Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms

Abstract

Tectonic dynamics and niche availability play intertwined roles in determining patterns of diversification. Such drivers explain the current distribution of many clades, whereas events such as the rise of angiosperms can have more specific impacts, such as on the diversification rates of herbivores. The Tortricidae, a diverse group of phytophagous moths, are ideal for testing the effects of these determinants on the diversification of herbivorous clades. To estimate ancestral areas and diversification patterns in Tortricidae, a complete tribal‐level dated tree was inferred using molecular markers (one mitochondrial and five nuclear) and calibrated using fossil constraints. We found that Tortricidae diverged from their sister group c. 120 Myr ago (Ma) and diversified c. 97 Ma, a timeframe synchronous with the rise of angiosperms in the Early–mid Cretaceous. Ancestral areas analysis, based on updated Wallace's biogeographical regions, supports the hypothesis of a Gondwanan origin of Tortricidae in the South American plate. We also detected an increase in speciation rate that coincided with the peak of angiosperm diversification in the Cretaceous. This in turn probably was further heightened by continental colonization of the Palaeotropics when angiosperms became dominant by the end of the Late Cretaceous.

Re-examining the rare and the lost: a review of fossil Tortricidae (Lepidoptera)

“…The monophyly of Chlidanotinae has been challenged, and two recent studies based on molecular data suggest that the subfamily is paraphyletic (Regier et al 2012;Fagua et al 2017). Although the subfamily of this fossil cannot be determined for certain, we conclude that it most likely is a tortricid.…”

“…Although the monophyly of Tortricidae is widely accepted and strongly supported by molecular and morphological data (Regier et al 2012;Heikkilä et al 2015;Fagua et al 2017), very few morphological characters have been recognized as diagnostic of this clade. Putative synapomorphies include one female character and three larval characters (Powell & Brown 2012): flat papillae anales facing ventrally in the female genitalia; presence of the D2 seta on a shared dorsal pinaculum on A9 in the larva; the presence of a morphologically distinct anal fork (secondarily lost in some internal feeders) in the larva; and the anterior, slightly dorsoanterior or ventroanterior position of the SD1 seta with regard to the spiracle on A8 in the larva.…”

“…Regier et al (2012) confirmed that Chlidanotinae (as monophyletic or paraphyletic) are the most basal tortricid lineage and recognized the division of the subfamily into three tribes: Polyorthini plus a monophyletic Hilarographini + Chlidanotini, with Polyorthini as the first lineage to diverge. Fagua et al (2017) also recovered a paraphyletic Chlidanotinae as the basal-most lineages of Tortricidae. Poinar and Brown (1993) exclude the moth from Chlidanotini and Hilarographini because of differences in the morphology diagnostic of these tribes (e.g., members of these tribes lack raised scales of the forewing and most have conspicuously thickened antennae).…”

“…comm.). Subsequently, in the divergence time analysis by Fagua et al (2017) this fossil was used to give a minimum age to Olethreutinae. To support this placement, Fagua et al (2017) mention "the presence of forewing veins M-stem and stem of R4+5".…”

“…Subsequently, in the divergence time analysis by Fagua et al (2017) this fossil was used to give a minimum age to Olethreutinae. To support this placement, Fagua et al (2017) mention "the presence of forewing veins M-stem and stem of R4+5". Although these authors acknowledge that this is a plesiomorphic trait among extant Tortricidae, they mention that it is found almost exclusively in Olethreutinae.…”