuous or raised substrates; latrines; road accessibility and driveability; territory borders; rendezvous sites; non-invasive sampling (DNA and stress hormones from scats).
- **Statistics or models:** Chi-square tests (observed versus expected scat numbers, with expected numbers based on distance surveyed and a correction for how many roads meet at a crossroads); Spearman rank correlations; K-means cluster analysis (“conglomerate analysis”) grouping crossroads by accessibility, driveability and scat number. Possible errors in the paper: (1) the abstract says 60.1% of scats were “at crossroads”, but Table 1 shows only 67 of 311 scats (21.5%) inside crossroads; 60.1% is crossroads plus their 30 m periphery together (187 of 311). (2) The main chi-square is printed as 1.170 (df 2, P<0.001); this cannot give P<0.001, and recalculating from Table 1 gives about 1170, so the decimal point is probably a typing error. (3) Scats per km at the periphery (1.67) and along the rest of the roads (0.15) do not quite match Table 1 (120 / 77.64 km = 1.55; 124 / 751.48 km = 0.17). (4) The methods call 837.2 km a “total area”; it is a distance.
- **Data type:** New field data: 14 surveys (2 to 3 days each) from May 1998 to March 2000; 837.2 km of roads and firebreaks surveyed in total, 99 crossroads, 311 wolf scats.

## Synopsis

### 1. What they did

Wolves mark their territories with urine and with secretions from glands, and scats probably play a part too. A scat left at a busy, prominent spot is more likely to be found by other wolves. People who know wolves say crossroads are good places to find their tracks, but how wolves use crossroads for scat marking had not been well documented.

The researchers studied a wolf population in Montes do Invernadeiro Natural Park, northwest Spain, from May 1998 to March 2000. Over 14 surveys of 2 to 3 days each, they searched 837.2 km of unsurfaced roads, asphalt roads and firebreaks for wolf scats, and mapped each one on a 1 km grid.

They compared how many scats lay:

- Inside crossroads: the area where two or more roads meet.

- On the periphery: the stretch of road up to 30 m from the centre of the crossroads.

- Along the rest of the roads.

They worked out how many scats would be expected in each zone if scats were spread at random, allowing for the extra chance that a wolf passes through a crossroads compared with any single point on a road. They also noted whether each scat lay on a conspicuous or raised object.

For each of 99 crossroads they recorded the number of roads meeting there, the kinds of road, the surface area, the altitude, the accessibility (how important the roads were as routes between areas, scored 1 to 3) and the driveability (how easily a vehicle could travel them, scored 1 to 3).

### 2. What they found

Where the scats were

- They found 311 scats: 67 inside crossroads, 120 on the periphery and 124 along the rest of the roads.

- Far more scats were inside crossroads and on their periphery than expected by chance. Inside crossroads there were 8.30 scats per km, compared with 1.67 on the periphery and 0.15 along the rest of the roads.

- Together, crossroads and their periphery held 60.1% of all scats. (The abstract calls this “at crossroads”.)

- The closer to the centre of a crossroads, the more scats. Even so, 57.8% of scats lay between 2 and 10 m from the centre.

- Grid squares with more crossroads held more scats.

What the scats were on

- At crossroads and their periphery, 72.1% of scats were on conspicuous or raised objects, compared with 45.6% along the rest of the roads.

Which crossroads were chosen

- Crossroads where four or more roads met held more scats on average (5.7) than those where three roads met (3.2).

- Crossroads with more scats were those with higher accessibility and driveability, and to a lesser extent those with a larger surface area. Altitude and the type of road made no difference.

- Grouping the crossroads gave three clear types: 12 with high accessibility, high driveability and many scats (6.83 on average); 24 in between (3.21 scats); and 63 with low accessibility, low driveability and very few scats (0.30).

### 3. What they concluded

- Wolves travelling along roads choose crossroads for their scats. Because wolves can arrive at a crossroads from several directions, a scat there is more likely to be found by other wolves. Scats on conspicuous objects add a visual signal to the scent.

- Wolves favour crossroads that link different mountains and important areas of their territory, such as the rendezvous site. Crossroads with low accessibility are hardly marked, probably because wolves rarely use those roads.

- The same crossroads were marked again and again. The authors suggest wolves are drawn to old scats and mark over them, so that piles of scats (latrines) build up.

- The most heavily marked crossroads were far apart and on mountain tops, where scent spreads better. The authors suggest this marks territory borders against intruders.

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

The study is a scat survey: the researchers searched roads and firebreaks for wolf scats and recorded where each lay and what it was on. The paper does not say how scats were identified as wolf scats, and no footprints or trailing were used.

### 5. Tracker or observer certification

Not reported. The paper does not say who found and identified the scats or how their identification was checked. Several people are thanked for help with the fieldwork and surveys.

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

- Because scats pile up at crossroads, crossroads give clear evidence that wolves are present. Knowing where wolves are and how many there are, which needs indirect methods, is essential for their conservation.

- Crossroads rich in scats could supply valuable material for non-invasive studies, such as DNA work and stress hormone (glucocorticoid) measurements.

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

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

- For anyone searching for wolf sign, the most connected and most easily travelled crossroads, and the first few metres around them, are where scats are most likely to be found, often on raised objects.

- The flip side: scat surveys done only on roads, or mostly at crossroads, will find many scats in a few places and may not show how wolves use the rest of their territory.

## Cite this summary

Lawrence, K. 2026. Summary of: The importance of crossroads in faecal marking behaviour of the wolves (Canis lupus) (Barja and de Miguel et al., 2004). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/2004-barja-de-miguel-importance-crossroads-faecal-marking/