Herbivory in canopy gaps created by a typhoon varies by understory plant leaf phenology

1. Availabilities of light and soil nitrogen for understory plants vary by extent of canopy gap formation through typhoon disturbance. We predicted that variation in resource availability and herbivore abundance in canopy gaps would affect herbivory through variation in leaf traits among plant species. We studied six understory species that expand their leaves before or after canopy closure in deciduous forests. We measured the availabilities of light, soil nitrogen, soil water content, and herbivore abundance in 20 canopy gaps (28.3-607.6 m(2)) formed by a typhoon and in four undisturbed stands. We also measured leaf traits and herbivory on understory plants. 2. The availabilities of light and soil nitrogen increased with increasing gap size. However, soil water content did not. The abundance of herbivorous insects (such as Lepidoptera and Orthoptera) increased with increasing gap size. 3. Concentrations of condensed tannins, total phenolics, and nitrogen in leaves and the leaf mass per area increased in late leaf expansion species with increasing gap size, whereas none of the leaf traits varied by gap size in early leaf expansion species. 4. Herbivory increased on early leaf expansion species with increasing gap size, but decreased on late leaf expansion species. In these late leaf expansion species, total phenolics and C : N ratio had negative relationships with herbivory. 5. These results suggested that after typhoon disturbance, increased herbivory on early leaf expansion species can be explained by increased herbivore abundance, whereas decreased herbivory on late leaf expansion species can be explained by variation in leaf traits.