Changes in asymbiotic, heterotrophic nitrogen fixation on leaf litter of Metrosideros polymorpha with long-term ecosystem development in Hawaii

We measured nitrogenase activity (acetylene reduction) of asymbiotic, heterotrophic, nitrogen-fixing bacteria on leaf litter from the tree Metrosideros polymorpha collected from six sites on the Hawaiian archipelago. At all sites M. polymorpha was the dominant tree, and its litter was the most abundant on the forest floor. The sites spanned a soil chronosequence of 300 to 4.1 million y. We estimated potential nitrogen fixation associated with this leaf litter to be highest at the youngest site (1.25 kg ha(-1) y(-1)), declining to between 0.05 and 0.22 kg ha(-1) y(-1) at the oldest four sites on the chronosequence. To investigate how the availability of weathered elements influences N fixation rates at different stages of soil development, we sampled M. polymorpha leaf litter from complete, factorial fertilization experiments located at the 300-y, 20,000-y and 4.1 million-y sites. At the youngest and oldest sites, nitrogenase activity on leaf litter increased significantly in the plots fertilized with phosphorus and "total" (all nutrients except N and P); no significant increases in nitrogenase activity were measured in leaf litter from treatments at the middle-aged site. The results suggest that the highest rates of N fixation are sustained during the "building" or early phase of ecosystem development when N is accumulating and inputs of geologically cycled (lithophilic) nutrients from weathering are substantial.