Climatic and physiological effects on leaf and tree-ring stable isotopes in California redwoods

Variation in the stable isotope composition of organic matter can provide important information about environmental change and biological responses to it. We analyzed the stable carbon (d13C) and oxygen (d18O) isotope ratios of leaves and of the cellulose from individual tree-rings of California's two redwood species to examine how these trees have responded to environmental variation and change in both time and space. Analyses of leaf d13C for both coast redwood (Sequoia sempervirens) and giant sequoia (Sequoiadendron giganteum) from throughout their geographical ranges show a marked gradient with tree height for trees of all sizes and ages but no clear difference among species or populations. The gradient is best explained by tree response to changes in both microenvironment and physiology that are known to change with height. In contrast, leaf d18O for both species showed no clear relationship with height but very clear differences between species and populations with giant sequoia displaying a much stronger inferred leaf-level response to the higher evaporative conditions present in the Sierra Nevada mountains as compared to the coast. Both species showed population-level differences with the driest and warmest sites most distinct from all of the others. Intra-annual analyses of d13C and d18O in tree-rings over a 21-year period (1974-1994) were also used to explore how climate and tree response to climate was recorded for both species. These analyses revealed unique (local) climatic effects and response to the climate for each species and population of both redwood species. Most pronounced was a significant increase in intrinsic Water Use Efficiency (iWUE) derived from d13C data over the study period in both species, and a distinct d18O response in relation to drought (e.g. 1976/1977) and to warmer days and nights and above-average precipitation (e.g., 1982-1985). Patterns of co-variation in d13C and d18O in both species suggest that during dry and also warm periods these trees appear to first down-regulate their water use and secondly their carbon fixation and that high evaporative conditions drive some of the most marked changes in both variables. This information should be useful for efforts to conserve and protect both redwood species under novel environmental conditions expected in the coming decades.