Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola)

Abstract

Segment 9 of male Raphidioptera, comprising tergite, sternite, gonocoxites, gonostyli and gonapophyses, is a benchmark for homologies in the male and female terminalia of the three Neuropterida orders Raphidioptera, Megaloptera and Neuroptera. The segments relating to genitalia are 9, 10 and 11 in males and 7, 8 and 9 in females. Results from holomorphological and recent molecular cladistic analyses of Neuropterida agree in supporting the sister‐group relationships between: (1) the Raphidioptera and the clade Megaloptera + Neuroptera, and (2) the suborder Nevrorthiformia and all other Neuroptera. The main discrepancy between the results of these studies is the nonmonophyly of the suborder Hemerobiiformia in the molecular analysis. The monophyly of the Megaloptera (which has been repeatedly questioned) is further corroborated by a hitherto overlooked ground pattern autapomorphy: the presence of eversible sacs within the complex of the fused gonocoxites 11 in Corydalidae and Sialidae. The recently discovered paired complex of gonocoxites 10 (parameres) in Nipponeurorthus (Nevrorthidae) indicates that the curious apex of sternite 9 of Nevrorthus and Austroneurorthus is the amalgamation of the sclerites of gonocoxites 10 with sternite 9, interpreted as synapomorphic. In the molecular study, the Nevrorthidae, Sisyridae and Osmylidae branch off in consecutive splitting events, a result that is supported by the analysis of male genital sclerites reported here. Extraordinary parallel apomorphies (e.g. excessive enlargement and modification of gonocoxites 10 ending in a thread‐like ‘penisfilum’) in derived representatives of Coniopterygidae, Berothidae, Rhachiberothidae and Mantispidae corroborate the dilarid clade of the morphological analysis and leads us to hypothesize a sister‐group relationship of the Coniopterygidae with the dilarid clade. A re‐interpretation of the tignum of Chrysopidae as gonocoxites 11 means that the structure previously called the gonarcus represents the fused gonocoxites 9. In Hemerobiidae, the corresponding sclerite is consequently also homologized as fused gonocoxites 9. The enlargement of the lateral wings of the gonocoxites in both families is interpreted as a synapomorphy. Excessive enlargement of gonostyli 11 in the Polystoechotid clade and Myrmeleontiformia supports a sister‐group relationship of these two clades. The occurrence of certain serial homologues of female genitalia structures (gonocoxites and gonapophyses), such as the digitiform processus together with the flat appendices in segment 8 of certain Myrmeleontidae, or the wart‐like processus together with the flat circular sclerites in segment 7 of certain Berothidae, as well as the presence of gonocoxites 8 as pseudosternites in certain Nemopteridae and Coniopterygidae, are probably character reversals. The digitiform processus of tergite 9 (pseudogonocoxites) in Rhachiberothidae and Austroberothella (Berothidae) are either independently developed acquisitions with a function in oviposition, or are homologous sclerites...

Sperm structure of some Neuroptera and phylogenetic considerations

“…In the last 10 years, different relationships among the Neuroptera families have been proposed based either on the cladistic analysis of some selected morphological characters (Aspöck et al 2001), molecular data (Aspöck et al 2003;Winterton 2003;Haring & Aspöck 2004), the genital sclerites (Aspöck & Aspöck 2008), or the morphology of the larva head (Beutel et al 2010). In particular, morphological studies suggested that Mantispidae would be placed in a clade together with Berothidae, Rhachiberothidae and Dilaridae (the dilarid clade).…”

“…In particular, morphological studies suggested that Mantispidae would be placed in a clade together with Berothidae, Rhachiberothidae and Dilaridae (the dilarid clade). Moreover, a sister-group relationship of Coniopterygidae + dilarid clade has been hypothesized (Aspöck et al 2001;Aspöck & Aspöck 2008;Zimmermann et al 2009;Beutel et al 2010). On the contrary, Hemerobiidae and Chrysopidae would have to be placed in a separate clade.…”

Ancient pinnate leaf mimesis among lacewings

“…The genus is noteworthy for the large body size, elongate forewings, long filiform antennae, protrudent compound eyes, undulating margins to the forewings, disruptive pattern of coloration resembling leaf pinnae, distinctly thickened MP branches resembling a leaf rachis, complicated venation, and an appendage close to the caudal segment (Fig. 1B) that resembles the gonapophysis lateralis of Osmylidae, a relatively primitive feature among Neuroptera (10). Although moderate-to small-sized lacewings are dominant among extant Neuroptera, such large species are frequently found in the Middle Jurassic, notably species of the Aetheogrammatidae, Kalligrammatidae, and Grammolingiidae (11)(12)(13).…”

“…Abdomen with 10 segments and with an appendix close to caudal segment. The appendix resembles the gonapophysis lateralis of Osmylidae, which represents a primitive character of Neuroptera (10).…”

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.…”