Insect herbivory on restored rainforest seedlings weakened by neighbours but unaffected by invasive coffee

Abstract

Restoration of degraded tropical forests is often impeded by invasive species. For regenerating native seedlings, the presence of invasives in the neighbourhood can alter insect herbivory patterns, ultimately shaping restoration trajectories; however, such indirect effects are rarely examined. Here, we investigated the effect of robusta coffee (Coffea canephora) - a shade-tolerant invasive species under closed-canopy secondary forests in the Western Ghats - on the incidence of herbivory (proportion of leaves with any sign of damage) and the extent of leaf damage (percentage leaf area consumed) in seedlings of 10 rainforest species in plots from which coffee plants were either weeded out or left intact. We further examined whether local neighbourhood densities of coffee and other saplings, and species leaf traits, explained herbivory patterns. Removal of invasive coffee did not influence herbivory incidence or the extent of damage across our focal species. However, the incidence of herbivory declined with increasing neighbourhood plant density, suggesting that neighbourhood plants provide a resource dilution effect. Both incidence and extent of herbivore damage were strongly species-specific and partly explained by leaf traits: greater leaf carbon content was correlated with lower herbivory incidence. Contrary to expectations, plants with resource-acquisitive traits (high leaf nitrogen and specific leaf area) experienced lower incidence of herbivory and extent of damage. Our findings suggest that in this system, the indirect effects via herbivores are perhaps not as important in influencing restoration as the more direct effects of invasive species, such as competition for resources.