The Social and Reproductive Challenges Faced by Free-Roaming Horse (Equus caballus) Stallions.
Abstract: In captivity, intact male horses, due to their sexual drive, are usually socially isolated from other horses. This lifestyle strongly contrasts with that experienced by horses living in free-roaming, feral, or semi-feral conditions, where adult stallions have several roles in their social group, with successful reproduction being their primary drive. Reproductive skew in wild populations is high; many stallions will fail to reproduce at all, while others achieve high levels of reproductive success, siring a large number of foals. Successful stallions are those with particular characteristics and abilities that facilitate harem formation and tenure, allowing them to successfully take over a harem or establish a new one, protect mares from rival stallions, employ appropriate social behaviour to maintain group cohesion, and avoid kin-mating, for example through kin recognition mechanisms. Whilst the life of free-living stallions is far from stress-free, they retain ancestral adaptations to selection pressures (such as predation and competition) exhibited by their natural environment over thousands of years. Here, we discuss the challenges faced by free-living horse stallions, the roles they play in social groups, and their resulting social needs. By understanding these pressures and how stallions react to them, we highlighted the importance of the social environment for the stallion. It is hoped that a better understanding of wild stallions' lives will lead to their needs being more clearly met in captivity, reducing stereotypical behaviour and improving welfare.
Publication Date: 2023-03-24 PubMed ID: 37048406PubMed Central: PMC10093049DOI: 10.3390/ani13071151Google Scholar: Lookup
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
- Journal Article
- 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.
This research paper explores the lifestyle of free-roaming horse stallions and the multitude of roles they assume within their social group, focusing on the key characteristics and abilities contributing to their reproductive success. Insight into the natural behavior and needs of these stallions could enhance their welfare in captivity.
Contrast Between Captivity and Free-Roaming Stallions
- In captivity, male horses are often isolated due to their sexual drive. This way of life sharply contrasts with that of their free-roaming counterparts.
- Free-roaming horse stallions, living under feral or semi-feral conditions, play various roles within their communities. Their primary drive is successful reproduction.
Reproductive Success Variation in Free-Roaming Stallions
- Reproductive skew is high among wild horse populations. Some stallions fail to reproduce entirely, while others sire a significant number of offspring.
- The distinctive traits and abilities of successful stallions have been honed through natural selection. These attributes enable them to establish or seize control of a harem, protect females from potential competitors, maintain group cohesion through appropriate social behaviour, and avoid mating with relatives through kin recognition mechanisms.
Stress and Adaptations in the Life of Free-Living Stallions
- Although the life of a free-roaming stallion is often fraught with stress, these horses are adapted to withstand and navigate the selection pressures of their natural environment, such as competition and predation.
Significance of the Social Environment for Stallions
- This paper stresses the importance of fully understanding the lives and needs of free-roaming stallions to facilitate their welfare in captivity.
- A complete comprehension of their social interactions, roles within groups, and the challenges they encounter could lead to a reduction in stereotypical behavior and an improved standard of welfare when these horses are in captivity.
Cite This Article
APA
Górecka-Bruzda A, Jaworska J, Stanley CR.
(2023).
The Social and Reproductive Challenges Faced by Free-Roaming Horse (Equus caballus) Stallions.
Animals (Basel), 13(7), 1151.
https://doi.org/10.3390/ani13071151 Publication
Researcher Affiliations
- Department of Animal Behaviour and Welfare, Institute of Genetics and Animal Biotechnology, Polish Academy of Sciences, 05-552 Magdalenka, Poland.
- Institute of Animal Reproduction and Food Research, Polish Academy of Sciences, Department of Gamete and Embryo Biology, 10-243 Olsztyn, Poland.
- Animal Behaviour & Welfare Research Group, Department of Biological Sciences, University of Chester, Chester CH1 4B, UK.
Conflict of Interest Statement
The authors declare no conflict of interest.
References
This article includes 172 references
- Bouissou MF. Androgens, aggressive behaviour and social relationships in higher mammals.. Horm Res 1983;18(1-3):43-61.
- Christensen JW, Ladewig J, Søndergaard E, Malmkvist J. Effects of individual versus group stabling on social behaviour in domestic stallions.. Appl. Anim. Behav. Sci. 2002;75:233–248.
- Christensen JW, Zharkikh T, Ladewig J, Yasinetskaya N. Social behaviour in stallion groups (Equus przewalskii and Equus caballus) kept under natural and domestic conditions.. Appl. Anim. Behav. Sci. 2002;76:11–20.
- Hartmann E, Søndergaard E, Keeling LJ. Keeping horses in groups: A review.. Appl. Anim. Behav. Sci. 2012;136:77–87.
- Van Dierendonck MC. The Importance of Social Relationships in Horses.. 2006.
- de Oliveira RA, Aurich C. Aspects of Breeding Stallion Management with Specific Focus on Animal Welfare.. J Equine Vet Sci 2021 Dec;107:103773.
- Burger D, Wedekind C, Wespi B, Imboden I, Meinecke-Tillmann S, Sieme H. The potential effects of social interactions on reproductive efficiency of stallions.. J. Equine Vet. Sci. 2012;32:455–457.
- McDonnell SM. Normal and abnormal sexual behavior.. Vet Clin North Am Equine Pract 1992 Apr;8(1):71-89.
- McDonnell SM. Reproductive behavior of stallions and mares: comparison of free-running and domestic in-hand breeding.. Anim Reprod Sci 2000 Jul 2;60-61:211-9.
- Kirkwood JK, Sainsbury AW. Ethics of interventions for the welfare of free-living wild animals.. Anim. Welf. 1996;5:235–244.
- Harley JJ, Stack JD, Braid H, McLennan KM, Stanley CR. Evaluation of the Feasibility, Reliability, and Repeatability of Welfare Indicators in Free-Roaming Horses: A Pilot Study.. Animals (Basel) 2021 Jul 2;11(7).
- Carenzi C, Verga M. Animal welfare: Review of the scientific concept and definition.. Italian J. Anim. Sci. 2009;8((Suppl. S1)):21–30.
- Duncan IJ. Gordon memorial lecture. Poultry welfare: science or subjectivity?. Br Poult Sci 2002 Dec;43(5 Suppl):643-52.
- Mignon-Grasteau S, Boissy A, Bouix J, Faure J-M, Fisher AD, Hinch GV, Jensen P, Le Neindre P, Mormède P, Prunet P. Genetics of adaptation and domestication in livestock.. Livest. Prod. Sci. 2005;93:3–14.
- Diamond J. Evolution, consequences and future of plant and animal domestication.. Nature 2002 Aug 8;418(6898):700-7.
- Górecka A, Jezierski T. Analiza wskaźników użytkowości rozpłodowej klaczy konika polskiego w latach 1956–1995.. Prac. Mat. Zoot. 1997;51:21–29.
- Górecka A, Jezierski TA, Słoniewski K. Relationships between sexual behaviour, dominant follicle area, uterus ultrasonic image and pregnancy rate in mares of two breeds differing in reproductive efficiency.. Anim Reprod Sci 2005 Jul;87(3-4):283-93.
- Bosh KA, Powell D, Shelton B, Zent W. Reproductive performance measures among Thoroughbred mares in central Kentucky, during the 2004 mating season.. Equine Vet J 2009 Dec;41(9):883-8.
- Lane EA, Bijnen ML, Osborne M, More SJ, Henderson IS, Duffy P, Crowe MA. Key Factors Affecting Reproductive Success of Thoroughbred Mares and Stallions on a Commercial Stud Farm.. Reprod Domest Anim 2016 Apr;51(2):181-7.
- Topczewska J, Krupa W. Impact of maintenance systems on the Hucul mares’ reproductive rates.. Arch. Anim. Breed. 2013;56:246–254.
- Davies Morel MC, Gunnarsson V. A survey of the fertility of Icelandic stallions.. Anim Reprod Sci 2000 Dec 1;64(1-2):49-64.
- Berger J. Organizational systems and dominance in feral horses in the Grand Canyon.. Behav. Ecol. Sociobiol. 1977;2:131–146.
- Boyd L, Keiper R. Behavioural ecology of feral horses.. 2005;pp. 55–82.
- Feh C. Relationships and communication in socially natural horse herds.. 2005;pp. 83–93.
- Krueger K. Social Ecology of Horses.. 2008;pp. 195–206.
- Linklater W. The Social and Spatial Organization of Horses.. 1998.
- Tyler SJ. The behaviour and social organization of the New Forest ponies.. Anim. Behav. Monogr. 1972;5:87–196.
- Klingel H. Social Behaviour of African Equidae.. Zool. Afr. 1972;7:175–185.
- Klingel H. A comparison of the social behaviour of the Equidae; Proceedings of the International Symposium the Behaviour of Ungulates and Its Relation to Management; Calgary, AB, Canada.. 2–5 November 1974; pp. 124–132.
- Feh C. Social behaviour and relationships of Przewalski horses in Dutch semi-reserves.. Appl. Anim. Behav. Sci. 1988;21:71–87.
- Hoffmann R. On the development of social behaviour in immature males of a feral horse population (Equus przewalskii f. caballus). Z. Saugetierkd. 1985;50:302–314.
- Bourjade M, Tatin L, King SRB, Feh C. Early reproductive success, preceding bachelor ranks and their behavioural correlates in young Przewalski’s stallions.. Ethol. Ecol. Evol. 2009;21:1–14.
- Gersick AS, Rubenstein DI. Physiology modulates social flexibility and collective behaviour in equids and other large ungulates.. Philos Trans R Soc Lond B Biol Sci 2017 Aug 19;372(1727).
- Feh C. Alliances and reproductive success in Camargue stallions.. Anim Behav 1999 Mar;57(3):705-713.
- Hampson BA, de Laat MA, Mills PC, Pollitt CC. Distances travelled by feral horses in 'outback' Australia.. Equine Vet J Suppl 2010 Nov;(38):582-6.
- McDonnell SM, Murray SC. Bachelor and harem stallion behavior and endocrinology.. Biol. Reprod. 1995;52:577–590.
- McDonnell SM, de Haviland JCS. Agonistic ethogram of the equid bachelor band.. Appl. Anim. Behav. Sci. 1995;43:147–188.
- Linnartz L, Linnartz-Nieuwdorp E. The social organisation of natural herds of koniks (Equus caballus): Subordinate stallions, rule or exception?. Lutra 2017;60:27–42.
- Hex SB, Tombak K, Rubenstein DI. A new classification of mammalian uni-male multi-female groups based on the fundamental principles governing inter-and intrasexual relationships.. Behav. Ecol. Sociobiol. 2021;75:1–32.
- Linklater WL, Cameron EZ. Tests for cooperative behaviour between stallions.. Anim Behav 2000 Dec;60(6):731-743.
- Linklater WL, Cameron EZ, Minot EO, Stafford KJ. Stallion harassment and the mating system of horses.. Anim Behav 1999 Aug;58(2):295-306.
- Keiper R. The Assateague Ponies.. 1985.
- Jaworska J, Jaworski Z, McDonnell SM, Górecka-Bruzda A. Harem stallion changes are not associated with diminished reproductive performance of females in semi-feral Konik polski horses (Equus caballus).. Theriogenology 2020 Jul 15;151:1-6.
- Stanley CR, Mettke-Hofmann C, Hager R, Shultz S. Social stability in semiferal ponies: Networks show interannual stability alongside seasonal flexibility.. Anim. Behav. 2018;136:175–184.
- Cameron EZ, Setsaas TH, Linklater WL. Social bonds between unrelated females increase reproductive success in feral horses.. Proc Natl Acad Sci U S A 2009 Aug 18;106(33):13850-3.
- Sigurjónsdóttir H, Snorrason S, van Dierendonck M, Thórhallsdóttir A. Social relationships in a group of horses without a mature stallion.. Behaviour 2003;140:783–804.
- Crowell-Davis SL, Houpt KA, Kane L. Play development in Welsh pony (Equus caballus) foals.. Appl. Anim. Behav. Sci. 1987;18:119–131.
- Šandlová K, Komárková M, Ceacero F. Daddy, daddy cool: stallion-foal relationships in a socially-natural herd of Exmoor ponies.. Anim Cogn 2020 Jul;23(4):781-793.
- Sigurjónsdóttir H, Haraldsson H. Significance of Group Composition for the Welfare of Pastured Horses.. Animals (Basel) 2019 Jan 5;9(1).
- McDonnell SM, Poulin A. Equid play ethogram.. Appl. Anim. Behav. Sci. 2002;78:263–290.
- Cameron EZ, Linklater WL. Individual mares bias investment in sons and daughters in relation to their condition.. Anim Behav 2000 Sep;60(3):359-367.
- Cameron EZ, Linklater WL, Stafford KJ, Minot EO. Maternal investment results in better foal condition through increased play behaviour in horses.. Anim. Behav. 2008;76:1511–1518.
- Gray ME. An infanticide attempt by a free-roaming feral stallion (Equus caballus).. Biol Lett 2009 Feb 23;5(1):23-5.
- Gray ME, Cameron EZ, Peacock MM, Thain DS, Kirchoff VS. Are low infidelity rates in feral horses due to infanticide?. Behav. Ecol. Soc. 2012;66:529–537.
- Duncan P. Foal killing by stallions.. Appl. Anim. Ethol. 1982;8:567–570.
- Boyd LE. The behaviour of Przewalski’s horses and its importance to their management.. Appl. Anim. Behav. Sci. 1991;29:301–318.
- Kolter L, Zimmermann W. Social behaviour of Przewalski horses (Equus p. przewalskii) in the Cologne Zoo and its consequences for management and housing.. Appl. Anim. Behav. Sci. 1988;21:117–145.
- Ryder OA, Massena R. A case of male infanticide in Equus przewalskii.. Appl. Anim. Behav. Sci. 1998;21:187–190.
- Zarkikh TL. The cases of infanticide in the Przewalskii horses in Askanja-Nova.. Vestnik Zool. 1999;11:80–82.
- Lema FJ, Ribeiro S, Palacios V. Observations of wolves hunting free-ranging horses in NW Iberia.. Carnivore Damage Prevention 2022;24:1–10.
- Van Duyne C, Ras E, De Vos AE, De Boer WF, Henkens RJ, Usukhjargal D. Wolf predation among reintroduced przewalski horses in Hustai National Park, Mongolia.. J. Wild. Manag. 2009;73:836–843.
- Andreasen AM, Stewart KM, Longland WS, Beckmann JP. Prey specialization by cougars on feral horses in a desert environment.. J. Wildl. Manag. 2021;85:1104–1120.
- Freitas J, Álvares F. Economic impact of wolf predation on free-ranging horses in Portugal.. Carniv. Damage Prev. News. 2021;23:37–47.
- Klingel H. Social organization and reproduction in equids.. J Reprod Fertil Suppl 1975 Oct;(23):7-11.
- Salter RE, Hudson RJ. Social organization of feral horses in western Canada.. Appl. Anim. Ethol. 1982;8:207–223.
- Berger J. Wild Horses of the Great Basin: Social Competition and Population Size.. 1986.
- Shultz S, Stanley CR. Mummy’s boys: Sex differential maternal-offspring bonds in semi-feral horses.. Behaviour 2012;149:251–274.
- Rutberg AT, Keiper RR. Proximate causes of natal dispersal in feral ponies: Some sex differences.. Anim. Behav. 1993;46:969–975.
- Jaworowska M. Die Fortpflanzung primitiven polnischen Pferden, die frei im Wald- schutzgebiet leben—Säugetierk. Mitteil.. 1981;29:46–71.
- Górecka-Bruzda A, Jaworska J, Siemieniuch M, Jaworski Z, Stanley CR, Wocławek-Potocka I, Lansade L. Human-controlled reproductive experience may contribute to incestuous behavior observed in reintroduced semi-feral stallions (Equuscaballus).. Theriogenology 2022 Mar 1;180:82-86.
- Kaseda Y, Khalil AM. Harem size reproductive success of stallions in Misaki feral horses.. Appl. Anim. Behav. Sci. 1996;47:163–173.
- Kaseda Y, Ogawa H, Khalil AM. Causes of natal dispersal and emigration and their effects on harem formation in Misaki feral horses.. Equine Vet J 1997 Jul;29(4):262-6.
- Heitor F, Vicente L. Dominance relationships and patterns of aggression in a bachelor group of Sorraia horses (Equus caballus). J. Ethol. 2010;28:35–44.
- Khalil AM, Murakami N. Factors affecting the harem formation process by young Misaki feral stallions.. J Vet Med Sci 1999 Jun;61(6):667-71.
- Tilson RL, Sweeny KA, Binczik GA, Reindl NJ. Buddies and bullies: Social structure of a bachelor group of Przewalski horses.. Appl. Anim. Behav. Sci. 1988;21:169–185.
- Khalil AM, Murakami N. Effect of natal dispersal on the reproductive strategies of the young Misaki feral stallions.. Appl. Anim. Behav. Sci. 1999;62:281–291.
- Medill SA. Sociality of Sable island Horses: Population, Group, and Individual Interactions.. 2018.
- D'Souza-Anjo M, Christensen BW, Brabender K, Zimmermann W, Kass PH, Schwarzenberger F. A comparison of fecal steroid metabolite concentrations between harem and bachelor stallions in a free-Ranging population of przewalski's horses (Equus ferus przewalskii).. Zoo Biol 2017 Mar;36(2):127-131.
- Donaldson MT, McDonnell SM, Schanbacher BJ, Lamb SV, McFarlane D, Beech J. Variation in plasma adrenocorticotropic hormone concentration and dexamethasone suppression test results with season, age, and sex in healthy ponies and horses.. J Vet Intern Med 2005 Mar-Apr;19(2):217-22.
- Jones MM, Nuñez CMV. Laissez-Faire Stallions? Males' Fecal Cortisol Metabolite Concentrations Do Not Vary with Increased Female Turnover in Feral Horses (Equus caballus).. Animals (Basel) 2023 Jan 3;13(1).
- Briefer Freymond S, Briefer EF, Von Niederhäusern R, Bachmann I. Pattern of social interactions after group integration: a possibility to keep stallions in group.. PLoS One 2013;8(1):e54688.
- Popescu S, Lazar EA, Borda C, Blaga Petrean A, Mitrănescu E. Changes in Management, Welfare, Emotional State, and Human-Related Docility in Stallions.. Animals (Basel) 2022 Oct 30;12(21).
- Gehlen H, Krumbach K, Thöne-Reineke C. Keeping Stallions in Groups-Species-Appropriate or Relevant to Animal Welfare?. Animals (Basel) 2021 May 4;11(5).
- Klingel H. Social organization of feral horses.. J Reprod Fertil Suppl 1982;32:89-95.
- Franke Stevens E. Instability of harems of feral horses in relation to season and presence of subordinate stallions.. Behaviour 1990;112:149–161.
- Kaseda Y, Khalil AM, Ogawa H. Harem stability and reproductive success of Misaki feral mares.. Equine Vet J 1995 Sep;27(5):368-72.
- Monard AM, Duncan P, Boy V. The proximate mechanism of natal dispersal in female horses.. Behaviour 1996;133:1095–1124.
- Feist JD, McCullough DR. Reproduction in feral horses.. J Reprod Fertil Suppl 1975 Oct;(23):13-8.
- Miller R. Male aggression, dominance and breeding behavior in Red Desert feral horses.. Z. Tierpsychol. 1981;57:340–351.
- Rutberg AT. Inter-group transfer in Assateague pony mares.. Anim. Behav. 1990;40:945–952.
- Burger D, Meuwly C, Marti E, Sieme H, Oberthür M, Janda J, Meinecke-Tillmann S, Wedekind C. MHC-correlated preferences in diestrous female horses (Equus caballus).. Theriogenology 2017 Feb;89:318-323.e1.
- Jaworska J, Jaworski Z, Tobolski D, Siemieniuch M, Janowski T, Górecka-Bruzda A. Selection of reproductive partners in semi-feral horses (Equus caballus) is not influenced by major histocompatibility complex (MHC): A field study.. Appl. Anim. Behav. Sci. 2020;226:104973.
- Jones MM, Proops L, Nuñez CM. Rising up to the challenge of their rivals: Mare infidelity intensifies stallion response to playback of aggressive conspecific vocalizations.. Appl. Anim. Behav. Sci. 2020;225:104949.
- Madosky JM, Rubenstein DI, Howard JJ, Stuska S. The effects of immunocontraception on harem fidelity in a feral horse (Equus caballus) population.. Appl. Anim. Behav. Sci. 2010;128:50–56.
- Granquist SM, Thorhallsdottir AG, Sigurjonsdottir H. The effect of stallions on social interactions in domestic and semi feral harems.. Appl. Anim. Behav. Sci. 2012;141:49–56.
- Duncan P. Time-budgets of Camargue horses II. Time-budgets of adult horses and weaned sub-adults.. Behaviour 1980;72:26–48.
- Linklater S. Spatial structure and range use by Kaimanawa wild horses (Equus caballus: Equidae). N. Zeal. J. Ecol. 2000;24:139.
- Pacheco MA, Herrera EA. Social structure of feral horses in the llanos of Venezuela.. J. Mammal. 1997;78:15–22.
- Feh C. Long-term paternity data in relation to different aspects of rank for camargue stallions, Equus caballus.. Anim. Behav. 1990;40:995–996.
- Pinto P, Hirata S. Does size matter? Examining the possible mechanisms of multi-stallion groups in horse societies.. Behav Processes 2020 Dec;181:104277.
- Heitor F, do Mar Oom M, Vicente L. Social relationships in a herd of Sorraia horses Part II. Factors affecting affiliative relationships and sexual behaviours.. Behav Processes 2006 Nov 1;73(3):231-9.
- Jezierski T, Górecka A. Changes in the horses heart rate during different levels of social isolation.. Anim. Sci. Pap. Rep. 2000;18:33–41.
- Houpt KA, Keiper R. The position of the stallion in the equine dominance hierarchy of feral and domestic ponies.. J. Anim. Sci. 1982;54:945–950.
- Krueger K, Flauger B, Farmer K, Hemelrijk C. Movement initiation in groups of feral horses.. Behav Processes 2014 Mar;103:91-101.
- Briard L, Deneubourg JL, Petit O. How stallions influence the dynamic of collective movements in two groups of domestic horses, from departure to arrival.. Behav Processes 2017 Sep;142:56-63.
- Grogan EH, McDonnell SM. Injuries and blemishes in a semi-feral herd of ponies.. J. Equine Vet. Sci. 2005;25:26–30.
- Górecka-Bruzda A, Jaworski Z, Jaworska J, Siemieniuch M. Welfare of Free-Roaming Horses: 70 Years of Experience with Konik Polski Breeding in Poland.. Animals (Basel) 2020 Jun 24;10(6).
- Jaworski Z. (University of Warmia and Mazury, Olsztyn, Poland). Personal Communication. 2022.
- Rabb MH, Thompson DL Jr, Barry BE, Colborn DR, Garza F Jr, Hehnke KE. Effects of sexual stimulation, with and without ejaculation, on serum concentrations of LH, FSH, testosterone, cortisol and prolactin in stallions.. J Anim Sci 1989 Oct;67(10):2724-9.
- Tamanini C, Giordano N, Bono G, Mattioli M. Plasma-concentration of testosterone, androstenedione, LH and estrone sulfate in male horses up to 7 years of age.. Arch. Wet. Ital. 1983;34:25–29.
- Aurich J, Wulf M, Ille N, Erber R, von Lewinski M, Palme R, Aurich C. Effects of season, age, sex, and housing on salivary cortisol concentrations in horses.. Domest Anim Endocrinol 2015 Jul;52:11-6.
- Ginther OJ, Lara A, Leoni M, Bergfelt DR. Herding and snaking by the harem stallion in domestic herds.. Theriogenology 2002 May;57(8):2139-46.
- Costa H, Fragoso S, Heitor F. The relevance of affiliative relationships in horses: Review and future directions.. Pet. Behav. Sci. 2019;8:11–26.
- Kimura R. Mutual grooming and preferred associate relationships in a band of free-ranging horses.. Appl. Anim. Behav. Sci. 1998;59:265–276.
- Feh C, de Mazières J. Grooming at a preferred site reduces heart rate in horses.. Anim. Behav. 1993;46:1191–1194.
- Rho JR, Srygley RB, Choe JC. Age-specific fertility rates of the Jeju pony (Equus caballus). J. Ethol. 2004;22:113–117.
- Asa CS, Goldfoot DA, Ginther OJ. Sociosexual behavior and the ovulatory cycle of ponies (Equus caballus) observed in harem groups.. Horm Behav 1979 Aug;13(1):49-65.
- Shimada M, Suzuki N. The Contribution of Mutual Grooming to Affiliative Relationships in a Feral Misaki Horse Herd.. Animals (Basel) 2020 Sep 3;10(9).
- Boyd LE. The Natality, Foal Survivorship, and Mare-Foal Behavior of Feral Horses in Wyoming’s Red Desert.. 1980.
- Šandlová K, Komárková M, Ceacero F. Oh, daddy where are thou: Social context modifies foals’ behaviour differently for colts and fillies.. J. Equine Vet. Sci. 2021;100:103543.
- Berger J, Cunningham C. INFLUENCE OF FAMILIARITY ON FREQUENCY OF INBREEDING IN WILD HORSES.. Evolution 1987 Jan;41(1):229-231.
- Bowling AT, Touchberry RW. Parentage of Great Basin feral horses.. J. Wildl. Manag. 1990;54:424–429.
- Pinto P, Mendonça RS, Hirata S. Examining the costs and benefits of male-male associations in a group-living equid.. Appl. Anim. Behav. Sci. 2022;2022:105660.
- Linklater WL, Cameron EZ, Stafford KJ, Minot EO. Removal experiments indicate that subordinate stallions are not helpers.. Behav Processes 2013 Mar;94:1-4.
- Reznick D, Nunney L, Tessier A. Big houses, big cars, superfleas and the costs of reproduction.. Trends Ecol Evol 2000 Oct 1;15(10):421-425.
- Speakman JR. The physiological costs of reproduction in small mammals.. Philos Trans R Soc Lond B Biol Sci 2008 Jan 27;363(1490):375-98.
- Clutton-Brock TH, Vincent AC. Sexual selection and the potential reproductive rates of males and females.. Nature 1991 May 2;351(6321):58-60.
- Hayward A, Gillooly JF. The cost of sex: quantifying energetic investment in gamete production by males and females.. PLoS One 2011 Jan 24;6(1):e16557.
- Jeannerat E, Marti E, Berney C, Janett F, Bollwein H, Sieme H, Burger D, Wedekind C. Stallion semen quality depends on major histocompatibility complex matching to teaser mare.. Mol Ecol 2018 Feb;27(4):1025-1035.
- Pusey A, Wolf M. Inbreeding avoidance in animals.. Trends Ecol Evol 1996 May;11(5):201-6.
- Duncan P, Feh C, Gleize JC, Malkas P, Scott AM. Reduction of inbreeding in a natural herd of horses.. Anim. Behav. 1984;32:520–527.
- Houpt KA. Formation and dissolution of the mare–foal bond.. Appl. Anim. Behav. Sci. 2002;78:319–328.
- Khalil A, Kaseda Y. Early experience affects developmental behaviour and timing of harem formation in Misaki horses.. Appl. Anim. Behav. Sci. 1998;59:253–263.
- Watts ET, Johnson CN, Carver S, Butler C, Harvey AM, Cameron EZ. Maternal protectiveness in feral horses: Responses to intraspecific and interspecific sources of risk.. Anim. Behav. 2020;159:1–11.
- Hrdy SB. Infanticide among animals—Review, classification, and examination of the implications for the reproductive strategies of females.. Ethol. Sociobiol. 1979;1:13–40.
- Bartoš L, Dušek A, Bartošová J, Pluháček J, Putman R. How to escape male infanticide: Mechanisms for avoiding or terminating pregnancy in mammals.. Mamm. Rev. 2021;51:143–153.
- Aurich C. Reproductive cycles of horses.. Anim Reprod Sci 2011 Apr;124(3-4):220-8.
- Feh C, Munkhtuya B. Male infanticide and paternity analyses in a socially natural herd of Przewalski's horses: sexual selection?. Behav Processes 2008 Jul;78(3):335-9.
- Bartoš L, Bartošová J, Pluháček J. Male-free environment prevents pregnancy disruption in domestic horse mares mated away of home.. Appl. Anim. Behav. Sci. 2018;200:67–70.
- Bartoš L, Bartošová J, Pluháček J. Pregnancy disruption in artificially inseminated domestic horse mares as a counterstrategy against potential infanticide.. J Anim Sci 2015 Nov;93(11):5465-8.
- Bartoš L, Bartošová J, Pluháček J, Šindelářová J. Promiscuous behavior disrupts pregnancy block in domestic horse mares.. Behav. Ecol. Sociobiol. 2011;65:1567–1572.
- Noronha LE, Antczak DF. Maternal immune responses to trophoblast: the contribution of the horse to pregnancy immunology.. Am J Reprod Immunol 2010 Oct;64(4):231-44.
- Crabtree JR, Chang Y, De Mestre AM. Clinical presentation, treatment and possible causes of persistent endometrial cups illustrated by two cases.. Equine Vet. Edu. 2012;24:251–259.
- Setchell PB. Domestication and reproduction.. Anim. Reprod. Sci. 1992;28:195–202.
- Hall LK, Larsen RT, Knight RN, McMillan BR. Feral horses influence both spatial and temporal patterns of water use by native ungulates in a semi-arid environment.. Ecosphere 2018;9:e02096.
- Nimmo DG, Miller KK, Adams R. Managing feral horses in Victoria: A study of community attitudes and perceptions.. Ecol. Manag. Restor. 2007;8:237–241.
- Cornelissen P. Large Herbivores as a Driving Force of Woodland-Grassland Cycles. The Mutual Interactions between the Population Dynamics of Large Herbivores and Vegetation Development in a Eutrophic Wetland.. 2017.
- Rubenstein DI, Hohmann ME. Parasites and Social Behavior of Island Feral Horses.. Oikos 1989;55:312–320.
- Bechert US, Turner JW Jr, Baker DL, Eckery DC, Bruemmer JE, Lyman CC, Fraker MA. Fertility control options for management of free-roaming horse populations.. Hum. Wildl. Interact. 2022;16:5.
- Kirkpatrick JF, Liu IM, Turner JW Jr, Naugle R, Keiper R. Long-term effects of porcine zonae pellucidae immunocontraception on ovarian function in feral horses (Equus caballus).. J Reprod Fertil 1992 Mar;94(2):437-44.
- Kirkpatrick JF, Lyda RO, Frank KM. Contraceptive vaccines for wildlife: a review.. Am J Reprod Immunol 2011 Jul;66(1):40-50.
- Powell DM. Preliminary evaluation of porcine zona pellucida (PZP) immunocontraception for behavioral effects in feral horses (Equus caballus).. J Appl Anim Welf Sci 1999;2(4):321-35.
- Ransom JI, Powers JG, Garbe HM, Oehler Sr MW, Terry M, Nett TM, Baker DL. Behavior of feral horses in response to culling and GnRH immunocontraception.. Appl. Anim. Behav. Sci. 2014;157:81–92.
- Nuñez CM, Adelman JS, Mason C, Rubenstein DI. Immunocontraception decreases group fidelity in a feral horse population during the non-breeding season.. Appl. Anim. Behav. Sci. 2009;117:74–83.
- King SR, Schoenecker KA, Cole MJ. Effect of adult male sterilization on the behavior and social associations of a feral polygynous ungulate: The horse.. Appl. Anim. Behav. Sci. 2022;249:105598.
- Eagle TC, Asa CS, Garrott RA, Plotka ED, Siniff DB, Tester JR. Efficacy of dominant male sterilization to reduce reproduction in feral horses.. Wildl. Soc. Bull. 1993;21:116–121.
- Berger J, Kock M, Cunningham C, Dodson N. Chemical restraint of wild horses: effects on reproduction and social structure.. J Wildl Dis 1983 Jul;19(3):265-8.
- Ponthier J, Rigaux G, Parrilla-Hernandez S, Egyptien S, Gatez C, Carrasco Leroy C, Deleuze S. Case Report: Suppression of Harem Stallion Behavior and Fertility Following Anti-Gonadotropin-Releasing Hormone Vaccination of a Captive Wild Przewalski's Horse (Equus ferus przewalskii).. Front Vet Sci 2020;7:569185.
- Vitoria A, Romero A, Fuente S, Barrachina L, Vazquez FJ. Application of a laparoscopic technique for vasectomy in standing horses.. Vet Rec 2019 Sep 21;185(11):345.
- Pozor MA, Macpherson ML, McDonnell SM, Nollin M, Roser JF, Love C, Runyon S, Thomas BF, Troedsson MH. Indenopyride derivative RTI-4587-073(l): a candidate for male contraception in stallions.. Theriogenology 2013 Dec;80(9):1006-16.
- Scully CM, Lee RL, Pielstick L, Medlock J, Patton KM, Collins GH, Kutzler MA. COMPARISON OF CHEMICAL AND SURGICAL VASECTOMY ON TESTICULAR ACTIVITY IN FREE-ROAMING HORSES (EQUUS CABALLUS).. J Zoo Wildl Med 2015 Dec;46(4):815-24.
- Palmer C, Pedersen HG, Sandøe P. Beyond castration and culling: Should we use non-surgical, pharmacological methods to control the sexual behavior and reproduction of animals?. J. Agric. Environ. Ethics. 2018;31:197–218.
- Lenz TR. The Unwanted Horse in the United States: An Overview of the Issue.. J. Equine Vet. Sci. 2009;29:253–258.
- Ward L, Lindsey S, Martin JM, Nicodemus M, Memili E. Challenges and opportunities in rising feral horse populations.. Prof. Anim. Sci. 2016;32:717–724.
- Lansade L, Lévy F, Parias C, Reigner F, Górecka-Bruzda A. Weaned horses, especially females, still prefer their dam after five months of separation.. Animal 2022 Oct;16(10):100636.
- Ladewig J., Søndergaard E., Christensen J.W. In: The Domestic Horse: The Origins, Development and Management of Its Behaviour. Mills D.S., McDonnell S.M., editors. Cambridge University Press; Cambridge, UK: 2005. pp. 139–149.
- Poyato-Bonilla J, Perdomo-González DI, Sánchez-Guerrero MJ, Varona L, Molina A, Casellas J, Valera M. Genetic inbreeding depression load for morphological traits and defects in the Pura Raza Española horse.. Genet Sel Evol 2020 Oct 20;52(1):62.
- Dini P, Bartels T, Revah I, Claes AN, Stout TAE, Daels P. A retrospective study on semen quality parameters from four different Dutch horse breeds with different levels of inbreeding.. Theriogenology 2020 Nov;157:18-23.
- Hill EW, Stoffel MA, McGivney BA, MacHugh DE, Pemberton JM. Inbreeding depression and the probability of racing in the Thoroughbred horse.. Proc Biol Sci 2022 Jun 29;289(1977):20220487.
- Poncet P-A, Bachmann I, Burkhardt R, Ehrbar B, Herrmann R, Friedli K, Leuenberger H, Lüth A, Montavon S, Pfammatter M. Ethical Reflections on the Dignity and Welfare of Horses and Other Equids—Pathways to Enhanced Protection.. 2022.
- Cerasoli F, Podaliri Vulpiani M, Saluti G, Conte A, Ricci M, Savini G, D'Alterio N. Assessment of Welfare in Groups of Horses with Different Management, Environments and Activities by Measuring Cortisol in Horsehair, Using Liquid Chromatography Coupled to Hybrid Orbitrap High-Resolution Mass Spectrometry.. Animals (Basel) 2022 Jul 6;12(14).
- Briard L, Deneubourg JL, Petit O. L’étalon, un cheval presque comme les autres: Statut et comportement social de l'étalon vivant en groupe familial. 2016;pp. 121–124.
Citations
This article has been cited 3 times.- Khan IU, Khairullah AR, Khan AY, Rehman AU, Mustofa I. Strategic approaches to improve equine breeding and stud farm outcomes. Vet World 2025 Feb;18(2):311-328.
- Lovász L, Sommer-Trembo C, Barth JMI, Scasta JD, Grancharova-Hill R, Lemoine RT, Kerekes V, Merckling L, Bouskila A, Svenning JC, Fages A. Rewilded horses in European nature conservation - a genetics, ethics, and welfare perspective. Biol Rev Camb Philos Soc 2025 Feb;100(1):407-427.
- Jastrzębska E, Siemieniuch M, Bizio A, Pietruszka J, Górecka-Bruzda A. Are Users Good Assessors of Social Dominance in Domestic Horses?. Animals (Basel) 2024 Jul 7;14(13).
Use Nutrition Calculator
Check if your horse's diet meets their nutrition requirements with our easy-to-use tool Check your horse's diet with our easy-to-use tool
Talk to a Nutritionist
Discuss your horse's feeding plan with our experts over a free phone consultation Discuss your horse's diet over a phone consultation
Submit Diet Evaluation
Get a customized feeding plan for your horse formulated by our equine nutritionists Get a custom feeding plan formulated by our nutritionists