An exact version of Life Table Response Experiment analysis, and the R package exactLTRE

Abstract

Abstract Matrix population models are frequently built and used by ecologists to analyse demography and elucidate the processes driving population growth or decline. Life Table Response Experiments (LTREs) are comparative analyses that decompose the realized difference or variance in population growth rate (λ) into contributions from the differences or variances in the vital rates (i.e. the matrix elements). Since their introduction, LTREs have been based on approximations and have not included biologically relevant interaction terms. We used the functional analysis of variance framework to derive an exact LTRE method, which calculates the exact response of λ to the difference or variance in a given vital rate, for all interactions among vital rates—including higher‐order interactions neglected by the classical methods. We used the publicly available COMADRE and COMPADRE databases to perform a meta‐analysis comparing the results of exact and classical LTRE methods. We analysed 186 and 1487 LTREs for animal and plant matrix population models, respectively. We found that the classical methods often had small errors, but that very high errors were possible. Overall error was related to the difference or variance in the matrices being analysed, consistent with the Taylor series basis of the classical method. Neglected interaction terms accounted for most of the errors in fixed design LTRE, highlighting the importance of two‐way interaction terms. For random design LTRE, errors in the contribution terms present in both classical and exact methods were comparable to errors due to neglected interaction terms. In most examples we analysed, evaluating exact contributions up to three‐way interaction terms was sufficient for interpreting 90% or more of the difference or variance in λ. Relative error, previously used to evaluate the accuracy of classical LTREs, is not a reliable metric of how closely the classical and exact methods agree. Error compensation between estimated contribution terms and neglected contribution terms can lead to low relative error despite faulty biological interpretation. Trade‐offs or negative covariances among matrix elements can lead to high relative error despite accurate biological interpretation. Exact LTRE provides reliable and accurate biological interpretation, and the R package exactLTRE makes the exact method accessible to ecologists.

Keywords

• demography
• life history
• Life Table Response Experiment
• matrix population model
• population growth rate