Shrub interaction with oak seedlings under high herbivory pressure: The role of ontogeny, drought, and plant competition

Abstract

Land use changes occurring during the last decades have caused new challenges in forest management of Mediterranean ecosystems, such as overpopulation of wild ungulates. This is causing serious impacts on forest regeneration, particularly increased seedling mortality of dominant tree species. This study aims to unravel whether oak seedlings could be facilitated by non-palatable shrubs under herbivory and drought stress to improve assisted oak regeneration. For this purpose, we planted one- and two-year-old seedlings of an evergreen oak (Quercus ilex L.) in a Mediterranean open shrubland, considering three factors: herbivore exposure (with or without herbivores), seedling age (one- vs. two-year-old seedlings), and shrub cover (seedlings planted under non-palatable shrubs Cistus ladanifer or in open microsites). We evaluated seedling survival and performance monthly along the growing season (February-September), and measured physiological traits in May, just prior to the drought season. Our results revealed that herbivores, mostly wild boar (Sus scrofa), were the main cause of mortality from February to late May, while drought was responsible of mortality from early June to September. Almost all seedlings died within the first year of plantation after a severe long summer. Leaf predawn water potential and leaf stomatal conductance of one-year-old seedlings were lower under shrubs than in open microsites, a negative effect of shrubs that was not evident in the two-year-old seedlings. Plant photosynthetic performance of two-year old seedlings was higher under shrubs than in open microsites. Interestingly, the net effect of shrubs as facilitators (measured monthly through chlorophyll content) was greater in areas with no herbivores, probably due to the strong difference in herbaceous plant cover between open areas and under shrubs. Our results suggest that facilitation of Q. ilex seedlings by C. ladanifer shrub is weak under herbivory pressure and strongly depends on seedling age as shrubs seem to mostly compete for water with one-year-old seedlings and alleviate photoinhibition in two-year-old seedlings. This study reveals the possible use of two-year-old seedlings under very specific conditions, but some important inconveniences may arise as they are more stressed in the open microsites (ungulate induced mortality was similar for one and two-year old oak seedlings) and require more resources (e.g., time, energy and water) before plantation than one-year-old seedlings. We conclude that the age of plant material must be taken into account when planning oak restoration practices that can potentially benefit from the facilitative effects of shrubs under moderate biotic and abiotic stress.