Article

Article name INTERSPECIFIC SCENT MARKING: AN EXPERIMENTAL TEST OF THE INFLUENCE OF ADDED LION AND SPOTTED HYENA SCATS ON LEOPARD BEHAVIOUR IN SOUTH AFRICA
Authors

Agathe S.S. Gervais, MSc Student in the Mammal Research Institute of the University of Pretoria (Lynwood Road, Hatfield, Pretoria, 0083, South Africa); iD ORCID: https://orcid.org/0000-0002-6065-8790; e-mail: agathegervais24@gmail.com
Alexander R. Braczkowski, PhD, Research fellow at Griffith University's Centre for Planetary Health and Food Security (Ring Road, Nathan Campus, Griffith University, Australia); iD ORCID: https://orcid.org/0000-0002-0099-7803; email: alexander.braczkowski@gmail.com
Michael J. Somers, PhD, Professor in the Mammal Research Institute of the University of Pretoria (Lynwood Road, Hatfield, Pretoria, 0083, South Africa); iD ORCID: https://orcid.org/0000-0002-5836-8823; e-mail: michael.somers@up.ac.za

Reference to article

Gervais A.S.S., Braczkowski A.R., Somers M.J. 2026. Macroinvertebrates of Arctic lakes from Kolguev Island (Nenets Autonomous Okrug, Russia): fauna and driving factors of assemblage diversity. Nature Conservation Research 11(4). https://dx.doi.org/10.24189/ncr.2026.028

Electronic Supplement 1. Additional information for the paper by Gervais et al. (2026) (Link)

Section Research articles
DOI https://dx.doi.org/10.24189/ncr.2026.028
Abstract

Solitary carnivores rely strongly on scent marking to communicate, and can be attracted or repelled by the scents of conspecifics or heterospecifics. We aimed to understand the effects of intra- and interspecific olfactory interactions on Panthera pardus (hereinafter – leopard), and to test whether heterospecific scents could be manipulated as a tool to deter leopards. We monitored leopard scent-marking activity in the Sabi Sands Game Reserve, South Africa, by camera-trapping 43 marking sites (1450 trap nights) over a 61-day period in the 2021 dry season, of which a 38-day experimental phase used 27 sites. We addressed three objectives: (1) characterising the intraspecific communicative functions of marking; (2) evaluating the influence of dominant-competitor scents on leopard behaviour; and (3) assessing the potential of these scents as a deterrent. To simulate the presence of Panthera leo (hereinafter – lion) and Crocuta crocuta (hereinafter – spotted hyena), we added freshly collected scats to treatment sites and compared leopard visit interval, stay duration, and behaviour duration with control sites (no scats) using generalised linear mixed-effects models (GLMEMs). At the intraspecific level, 40% of consecutive visits to marking sites involved leopards of different sexes. Contrary to our prediction of male-dominated marking, females scent marked significantly more than males, suggesting that leopard scent marks serve both mating and territorial-advertisement functions. Contrary to our prediction of avoidance, leopard visit frequency was unchanged at sites with lion scats, although stay duration was significantly longer, suggesting a more complex olfactory response than simple avoidance. Our results support the prediction that leopards use scent-marking sites to gather information about neighbouring conspecifics and lions and to signal their own presence. We did not find strong evidence that lion or spotted hyena scats can deter leopards or be used as a wildlife-management tool. These findings highlight the role of olfactory communication in territorial behaviour and both its potential and its limitations for conservation.

Keywords

behavioural ecology, dear enemy effect, generalised linear mixed-effects models, intraguild interactions, translocated faeces

Artice information

Received: 08.10.2024. Revised: 10.07.2026. Accepted: 31.07.2026.

The full text of the article
References

Allen M.L., Wittmer H.U., Houghtaling P., Smith J., Elbroch L.M., 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

Allen M.L., Yovovich V., Wilmers C.C. 2016a. Evaluating the responses of a territorial solitary carnivore to potential mates and competitors. Scientific Reports 6: 27257. DOI: 10.1038/srep27257

Allen M.L., Wittmer H.U., Setiawan E., Jaffe S., Marshall A.J. 2016b. Scent marking in Sunda clouded leopards (Neofelis diardi): novel observations close a key gap in understanding felid communication behaviours. Scientific Reports 6: 35433. DOI: 10.1038/srep35433

Apps P.J. 2013. Are mammal olfactory signals hiding right under our noses?. Naturwissenschaften 100(6): 487–506. DOI: 10.1007/s00114-013-1054-1

Apps P.J., Mmualefe L., Jordan N.R., Golabek K.A., McNutt J.W. 2014. The “tomcat compound” 3-mercapto-3-methylbutanol occurs in the urine of free-ranging leopards but not in African lions or cheetahs. Biochemical Systematics and Ecology 53: 17–19. DOI: 10.1016/j.bse.2013.12.013

Arnold J., Soulsbury C.D., Harris S. 2011. Spatial and behavioral changes by red foxes (Vulpes vulpes) in response to artificial territory intrusion. Canadian Journal of Zoology 89(9): 808–815. DOI: 10.1139/z11-069

Asa C.S. 1993. Relative contributions of urine and anal-sac secretions in scent marks of large felids. American Zoologist 33(2): 167–172. DOI: 10.1093/icb/33.2.167

Ausband D.E., Mitchell M.S., Bassing S.B., White C. 2013. No trespassing: using a biofence to manipulate wolf movements. Wildlife Research 40(3): 207–216. DOI: 10.1071/WR12176

Bailey T.N. 1993. The African leopard: Ecology and behavior of a solitary felid. New York: Columbia University Press. 429 p.

Balme G.A., Hunter L.T. 2013. Why leopards commit infanticide. Animal Behaviour 86(4): 791–799. DOI: 10.1016/j.anbehav.2013.07.019

Balme G.A., Slotow R., Hunter L.T.B. 2009. Impact of conservation interventions on the dynamics and persistence of a persecuted leopard (Panthera pardus) population. Biological Conservation 142(11): 2681–2690. DOI: 10.1016/j.biocon.2009.06.020

Balme G.A., Hunter L.T., Goodman P., Ferguson H., Craigie J., Slotow R. 2010. An adaptive management approach to trophy hunting of leopards (Panthera pardus): A case study from KwaZulu-Natal, South Africa. In: D.W. Macdonald, A.J. Loveridge (Eds.): Biology and conservation of wild felids. Oxford: Oxford University Press. P. 341–352.

Balme G.A., Hunter L., Braczkowski A.R. 2012. Applicability of age-based hunting regulations for African leopards. PLoS ONE 7(4): e35209. DOI: 10.1371/journal.pone.0035209

Balme G.A., Batchelor A., de Woronin Britz N., Seymour G., Grover M., Hes L., Macdonald D.W., Hunter L.T.B. 2013. Reproductive success of female leopards Panthera pardus: the importance of top-down processes. Mammal Review 43(3): 221–237. DOI: 10.1111/j.1365-2907.2012.00219.x

Balme G.A., Pitman R.T., Robinson H.S., Miller J.R.B., Funston P.J., Hunter L.T.B. 2017. Leopard distribution and abundance is unaffected by interference competition with lions. Behavioral Ecology 28(5): 1348–1358. DOI: 10.1093/beheco/arx098

Balme G., Rogan M., Thomas L., Pitman R., Mann G., Whittington-Jones G., Midlane N., Broodryk M., Broodryk K., Campbell M., Alkema M., Wright D., Hunter L. 2019. Big cats at large: Density, structure, and spatio-temporal patterns of a leopard population free of anthropogenic mortality. Population Ecology 61(3): 256–267. DOI: 10.1002/1438-390X.1023

Banks P.B., Daly A., Bytheway J.P. 2016. Predator odours attract other predators, creating an olfactory web of information. Biology Letters 12(5): 20151053. DOI: 10.1098/rsbl.2015.1053

Bates D., Mächler M., Bolker B., Walker S. 2015. Fitting linear mixed-effects models using lme4. Journal of Statistical Software 67(1): 1–48. DOI: 10.18637/jss.v067.i01

Bolker B.M., Brooks M.E., Clark C.J., Geange S.W., Poulsen J.R., Stevens M.H.H., White J.S.S. 2009. Generalized linear mixed models: a practical guide for ecology and evolution. Trends in Ecology and Evolution 24(3): 127–135. DOI: 10.1016/j.tree.2008.10.008

Bothma J. du P. 1998. A review of the ecology of the southern Kalahari leopard. Transactions of the Royal Society of South Africa 53(2): 257–266. DOI: 10.1080/00359199809520392

Bothma J. du P., Coertze R.J. 2004. Scent-marking frequency in southern Kalahari leopards. South African Journal of Wildlife Research 34(2): 163–169. DOI: 10.10520/EJC117190

Bothma J. du P., le Riche E.A.N. 1984. Aspects of the ecology and the behaviour of the leopard Panthera pardus in the Kalahari desert. Koedoe 27: 259–279. DOI: 10.4102/koedoe.v27i2.585

Bothma J. du P., le Riche E.A.N. 1995. Evidence of the use of rubbing, scent-marking and scratching-posts by Kalahari leopards. Journal of Arid Environments 29(4): 511–517. DOI: 10.1016/S0140-1963(95)80023-9

Cornhill K.L., Kerley G.I.H. 2020a. Cheetah communication at scent-marking sites can be inhibited or delayed by predators. Behavioral Ecology and Sociobiology 74: 21. DOI: 10.1007/s00265-020-2802-9

Cornhill K.L., Kerley G.I.H. 2020b. Cheetah behaviour at scent-marking sites indicates differential use by sex and social rank. Ethology 126(10): 976–986. DOI: 10.1111/eth.13071

du Preez B.D. 2014. The impact of intraguild competition with lion Panthera leo on leopard Panthera pardus behavioural ecology. PhD Thesis. Oxford: University of Oxford. 262 p.

Eisenberg J.F. 1970. A splendid predator does its own thing untroubled by man. Smithsonian 1(6): 48–53.

Eklund A., López-Bao J.V., Tourani M., Chapron G., Frank J. 2017. Limited evidence on the effectiveness of interventions to reduce livestock predation by large carnivores. Scientific Reports 7: 2097. DOI: 10.1038/s41598-017-02323-w

Fattebert J., Dickerson T., Balme G., Slotow R., Hunter L. 2013. Long-distance natal dispersal in leopard reveals potential for a three-country metapopulation. South African Journal of Wildlife Research 43(1): 61–67. DOI: 10.3957/056.043.0108

Ferrero D.M., Lemon J.K., Fluegge D., Pashkovski S.L., Korzan W.J., Datta S.R., Spehr M., Fendt M., Liberles S.D. 2011. Detection and avoidance of a carnivore odor by prey. Proceedings of the National Academy of Sciences of the United States of America 108(27): 11235–11240. DOI: 10.1073/pnas.1103317108

Fisher J. 1954. Evolution and bird sociality. In: J. Huxley, A.C. Hardy, E.B. Ford (Eds.): Evolution as a process. London: Allen & Unwin. P. 71–83.

Fortescue A.K. 1997. The use of remote sensing and geographic information system (GIS) techniques, to interpret savanna ecosystem patterns in the Sabi Sand game reserve, Mpumalanga province. PhD Thesis. Grahamstown: Rhodes University. 140 p.

Friedmann Y., Traylor-Holzer K. 2008. Leopard (Panthera pardus) case study. WG-5 Mammals, 4. Johannesburg: CITES. 25 p.

Garvey P.M., Glen A.S., Pech R.P. 2016. Dominant predator odour triggers caution and eavesdropping behaviour in a mammalian mesopredator. Behavioral Ecology and Sociobiology 70(4): 481–492. DOI: 10.1007/s00265-016-2063-9

Garvey P.M., Glen A.S., Clout M.N., Wyse S.V., Nichols M., Pech R.P. 2017. Exploiting interspecific olfactory communication to monitor predators. Ecological Applications 27(2): 389–402. DOI: 10.1002/eap.1483

Grimbeek A.M. 1992. The ecology of the leopard (Panthera pardus) in the Waterberg. PhD Thesis. Pretoria: University of Pretoria. 166 p.

Hayward M.W., Slotow R. 2009. Temporal partitioning of activity in large African carnivores: tests of multiple hypotheses. South African Journal of Wildlife Research 39(2): 109–125. DOI: 10.3957/056.039.0207

Jackson C.R., McNutt J.W., Apps P.J. 2012. Managing the ranging behaviour of African wild dogs (Lycaon pictus) using translocated scent marks. Wildlife Research 39(1): 31–34. DOI: 10.1071/WR11070

Jacobson A.P., Gerngross P., Lemeris J.R., Schoonover R.F., Anco C., Breitenmoser-Würsten C., Durant S.M., Farhadinia M.S., Henschel P., Kamler J.F., Laguardia A., Rostro-García S., Stein A.B., Dollar L. 2016. Leopard (Panthera pardus) status, distribution, and the research efforts across its range. PeerJ 4: e1974. DOI: 10.7717/peerj.1974

Jones M.E., Apfelbach R., Banks P.B., Cameron E.Z., Dickman C.R., Frank A., McLean S., McGregor I.S., Müller-Schwarze D., Parsons M.H., Sparrow E., Blumstein D.T. 2016. A nose for death: Integrating trophic and informational networks for conservation and management. Frontiers in Ecology and Evolution 4: 124. DOI: 10.3389/fevo.2016.00124

Kennel C.F. 2021. The gathering anthropocene crisis. Anthropocene Review 8(1): 83–95. DOI: 10.1177/2053019620957355

Laundré J.W., Hernández L., Altendorf K.B. 2001. Wolves, elk, and bison: reestablishing the “landscape of fear” in Yellowstone National Park, U.S.A. Canadian Journal of Zoology 79(8): 1401–1409. DOI: 10.1139/z01-094

le Roex N., Mann G.K.H., Hunter L.T.B., Balme G.A. 2022. Relaxed territoriality amid female trickery in a solitary carnivore. Animal Behaviour 194: 225–231. DOI: 10.1016/j.anbehav.2022.09.022

Lennox R.J., Gallagher A.J., Ritchie E.G., Cooke S.J. 2018. Evaluating the efficacy of predator removal in a conflict-prone world. Biological Conservation 224: 277–289. DOI: 10.1016/j.biocon.2018.05.003

MacDonald D.W. 1980. Patterns of scent marking with urine and faeces amongst carnivore communities. Symposia of the Zoological Society of London 45: 107–139.

Melzheimer J., Heinrich S.K., Wasiolka B., Mueller R., Thalwitzer S., Palmegiani I., Weigold A., Portas R., Roeder R., Krofel M., Hofer H., Wachter B. 2020. Communication hubs of an asocial cat are the source of a human-carnivore conflict and key to its solution. Proceedings of the National Academy of Sciences of the United States of America 117(52): 33325–33333. DOI: 10.1073/pnas.2002487117

Miller J.R.B., Pitman R.T., Mann G.K.H., Fuller A.K., Balme G.A. 2018. Lions and leopards coexist without spatial, temporal or demographic effects of interspecific competition. Journal of Animal Ecology 87(6): 1709–1726. DOI: 10.1111/1365-2656.12883

Morrison M.L., Block W.M., Strickland M.D., Collier B.A., Peterson M.J. 2008. Wildlife Study Design. 2nd ed. New York: Springer. 587 p.

Palomares F., Caro T.M. 1999. Interspecific killing among mammalian carnivores. American Naturalist 153(5): 492–508. DOI: 10.1086/303189

Pirie T.J., Thomas R.L., Reilly B.K., Fellowes M.D. 2014. Social interactions between a male leopard (Panthera pardus) and two generations of his offspring. African Journal of Ecology 52(4): 574–576. DOI: 10.1111/aje.12154

R Core Team. 2017. R: A language and environment for statistical computing (ver. 3.4.3). Vienna: R Foundation for Statistical Computing. Available from https://www.R-project.org

Rafiq K., Jordan N.R., Meloro C., Wilson A.M., Hayward M.W., Wich S.A., McNutt J.W. 2020. Scent-marking strategies of a solitary carnivore: boundary and road scent marking in the leopard. Animal Behaviour 161: 115–126. DOI: 10.1016/j.anbehav.2019.12.016

Reiger I. 1979. Scent rubbing in carnivores. Carnivore 2: 17–25.

Rutherford M.C., Mucina L., Lotter M.C., Bredenkamp G.J., Smit J.H.L., Scott-Shaw R., Hoare D.B., Goodman P.S., Besuidenhout H., Scott L., Ellis F., Powrie L.W., Siebert F., Mostert T.H., Henning B.J., Venter C.E., Camp K.G.T., Siebert S.J., Matthews W.S., Burrows J.E., Dobson L., van Rooyen N., Smidt E., Winter P.J.D., du Preez P.J., Ward R.A., Williamson S., Hurter P.J.H. 2006. Savanna Biome. In: L. Mucina, M.C. Rutherford (Eds.): The Vegetation of South Africa, Lesotho and Swaziland. Pretoria: South African National Biodiversity Institute. P. 440–529.

Sadleir R.M.F.S. 1966. Notes on reproduction in the larger Felidae. International Zoo Yearbook 6(1): 184–187. DOI: 10.1111/j.1748-1090.1966.tb01746.x

Schmidtz D., Willott E. 2012. Private landowners cooperate to sustain wildlife habitat: the case of the Sabi Sand game reserve. Journal of Applied Corporate Finance 24(2): 78–85. DOI: 10.1111/j.1745-6622.2012.00382.x

Smith J.L.D., McDougal C., Miquelle D. 1989. Scent marking in free-ranging tigers, Panthera tigris. Animal Behaviour 37(1): 1–10. DOI: 10.1016/0003-3472(89)90001-8

Smith T., Fitchett J.M. 2020. Drought challenges for nature tourism in the Sabi Sands Game Reserve in the eastern region of South Africa. African Journal of Range and Forage Science 37(1): 107–117. DOI: 10.2989/10220119.2019.1700162

Somers M., Rasa O.A.E., Apps P.J. 1990. Marking behaviour and dominance in Suni antelope (Neotragus moschatus). Zeitschrift für Säugetierkunde 55(5): 340–352.

Soso S.B., Koziel J.A. 2017. Characterizing the scent and chemical composition of Panthera leo marking fluid using solid-phase microextraction and multidimensional gas chromatography-mass spectrometry-olfactometry. Scientific Reports 7(1): 5137. DOI: 10.1038/s41598-017-04973-2

Sunquist M.E., Sunquist F. 2002. Leopard. In: Wild Cats of the World. Chicago: University of Chicago Press. P. 318–342.

Tobler M. 2015. Camera Base version 1.7 user guide. Available from http://www.atrium-biodiversity.org

Webster M.M., Rutz C. 2020. How STRANGE are your study animals?. Nature 582(7812): 337–340. DOI: 10.1038/d41586-020-01751-5

Wemmer C., Scow K. 1977. Communication in the Felidae with emphasis on scent marking and contact patterns. In: T.A. Sebeok (Ed.): How animals communicate. Bloomington: Indiana University Press. P. 749–766.

Zuur A.F., Ieno E.N., Walker N., Saveliev A.A., Smith G.M. 2009. Mixed effects models and extensions in ecology with R. New York: Springer. 574 p.