Uller, C., & Lewis, J. (2009). Horses (Equus caballus) select the greater of two quantities in small numerical contrasts. Anim. Cogn., 12(5), 733–738.
Abstract: Abstract The ability to select the greater numerosity over another in small sets seems to stem from the calculation of which set contains more, and has been taken as evidence of a primordial representation at the roots of the primate numerical system. We tested 56 horses (Equus caballus) in a paradigm previously used with human infants and nonhuman primates. Horses saw two quantities paired in contrasts—2 versus 1, 3 versus 2, 6 versus 4 and a control for volume, 2 versus 1 big—and had to make a choice by snout touching the container holding the numerosity selected. The horses spontaneously selected the greater of the two quantities when the numerosities were small. These results add to evidence showing spontaneous quantity assessment in a variety of species.
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Krueger, K., Flauger, B., Farmer, K., & Maros, K. (2011). Horses (Equus caballus) use human local enhancement cues and adjust to human attention. Anim. Cogn., 14(2), 187–201.
Abstract: This study evaluates the horse (Equus caballus) use of human local enhancement cues and reaction to human attention when making feeding decisions. The superior performance of dogs in observing human states of attention suggests this ability evolved with domestication. However, some species show an improved ability to read human cues through socialization and training. We observed 60 horses approach a bucket with feed in a three-way object-choice task when confronted with (a) an unfamiliar or (b) a familiar person in 4 different situations: (1) squatting behind the bucket, facing the horse (2) standing behind the bucket, facing the horse (3) standing behind the bucket in a back-turned position, gazing away from the horse and (4) standing a few meters from the bucket in a distant, back-turned position, again gazing away from the horse. Additionally, postures 1 and 2 were tested both with the person looking permanently at the horse and with the person alternating their gaze between the horse and the bucket. When the person remained behind the correct bucket, it was chosen significantly above chance. However, when the test person was turned and distant from the buckets, the horses’ performance deteriorated. In the turned person situations, the horses approached a familiar person and walked towards their focus of attention significantly more often than with an unfamiliar person. Additionally, in the squatting and standing person situations, some horses approached the person before approaching the correct bucket. This happened more with a familiar person. We therefore conclude that horses can use humans as a local enhancement cue independently of their body posture or gaze consistency when the persons remain close to the food source and that horses seem to orientate on the attention of familiar more than of unfamiliar persons. We suggest that socialization and training improve the ability of horses to read human cues.
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Krueger, K., Farmer, K., & Heinze, J. (2014). The effects of age, rank and neophobia on social learning in horses. Anim. Cogn., 17(3), 645–655.
Abstract: Social learning is said to meet the demands of complex environments in which individuals compete over resources and co-operate to share resources. Horses (Equus caballus) were thought to lack social learning skills because they feed on homogenously distributed resources with few reasons for conflict. However, the horse’s social environment is complex, which raises the possibility that its capacity for social transfer of feeding behaviour has been underestimated. We conducted a social learning experiment using 30 socially kept horses of different ages. Five horses, one from each group, were chosen as demonstrators, and the remaining 25 horses were designated observers. Observers from each group were allowed to watch their group demonstrator opening a feeding apparatus. We found that young, low ranking, and more exploratory horses learned by observing older members of their own group, and the older the horse, the more slowly it appeared to learn. Social learning may be an adaptive specialisation to the social environment. Older animals may avoid the potential costs of acquiring complex and potentially disadvantageous feeding behaviours from younger group members. We argue that horses show social learning in the context of their social ecology, and that research procedures must take such contexts into account. Misconceptions about the horse’s sociality may have hampered earlier studies.
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Muller, C. E., & Uden, P. (2007). Preference of horses for grass conserved as hay, haylage or silage. Animal Feed Science and Technology, 132(1-2), 66–78.
Abstract: The purpose of this experiment was to determine if different methods of forage conservation influenced horse preference for conserved forages. Silage, haylage with two different dry matter (DM) levels and hay was produced from the same grass crop at the same botanical maturity stage. Four horses were simultaneously offered the four forages (1 kg DM of each forage) once daily for four consecutive experimental periods, each period consisting of 5 days. On each experimental day, the horses were observed for 2 h and their first choice, eating time and forage consumption was registered for every forage. The number of times each horse depleted individual forages and the number of times each horse tasted or smelled a forage, but left it in favour of another forage, was also recorded. Silage had the highest rate of consumption (0.90 kg DM/day, S.D. 0.14) and longest eating time (28.4 min/day, S.D. 5.16). Hay had the lowest rate of consumption (0.23 kg DM/day, S.D. 0.14) and shortest eating time (6.8 min/day, S.D. 4.08), while the haylages were intermediate. Silage was the first choice 72 of 84 times (85%). Hay was never completely consumed and silage was never left in favour of another forage after smelling or tasting it. We conclude that the forage conservation methods had an impact on horse preference in favour of silage, even if the reason for silage preference remains to be explained.
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Ishida, N., Hirano, T., & Mukoyama, H. (1994). Detection of aberrant alleles in the D-loop region of equine mitochondrial DNA by single-strand conformation polymorphism (SSCP) analysis. Anim Genet, 25(4), 287.
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Momozawa, Y., Takeuchi, Y., Tozaki, T., Kikusui, T., Hasegawa, T., Raudsepp, T., et al. (2007). SNP detection and radiation hybrid mapping in horses of nine candidate genes for temperament. Anim Genet, 38(1), 81–83.
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Breen, M., Downs, P., Irvin, Z., & Bell, K. (1994). Intrageneric amplification of horse microsatellite markers with emphasis on the Przewalski's horse (E. przewalskii). Anim Genet, 25(6), 401–405.
Abstract: Primer sequences flanking 13 microsatellite loci isolated from the domestic horse (E. caballus) were successfully used to amplify homologous loci in the Przewalski's horse (E. przewalskii). The results demonstrate that the level of polymorphism at all 13 loci in the Przewalski's horse was comparable to that in the domestic horse and the overall exclusion probability in the Przewalski's horse was calculated to be 0.9994. The results suggest that it should be possible to use E. caballus-derived microsatellite markers to provide parentage verification and additional valuable information to the captive management of E. przewalskii. The ability to amplify corresponding loci in the remaining five species of the genus was also confirmed, illustrating the general application of markers isolated from the domestic horse to the evaluation of polymorphism in the other six species of the genus.
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Wallner, B., Brem, G., Muller, M., & Achmann, R. (2003). Fixed nucleotide differences on the Y chromosome indicate clear divergence between Equus przewalskii and Equus caballus. Anim Genet, 34(6), 453–456.
Abstract: The phylogenetic relationship between Equus przewalskii and E. caballus is often a matter of debate. Although these taxa have different chromosome numbers, they do not form monophyletic clades in a phylogenetic tree based on mtDNA sequences. Here we report sequence variation from five newly identified Y chromosome regions of the horse. Two fixed nucleotide differences on the Y chromosome clearly display Przewalski's horse and domestic horse as sister taxa. At both positions the Przewalski's horse haplotype shows the ancestral state, in common with the members of the zebra/ass lineage. We discuss the factors that may have led to the differences in mtDNA and Y-chromosomal observations.
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Oakenfull, E. A., & Ryder, O. A. (1998). Mitochondrial control region and 12S rRNA variation in Przewalski's horse (Equus przewalskii). Anim Genet, 29(6), 456–459.
Abstract: Variation in the control region and the 12S rRNA gene of all surviving mitochondrial lineages of Przewalski's horse was investigated. Variation is low despite the present day population being descended from 13 individuals probably representing animals from three different regions of its range. Phylogenetic comparison of these sequences, with sequences for the domestic horse, does not resolve the ancestral status of either horse.
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Aberle, K. S., Hamann, H., Drögemüller, C., & Distl, O. (2004). Genetic diversity in German draught horse breeds compared with a group of primitive, riding and wild horses by means of microsatellite DNA markers. Anim. Gen., 35(4), 270–277.
Abstract: Summary We compared the genetic diversity and distance among six German draught horse breeds to wild (Przewalski's Horse), primitive (Icelandic Horse, Sorraia Horse, Exmoor Pony) or riding horse breeds (Hanoverian Warmblood, Arabian) by means of genotypic information from 30 microsatellite loci. The draught horse breeds included the South German Coldblood, Rhenish German Draught Horse, Mecklenburg Coldblood, Saxon Thuringa Coldblood, Black Forest Horse and Schleswig Draught Horse. Despite large differences in population sizes, the average observed heterozygosity (Ho) differed little among the heavy horse breeds (0.64�0.71), but was considerably lower than in the Hanoverian Warmblood or Icelandic Horse population. The mean number of alleles (NA) decreased more markedly with declining population sizes of German draught horse breeds (5.2�6.3) but did not reach the values of Hanoverian Warmblood (NA = 6.7). The coefficient of differentiation among the heavy horse breeds showed 11.6% of the diversity between the heavy horse breeds, as opposed to 21.2% between the other horse populations. The differentiation test revealed highly significant genetic differences among all draught horse breeds except the Mecklenburg and Saxon Thuringa Coldbloods. The Schleswig Draught Horse was the most distinct draught horse breed. In conclusion, the study demonstrated a clear distinction among the German draught horse breeds and even among breeds with a very short history of divergence like Rhenish German Draught Horse and its East German subpopulations Mecklenburg and Saxon Thuringa Coldblood.
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