Abstract
Widely known for pest species that include major modulators of temperateforests, the genus Choristoneura is part of the species-rich tribe Archipini of leafrollermoths (Tortricidae). Delimitation of the genus has remained unresolved because nophylogeny has included species endemic to Africa and studies have often omitted thetype species of the genus. Further taxonomic confusion has been generated by thetransfer of Archips occidentalis (Walsingham) to Choristoneura, creating a homonymwith Choristoneura occidentalis Freeman, an important defoliator of North Americanforests. To define the limits of the genus, we reconstructed a phylogeny using DNAsequences for mitochondrial cytochrome oxidase subunit I and nuclear ribosomal 28Sgenes. Our ingroup included 23 Choristoneura species-level taxa, complemented by alarge sample of outgroups comprising 82 species of Archipini and other Tortricidae.We generated a time-calibrated tree using fossil and secondary calibrations and weinferred biogeographic and diversification processes in Choristoneura. Our analysisrecovered the genus as polyphyletic, with Archips occidentalis, Choristoneura simonyiand Choristoneura evanidana excluded from the main clade. Based on the recoveredphylogenies and a redefinition, we restrict Choristoneura primarily to species with anorthern hemisphere distribution. Our analysis supports A. occidentalis as the sistergroup of Cacoecimorpha pronubana, C. simonyi as the sister of ‘Xenotemna’ pallorana,and C. evanidana as the sister of Archips purpurana. A new combination is proposed:Archips evanidana comb.n.; the availability of ‘Xenotemna’ as a valid name is discussedand A. occidentalis is considered as an orphaned name within the Archipini. Wefound support for a Holarctic origin of Choristoneura about 23 Ma, followed by earlydivergence in the Palearctic region. The main divergence occurred at 16Ma, with oneclade in the Nearctic and another in the Palearctic. Subsequent cladogenetic eventswere synchronous and related to herbivorous specialization, with each clade dividedinto coniferophagous and polyphagous lineages. Their specialization as conifer feederstemporally matched the expansion of boreal forest during the Miocene.