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July 6, 2026acta ethologica0 citationsOpen Access

An evaluation of Jarman’s hypothesis using grouping behaviour of three sympatric forest-dwelling ungulates in a subtropical cloud forest in Taiwan

HTHayato TakadaYLYu‑Jen LiangNSNick Ching‑Min Sun

Key Points

  • This research investigates if Jarman's hypothesis applies to forest-dwelling ungulates in subtropical cloud forests.
  • Evaluated grouping behaviour of three ungulate species: sambar, Reeves’ muntjac, and Formosan serow.
  • Conducted observational studies in Yushan National Park, Taiwan over an extended period.
  • Assessed group sizes in relation to their diets and other ecological factors.
  • No significant differences in group size among the three ungulate species observed.
  • Each species primarily exhibits solitary behaviour throughout the year.
  • Female groups only formed in sambar, linked to their grazer diet and body size.

Abstract

Jarman’s hypothesis maintains that larger-bodied grazers that feed mainly on grass-like graminoids tend to form large groups, and smaller-bodied browsers that feed mainly on non-graminoids tend to be solitary. Whether this hypothesis held true for forest-dwelling ungulates in subtropical zones had not been investigated. We evaluate Jarman’s hypothesis by investigating grouping behaviour of three sympatric subtropical forest-dwelling ungulates—sambar (a larger-bodied grazer), and Reeves’ muntjac and Formosan serow (smaller-bodied browsers)—in Yushan National Park, central Taiwan. We found no differences in group size among these species, with each being mainly solitary throughout the year. This suggest that grazer-like diets and large body size do not promote increased group size in this forest habitat. Subtropical cloud forests provide many opportunities for animals to conceal themselves, rendering antipredator benefits associated with being in groups of limited value. Female groups—the presumed foundation of gregarious societies—occurred only in sambar, suggesting that a grazer-like diet and large body size provide the basis for group formation. Highly productive and uniformly distributed graminoids—the staple food of sambar, and thought to mitigate intraspecific food competition—possibly explain why female group formation occurred only in sambars. These findings suggest that the direct selective pressure for increased group size in open habitat where places for animals to conceal themselves are few is predation, rather than the need to mitigate intraspecific competition for food resources.

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Cite This Study

Takada et al. (2026) studied this question.

synapsesocial.com/papers/6a4b45b2997070ff83b5b4b3https://doi.org/10.1007/s10211-026-00500-7
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