# Top marks from top dogs: tandem marking and pair bond advertisement in African wild dogs

> When a dominant African wild dog leaves a urine mark, their partner often marks right on top of it. Watching 13 packs in northern Botswana, researchers found that who ends up “on top” depends on whether the pair are a true breeding couple or a brother and sister who inherited the pack. Sister-led pairs kept more of their marks apart, which hints that their scent may be a message to outside males.

Summary by Kersey Lawrence, Original Wisdom. Updated 9 November 2026. Page: https://www.originalwisdom.com/tracking-literature/2014-jordan-apps-top-marks-top-dogs/

## Citation and access

- **Authors:** Neil R. Jordan, Peter J. Apps, Krystyna A. Golabek and John W. McNutt
- **Year published:** 2014
- **Journal or source:** Animal Behaviour
- **Full citation:** Jordan, N. R., Apps, P. J., Golabek, K. A. and McNutt, J. W. 2014. Top marks from top dogs: tandem marking and pair bond advertisement in African wild dogs. Animal Behaviour 88: 211–217.
- **DOI:** 10.1016/j.anbehav.2013.12.001
- **Access status:** Publisher link only
- **Where to find it:** https://doi.org/10.1016/j.anbehav.2013.12.001

## Tags

- **Species:** Mammals > African wild dog
- **Region:** Africa > Botswana
- **Behaviour and ecology:** Behaviour; Communication; Denning
- **Methods and technology:** Akaike Information Criterion (AIC); Binomial test; Chi-square test; Direct observation; Radio telemetry
- **Data type:** New field data
- **Observer qualification:** Not needed
- **Discipline:** Behaviour and ecology
- **Year published:** 2014

## Detailed keywords

- **Species:** Studied: African wild dog (Lycaon pictus), the dominant breeding pair in each of 13 packs. Other species discussed from the literature only: grey wolf (Canis lupus), coyote (Canis latrans), domestic dog (Canis familiaris), golden jackal (Canis aureus), red fox (Vulpes vulpes), bush dog (Speothos venaticus), Ethiopian wolf (Canis simensis), meerkat (Suricata suricatta), banded mongoose (Mungos mungo), leopard (Panthera pardus), Kirk’s dikdik (Madoqua kirkii), klipspringer (Oreotragus oreotragus), oribi (Ourebia ourebi), plains zebra (Equus burchelli), meadow vole (Microtus pennsylvanicus), prairie vole (Microtus ochrogaster), golden hamster (Mesocricetus auratus) and house mouse (Mus domesticus).
- **Location and habitat:** Northern Botswana: a study area of about 2600 km² at about 950 m elevation, bordered by the Okavango Delta and including the Moremi Game Reserve and Wildlife Management Areas (including land of the Sankuyo community).
- **Topics:** scent marking with urine; tandem marking (both members of a pair marking the same spot in quick succession); overmarking; the “top mark” (the most recent mark at a site); pair bond formation, maintenance and advertisement; mate guarding; relatedness of the dominant pair (full siblings versus unrelated pairs); self-advertisement to attract immigrant mates; “test and assess” strategy; dispersal by both sexes; leg postures (cocked leg, raised leg, handstand); seasonal patterns, including the breeding period and denning season; individual identification from coat pattern, ear notches and tail stripes.
- **Statistics or models:** Generalized linear mixed models (GLMMs) with binomial errors and logit link, with pack, individual and site as random terms; model selection by Akaike’s information criterion (AIC, AICc) and Akaike weights; model averaging (MuMIn package in R); binomial tests of proportions with continuity correction; chi-square tests; Welch two-sample t test. Possible inconsistencies in the paper: (1) the number of sites with unrelated pairs is given as 418 and as 417 in the text (85 related plus 418 makes 503, not 502), and as 427 in the Figure 1 caption; (2) Figure 2 gives 501 sites and 153 female-initiated bouts, against 502 sites and 154 female-initiated sites in the text; (3) the dominant marks that never became tandem marks add up to 193 sites (21 related plus 172 unrelated), against 237 single marks reported earlier; (4) the methods name “initiator identity” as a random term, while Table 1 lists “investigator identity” (N = 21).
- **Data type:** New field data: observations of 13 free-ranging packs, May 2011 to September 2012, in a population studied since 1989.

## Synopsis

### 1. What they did

In many pair-bonded canids and antelopes, both members of a pair scent-mark the same spot, one on top of the other. This is called tandem marking. It is usually thought to help a pair bond form, or to advertise to others that a mated pair holds the territory. The authors wanted to know who marks first and who ends up “on top”, because the top mark at a site seems to matter most to the animals that visit it later.

They watched the dominant pairs of 13 free-ranging African wild dog packs in northern Botswana between May 2011 and September 2012. Almost all the dogs (95.5% of 112) were of known age and origin, and each was recognized by their unique coat pattern, ear notches and tail stripes. One to four dogs per pack wore radio collars so the packs could be found from the air and from a vehicle.

From a vehicle, the observers recorded on video or datasheets every urine mark the dominant pair made with a leg posture (cocked leg, raised leg or handstand), every time a dog sniffed or licked a mark, and every overmark, until the pack left the site. Squat urinations and defecations were left out, as earlier work suggests they are not signals.

Two of the 13 dominant pairs were full siblings: after a dominant died, a brother or sister of the survivor took over the role. These “related” pairs have no breeding opportunity inside the pack, so the authors compared them with the 11 “unrelated” pairs.

### 2. What they found

Who marks and who ends on top

- The dominant pairs marked at 502 sites. The dominant male started marking at 69.32% of sites and left the top mark at 80.29%.

- Marks started by females were much more likely to be tandem-marked (85.06% of 154) than marks started by males (40.23% of 348).

- Males always tandem-marked female marks that they sniffed. Females sometimes sniffed or passed close to a male’s mark and walked away without marking, which suggests an active choice.

- Both sexes were more likely to overmark a site where their partner’s scent was on top than one where their own scent was already on top. Marks that were already layered (double or triple marks) were sniffed and overmarked at similar rates to single marks, which shows that the top scent is what counts.

Related versus unrelated pairs

- Marks were more likely to become tandem marks in related pairs (75.29% of 85 sites) than in unrelated pairs (60.05% of 418 sites).

- Males were on top at the end of marking more often in unrelated pairs (82.73% of sites) than in related pairs (68.24%).

- In related pairs, females left a much larger share of the marks that were never tandem-marked (42.86% of 21, against 14.53% of 172 in unrelated pairs). Males did not pass close to the female marks they missed, so these females seem to have placed marks where the males would not find them.

- Marking bouts were about twice as long in related pairs (an average of 4.20 marks per bout) as in unrelated pairs (2.41 marks per bout).

Through the year

- Every month, a higher proportion of female-started marks than male-started marks were tandem-marked.

- In June and July, when the dominant female stayed near the den, almost all of her marks were tandem-marked, and almost none of the male’s were.

- Males tended to tandem-mark more during the female’s short receptive period in April.

### 3. What they concluded

- Dominant African wild dogs pay attention to which scent is on top of a marking site and are motivated to put their own scent in that top position.

- Males were more likely to cover their partner’s marks than the other way round, which fits the idea that advertising the pair bond, and guarding a mate, matters more to males. The authors link this to males being more at risk: male wild dogs disperse farther, so resident packs may meet roaming males more often than roaming females.

- In unrelated pairs, females may use a “test and assess” strategy: by marking a lot, they show whether their male can keep covering their scent, and the pattern advertises both a working pair bond and a high-quality male who is not available.

- In related pairs, where a working pair bond would not help, females seem to advertise themselves, perhaps to attract an unrelated male from outside the pack. They kept more of their marks apart from the male’s, and marking bouts were longer, perhaps from competitive overmarking within the pair.

- It is unclear why related males keep marking as if advertising a pair bond. Experiments presenting single and tandem marks to potential immigrants are needed.

- The results rest on only two related pairs, so more work is needed to test these ideas.

### 6. Other technologies, skills or innovations

- Radio collars (Vectronic and Sirtrack), located by radio tracking from the air and from a vehicle. Dogs were darted from a vehicle and immobilized to fit the collars; the authors report no negative effects observed.

- Individual identification from drawings and photographs of each dog’s coat pattern, ear notches and tail stripes.

- Video and datasheet recording by critical incident sampling (recording every marking event as it happens).

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

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

- The study did not use tracking, but it shows how much information can sit in one urine mark site. A site that looks like a single wet patch may hold several layered marks from both members of the dominant pair, and in wild dogs the top mark is usually the dominant male’s.

- The leg postures described (cocked leg, raised leg and handstand) may help trackers who find urine marks to think about which animal and what kind of signal left them.

## Cite this summary

Lawrence, K. 2026. Summary of: Top marks from top dogs: tandem marking and pair bond advertisement in African wild dogs (Jordan and Apps et al., 2014). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/2014-jordan-apps-top-marks-top-dogs/