Origin of the Hawaiian rainforest ecosystem and its evolution in long-term primary succession.

Born among volcanoes in the north central Pacific about 4 million years ago, the Hawaiian rainforest became assembled from spores of algae, fungi, lichens, bryophytes, ferns and from seeds of about 275 flowering plants that over the millenia evolved into ca. 1000 endemic species. Outstanding among the forest builders were the tree ferns (Cibotium spp.) and the 'Ōhi'a lehua trees (Metrosideros spp.), which still dominate the Hawaiian rainforest ecosystem today. The structure of this forest is simple. The canopy in closed mature rainforests is dominated by cohorts of Metrosideros polymorpha and the undergrowth by tree fern species of Cibotium. When a new lava flow cuts through this forest, kipuka are formed, i.e. islands of remnant vegetation. On the new volcanic substrate, the assemblage of plant life-forms is similar as during the evolution of this system. In open juvenile forests, a mat-forming fern, the uluhe fern (Dicranopteris lineraris) becomes established. It inhibits further regeneration of the dominant 'Ōhi'a tree, thereby reinforcing the cohort structure of the canopy guild. In the later part of its life cycle, the canopy guild breaks down often in synchrony. The trigger is hypothesized to be a climatic perturbation. After that disturbance the forest becomes reestablished in about 30-40 yr. As the volcanic surfaces age, they go from a mesotrophic to a eutrophic phase, reaching a biophilic nutrient climax by about 1-25 K yr. Thereafter, a regressive oligotrophic phase follows; the soils become exhausted of nutrients. The shield volcanoes break down. Marginally, forest habitats change into bogs and stream ecosystems. The broader 'Ōhi'a rainforest redeveloping in the more dissected landscapes of the older islands looses stature, often forming large gaps that are invaded by the aluminum tolerant uluhe fern. The 'Ōhi'a trees still thrive on soils rejuvenated from landslides and from Asian dust on the oldest (5 million year old) island Kaua'i but their stature and living biomass is greatly diminished. [ABSTRACT FROM AUTHOR]