Abstract

BackgroundSkippers (Family: Hesperiidae) are a large group of butterflies with ca. 4000 species under 567 genera. The lack of a time-calibrated higher-level phylogeny of the group has precluded understanding of its evolutionary past. We here use a 10-gene dataset to reconstruct the most comprehensive time-calibrated phylogeny of the group, and explore factors that affected the diversification of these butterflies.ResultsAncestral state reconstructions show that the early hesperiid lineages utilized dicots as larval hostplants. The ability to feed on monocots evolved once at the K-Pg boundary (ca. 65 million years ago (Mya)), and allowed monocot-feeders to diversify much faster on average than dicot-feeders. The increased diversification rate of the monocot-feeding clade is specifically attributed to rate shifts in two of its descendant lineages. The first rate shift, a four-fold increase compared to background rates, happened ca. 50 Mya, soon after the Paleocene-Eocene thermal maximum, in a lineage of the subfamily Hesperiinae that mostly fed on forest monocots. The second rate shift happened ca. 40 Mya in a grass-feeding lineage of Hesperiinae when open-habitat grasslands appeared in the Neotropics owing to gradual cooling of the atmospheric temperature.ConclusionsThe evolution of monocot feeding strongly influenced diversification of skippers. We hypothesize that although monocot feeding was an intrinsic trait that allowed exploration of novel niches, the lack of extensive availability of monocots comprised an extrinsic limitation for niche exploration. The shifts in diversification rate coincided with paleoclimatic events during which grasses and forest monocots were diversified.

Highlights

  • Skippers (Family: Hesperiidae) are a large group of butterflies with ca. 4000 species under 567 genera

  • To test the hypothesis that the rate shifts have occurred in response to ecological opportunity, we investigate whether (i) rate shifts are associated with feeding preferences of the lineages, (ii) the K-Pg extinction had an impact on diversification rate, (iii) the diversification pattern reflects the biogeographic distribution of lineages, and (iv) paleoclimate has influenced diversification rate

  • Diversification rates and pattern The Maximum Clade Credibility (MCC) tree indicates that Hesperiidae began diversifying in the late Cretaceous ca. 82 Mya, when Coeliadinae diverged from the rest of the family

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Summary

Introduction

Skippers (Family: Hesperiidae) are a large group of butterflies with ca. 4000 species under 567 genera. The lack of a time-calibrated higher-level phylogeny of the group has precluded understanding of its evolutionary past. We here use a 10-gene dataset to reconstruct the most comprehensive time-calibrated phylogeny of the group, and explore factors that affected the diversification of these butterflies. The remarkable diversity of life is often attributed to either the gradual accumulation of species over long periods of time [1, 2] or to dramatic changes in diversification rates across lineages and time [3, 4]. Paleontological evidence [3] and phylogenetic comparisons [4] across the tree of life predict the latter scenario as the major cause, where the fluctuations in speciation and extinction rates explain the historical pattern of lineage accumulation. Herbivorous insects comprise about half of all terrestrial eukaryotic species [13], exemplifying the importance of insect-plant interactions in

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