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Why wait to mark? Possible reasons behind latency from olfactory exploration to overmarking in four African equid species

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Abstract

Whereas most studies on overmarking in mammals analysed the rate of overmarking, that those investigate time between exploration of an olfactory stimulus and the response to it remain less common, with inconsistent results. We examined the latency in time between elimination by the sender and sniffing by the receiver, and from sniffing and overmarking, in four captive African equid species to explore differences among species, and among age and sex classes. We investigated these latency time periods in light of three potential hypotheses explaining overmarking behaviour in equids: social bonds, group cohesion, and intrasexual competition. Analysing 1684 events of sniffing and 719 of overmarking among 130 individuals, we found that (i) the time from elimination to overmarking was shorter among female friends and in parent–offspring dyads, proving support to the social bond hypothesis; (ii) intraspecific differences in time periods do not reflect the social organisation of species, thus not supporting the group cohesion hypothesis; (iii) males were more attracted to elimination of conspecifics than females, and female’s eliminations were inspected longer, in line with the sexual competition hypothesis and/or reproductive behaviour. In addition, we found that the younger foals came to sniff eliminations faster than older ones, and in larger groups foals devoted longer time to sniffing the elimination before overmarking. We concluded that examination of the elimination could be driven by motivations other than the decision to overmark. Whereas overmarking serves to express bonds to a familiar individual, the latency of overmarking reflects more reproductive interests.

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References

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

    Article  CAS  PubMed  Google Scholar 

  • Black-Décima P, Santana M (2011) Olfactory communication and counter-marking in brown brocket deer Mazama gouazoubira. Acta Theriol 56:179–187

    Article  Google Scholar 

  • Blank D, Ruckstuhl K, Yang WK (2015) The economics of scent marking with urine and feces in goitered gazelle (Gazella subgutturosa). Mammal Res 60:51–60

    Article  Google Scholar 

  • Boyd L, Kasman L (1986) The marking behavior of male Przewalski’s horses. In: Duvall D, Muller-Schwarze D, Silverstein RM (eds) Chemical signals in vertebrates, ecology, evolution, and comparative biology, vol IV. Plenum Press, New York, pp 623–626

    Chapter  Google Scholar 

  • Brashares JS, Arcese P (1999) Scent marking in a territorial African antelope: II. The economics of marking with faeces. Anim Behav 57:11–17

    Article  CAS  PubMed  Google Scholar 

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

    Google Scholar 

  • Cinková I, Policht R (2015) Discrimination of familiarity and sex from chemical cues in the dung by wild southern white rhinoceros. Anim Cogn 18:385–392

    Article  PubMed  Google Scholar 

  • Dreier S, van Zweden JS, D’Ettorre P (2007) Long-term memory of individual identity in ant queens. Biol Lett 3:459–462

    Article  PubMed  PubMed Central  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 

  • Ferkin MH, Leonard ST, Bartos K, Schmick MK (2001) Meadow voles and prairie voles differ in the length of time they prefer the top-scent donor of an over-mark. Ethology 107:1099–1114

    Article  Google Scholar 

  • Ferkin MH, Lee DN, Leonard ST (2004) The reproductive state of female voles affects their scent marking behavior and the responses of male conspecifics to such marks. Ethology 110:257–272

    Article  Google Scholar 

  • Font E, Barbosa D, Sampedro C, Carazo P (2012) Social behavior, chemical communication, and adult neurogenesis: studies of scent mark function in Podarcis wall lizards. Gen Comp Endocrinol 177:9–17

    Article  CAS  PubMed  Google Scholar 

  • Ghosal R, Seshagiri PB, Sukumar R (2012) Dung as a potential medium for inter-sexual chemical signaling in Asian elephants (Elephas maximus). Behav Proc 91:15–21

    Article  Google Scholar 

  • Ginsberg JR (1989) The ecology of female behaviour and male reproductive success in the Grevy’s zebra, Equus grevyi. In: Jewell PA, Maloiy GMO (eds) Symposia of the zoological society of London. Clarendon Press, pp 89–110

    Google Scholar 

  • Heymann EW (1998) Sex differences in olfactory communication in a primate, the moustached tamarin, Saguinus mystax (Callitrichinae). Behav Ecol Sociobiol 43:37–45

    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 

  • Jezierski T, Jaworski Z, Sobczynska M, Kaminska B, Gorecka-Bruzda A, Walczak M (2015) Excreta-mediated olfactory communication in Konik stallions: a preliminary study. J Vet Behav Clin Appl Res 10:353–364

    Article  Google Scholar 

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

    Article  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 (2007) Scent-marking investment is determined by sex and breeding status in meerkats. Anim Behav 74:531–540

    Article  Google Scholar 

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

    Article  Google Scholar 

  • Joubert E (1972) The social organisation and associated behaviour in the Hartmann zebra Equus zebra hartmannae. Madoqua Ser I 6:17–56

    Google Scholar 

  • Kappeler PM (1998) To whom it may concern: the transmission and function of chemical signals in Lemur catta. Behav Ecol Sociobiol 42:411–421

    Article  Google Scholar 

  • Kimura R (2000) Relationship of the type of social organization to scent-marking and mutual-grromin behaviour in Grevy’s (Equus grevyi) and Grant’s zebra (Equus burchelli bohmi). J Equi 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 (1969) The social organization and population ecology of the plains zebra (Equus quagga). Zool Afr 4:249–263

    Article  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

  • Klingel H (1977) Observations on social organization and behaviour of African and Asiatic wild assess (Equus africanus and E. hemionus). Z Tierpsychol 44:323–331

    Article  CAS  PubMed  Google Scholar 

  • Krueger K, Flauger B (2011) Olfactory recognition of individual competitors by means of faeces in horse (Equus caballus). Anim Cogn 14:245–257

    Article  PubMed  Google Scholar 

  • Leclaire S, Strandh M, Mardon J, Westerdahl H, Bonadonna F (2017) Odour-based discrimination of similarity at the major histocompatibility complex in birds. Proc Roy Soc B. https://doi.org/10.1098/rspb.2016.2466

    Article  Google Scholar 

  • Linklater WL, Mayer K, Swaisgood RR (2013) Chemical signals of age, sex and identity in black rhinoceros. Anim Behav 85:671–677

    Article  Google Scholar 

  • Marinier SL, Alexander AJ, Waring GH (1988) Flehmen behaviour in the domestic horse: Discrimination of conspecific odours. Appl Anim Behav Sci 19:227–237

    Article  Google Scholar 

  • Marneweck C, Jurgens A, Shrader AM (2017) Dung odours signal sex, age, territorial and oestrous state in white rhinos. Proc Roy Soc B. https://doi.org/10.1098/rspb.2016.2376

    Article  Google Scholar 

  • Marneweck C, Jurgens A, Shrader AM (2018) The role of middens in white rhino olfactory communication. Anim Behav 140:7–18

    Article  Google Scholar 

  • Marneweck C, Jürgens A, Shrader AM (2019) The first report of urine overmarking of pro-oestrus female dung by a male white rhino. Afr Zool 54:175–179

    Article  Google Scholar 

  • Meyer JM, Goodwin TE, Schulte BA (2008) Intrasexual chemical communication and social responses of captive female African elephants, Loxodonta africana. Anim Behav 76:163–174

    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 

  • Penzhorn BL (1984) Observations on mortality of free-ranging Cape Mountain zebras Equus zebra zebra. S Afr J Wild Res 14:89–90

    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 

  • Pluháček J, Tučková V, King SRB (2019a) Overmarking behaviour of zebra males: no scent masking, but a group cohesion function across three species. Behav Ecol Sociobiol 73:137

    Article  Google Scholar 

  • Pluháček J, Tučková V, Šárová R, King SRB (2019b) Test of four hypotheses to explain the function of overmarking in foals of four equid species. Anim Cogn 22:231–241

    Article  PubMed  Google Scholar 

  • Pluháček J, Tučková V, Šárová R, King SRB (2020) Effect of social organisation on interspecific differences in overmarking behaviour of foals in African equids. Anim Cogn 23:131–140

    Article  PubMed  Google Scholar 

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

    Article  CAS  PubMed  Google Scholar 

  • Steiner CC, Ryder OA (2011) Molecular phylogeny and evolution of the Perissodactyla. Zool J Lin Soc 163:1289–1303

    Article  Google Scholar 

  • Steiner CC, Mitelberg A, Tursi R, Ryder OA (2012) Molecular phylogeny of extant equids and effects of ancestral polymorphism in resolving species-level phylogenies. Mol Phyl Evol 65:573–581

    Article  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 Monographs 5:85–196

    Google Scholar 

  • Wolff JO, Mech SG, Thomas SA (2002) Scent marking in female prairie voles: a test of alternative hypotheses. Ethol 108:483–494

    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. We highly appreciated the comments of an anonymous reviewer and Kazuhiro Goto that helped to improve the manuscript. This work was supported by the Ministry of Agriculture of the Czech Republic (MZE-RO0718) and by grant no. 2011/008 of the Student Grant Agency of the University of South Bohemia.

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Authors and Affiliations

  1. Department of Ethology, Institute of Animal Science, Přátelství 815, Uhříněves, 10400, Prague, Czech Republic

    Jan Pluháček & Radka Šárová

  2. Ostrava Zoo, Michálkovická 2081/197, 71000, Ostrava, Czech Republic

    Jan Pluháček

  3. Departement of Biology and Ecology, Faculty of Science, University of Ostrava, Chittussiho 10, 71000, Ostrava, Czech Republic

    Jan Pluháček

  4. Department of Zoology, Faculty of Science, University of South Bohemia, Branišovská 1760, 37005, České Budějovice, Czech Republic

    Vladimíra Tučková

  5. Natural Resource Ecology Laboratory, Warner College of Natural Resources, Colorado State University, Campus Delivery 1499, Fort Collins, USA

    Sarah R. B. King

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  1. Jan Pluháček
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Correspondence to Jan Pluháček.

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Pluháček, J., Tučková, V., Šárová, R. et al. Why wait to mark? Possible reasons behind latency from olfactory exploration to overmarking in four African equid species. Anim Cogn 25, 1443–1452 (2022). https://doi.org/10.1007/s10071-022-01625-5

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