Bryophyte and lichen biomass and nitrogen fixation in a high elevation cloud forest in Cerro de La Muerte, Costa Rica.

Cloud forests have been found to lose more nitrogen in stream discharge than they gain from atmospheric deposition. They also support a large diversity and biomass of tree epiphytes, predominately composed of cryptogams. Since cryptogam epiphytes harbor nitrogen fixing cyanobacteria, they may help make up for the nitrogen loss from ecosystems. We assessed cryptogam biomass on the ground, boles and branches in Quercus costaricensis dominated stands near the tree line in the Cordillera de Talamanca, Costa Rica. Nitrogen fixation was assayed using 15N2 uptake. Total cryptogam biomass was 2 977 kg ha-1, with 67% being found on the lower branches. Bryophytes and chlorolichens made up 53% and 44%, respectively, of the biomass. Half of the bryophyte mass was composed of the liverwort Plagiochila heterophylla, and 66% of the chlorolichen of Lobariella pallida. There were no significant differences in nitrogen fixation rates between the cryptogam species, with a mean rate of 5.04 µg N g-1 day-1 during the predominantly wet condition in the forest. The overall nitrogen input from fixation was 6.1 kg N ha-1 year-1, of which 78% came from bryophytes, 18% from chlorolichens, and 4% from cyanolichens. Only 2.0% of the fixation occurred in cryptogams on the ground, whereas 67%, 24%, and 7% occurred on the lower branches, boles, and upper branches, respectively. These results show that tree epiphytes constitute a significant source of nitrogen for these forests, due to the trees' large surface area, and can make up for the nitrogen lost from these ecosystems.

Nitrogen-Fixing Bacteria Associated with Peltigera Cyanolichens and Cladonia Chlorolichens.

Lichens have been extensively studied and described; however, recent evidence suggests that members of the bacterial community associated with them could contribute new functions to the symbiotic interaction. In this work, we compare the nitrogen-fixing guild associated with bipartite terricolous lichens with different types of photobiont: Peltigera cyanolichens and Cladonia chlorolichens. Since cyanobacteria contribute nitrogen to the symbiosis, we propose that chlorolichens have more diverse bacteria with the ability to fix nitrogen compared to cyanolichens. In addition, since part of these bacteria could be recruited from the substrate where lichens grow, we propose that thalli and substrates share some bacteria in common. The structure of the nitrogen-fixing guild in the lichen and substrate bacterial communities of both lichens was determined by terminal restriction fragment length polymorphism (TRFLP) of the nifH gene. Multivariate analyses showed that the nitrogen-fixing bacteria associated with both types of lichen were distinguishable from those present in their substrates. Likewise, the structure of the nitrogen-fixing bacteria present in the cyanolichens was different from that of chlorolichens. Finally, the diversity of this bacterial guild calculated using the Shannon index confirms the hypothesis that chlorolichens have a higher diversity of nitrogen-fixing bacteria than cyanolichens.