Isotopic characteristics of canopies in simulated leaf assemblages

The geologic history of closed-canopy forests is of great interest to paleoecologists and paleoclimatologists alike. Closed canopies have pronounced effects on local, continental and global rainfall and temperature patterns. Although evidence for canopy closure is difficult to reconstruct from the fossil record, the characteristic isotope gradients of the canopy effect could be preserved in leaves and proxy biomarkers. To assess this, we employed new carbon isotopic data for leaves collected in diverse light environments within a deciduous, temperate forest (Maryland, USA) and for leaves from a perennially closed canopy, moist tropical forest (Bosque Protector San Lorenzo, Panama). In the tropical forest, leaf carbon isotope values range 10 parts per thousand, with higher delta C-13(leaf) values occurring both in upper reaches of the canopy, and with higher light exposure and lower humidity. Leaf fractionation (Delta(leaf)) varied negatively with height and light and positively with humidity. Vertical C-13 enrichment in leaves largely reflects changes in Delta(leaf), and does not trend with delta C-13 of CO2 within the canopy. At the site in Maryland, leaves express a more modest delta C-13 range (similar to 6 parts per thousand), with a clear trend that follows both light and leaf height. Using a model we simulate leaf assemblage isotope patterns from canopy data binned by elevation. The re-sampling (bootstrap) model determined both the mean and range of carbon isotope values for simulated leaf assemblages ranging in size from 10 to over 1000 leaves. For the tropical forest data, the canopy's isotope range is captured with 50 or more randomly sampled leaves. Thus, with a sufficient number of fossil leaves it is possible to distinguish isotopic gradients in an ancient closed canopy forest from those in an open forest. For very large leaf assemblages, mean isotopic values approximate the delta C-13 of carbon contributed by leaves to soil and are similar to observed delta C-13(litter) values at forested sites within Panama, including the site where leaves were sampled. The model predicts a persistent similar to 1 parts per thousand difference in delta C-13(litter) for the two sites which is consistent with higher water availability in the tropical forests. This work provides a new framework for linking contemporary ecological observations to the geochemical record using flux-weighted isotope data and lends insights to the effect of forest architecture on organic and isotopic records of ancient terrestrial ecosystems. (C) 2014 Elsevier Ltd. All rights reserved.