Animal Cognition

, Volume 17, Issue 3, pp 645–655 | Cite as

The effects of age, rank and neophobia on social learning in horses

  • Konstanze KruegerEmail author
  • Kate Farmer
  • Jürgen Heinze
Original Paper


Social learning is said to meet the demands of complex environments in which individuals compete over resources and cooperate 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.


Horse Social learning Sociality Ecology Social relationships 



We thank Michael Tomasello, Kurt Kotrschal, Christian Schloegel, Richard Byrne, Christine Nicol and Becky Hothersall for helpful discussions, Knut Krüger for helping with statistical analyses, Gudrun Schneider, Andreas Sailer, Anja Schlecht, Magdalena Schneider, and Stephanie Leopold for assistance at the experiments, as well as Sonja und Andreas Nieling, Karin Stadtherr, Tobias Knoll, Alexandra Stupperich, Elisabeth and Marion Klein, Nina Streek, Christina Bachfisch and Andrea Schöpe for providing us with horses. Finally, we would like to thank the Dr. Peter Deubner Stiftung for financial support.

Supplementary material

10071_2013_696_MOESM1_ESM.pdf (58 kb)
Supplementary material 1 (PDF 57 kb)
10071_2013_696_MOESM2_ESM.pdf (99 kb)
Supplementary material 2 (PDF 98 kb)

Video ESM 3: Demonstrator Training

The training phase of independent demonstration is reached. The demonstrator is first led to the experimental apparatus and then approaches the apparatus and opens the drawer independently by itself (AVI 151 Mb)

Video ESM 4: Successful learner

A learner horse pulls the rope, opens and feeds from the drawer after demonstration (AVI 143 Mb)

10071_2013_696_MOESM5_ESM.pdf (31 kb)
Supplementary material 5 (PDF 31 kb)
10071_2013_696_MOESM6_ESM.pdf (22 kb)
Supplementary material 6 (PDF 22 kb)


  1. Baer KL, Potter GD, Friend TH, Beaver BV (1983) Observation effects on learning in horses. Appl Anim Ethol 11:123–129. doi: 10.1016/0304-3762(83)90121-9 CrossRefGoogle Scholar
  2. Baker AEM, Crawford BH (1986) Observational learning in horses. Appl Anim Behav Sci 15:7–13. doi: 10.1016/0168-1591(86)90017-1 CrossRefGoogle Scholar
  3. Berger J (1986) Wild horses of the Great Basin. University of Chicago Press, ChicagoGoogle Scholar
  4. Boogert NJ, Reader SM, Laland KN (2006) The relation between social rank, neophobia and individual learning in starlings. Anim Behav 72:1229–1239. doi: 10.1016/j.anbehav.2006.02.021 CrossRefGoogle Scholar
  5. Byrne RW (2002) Imitation of novel complex actions: What does the evidence from animals mean? In: Snowdon CT, Roper TJ, Rosenblatt JS (eds) Advances in the study of behavior. Academic Press, San Diego, pp 77–105Google Scholar
  6. Byrne RW, Russon AE (1998) Learning by imitation: a hierarchical approach. Behav Brain Sci 21:667–721PubMedGoogle Scholar
  7. Byrne R, Whiten A (1988) Machiavellian intelligence. Oxford Univ Press, OxfordGoogle Scholar
  8. Call J (2006) Inferences by exclusion in the great apes: the effect of age and species. Anim Cogn 9:393–403. doi: 10.1007/s10071-006-0037-4 PubMedCrossRefGoogle Scholar
  9. Cameron EZ, Setsaas TH, Linklater WL (2009) Social bonds between unrelated females increase reproductive success in feral horses. Proc Natl Acad Sci USA 106:13850–13853. doi: 10.1126/science.1088580 PubMedCentralPubMedCrossRefGoogle Scholar
  10. Carroll J, Murphy CJ, Neitz M, Hoeve JN, Neitz J (2001) Photopigment basis for dichromatic color vision in the horse. J Vis 1:80–87. doi: 10.1167/1.2.2 PubMedCrossRefGoogle Scholar
  11. Chalmeau R, Gallo A (1993) Social constraints determine what is learned in the chimpanzee. Behav Process 28:173–179. doi: 10.1016/0376-6357(93)90090-E CrossRefGoogle Scholar
  12. Christensen JW, Keeling LJ, Nielsen BL (2005) Responses of horses to novel visual, olfactory and auditory stimuli. Appl Anim Behav Sci 93:53–65. doi: 10.1016/j.applanim.2005.06.017 CrossRefGoogle Scholar
  13. Christensen JW, Rundgren M, Olsson K (2006) Training methods for horses: habituation to a frightening stimulus. Equine Vet J 38:439–443PubMedCrossRefGoogle Scholar
  14. Christensen JW, Malmkvist J, Nielsen BL, Keeling LJ (2008) Effects of a calm companion on fear reactions in naive test horses. Equine Vet J 40:46–50. doi: 10.2746/042516408X245171 PubMedCrossRefGoogle Scholar
  15. Clarke JV, Nicol CJ, Jones R, McGreevy PD (1996) Effects of observational learning on food selection in horses. Appl Anim Behav Sci 50:177–184. doi: 10.1016/0168-1591(96)01071-4 CrossRefGoogle Scholar
  16. Coussi-Korbel S, Fragaszy DM (1995) On the relation between social dynamics and social learning. Anim Behav 50:1441–1453. doi: 10.1016/0003-3472(95)80001-8 CrossRefGoogle Scholar
  17. Cozzi A, Sighieri C, Gazzano A, Nicol CJ, Baragli P (2010) Post-conflict friendly reunion in a permanent group of horses (Equus caballus). Behav Process 85:185–190. doi: 10.1016/j.beproc.2010.07.007 CrossRefGoogle Scholar
  18. Devenport JA, Patterson MR, Devenport LD (2005) Dynamic averaging and foraging decisions in horses (Equus callabus). J Comp Psychol 119:352–358. doi: 10.1037/0735-7036.119.3.352 PubMedCrossRefGoogle Scholar
  19. Dunbar R (2003) Evolution of the social brain. Science 302:1160–1161. doi: 10.1126/science.1092116 PubMedCrossRefGoogle Scholar
  20. Dunbar RIM, Shultz S (2007) Evolution in the social brain. Science 317:1344–1347. doi: 10.1126/science.1145463 PubMedCrossRefGoogle Scholar
  21. Feist JD, McCullough DR (1976) Behavior patterns and communication in feral horses. Z Tierpsychol 41:337–371. doi: 10.1111/j.1439-0310.1976.tb00947.x PubMedCrossRefGoogle Scholar
  22. Galef BG (1995) Why behaviour patterns that animals learn socially are locally adaptive. Anim Behav 49:1325–1334. doi: 10.1006/anbe.1995.0164 CrossRefGoogle Scholar
  23. Galef BG Jr, Giraldeau LA (2001) Social influences on foraging in vertebrates: causal mechanisms and adaptive functions. Anim Behav 61:3–15. doi: 10.1006/anbe.2000.1557 PubMedCrossRefGoogle Scholar
  24. Geisbauer G, Griebel U, Schmid A, Timney B (2004) Brightness discrimination and neutral point. Can J Zool 82:660–670. doi: 10.1139/Z04-026 CrossRefGoogle Scholar
  25. Giraldeau L-A, Lefebvre L (1987) Scrounging prevents cultural transmission of food-finding behaviour in pigeons. Anim Behav 35:387–394. doi: 10.1016/S0003-3472(87)80262-2 CrossRefGoogle Scholar
  26. Giraldeau L-A, Valone TJ, Templeton JJ (2002) Potential disadvantages of using socially acquired information. Phil Trans Biol Sci 357:1559–1566. doi: 10.1098/rstb.2002.1065 CrossRefGoogle Scholar
  27. Greco BJ, Brown TK, Andrews JRM, Swaisgood RR, Caine NG (2013) Social learning in captive African elephants (Loxodonta africana africana). Anim Cogn 16:459–469. doi: 10.1007/s10071-012-0586-7 PubMedCrossRefGoogle Scholar
  28. Guttridge TL, Dijk S, Stamhuis EJ, Krause J, Gruber SH, Brown C (2013) Social learning in juvenile lemon sharks, Negaprion brevirostris. Anim Cogn 16:55–64. doi: 10.1007/s10071-012-0550-6 PubMedCrossRefGoogle Scholar
  29. Hall CA, Cassaday HJ, Derrington AM (2003) The effect of stimulus height on visual discrimination in horses. J Anim Sci 81:1715–1720PubMedGoogle Scholar
  30. Hanggi EB, Ingersoll JF (2009) Long-term memory for categories and concepts in horses (Equus caballus). Anim Cogn 13:451–462. doi: 10.1007/s10071-008-0205-9 CrossRefGoogle Scholar
  31. Hanggi EB, Ingersoll JF, Waggoner TL (2007) Color vision in horses (Equus caballus): deficiencies identified using a pseudoisochromatic plate test. J Comp Psychol 121:65–72. doi: 10.1037/0735-7036.121.1.65 PubMedCrossRefGoogle Scholar
  32. Hare B, Call J, Tomasello M (2001) Do chimpanzees know what conspecifics know? Anim Behav 61:139–151. doi: 10.1006/anbe 2000.1518PubMedCrossRefGoogle Scholar
  33. Hemelrijk CK, Wantia J, Gygax L (2005) The construction of dominance order: comparing performance of five methods using an individual-based model. Behaviour 142:1043–1064. doi: 10.1163/156853905774405290 CrossRefGoogle Scholar
  34. Heyes CM (1994) Social learning in animals: categories and mechanisms. Biol Rev 69:207–231. doi: 10.1111/j.1469-185X.1994.tb01506.x PubMedCrossRefGoogle Scholar
  35. Heyes CM (2002) Transformation and associative theories of imitation. In: Dautenhahn K, Nehaniv CL (eds) Imitation in animals and artefacts. MIT Press, Cambridge, pp 501–523Google Scholar
  36. Heyes C, Galef BG (1996) Social learning in animals: the roots of culture. Academic Press Inc., San DiegoGoogle Scholar
  37. Hopewell L, Leaver L, Lea S, Wills A (2010) Grey squirrels (Sciurus carolinensis) show a feature-negative effect specific to social learning. Anim Cogn 13:219–227. doi: 10.1007/s10071-009-0259-3 PubMedCrossRefGoogle Scholar
  38. Houpt KA, Law K, Martinisi V (1978) Dominance hierarchies in domestic horses. Appl Anim Ethol 4:273–283. doi: 10.1016/0304-3762(78)90117-7 CrossRefGoogle Scholar
  39. Imanishi K (1957) Identification: A process of enculturation in the subhuman society of Macaca fuscata. Primates 1, 1–29–29. doi:  10.1007/BF01667196
  40. Kerth G (2010) Group decision-making in animal societies. In: Kappeler P (ed) Animal behaviour: evolution and mechanisms. Springer, Berlin Heidelberg, pp 241–265CrossRefGoogle Scholar
  41. König von Borstel U, Euent S, Graf P, König S, Gauly M (2011) Equine behaviour and heart rate in temperament tests with or without rider or handler. Physiol Behav 104:454–463. doi: 10.1016/j.physbeh.2011.05.010 PubMedCrossRefGoogle Scholar
  42. Krueger K, Flauger B (2008) Social feeding decisions in horses (Equus caballus). Behav Process 78:76–83. doi: 10.1016/j.beproc.2008.01.009 CrossRefGoogle Scholar
  43. Krueger K, Flauger B (2011) Olfactory recognition of individual competitors by means of faeces in horse (Equus caballus). Anim Cogn 14:245–257. doi: 10.1007/s10071-010-0358-1 PubMedCrossRefGoogle Scholar
  44. Krueger K, Heinze J (2008) Horse sense: social status of horses (Equus caballus) affects their likelihood of copying other horses` behavior. Anim Cogn 11:431–439. doi: 10.1007/s10071-007-0133-0 PubMedCrossRefGoogle Scholar
  45. Krueger K, Flauger B, Farmer K, Maros K (2011) Horses (Equus caballus) use human local enhancement cues and adjust to human attention. Anim Cogn 14:187–201. doi: 10.1007/s10071-010-0352-7 PubMedCrossRefGoogle Scholar
  46. Laland KN (2004) Social learning strategies. Learn Behav 32:4–14PubMedCrossRefGoogle Scholar
  47. Le Scolan N, Hausberger M, Wolff A (1997) Stability over situations in temperamental traits of horses as revealed by experimental and scoring approaches. Behav Process 41:257–266. doi: 10.1016/S0376-6357(97)00052-1 CrossRefGoogle Scholar
  48. Lefebvre L, Giraldeau LA (1996) Is social learning an adaptive specialisation? In: Heyes CM, Galef BG Jr (eds) Social learning in animals: the root of culture. Academic Press, San Diego, pp 107–128CrossRefGoogle Scholar
  49. Lindberg AC, Kelland A, Nicol CJ (1999) Effects of observational learning on acquisition of an operant response in horses. Appl Anim Behav Sci 61:187–199. doi: 10.1016/S0168-1591(98)00184-1 CrossRefGoogle Scholar
  50. Linklater WL (2000) Adaptive explanation in socio-ecology: lessons from the Equidae. Biol Rev 75:1–20. doi: 10.1111/j.1469-185X.1999.tb00039.x PubMedCrossRefGoogle Scholar
  51. Macuda T, Timney B (1999) Luminance and chromatic discrimination in the horse (Equus caballus). Behav Process 44:301–307CrossRefGoogle Scholar
  52. McDonnell SM (2003) The equid ethogram: A practical field guide to horse behavior. Eclipse Press, Lexington, KentuckyGoogle Scholar
  53. McDonnell SM, Haviland JCS (1995) Agonistic ethogram of the equid bachelor band. Appl Anim Behav Sci 43:147–188. doi: 10.1016/0168-1591(94)00550-X CrossRefGoogle Scholar
  54. Mersmann D, Tomasello M, Call J, Kaminski J, Taborsky M (2011) Simple mechanisms can explain social learning in domestic dogs (Canis familiaris). Ethology 117:675–690. doi: 10.1111/j.1439-0310.2011.01919.x CrossRefGoogle Scholar
  55. Mettke-Hofmann C, Winkler H, Leisler B (2002) The significance of ecological factors for exploration and neophobia in parrots. Ethology 108:249–272. doi: 10.1046/j.1439-0310.2002.00773.x CrossRefGoogle Scholar
  56. Momozawa Y, Ono T, Sato F, Kikusui T, Takeuchi Y, Mori Y, Kusunose R (2003) Assessment of equine temperament by a questionnaire survey to caretakers and evaluation of its reliability by simultaneous behavior test. Appl Anim Behav Sci 84:127–138CrossRefGoogle Scholar
  57. Momozawa Y, Kusunose R, Kikusui T, Takeuchi Y, Mori Y (2005) Assessment of equine temperament questionnaire by comparing factor structure between two separate surveys. Appl Anim Behav Sci 92:77–84. doi: 10.1016/j.applanim.2004.11.006 CrossRefGoogle Scholar
  58. Murphy LB (1978) The practical problems of recognizing and measuring fear and exploration behaviour in the domestic fowl. Anim Behav 26(Part 2):422–431. doi: 10.1016/0003-3472(78)90059-3 CrossRefGoogle Scholar
  59. Murphy J, Arkins S (2007) Equine learning behaviour. Behav Process 76:1–13. doi: 10.1016/j.beproc.2006.06.009 CrossRefGoogle Scholar
  60. Nicol CJ (2002) Equine learning: progress and suggestions for future research. Appl Anim Behav Sci 78:193–208. doi: 10.1016/S0168-1591(02)00093-X CrossRefGoogle Scholar
  61. Nicol CJ, Pope SJ (1999) The effects of demonstrator social status and prior foraging success on social learning in laying hens. Anim Behav 57:163–171. doi: 10.1006/anbe.1998.0920 PubMedCrossRefGoogle Scholar
  62. Pongrácz P, Miklósi Á, Kubinyi E, Gurobi K, Topál J, Csányi V (2001) Social learning in dogs: the effect of a human demonstrator on the performance of dogs in a detour task. Anim Behav 62:1109–1117CrossRefGoogle Scholar
  63. Pongrácz P, Vida V, Bánhegyi P, Miklósi Á (2008) How does dominance rank status affect individual and social learning performance in the dog (Canis familiaris)? Anim Cogn 11:75–82. doi: 10.1007/s10071-007-0090-7 PubMedCrossRefGoogle Scholar
  64. Proops L, McComb K, Reby D (2009) Cross-modal individual recognition in domestic horses (Equus caballus). Proc Natl Acad Sci USA 106:947–951. doi: 10.1073/pnas.0809127105 PubMedCentralPubMedCrossRefGoogle Scholar
  65. Reader SM, Laland KN (2002) Social intelligence, innovation, and enhanced brain size in primates. Proc Natl Acad Sci USA 99:4436–4441. doi: 10.1073/pnas.062041299 PubMedCentralPubMedCrossRefGoogle Scholar
  66. Russon AE, Galdikas BMF (1995) Constraints on great apes’ imitation: model and action selectivity in rehabilitant orangutan (Pongo pygmaeus) imitation. J Comp Psychol 109:5–17. doi: 10.1037/0735-7036.109.1.5 PubMedCrossRefGoogle Scholar
  67. Schmidt M, Lipson H (2009) Distilling free-form natural laws from experimental data. Science 324:81–85. doi: 10.1126/science.1165893 PubMedCrossRefGoogle Scholar
  68. Schneider G, Krueger K (2012) Third-party interventions keep social partners from exchanging affiliative interactions with others. Anim Behav 83:377–387. doi: 10.1016/j.anbehav.2011.11.007 CrossRefGoogle Scholar
  69. Sherry DF, Galef BG Jr (1984) Cultural transmission without imitation: milk bottle opening by birds. Anim Behav 32:937–938. doi: 10.1016/S0003-3472(84)80185-2 CrossRefGoogle Scholar
  70. R Development Core Team (2012) R: A language and environment for statistical computing.
  71. Tomasello M (1990) Cultural transmission in the tool use and communicatory signalling of chimpanzees? In: Parker ST, Gibson KR (eds) Language and intelligence in monkeys and apes. Cambridge University Press, Cambridge, pp 274–311CrossRefGoogle Scholar
  72. Tyler SJ (1972) The behaviour and social organisation of the new forest ponies. Anim Behav Monogr 5:85–196CrossRefGoogle Scholar
  73. Valone TJ, Templeton JJ (2002) Public information for the assessment of quality: a widespread social phenomenon. Phil Trans Biol Sci 357:1549–1557CrossRefGoogle Scholar
  74. van Schaik CP (2010) Social learning and culture in animals. In: Kappeler P (ed) Animal behaviour: evolution and mechanisms. Springer, Berlin Heidelberg, pp 623–653CrossRefGoogle Scholar
  75. Vidament M, Yvon JM, Roche H, Le Bon H, Menard C, Delfosse A, Dubois C, Sabot J, Lansade L (2012) Temperament of stallions: relation with age, breed and level of riders. Proc 2 Int Equine Sci MtgGoogle Scholar
  76. Whiten A, Ham R (1992) On the nature and evolution of imitation in the animal kingdom: reappraisal of a century of research. Adv Study Behav 21:239–283CrossRefGoogle Scholar
  77. Whiten A, Horner V, Litchfield CA, Marshall-Pescini S (2004) How do apes ape? Learn Behav 32:36–52PubMedCrossRefGoogle Scholar
  78. Wilson SD, Clark AB, Coleman K, Dearstyne T (1994) Shyness and boldness in humans and other animals. Trends Ecol Evol 9:442–446. doi: 10.1016/0169-5347(94)90134-1 CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  • Konstanze Krueger
    • 1
    • 2
    Email author
  • Kate Farmer
    • 3
  • Jürgen Heinze
    • 1
  1. 1.Biology 1University of RegensburgRegensburgGermany
  2. 2.Nuertingen-Geislingen UniversityNürtingenGermany
  3. 3.Centre for Social Learning and Cognitive Evolution, School of PsychologyUniversity of St. AndrewsSt. AndrewsScotland, UK

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