The Roots of Diversity: Below Ground Species Richness and Rooting Distributions in a Tropical Forest Revealed by DNA Barcodes and Inverse Modeling

F. Andrew Jones is with the Smithsonian Tropical Research Institute, David L. Erickson is with the Smithsonian Institution, Moises A. Bernal is with the Smithsonian Tropical Research Institute and UT Austin, Eldredge Bermingham is with the Smithsonian Tropical Research Institute, W. John Kress is with the Smithsonian Institution, Edward Allen Herre is with the Smithsonian Tropical Research Institute, Helene C. Muller-Landau is with the Smithsonian Tropical Research Institute, Benjamin L. Turner is with the Smithsonian Tropical Research Institute.Background -- Plants interact with each other, nutrients, and microbial communities in soils through extensive root networks. Understanding these below ground interactions has been difficult in natural systems, particularly those with high plant species diversity where morphological identification of fine roots is difficult. We combine DNA-based root identification with a DNA barcode database and above ground stem locations in a floristically diverse lowland tropical wet forest on Barro Colorado Island, Panama, where all trees and lianas >1 cm diameter have been mapped to investigate richness patterns below ground and model rooting distributions. Methodology/Principal Findings -- DNA barcode loci, particularly the cpDNA locus trnH-psba, can be used to identify fine and small coarse roots to species. We recovered 33 species of roots from 117 fragments sequenced from 12 soil cores. Despite limited sampling, we recovered a high proportion of the known species in the focal hectare, representing approximately 14% of the measured woody plant richness. This high value is emphasized by the fact that we would need to sample on average 13 m2 at the seedling layer and 45 m2 for woody plants >1 cm diameter to obtain the same number of species above ground. Results from inverse models parameterized with the locations and sizes of adults and the species identifications of roots and sampling locations indicates a high potential for distal underground interactions among plants. Conclusions -- DNA barcoding techniques coupled with modeling approaches should be broadly applicable to studying root distributions in any mapped vegetation plot. We discuss the implications of our results and outline how second-generation sequencing technology and environmental sampling can be combined to increase our understanding of how root distributions influence the potential for plant interactions in natural ecosystems.FAJ acknowledges the support of a Tupper postdoctoral fellowship in tropical biology and the National Science Foundation (DEB 0453665). Funding was provided by the Smithsonian Institution Global Earth Observatory, the Smithsonian Tropical Research Institute/Center for Tropical Forest Sciences endowment fund, and the Smithsonian Tropical Research Institute/Frank Levinson fund. We would like to thank Autoridad Nacional del Ambiente and the Smithsonian Tropical Research Institute for processing research permits. We thank S. Hubbell and R. Condit for access to plot data, S. Schnitzer for liana census data (NSF DEB 0613666), and L. Comita and S. Hubbell for access to seedling data (NSF DEB 0075102 and DEB 0823728). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.Marine Scienc