Restoring ecological function in temperate grassy woodlands: manipulating soil nutrients, exotic annuals and native perennial grasses through carbon supplements and spring burns

1. Ecological invasions are often associated with persistent changes to underlying ecological processes. Restoration of invaded communities is dependent on manipulation of these processes to favour the target species composition and impart resistance to further invasion. We applied these principles to extensively degraded grassy woodlands in temperate agricultural regions of Australia, where widespread invasion by mediterranean annuals is related to altered ecological processes such as soil nutrient cycling. 2. We investigated carbon supplementation and spring burns, in association with re-establishment of native perennial grasses, as potential management tools for manipulating nitrogen cycling, soil seed banks and establishment conditions in degraded woodland understoreys. Through these mechanisms we aimed to enhance native cover and increase resistance to invasion by exotic annuals. 3. In two contrasting degraded remnants, repeated sucrose applications temporarily reduced soil nitrate to inferred pre-European levels, which dramatically reduced growth of exotic annuals and enhanced native perennial abundance. Repeated spring burns did not reduce soil nitrate but reduced exotic annual grasses through effects on soil seed banks and/or establishment conditions. 4. Spring burns and carbon supplements both significantly enhanced establishment of Themeda australis, a dominant tussock grass prior to European settlement. Combinations of T. australis seed addition and either spring burning or carbon supplements enhanced native plant abundance more effectively than treatments without seed addition. 5. Within 18 months of their establishment, T. australis seedlings significantly reduced soil nitrate in some treatments, providing a preliminary indication that re-establishment of a dense sward of this species may restore ecosystem function to a low-nitrate state that favours native perennials over exotic annuals. 6. Synthesis and applications. Ecological restoration can be viewed as targeted intervention in species-environment interactions, whereby ecological conditions are manipulated to enhance establishment or vigour of key species, and these species in turn help restore ecological processes that favour the target species composition. In grassy ecosystems re-establishing a perennial sward of appropriate native tussock grasses may be critical for restoring pre-disturbance nitrogen cycles and improving resistance to invasion by exotic annuals. Carbon supplements and spring burns facilitate this process through complementary mechanisms.