{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,25]],"date-time":"2026-06-25T18:47:02Z","timestamp":1782413222107,"version":"3.54.5"},"reference-count":64,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2025,2,22]],"date-time":"2025-02-22T00:00:00Z","timestamp":1740182400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>This paper challenges the prevalent characterization of domesticated horses as prey species that inherently view humans as predators. Drawing on evolutionary, ethological, and cognitive evidence, we propose the \u201cmutualistic coevolution hypothesis\u201d, which posits that horses and humans have evolved a partnership marked by cooperation rather than fear. We critically assess the \u201cprey hypothesis\u201d, emphasizing a predator\u2013prey model, which dominates equine training and the literature, and we argue that it inadequately explains horses\u2019 morphology, behaviors, and cognitive capacities. Comparative studies on horses\u2019 socio-cognitive skills suggest that domestication has fostered emotional, behavioral, and cognitive adaptations supporting a human\u2013horse bond. This review examines evidence from archaeological findings and experimental research on horses\u2019 responsiveness to human gestures, emotions, and social cues, underscoring their complex cognition and capacity for collaboration. Furthermore, morphological and behavioral analyses reveal inconsistencies in using orbital orientation or predation-related traits as evidence for categorizing horses as prey species. By emphasizing the coevolutionary dynamics underlying human\u2013horse interactions, we advocate for replacing traditional training models centered on fear and submission with approaches that leverage horses\u2019 mutualistic and social nature. This perspective offers insights for enhancing horse welfare and improving human\u2013equine relationships.<\/jats:p>","DOI":"10.3390\/ani15050641","type":"journal-article","created":{"date-parts":[[2025,2,24]],"date-time":"2025-02-24T10:52:47Z","timestamp":1740394367000},"page":"641","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Nay to Prey: Challenging the View of Horses as a \u201cPrey\u201d Species"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3033-6686","authenticated-orcid":false,"given":"Netzin G.","family":"Steklis","sequence":"first","affiliation":[{"name":"Human-Animal Interaction Research Initiative, School of Animal and Comparative Biomedical Sciences, University of Arizona, Tucson, AZ 85721, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mateo","family":"Pe\u00f1aherrera-Aguirre","sequence":"additional","affiliation":[{"name":"Human-Animal Interaction Research Initiative, School of Animal and Comparative Biomedical Sciences, University of Arizona, Tucson, AZ 85721, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Horst Dieter","family":"Steklis","sequence":"additional","affiliation":[{"name":"Human-Animal Interaction Research Initiative, School of Animal and Comparative Biomedical Sciences, University of Arizona, Tucson, AZ 85721, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,2,22]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Taylor, R.J. (1984). Predation, Springer Science & Business Media.","DOI":"10.1007\/978-94-009-5554-7"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1163\/156853008X291417","article-title":"Talking about horses: Control and freedom in the world of \u201cnatural horsemanship\u201d","volume":"16","author":"Birke","year":"2008","journal-title":"Soc. Anim."},{"key":"ref_3","first-page":"1232","article-title":"The revolution in horsemanship","volume":"216","author":"Miller","year":"2000","journal-title":"JAVMA"},{"key":"ref_4","unstructured":"Waran, N. (2002). Horse behaviour: Evolution, domestication and feralization. The Welfare of Horses, Kluwer."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Dalla Costa, E., Minero, M., Lebelt, D., Stucke, D., Canali, E., and Leach, M.C. (2014). Development of the Horse Grimace Scale (HGS) as a Pain Assessment Tool in Horses Undergoing Routine Castration. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0092281"},{"key":"ref_6","unstructured":"Parelli, P. (1993). Natural Horse-Manship, Publisher\u2019s Press."},{"key":"ref_7","unstructured":"Parelli, P. (2024, December 13). Horses and Humans Have Mutual Responsibilities. Available online: https:\/\/shopus.parelli.com\/blogs\/savvystation\/principle-number-4-horses-and-humans-have-mutual-responsibilities."},{"key":"ref_8","unstructured":"Johnson, E. (2024, December 15). Horse Behavior and Psychology (Part 2). Available online: https:\/\/equisearch.com\/articles\/horse-behavior-and-psychology-part-ii\/."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.tvjl.2009.03.005","article-title":"The horse\u2013human dyad: Can we align horse training and handling activities with the equid social ethogram?","volume":"181","author":"McGreevy","year":"2009","journal-title":"Vet. J."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Waran, N., McGreevy, P., and Casey, R.A. (2007). Training Methods and Horse Welfare. The Welfare of Horses, Springer.","DOI":"10.1007\/978-0-306-48215-1"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1861","DOI":"10.1016\/S0042-6989(98)00276-4","article-title":"Local and global stereopsis in the horse","volume":"39","author":"Timney","year":"1999","journal-title":"Vis. Res."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1159\/000108621","article-title":"Ecomorphology of orbit orientation and the adaptive significance of binocular vision in primates and other mammals","volume":"71","author":"Heesy","year":"2007","journal-title":"Brain Behav. Evol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"622","DOI":"10.1111\/joa.12953","article-title":"Taking a look into the orbit of mammalian carnivorans","volume":"234","author":"Pastor","year":"2019","journal-title":"J. Anat."},{"key":"ref_14","unstructured":"Heesy, C.P. (2003). The eVolution of Orbit Orientation in Mammals and the Function of the Primate Postorbital Bar. [Ph.D. Thesis, State University of New York]."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1006\/jhev.1995.1057","article-title":"Allometric and functional influences on primate orbit orientation and the origins of the Anthropoidea","volume":"29","author":"Ross","year":"1995","journal-title":"J. Hum. Evol."},{"key":"ref_16","unstructured":"Orme, D., Freckleton, R., Thomas, G., Petzoldt, T., Fritz, S., Isaac, N., and Pearse, W. (2013). The Caper Package: Comparative Analysis of Phylogenetics and Evolution in R, R Foundation for Stastical Computing. R Package Version."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1007\/BF00346285","article-title":"Distribution of retinal ganglion cells in five mammalian species (pig, sheep, ox, horse, dog)","volume":"150","author":"Hebel","year":"1976","journal-title":"Anat. Embryol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"e1500391","DOI":"10.1126\/sciadv.1500391","article-title":"Why do animal eyes have pupils of different shapes?","volume":"1","author":"Banks","year":"2015","journal-title":"Sci. Adv."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"0112","DOI":"10.1038\/s41559-017-0112","article-title":"Primate brain size is predicted by diet but not sociality","volume":"1","author":"DeCasien","year":"2017","journal-title":"Nat. Ecol. Evol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"10113","DOI":"10.1073\/pnas.0401955101","article-title":"Binocularity and brain evolution in primates","volume":"101","author":"Barton","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Denion, E., Hitier, M., Guyader, V., Dugu\u00e9, A.E., and Mouriaux, F. (2015). Unique human orbital morphology compared with that of apes. Sci. Rep., 5.","DOI":"10.1038\/srep11528"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1146\/annurev-vision-093019-113212","article-title":"Binocular vision and stereopsis across the animal kingdom","volume":"7","author":"Read","year":"2021","journal-title":"Annu. Rev. Vis. Sci."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Janczarek, I., Stachurska, A., K\u0119dzierski, W., Wi\u015bniewska, A., Ry\u017cak, M., and Kozio\u0142, A. (2020). The intensity of physiological and behavioral responses of horses to predator vocalizations. BMC Vet. Res., 16.","DOI":"10.1186\/s12917-020-02643-6"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/j.applanim.2007.08.003","article-title":"Predator odour per se does not frighten domestic horses","volume":"112","author":"Christensen","year":"2008","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1332","DOI":"10.1126\/science.1168594","article-title":"The earliest horse harnessing and milking","volume":"323","author":"Outram","year":"2009","journal-title":"Science"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1038\/s41586-021-03798-4","article-title":"Dairying enabled early bronze age Yamnaya steppe expansions","volume":"598","author":"Wilkin","year":"2021","journal-title":"Nature"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Anthony, D.W. (2009). The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World, Princeton University Press.","DOI":"10.1515\/9781400831104"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.applanim.2007.04.015","article-title":"A review of the human-horse relationship","volume":"109","author":"Hausberger","year":"2008","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1007\/s10071-009-0223-2","article-title":"How horses (Equus caballus) see the world: Humans as significant \u201cobjects\u201d","volume":"12","author":"Fureix","year":"2009","journal-title":"Anim. Cogn."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1016\/j.anbehav.2009.12.037","article-title":"Positive interactions lead to lasting positive memories in horses, Equus caballus","volume":"79","author":"Sankey","year":"2010","journal-title":"Anim. Behav."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1007\/s10071-009-0257-5","article-title":"Attributing attention: The use of human-given cues by domestic horses (Equus caballus)","volume":"13","author":"Proops","year":"2010","journal-title":"Anim. Cogn."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Proops, L., Rayner, J., Taylor, A.M., and McComb, K. (2013). The responses of young domestic horses to human-given cues. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0067000"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1007\/s10071-008-0136-5","article-title":"Comprehension of human pointing gestures in horses (Equus caballus)","volume":"11","author":"Maros","year":"2008","journal-title":"Anim. Cogn."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"103906","DOI":"10.1016\/j.beproc.2019.103906","article-title":"Horses prefer to solicit a person who previously observed a food-hiding process to access this food: A possible indication of attentional state attribution","volume":"166","author":"Ringhofer","year":"2019","journal-title":"Behav. Process."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"947","DOI":"10.1073\/pnas.0809127105","article-title":"Cross-modal individual recognition in domestic horses (Equus caballus)","volume":"106","author":"Proops","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Lansade, L., Colson, V., Parias, C., Reigner, F., Bertin, A., and Calandreau, L. (2020). Human face recognition in horses: Data in favor of a holistic process. Front. Psychol., 11.","DOI":"10.3389\/fpsyg.2020.575808"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Smith, A.V., Proops, L., Grounds, K., Wathan, J., and McComb, K. (2016). Functionally relevant responses to human facial expressions of emotion in the domestic horse (Equus caballus). Biol. Lett., 12.","DOI":"10.1098\/rsbl.2015.0907"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1283","DOI":"10.1007\/s10071-023-01775-0","article-title":"What drives horse success at following human-given cues? An investigation of handler familiarity and living conditions","volume":"26","author":"Liehrmann","year":"2023","journal-title":"Anim. Cogn."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.applanim.2017.05.020","article-title":"Longitudinal study on human-related behaviour in horses\u2014Can horses (Equus caballus) be de-domesticated?","volume":"195","author":"Jaworski","year":"2017","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1093\/emph\/eov015","article-title":"The evolution of capture myopathy in hooved mammals: A model for human stress cardiomyopathy?","volume":"2015","author":"Blumstein","year":"2015","journal-title":"Evol. Med. Public Health"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Steklis, N.G., Pe\u00f1aherrera-Aguirre, M., Steklis, H.D., and Herrera, I. (2024). Why Were Zebras Not Domesticated? A Review of Domesticability Traits and Tests of Their Role in Ungulate Domestications with Macroevolutionary Models. Animals, 14.","DOI":"10.3390\/ani14162355"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1091","DOI":"10.1002\/jwmg.21995","article-title":"Ecological interactions involving feral horses and predators: Review with implications for biodiversity conservation","volume":"85","author":"Boyce","year":"2021","journal-title":"J. Wildl. Manag."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1037\/a0039677","article-title":"Evolutionary constraints on equid domestication: Comparison of flight initiation distances of wild horses (Equus caballus ferus) and plains zebras (Equus quagga)","volume":"129","author":"Brubaker","year":"2015","journal-title":"J. Comp. Psychol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1415","DOI":"10.1111\/evo.12356","article-title":"Ecological drivers of antipredator defenses in carnivores","volume":"68","author":"Stankowich","year":"2014","journal-title":"Evolution"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1093\/beheco\/arab048","article-title":"A meta-analysis of the group-size effect on vigilance in mammals","volume":"32","author":"Beauchamp","year":"2021","journal-title":"Behav. Ecol."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Butler, J.R., Linnell, J.D., Morrant, D., Athreya, V., Lescureux, N., and McKeown, A. (2014). Dog eat dog, cat eat dog: Social-ecological dimensions of dog predation by wild carnivores. Free-Ranging Dogs and Wildlife Conservation, Oxford University Press.","DOI":"10.1093\/acprof:osobl\/9780199663217.003.0005"},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Dagg, A.I. (2011). Animal Friendships, Cambridge University Press.","DOI":"10.1017\/CBO9780511794155"},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Menchetti, L., Calipari, S., Mariti, C., Gazzano, A., and Diverio, S. (2020). Cats and dogs: Best friends or deadly enemies? What the owners of cats and dogs living in the same household think about their relationship with people and other pets. PLoS ONE, 15.","DOI":"10.1371\/journal.pone.0237822"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1046\/j.1439-0442.2002.jv416.x","article-title":"Dog bite wounds in dogs and cats: A retrospective study of 196 cases","volume":"49","author":"Shamir","year":"2002","journal-title":"J. Vet. Med. Ser. A"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/j.applanim.2005.02.005","article-title":"Heart rate and behavioural responses of dogs in the Ainsworth\u2019s Strange Situation: A pilot study","volume":"94","author":"Palestrini","year":"2005","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Spotte, S. (2012). Societies of Wolves and Free-Ranging Dogs, Cambridge University Press.","DOI":"10.1017\/CBO9781139057769"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"20211080","DOI":"10.1098\/rspb.2021.1080","article-title":"Killing conspecific adults in mammals","volume":"288","year":"2021","journal-title":"Proc. R. Soc. B"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1007\/s10071-021-01557-6","article-title":"Cognition and the human\u2013animal relationship: A review of the sociocognitive skills of domestic mammals toward humans","volume":"25","author":"Jardat","year":"2022","journal-title":"Anim. Cogn."},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Ferreira, V.H.B., Lansade, L., Calandreau, L., Cunha, F., and Jensen, P. (2023). Are domesticated animals dumber than their wild relatives? A comprehensive review on the domestication effects on animal cognitive performance. Neurosci. Biobehav. Rev., 154.","DOI":"10.1016\/j.neubiorev.2023.105407"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1007\/s40806-023-00360-5","article-title":"The relation of clade-specific biophilia to the construct of animality","volume":"9","author":"Figueredo","year":"2023","journal-title":"Evol. Psychol. Sci."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"112612","DOI":"10.1016\/j.paid.2024.112612","article-title":"Dimensions of Animality: Expanding nomological breadth and controlling phenotypic similarity","volume":"223","author":"Steklis","year":"2024","journal-title":"Personal. Individ. Differ."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1007\/s10071-005-0008-1","article-title":"A comparative analysis of animals\u2019 understanding of the human pointing gesture","volume":"9","author":"Soproni","year":"2006","journal-title":"Anim. Cogn."},{"key":"ref_58","unstructured":"Heleski, C., McLean, A., Swanson, J., and Grandin, T. (2009). Practical methods for improving the welfare of horses, donkeys and other working draught animals in developing areas. Improving Animal Welfare: A Practical Approach, CABI."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Bonsi, M., Anderson, N.E., and Carder, G. (2023). The Socioeconomic Impact of Diseases of Working Equids in Low and Middle-Income Countries: A Critical Review. Animals, 13.","DOI":"10.3390\/ani13243865"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1080\/10888705.2023.2228029","article-title":"Belief in Animal Sentience and Affective Owner Attitudes are linked to Positive Working Equid Welfare across Six Countries","volume":"28","author":"Haddy","year":"2023","journal-title":"J. Appl. Anim. Welf. Sci."},{"key":"ref_61","unstructured":"Alberta Equine Welfare Group (2008). Alberta Horse Welfare Report, Alberta Equine Welfare Group. Alberta Farm and Animal Care."},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Choi, Y., Joo, S., and Chun, M.S. (2024). A Legal Ban on Dog Meat Production: Political Decision-Making for an Ethical Community. Animals, 14.","DOI":"10.3390\/ani14152269"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1163\/15685306-12341471","article-title":"Dog \u201cmeat\u201d consumption in China: A survey of the controversial eating habit in two cities","volume":"25","author":"Li","year":"2017","journal-title":"Soc. Anim."},{"key":"ref_64","unstructured":"Ho, L.S.T., Ane, C., Lachlan, R., Tarpinian, K., Feldman, R., Yu, Q., van der Bijl, W., Maspons, J., Vos, R., and Bastide, P. (2024, December 13). Package \u2018Phylolm\u2019. Available online: http:\/\/cran.r-project.org\/web\/packages\/phylolm\/index.html."}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/15\/5\/641\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T16:40:32Z","timestamp":1760028032000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/15\/5\/641"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,2,22]]},"references-count":64,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2025,3]]}},"alternative-id":["ani15050641"],"URL":"https:\/\/doi.org\/10.3390\/ani15050641","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,2,22]]}}}