Overmarking behaviour of zebra males: no scent masking, but a group cohesion function across three species

  • Jan PluháčekEmail author
  • Vladimíra Tučková
  • Sarah R. B. King
Original Article


Among many hypotheses in the literature that explain overmarking in mammals, most studies favour a sexual selection hypothesis. However, results in the literature are conflicting. In this study, we tested two hypotheses that could explain overmarking by males: (i) as a part of sexual selection, more specifically to mask scent of receptive females and (ii) as a form of communication serving to aid group cohesion. We observed each of the three zebra species in eight different herds at four zoos. In total, we recorded 1395 eliminations (760 defecations, 635 urinations) performed by 78 individuals including 8 stallions. Stallions investigated 248 eliminations and overmarked 124. The rate of overmarking by stallions was higher than those of all other sex and age categories. Stallions of all species overmarked all age and sex categories, except Grévy’s zebra stallions did not overmark foal eliminations. In contrast to our first hypothesis, when attracted to the elimination, stallions overmarked non-oestrus females more often than oestrus ones. Thus, our results did not support the hypothesis that overmarking by males has a sexual selection function, but it could be explained by the group cohesion hypothesis. Based on our results, it seems that overmarking by equid males plays a greater role in intra-specific communication than in intra-sexual competition. In addition, this behaviour might play different roles in different species based on their social organisation.

Significance statement

Overmarking is ubiquitous among terrestrial mammals; however, the function of this behaviour has not been fully explained. In addition, previous studies came to differing conclusions. In this study, we tested two hypotheses possibly explaining overmarking by adult males in three equid species. We found that adult males overmarked at a much higher rate than individuals of all other sex and age categories. In contrast to many other studies, our results did not support the hypothesis that overmarking by males has a sexual selection function, but it could be explained by a group cohesion hypothesis.


Competition Defecation Equus Marking behaviour Olfactory communication Scent marking 



The authors gratefully acknowledge the assistance of the Zoological Garden staff at Dvůr Králové, Brno, Liberec, Ostrava, and Ústí nad Labem, in particular Luděk Čulík, Markéta Čulíková, Aleš Kopecký, Jiří Soumar, Miroslava Kubelková, Miroslava Doležalová, Pavel Moucha, Radek Hlávka, Lubomír Melichar, Jiří Vítek, Petra Padalíková, Pavel Král, Jaroslav Novák, Rostislav Střižík, Eva Zajoncová, and Lenka Málcová. We thank Radka Šárová, Luděk Bartoš, and Jitka Bartošová for their helpful comments on the earlier draft of the manuscript. The manuscript was highly improved by comments of four anonymous reviewers.

Funding information

This work was supported by the Ministry of Agriculture of the Czech Republic (MZE-RO0719) and by grant no. 2011/008 of the Student Grant Agency of the University of South Bohemia.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

All applicable international, national, and institutional guidelines for the care and use of animals were followed. This research was carried out in line with all legislation of European Union and the Czech Republic. We received all appropriate permissions from the zoo authorities where the observations were performed. As this was an observational study with no manipulation or disturbance of the animals no animal care and use protocols were required.

Supplementary material

265_2019_2744_MOESM1_ESM.docx (25 kb)
ESM 1 (DOCX 24 kb)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  1. 1.Department of EthologyInstitute of Animal SciencePragueCzech Republic
  2. 2.Ostrava ZooOstravaCzech Republic
  3. 3.Departement of Biology and Ecology, Faculty of ScienceUniversity of OstravaOstravaCzech Republic
  4. 4.Department of Zoology, Faculty of ScienceUniversity of South BohemiaCeske BudejoviceCzech Republic
  5. 5.Natural Resource Ecology Laboratory, Warner College of Natural ResourcesColorado State UniversityFort CollinsUSA

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