Changes in understory species occurrence of a secondary broadleaved forest after mass mortality of oak trees under deer foraging pressure

The epidemic of mass mortality of oak trees by Japanese oak wilt has affected secondary deciduous broadleaved forests that have been used as coppices in Japan. The dieback of oak trees formed gaps in the crown that would be expected to enhance the regeneration of shade-intolerant pioneer species. However, foraging by sika deer Cervus nippon has also affected forest vegetation, and the compound effects of both on forest regeneration should be considered when they simultaneously occur. A field study was conducted in Kyôto City, Japan to investigate how these compound effects affected the vegetation of the understory layer of these forests. The presence/absence of seedlings and saplings was observed for 200 quadrats sized 5 m ×5 m for each species in 1992, before the mass mortality and deer encroachment, and in 2014 after these effects. A hierarchical Bayesian model was constructed to explain the occurrence, survival, and colonization of each species with their responses to the gaps that were created, expanded, or affected by the mass mortality of Quercus serrata trees. The species that occurred most frequently in 1992, Eurya japonica, Quercus glauca, and Cleyera japonica, also had the highest survival probabilities. Deer-unpalatable species such as Symplocos prunifolia and Triadica sebifera had higher colonization rates in the gaps, while the deer-palatable species Aucuba japonica had the smallest survival probability. The gaps thus promoted the colonization of deer-unpalatable plant species such as Symplocos prunifolia and Triadica sebifera. In the future, such deer-unpalatable species may dominate gaps that were created, expanded, or affected by the mass mortality of oak trees.