Patterns of secondary invasion in the understory of exotic, invasive timber stands

Current climate and land cover change threaten global mountaintops with increased spread of invasive species. Long-established plantations of exotic and invasive trees on these mountaintops can alter their surroundings, further increasing invader-facilitated or secondary invasion. Identifying the ecological conditions that promote such specific associations can help develop better management interventions. The Western Ghatss Shola Sky Islands (>1400m MSL) host vast stretches of exotic and invasive tree plantations that sustain colonisation of other invasive woody, herbaceous and fern species in their understories. Here we analysed vegetation and landscape variables from 232 systematically-placed plots in randomly selected grids using NMDS and Phi Coefficient approaches, to examine patterns of association (positive interactions) between secondary understory invasive species with specific exotic and invasive overstory species. We also conducted GLMM with zero inflation to determine the influence of environmental variables where such associations occur. We find that secondary invasion of multiple species under the canopy of other exotic invasives is widespread across the Shola Sky Islands. Stands of Eucalyptus host the colonisation of 70% of non-native invasive species surveyed across the Shola Sky Islands. In particular, Lantana camara invasion is strongly associated with Eucalyptus stands. We also found that climatic variables affect the colonisation of understorey woody invasive species, while invasion by exotic herbaceous species is associated with the density of road networks.. Canopy cover impacts all invasives negatively, while incidence of fire was negatively associated with invasion by Lantana spp and the Pteridium spp. While the restoration of natural habitats largely targets the highly invasive Acacia, less invasive Eucalyptus and Pine are often not included. Our study suggests that retaining such exotic species in natural habitats, particularly protected areas, can hinder ongoing restoration efforts by facilitating further invasions by multiple woody and herbaceous species.