Overmarking remains an unstudied topic in juvenile mammals. We have previously documented a very high rate of overmarking by foals in four captive African equid species: mountain zebra (Equus zebra), plains zebra (Equus quagga), Grévy's zebra (Equus grevyi), and African wild ass (Equus africanus). African equids vary interspecifically in their social organisation. Since differences in social organisation affect many mammalian behaviours, in this study we investigated interspecific differences in overmarking behaviour of foals, analysing only cases where elimination of any other individual was explored by a foal. We hypothesised that the pattern of overmarking by foals should reflect either differences in social organisation of the species or phylogenetic relations among them. We found that in all species very young foals explored mostly maternal eliminations, and this preference declined with increasing age of the foal and reflected the social organisation of the species; the highest overmarking rate was in species with high intragroup aggression (mountain zebra) and lowest in species with low intragroup aggression and which form crèches (African wild ass). Similarly, the rate of overmarking of the mother, as opposed to other herdmates, was associated with social organisation of the respective species. Thus, we found interspecific differences in overmarking by foals, which were associated with variability in social organisation. Since we also revealed differences between African wild ass and zebra behaviour in early stages of ontogeny, we cannot refute the effect of phylogeny on overmarking behaviour. Additionally, our results supported the identity sharing hypothesis as an explanation of overmarking.

Department of Biology and Ecology Faculty of Science University of Ostrava Chittussiho 10 710 00 Ostrava Czech Republic

Department of Ethology Institute of Animal Science Přátelství 815 104 00 Praha Uhříněves Czech Republic

Department of Zoology Faculty of Science University of South Bohemia Branišovská 1760 370 05 České Budějovice Czech Republic

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

Ostrava Zoo Michálkovická 2081 197 710 00 Ostrava Czech Republic

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

Bekoff M, Wells MC (1981) Behavioral budgeting by wild coyotes—the influence of food resources and social organisation. Anim Behav 29:794–801

Berger J (1988) Social systems, resources and phylogenetic inertia: an experimental test and its limitations. In: Slobodchikoff CN (ed) The ecology of social behaviour. Academic Press, London, pp 157–186

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

Brinck C, Erlinge S, Sandell M (1983) Anal sac secretion in mustelids—a comparison. J Chem Ecol 9:727–745 PubMed

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

Caine NG, Marra SL (1988) Vigilance and social organisation in two species of primates. Anim Behav 36:897–904

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

Ferkin MH (1999) Meadow voles (Microtus pennsylvanicus, Arvicolidae) over-mark and adjacent-mark the scent marks of same-sex conspecifics. Ethology 105:825–837

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

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

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

Ganslosser A, Dellert B (1997) Experimental alternations of food distribution in two species of captive equids (Equus burchelli and Equus hemionus). Ethol Ecol Evol 9:1–17

Ginsberg JR (1989) The ecology of female behaviour and male reproductive success in the Grevy’s zebra, Equus grevyi. In: Maloiy GMO (ed) Jewell PA. Symp Zool Soc Lond Clarendon Press, Oxford, pp 89–110

Isbell LA, Young TP (1993) Social and ecological influences on activity budgets of vervet monkeys, and their implications for group living. Behavl Ecol Sociobiol 32:377–385

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

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

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

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

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

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

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 Equin Sci 11:91–98

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

Klingel H (1972) Social behaviour of African Equidae. Zool Afr 7:175–185

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

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 PubMed

Koomen R, Herrmann E (2018) The effects of social context and food abundance on chimpanzee feeding competition. Am J Primatol 80:e22734

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

Krüger K, Gaillard C, Stranzinger G, Rieder S (2005) Phylogenetic analysis and species allocation of individual equids using microsatellite data. J Anim Breed Gen 122:78–86

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

Lott DF (1991) Intraspecific variation in the social systems of wild vertebrates. Cambridge University Press, Cambridge

Macdonald DW (1980) Patterns of scent marking with urine and faeces amongst carnivore communities. Symp Zool Soc Lond 45:107–139

Macdonald DW (1985) The carnivores: order Carnivora. In: Brown RE, Macdonald DW (eds) Social odours in mammals, vol 2. Clarendon Press, Oxford, pp 619–722

Maestripieri D (1994) Mother-infant relationships in three species of macaques (Macaca mulatta, M. nemestrina, M. arctoides) II. The social environment. Behaviour 131:97–113

McGuire B, Novak M (1984) A comparison of maternal behavior in the meadow vole (Microtus pennsylvanicus), prairie vole (Microtus ochrogaster) and pine vole (Microtus pinetorum). Anim Behav 32:1132–1141

McGuire B, Vermeylen F, Bemis WE (2011) Suckling behaviour in three species of voles. Behaviour 148:551–573

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

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

Moehlman PD, Kebede F, Yohannes H (2013) Equus africanus African wild ass. In: Kingdon J, Hoffmann M (eds) Mammals of Africa, vol V. carnivores, pangolines, equids and rhinoceros. Bloomsbury, London, pp 422–428

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

Oakenfull EA, Lim HN, Ryder OA (2000) A survey of equid mitochondrial DNA: implications for the evolution, genetic diversity and conservation of Equus. Conserv Gen 1:341–355

Olléová M, Pluháček J, King SRB (2012) Effect of social system on allosuckling and adoption in zebras. J Zool 288:127–134

Palombit RA (2015) Infanticide as sexual conflict: coevolution of male strategies and female counterstrategies. Cold Spring Harbor Perspect Biol 7:a017640

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

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

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

Pluháček J, Olléová M, Bartošová J, Bartoš L (2012) Effect of ecological adaptation on suckling behaviour in three zebra species. Behaviour 149:1395–1411

Pluháček J, Olléová M, Bartoš L, Bartošová J (2014) Time spent suckling is affected by different social organization in three zebra species. J Zool 292:10–17

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

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

Policht R, Karadžos A, Frynta D (2011) Comparative analysis of long-range calls in equid stallions (Equidae): are acoustic parameters related to social organization? Afr Zool 46:18–26

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

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

Rubenstein DI (1989) The life history and social organization in arid adapted ungulates. J Arid Environ 17:145–156

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

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

Sundaresan SR, Fischhoff IR, Dushoff J, Rubenstein DI (2007) Network metrics reveal differences in social organization between two fission-fusion species, Grevy’s zebra and onager. Oecologia 151:140–149 PubMed

Thierry B (1985) Social development in three species of macaques (Macaca mulatta, M. fascicularis, M. tonkeana): a preliminary report on the first ten weeks of life. Behav Proc 11:89–95

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

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

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

Vilstrup JT, Seguin-Orlando A, Stiller M, Ginolhac A, Raghavan M, Nielsen SCA, Weinstock J, Froese D, Vasiliev SK, Ovodov ND, Clary J, Helgen KM, Fleischer RC, Cooper A, Shapiro B, Orlando L (2013) Mitochondrial phylogenomics of modern and ancient equids. Plos One 8:e55950 PubMed PMC

Williams SD (2013) Equus grevyi Grévy’s zebra. In: Kingdon J, Hoffmann M (eds) Mammals of Africa, vol V. Carnivores, pangolines, equids and rhinoceros. Bloomsbury, London, pp 422–428

Woodward JRL, Schmick MK, Ferkin MH (1999) Response of prairie voles, Microtus ochrogaster (Rodentia, Arvicolidae) to scent over-marks of two same-sex conspecifics: a test of the scent masking hypothesis. Ethology 105:1009–1017

Woodward JRL, Bartos K, Ferkin MH (2000) Meadow voles (Microtus pennsylvanicus) and prairie voles (Microtus ochrogaster) differ in their responses to over-marks from opposite- and same-sex conspecifics. Ethology 106:979–992

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

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