Nonnative ungulates can alter the structure and function of forest ecosystems. Feral pigs in particular pose a substantial threat to native plant communities throughout their global range. Hawaiian forests are exceptionally vulnerable to feral pig activity because native vegetation evolved in the absence of large mammalian herbivores. A common approach for conserving and restoring forests in Hawaii is fencing and removal of feral pigs. The extent of native plant community recovery and nonnative plant invasion following pig removal, however, is largely unknown. Our objective was to quantify changes in native and nonnative understory vegetation over a 16 yr period in adjacent fenced (pig-free) vs. unfenced (pig-present) Hawaiian montane wet forest. Native and nonnative understory vegetation responded strongly to feral pig removal. Density of native woody plants rooted in mineral soil increased sixfold in pig-free sites over 16 yr, whereas establishment was almost exclusively restricted to epiphytes in pig-present sites. Stem density of young tree ferns increased significantly (51.2%) in pig-free, but not pig-present sites. Herbaceous cover decreased over time in pig-present sites (67.9%). In both treatments, number of species remained constant and native woody plant establishment was limited to commonly occurring species. The nonnative invasive shrub, Psidium cattleianum, responded positively to release from pig disturbance with a fivefold increase in density in pig-free sites. These results suggest that while common native understory plants recover within 16 yr of pig removal, control of nonnative plants and outplanting of rarer native species are necessary components of sustainable conservation and restoration efforts in these forests.
The impact that an exotic species can have on the composition of the community it enters is a function of its abundance, its particular species traits and characteristics of the recipient community. In this study we examined species composition in 14 sites burned in fires fuelled by non‐indigenous C4 grasses in Hawaii Volcanoes National Park, Hawaii. We considered fire intensity, time since fire, climatic zone of site, unburned grass cover, unburned native cover and identity of the most abundant exotic grass in the adjacent unburned site as potential predictor variables of the impact of fire upon native species. We found that climatic zone was the single best variable for explaining variation in native cover among burned sites and between burned and unburned pairs. Fire in the eastern coastal lowlands had a very small effect on native plant cover and often stimulated native species regeneration, whereas fire in the seasonal submontane zone consistently caused a decline in native species cover and almost no species were fire tolerant. The dominant shrub, Styphelia tameiameia, in particular was fire intolerant. The number of years since fire, fire intensity and native cover in reference sites were not significantly correlated with native species cover in burned sites. The particular species of grass that carried the fire did however, have a significant effect on native species recovery. Where the African grass Melinis minutiflora was a dominant or codominant species, fire impacts were more severe than where it was absent regardless of climate zone. Overall, the impacts of exotic grass‐fuelled fires on native species composition and cover in seasonally dry Hawaiian ecosystems was context specific. This specificity is best explained by differences between the climatic zones in which fire occurred. Elevation was the main physical variable that differed among the climatic zones and it alone could explain a large percentage of the variation in native cover among sites. Rainfall, by contrast, did not vary systematically with elevation. Elevation is associated with differences in composition of the native species assemblages. In the coastal lowlands, the native grass Heteropogon contortus, was largely responsible for positive changes in native cover after fire although other native species also increased. Like the exotic grasses, this species is a perennial C4 grass. It is lacking in the submontane zone and there are no comparable native species there and almost all native species in the submontane zone were reduced by fire. The lack of fire tolerant species in the submontane zone thus clearly contributes to the devastating impact of fire upon native cover there.
When invasive woody plants become dominant, they present an extreme challenge for restoration of native plant communities. Invasive Morella faya (fire tree) forms extensive, nearly monospecific stands in wet and mesic forests on the Island of Hawai'i. We used logging, girdling, and selective girdling over time (incremental girdling) to kill stands of M. faya at different rates, with the objective of identifying a method that best promotes native forest re-establishment. We hypothesized that rapid canopy opening by logging would lead to establishment of fast-growing, non-native invaders, but that slower death of M. faya by girdling or incremental girdling would increase the establishment by native plants adapted to partial shade conditions. After applying the M. faya treatments, seed banks, seed rain, and plant recruitment were monitored over 3 years. Different plant communities developed in response to the treatments. Increased light and nitrogen availability in the logged treatment were associated with invasion by non-native species. Native species, including the dominant native forest tree, (Metrosideros polymorpha) and tree fern (Cibotium glaucum), established most frequently in the girdle and incremental girdle treatments, but shortlived non-native species were more abundant than native species. A diverse native forest is unlikely to develop following any of the treatments due to seed limitation for many native species, but girdling and incremental girdling promoted natural establishment of major components of native Hawaiian forest. Girdling may be an effective general strategy for reestablishing native vegetation in areas dominated by woody plant invaders.
Fire tree (Morella faya) has invaded extensive areas of wet and mesic forest on the Island of Hawai'i, forming nearly monospecific stands. Our objective was to identify a method of controlling M. faya, which would allow native plants to establish while minimizing establishment by invasive plants. Treatments (logging all trees, trees left standing but girdled, and incremental girdling over 20 months) were selected to kill M. faya stands at different rates, thereby creating different conditions for species establishment. Leaf litter was either removed or left in place; seeds and seedlings of three native pioneer species, three native forest species, and three alien invasive species were then added to determine their ability to establish. Native pioneer species established best in the log and girdle treatments, whereas seedling emergence of native forest species was higher in the girdle and incremental girdle treatments. Seedlings of invasive species emerged faster than the natives, but each of them responded differently to the stand treatments. Leaf litter reduced seedling emergence for all species, with small-seeded species (<1 mg/seed) affected most under low light conditions. No single method eliminated all invaders, but girdling of M. faya provided suitable conditions for most native species. If combined with selective removal of the most disruptive alien species and native seed additions, girdling could be an effective general strategy for restoring native forests that have been overwhelmed by woody invaders.
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