Hares promote seed dispersal and seedling establishment after volcanic eruptions

• Seeds are dispersed everywhere by pellets in hare on a volcano.
• Pellets promote delayed seed germination.
• The relationships between hare and plant are likely to be mutually evolved.

Although seed dispersal through animal guts (endozoochory) is a process that determines plant establishment, the behaviour of carriers mean that the seeds are not always dispersed to suitable habitats for germination. The germinable seeds of Gaultheria miqueliana were stored in the pellets of a hare (Lepus timidus ainu) on Mount Koma in northern Japan. To clarify the roles of hares in seed dispersal and germination, field censuses and laboratory experiments were conducted. The field observations were conducted on pellets and seeds in four habitats (bare ground, G. miqueliana shrub patch, Salix reinii patch, and Larix kaempferi understory), and the laboratory experiments were conducted on seed germination with different light, water potential and cold stratification treatments. The laboratory experiments confirmed that seed germination began a few weeks after the sowing of seeds, independent of cold stratification, when light was sufficient and the water potential was low. The seeds did not germinate at high water potential. The pellets were gradually degraded in situ. More seeds germinated from crushed than from intact pellets. Therefore, over the long term, seeds germinated when exposed to light due to the degradation of pellets. The pellets were proportionally dispersed among the four studied habitats. More seeds sown in the field germinated more in shaded habitats, such as in the Gaultheria patch and the Larix understory, and seeds did not germinate on bare ground, where drought often occurred. Thus, the hares had two roles in the dispersal and germination of seeds: (1) the expansion of G. miqueliana populations through seed dispersal to various habitats and (2) the facilitation of delayed seed germination to avoid risks of hazards such as drought. The relationships between small mammals represented by the hare and the shrubs that produce berries are likely to be more mutually evolved than was previously thought.