Abstract

As phylogenetically controlled experimental designs become increasingly common in ecology, the need arises for a standardized statistical treatment of these datasets. Phylogenetically paired designs circumvent the need for resolved phylogenies and have been used to compare species groups, particularly in the areas of invasion biology and adaptation. Despite the widespread use of this approach, the statistical analysis of paired designs has not been critically evaluated. We propose a mixed model approach that includes random effects for pair and species. These random effects introduce a “two-layer” compound symmetry variance structure that captures both the correlations between observations on related species within a pair as well as the correlations between the repeated measurements within species. We conducted a simulation study to assess the effect of model misspecification on Type I and II error rates. We also provide an illustrative example with data containing taxonomically similar species and several outcome variables of interest. We found that a mixed model with species and pair as random effects performed better in these phylogenetically explicit simulations than two commonly used reference models (no or single random effect) by optimizing Type I error rates and power. The proposed mixed model produces acceptable Type I and II error rates despite the absence of a phylogenetic tree. This design can be generalized to a variety of datasets to analyze repeated measurements in clusters of related subjects/species.

Highlights

  • In the last decade, the number of phylogenetically controlled experimental designs and statistical analyses has increased dramatically in the field of ecology (e.g., Agrawal et al 2005; Funk and Vitousek 2007; Heard and Sax 2013)

  • When conventional statistics are applied to comparative data, the overestimate of independent observations leads to inflated Type I error rates

  • We found that the two reference models, Model 1 and Model 2, had inflated Type I error rates (Fig. 1)

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Summary

Introduction

The number of phylogenetically controlled experimental designs and statistical analyses has increased dramatically in the field of ecology (e.g., Agrawal et al 2005; Funk and Vitousek 2007; Heard and Sax 2013). Controlling for phylogenetic relatedness among a suite of species is important because similarities in traits or responses among species may result from biological and ecological factors or may be strongly affected by shared evolutionary history. Adding phylogenetic information into analyses or experimental designs is essential to tease apart the influence of ecological and evolutionary factors on our trait or response of interest. The leaf economics spectrum identifies strong correlations among leaf metabolic processes and structure across a broad taxonomic range of species resulting from biophysical constraints on leaves (Reich et al 1997). Using a phylogenetically controlled analysis, Ackerly and Reich (1999) found that, overall, these correlations were not an artifact of shared evolutionary history at either end of the leaf economics spectrum (e.g., thick-leaved species with low photosynthetic rates occur in closely related genera). The correlation between leaf life span and leaf area was driven by large differences in these traits between angiosperms and conifers – there was very little variation in these traits within these plant groups (Ackerly and Reich 1999)

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