Effects of invasive rats and burrowing seabirds on seeds and seedlings on New Zealand islands.

Rats (Rattus rattus, Rattus norvegicus, Rattus exulans) are important invaders on islands. They alter vegetation indirectly by preying on burrowing seabirds. These seabirds affect vegetation through nutrient inputs from sea to land and physical disturbance through trampling and burrowing. Rats also directly affect vegetation though consumption of seeds and seedlings. Seedling communities on northern New Zealand islands differ in composition and densities among islands which have never been invaded by rats, are currently invaded by rats, or from which rats have been eradicated. We conducted experimental investigations to determine the mechanisms driving these patterns. When the physical disturbance of seabirds was removed, in soils collected from islands and inside exclosures, seedling densities increased with seabird burrow density. For example, seedling densities inside exclosures were 10 times greater than those outside. Thus the negative effects of seabirds on seedlings, by trampling and uprooting, overwhelm the potentially beneficial effects of high levels of seed germination, seedling emergence, and possibly seed production, which result from seed burial and nutrient additions. Potential seedling density was reduced on an island where rats were present, germination of seeds from soils of this island was approximately half that found on other islands, but on this island seedling density inside exclosures was 7 times the density outside. Although the total negative effects of seabirds and rats on seedling densities are similar (reduced seedling density), the differences in mechanisms and life stages affected result in very different filters on the plant community.

Invasive rats alter woody seedling composition on seabird-dominated islands in New Zealand.

Invasive rats (Rattus rattus, R. norvegicus, R. exulans) have large impacts on island habitats through both direct and indirect effects on plants. Rats affect vegetation by extirpating burrowing seabirds through consumption of eggs, chicks, and adults. These seabirds serve as ecosystem engineers, affecting plant communities by burying and trampling seeds and seedlings, and by altering microclimate. Rats also directly affect plant communities by consuming seeds and seedlings. We studied the direct and indirect impacts of rats on the seedlings of woody plants on 21 islands in northern New Zealand. We compared seedling densities and richness on islands which differed in status with respect to rats: nine islands where rats never invaded, seven islands where rats were present at the time of our study, and five islands where rats were either eradicated or where populations were likely to be small as a result of repeated eradications and re-invasions. In addition, we compared plots from a subset of the 21 islands with different burrow densities to examine the effects of burrowing seabirds on plants while controlling for other factors that differ between islands. We categorized plant communities by species composition and seedling density in a cluster analysis. We found that burrow densities explained more variation in seedling communities than rat status. In areas with high seabird burrow density seedling densities were low, especially for the smallest seedlings. Species richness and diversity of seedlings, but not seedling density, were most influenced by changes in microclimate induced by seabirds. Islands where rats had been eradicated or that had low rat populations had the lowest diversity and richness of seedlings (and adults), but the highest seedling density. Seedling communities on these islands were dominated by Pseudopanax lessonii and Coprosma macrocarpa. This indicates lasting effects of rats that may prevent islands from returning to pre-invasion states.

Above- and below-ground impacts of introduced predators in seabird-dominated island ecosystems.

Predators often exert multi-trophic cascading effects in terrestrial ecosystems. However, how such predation may indirectly impact interactions between above- and below-ground biota is poorly understood, despite the functional importance of these interactions. Comparison of rat-free and rat-invaded offshore islands in New Zealand revealed that predation of seabirds by introduced rats reduced forest soil fertility by disrupting sea-to-land nutrient transport by seabirds, and that fertility reduction in turn led to wide-ranging cascading effects on belowground organisms and the ecosystem processes they drive. Our data further suggest that some effects on the belowground food web were attributable to changes in aboveground plant nutrients and biomass, which were themselves related to reduced soil disturbance and fertility on invaded islands. These results demonstrate that, by disrupting across-ecosystem nutrient subsidies, predators can indirectly induce strong shifts in both above- and below-ground biota via multiple pathways, and in doing so, act as major ecosystem drivers.