# The functional role of scent marking in the social organization of large sympatric neotropical felids

> In a Belizean rainforest, pumas and jaguars share the same trails but leave very different messages. By walking the trails for scrapes and filming the busiest marking spots, researchers found that a few resident male pumas made most of the scrapes and scraped even more when rivals or females had passed by, while jaguars mostly sprayed and left little to see. Peccaries, ocelots and even a tayra stopped to check the big cats’ marks.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/2019-wooldridge-foster-functional-role-scent-marking/

## Citation and access

- **Authors:** Rebecca L. Wooldridge, Rebecca J. Foster and Bart J. Harmsen
- **Year published:** 2019
- **Journal or source:** Journal of Mammalogy
- **Full citation:** Wooldridge, R. L., Foster, R. J. and Harmsen, B. J. 2019. The functional role of scent marking in the social organization of large sympatric neotropical felids. Journal of Mammalogy 100(2): 445–453.
- **DOI:** 10.1093/jmammal/gyz055
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.1093/jmammal/gyz055

## Tags

- **Species:** Mammals > Jaguar; Mammals > Mountain lion; Mammals > Ocelot
- **Region:** North America > Belize
- **Tracking:** Individual identification; Sign surveys; Spoor age estimation; Tracking; Tracks and signs identification
- **Behaviour and ecology:** Behaviour; Carnivore coexistence; Communication; Intraguild interactions; Predator avoidance
- **Methods and technology:** Camera traps; Chi-square test; Global Positioning System (GPS) collars; Mann–Whitney U test
- **Data type:** New field data
- **Observer qualification:** Not reported
- **Discipline:** Behaviour and ecology; Wildlife tracking
- **Year published:** 2019

## Detailed keywords

- **Species:** Studied: puma (Puma concolor), also called mountain lion, and jaguar (Panthera onca); ocelot (Leopardus pardalis) scent marks and the animals visiting them were also recorded. Recorded at scrape clusters within 28 days of a scent mark: margay (Leopardus wiedii, spelled “weidii” in the paper’s table), jaguarundi (Puma yaguaroundi), gray fox (Urocyon cinereoargenteus), coatimundi (Nasua nasua), raccoon (Procyon lotor), tayra (Eira barbara), greater grison (Galictis vittata), white-lipped peccary (Tayassu pecari), collared peccary (Pecari tajacu), red brocket deer (Mazama americana), tapir (Tapirus terrestris) and northern tamandua (Tamandua mexicana). Detected but left out of the analysis because foraging could not be told apart from investigation: paca (Paca cuniculus), nine-banded armadillo (Dasypus novemcinctus), hog-nosed skunk (Conepatus leuconotus) and common opossum (Didelphis virginiana). Botfly (Dermatobia hominis) swellings helped identify individual pumas.
- **Location and habitat:** About 10 km² in the eastern half of the Cockscomb Basin Wildlife Sanctuary, Belize, Central America (16°47′N, 88°36′W), part of the Maya Mountain Massif. Lowland subtropical moist broadleaf forest (425 km² sanctuary, logged until the 1980s, protected since 1986), 50 to 1,133 m elevation, about 2,700 mm rain a year. Surveys ran along three former logging roads, now tourist trails without vehicles, 2 to 5 m wide and 2.7 to 8.6 km long.
- **Topics:** scent marking; scrapes; spraying; rubbing and rolling; urine marking; olfactory investigation (sniffing) and flehmen response; scrape clusters; scrape degradation and how long scrapes stay visible; social organization and dominance hierarchy; resident versus transient males; return intervals to trails; female mate choice and courtship; eavesdropping on scent marks by other species; interspecific investigation; scent transference (a tayra rubbing in puma scat); sympatric large felids; avoidance between jaguars and pumas.
- **Statistics or models:** Mann–Whitney U test (scrape rates of male pumas versus male jaguars); generalized linear mixed models with a Poisson distribution and log link in the R package lme4, with individual and observation-level random effects for overdispersion (checked with blmeco); likelihood ratio tests against null models (lmtest); chi-square tests, and Fisher’s exact tests for small samples; a binomial general linear model for female mate choice. R version 3.3.2, significance at P < 0.05. Inconsistencies in the paper: (1) the text says 10 species were detected within 28 days of ocelot scent marks, but Table 3 lists 9; (2) the text gives the female-presence effect on male puma return interval as P < 0.05, while the Figure 2 caption gives P = 0.05; (3) the Discussion says ocelots may use larger cats’ marks “to avoid larger conspecifics”, where other species (not conspecifics) appear to be meant.
- **Data type:** New field data. Daily scrape surveys for 20 consecutive days and weekly surveys for 13 weeks, May to September 2016; 20 camera traps (2,060 trap-nights) May to September 2016; video traps at scrape clusters May 2016 to July 2017 (9,420 trap-nights). In all, 50 locations and 9,911 functional trap days.

## Synopsis

### 1. What they did

Solitary cats that live far apart, like pumas and jaguars, keep in touch through scent marks: scrapes on the ground, urine sprayed on vegetation, and rubbing or rolling. Most of what was known about how cats use these marks came from pumas and Eurasian lynx in the Northern Hemisphere. This study looked at a tropical forest where two large cats, pumas and the larger jaguars, share the same trails.

In the Cockscomb Basin Wildlife Sanctuary, Belize, the researchers:

- Searched trails for scrapes. Two researchers walked either side of three trails scanning the ground, daily for 20 days and weekly for 13 weeks (May to September 2016). They recorded each scrape’s size and location and marked it with small sticks so it was not counted twice.

- Measured how long scrapes stay visible. They photographed 11 scrapes for up to 35 days and scored how visible the bare soil and the raked pile remained.

- Linked scrapes to individual cats. Twenty camera traps about 700 m apart identified which cats used each 700 m stretch of trail. Jaguars were identified by their rosette patterns with the HotSpotter software; pumas by scars, markings, botfly swellings and Global Positioning System (GPS) collars.

- Filmed scrape clusters. Video cameras at places with more than three scrapes in 9 m² recorded how cats and other animals marked and investigated, from May 2016 to July 2017.

They then tested whether male pumas and jaguars returned to trails sooner, or scraped more, when other cats of their species had been there in the previous 28 days, and whether female pumas paired with the males that scraped most.

### 2. What they found

Scrapes fade within weeks

- After 7 days, median soil visibility was 40%. After 28 days no bare soil was visible in any monitored scrape, and the pile was only just visible in 27%. The authors therefore assumed scrapes stop working as signals after 28 days.

Who marked, and how

- They identified 22 jaguars (seven males, nine females, five juveniles, one of unknown sex) and 16 pumas (10 males, six females).

- They found 703 scrapes. Of the 224 that could be linked to individuals, 88% were made by two resident male pumas.

- Male pumas scraped far more than male jaguars: about 5.1 scrapes per km against 0.4.

- On video, pumas mostly scraped (85% of their 221 marking events) or rubbed and rolled (14%); two female pumas urinated on male scrapes. Jaguars mostly sprayed vegetation (four of six marking events), and all nine ocelot marks were sprays.

- Only a few individuals marked. Resident male pumas marked on 36% of visits to scrape clusters, transient males on 11%, and females on 6%. Only two of the seven male jaguars were filmed marking.

Responding to others

- Male pumas came back to a trail stretch about 6 days sooner when another male had been there in the previous 28 days, and about 4.5 days sooner after a female.

- Male jaguars came back about 7.3 days sooner after another male, but not after a female.

- Male pumas scraped about three times as much after another male had been present, and about twice as much after a female. They also sniffed more often.

Mate choice

- One resident male puma was filmed travelling and courting with three different females. He scraped more (7.1 scrapes per 700 m) than males not seen with females (1.5).

- Two resident male jaguars, the only male jaguars filmed marking, were each seen courting the same female.

Other species

- Nine species investigated scent marks. Smaller species investigated the marks of larger ones (a puma and ocelots sniffed jaguar marks), but jaguars never investigated puma or ocelot marks, and no animal counter-marked another species’ mark.

- Collared peccaries, prey of both big cats, had the highest investigation rate of puma marks among other species.

- A tayra rubbed their head and neck in puma scat.

### 3. What they concluded

- Jaguars and pumas mark in different ways: jaguars mainly spray, pumas mainly scrape. Pumas scrape in the tropics as they do in temperate areas, which suggests scraping is used across their range whatever the ecosystem. Jaguar marking was probably underestimated, because sprays and rubbing leave little to see.

- Marking in these cats appears tied to a dominance hierarchy. Transient males seemed to avoid marking on the trails, perhaps so resident males would not notice them, and marked only when a female had recently been there.

- Because male pumas scraped more when rivals and potential mates were around, the authors infer that scraping signals dominance to both. Females may choose the males that scrape most, which may act as an honest signal of quality and could lower the risk of infanticide.

- Jaguars and pumas seem to be constantly assessing, and reacting to, the presence and marks of competitors and potential mates, which helps maintain complex social structures.

- Smaller species may use larger cats’ marks to avoid them, and prey or smaller carnivores may even pick up a predator’s scent as protection. Big cat scent marks may play an important role in the wider forest community.

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

The study rests on a sign survey. Two researchers walked the trails and identified scrapes by their raked pile of leaves and soil at one end and a patch of bare soil, often split by a ridge in the middle. They measured and mapped each scrape and tracked how its visible features faded over time, to judge how old a scrape could be and still be found. Scrapes were assigned to individual cats by matching their location with camera-trap detections on the same 700 m stretch of trail, not by footprints. No trailing was reported.

### 5. Tracker or observer certification

Not reported. Who identified the scrapes and how experienced they were is not described; E. Sanchez and S. Puls are thanked for help with the scrape surveys. Both jaguars and pumas make scrapes, so the species and individual behind each scrape were inferred from camera traps rather than from the sign itself.

### 6. Other technologies, skills or innovations

- Pantheracam V4 and V6 camera traps along the trails, and Browning Strike Force video traps at scrape clusters, recording up to 30 seconds by day and 10 seconds at night.

- HotSpotter pattern-recognition software to identify individual jaguars from their rosettes.

- A simple scoring method for scrape ageing: soil visibility as a percentage of the original, and pile visibility from 0 to 5.

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

- The paper does not draw conservation implications directly. It suggests that big cat scent marks may matter to the wider forest community, as other carnivores and prey species investigate them.

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

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

- The scrape-ageing results give trackers a field benchmark from this forest: bare soil had disappeared from half the scrapes by 14 days and from every monitored scrape by four weeks, while the raked pile lasted longer.

- Pumas and jaguars leave very different amounts of visible sign. A survey that counts only scrapes would mostly record pumas and miss most jaguar marking, which is sprayed on vegetation.

## Cite this summary

Lawrence, K. 2026. Summary of: The functional role of scent marking in the social organization of large sympatric neotropical felids (Wooldridge and Foster et al., 2019). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/2019-wooldridge-foster-functional-role-scent-marking/