Phylogeny, classification and divergence times of pygmy leafmining moths (Lepidoptera: Nepticulidae): the earliest lepidopteran radiation on Angiosperms?

Publication Type:Journal Article
Year of Publication:2016
Authors:C. Doorenweerd, van Nieukerken, E. J., Hoare, R. J. B.
Journal:Systematic Entomology
Date Published:2016-10-31
Keywords:Acalyptris, Acalyptris platani group, Acalyptris psammophricta group, Acalyptris scirpi group, Acalyptris staticis group, Areticulata, Bohemannia, Bohemannia pulverosella, Casanovula, CRETACEOUS, Ectoedemia, Ectoedemia angulifasciella, Ectoedemia angulifasciella group, Ectoedemia commiphorella group, Ectoedemia occultella, Ectoedemia populella group, Ectoedemia quadrinotata, Ectoedemia subbimaculella group, Enteucha, Enteucha basidactyla, Enteucha gilvafascia, Etainia, Etainia obtusa, Fomoria, Fomoria groschkei group, Fomoria vannifera, Fomoria vannifera group, Fomoria viridissimella, Fomoria weaveri group, glaucolepis, Glaucolepis argentosa, Glaucolepis headleyella group, Glaucolepis oishiella, Glaucolepis raikhonae, Glaucolepis raikhonae group, Glaucolepis saccharella, Glaucolepis saccharella group, Hesperolyra, Leafmines, Levarchama, Menurella, Menurella xenadelpha, Molecular clock, Molecular phylogenetics, Morphology, Muhabbetana, Muhabbetana grandinosa, Neotrifurcula, Nepticulidae, Opostega, Opostega salaciella, Opostegidae, Opostegoides, Ozadelpha, Ozadelpha conostegiae, Ozadelpha EvN4680, Ozadelpha guajavae, Parafomoria, Pectinivalva, phylogeny, Pseudopostega, Pygmy moths, Roscidotoga, Simplimorpha, Simplimorpha lanceifoliella, Simplimorpha promissa, Stigmella, Stigmella anomalella group, Stigmella aurella group, Stigmella betulicola group, Stigmella epicosma group, Stigmella floslactella group, Stigmella hybnerella group, Stigmella intronia, Stigmella irregularis group, Stigmella lapponica group, Stigmella lemniscella group, Stigmella malella, Stigmella naturnella, Stigmella ogygia group, Stigmella oxyacanthella group, Stigmella paliurella group, Stigmella paradoxa group, Stigmella prunifoliella group, Stigmella quercipulchella group, Stigmella ruficapitella group, Stigmella saginella group, Stigmella salicis group, Stigmella sanguisorbae group, Stigmella slingerlandella, Stigmella tricentra, Stigmella ulmivora group, Stigmella ultima group, Trifurcula, Trifurcula cryptella group, Trifurcula pallidella group, Trifurcula subnitidella group, Varius, Zimmermannia

Nepticulidae is one of the early diverging Lepidoptera lineages, and currently comprises over 850 described species. The larvae of the vast majority of the species are leafminers on Angiosperms and highly monophagous, which has led to persistent ideas on coevolution with their plant hosts. We here present a molecular phylogeny based on eight gene fragments from 355 species, representing 20 out of 22 extant Nepticulidae genera. Using two fossil calibration points, we performed molecular dating to place the origin of the family in the Early Cretaceous, before the main Angiosperm diversification. Based on our results we propose a new classification, abandoning all ranks between family and genus, as well as subgenera to allow for a stable classification. The position of Enteucha within Nepticulidae remains somewhat ambiguous, and the species rich cosmopolitan genus Stigmella, with nearly half of all described Nepticulidae, requires further study. Ectoedemia, Zimmermannia, Acalyptris, Etainia, Parafomoria, Muhabbetana and Fomoria appear to have diversified in a relatively short evolutionary period, leading to short branches in the molecular phylogeny and unclear suprageneric relations. Otherwise supports throughout the phylogeny are mostly high and the species groups, genera and higher clades are discussed in respect to their supporting morphological and life history characters. Wing venation characters are confirmed to mostly be reliable and relevant for classification, but some other characters previously used require re-interpretation. The species groups of most genera are recovered, but only partly so in the large genus Stigmella. The molecular dating results are compared with existing knowledge on the timing of the Angiosperm radiation and reveal that the diversification of Nepticulidae could largely have been contemporaneous with their hosts, although some of the genera restricted to a single plant family appear to have begun to diversify before their hosts. This is the first published update on the full classification of Nepticulidae in over 15 years, and the first time that more than two Nepticulidae taxa are included in molecular dating estimates.

Short Title:Syst. Ent.
Taxonomic name: 
Acalyptris (Nepticuloidea), Acalyptris platani group (Nepticuloidea), Acalyptris scirpi group (Nepticuloidea), Acalyptris staticis group (Nepticuloidea), Areticulata (Nepticuloidea), Bohemannia (Nepticuloidea), Bohemannia pulverosella (Nepticuloidea), Casanovula (Nepticuloidea), Ectoedemia (Nepticuloidea), Ectoedemia angulifasciella (Nepticuloidea), Ectoedemia angulifasciella group (Nepticuloidea), Ectoedemia commiphorella group (Nepticuloidea), Ectoedemia occultella (Nepticuloidea), Ectoedemia populella group (Nepticuloidea), Ectoedemia quadrinotata (Nepticuloidea), Ectoedemia subbimaculella group (Nepticuloidea), Enteucha (Nepticuloidea), Enteucha basidactyla (Nepticuloidea), Enteucha gilvafascia (Nepticuloidea), Etainia (Nepticuloidea), Etainia obtusa (Nepticuloidea), Fomoria (Nepticuloidea), Fomoria groschkei group (Nepticuloidea), Fomoria vannifera (Nepticuloidea), Fomoria vannifera group (Nepticuloidea), Fomoria viridissimella (Nepticuloidea), Fomoria weaveri group (Nepticuloidea), Glaucolepis (Nepticuloidea), Glaucolepis argentosa (Nepticuloidea), Glaucolepis headleyella group (Nepticuloidea), Glaucolepis oishiella (Nepticuloidea), Glaucolepis raikhonae (Nepticuloidea), Glaucolepis raikhonae group (Nepticuloidea), Glaucolepis saccharella (Nepticuloidea), Glaucolepis saccharella group (Nepticuloidea), Hesperolyra (Nepticuloidea), Levarchama (Nepticuloidea), Menurella (Nepticuloidea), Menurella xenadelpha (Nepticuloidea), Muhabbetana (Nepticuloidea), Muhabbetana grandinosa (Nepticuloidea), Neotrifurcula (Nepticuloidea), Nepticulidae (Nepticuloidea), Opostega (Nepticuloidea), Opostega salaciella (Nepticuloidea), Opostegidae (Nepticuloidea), Opostegoides (Nepticuloidea), Ozadelpha (Nepticuloidea), Ozadelpha conostegiae (Nepticuloidea), Ozadelpha EvN4680 (Nepticuloidea), Ozadelpha guajavae (Nepticuloidea), Parafomoria (Nepticuloidea), Parafomoria (Nepticuloidea), Pectinivalva (Nepticuloidea), Pseudopostega (Nepticuloidea), Roscidotoga (Nepticuloidea), Simplimorpha (Nepticuloidea), Simplimorpha lanceifoliella (Nepticuloidea), Simplimorpha promissa (Nepticuloidea), Stigmella (Nepticuloidea), Stigmella anomalella group (Nepticuloidea), Stigmella aurella group (Nepticuloidea), Stigmella betulicola group (Nepticuloidea), Stigmella epicosma group (Nepticuloidea), Stigmella floslactella group (Nepticuloidea), Stigmella hybnerella group (Nepticuloidea), Stigmella intronia (Nepticuloidea), Stigmella irregularis group (Nepticuloidea), Stigmella lapponica group (Nepticuloidea), Stigmella lemniscella group (Nepticuloidea), Stigmella malella (Nepticuloidea), Stigmella naturnella (Nepticuloidea), Stigmella ogygia group (Nepticuloidea), Stigmella oxyacanthella group (Nepticuloidea), Stigmella paliurella group (Nepticuloidea), Stigmella paradoxa group (Nepticuloidea), Stigmella prunifoliella group (Nepticuloidea), Stigmella quercipulchella group (Nepticuloidea), Stigmella saginella group (Nepticuloidea), Stigmella salicis group (Nepticuloidea), Stigmella sanguisorbae group (Nepticuloidea), Stigmella slingerlandella (Nepticuloidea), Stigmella tricentra (Nepticuloidea), Stigmella ulmivora group (Nepticuloidea), Stigmella ultima group (Nepticuloidea), Trifurcula (Nepticuloidea), Trifurcula cryptella group (Nepticuloidea), Trifurcula pallidella group (Nepticuloidea), Trifurcula subnitidella group (Nepticuloidea), Varius (Nepticuloidea), Zimmermannia (Nepticuloidea)
Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith