# The Role of Scent Marking in Mate Selection by Female Pumas (Puma concolor)

> A small pile of scraped-up soil can be a puma’s dating profile. Filming 30 community scrape sites in California’s Santa Cruz Mountains, researchers found that females visit in short bursts that seem to match their time in heat, and tend to choose the older, heavier males that visit and scrape the most, likely the dominant residents. Some females mated with more than one male, which may help protect their kittens.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/2015-allen-wittmer-role-scent-marking-mate/

## Citation and access

- **Authors:** Maximilian L. Allen, Heiko U. Wittmer, Paul Houghtaling, Justine Smith, L. Mark Elbroch and Christopher C. Wilmers
- **Year published:** 2015
- **Journal or source:** PLOS ONE
- **Full citation:** Allen, M. L., Wittmer, H. U., Houghtaling, P., Smith, J., Elbroch, L. M. and Wilmers, C. C. 2015. The role of scent marking in mate selection by female pumas (Puma concolor). PLOS ONE 10(10): e0139087.
- **DOI:** 10.1371/journal.pone.0139087
- **Access status:** Open access (CC BY 4.0)
- **PDF:** https://www.originalwisdom.com/wp-content/uploads/2026/11/2015_Allen_and_Wittmer_etal_the-role-of-scent-marking-in-mate-selection-by-female-pumas.pdf

## Tags

- **Species:** Mammals > Mountain lion
- **Region:** North America > United States
- **Tracking:** Tracking; Tracks and signs identification
- **Behaviour and ecology:** Behaviour; Communication; Home range; Movement
- **Methods and technology:** Akaike Information Criterion (AIC); Analysis of variance (ANOVA); Camera traps; Global Positioning System (GPS) collars
- **Data type:** New field data
- **Observer qualification:** Not reported
- **Discipline:** Behaviour and ecology; Wildlife tracking
- **Year published:** 2015

## Detailed keywords

- **Species:** Studied: puma (Puma concolor), also called mountain lion: 39 GPS-collared pumas (20 females, 19 males), plus uncollared pumas recorded on camera. Also scent marking at the community scrapes but not studied: coyote (Canis latrans), bobcat (Lynx rufus), gray fox (Urocyon cinereoargenteus) and striped skunk (Mephitis mephitis).
- **Location and habitat:** Santa Cruz Mountains, California, United States: a 1,700 km² study area in parts of Santa Cruz, San Mateo and Santa Clara counties, bounded by the Pacific Ocean, the cities of San Jose and Santa Cruz, and Highway 101, and cut through by Highway 17. Public and private land, including California State Parks, Midpeninsula Regional Open Space District, Peninsula Open Space Trust and Santa Clara County Parks.
- **Topics:** mate selection (mate choice); scent marking; scraping; community scrapes as “billboards”; olfactory investigation; flehmen response; caterwauling; visitation bouts and oestrus; male residency and dominance; body mass and age of males; forays outside the home range; territorial transgression by non-resident males; polyandry (females mating with more than one male); infanticide avoidance; paternity confusion.
- **Statistics or models:** Mixed-model analysis of variance (ANOVA) with individual as a random effect (nlme package in R); 95% local convex hull (LoCoH) home ranges in ArcMap; two-tailed t-test assuming equal variances; stepwise Akaike Information Criterion (AIC) model selection (MASS package), with C-statistics and AIC weights. Inconsistencies in the paper: the abstract says 29 community scrapes, the methods say 30 community scrape areas; the abstract says “at least 14%” of females mated with more than one male, the results say 2 of 17 females (12%), and the discussion says 14.2% of the consorting documented (2 of 23 consorting events would be about 9%). The results cite “Video 5” for consorting, while the supporting video of consorting is S3 Video. In Table 2, the best 90-day model (total scrapes alone) has a lower C-statistic (0.72) than the second model (0.81), although it has the best AIC score.
- **Data type:** New field data: camera monitoring of community scrapes and GPS collar data, May 2011 to July 2013 (home ranges for July 2011 to June 2012 and July 2012 to June 2013). Data available at Dryad (doi:10.5061/dryad.6d5h2).

## Synopsis

### 1. What they did

Choosing a mate is hard to study in solitary carnivores, because they are secretive and rarely meet. Pumas mostly communicate through scent marking. The commonest form is scraping: the puma pushes up a small mound of soil and leaves with their hind feet, and sometimes urinates or defecates on it. Places where scrapes are clumped (at least 3 in 9 m²) and used by more than one puma are called community scrapes. The authors asked how female pumas use these sites to choose a mate.

They tested three ideas:

- Males would visit community scrapes regularly, to advertise to females and mark their territory, while females would visit irregularly, staying away while raising kittens and visiting when in oestrus (in heat).

- Females would make fewer trips outside their usual home range than males, since they mate only about once every two years, while males look for every chance to mate.

- Females would choose males that seem to be residents, which males advertise by how often they visit the scrapes. Because older, heavier males tend to be residents, females might also select for age and mass.

From May 2011 to July 2013 in the Santa Cruz Mountains of California, they:

- Collared pumas. They fitted Global Positioning System (GPS) collars to 39 pumas (20 females and 19 males), taking a location about every 4 hours.

- Found community scrapes. A computer method picked out places that collared pumas visited repeatedly with at least 7 days between visits (to leave out kill sites). Field teams checked these places, and found more scrapes during other site visits. In all they documented 299 community scrapes.

- Filmed them. They set one motion-triggered infrared video camera about 3 m from each of a selection of active community scrapes, recording 60-second clips. After pooling cameras less than 1 km apart and dropping sites watched for under 3 months, they had 30 community scrape areas, each watched for an average of 501 days.

- Identified the pumas on video. Collared pumas could be matched to their mass and age at capture. Uncollared pumas were sexed and aged from their body, and identified where possible from spot patterns on the inside of the upper leg, scars, kinked tails and old injuries. Young pumas and females with kittens were left out.

- Analysed visits and mate choice. They compared how often males and females returned to the same scrape, counted trips outside each puma’s home range, and tested which male traits and behaviours best predicted which male a female consorted with.

### 2. What they found

Visits to the community scrapes

- They watched 676 visits by adult males and 179 visits by adult females without kittens.

- Males visited regularly all year, on average every 17.7 days.

- Females visited in short bouts (40 bouts, plus 28 single visits). A bout lasted about 32 days, with about 3.7 visits, roughly every 12 days. Bouts happened at any time of year but were about a year apart (on average 367 days).

- During their bouts, females visited 1.5 times more often than males. Overall, males visited about twice as often as females, but this difference was not statistically significant.

- Females caterwauled (gave a loud, carrying call) in 28.2% of their visitation bouts.

Trips outside the home range

- Males made 6.7 trips outside their home range per year (range 2 to 12), significantly more than females at 2.0 (range 0 to 5).

- Three times, a male turned up at a scrape outside their home range for the first time during a female’s bout, then visited less and less once the bout ended.

Choosing a mate

- They recorded 23 consorting events (travelling together, or visiting the same scrape within 2 hours) involving 17 females.

- On camera, each female chose from an average of 1.7 males (range 1 to 3). This matched the number of collared males whose home ranges overlapped each female’s (1.7 and 2.0 in the two years).

- All chosen males of known age (8) were 3 to 7 years old.

- During a female’s bout, the best model of which male was chosen included how often the male visited, together with the male’s mass and age.

- Over the 90 days before consorting (about the length of a puma pregnancy), the best predictor was the total number of scrapes the male had made.

- Two of the 17 females (12%) consorted with two different males in a single bout. A genetic study in the same area found two fathers in one of eight litters.

- Three times, a non-resident male consorted with a female at a scrape that only one male had used before.

### 3. What they concluded

- Community scrapes work like “billboards”, where males advertise themselves and females choose between them.

- Females use several cues together. While actively choosing, they favour males that visit often and are heavier and older. Over the longer run, the male that scrapes the most is likely the dominant resident male, since young pumas rarely scrape. Choosing that male likely lowers the risk of the kittens later being killed by a male (infanticide).

- Female visits seem to match oestrus, with bouts that may cover two oestrus cycles. The authors caution that most female data came from uncollared pumas, which are harder to identify.

- Scent marking does not fully keep other males out. Males leave their home ranges more than once thought, possibly drawn by caterwauling females. When a female chooses a trespassing male, more genetic variety in the kittens may matter more than residency.

- Some females mate with more than one male. This may confuse paternity and so reduce infanticide, and may also increase genetic variety in the kittens.

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

No trailing was reported. The study rests on a sign: the scrape. Field teams visited places that GPS data showed collared pumas returned to again and again, and recognised community scrapes from the clumped scrapes there. They also found further scrapes during opportunistic site visits. They counted how many scrapes each puma made on video. Hounds were used to help capture the pumas.

### 5. Tracker or observer certification

Not reported. No observer qualification is described for finding and recognising scrapes in the field or for identifying individual pumas on video. The authors themselves note that identifying uncollared individuals is often unreliable and that female visit rates should be read with caution.

### 6. Other technologies, skills or innovations

- GPS collars (Vectronics Aerospace GPS Plus 1D), with a location about every 4 hours and a mean success rate of 76%.

- Motion-triggered infrared video cameras (Bushnell TrophyCam) recording 60-second clips, serviced every 1 to 3 months.

- A GPS-based method to pick out repeatedly visited places that might hold community scrapes, while leaving out kill sites.

- Identifying uncollared pumas on video from spot patterns on the inner upper leg, scars, tail kinks and old injuries.

- Ageing captured pumas by gum line recession.

- 95% local convex hull (LoCoH) home ranges, used to measure home range overlap and trips outside the home range.

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

The paper does not discuss conservation directly. It describes community scrapes as playing an integral role in puma communication and mate selection, and suggests they may matter even more for males to advertise dominance in species whose populations are clumped.

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

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

- Scrapes are among the commonest puma signs. This study gives them meaning: at a community scrape, a fresh scrape is most likely from a resident adult male, while repeated female visits over a few weeks may mark a female in oestrus.

- GPS showed where pumas kept returning, but people on the ground had to find and recognise the scrapes there. Sign interpretation was key to setting up the whole study.

## Cite this summary

Lawrence, K. 2026. Summary of: The Role of Scent Marking in Mate Selection by Female Pumas (Puma concolor) (Allen and Wittmer et al., 2015). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/2015-allen-wittmer-role-scent-marking-mate/