title: The phylogeny of brown lacewings (Neuroptera: Hemerobiidae) reveals multiple reductions in wing venation.
abstract: The last time the phylogenetic relationships among members of the family Hemerobiidae were studied quantitatively was over 12 years ago and based exclusively on morphology. Our study builds upon this morphological evidence by adding sequence data from three gene loci to provide a total evidence phylogeny of brown lacewings (Neuroptera: Hemerobiidae). Thirty-seven species representing nineteen Hemerobiidae genera were compared with outgroups from the families Ithonidae, Psychopsidae and Chrysopidae in Bayesian and parsimony analyses using a single nuclear gene (CAD) and two mitochondrial (16S rDNA and Cytochrome Oxidase I) genes. We compare divergence time estimates of Hemerobiidae cladogenesis under the two most commonly used relaxed clock models and discuss the evolution of wing venation in the family.
  
  
  
  We recovered a phylogeny largely incongruent with previously published morphological studies, although all but two subfamilies (i.e., Notiobiellinae and Drepanacrinae) were recovered as monophyletic. We found the subfamily Drepanacrinae paraphyletic with respect to Psychobiellinae, and Notiobiellinae to be polyphyletic. We thus offer a revised concept of Notiobiellinae, comprising only Notiobiella Banks, and erect a new subfamily Zachobiellinae including the remaining genera previously placed in Notiobiellinae. Psychobiellinae is synonymized with Drepanacrinae. Unlike the previous hypothesis that proposed a remarkably laddered topology, our tree suggests that hemerobiids diverged as three main clades. Moreover, in contrast to the vein proliferation hypothesis, we found that hemerobiids have instead undergone multiple reductions in the number of radial veins, this scenario questions the relevance of this character as diagnostic of various subfamilies
  
  
  
  Our phylogenetic hypothesis and divergence times analysis suggest that extant hemerobiids originated around the end of the Triassic and evolved as three distinct clades that diverged from one another during the Late Jurassic to Early Cretaceous. Contrary to earlier phylogenetic hypotheses, Carobius Banks (Carobiinae) is sister to the previously unplaced genus Notherobius New in a clade more closely related to Sympherobiinae, Megalominae and Zachobiellinae subfam. nov. The addition of taxa which are not available for DNA sequencing should be the focus of future studies, especially Adelphohemerobius Oswald, which is particularly important to test our inferences regarding the evolution of wing venation in Hemerobiidae.
authors.affiliation: California Department of Food & Agriculture, California State Collection of Arthropods, 3294 Meadowview Rd, Sacramento, CA, USA.
authors.forename: Ivonne J
authors.initials: IJ
authors.lastname: Garzón-Orduña
authors.affiliation: Instituto de Ciencias Básicas e Ingeniería, Universidad Autónoma del Estado de Hidalgo, Pachuca, Hidalgo, Mexico.
authors.forename: Imelda
authors.initials: I
authors.lastname: Menchaca-Armenta
authors.affiliation: Instituto de Ciencias Básicas e Ingeniería, Universidad Autónoma del Estado de Hidalgo, Pachuca, Hidalgo, Mexico.|Departamento de Zoología, Instituto de Biología, UNAM, Ciudad Universitaria, México, DF, Mexico.
authors.forename: Atilano
authors.initials: A
authors.lastname: Contreras-Ramos
authors.affiliation: Department of Entomology, China Agricultural University, Beijing, 100193, China.
authors.forename: Xingyue
authors.initials: X
authors.lastname: Liu
authors.affiliation: California Department of Food & Agriculture, California State Collection of Arthropods, 3294 Meadowview Rd, Sacramento, CA, USA. wintertonshaun@gmail.com.
authors.forename: Shaun L
authors.initials: SL
authors.lastname: Winterton
mesh_terms: D000818:Animals; D001499:Bayes Theorem; D005075:Biological Evolution; D007313:Insecta / Q000033:anatomy & histology* / Q000145:classification / Q000235:genetics*; D010802:Phylogeny; D014921:Wings, Animal / Q000033:anatomy & histology*