Abstract
Ecosystem engineers that modify the soil and ground-layer properties exert a strong influence on vegetation communities in ecosystems worldwide. Understanding the interactions between animal engineers and vegetation is challenging when in the presence of large herbivores, as many vegetation communities are simultaneously affected by both engineering and herbivory. The superb lyrebird Menura novaehollandiae, an ecosystem engineer in wet forests of south-eastern Australia, extensively modifies litter and soil on the forest floor. The aim of this study was to disentangle the impacts of engineering by lyrebirds and herbivory by large mammals on the composition and structure of ground-layer vegetation. We carried out a 2-year, manipulative exclusion experiment in the Central Highlands of Victoria, Australia. We compared three treatments: fenced plots with simulated lyrebird foraging; fenced plots excluding herbivores and lyrebirds; and open controls. This design allowed assessment of the relative impacts of engineering and herbivory on germination rates, seedling density, vegetation cover and structure, and community composition. Engineering by lyrebirds enhanced the germination of seeds in the litter layer. After 2 years, more than double the number of germinants were present in "engineered" than "non-engineered" plots. Engineering did not affect the density of seedlings, but herbivory had strong detrimental effects. Herbivory also reduced the floristic richness and structural complexity (<0.5 m) of forest vegetation, including the cover of herbs. Neither process altered the floristic composition of the vegetation within the 2-year study period. Ecosystem engineering by lyrebirds and herbivory by large mammals both influence the structure of forest-floor vegetation. The twofold increase in seeds stimulated to germinate by engineering may contribute to the evolutionary adaptation of plants by allowing greater phenotypic expression and selection than would otherwise occur. Over long timescales, engineering and herbivory likely combine to maintain a more-open forest floor conducive to ongoing ecosystem engineering by lyrebirds.