Abstract

Geographical body size variation has long interested evolutionary biologists, and a range of mechanisms have been proposed to explain the observed patterns. It is considered to be more puzzling in ectotherms than in endotherms, and integrative approaches are necessary for testing non-exclusive alternative mechanisms. Using lacertid lizards as a model, we adopted an integrative approach, testing different hypotheses for both sexes while incorporating temporal, spatial, and phylogenetic autocorrelation at the individual level. We used data on the Spanish Sand Racer species group from a field survey to disentangle different sources of body size variation through environmental and individual genetic data, while accounting for temporal and spatial autocorrelation. A variation partitioning method was applied to separate independent and shared components of ecology and phylogeny, and estimated their significance. Then, we fed-back our models by controlling for relevant independent components. The pattern was consistent with the geographical Bergmann's cline and the experimental temperature-size rule: adults were larger at lower temperatures (and/or higher elevations). This result was confirmed with additional multi-year independent data-set derived from the literature. Variation partitioning showed no sex differences in phylogenetic inertia but showed sex differences in the independent component of ecology; primarily due to growth differences. Interestingly, only after controlling for independent components did primary productivity also emerge as an important predictor explaining size variation in both sexes. This study highlights the importance of integrating individual-based genetic information, relevant ecological parameters, and temporal and spatial autocorrelation in sex-specific models to detect potentially important hidden effects. Our individual-based approach devoted to extract and control for independent components was useful to reveal hidden effects linked with alternative non-exclusive hypothesis, such as those of primary productivity. Also, including measurement date allowed disentangling and controlling for short-term temporal autocorrelation reflecting sex-specific growth plasticity.

Highlights

  • Spatial and temporal variation in phenotypic traits is a central issue in ecology and evolution

  • After separating the shared and independent components of phylogeny and ecology, we found a significant effect of phylogeny

  • The effect of the independent component of ecology was much stronger in females than in males because females were much more influenced by the course of the season

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Summary

Introduction

Spatial and temporal variation in phenotypic traits is a central issue in ecology and evolution. Geographical variation of body size along environmental gradients has been studied for more than 160 years [1]. This topic is currently the focus of intense interest, including in the context of the response of organisms to environmental change [2], and a major challenge is disentangling among (phylo)genetic and ecological factors [3, 4]. A positive association of body size with latitude and/or elevation, or a negative association with temperature, the so-called Bergmann’s rule [1], has been frequently observed An adaptive thermoregulatory explanation for this pattern was originally proposed by Bergmann [1] for which the areavolume ratio of larger sizes facilitates heat conservation in colder environments. It has been suggested that other nonexclusive mechanisms can be involved and even confounded with thermoregulatory mechanisms ([5,6,7] and references therein)

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