Owlflies are derived antlions: anchored phylogenomics supports a new phylogeny and classification of Myrmeleontidae (Neuroptera)

Abstract

The first phylogenomic analysis of the antlions is presented, based on 325 genes captured using anchored hybrid enrichment. A concatenated matrix including 207 species of Myrmeleontoidea (170 Myrmeleontidae) was analysed under maximum likelihood and Bayesian inference. Both Myrmeleontidae (antlions) and Ascalaphidae (owlflies) were recovered as paraphyletic with respect to each other. The majority of the subfamilies traditionally assigned to both Myrmeleontidae and Ascalaphidae were also recovered as paraphyletic. By contrast, all traditional antlion tribes were recovered as monophyletic (except Brachynemurini), but most subtribes were found to be paraphyletic. When compared with the traditional classification of Myrmeleontidae, our results do not support the current taxonomy. Therefore, based on our phylogenomic results, we propose a new classification for the antlions, which synonymizes Ascalaphidae with Myrmeleontidae and divides the family into four subfamilies (Ascalaphinae, Myrmeleontinae, Dendroleontinae and Nemoleontinae) and 17 tribes. We also highlight the most pressing issues in antlion systematics and indicate taxa that need further taxonomic and phylogenetic attention. Finally, we present a comprehensive table placing all extant genera of antlions and owlflies in our new proposed classification, including details on the number of species, distribution and notes on the likely monophyly of each genus.

Total‐evidence phylogeny of the owlflies (Neuroptera, Ascalaphidae) supports a new higher‐level classification

“…Although the results obtained in the current study are based on much less data than those of Machado et al (), the topology presented here (Figure , based on the Bayesian phylogeny) is taken as a better estimate of relationships for the Ascalaphidae and closely allied antlions, for the following reasons. The final topology here of a monophyletic Palparine + (Stilbopteryginae + Ascalaphidae) is congruent with that obtained by Machado et al (). The present study included a much richer taxon sampling for the ingroup. Key higher‐level clades were recovered as monophyletic in every analysis (Figs. –), including all major groups within the owlflies, despite their relationships to one another changing slightly. These same nodes, generally, exhibited high support in each analysis (posterior probabilities, bootstrap values, decay indices). The result of a monophyletic Ascalaphidae independent of Stilbopteryginae is congruent with previous hypotheses and numerous analyses based on either morphology or molecular data (Badano, Aspöck, Aspöck, & Haring, ; Badano, Aspöck, Aspöck, & Cerretti, ; Henry, ; Mansell, ; Michel et al, ; New, ; Stange, , ). The result of Stilbopteryginae as an independent lineage immediately adjacent to Ascalaphidae is congruent with other recent studies based on extensive morphological (Badano, Aspöck, Aspöck, & Cerretti, ) and molecular data (Michel et al, ). Examination of the results of the morphology‐only cladogram (Figure ) in the current study reveals that while it supports a monophyletic Ascalaphidae, and a sister group relationship for the Stilbopteryginae, Bremer supports are very low or nil, suggesting that most of the signal for relationships recovered in the combined data analyses comes from the molecular data, and is not being overpowered by the morphological data. Although the morphology‐based cladistic analysis (Figure ) had weak support, and exhibited numerous unresolved clades, its sequence of nesting of major groups nevertheless agreed with that of the Bayesian analysis (Figure ).…”

“…Until Jones ( and the present work) and Machado et al (), the only previous author to have attempted any sort of detailed, tree‐based analysis of relationships within the family Ascalaphidae was Henry (, ). Henry () presented a simple dendrogram (Figure ) that optimized several characters: (a) evolution and loss of repagula (“barriers”: defined as abortive eggs laid below egg masses on twigs in some owlflies), from abortive eggs with trophic functions, to abortive eggs with barrier function, to ant‐repelling repagula, and then lost; (b) split eyes; and (c) ovariole number.…”

“…| 763 JONES amount of phylogenetic data (Aspöck et al, 2001;Badano, Aspöck, Aspöck, & Haring, 2017;Badano, Aspöck, Aspöck, & Cerretti, 2017;Gao et al, 2018;Henry, 1978c;Jones, 2014;Lan et al, 2016;Machado et al, 2018;Michel et al, 2017;Riek, 1976;Song et al, 2019Song et al, , 2018Stange, 1994;Wang et al, 2017;Winterton et al, 2010Winterton et al, , 2018Zhang & Yang, 2017).…”

Diversity of lacewing assemblages (Neuropterida:Neuroptera) in different forest habitats and agricultural areas in the East Mediterranean area of Turkey

“…The subfamily Ascalaphinae (Myrmeleontidae) was analyzed as a separate taxon from the other three subfamilies of Myrmeleontidae, since it was included in Myrmeleontidae recently (Machado et al . ).…”

“…Ascalaphinae, Chrysopidae, Coniopterygidae, Myrmeleontidae (except Ascalaphinae), and Nemopteridae to test whether geographical distance affected species composition. The subfamily Ascalaphinae (Myrmeleontidae) was analyzed as a separate taxon from the other three subfamilies of Myrmeleontidae, since it was included in Myrmeleontidae recently (Machado et al 2019).…”