The phylogeny of brown lacewings (Neuroptera: Hemerobiidae) reveals multiple reductions in wing venation

Abstract

BackgroundThe 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.ResultsWe 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 subfamiliesConclusionsOur 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.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0746-5) contains supplementary material, which is available to authorized users.

Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera)

“…Carobinae and Sympherobinae are recovered as sister groups, as found in Garzón‐Orduña et al . () as well as the close relationship among Notiobiellinae, Hemerobiinae, Drepanacrinae and Psychobiellinae. The sister‐group relationship between Microminae and Drepanepteryginae was recovered here with strong support, as indicated previously by Oswald () and Garzón‐Orduña et al .…”

“…Internal relationships within Hemerobiidae found here overlap in part with both previous quantitative studies on the group using morphology (Oswald, ) and DNA sequences combined with morphology (Garzón‐Orduña et al ., ). Here we recovered Zachobiella Banks (Zachobiellinae) as sister to the rest of Hemerobiidae, a result not found in previous studies.…”

Evolution of green lacewings (Neuroptera: Chrysopidae): a molecular supermatrix approach

“…By incorporating DNA sequences from previous studies and new sequences as part of this study, we sampled 84 species of Chrysopidae in 51 genera, representing all tribes, subfamilies and previously recognized groups of genera (Brooks, ), in the most comprehensive analysis of the family to date (Table S5). Where possible, we included multiple exemplar species per genus to ensure adequate sampling of respective lineages, as done in previous studies in related lacewing families (Garzón‐Orduña et al ., ; Liu et al ., ; Badano et al ., ; Winterton et al ., , 2018). The highest proportion of sampled genera came from Nothochrysinae (66% of the genera in the family), followed by Chrysopinae (62% including Nothancyla ), and only a single genus (with multiple species) sampled for Apochrysinae (17%).…”

Evolution of green lacewings (Neuroptera: Chrysopidae): an anchored phylogenomics approach

“…Hemerobiidae has long been considered the sister family to Chrysopidae, based primarily on the morphological similarity of their larval stages (including a trumpet‐shaped empodium in at least the first instar). This phylogenetic association has been recovered in some quantitative studies using both morphology and DNA sequence data (Winterton et al ., , ; Garzón‐Orduña et al ., ; Wang et al ., ), while other published studies using a variety of data sources have also recovered Hemerobiidae in other locations within Neuroptera, and not sister to Chrysopidae (e.g. Winterton, ; Yang et al ., ).…”

“…Recent studies have begun resolving higher‐level relationships within Neuroptera using large amounts of DNA sequence data, resulting in progress in understanding evolution of the order at all levels (e.g. Winterton et al ., , ; Liu et al ., ; Shi et al ., ; Garzón‐Orduña et al ., , 2017; Wang et al ., ; Bakkes et al ., ; Machado et al ., ). Here, we present the first large‐scale genomic approach to understanding green lacewing phylogeny, in this instance using anchored hybrid enrichment data sequenced for 82 species, representing 50 genera of chrysopids.…”