Funding for this study was provided by a student award from the Boston University Biogeoscience program and from the National Science Foundation (NSF-DDIG DEB-1406521, NSF DEB-1149929, 1406521, 1114804, and 1637685). PS was in part supported by the Watershed Function Scientific Focus Area funded by the U.S. Department of Energy, Office of Science, Office of Biological and Environmental Research under Award Number DE-AC02-05CH11231. JD acknowledges support from the Fundação para Ciência e Tecnologia (IF/00950/2014). Hubbard Brook is a Long-Term Ecological Research site supported with funding from the National Science Foundation. This research is a contribution to the Hubbard Brook Ecosystem Study, administered by the USDA Forest Service, Northeast Forest Experiment Station, Radnor, Pennsylvania.