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Test of four hypotheses to explain the function of overmarking in foals of four equid species

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Abstract

Overmarking occurs when one individual places its scent mark directly on top of the scent mark of another individual. Although it is almost ubiquitous among terrestrial mammals, we know little about the function of overmarking. In addition, almost all studies on mammalian overmarking behaviour dealt with adult individuals. Reports on this behaviour in juveniles are extremely rare, yet may elucidate the function of this behaviour. We tested four mutually non-exclusive hypotheses which might explain this behaviour in juveniles: (1) conceal the individual’s scent identity, (2) announcement of association with other group members, especially the mother—i.e., sharing identity with the mother, (3) to prevent the next conception of the mother, i.e., parent-offspring conflict, and (4) an early expression of male sexual behaviour. We observed 43 foals (out of 108 individuals) from all African equid species (Equus africanus, E. grevyi, E. quagga, E. zebra) in five zoos. In total, we recorded 3340 eliminations; 260 of these events were overmarked by 38 individual foals representing all species. This represents one of the highest rates of overmarking ever recorded by mammalian juveniles. Foals of all species except African wild ass overmarked the mother more often than another herdmate: with male foals overmarked at a higher rate than female foals. Mothers preferred to overmark foals, but not exclusively their own foal. Our results provide support for the hypotheses that overmarking serves to share identity between foal and mother, and that it is an early expression of male sexual behaviour.

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References

  • Altmann J (1974) Observational study of behaviour: sampling methods. Behaviour 49:227–267

    Article  CAS  Google Scholar 

  • Brown RE, Macdonald DW (1985) Social odours in mammals, vol 2. Clarendon Press, Oxford

    Google Scholar 

  • Crowell-Davis SL, Houpt KA (1985) The ontogeny of flehmen in horses. Anim Behav 33:739–745

    Article  Google Scholar 

  • D’Aniello B, Semin GR, Alterisio A, Aria M, Scandurra A (2018) Interspecies transmission of emotional information via chemosignals: from humans to dogs (Canis lupus familiaris). Anim Cogn 21:67–78

    Article  PubMed  Google Scholar 

  • Feist JD, McCullough D (1976) Behaviour patterns and communication in feral horses. Zeit Tierpsychol 41:337–371

    Article  CAS  Google Scholar 

  • Ferkin MH, Pierce AA (2007) Perspectives on over-marking: is it good to be on top? J Ethol 25:107–116

    Article  Google Scholar 

  • Gosling LM (1982) A reassessment of the function of scent-marking in territories. Zeit Tierpsychol 60:89–118

    Article  Google Scholar 

  • Gosling LM (1985) The even-toed ungulates: order Artiodactyla. In: Brown RE, Macdonald DW (eds) Social odours in mammals, vol 2, Clarendon Press, Oxford, pp 550–618

    Google Scholar 

  • Grange S, Duncan P, Gaillard JM, Sinclair ARE, Gogan PJP, Packer C, Hofer H, East M (2004) What limits the Serengeti zebra population? Oecologia 140:523–532

    Article  PubMed  Google Scholar 

  • Gray ME, Cameron EZ, Peacock MM, Thain DS, Kirchoff VS (2012) Are low infidelity rates in feral horses due to infanticide? Behav Ecol Sociobiol 66:529–537

    Article  Google Scholar 

  • Hothersall B, Harris P, Sortoft L, Nicol CJ (2010) Discrimination between conspecific odour samples in the horse (Equus caballus). Appl Anim Behav Sci 126:37–44

    Article  Google Scholar 

  • Hughes NK, Kelley JL, Banks PB (2012) Dangerous liaisons: the predation risks of receiving social signals. Ecol Lett 15:1326–1339

    Article  PubMed  Google Scholar 

  • Hurst JL (1990a) Urine marking in populations of wild house mice Mus domesticus Rutty 2. Communications between females. Anim Behav 40:223–232

    Article  Google Scholar 

  • Hurst JL (1990b) Urine marking in populations of wild house mice Mus domesticus Rutty. 1. Communication between males. Anim Behav 40:209–222

    Article  Google Scholar 

  • Hurst JL (1990c) Urine marking in populations of wild house mice Mus domesticus Rutty. 3. Communication between sexes. Anim Behav 40:233–243

    Article  Google Scholar 

  • Johnson RP (1973) Scent marking in mammals. Anim Behav 21:521–535

    Article  Google Scholar 

  • Johnston RE (2008) Individual odors and social communication: Individual recognition, kin recognition, and scent over-marking. In: Brockmann HJ, Roper TJ, Naguib M, Wynne-Edwards KE, Barnard C, Mitani JC (eds) Advances in the Study of Behavior, vol 38, Academic Press, New York, pp 439–505

    Google Scholar 

  • Johnston RE, Chiang G, Tung C (1994) The information in scent over-marks of golden hamsters. Anim Behav 48:323–330

    Article  Google Scholar 

  • Jordan NR, Manser MB, Mwanguhya F, Kyabulima S, Ruedi P, Cant MA (2011a) Scent marking in wild banded mongooses: 1. Sex-specific scents and overmarking. Anim Behav 81:31–42

    Article  Google Scholar 

  • Jordan NR, Mwanguhya F, Furrer RD, Kyabulima S, Ruedi P, Cant MA (2011b) Scent marking in wild banded mongooses: 2. Intrasexual overmarking and competition between males. Anim Behav 81:43–50

    Article  Google Scholar 

  • Jordan NR, Mwanguhya F, Kyabulima S, Ruedi P, Hodge SJ, Cant MA (2011c) Scent marking in wild banded mongooses: 3. Intrasexual overmarking in females. Anim Behav 81:51–60

    Article  Google Scholar 

  • Kimura R (2000) Relationship of the type of social organization to scent-marking and mutual-grooming behaviour in Grevy’s (Equus grevyi) and Grant’s zebra (Equus burchelli bohmi). J Equine Sci 11:91–98

    Article  Google Scholar 

  • Kimura R (2001) Volatile substances in feces, urine and urine/marked feces of feral horses. Can J Anim Sci 81:411–420

    Article  CAS  Google Scholar 

  • King SRB, Gurnell J (2007) Scent-marking behaviour by stallion: an assessment of function in a reintroduced population of Przewalski horses (Equus ferus przewalskii). J Zool 272:30–36

    Article  Google Scholar 

  • Klingel H (1974) Soziale Organisation und Verhalten des Grevy-Zebras (Equus grevyi). Zeit Tierpsychol 36:36–70

    Google Scholar 

  • Klingel H (1975) A comparison of the social behaviour of the Equidae. In: Geist V, Walther F (eds) The papers of an international symposium on the behaviour of ungulates and its relation to management. IUCN, pp 124–132

  • Koivula M, Korpimäki E (2001) Do scent marks increase predation risk of microtine rodents? Oikos 95:275–281

    Article  Google Scholar 

  • Lanata A, Nardelli M, Valenza G, Baragli P, D’Aniello B, Alterisio A, Scandurra A, Semin GR, Scilingo EP (2018) A case for the interspecies transfer of emotions: a preliminary investigation on how humans odors modify reactions of the autonomic nervous system in horses. 40th Annual International Conference of the IEEE Engineering in Medicine and Biology Society (EMBC):522–525

  • Liang KY, Zeger SL (1986) Longitudinal data analysis using generalized linear models. Biometrika 73:13–22

    Article  Google Scholar 

  • Miquelle DG (1991) Are moose mice—the function of scent urination in moose. Am Nat 138:460–477

    Article  Google Scholar 

  • Moehlman PD (1985) The odd-toed ungulates: order Perrisodactyla. In: Brown RE, Macdonald DW (eds) Social odours in mammals, vol 2, Clarendon Press, Oxford, pp 531–549

    Google Scholar 

  • Moehlman PD (1998) Behavioral patterns and communication in feral asses (Equus africanus). Appl Anim Behav Sci 60:125–169

    Article  Google Scholar 

  • Müller-Schwarze D (2006) The chemical ecology of vertebrates. Cambridge University Press, Cambridge

    Book  Google Scholar 

  • Oom MdM, Reis MS (1986) Ecology, social organization and behaviour of the feral horses in the Peneda-Geres national park. Arquivos do Museu Bocage serie A 3:169–195

    Google Scholar 

  • Pastro LA, Banks PB (2006) Foraging responses of wild house mice to accumulations of conspecific odor as a predation risk. Behav Ecol Sociobiol 60:101–107

    Article  Google Scholar 

  • Penzhorn BL (1984) A long-term study of social organization and behaviour of Cape mountain zebra (Equus zebra zebra). Zeit Tierpsychol 64:97–146

    Article  Google Scholar 

  • Pluháček J, Bartoš L, Víchová J (2006) Variation in incidence of male infanticide within subspecies of plains zebra (Equus burchelli). J Mammal 87:35–40

    Article  Google Scholar 

  • Pluháček J, Bartoš L, Doležalová M, Bartošová-Víchová J (2007) Sex of the foetus determines the time of weaning of the previous offspring of captive plains zebra (Equus burchelli). Appl Anim Behav Sci 105:192–204

    Article  Google Scholar 

  • Porton I (1983) Bush dog urine-marking—its role in pair forming and maintenance. Anim Behav 31:1061–1069

    Article  Google Scholar 

  • Ralls K (1971) Mammalian scent marking. Science 171:443–449

    Article  CAS  PubMed  Google Scholar 

  • Roberts SC (2007) Scent marking. In: Wolff JO, Sherman PW (eds) Rodent societies: an ecological and evolutionary perspective. University of Chicago Press, Chicago, pp 255–266

    Google Scholar 

  • Rothman RJ, Mech LD (1979) Scent marking in lone wolves and newly formed pairs. Anim Behav 27:750–760

    Article  Google Scholar 

  • Schilder MBH (1992) Stability and dynamics of group composition in a herd of captive plains zebra. Ethology 90:145–154

    Google Scholar 

  • Siniscalchi M, d’Ingeo S, Minunno M, Quaranta A (2018) Communication in dogs. Animals 8:131

    Article  Google Scholar 

  • Trivers RL (1974) Parent-offspring conflict. Am Zool 14:249–264

    Google Scholar 

  • Tučková V, Šárová R, Bartošová J, King SRB, Pluháček J (2018) Overmarking by adult females in four equid species: social bonds and group cohesion. J Zool 306:180–188

    Article  Google Scholar 

  • Turner JW, Perkins A, Kirkpatrick JF (1981) Elimination marking behavior in feral horses. Can J Zool 59:1561–1566

    Article  Google Scholar 

  • Tyler SJ (1972) The behaviour and social organisation of the new forest ponies. Anim Behav Monogr 5:85–196

    Article  Google Scholar 

  • Viitala J, Korpimaki E, Palokangas P, Koivula M (1995) Attraction of kestrels to vole scent marks visible in ultraviolet-light. Nature 373:425–427

    Article  CAS  Google Scholar 

  • Wright S (1922) Coefficients of inbreeding and relationship. Am Nat 56:330–338

    Article  Google Scholar 

  • Wronski T, Apio A, Plath M, Ziege M (2013) Sex difference in the communicatory significance of localized defecation sites in Arabian gazelles (Gazella arabica). J Ethol 31:129–140

    Article  Google Scholar 

Download references

Acknowledgements

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. 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.

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Pluháček, J., Tučková, V., King, S.R.B. et al. Test of four hypotheses to explain the function of overmarking in foals of four equid species. Anim Cogn 22, 231–241 (2019). https://doi.org/10.1007/s10071-019-01239-4

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