Abstract
Biological nitrogen (N) fixation is the primary source of “new” N to unmanaged ecosystems, and recent analyses suggest that terrestrial N inputs via free-living N fixation may be more important than previously assumed. This may be particularly true in some tropical rain forests, where free-living fixation could outpace symbiotic N fixation to represent the dominant source of new N inputs. However, our understanding of the controls over free-living N fixation in tropical rain forests remains poor, which directly constrains our ability to predict how N cycling will respond to changing environmental conditions. Although both phosphorus (P) and molybdenum (Mo) availability have been shown to limit free-living N fixation rates in the tropics, few studies have simultaneously explored P versus Mo limitation or the potential importance of P × Mo interactions. Here, an archived set of foliar, litter, and soil samples from a Costa Rican tropical rain forest provided an opportunity to simultaneously assess the relative strength of P versus Mo relationships with free-living N fixation rates. We also conducted a short-term, full-factorial (P × Mo) litter incubation experiment to directly assess nutrient limitation, allowing us to explore P and Mo controls over free-living N fixation rates using both observational and experimental approaches. We previously showed that N fixation rates were positively correlated with P concentrations in all substrates and, using the archived samples, we now show that Mo concentrations correlated with N fixation only in canopy leaves (where total Mo concentrations were extremely low). Likewise, fertilization with P alone (and not Mo) stimulated leaf litter N fixation rates. Thus, our results suggest that P availability dominantly controls free-living N fixation at this site, and when taken with data from other studies, our results suggest that attempts to identify “the nutrient” that limits N fixation in “the tropics” may be misguided. Rather, nutrient controls over free-living N fixation appear to be more nuanced—and the true nature of nutrient limitation to N fixation likely varies over a variety of scales across the vast tropical rain forest biome.
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Acknowledgments
We thank A. Barron, L. Hedin, W. Bowman, J. Neff, T. Seastedt, D. Nemergut, and S. Schmidt for discussions that helped shape this research and to two anonymous reviewers whose suggestions significantly improved the paper. We thank W. Wieder and A. Vega for help with sample collection; J. Feis and N. Ascarrunz for laboratory assistance; P. Vitousek and B. Houlton for advice on a previous version of the manuscript; John Zobitz and Dan Liptzin for statistical advice; and H. and M. Michaud, F. Campos and the Organization for Tropical Studies (OTS) and Ministerio de Ambiente y Energia (MINAE) in Costa Rica for logistical support. We appreciate the help of J. Drexler, F. Luiszer and the CU Boulder Laboratory for Environmental and Geological Studies for their assistance with molybdenum extraction and analysis. Any use of trade names is for descriptive purposes only and does not imply endorsement by the U.S. Government. This work was supported by grants from the National Science Foundation and the Andrew W. Mellon Foundation to CC, AT, and SR.
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Reed, S.C., Cleveland, C.C. & Townsend, A.R. Relationships among phosphorus, molybdenum and free-living nitrogen fixation in tropical rain forests: results from observational and experimental analyses. Biogeochemistry 114, 135–147 (2013). https://doi.org/10.1007/s10533-013-9835-3
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DOI: https://doi.org/10.1007/s10533-013-9835-3