The impact of herbivores on the natural regeneration of temperate deciduous woodland
Many studies have reported deficient or intermittent patterns of natural regeneration in temperate deciduous woodland. The present study aimed to assess the relative impact of herbivore-mediated plant mortality on the natural regeneration dynamics of representative tree species (Acer pseudoplatanus, Betula pendula, Fraxinus excelsior, Ilex aquifolium, Sorbus aucuparia, Taxus baccata and Ulmus glabra) in representative temperate deciduous woodland (Shipley Wood and Derwent Gorge, County Durham). Sapling density and the density, spatial association and size distribution of adult tree populations varied significantly between tree species and study sites, reflecting contrasting patterns of current and historical regeneration. Rates of post-dispersal seed predation and seedling herbivory were quantified using field-based 'cafeteria' trials. Rodents were the principal agents responsible for seed predation, whereas seedling herbivory was attributable to a mixed suite of herbivores including invertebrates, rodents and larger mammals. Rates of seed predation and seedling herbivory varied significantly between tree species, most likely reflecting individualistic, trade-off responses to chemical and physical attributes. Fine-scale spatial variation most likely reflected the preferential foraging of rodents beneath protective vegetation cover. There was no consistent evidence to support the hypothesis that dispersed seeds may escape disproportionately high offspring mortality beneath parent plants resulting from increased herbivore activity. Natural seedling density varied significantly between years and between tree species, according to the abundance of viable seeds produced by conspecific adult trees. Although seedling survivorship varied significantly between species, the survivorship of each species was similar between years and between cohorts of the same year. Canopy cover, field layer cover or correlated factors were significant determinants of seed germination and seedling emergence, establishment and survival, according to age- and species-specific tolerances. In relative terms, patterns of natural regeneration were primarily herbivore- limited (Acer and Taxus), microsite-limited (Betula, Fraxinus, Ilex and Ulmus) or limited by herbivores and microsites (Sorbus). The availability of viable seeds may have also limited the recruitment of Ilex, Sorbus, Taxus and Ulmus. Vegetative expansion, mast seeding, seed bank regeneration and repeated, prolonged reproduction may have reduced the actual impact of herbivory on natural regeneration, such that long-lived iteroparous tree species were unlikely to have been critically dependent on current recruitment.