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Proceedings. Biological sciences2026; 293(2063); 20252468; doi: 10.1098/rspb.2025.2468

Spatial strategies in non-territorial societies: how feral horses maintain boundaries with other groups.

Abstract: Encounters between competitive individuals or groups are common in social animals and can involve costly aggression; thus, animals often employ strategies to minimize direct conflict. However, research on whether and how animals adjust their group spatial structure when they encounter or spatially co-occur with a different group remains limited. We investigated how non-territorial units in feral horse multilevel societies manage spatial encounters with neighbouring units. We observed 25 reproductive units in northern Portugal, using drones, and employed statistical analyses to quantify spatial dynamics. We found that horses actively adjusted their spatial formations depending on the proximity of other units. Units became more circular and cohesive when other units approached, but when extremely close, they elongated their shapes to avoid boundary crossings. These adjustments indicate that horses maintain flexible unit boundaries to prevent mixing with other units. A notable exception was a particular pair of units that frequently crossed boundaries and intermixed, representing a unique social level within the horse multilevel society. Overall, our findings indicate that feral horses respond adaptively to inter-unit encounters by balancing the benefits of aggregation with the need to reduce direct overlap, providing new insights into the spatial organization of non-territorial societies.
Publication Date: 2026-01-21 PubMed ID: 41558789DOI: 10.1098/rspb.2025.2468Google Scholar: Lookup
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  • Journal Article

Summary

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Overview

  • This study examined how non-territorial groups of feral horses manage space and avoid conflict when they come close to other groups.
  • Researchers found that horses change their group shape and spacing in response to other groups nearby, helping them maintain boundaries without physical aggression.

Introduction to the Research Topic

  • Social animals often face encounters with other individuals or groups, which can lead to costly aggressive interactions.
  • To reduce conflict, animals use spatial strategies to maintain group boundaries and minimize direct confrontations.
  • Despite this, little is known about how non-territorial animals adjust the spatial structure of their groups during encounters with neighboring groups.
  • Feral horses live in multilevel societies but are generally non-territorial, making them a valuable model for studying group spatial dynamics and boundary maintenance.

Methods and Data Collection

  • The researchers observed 25 reproductive units of feral horses in northern Portugal.
  • They used drone technology to accurately capture spatial formations and movements of these units from a bird’s-eye view.
  • Statistical analyses were applied to the spatial data to quantify how group shapes and formations changed depending on the proximity of other horse units.

Key Findings on Spatial Adjustments

  • When other horse units were approaching, the groups became more circular and tightly cohesive, likely to reinforce group unity and boundary clarity.
  • As the units got extremely close, they shifted to more elongated shapes, which helped avoid physical boundary crossing and potential conflicts.
  • These flexible spatial adjustments suggest that feral horses actively work to maintain clear boundaries between groups without resorting to aggressive territorial defense.

Social Exceptions and Multilevel Society Implications

  • A notable exception was one pair of units that frequently crossed boundaries and intermixed, showing a higher level of social integration.
  • This behavior represents an additional, unique social level within the horse multilevel society, indicating variability in social and spatial relationships.

Conclusions and Broader Significance

  • The study highlights how non-territorial animals like feral horses manage spatial organization through adaptive strategies rather than by defending fixed territories.
  • Flexible changes in group shape and cohesion strike a balance between staying aggregated (for social benefits and safety) and avoiding overlap (to reduce conflict).
  • These insights enrich understanding of spatial ecology and social behavior in animals without rigid territorial boundaries, helping to explain how complex social systems can be maintained peacefully.

Cite This Article

APA
Maeda T, Inoue S, Ringhofer M, Hirata S, Yamamoto S. (2026). Spatial strategies in non-territorial societies: how feral horses maintain boundaries with other groups. Proc Biol Sci, 293(2063), 20252468. https://doi.org/10.1098/rspb.2025.2468

Publication

ISSN: 1471-2954
NlmUniqueID: 101245157
Country: England
Language: English
Volume: 293
Issue: 2063
PII: 20252468

Researcher Affiliations

Maeda, Tamao
  • Research Center for Integrative Evolutionary Science, The Graduate University for Advanced Studies SOKENDAI, Kanagawa, Japan.
  • Wildlife Research Center, Kyoto University, Kyoto, Japan.
Inoue, Sota
  • Institute for Advanced Research, Nagoya University, Aichi, Japan.
  • Graduate School for Environmental Studies, Nagoya University, Aichi, Japan.
  • Graduate School of Information Science and Technology, The University of Osaka, Osaka, Japan.
Ringhofer, Monamie
  • Faculty of Life and Environmental Sciences, Department of Animal Sciences, Teikyo University of Science, Tokyo, Japan.
Hirata, Satoshi
  • Wildlife Research Center, Kyoto University, Kyoto, Japan.
Yamamoto, Shinya
  • Institute for Advanced Study, Kyoto University, Kyoto, Japan.
  • Institute for the Future of Human Society, Kyoto University, Kyoto, Japan.

MeSH Terms

  • Animals
  • Horses / physiology
  • Social Behavior
  • Portugal
  • Male
  • Female
  • Behavior, Animal

Grant Funding

  • Japan Science and Technology Agency
  • Japan Society for the Promotion of Science

Citations

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