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.

• Keywords
• Equus, Group cohesion, Intraspecific differences, Marking behaviour, Olfactory communication, Sexual competition, Social bonds, Zebra,
• MeSH
• Smell * physiology   MeSH
• Horses MeSH
• Mammals * MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Female MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH

In spite of a highly developed olfactory apparatus of horses, implying a high adaptive value, research on equine olfaction is sparse. Our limited knowledge on equine olfaction poses a risk that horse behavior does not match human expectations, as horses might react fearful when exposed to certain odors, which humans do not consider as frightening. The benefit of acquiring more knowledge of equine olfaction is therefore twofold; (1) it can aid the understanding of horse behavior and hence reduce the risk of dangerous situations, and (2) there may be unexplored potential of using odors in several practical situations where humans interact with horses. This study investigated behavior and olfactory sensitivity of 35 Icelandic horses who were presented with four odors: peppermint, orange, lavender and cedar wood in a Habituation/Dishabituation paradigm. The response variables were sniffing duration per presentation and behavioral reaction (licking, biting, snorting, and backing), and data were analyzed for potential effects of age, sex and pregnancy. Results showed that habituation occurred between successive odor presentations (1st vs. 2nd and 2nd vs. 3rd presentations: P < 0.001), and dishabituation occurred when a new odor was presented (1st vs. 3rd presentations: P < 0.001). Horses were thus able to detect and distinguish between all four odors, but expressed significantly longer sniffing duration when exposed to peppermint (peppermint vs. orange, lavender and cedar wood: P < 0.001). More horses expressed licking when presented to peppermint compared to cedar wood and lavender (P = 0.0068). Pregnant mares sniffed odors less than non-pregnant mares (P = 0.030), young horses (age 0-5 years) sniffed cedar wood for longer than old horses (P = 0.030), whereas sex had no effect (P > 0.050). The results show that horses' odor exploration behavior and interest in odors varies with age and pregnancy and that horses naïve to the taste of a substrate, may be able to link smell with taste, which has not been described before. These results can aid our understanding of horses' behavioral reactions to odors, and in the future, it may be possible to relate these to the physiology and health of horses.

• Keywords
• enrichment, horse-human relationship, nose-work, olfaction, scent, sensory ability, smell, trigeminal nerve,
• Publication type
• Journal Article MeSH

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.

• Keywords
• African wild ass, Equus, Interspecific differences, Marking behaviour, Olfactory communication, Social organisation, Zebra,
• MeSH
• Aggression MeSH
• Equidae * MeSH
• Phylogeny MeSH
• Horses MeSH
• Humans MeSH
• Mothers * MeSH
• Animals MeSH
• Check Tag
• Humans MeSH
• Female MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH