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. With increasing awareness of the consequences of climate change for globalecosystems, the focus and application of tree ring research have shifted toreconstruction of long-term climate-related trends in tree growth.Contemporary methods for estimating and removing biological growth trendsfrom tree ring series (standardization) are ill-adapted to shade-tolerantspecies, leading to biases in the resultant chronologies. Further, manymethods, including regional curve standardization (RCS), encountersignificant limitations for species in which accurate age estimation isdifficult. In this study we present and test two tree ring standardizationmodels that integrate tree size in the year of ring formation into theestimation of the biological growth trend. The first method, dubbedsize-deterministic standardization (SDS), uses tree diameter as the solepredictor of the growth trend. The second method includes the combined(COMB) effects of age and diameter. We show that both the SDS and COMBmethods reproduce long-term trends in simulated tree ring data better thanconventional methods; this result is consistent across multiple species.Further, when applied to real tree ring data, the SDS and COMB modelsreproduce long-term, time-related trends as reliably as traditional RCS and more reliably than other common standardization methods (i.e. C-method, basal area increments,conservative detrending). We recommend the inclusion of tree size in theyear of ring formation in future tree ring standardization models,particularly when dealing with shade-tolerant species, as it does notcompromise model accuracy and allows for the inclusion of unaged trees.

Trees do not always act their age: size-deterministic tree ring standardization for long-term trend estimation in shade-tolerant trees