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Animal cognition2025; 28(1); 96; doi: 10.1007/s10071-025-02016-2

A systematic review on the effect of individual characteristics and management practices on equine cognition.

Abstract: Equine cognition is relevant to the many roles that horses serve in society, such as leisure riding, competitions, or even animal-assisted therapy. Equine cognitive abilities have been explored in recent years. However, gaining an overview of horse cognition is challenging due to the broad range of abilities studied and the diverse methodologies employed. In addition, the subjects of existing equine cognition studies vary greatly in contextual factors such as their breed, age, sex, and management conditions - each of which may influence test performance in the following cognitive categories: Discrimination Learning; Learning Sets, Categorisation and Concept Formation; Spatial Cognition; Social Learning; and Memory. The aims of this review were (1) to establish whether contextual information was provided in research articles on horse cognition, (2) to tabulate information on the characteristics, housing, and management of subjects used in different categories of cognitive test, (3) to provide an overview of cognitive abilities demonstrated by horses, i.e., the results obtained in cognitive tests, with a specific emphasis on the contextual factors shaping them. The results of this review highlighted important points for future research. Better reporting of subject characteristics in scientific publications would enable investigation of the factors which shape horses' cognitive abilities, and the use of standardized methods and procedures across studies would facilitate future comparative work.
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Publication Date: 2025-11-26 PubMed ID: 41296132PubMed Central: PMC12657588DOI: 10.1007/s10071-025-02016-2Google Scholar: Lookup
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  • Journal Article
  • Systematic Review

Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

Overview

  • This systematic review investigates how individual horse characteristics and management practices affect their cognitive abilities.
  • The review aims to assess the reporting of contextual factors in existing horse cognition research and to summarize cognitive test results considering these factors.

Introduction and Importance of Equine Cognition

  • Horses play diverse roles in society including leisure riding, competitions, and animal-assisted therapy.
  • Understanding equine cognition—how horses think, learn, and remember—is important for improving their welfare and training methods.
  • Research into equine cognitive abilities has expanded recently but varies widely in methods and subjects used.

Challenges in Reviewing Equine Cognition Studies

  • The reviewed studies differ greatly in the cognitive abilities tested, such as discrimination learning, spatial cognition, social learning, and memory.
  • Subjects in these studies vary in breed, age, sex, housing, and management conditions, all of which can influence cognitive test outcomes.
  • The diversity and inconsistency in sample characteristics and study methods make it hard to draw overarching conclusions about horse cognition.

Aims of the Systematic Review

  • Determine whether research articles on horse cognition provide detailed contextual information about the subjects.
  • Collect and tabulate data on individual horse characteristics and management/housing conditions linked to different cognitive tests.
  • Summarize and provide an overview of horses’ demonstrated cognitive abilities along with emphasis on how contextual factors shape these abilities.

Key Findings and Recommendations

  • Many studies lack adequate reporting of subject characteristics like breed, age, sex, and management conditions.
  • Incomplete reporting limits understanding of how these contextual factors impact cognitive performance in horses.
  • There is considerable variation in methodologies and procedures used across studies, further complicating comparisons and meta-analyses.
  • To advance the field, future research should improve the transparency and consistency of reporting horse characteristics and management practices.
  • Standardizing cognitive testing methods will enable more reliable comparisons of findings from different studies.
  • Understanding how individual and contextual factors influence cognition can improve training, welfare practices, and applications such as therapy and sport.

Conclusion

  • This review highlights the need for better standardized research approaches in equine cognition.
  • Improved documentation of horse-specific factors and housing/management conditions is critical.
  • Such efforts will allow researchers to more accurately evaluate horses’ cognitive abilities and the variables affecting them.
  • The findings ultimately support enhanced horse care and more effective use in their various societal roles.

Cite This Article

APA
(2025). A systematic review on the effect of individual characteristics and management practices on equine cognition. Anim Cogn, 28(1), 96. https://doi.org/10.1007/s10071-025-02016-2

Publication

Animal cognition
ISSN: 1435-9456
NlmUniqueID: 9814573
Country: Germany
Language: English
Volume: 28
Issue: 1
Pages: 96
PII: 96

Researcher Affiliations

MeSH Terms

  • Animals
  • Horses / psychology
  • Horses / physiology
  • Cognition
  • Animal Husbandry / methods

Conflict of Interest Statement

Declarations. Competing interests: The authors declare no competing interests. Animal ethics and consent to participate declarations: not applicable. Clinical trial number: not applicable.

References

This article includes 215 references
  1. Ahrendt LP, Christensen JW, Ladewig J. The ability of horses to learn an instrumental task through social observation.. Appl Anim Behav Sci 139:105–113.
    doi: 10.1016/j.applanim.2012.03.004google scholar: lookup
  2. Alberghina D, Bray EE, Buchsbaum D, Byosiere SE, Espinosa J, Gnanadesikan GE, Guran CNA, Hare E, Horschler DJ, Huber L, Kuhlmeier VA, MacLean EL, Pelgrim MH, Perez B, Ravid-Schurr D, Rothkoff L, Sexton CL, Silver ZA, Stevens JR. Manydogs project: a big team science approach to investigating canine behavior and cognition.. Comp Cogn Behav Rev 18:59–77.
    doi: 10.3819/CCBR.2023.180004google scholar: lookup
  3. Albiach-Serrano A, Bräuer J, Cacchione T, Zickert N, Amici F. The effect of domestication and ontogeny in swine cognition ( and ).. Appl Anim Behav Sci 141:25–35.
    doi: 10.1016/j.applanim.2012.07.005google scholar: lookup
  4. Alexander GE, Ryan L, Bowers D, Foster TC, Bizon JL, Geldmacher DS, Glisky EL. Characterizing cognitive aging in humans with links to animal models.. Front Aging Neurosci 4:21.
    doi: 10.3389/fnagi.2012.00021pmc: PMC3439638pubmed: 22988439google scholar: lookup
  5. Alves J, de Sá Couto-Pereira N, de Lima RMS, Quillfeldt JA, Dalmaz C. Effects of early life adversities upon memory processes and cognition in rodent models.. Neuroscience 497:282–307.
    doi: 10.1016/j.neuroscience.2022.04.023pubmed: 35525496google scholar: lookup
  6. Arts JWM, van der Staay FJ, Ekkel ED. Working and reference memory of pigs in the spatial holeboard discrimination task.. Behav Brain Res 205:303–306.
    doi: 10.1016/j.bbr.2009.06.014pubmed: 19539660google scholar: lookup
  7. Asher L, Friel M, Griffin K, Collins LM. Mood and personality interact to determine cognitive biases in pigs.. Biol Lett 12:20160402.
    doi: 10.1098/rsbl.2016.0402pmc: PMC5134031pubmed: 27852940google scholar: lookup
  8. Balestrieri A, Corona S, Gazzola A, Romano A. Numerical discrimination in domestic horses: a spontaneous binary choice test.. Behav 161:677–694.
    doi: 10.1163/1568539X-bja10281google scholar: lookup
  9. Baragli P, Vitale V, Paoletti E, Sighieri C, Reddon AR. Detour behaviour in horses ().. J Ethol 29:227–234.
    doi: 10.1007/s10164-010-0246-9google scholar: lookup
  10. Baragli P, Demuru E, Scopa C, Palagi E. Are horses capable of mirror self-recognition? A pilot study.. PLoS ONE 12:e0176717.
    doi: 10.1371/journal.pone.0176717pmc: PMC5433687pubmed: 28510577google scholar: lookup
  11. Baragli P, Vitale V, Sighieri C, Lanata A, Palagi E, Reddon AR. Consistency and flexibility in solving spatial tasks: different horses show different cognitive styles.. Sci Rep 7:16557.
    doi: 10.1038/s41598-017-16729-zpmc: PMC5707407pubmed: 29185468google scholar: lookup
  12. Beery AK, Zucker I. Sex bias in neuroscience and biomedical research.. Neurosci Biobehav Rev 35(3):565–572.
    doi: 10.1016/j.neubiorev.2010.07.002pmc: PMC3008499pubmed: 20620164google scholar: lookup
  13. Beilharz RG, Nitter G. The missing e: the role of the environment in evolution and animal breeding.. J Anim Breed Genet 115:439–453.
    doi: 10.1111/j.1439-0388.1998.tb00366.xgoogle scholar: lookup
  14. Beilharz RG, Luxford BG, Wilkinson JL. Quantitative genetics and evolution: is our understanding of genetics sufficient to explain evolution?. J Anim Breed Genet 110:161–170.
    doi: 10.1111/j.1439-0388.1993.tb00728.xpubmed: 21395715google scholar: lookup
  15. Bensky MK, Gosling SD, Sinn DL. Chapter Five - The World from a Dog’s Point of View: A Review and Synthesis of Dog Cognition Research.. .
  16. Bernauer K, Kollross H, Schuetz A, Farmer K, Krueger K. How do horses () learn from observing human action?. Anim Cogn 23:1–9.
    doi: 10.1007/s10071-019-01310-0pubmed: 31531748google scholar: lookup
  17. Bodden C, von Kortzfleisch VT, Karwinkel F, Kaiser S, Sachser N, Richter SH. Heterogenising study samples across testing time improves reproducibility of behavioural data. Sci Rep 9:8247.
    doi: 10.1038/s41598-019-44705-2pmc: PMC6547843pubmed: 31160667google scholar: lookup
  18. Bolhuis JE, Schouten WGP, de Leeuw JA, Schrama JW, Wiegant VM. Individual coping characteristics, rearing conditions and behavioural flexibility in pigs. Behav Brain Res 152:351–360.
    doi: 10.1016/j.bbr.2003.10.024pubmed: 15196803google scholar: lookup
  19. Bolhuis JE, Oostindjer M, Hoeks C, De Haas E, Bartels A, Ooms M, Kemp B. Working and reference memory of pigs (Sus scrofa domesticus) in a holeboard Spatial discrimination task: the influence of environmental enrichment. Anim Cogn 16:845–850.
    doi: 10.1007/s10071-013-0646-7pubmed: 23740471google scholar: lookup
  20. Bornstein MH. Sensitive periods in development: structural characteristics and causal interpretations. Psychol Bull 105:179–197.
    doi: 10.1037/0033-2909.105.2.179pubmed: 2648441google scholar: lookup
  21. Bowling AT, Zimmermann W, Ryder O, Penado C, Peto S, Chemnick L, Yasinetskaya N, Zharkikh T. Genetic variation in przewalski’s horses, with special focus on the last wild caught mare, 231 Orlitza III. Cytogenet Genome Res 102:226–234.
    doi: 10.1159/000075754pubmed: 14970708google scholar: lookup
  22. Bräuer J, Call J, Tomasello M. Visual perspective taking in dogs () in the presence of barriers. Appl Anim Behav Sci 88:299–317.
    doi: 10.1016/j.applanim.2004.03.004google scholar: lookup
  23. Bray EE, Gruen ME, Gnanadesikan GE, Horschler DJ, Levy KM, Kennedy BS, Hare BA, MacLean EL. Cognitive characteristics of 8- to 10-week-old assistance dog puppies. Anim Behav 166:193–206.
    doi: 10.1016/j.anbehav.2020.05.019pmc: PMC7384752pubmed: 32719570google scholar: lookup
  24. Bray EE, Gruen ME, Gnanadesikan GE, Horschler DJ, Levy KM, Kennedy BS, Hare BA, MacLean EL. Dog cognitive development: a longitudinal study across the first 2 years of life. Anim Cogn 24:311–328.
    doi: 10.1007/s10071-020-01443-7pmc: PMC8035344pubmed: 33113034google scholar: lookup
  25. Briefer Freymond S, Ruet A, Grivaz M, Fuentes C, Zuberbühler K, Bachmann I, Briefer EF. Stereotypic horses (Equus caballus) are not cognitively impaired. Anim Cogn 22:17–33.
    doi: 10.1007/10071-018-1217-8pubmed: 30328528google scholar: lookup
  26. Briefer Freymond S, Beuret S, Ruet A, Zuberbühler K, Bachmann I, Briefer EF. Stereotypic behaviour in horses lowers stress but not spatial learning performance. Appl Anim Behav Sci 232:105099.
    doi: 10.1016/j.applanim.2020.105099google scholar: lookup
  27. Brubaker L, Udell MAR. Cognition and learning in horses (): what we know and why we should ask more. Behav Process 126:121–131.
    doi: 10.1016/j.beproc.2016.03.017google scholar: lookup
  28. Bugnyar T, Reber SA, Buckner C. Ravens attribute visual access to unseen competitors. Nat Commun 7:10506.
    doi: 10.1038/ncomms10506pmc: PMC4740864pubmed: 26835849google scholar: lookup
  29. Burla JB, Siegwart J, Nawroth C. Human demonstration does not facilitate the performance of horses () in a spatial problem-solving task. Animals 8:96.
    doi: 10.3390/ani8060096pmc: PMC6025305pubmed: 29899296google scholar: lookup
  30. Bushby EV, Friel M, Goold C, Gray H, Smith L, Collins LM. Factors influencing individual variation in farm animal cognition and how to account for these statistically. Front Vet Sci 5:193.
    doi: 10.3389/fvets.2018.00193pmc: PMC6107851pubmed: 30175105google scholar: lookup
  31. Byosiere SE, Mundry R, Range F, Virányi Z. Selective responding to human ostensive communication is an early developing capacity of domestic dogs.. Dev Sci 2023;26:e13361.
    doi: 10.1111/desc.13361pubmed: 36545915google scholar: lookup
  32. Chapagain D, Range F, Huber L, Virányi Z. Cognitive aging in dogs.. Gerontology 2017;64:165–171.
    doi: 10.1159/000481621pmc: PMC5841136pubmed: 29065419google scholar: lookup
  33. Chaya L, Cowan E, McGuire B. A note on the relationship between time spent in turnout and behaviour during turnout in horses ().. Appl Anim Behav Sci 2006;98:155–160.
    doi: 10.1016/j.applanim.2005.08.020google scholar: lookup
  34. Christensen JW, Zharkikh T, Ladewig J, Yasinetskaya N. Social behaviour in stallion groups ( and ) kept under natural and domestic conditions.. Appl Anim Behav Sci 2002;76:11–20.
    doi: 10.1016/S0168-1591(01)00208-8google scholar: lookup
  35. Christensen JW, Zharkikh T, Ladewig J. Do horses generalise between objects during habituation?. Appl Anim Behav Sci 2008;114:509–520.
    doi: 10.1016/j.applanim.2008.03.007google scholar: lookup
  36. Christensen JW, Ahrendt LP, Malmkvist J, Nicol C. Exploratory behaviour towards novel objects is associated with enhanced learning in young horses.. Sci Rep 2021;11:1428.
    doi: 10.1038/s41598-020-80833-wpmc: PMC7809405pubmed: 33446827google scholar: lookup
  37. Clarke JV, Nicol CJ, Jones R, McGreevy PD. Effects of observational learning on food selection in horses.. Appl Anim Behav Sci 1996;50:177–184.
    doi: 10.1016/0168-1591(96)01071-4google scholar: lookup
  38. Craig M, Rand J, Mesch R, Shyan-Norwalt M, Morton J, Flickinger E. Domestic dogs () and the radial arm maze: spatial memory and serial position effects.. J Comp Psychol 2012;126(3):233–242.
    doi: 10.1037/a0025929pubmed: 22905996google scholar: lookup
  39. Cruz-Becerra D, Burunat-Gutiérrez E, HernánDez-Barrios A, Pérez-Acosta AM. Auditive discrimination of equine gaits by parade horses.. Univ Psychol 2009;8:507–518.
  40. d’Ingeo S, Quaranta A, Siniscalchi M, Stomp M, Coste C, Bagnard C, Hausberger M, Cousillas H. Horses associate individual human voices with the valence of past interactions: a behavioural and electrophysiological study.. Sci Rep 2019;9:11568.
    doi: 10.1038/s41598-019-47960-5pmc: PMC6689011pubmed: 31399629google scholar: lookup
  41. Dobos P, Pongrácz P. Would you detour with me? Association between functional breed selection and social learning in dogs sheds light on elements of Dog–Human cooperation.. Animals 2023;13(12):2001.
    doi: 10.3390/ani13122001pmc: PMC10295277pubmed: 37370511google scholar: lookup
  42. Doré FY. Object permanence in adult cats (Felis catus).. J Comp Psychol 1986;100:340–347.
    doi: 10.1037/0735-7036.100.4.340google scholar: lookup
  43. Dougherty DM, Lewis P. Stimulus-generalization, discrimination-learning, and peak shift in horses.. J Exp Anal Behav 1991;56:97–104.
    doi: 10.1901/jeab.1991.56-97pmc: PMC1323085pubmed: 1940765google scholar: lookup
  44. Dudde A, Krause ET, Matthews LR, Schrader L. More than eggs – relationship between productivity and learning in laying hens.. Front Psychol 2018;9:2000.
    doi: 10.3389/fpsyg.2018.02000pmc: PMC6212530pubmed: 30416464google scholar: lookup
  45. Dumas C. Object permanence in cats (): an ecological approach to the study of invisible displacements.. J Comp Psychol 1992;106:404–410.
    doi: 10.1037/0735-7036.106.4.404pubmed: 1451424google scholar: lookup
  46. Dunbar RIM. The social brain hypothesis.. Evol Anthropol 1998;6:178–190.
    doi: 10.1002/(SICI)1520-6505(1998)6:5<178::AID-EVAN5>3.0.CO;2-8google scholar: lookup
  47. Duranton C, Range F, Virányi Z. Do pet dogs () follow ostensive and non-ostensive human gaze to distant space and to objects?. R Soc Open Sci 2017;4:170349.
    doi: 10.1098/rsos.170349pmc: PMC5541559pubmed: 28791164google scholar: lookup
  48. Endenburg N. Perceptions and attitudes towards horses in European societies.. Equine Vet J 1999;28(Suppl):38–41.
    doi: 10.1111/j.2042-3306.1999.tb05154.xgoogle scholar: lookup
  49. Erhard HW, Boissy A, Rae MT, Rhind SM. Effects of prenatal undernutrition on emotional reactivity and cognitive flexibility in adult sheep.. Behav Brain Res 2004;151:25–35.
    doi: 10.1016/j.bbr.2003.08.003pubmed: 15084418google scholar: lookup
  50. Erickson CA, Barnes CA. The neurobiology of memory changes in normal aging.. Exp Gerontol 2003;38:61–69.
    doi: 10.1016/s0531-5565(02)00160-2pubmed: 12543262google scholar: lookup
  51. Esch L, Wöhr C, Erhard M, Krüger K. Horses’ () laterality, stress hormones, and task related behavior in innovative problem-solving.. Animals 2019;9:265.
    doi: 10.3390/ani9050265pmc: PMC6562608pubmed: 31121937google scholar: lookup
  52. Evans L, Cameron-Whytock H, Ijichi C. Eye understand: physiological measures as novel predictors of adaptive learning in horses.. Appl Anim Behav Sci 2024;271:106152.
    doi: 10.1016/j.applanim.2023.106152google scholar: lookup
  53. Evans L, Cameron-Whytock H, Ijichi C. Whoa, no-go: evidence consistent with model-based strategy use in horses during an inhibitory task.. Appl Anim Behav Sci 2024;277:106339.
    doi: 10.1016/j.applanim.2024.106339google scholar: lookup
  54. Fiske JC, Potter GD. Discrimination reversal-learning in yearling horses.. J Anim Sci 1979;49:583–588.
    doi: 10.2527/jas1979.492583xgoogle scholar: lookup
  55. Flannery B. Relational discrimination learning in horses.. Appl Anim Behav Sci 1997;54:267–280.
    doi: 10.1016/S0168-1591(97)00006-3google scholar: lookup
  56. Freedman DG, King JA, Elliot O. Critical period in the social development of dogs.. Science 1961;133:1016–1017.
    doi: 10.1126/science.133.3457.1016pubmed: 13701603google scholar: lookup
  57. Fureix C, Jego P, Henry S, Lansade L, Hausberger M. Towards an ethological animal model of depression? A study on horses.. PLoS ONE 2012;7:e39280.
    doi: 10.1371/journal.pone.0039280pmc: PMC3386251pubmed: 22761752google scholar: lookup
  58. Gabor V, Gerken M. Horses use procedural learning rather than conceptual learning to solve matching to sample.. Appl Anim Behav Sci 2010;126:119–124.
    doi: 10.1016/j.applanim.2010.06.006google scholar: lookup
  59. Gabor V, Gerken M. Cognitive testing in horses using a computer based apparatus.. Appl Anim Behav Sci 2012;139:242–250.
    doi: 10.1016/j.applanim.2012.04.010google scholar: lookup
  60. Gabor V, Gerken M. Shetland ponies () show quantity discrimination in a matching-to-sample design.. Anim Cogn 2014;17:1233–1243.
    doi: 10.1007/s10071-014-0753-0pubmed: 24831887google scholar: lookup
  61. Gabor V, Gerken M. Study into long-term memory of a complex learning task in Shetland ponies ().. Appl Anim Behav Sci 2018;198:60–66.
    doi: 10.1016/j.applanim.2017.09.010google scholar: lookup
  62. Gagnon S, Doré FY. Cross-sectional study of object permanence in domestic puppies ().. J Comp Psychol 1994;108:220–232.
    doi: 10.1037/0735-7036.108.3.220pubmed: 7924252google scholar: lookup
  63. Gerencsér L, Pérez Fraga P, Lovas M, Újváry D, Andics A. Comparing interspecific socio-communicative skills of socialized juvenile dogs and miniature pigs.. Anim Cogn 2019;22:917–929.
    doi: 10.1007/s10071-019-01284-zpmc: PMC6834752pubmed: 31256339google scholar: lookup
  64. Gieling ET, Nordquist RE, van der Staay FJ. Assessing learning and memory in pigs.. Anim Cogn 2011;14:151–173.
    doi: 10.1007/s10071-010-0364-3pmc: PMC3040303pubmed: 21203792google scholar: lookup
  65. Goulet S, Doré FY, Rousseau R. Object permanence and working memory in cats ().. J Exp Psychol Anim Behav Process 1994;20:347–365.
    doi: 10.1037/0097-7403.20.4.347pubmed: 7964519google scholar: lookup
  66. Gouyet C, Ringhofer M, Yamamoto S. Horses cross-modally recognize women and men.. Sci Rep 2023;13:3864.
    doi: 10.1038/s41598-023-30830-6pmc: PMC9995451pubmed: 36890162google scholar: lookup
  67. Greening L, Downing J, Amiouny D, Lekang L, McBride S. The effect of altering routine husbandry factors on sleep duration and memory consolidation in the horse.. Appl Anim Behav Sci 2021;236:105229.
    doi: 10.1016/j.applanim.2021.105229google scholar: lookup
  68. Haag EL, Rudman R, Houpt KA. Avoidance, maze-learning and social-dominance in ponies.. J Anim Sci 1980;50:329–335.
    doi: 10.2527/jas1980.502329xgoogle scholar: lookup
  69. Haemmerli S, Thill C, Amici F, Cacchione T. Domestic horses () fail to intuitively reason about object properties like solidity and weight.. Anim Cogn 2018;21:441–446.
    doi: 10.1007/s10071-018-1177-zpubmed: 29525917google scholar: lookup
  70. Hall C, Goodwin D, Heleski C, Randle H, Waran N. Is there evidence of learned helplessness in horses?. J Appl Anim Welf Sci 2008;11:249–266.
    doi: 10.1080/10888700802101130pubmed: 18569222google scholar: lookup
  71. Hammell DL, Kratzer DD, Bramble WJ. Avoidance and maze learning in pigs.. J Anim Sci 1975;40:573–579.
    doi: 10.2527/jas1975.403573xpubmed: 1116978google scholar: lookup
  72. Hanggi EB. Categorization learning in horses ().. J Comp Psychol 1999;113:243–252.
    doi: 10.1037/0735-7036.113.3.243google scholar: lookup
  73. Hanggi EB. Discrimination learning based on relative size concepts in horses ().. Appl Anim Behav Sci 2003;83:201–213.
    doi: 10.1016/S0168-1591(03)00136-9google scholar: lookup
  74. Hanggi EB. Rotated object recognition in four domestic horses ().. J Equine Vet Sci 2010;30:175–186.
    doi: 10.1016/j.jevs.2010.02.003google scholar: lookup
  75. Hanggi EB. Short-term memory testing in domestic horses: experimental design plays a role.. J Equine Vet Sci 2010;30:617–623.
    doi: 10.1016/j.jevs.2010.10.004google scholar: lookup
  76. Hanggi EB, Ingersoll JF. Long-term memory for categories and concepts in horses ().. Anim Cogn 2009;12:451–462.
    doi: 10.1007/s10071-008-0205-9pubmed: 19148689google scholar: lookup
  77. Hausberger M, Stomp M, Sankey C, Brajon S, Lunel C, Henry S. Mutual interactions between cognition and welfare: the horse as an animal model.. Neurosci Biobehav Rev 2019;107:540–559.
    doi: 10.1016/j.neubiorev.2019.08.022pubmed: 31491471google scholar: lookup
  78. Helton WS, Helton ND. Physical size matters in the domestic dog’s () ability to use human pointing cues.. Behav Processes 2010;85:77–79.
    doi: 10.1016/j.beproc.2010.05.008pubmed: 20553825google scholar: lookup
  79. Henriksson J, Sauveroche M, Roth LSV. Effects of size and personality on social learning and human-directed behaviour in horses ().. Anim Cogn 2019;22:1001–1011.
    doi: 10.1007/s10071-019-01291-0pmc: PMC6834737pubmed: 31312981google scholar: lookup
  80. Hernandez CE, Harding JE, Oliver MH, Bloomfield FH, Held SDE, Matthews LR. Effects of litter size, sex and periconceptional ewe nutrition on side preference and cognitive flexibility in the offspring.. Behav Brain Res 2009;204:82–87.
    doi: 10.1016/j.bbr.2009.05.019pubmed: 19467268google scholar: lookup
  81. Hothersall B, Harris P, Sörtoft L, Nicol CJ. Discrimination between conspecific odour samples in the horse. Appl Anim Behav Sci 126:37–44.
    doi: 10.1016/j.applanim.2010.05.002google scholar: lookup
  82. Humphrey NK. The social function of intellect. .
  83. Janczarek I, Stachurska A, Pieszka M, Dracz K, Tkaczyk E, Kedzierski W, Łuszczynski J. Effect of fearfulness and cortisol reactivity to stress on the Spatial learning performance in mountain primitive horses. J Vet Behav 60:10–17.
    doi: 10.1016/j.jveb.2022.10.007google scholar: lookup
  84. Jardat P, Calandreau L, Ferreira V, Gouyet C, Parias C, Reigner F, Lansade L. Pet-directed speech improves horses’ attention toward humans. Sci Rep 12:4297.
    doi: 10.1038/s41598-022-08109-zpmc: PMC8917202pubmed: 35277552google scholar: lookup
  85. Jardat P, Liehrmann O, Reigner F, Parias C, Calandreau L, Lansade L. Horses discriminate between human facial and vocal expressions of sadness and joy. Anim Cogn 26:1733–1742.
    doi: 10.1007/s10071-023-01817-7pubmed: 37543956google scholar: lookup
  86. Jardat P, Ringhofer M, Yamamoto S, Gouyet C, Degrande R, Parias C, Reigner F, Calandreau L, Lansade L. Horses form cross-modal representations of adults and children. Anim Cogn 26:369–377.
    doi: 10.1007/s10071-022-01667-9pubmed: 35962844google scholar: lookup
  87. Jardat P, Menard-Peroy Z, Parias C, Reigner F, Calandreau L, Lansade L. Horses can learn to identify joy and sadness against other basic emotions from human facial expressions. Behav Process 220:105081.
    doi: 10.1016/j.beproc.2024.105081google scholar: lookup
  88. Kabadayi C, Bobrowicz K, Osvath M. The detour paradigm in animal cognition. Anim Cogn 21:21–35.
    doi: 10.1007/s10071-017-1152-0pmc: PMC5756264pubmed: 29234898google scholar: lookup
  89. Kaminski J, Schulz L, Tomasello M. How dogs know when communication is intended for them. Dev Sci 15:222–232.
    doi: 10.1111/j.1467-7687.2011.01120.xpubmed: 22356178google scholar: lookup
  90. Kappel S, Ramirez Montes De Oca MA, Collins S, Herborn K, Mendl M, Fureix C. Do you see what i see? Testing horses’ ability to recognise real–life objects from 2D computer projections. Anim Cogn 26:1147–1159.
    doi: 10.1007/s10071-023-01761-6pmc: PMC9980859pubmed: 36864246google scholar: lookup
  91. von Kortzfleisch VT, Ambrée O, Karp NA, Meyer N, Novak J, Palme R, Rosso M, Touma C, Würbel H, Kaiser S, Sachser N, Richter SH. Do multiple experimenters improve the reproducibility of animal studies?. PLoS Biol 20:e3001564.
    doi: 10.1371/journal.pbio.3001564pmc: PMC9070896pubmed: 35511779google scholar: lookup
  92. Koszałka A, Lustyk K, Pytka K. Sex-dependent differences in animal cognition. Neurosci Biobehav Rev 153:105374.
    doi: 10.1016/j.neubiorev.2023.105374pubmed: 37634555google scholar: lookup
  93. Krachun C, Lurz R, Mahovetz LM, Hopkins WD. Mirror self-recognition and its relationship to social cognition in chimpanzees. Anim Cogn 22:1171–1183.
    doi: 10.1007/s10071-019-01309-7pubmed: 31542841google scholar: lookup
  94. Krause A, Kreiser M, Puppe B, Tuchscherer A, Düpjan S. The effect of age on discrimination learning and self-control in a marshmallow test for pigs. Sci Rep 11:18287.
    doi: 10.1038/s41598-021-97770-xpmc: PMC8440626pubmed: 34521956google scholar: lookup
  95. Krüger K, Farmer K, Heinze J. The effects of age, rank and neophobia on social learning in horses. Anim Cogn 17:645–655.
    doi: 10.1007/s10071-013-0696-xpubmed: 24170136google scholar: lookup
  96. Kundey SMA, Strandell B, Mathis H, Rowan JD. Learning of monotonic and nonmonotonic sequences in domesticated horses () and chickens (). Learn Motiv 41:213–223.
    doi: 10.1016/j.lmot.2010.04.006google scholar: lookup
  97. Lampe JF, Andre J. Cross-modal recognition of human individuals in domestic horses (). Anim Cogn 15:623–630.
    doi: 10.1007/s10071-012-0490-1pubmed: 22526687google scholar: lookup
  98. Lampe M, Bräuer J, Kaminski J, Virányi Z. The effects of domestication and ontogeny on cognition in dogs and wolves. Sci Rep 7:11690.
    doi: 10.1038/s41598-017-12055-6pmc: PMC5601471pubmed: 28916808google scholar: lookup
  99. Lambert M, Farrar B, Garcia-Pelegrin E, Reber S, Miller R. ManyBirds: A multi-site collaborative Open Science approach to avian cognition and behavior research. Anim Behav Cogn 9:133–152.
    doi: 10.26451/abc.09.01.11.2022google scholar: lookup
  100. Lansade L, Colson V, Parias C, Trösch M, Reigner F, Calandreau L. Female horses spontaneously identify a photograph of their keeper, last seen six months previously. Sci Rep 10:6302.
    doi: 10.1038/s41598-020-62940-wpmc: PMC7156667pubmed: 32286345google scholar: lookup
  101. Lansade L, Colson V, Parias C, Reigner F, Bertin A, Calandreau L. Human face recognition in horses: data in favor of a holistic process. Front Psychol 11:575808.
    doi: 10.3389/fpsyg.2020.575808pmc: PMC7522352pubmed: 33041946google scholar: lookup
  102. Lemaire BS, Rugani R, Regolin L, Vallortigara G. Response of male and female domestic chicks to change in the number (quantity) of imprinting objects. Learn Behav 49:54–66.
    doi: 10.3758/s13420-020-00446-1google scholar: lookup
  103. Le Scolan N, Hausberger M, Wolff A. Stability over situations in temperamental traits of horses as revealed by experimental and scoring approaches. Behav Processes 41:257–266.
    doi: 10.1016/S0376-6357(97)00052-1pubmed: 24896858google scholar: lookup
  104. Lesimple C, Gautier E, Benhajali H, Rochais C, Lunel C, Bensaïd S, Khalloufi A, Henry S, Hausberger M. Stall architecture influences horses’ behaviour and the prevalence and type of stereotypies. Appl Anim Behav Sci 219:104833.
    doi: 10.1016/j.applanim.2019.104833google scholar: lookup
  105. Levine MA. Domestication and early history of the horse. .
  106. Lindberg AC, Kelland A, Nicol CJ. 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)00184-1google scholar: lookup
  107. Lindqvist C, Jensen P. Domestication and stress effects on contrafreeloading and spatial learning performance in red jungle fowl () and white leghorn layers. Behav Processes 81:80–84.
    doi: 10.1016/j.beproc.2009.02.005pubmed: 19429200google scholar: lookup
  108. Lovrovich P, Sighieri C, Baragli P. Following human-given cues or not? Horses () get smarter and change strategy in a delayed three choice task. Appl Anim Behav Sci 166:80–88.
    doi: 10.1016/j.applanim.2015.02.017google scholar: lookup
  109. Macpherson K, Roberts WA. Spatial memory in dogs () on a radial maze. J Comp Psychol 124(1):47–56.
    doi: 10.1037/a0018084pubmed: 20175596google scholar: lookup
  110. Mader DR, Price EO. Discrimination-learning in horses – Effects of breed, age and social-dominance. J Anim Sci 50:962–965.
    doi: 10.2527/jas1980.505962xpubmed: 7390949google scholar: lookup
  111. Mal ME, McCall CA, Newland C, Cummins KA. Evaluation of a one-trial learning apparatus to test learning ability in weanling horses. Appl Anim Behav Sci 35:305–311.
    doi: 10.1016/0168-1591(93)90082-Zgoogle scholar: lookup
  112. Maney DL. Perils and pitfalls of reporting sex differences. Philos Trans R Soc Lond B Biol Sci 371:20150119.
    doi: 10.1098/rstb.2015.0119pmc: PMC4785904pubmed: 26833839google scholar: lookup
  113. ManyBabies Consortium. Quantifying sources of variability in infancy research using the infant-directed-speech preference. Adv Methods Pract Psychol Sci 3:24–52.
    doi: 10.1177/2515245919900809google scholar: lookup
  114. Marinier SL, Alexander AJ, Waring GH. Flehmen behaviour in the domestic horse: discrimination of conspecific odours. Appl Anim Behav Sci 19:227–237.
    doi: 10.1016/0168-1591(88)90003-2google scholar: lookup
  115. Marshall-Pescini S, Virányi Z, Range F. The effect of domestication on inhibitory control: wolves and dogs compared. PLoS ONE 10:e0118469.
    doi: 10.1371/journal.pone.0118469pmc: PMC4340790pubmed: 25714840google scholar: lookup
  116. Marshall-Pescini S, Rao A, Virányi Z, Range F. The role of domestication and experience in ‘looking back’ towards humans in an unsolvable task. Sci Rep 7:46636.
    doi: 10.1038/srep46636pmc: PMC5395970pubmed: 28422169google scholar: lookup
  117. Mason MA, Briefer EF, Semple S, McElligott AG. Goat Emotions, Cognition, and personality. .
  118. McCall CA. The effect of body condition of horses on discrimination learning abilities. Appl Anim Behav Sci 22:327–334.
    doi: 10.1016/0168-1591(89)90027-0google scholar: lookup
  119. McDermott LM, Ebmeier KP. A meta-analysis of depression severity and cognitive function. J Affect Disord 119:1–8.
    doi: 10.1016/j.jad.2009.04.022pubmed: 19428120google scholar: lookup
  120. McLean AN. Short-term spatial memory in the domestic horse. Appl Anim Behav Sci 85:93–105.
    doi: 10.1016/j.applanim.2003.09.009google scholar: lookup
  121. McVey A, Wilkinson A, Mills DS. Social learning in horses: the effect of using a group leader demonstrator on the performance of familiar conspecifics in a detour task. Appl Anim Behav Sci 209:47–54.
    doi: 10.1016/j.applanim.2018.08.015google scholar: lookup
  122. Mejdell CM, Buvik T, Jørgensen GHM, Bøe KE. Horses can learn to use symbols to communicate their preferences. Appl Anim Behav Sci 184:66–73.
    doi: 10.1016/j.applanim.2016.07.014google scholar: lookup
  123. Miller HC, Gipson CD, Vaughan A, Rayburn-Reeves R, Zentall TR. Object permanence in dogs: invisible displacement in a rotation task. Psychon Bull Rev 16:150–155.
    doi: 10.3758/PBR.16.1.150pubmed: 19145026google scholar: lookup
  124. Miyashita Y, Nakajima S, Imada H. Differential outcome effect in the horse. J Exp Anal Behav 74:245–253.
    doi: 10.1901/jeab.2000.74-245pmc: PMC1284794pubmed: 11029025google scholar: lookup
  125. Nakamura K, Takimoto-Inose A, Hasegawa T. Cross-modal perception of human emotion in domestic horses (). Sci Rep 8:8660.
    doi: 10.1038/s41598-018-26892-6pmc: PMC6013457pubmed: 29930289google scholar: lookup
  126. Nawroth C, Ebersbach M, von Borell E. Juvenile domestic pigs () use human-given cues in an object choice task. Anim Cogn 17:701–713.
    doi: 10.1007/s10071-013-0702-3pubmed: 24197275google scholar: lookup
  127. Nawroth C, von Borell E, Langbein J. Object permanence in the Dwarf goat (): perseveration errors and the tracking of complex movements of hidden objects. Appl Anim Behav Sci 167:20–26.
    doi: 10.1016/j.applanim.2015.03.010google scholar: lookup
  128. Nicol CJ. Equine learning: progress and suggestions for future research. Appl Anim Behav Sci 78:193–208.
    doi: 10.1016/S0168-1591(02)00093-Xgoogle scholar: lookup
  129. Nicol CJ, Pope SJ. Social learning in sibling pigs. Appl Anim Behav Sci 40:31–43.
    doi: 10.1016/0168-1591(94)90085-Xgoogle scholar: lookup
  130. Oostindjer M, Bolhuis JE, Mendl M, Held S, van den Brand H, Kemp B. Learning how to eat like a pig: effectiveness of mechanisms for vertical social learning in piglets. Anim Behav 82:503–511.
    doi: 10.1016/j.anbehav.2011.05.031google scholar: lookup
  131. Orlando L. Ancient genomes reveal unexpected horse domestication and management dynamics. BioEssays 42:e1900164.
    doi: 10.1002/bies.201900164pubmed: 31808562google scholar: lookup
  132. Osthaus B, Lea SEG, Slater AM. Dogs () fail to show understanding of means-end connections in a string-pulling task. Anim Cogn 8:37–47.
    doi: 10.1007/s10071-004-0230-2pubmed: 15338446google scholar: lookup
  133. Osthaus B, Proops L, Hocking I, Burden F. Spatial cognition and perseveration by horses, donkeys and mules in a simple a-not-b detour task. Anim Cogn 16:301–305.
    doi: 10.1007/s10071-012-0589-4pubmed: 23271641google scholar: lookup
  134. Overmier JB, Seligman MEP. Effects of inescapable shock upon subsequent escape and avoidance learning. J Comp Physiol Psychol 63:28–33.
    doi: 10.1037/h0024166pubmed: 6029715google scholar: lookup
  135. Pasnak R, Kurkjian M, Triana E. Assessment of stage 6 object permanence. Bull Psychon Soc 26:368–370.
    doi: 10.3758/BF03337685google scholar: lookup
  136. Péron F, Ward R, Burman O. Horses () discriminate body odour cues from conspecifics. Anim Cogn 17:1007–1011.
    doi: 10.1007/s10071-013-0717-9pubmed: 24305997google scholar: lookup
  137. Pietrzak S, Próchniak T, Osińska K. The influence of certain factors on the results obtained by horses classified in eventing ranks of the International Federation for Equestrian Sports. J Anim Sci Biol Bioeconomy 31:34–39.
  138. Piotti P, Szabó D, Bognár Z, Egerer A, Hulsbosch P, Carson RS, Kubinyi E. Effect of age on discrimination learning, reversal learning, and cognitive bias in family dogs. Learn Behav 46:537–553.
    doi: 10.3758/s13420-018-0357-7pmc: PMC6328011pubmed: 30251103google scholar: lookup
  139. Piotti P, Piseddu A, Aguzzoli E, Sommese A, Kubinyi E. Two valid and reliable tests for monitoring age-related memory performance and neophobia differences in dogs. Sci Rep 12:16175.
    doi: 10.1038/s41598-022-19918-7pmc: PMC9519567pubmed: 36171232google scholar: lookup
  140. Pongrácz P, Miklósi A, Timár-Geng K, Csányi V. Verbal attention getting as a key factor in social learning between dog () and human. J Comp Psychol 118:375–383.
    doi: 10.1037/0735-7036.118.4.375pubmed: 15584774google scholar: lookup
  141. Pongrácz P, Miklósi Á, Vida V, Csányi V. The pet dogs ability for learning from a human demonstrator in a detour task is independent from the breed and age. Appl Anim Behav Sci 90:309–323.
    doi: 10.1016/j.applanim.2004.08.004google scholar: lookup
  142. Pongrácz P, Szapu JS, Faragó T. Cats () read human gaze for referential information. Intelligence 74:43–52.
    doi: 10.1016/j.intell.2018.11.001google scholar: lookup
  143. Price EO. Behavioral aspects of animal domestication. Q Rev Biol 59:1–32.
    doi: 10.1086/413673google scholar: lookup
  144. Proops L, McComb K. Attributing attention: the use of human-given cues by domestic horses (). Anim Cogn 13:197–205.
    doi: 10.1007/s10071-009-0257-5pubmed: 19588176google scholar: lookup
  145. Proops L, McComb K. Cross-modal individual recognition in domestic horses () extends to familiar humans. Proc R Soc Lond B Biol Sci 279:3131–3138.
    doi: 10.1098/rspb.2012.0626google scholar: lookup
  146. Proops L, McComb K, Reby D. Cross-modal individual recognition in domestic horses (). Proc Natl Acad Sci U S A 106:947–951.
    doi: 10.1073/pnas.0809127105pmc: PMC2630083pubmed: 19075246google scholar: lookup
  147. Proops L, Walton M, McComb K. The use of human-given cues by domestic horses, , during an object choice task. Anim Behav 79:1205–1209.
    doi: 10.1016/j.anbehav.2010.02.015google scholar: lookup
  148. Ragonese G, Baragli P, Mariti C, Gazzano A, Lanata A, Ferlazzo A, Fazio E, Cravana C. Interspecific two-dimensional visual discrimination of faces in horses (). PLoS ONE 16:e0247310.
    doi: 10.1371/journal.pone.0247310pmc: PMC7894942pubmed: 33606816google scholar: lookup
  149. Range F, Bugnyar T, Schlögl C, Kotrschal K. Individual and sex differences in learning abilities in Ravens. Behav Process 73:100–106.
    doi: 10.1016/j.beproc.2006.04.002google scholar: lookup
  150. Range F, Möslinger H, Virányi Z. Domestication has not affected the understanding of means-end connections in dogs. Anim Cogn 15:597–607.
    doi: 10.1007/s10071-012-0488-8pmc: PMC4306441pubmed: 22460629google scholar: lookup
  151. Range F, Marshall-Pescini S, Kratz C, Virányi Z. Wolves lead and dogs follow, but they both cooperate with humans. Sci Rep 9:3796.
    doi: 10.1038/s41598-019-40468-ypmc: PMC6405935pubmed: 30846770google scholar: lookup
  152. Regolin L, Rugani R, Pagni P, Vallortigara G. Delayed search for social and nonsocial goals by young domestic chicks, domesticus. Anim Behav 70:855–864.
    doi: 10.1016/j.anbehav.2005.01.014google scholar: lookup
  153. Ringhofer M, Yamamoto S. Domestic horses send signals to humans when they face with an unsolvable task. Anim Cogn 20:397–405.
    doi: 10.1007/s10071-016-1056-4pubmed: 27885519google scholar: lookup
  154. Rorvang MV, Ahrendt LP, Christensen JW. A trained demonstrator has a calming effect on Naive horses when crossing a novel surface. Appl Anim Behav Sci 171:117–120.
    doi: 10.1016/j.applanim.2015.08.008google scholar: lookup
  155. Rorvang MV, Ahrendt LP, Christensen JW. Horses fail to use social learning when solving Spatial detour tasks. Anim Cogn 18:847–854.
    doi: 10.1007/s10071-015-0852-6pubmed: 25716720google scholar: lookup
  156. Rorvang MV, Nielsen TB, Christensen JW. Horses failed to learn from humans by observation. Animals 10:221.
    doi: 10.3390/ani10020221pmc: PMC7070367pubmed: 32013218google scholar: lookup
  157. Rørvang MV, Ničová K, Sassner H, Nawroth C. Horses’ () ability to solve visible but not invisible displacement tasks is associated with frustration behavior and heart rate. Front Behav Neurosci 15:792035.
    doi: 10.3389/fnbeh.2021.792035pmc: PMC8693624pubmed: 34955782google scholar: lookup
  158. Rosenberger K, Simmler M, Nawroth C, Langbein J, Keil N. Goats work for food in a contrafreeloading task. Sci Rep 10:22336.
    doi: 10.1038/s41598-020-78931-wpmc: PMC7752918pubmed: 33349649google scholar: lookup
  159. Rugani R, Regolin L, Vallortigara G. Imprinted numbers: newborn chicks’ sensitivity to number vs. continuous extent of objects they have been reared with. Dev Sci 13:790–797.
    doi: 10.1111/j.1467-7687.2009.00936.xpubmed: 20712745google scholar: lookup
  160. Rugani R, Zhang Y, Scarsi B, Regolin L. Hybro chicks outperform Ross308 in a numerical-ordinal task. Cognitive and behavioral comparisons between two broiler strains of newborn domestic chicks (Gallus gallus). Poult Sci 103:103571.
    doi: 10.1016/j.psj.2023.103148google scholar: lookup
  161. Rutkauskaite A, Jensen P. Domestication effects on social information transfer in chickens. Anim Cogn 25:1473–1478.
    doi: 10.1007/s10071-022-01628-2pmc: PMC9652291pubmed: 35507205google scholar: lookup
  162. Sabiniewicz A, Tarnowska K, Swiatek R, Sorokowski P, Laska M. Olfactory-based interspecific recognition of human emotions: horses (Equus ferus caballus) can recognize fear and happiness body odour from humans (Homo sapiens). Appl Anim Behav Sci 230:105072.
    doi: 10.1016/j.applanim.2020.105072google scholar: lookup
  163. Salvin HE, McGreevy PD, Sachdev PS, Valenzuela MJ. The canine sand maze: An appetitive spatial memory paradigm sensitive to age-related change in dogs. J Exp Anal Behav 95:109–118.
    doi: 10.1901/jeab.2011.95-109google scholar: lookup
  164. Sankey C, Richard-Yris MA, Leroy H, Henry S, Hausberger M. Positive interactions lead to lasting positive memories in horses, Equus caballus. Anim Behav 79:869–875.
    doi: 10.1016/j.anbehav.2009.12.037google scholar: lookup
  165. Sappington BF, Goldman L. Discrimination-learning and concept-formation in the Arabian horse. J Anim Sci 72:3080–3087.
    doi: 10.2527/1994.72123080xpubmed: 7759356google scholar: lookup
  166. Sappington BKF, McCall CA, Coleman DA, Kuhlers DL, Lishak RS. A preliminary study of the relationship between discrimination reversal learning and performance tasks in yearling and 2-year-old horses. Appl Anim Behav Sci 53:157–166.
    doi: 10.1016/S0168-1591(96)01157-4google scholar: lookup
  167. van Schaik CP, Burkart JM. Social learning and evolution: the cultural intelligence hypothesis. Philos Trans R Soc Lond B Biol Sci 366:1008–1016.
    doi: 10.1098/rstb.2010.0304pmc: PMC3049085pubmed: 21357223google scholar: lookup
  168. Schloegl C, Kotrschal K, Bugnyar T. Gaze following in common ravens, : ontogeny and habituation. Anim Behav 74:769–778.
    doi: 10.1016/j.anbehav.2006.08.017google scholar: lookup
  169. Schubert CL, Ryckewaert B, Pereira C, Matsuzawa T. Garrano horses perceive letters of the alphabet on a touchscreen system: a pilot study. Animals 12:3514.
    doi: 10.3390/ani12243514pmc: PMC9774258pubmed: 36552434google scholar: lookup
  170. Schuetz A, Farmer K, Krueger K. Social learning across species: horses () learn from humans by observation. Anim Cogn 20:567–573.
    doi: 10.1007/s10071-016-1060-8pubmed: 27866286google scholar: lookup
  171. . Action anticipation based on an agent’s epistemic state in toddlers and adults. .
    doi: 10.31234/osf.io/x4jbmgoogle scholar: lookup
  172. Seligman ME, Maier SF. Failure to escape traumatic shock. J Exp Psychol 74:1–9.
    doi: 10.1037/h0024514pubmed: 6032570google scholar: lookup
  173. Smith AV, Proops L, Grounds K, Wathan J, Scott SK, McComb K. Domestic horses () discriminate between negative and positive human nonverbal vocalisations. Sci Rep 8:13052.
    doi: 10.1038/s41598-018-30777-zpmc: PMC6115467pubmed: 30158532google scholar: lookup
  174. Sobotková E, Mikule V, Kuřitková D, Jiskrová I, Sládek L. Analysis of the current situation in international show jumping and assessment of the influence of the proportion of Thoroughbred in the pedigree, horse demographics and sport season on the performance of horses. J Vet Behav 47:12–21.
    doi: 10.1016/j.jveb.2021.09.010google scholar: lookup
  175. Søndergaard E, Ladewig J. Group housing exerts a positive effect on the behaviour of young horses during training. Appl Anim Behav Sci 87:105–118.
    doi: 10.1016/j.applanim.2003.12.010google scholar: lookup
  176. Spence KL, O’Sullivan TL, Poljak Z, Greer AL. A longitudinal study describing horse demographics and movements during a competition season in Ontario, Canada. Can Vet J 59:783–790.
    doi: 10.1186/1746-6148-8-43pmc: PMC6005130pubmed: 30026628google scholar: lookup
  177. Spence KL, O’Sullivan TL, Poljak Z, Greer AL. A longitudinal study describing horse demographics and movements during a competition season in Ontario, Canada. The Canadian Veterinary Journal 59:783–790.
  178. Stone SM. Human facial discrimination in horses: can they tell us apart?. Anim Cogn 13:51–61.
    doi: 10.1007/s10071-009-0244-xpubmed: 19533185google scholar: lookup
  179. Szabó D, Gee NR, Miklósi Á. Natural or pathologic? Discrepancies in the study of behavioral and cognitive signs in aging family dogs. J Vet Behav 11:86–98.
    doi: 10.1016/j.jveb.2015.08.003google scholar: lookup
  180. Tapp PD, Siwak C, Estrada J, Head E, Muggenburg BA, Cotman CW, Milgram NW. Size and reversal learning in the beagle dog as a measure of executive function and inhibitory control in aging. Learn Mem 10:64–73.
    doi: 10.1101/lm.54403pmc: PMC196651pubmed: 12551965google scholar: lookup
  181. Tapp PD, Siwak CT, Estrada J, Holowachuk D, Milgram NW. Effects of age on measures of complex working memory span in the beagle dog () using two versions of a spatial list learning paradigm.. Learn Mem 10:148–160.
    doi: 10.1101/lm.56503pmc: PMC196663pubmed: 12663753google scholar: lookup
  182. Tapp PD, Siwak CT, Head E, Cotman CW, Murphey H, Muggenburg BA, Ikeda-Douglas C, Milgram NW. Concept abstraction in the aging dog: development of a protocol using successive discrimination and size concept tasks.. Behav Brain Res 153:199–210.
    doi: 10.1016/j.bbr.2003.12.003pubmed: 15219721google scholar: lookup
  183. Téglás E, Gergely A, Kupán K, Miklósi Á, Topál J. Dogs’ gaze following is tuned to human communicative signals.. Curr Biol 22:209–212.
    doi: 10.1016/j.cub.2011.12.018pubmed: 22226744google scholar: lookup
  184. Toinon C, Waiblinger S, Rault JL. Maternal deprivation affects goat kids’ stress coping behaviour.. Physiol Behav 239:113494.
    doi: 10.1016/j.physbeh.2021.113494pubmed: 34116050google scholar: lookup
  185. Tommasi L, Vallortigara G, Zanforlin M. Young chickens learn to localize the centre of a spatial environment.. Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology 180:567–572.
    doi: 10.1007/s003590050073pubmed: 9163931google scholar: lookup
  186. Tomonaga M, Kumazaki K, Camus F, Nicod S, Pereira C, Matsuzawa T. A horse’s eye view: size and shape discrimination compared with other mammals.. Biol Lett 11:20150701.
    doi: 10.1098/rsbl.2015.0701pmc: PMC4685539pubmed: 26601679google scholar: lookup
  187. Triana E, Pasnak R. Object permanence in cats and dogs.. Anim Learn Behav 9:135–139.
    doi: 10.3758/BF03212035google scholar: lookup
  188. Trösch M, Cuzol F, Parias C, Calandreau L, Nowak R, Lansade L. Horses categorize human emotions cross-modally based on facial expression and non-verbal vocalizations.. Animals 9:862.
    doi: 10.3390/ani9110862pmc: PMC6912773pubmed: 31653088google scholar: lookup
  189. Trösch M, Ringhofer M, Yamamoto S, Lemarchand J, Parias C, Lormant F, Lansade L. Horses prefer to solicit a person who previously observed a food-hiding process to access this food: a possible indication of attentional state attribution.. Behav Process 166:103906.
    doi: 10.1016/j.beproc.2019.103906google scholar: lookup
  190. Trösch M, Bertin E, Calandreau L, Nowak R, Lansade L. Unwilling or willing but unable: can horses interpret human actions as goal directed?. Anim Cogn 23:1035–1040.
    doi: 10.1007/s10071-020-01396-xpubmed: 32449047google scholar: lookup
  191. Trösch M, Flamand A, Chasles M, Nowak R, Calandreau L, Lansade L. Horses solve visible but not invisible displacement tasks in an object permanence paradigm.. Front Psychol 11:562989.
    doi: 10.3389/fpsyg.2020.562989pmc: PMC7552213pubmed: 33117229google scholar: lookup
  192. Trösch M, Pellon S, Cuzol F, Parias C, Nowak R, Calandreau L, Lansade L. Horses feel emotions when they watch positive and negative horse-human interactions in a video and transpose what they saw to real life.. Anim Cogn 23:643–653.
    doi: 10.1007/s10071-020-01369-0pubmed: 32162112google scholar: lookup
  193. Uller C, Lewis J. Horses () select the greater of two quantities in small numerical contrasts.. Anim Cogn 12:733–738.
    doi: 10.1007/s10071-009-0225-0pubmed: 19387706google scholar: lookup
  194. Valenchon M, Levy F, Fortin M, Leterrier C, Lansade L. Stress and temperament affect working memory performance for disappearing food in horses, .. Anim Behav 86:1233–1240.
    doi: 10.1016/j.anbehav.2013.09.026google scholar: lookup
  195. Valenchon M, Levy F, Gorecka-Bruzda A, Calandreau L, Lansade L. Characterization of long-term memory, resistance to extinction, and influence of temperament during two instrumental tasks in horses.. Anim Cogn 16:1001–1006.
    doi: 10.1007/s10071-013-0648-5pubmed: 23743707google scholar: lookup
  196. Vallortigara G. Core knowledge as a neuro-ethologist views it.. Behav Brain Sci 47:e144.
    doi: 10.1017/S0140525X23003035pubmed: 38934426google scholar: lookup
  197. Vallortigara G, Regolin L, Rigoni M, Zanforlin M. Delayed search for a concealed imprinted object in the domestic chick.. Anim Cogn 1:17–24.
    doi: 10.1007/s100710050003google scholar: lookup
  198. Veissier I. Observational learning in cattle.. Appl Anim Behav Sci 35:235–243.
    doi: 10.1016/0168-1591(93)90139-Ggoogle scholar: lookup
  199. Veit A, Wondrak M, Huber L. Object movement re-enactment in free-ranging Kune Kune piglets.. Anim Behav 132:49–59.
    doi: 10.1016/j.anbehav.2017.08.004google scholar: lookup
  200. Veit A, Weißhaupt S, Bruat A, Wondrak M, Huber L. Emulative learning of a two-step task in free-ranging domestic pigs.. Anim Cogn 26:929–942.
    doi: 10.1007/s10071-022-01740-3pmc: PMC10066142pubmed: 36652043google scholar: lookup
  201. Versace E, Martinho-Truswell A, Kacelnik A, Vallortigara G. Priors in animal and artificial intelligence: where does learning begin?. Trends Cogn Sci 22:963–965.
    doi: 10.1016/j.tics.2018.07.005pubmed: 30097305google scholar: lookup
  202. Vitale Shreve KR, Udell MAR. What’s inside your cat’s head? A review of cat () cognition research past, present and future.. Anim Cogn 18:1195–1206.
    doi: 10.1007/s10071-015-0897-6pubmed: 26154131google scholar: lookup
  203. Wallis LJ, Virányi Z, Müller CA, Serisier S, Huber L, Range F. Aging effects on discrimination learning, logical reasoning and memory in pet dogs.. Age 38:6.
    doi: 10.1007/s11357-015-9866-xpmc: PMC5005891pubmed: 26728398google scholar: lookup
  204. Wathan J, Proops L, Grounds K, McComb K. Horses discriminate between facial expressions of conspecifics.. Sci Rep 6:38322.
    doi: 10.1038/srep38322pmc: PMC5171796pubmed: 27995958google scholar: lookup
  205. Watowich MM, MacLean EL, Hare B, Call J, Kaminski J, Miklósi Á, Snyder-Mackler N. Age influences domestic dog cognitive performance independent of average breed lifespan.. Anim Cogn 23:795–805.
    doi: 10.1007/s10071-020-01385-0pmc: PMC7384235pubmed: 32356029google scholar: lookup
  206. Webster MM, Rutz C. How strange are your study animals?. Nature 582:337–340.
    doi: 10.1038/d41586-020-01751-5pubmed: 32541916google scholar: lookup
  207. Whishaw IQ, Burke CJ. Memory for surface objects in an arena by the horse () under saddle: evidence for dual process theory of spatial representation.. Behav Processes 189:104442.
    doi: 10.1016/j.beproc.2021.104442pubmed: 34116138google scholar: lookup
  208. Whiten A, Byrne RW. Tactical deception in primates.. Behav Brain Sci 11:233–244.
    doi: 10.1017/S0140525X00049682google scholar: lookup
  209. Whitt E, Douglas M, Osthaus B, Hocking I. Domestic cats () do not show causal understanding in a string-pulling task.. Anim Cogn 12:739–743.
    doi: 10.1007/s10071-009-0228-xpubmed: 19449193google scholar: lookup
  210. Wilkins AS, Wrangham RW, Fitch WT. The domestication syndrome in mammals: a unified explanation based on neural crest cell behavior and genetics.. Genetics 197:795–808.
    doi: 10.1534/genetics.114.165423pmc: PMC4096361pubmed: 25024034google scholar: lookup
  211. Wobber V, Hare B, Koler-Matznick J, Wrangham R, Tomasello M. Breed differences in domestic dogs’ () comprehension of human communicative signals.. Interaction Studies. Social Behaviour and Communication in Biological and Artificial Systems 10:206–224.
    doi: 10.1075/is.10.2.06wobgoogle scholar: lookup
  212. Wolff A, Hausberger M. Learning and memorisation of two different tasks in horses: the effects of age, sex and sire.. Appl Anim Behav Sci 46:137–143.
    doi: 10.1016/0168-1591(95)00659-1google scholar: lookup
  213. Würbel H, Garner J. Refinement of rodent research through environmental enrichment and systematic randomization.. NC3Rs 9:1–9.
  214. Wylie CE, Ireland JL, Collins SN, Verheyen KLP, Newton JR. Demographics and management practices of horses and ponies in Great Britain: a cross-sectional study.. Res Vet Sci 95:410–417.
    doi: 10.1016/j.rvsc.2013.05.004pubmed: 23735292google scholar: lookup
  215. Zobel G, Nawroth C. Current state of knowledge on the cognitive capacities of goats and its potential to inform species-specific enrichment.. Small Rumin Res 192:106208.
    doi: 10.1016/j.smallrumres.2020.106208google scholar: lookup

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