Fore wing of Permithone belmontensis, from Kukalová-Peck (1991).

Belongs within: Holometabola.
Contains: Raphidioptera, Megaloptera, Coniopterygidae, Euneuroptera.

The Neuropterida are a clade of largely predatory insects whose wings are often densely veined. Members are united by possession of a medially divided metapostnotum, the first abdominal tergum having a caudally bifid longitudinal sulcus, the fusion of the gonoplacs in the ovipositor, and a proventriculus with an unpaired diverticulum (Grimaldi & Engel 2005). Modern Neuroptera have been divided between three orders, the Raphidioptera, Megaloptera and Neuroptera. The Neuroptera, lacewings, are characterised by having the ninth gonocoxites associated with the gonarcus, and larvae with sucking mouthparts in which the maxillae and mandibles form a tube (Grimaldi & Engel 2005). Relationships between the three orders have been subject to debate but Megaloptera and Neuroptera may be united in a clade Eidoneuroptera by the integration of the larval cardines into the head capsule and the structure of the male genitalia. Characters shared between Megaloptera and Raphidioptera such as chewing mandibles in the larvae and exarate (mobile) pupae are likely to be plesiomorphies of the clade as a whole.

Potential fossil representatives of the Neuropterida include the Protoneuroptera that may represent stem-representatives of the Eidoneuroptera and/or Neuroptera alone (Engel et al. 2018). The Permian–Jurassic Glosselytrodea were generally small insects with subequal pairs of wings bearing numerous crossveins. The fore wings were tegminous and possessed an expanded precostal area forming a prominent bulge at the wing base (Grimaldi & Engel 2005).

<==Neuropterida [Hemerobini, Myrmeleontidea, Neuropteroidea] GE05
    |  i. s.: Ororaphidia megalocephala WHW10
    |--Glosselytrodea [Jurinida] GE05
    |    |--Archoglossopterum R02 [Archoglossopteridae GE05]
    |    |--Uskatelytrum R02 [Uskatelytridae GE05]
    |    |--Mongolojurina RJ93 [Polycytellidae GE05]
    |    |    `--M. altaica Ponomarenko 1988 RJ93
    |    |--Glosselytron R02 [Glosselytridae GE05]
    |    |    `--G. martynovae R02
    |    |--Glossopterum R02 [Glossopteridae GE05]
    |    |    `--G. martynovae R02
    |    `--Jurinidae GE05
    |         |--Jurina marginata GE05
    |         `--Eoglosselytrum KN13
    |              |--E. perfectum R02
    |              `--E. perplexa (Riek 1953) KN13
    `--+--+--Parasialidae EWB18
       |  |    |--Parasialis latipennis MW15, P02
       |  |    `--Sonjanasialis MW15
       |  `--+--Nanosialidae EWB18
       |     `--Raphidioptera GE05
       `--+--Protoneuroptera EWB18
          |    |--Permoberotha GE05 [Permoberothidae EWB18, Protoneuroptera]
          |    |    `--P. villosa GE05
          |    `--Permithonidae [Permopsychopidae] EWB18
          |         |--Sylvasenex lacrimabundus P02
          |         |--Permithonopsis enormis P02
          |         |--Permosisyra paurovenosa P02
          |         |--Permopsychops belmontensis Tillyard 1926 [incl. Permithone venosa Davis 1943] F71
          |         |--Permipsythone panfilovi RJ93
          |         |--Tschekardithonopsis oblivius B11
          |         `--Permithone K-P91
          |              |--P. belmontensis Tillyard 1922 [incl. Permosmylus pincombeae Tillyard 1926] F71
          |              |--P. neoxenus Riek 1953 F71
          |              `--P. oliarcoides Tillyard 1926 F71
          `--Eidoneuroptera EWB18
               |--Megaloptera GE05
               `--Neuroptera [Euneuropteroidea, Hemerobiiformia, Myrmeleontida, Planipennia] GE05
                    |  i. s.: Mesopolystoechotidae REL02
                    |           |--Mesopolystoechus Martynov 1937 REL02
                    |           `--Megapolystoechus magnificus RJ93
                    |         Archegetes neuropterum W13
                    |         Megapolystoechotes Tillyard 1933 REL02
                    |         Cratosisyrops gongazai Martins-Neto 1997 NM03
                    |         Grammosmylidae GE05
                    |         Barrowiella Smithers 1984 T13
                    |           `--B. butleri Smithers 1984 T13
                    |         Sialidopsis [Sialidopsidae] R70
                    |           `--S. kargalensis R70
                    |         Palaemerobiidae RJ93
                    |         Vatiga illudens M99
                    |         Chrysoleonites P02
                    |--Coniopterygidae EWB18
                    `--Euneuroptera EWB18

*Type species of generic name indicated


[B11] Bashkuev, A. S. 2011. Nedubroviidae, a new family of Mecoptera: the first Paleozoic long-proboscid scorpionflies. Zootaxa 2895: 47–57.

[EWB18] Engel, M. S., S. L. Winterton & L. C. V. Breitkreuz. 2018. Phylogeny and evolution of Neuropterida: where have wings of lace taken us? Annual Review of Entomology 63: 531–551.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[KN13] Kirejtshuk, A. G. & A. Nel. 2013. Skleroptera, a new order of holometabolous insects (Insecta) from the Carboniferous. Zoosystematica Rossica 22 (2): 247–257.

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[MW15] McKenna, D. D., A. L. Wild, K. Kanda, C. L. Bellamy, R. G. Beutel, M. S. Caterino, C. W. Farnum, D. C. Hawks, M. A. Ivie, M. L. Jameson, R. A. B. Leschen, A. E. Marvaldi, J. V. McHugh, A. F. Newton, J. A. Robertson, M. K. Thayer, M. F. Whiting, J. F. Lawrence, A. Ślipiński, D. R. Maddison & B. D. Farrell. 2015. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40 (4): 835–880.

[M99] Moraes, G. J. de. 1999. Pest status of the cassava green mite in Brazil and strategies for its control. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 287–291. Ohio Biological Survey: Columbus (Ohio).

[NM03] Nel, A., J.-J. Menier, A. Waller, G. Hodebert & G. de Ploëg. 2003. New fossil spongilla-flies from the lowermost Eocene amber of France (Insecta, Neuroptera, Sisyridae). Geodiversitas 25 (1): 109–117.

[P02] Ponomarenko, A. G. 2002. Superorder Myrmeleontidea Latreille, 1802 (=Neuropteroidea Handlirsch, 1903). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 176–189. Kluwer Academic Publishers: Dordrecht.

[R02] Rasnitsyn, A. P. 2002. Order Jurinida M. Zalessky, 1928 (=Glosselytrodea Martynov, 1938). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 189–192. Kluwer Academic Publishers: Dordrecht.

[REL02] Ren, D., M. S. Engel & W. Lü. 2002. New giant lacewings from the Middle Jurassic of Inner Mongolia, China (Neuroptera: Polystoechotidae). Journal of the Kansas Entomological Society 75 (3): 188–193.

[R70] Riek, E. F. 1970. Fossil history. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 168–186. Melbourne University Press.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[T13] Taylor, C. K. 2013. Annotated bibliography for Barrow Island terrestrial invertebrates. Records of the Western Australian Museum Supplement 83: 135–144.

[WHW10] Winterton, S. L., N. B. Hardy & B. M. Wiegmann. 2010. On wings of lace: phylogeny and Bayesian divergence time estimates of Neuropterida (Insecta) based on morphological and molecular data. Systematic Entomology 35: 349–378.

[W13] Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press: Princeton (New Jersey).

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