An experimental study of the impacts of understorey forest vegetation and herbivory by red deer and rodents on seedling establishment and species composition in Waitutu Forest, New Zealand

Introduced mammalian herbivores are changing the structure and composition of New Zealand’s forest ecosystems and may modify forest succession after natural disturbances. We studied how introduced ungulates (red deer and feral pigs) and rodents (rats and house mice) affected the rate of recovery (i.e. the engineering resilience) of the forest understorey following artificial disturbance. We imposed disturbances by clearing understorey vegetation dominated by Blechnum ferns in forests on relatively fertile alluvium and elevated infertile marine terraces, and recorded recovery of vegetation (seedling establishment, species composition, cover and volume) in herbivore exclosures and controls. Seedlings quickly established on cleared plots: after 2 years, numbers of woody seedlings and ground cover of vascular plants relative to initial values were similar on cleared and uncleared treatments. Volume of plant biomass <2 m remained low on cleared subplots. Ungulates significantly reduced the re-establishment of woody seedlings ≥ 10 cm tall: only one seedling reached this height outside exclosures, compared with 29 seedlings inside. The number of seedlings <10 cm tall, expressed relative to numbers present pre-clearing, was not significantly affected by ungulates. The species composition of regenerating vegetation was more similar (Jaccard index) to pre-clearing understorey vegetation inside ungulate exclosures than outside. No consistent effect of rodents (primarily house mice) on seedling establishment or species composition was detected after 2 years, and rodent exclosures did not significantly affect survival of seedlings (Griselinia littoralis and Aristotelia serrata) planted as an index of rodent herbivory pressure. No significant differences in vegetation recovery were apparent between forest types. Rapid seedling recruitment in the absence of understorey vegetation and the presence of herbivores provided evidence that understorey vegetation competes with seedlings for light. Ungulate effects were consistent with other experiments that showed herbivores reduced the rate and altered the trajectory of vegetation regrowth after disturbance.