Explaining global insect species richness:lessons from a decade of macroevolutionary entomology

The last ten years have seen more research on insect macroevolution than all the previous years combined. Here I summarize and criticise the claims that have been made by comparative phylogenetic and fossil studies, and identify some future opportunities. We know the fossil record and phylogeny of insects much better than we did ten years ago. We cannot simply ascribe the richness of insects, or their subtaxa, to either age or diversification rate. There is evidence that fossil family richness peaked much earlier than previously suspected. Phylogenetic evidence however suggests that species-level net diversification rates are accelerating, though this is highly variable across taxa, implying ongoing changes in global taxonomic composition. Although there is evidence that wings and metamorphosis have had some macroevolutionary effects, the most definitive broad phylogenetic study does not suggest that they directly elevated net diversification of species. There is little evidence that insect body size influences net diversification rate. Compared to other phyla, arthropod richness, of which insects comprise the major part, is best explained by non-marine habit, presence of parasitic lifestyles, a skeleton, vision and dioecy. Herbivory cannot yet robustly be said to increase diversification over other diets across all insects: there are contrary analyses, and effects differ in different taxa. Many phylogenetic studies now document how it sometimes does: from co-speciation, to diffuse coevolution with host shifting. The last decade has shown that climate change and biogeographic processes are likely important in generating or limiting insect diversification, but there is a need for greater statistical rigour in such studies. There is also a need to understand the validity of some widely used statistical methods better, and to make better use of the data and methods that exist. Macroevolutionary entomology could greatly benefit from online data integration platforms to facilitate analyses of broader scope.