How has the environment shaped geographical patterns of insect body sizes? A test of hypotheses using sphingid moths

Abstract

Aim: We mapped the geographical pattern of body sizes in sphingid moths and investigated latitudinal clines. We tested hypotheses concerning their possible environmental control, that is, effects of temperature (negative: temperature size rule or Bergmann's rule; positive: converse Bergmann rule), food availability, robustness to starvation during extreme weather and seasonality. Location: Old World and Australia/Pacific region. Methods: Body size data of 950 sphingid species were compiled and related to their distribution maps. Focusing on body length, we mapped the median and maximum size of all species occurring in 100 km grid cells. In a comparative approach, we tested the predictions from explanatory hypotheses by correlating species' size to the average environmental conditions encountered throughout their range, under univariate and multivariate models. We accounted for phylogeny by stepwise inclusion of phylogenetically informed taxonomic classifications into hierarchical random‐intercept mixed models. Results: Median body sizes showed a distinctive geographical pattern, with large species in the Middle East and the Asian tropics, and smaller species in temperate regions and the Afrotropics. Absolute latitude explained very little body size variation, but there was a latitudinal cline of maximum size. Species' median size was correlated with net primary productivity, supporting the food availability hypothesis, whereas support for other hypotheses was weak. Environmental correlations contributed much less (i.e. <10%) to explaining overall size variation than phylogeny (inclusion of which led to models explaining >70% of variability). Main conclusion: The intuitive impression of larger species in the tropics is shaped by larger size maxima. Median body sizes are only very weakly related to latitude. Most of the geographical variation in body size in sphingid moths is explained by their phylogenetic past. NPP and forest cover correlate positively with the body size, which supports the idea that food availability allowed the evolution of larger sizes.