Herding mechanisms to maintain the cohesion of a harem group: two interaction phases during herding

  • Monamie RinghoferEmail author
  • Clark Kendrick Go
  • Sota Inoue
  • Renata S. Mendonça
  • Satoshi Hirata
  • Takatomi Kubo
  • Kazushi Ikeda
  • Shinya Yamamoto


In animal groups, individual interactions achieve coordinated movements to maintain cohesion. In horse harem groups, herding is a behavior in which males chase females from behind; it is considered to assist with group cohesiveness. However, the mechanisms by which the individuals move to maintain group cohesion are unknown. We applied novel non-invasive methods of drone filming and video tracking to observe horse movements in the field with high temporal and spatial resolution. We tracked all group members and drew trajectories. We analyzed the movements of females and found two phases of interactions based on their timing of movement initiation. The females that moved first were those nearest to the herding male, while the movement initiation of the later females was determined by the distance from the nearest moving female, not by the distance from the herding male. These interactions are unique among animal group movements and might represent a herding mechanism responsible for maintaining group cohesion. This might be due to long-term stable relationships within a harem group and strong social bonds between females. This study showed that the combination of drone filming and video tracking is a useful method for analyzing the movements of animals simultaneously in high resolution.


Drone Video tracking Herding Interaction of individuals Movement Horses 



Special thanks to Viana do Castelo city for supporting our project. We are also grateful to Carlos Pereira and Tetsuro Matsuzawa for managing the field work. We also thank Agostinho Costinha, Lourenço Almada, and the villagers in Montaria for their support during our stay. We would like to thank Editage ( for English language editing.


The study was financially supported by the Japan Society for the Promotion of Science (JSPS) core-to-core CCSN and JSPS-LGP-U04, and Ministry of Education, Culture, Sports, Science and Technology (MEXT) grant No.16H06283 to Tetsuro Matsuzawa, KAKENHI grant Nos. 15H01619, 15H05309, 17H0582, 19H00629,  and Kyoto University SPIRITS grant to Shinya Yamamoto, and KAKENHI grant No. 18K18342 to Monamie Ringhofer.

Compliance with ethical standards

Ethical approval

All applicable international, national, and institutional guidelines for the care and use of animals were followed. The field observations complied with the guidelines for animal studies in the wild issued by the Wildlife Research Center of Kyoto University, Japan.

Conflict of interest

The authors declared that they have no competing interests.

Supplementary material

10164_2019_622_MOESM1_ESM.docx (26 kb)
Supplementary file1 (DOCX 25 kb)
10164_2019_622_MOESM2_ESM.mp4 (20.9 mb)
Supplementary file2 (MP4 21390 kb)


  1. Bourjade M, Thierry B, Hausberger M, Petit O (2015) Is leadership a reliable concept in animals? An empirical study in the horse. PLoS ONE 10:e0126344. CrossRefPubMedPubMedCentralGoogle Scholar
  2. Briard L, Dorn C, Petit O (2015) Personality and affinities play a key role in the organisation of collective movements in a group of domestic horses. Ethology 121:888–902. CrossRefGoogle Scholar
  3. Cameron EZ, Setsaas TH, Linklater WL (2009) Social bonds between unrelated females increase reproductive success in feral horses. Proc Natl Acad Sci USA 106:13850–13853. CrossRefPubMedGoogle Scholar
  4. Christie KS, Gilbert SL, Brown CL, Hatfield M, Hanson L (2016) Unmanned aircraft systems in wildlife research: current and future applications of a transformative technology. Front Ecol Environ 14:241–251. CrossRefGoogle Scholar
  5. Farine DR, Strandburg-Peshkin A, Berger-Wolf T, Ziebart B, Brugere I, Li J, Crofoot MC (2016) Both nearest neighbours and long-term affiliates predict individual locations during collective movement in wild baboons. Sci Rep 6:27704. CrossRefPubMedPubMedCentralGoogle Scholar
  6. Farine DR, Strandburg-Peshkin A, Couzin ID, Berger-Wolf TY, Crofoot MC (2017) Individual variation in local interaction rules can explain emergent patterns of spatial organization in wild baboons. Proc Biol Soc. CrossRefGoogle Scholar
  7. Feh C (2005) Relationships and communication in socially natural horse herds. In: Mills D, McDonnell S (eds) The domestic horse. Cambridge University Press, New York, pp 83–93Google Scholar
  8. Fischhoff IR, Sundaresan SR, Cordingley J, Larkin HM, Sellier MJ, Rubenstein DI (2007) Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii. Anim Behav 73:825–831. CrossRefGoogle Scholar
  9. Franks NR, Richardson T (2006) Teaching in tandem-running ants. Nature 439:153. CrossRefPubMedGoogle Scholar
  10. Herbert-Read JE (2016) Understanding how animal groups achieve coordinated movement. J Exp Biol 219:2971–2983. CrossRefPubMedPubMedCentralGoogle Scholar
  11. Herbert-Read JE, Perna A, Mann RP, Schaerf TM, Sumpter DJ, Ward AJ (2011) Inferring the rules of interaction of shoaling fish. Proc Natl Acad Sci USA 108:18726–18731. CrossRefPubMedGoogle Scholar
  12. Inoue S, Yamamoto S, Ringhofer M, Mendonça RS, Pereira C, Hirata S (2018) Spatial positioning of individuals in a group of feral horses: a case study using drone technology. Mamm Res 64:249–259. CrossRefGoogle Scholar
  13. Jolles JW, Boogert NJ, Sridhar VH, Couzin ID, Manica A (2017) Consistent individual differences drive collective behavior and group functioning of schooling fish. Curr Biol 27(18):2862–2868. CrossRefPubMedPubMedCentralGoogle Scholar
  14. Katz Y, Tunstrøm K, Ioannou C, Huepe C, Couzin ID (2011) Inferring the structure and dynamics of interactions in schooling fish. Proc Natl Acad Sci USA 108:18720–18725. CrossRefPubMedGoogle Scholar
  15. King AJ, Sueur C, Huchard E, Cowlishaw G (2011) A rule-of-thumb based on social affiliation explains collective movements in desert baboons. Anim Behav 82:1337–1345. CrossRefGoogle Scholar
  16. Klingel H (1974) A comparison of the social behaviour of the Equidae. International Union for the Conservation of Nature and Natural Resources, MorgesGoogle Scholar
  17. Krueger K, Flauger B, Farmer K, Hemelrijk C (2014) Movement initiation in groups of feral horses. Behav Proc 103:91–101. CrossRefGoogle Scholar
  18. Lukeman R, Li YX, Edelstein-Keshet L (2010) Inferring individual rules from collective behavior. Proc Natl Acad Sci USA 107:12576–12580. CrossRefPubMedGoogle Scholar
  19. Matsuzawa T (2017) Horse cognition and behavior from the perspective of primatology. Primates 58:473–477. CrossRefPubMedGoogle Scholar
  20. Melletti M, Penteriani V, Mirabile M, Boitani L (2007) Some behavioral aspects of forest buffalo (Syncerus caffer nanus): from herd to individual. J Mammal 88:1312–1318. CrossRefGoogle Scholar
  21. Muro C, Escobedo R, Spector L, Coppinger RP (2011) Wolf-pack (Canis lupus) hunting strategies emerge from simple rules in computational simulations. Behav Proc 88:192–197. CrossRefGoogle Scholar
  22. Nagy M, Akos Z, Biro D, Vicsek T (2010) Hierarchical group dynamics in pigeon flocks. Nature 464:890–894. CrossRefPubMedGoogle Scholar
  23. Ramos A, Petit O, Longour P, Pasquaretta C, Sueur C (2015) Collective decision making during group movements in European bison, Bison bonasus. Anim Behav 109:149–160. CrossRefGoogle Scholar
  24. Reinhardt V (1983) Movement orders and leadership in a semi-wild cattle herd. Behaviour 83:251–264CrossRefGoogle Scholar
  25. Ringhofer M, Inoue S, Mendonça RS, Pereira C, Matsuzawa T, Hirata S, Yamamoto S (2017) Comparison of the social systems of primates and feral horses: data from a newly established horse research site on Serra D’Arga, northern Portugal. Primates 58:479–484. CrossRefPubMedGoogle Scholar
  26. Rubenstein DI (1994) The ecology of female social behavior in horses, zebras, and asses. In: Jarman P, Rossiter A (eds) Animal societies: individuals, interactions, and organization. Kyoto University Press, Kyoto, pp 13–28Google Scholar
  27. Šárová R, Špinka M, Panamá JLA, Šimeček P (2010) Graded leadership by dominant animals in a herd of female beef cattle on pasture. Anim Behav 79:1037–1045. CrossRefGoogle Scholar
  28. Squires VR, Daws GT (1975) Leadership and dominance relationships in Merino and Border Leicester sheep. App Anim Ethol 1:263–274CrossRefGoogle Scholar
  29. Stanley CR, Mettke-Hofmann C, Hager R, Shultz S (2018) Social stability in semiferal ponies: networks show interannual stability alongside seasonal flexibility. Anim Behav 136:175–184. CrossRefGoogle Scholar
  30. Strandburg-Peshkin A, Farine DR, Couzin ID, Crofoot MC (2015) Shared decision-making drives collective movement in wild baboons. Science 348:1358–1361. CrossRefPubMedPubMedCentralGoogle Scholar
  31. Strandburg-Peshkin A, Farine DR, Crofoot MC, Couzin ID (2017) Habitat and social factors shape individual decisions and emergent group structure during baboon collective movement. Elife 6:e19505. CrossRefPubMedPubMedCentralGoogle Scholar
  32. Strandburg-Peshkin A, Papageorgiou D, Crofoot MC, Farine DR (2018) Inferring influence and leadership in moving animal groups. Philos Trans R Soc B 373:20170006. CrossRefGoogle Scholar
  33. Strömbom D, Mann RP, Wilson AM, Hailes S, Morton AJ, Sumpter DJ, King AJ (2014) Solving the shepherding problem: heuristics for herding autonomous, interacting agents. J R Soc Interface. CrossRefPubMedPubMedCentralGoogle Scholar
  34. Sueur C, Kuntz C, Debergue E, Keller B, Robic F, Siegwalt-Baudin F, Richer C, Ramos A, Pelé M (2018) Leadership linked to group composition in Highland cattle (Bos taurus): Implications for livestock management. App Anim Behav Sci 198:9–18. CrossRefGoogle Scholar
  35. Syme GJ, Syme LA (1975) The concept of spatial leadership in farm animals: an experiment with sheep. Anim Behav 23:921–925CrossRefGoogle Scholar
  36. Torney CJ, Lamont M, Debell L, Angohiatok RJ, Leclerc L-M, Berdahl AM (2018) Inferring the rules of social interaction in migrating caribou. Philos Trans R Soc B 373:20170385. CrossRefGoogle Scholar

Copyright information

© Japan Ethological Society 2019

Authors and Affiliations

  1. 1.Institute for Advanced StudyKyoto UniversityKyotoJapan
  2. 2.Nara Institute of Science and TechnologyNaraJapan
  3. 3.Wildlife Research CenterKyoto UniversityKyotoJapan

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