On wings of lace: phylogeny and Bayesian divergence time estimates of Neuropterida (Insecta) based on morphological and molecular data

Abstract

Neuropterida comprise the holometabolan orders Neuroptera (lacewings, antlions and relatives), Megaloptera (alderflies, dobsonflies) and Raphidioptera (snakeflies) as a monophyletic group sister to Coleoptera (beetles). The higher‐level phylogenetic relationships among these groups, as well as the family‐level hierarchy of Neuroptera, have to date proved difficult to reconstruct. We used morphological data and multi‐locus DNA sequence data to infer Neuropterida relationships. Nucleotide sequences were obtained for fragments of two nuclear genes (CAD, 18S rDNA) and two mitochondrial genes (COI, 16S rDNA) for 69 exemplars representing all recently recognized families of Neuropterida as well as outgroup exemplars from Coleoptera. The joint posterior probability of phylogeny and divergence times was estimated using a Bayesian relaxed‐clock inference method to establish a temporal sequence of cladogenesis for the group over geological time. Megaloptera were found to be paraphyletic with respect to the rest of Neuropterida, calling into question the validity of the ordinal status for Megaloptera as presently defined. Ordinal relationships were weakly supported, and monophyly of Megaloptera was not recovered in any total‐evidence analysis; Corydalidae were frequently recovered as sister to Raphidioptera. Only in relaxed‐clock inferences were Raphidioptera and a paraphyletic Megaloptera recovered with strong support as a monophyletic group sister to Neuroptera. A monophyletic Neuroptera diverged from a common Raphidioptera + ‘Megaloptera’ ancestor during the Late Carboniferous. Contrary to some previous hypotheses, Coniopterygidae, not Nevrorthidae, were recovered as sister to the rest of Neuroptera, with Nevrorthidae recovered with Osmylidae and Sisyridae. The monophyly of the universally recognized Myrmeleontiformia was confirmed, with an origin in the mid‐Triassic, but a monophyletic Hemerobiiformia was not recovered in any analysis. Dilaridae were not closely related to the clade comprising Mantispidae and Berothidae, and diverged earlier than proposed previously. The phylogenetic status and taxonomic composition of Polystoechotidae and Ithonidae are in need of re‐evaluation, as Oliarces Carpenter (presently Ithonidae) was placed well within the present circumscription of Polystoechotidae.

A new genus of dustywings (Neuroptera: Coniopterygidae) in Late Cretaceous Vendean amber

“…Nevertheless, Garnaconis n. gen. has the hind wing base of Rs very close to that of M, which is a character present in Aleuropteryginae and in Flintoconis Sziráki, second brucheiserine genus, while Brucheiser Navás, has highly modified fore and hind wing venation delicate to interpret (Riek, 1975). The polarity of this character remains controversial because even the sister-group relationships of Coniopterygidae within the Neuroptera remain debatable: Aspöck, Plant, and Nemeschkal (2001) supported a 'Coniopterygidae + Sisyridae' clade, while Haring and Aspöck (2004) and Aspöck and Aspöck (2008) supported a 'Coniopterygidae + dilarid clade' (see summary in Aspöck & Aspöck, 2007); Winterton, Hardy, and Wiegmann (2010) found Coniopterygidae as sister group of all other Neuroptera; Beutel, Friedrich, and Aspöck (2010) considered that the position of this family remains uncertain; Zimmermann and others (2011) considered them as sister group to the clade (Mantispidae + (Dilaridae + (Rhachiberothidae + Berothidae))); while Aspöck, Haring, & Aspöck (2012) proposed them as sister group of the (Dilaridae + (Mantispidae + (Rhachiberothidae + Berothidae))). Note that this last hypothesis, as for the sisyrid or dilarid hypotheses, is congruent with a basal position of Rs as a plesiomorphy for the Coniopterygidae.…”

Contribution to the knowledge of Mantispoidea, Osmyloidea, and Myrmeleontoidea with new records for the Neuroptera fauna of northwestern Turkey

“…Dilaridae is a small family of the superfamily Osmyloidea (Winterton et al, 2010). D. turcicus adults were found in deciduous and coniferous forests in Bursa Province.…”

“…The semiaquatic larvae of Osmylus fulvicephalus (Scop. ), which is the most common species throughout Europe, prey on small arthropods (Gillott, 2005;Winterton et al, 2010).…”

The Nevrorthidae, mistaken at all times: phylogeny and review of present knowledge (Holometabola, Neuropterida, Neuroptera)

“…In the course of further molecular analyses, mentioned above, Nevrorthidae was retrieved either as a sister group to Sisyridae and Osmylidae and all three constituted a monophylum (Winterton et al 2010), or Nevrorthidae and Sisyridae alone constituted the sister group to the rest of Neuroptera except Coniopterygidae (Wang et al 2016).…”

“…, Haring and U. Aspöck 2004, U. Aspöck and H. Aspöck 2008a, Beutel et al 2010, Winterton et al 2010, Zimmermann et al 2011, Randolf et al 2013, Randolf et al 2014, Wang et al 2016, several hypotheses, which have been catalysed via Nevrorthidae, are of general significance regarding Neuropterida:…”