Leaf aging of Amazonian canopy trees as revealed by spectral and physiochemical measurements.

Leaf aging is a fundamental driver of changes in leaf traits, thereby regulating ecosystem processes and remotely sensed canopy dynamics. We explore leaf reflectance as a tool to monitor leaf age and develop a spectra-based partial least squares regression (PLSR) model to predict age using data from a phenological study of 1099 leaves from 12 lowland Amazonian canopy trees in southern Peru. Results demonstrated monotonic decreases in leaf water (LWC) and phosphorus (P ) contents and an increase in leaf mass per unit area (LMA) with age across trees; leaf nitrogen (N ) and carbon (C ) contents showed monotonic but tree-specific age responses. We observed large age-related variation in leaf spectra across trees. A spectra-based model was more accurate in predicting leaf age (R  = 0.86; percent root mean square error (%RMSE) = 33) compared with trait-based models using single (R  = 0.07-0.73; %RMSE = 7-38) and multiple (R  = 0.76; %RMSE = 28) predictors. Spectra- and trait-based models established a physiochemical basis for the spectral age model. Vegetation indices (VIs) including the normalized difference vegetation index (NDVI), enhanced vegetation index 2 (EVI2), normalized difference water index (NDWI) and photosynthetic reflectance index (PRI) were all age-dependent. This study highlights the importance of leaf age as a mediator of leaf traits, provides evidence of age-related leaf reflectance changes that have important impacts on VIs used to monitor canopy dynamics and productivity and proposes a new approach to predicting and monitoring leaf age with important implications for remote sensing.