BRYOCARB : A process-based model of thallose liverwort carbon isotope fractionation in response to CO2, O2, light and temperature

Abstract Evidence from laboratory experiments indicates that fractionation against the heavy stable isotope of carbon (Δ 13 C) by bryophytes (liverworts and mosses) is strongly dependent on atmospheric CO 2 . This physiological response may therefore provide the basis for developing a new terrestrial CO 2 proxy [Fletcher, B.J., Beerling, D.J., Brentnall, S.J., Royer, D.L., 2005. Fossil bryophytes as recorders of ancient CO 2 levels: experimental evidence and a Cretaceous case study. Global Biogeochem. Cycles 19 , GB3012]. Here, we establish a theoretical basis for the proxy by developing an extended model of bryophyte carbon isotope fractionation (BRYOCARB) that integrates the biochemical theory of photosynthetic CO 2 assimilation with controls on CO 2 supply by diffusion from the atmosphere. The BRYOCARB model is evaluated against measurements of the response of liverwort photosynthesis and Δ 13 C to variations in atmospheric O 2 , temperature and irradiance at different CO 2 concentrations. We show that the bryophyte proxy is at least as sensitive to variations in atmosphere CO 2 as the two other leading carbon isotope-based approaches to estimating palaeo-CO 2 levels ( δ 13 C of phytoplankton and of paleosols). Mathematical inversion of BRYOCARB provides a mechanistic means of estimating atmospheric CO 2 levels from fossil bryophyte carbon that can explicitly account for the effects of past differences in O 2 and climate.