Raphidiidae

Raphidia ophiopsis, copyright Guido Bohne.

Belongs within: Raphidioptera.

The Raphidiidae are a group of snakeflies characterised by the presence of three ocelli and a tube-like pronotum laterally covering the pleural region (Aspöck & Aspöck 1991).

<==Raphidiidae
    |--Agulla Navás 1914 VM20, A02
    |    |--A. adnixa K67
    |    |--A. astuta RD77
    |    `--A. bicolor WHW10
    `--+--Mongoloraphidia VM20
       |    |--M. eklipes GE05
       |    |--M. martynovae WHW10
       |    `--M. sororcula VM20
       `--+--+--+--Ohmella VM20
          |  |  |    |--O. baetica VM20
          |  |  |    |    |--O. b. baetica VM20
          |  |  |    |    `--O. b. bolivari VM20
          |  |  |    `--O. coffini Nel 1993 E02
          |  |  `--Atlantoraphidia maculicollis VM20
          |  `--+--Xanthostigma VM20
          |     |    |--X. gobicola VM20
          |     |    `--X. xanthostigma PK17
          |     `--+--Puncha Navás 1915 VM20, S63
          |        |    `--P. ratzeburgi VM20
          |        `--Venustoraphidia nigricollis VM20
          `--+--+--+--Subilla Navás 1916 VM20, S63
             |  |  |    |--S. confinis VM20
             |  |  |    |--S. schneideri S63
             |  |  |    |--S. sericea S63
             |  |  |    `--S. sulfuricosta Steinmann 1963 S63
             |  |  `--Phaeostigma VM20
             |  |       |--P. divina VM20
             |  |       |--P. major VM20
             |  |       `--P. robusta AA91
             |  `--+--Turcoraphidia amara VM20
             |     `--+--Parvoraphidia microstigma VM20
             |        `--Ornatoraphidia flavilabris VM20
             `--+--Dichrostigma flavipes VM20
                `--Raphidia Linnaeus 1758 E02 (see below for synonymy)
                     |--*R. ophiopsis Linnaeus 1758 E02
                     |--R. baltica Carpenter 1956 AA04
                     |--R. creedei Carpenter 1936 E02
                     |--R. elegans (Cockerell 1907) [=*Megaraphidia elegans] E02
                     |--R. euxina Navas 1915 [=R. (*Pretzmannia) euxina] E02
                     |--R. exhumata Cockerell 1909 [incl. R. pulveris Cockerell 1914] E02
                     |--R. flaviceps K01
                     |--R. funerata GE05
                     |--R. mediterranea VM20
                     |--R. mortua Rohwer 1909 E02
                     |--R. notata RD77
                     |--R. nuchensis Aspöck, Aspöck & Martynova 1968 [=R. (*Aserbeidshanoraphidia) nuchensis] E02
                     |--R. oblita A99
                     |--R. palaeoformis Aspöck & Aspöck 1968 [=R. (*Nigroraphidia) palaeoformis] E02
                     |--R. tranquilla Scudder 1890 [=Archiraphidia tranquilla; incl. Inocellia somnolenta Scudder 1890] E02
                     |--R. tumulata (Scudder 1890) (see below for synonymy) E02
                     `--R. veterana (Scudder 1890) [=Inocellia veterana, *Dictyoraphidia veterana] E02

Raphidiidae incertae sedis:
  Succinoraphidia Aspöck & Aspöck 2004 [Succinoraphidiinae] AA04
    `--*S. exhibens Aspöck & Aspöck 2004 AA04
  Lesna Navás 1915 S63
    |--L. adanana S63
    |--L. belinayi S63
    |--L. biroi S63
    |--L. caucasica S63
    |--L. davidi S63
    |--L. hungarica S63
    |--L. laticaput Steinmann 1963 S63
    |--L. longicauda S63
    |--L. lestica Steinmann 1963 S63
    |--L. major S63
    |--L. navasi Steinmann 1963 S63
    |--L. notata S63
    |--L. parnassia S63
    |--L. pilicollis S63
    `--L. stigmata Steinmann 1963 S63
  Glavia Navás 1916 S63
  Alena Navás 1916 S63
  Rhaphidilla Navás 1915 S63
  Navasana Steinmann 1963 S63
    `--*N. perumbrata Steinmann 1963 S63
  Harraphidia Steinmann 1963 S63
    `--*H. harpyia Steinmann 1963 S63

Raphidia Linnaeus 1758 E02 [=Rhaphidia Burmeister 1839 S63; incl. Archiraphidia Handlirsch 1910 E02, Aserbeidshanoraphidia Aspöck & Aspöck 1968 E02, Dictyoraphidia Handlirsch 1910 E02, Megaraphidia Cockerell 1907 E02, Nigroraphidia Aspöck & Aspöck 1968 E02, Pretzmannia Aspöck & Aspöck 1968 E02; Raphidiinae]

Raphidia tumulata (Scudder 1890) [=Inocellia tumulata, *Archiraphidia tumulata; incl. Inocellia eventa Scudder 1890, Archiraphidia eventa] E02

*Type species of generic name indicated

REFERENCES

[A02] Aspöck, U. 2002. Phylogeny of the Neuropterida (Insecta: Holometabola). Zoologica Scripta 31: 51–55.

[AA91] Aspöck, H., & U. Aspöck. 1991. Raphidioptera (snake-flies, camelneck-flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 521–524. Melbourne University Press: Carlton (Victoria).

[AA04] Aspöck, U., & H. Aspöck. 2004. Two significant new snakeflies from Baltic amber, with discussion on autapomorphies of the order and its included taxa (Raphidioptera). Systematic Entomology 29: 11–19.

[A99] Ax, P. 1999. Das System der Metazoa II. Ein Lehrbuch der phylogenetischen Systematik. Gustav Fisher Verlag: Stuttgart (transl. 2000. Multicellular Animals: The phylogenetic system of the Metazoa vol. 2. Springer).

[E02] Engel, M. S. 2002. The smallest snakefly (Raphidioptera: Mesoraphidiidae): a new species in Cretaceous amber from Myanmar, with a catalogue of fossil snakeflies. American Museum Novitates 3363: 1–22.

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

[K01] Klapálek, F. 1901. Neuropteroidák [Neuropteroiden]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 203–221. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[K67] Koch, L. E. 1967. The genus Chasmoptera (Neuroptera: Nemopteridae), with the description of a new species from Western Australia. Proceedings of the Royal Entomological Society, London (B) 36 (9–10): 137–146.

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[S63] Steinmann, H. 1963. Raphidiopterological studies I. Navasana gen. n. from Hungary, Harraphidia gen. n. from Morocco, new Lesna Nav. and Subilla Nav. species from Europe. Acta Zoologica Academiae Scientiarum Hungaricae 9 (1–2): 183–198.

[VM20] Vasilikopoulos, A., B. Misof, K. Meusemann, D. Lieberz, T. Flouri, R. G. Beutel, O. Niehuis, T. Wappler, J. Rust, R. S. Peters, A. Donath, L. Podsiadlowski, C. Mayer, D. Bartel, A. Böhm, S. Liu, P. Kapli, C. Greve, J. E. Jepson, X. Liu, X. Zhou, H. Aspöck & U. Aspöck. 2020. An integrative phylogenomic approach to elucidate the evolutionary history and divergence times of Neuropterida (Insecta: Holometabola). BMC Evolutionary Biology 20: 64.

[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.

Raphidioptera

Fibla hesperica, copyright Thijs Valkenburg.

Belongs within: Neuropterida.
Contains: Baissopteridae, Mesoraphidiidae, Raphidiidae.

The Raphidioptera, snakeflies, are a Holarctic group of predatory insects characterised by a remarkably elongate pronotum giving a appearance of a distinct 'neck'.

Characters (from Aspöck & Aspöck 1991): Mandibulate, endopterygote Neoptera; antennae multisegmented; compound eyes always present; ocelli present only in Raphidiidae; prothorax elongate; with two pairs of subequal wings; ovipositor long. Larvae terrestrial; with biting mouth-parts and lateral ocelli. Pupae decticous.

<==Raphidioptera [Raphidiida, Raphidiodea]
    |  i. s.: Archiinocellia Handlirsch 1910 E02
    |           `--*A. oligoneura Handlirsch 1910 E02
    |         Arariperaphidia Martins-Neto & Vulcano 1989 E02
    |           `--*A. rochai Martins-Neto & Vulcano 1989 E02
    |         Erma Navás 1918 [Ermidae] S63
    |--Priscaenigmatomorpha EWB18
    |    |--Chrysoraphidiidae EWB18
    |    `--Priscaenigmatidae AA04
    |         |--Priscaenigma Whalley 1985 AA04
    |         |    `--*P. obtusa Whalley 1985 E02
    |         `--Hondelagia Bode 1953 E02
    |              `--*H. reticulata Bode 1953 [=Mesoraphidia reticulata] E02
    `--Raphidiomorpha E02
         |--Juroraphidiidae EWB18
         `--+--Baissopteridae GE05
            `--+--Mesoraphidiidae GE05
               |--Alloraphidiidae GE05
               |    |--Pararaphidia Willmann 1994 E02
               |    |    |--P. deserta (Ponomarenko 1988) [=Alloraphidia deserta] E02
               |    |    `--P. vitimica (Martynova 1961) [=Proraphidia vitimica, Alloraphidia vitimica] E02
               |    |--Archeraphidia Ponomarenko 1988 E02
               |    |    |--*A. yakowlewi Ponomarenko 1988 E02
               |    |    |--A. baisensis Ponomarenko 1993 E02
               |    |    `--A. hosbayari Ponomarenko 1988 E02
               |    `--Alloraphidia Carpenter 1967 E02
               |         |--*A. dorfi Carpenter 1967 E02
               |         |--A. anomala Ren 1997 E02
               |         |--A. asiatica Ponomarenko 1993 E02
               |         `--A. petrosa Ponomarenko 1988 E02
               `--Raphidioidea [Neoraphidioptera, Raphidiina] AA04
                    |--Raphidiidae AA04
                    `--Inocelliidae AA04
                         |  i. s.: Parainocellia Aspöck & Aspöck 1968 AA04
                         |           |--P. bicolor WHW10
                         |           `--P. ressli AA91
                         |         Electrinocellia Engel 1995 E02
                         |           `--*E. peculiaris (Carpenter 1956) [=Inocellia peculiaris] E02
                         |         Succinofibla Aspöck & Aspöck 2004 AA04
                         |           `--*S. aperta Aspöck & Aspöck 2004 AA04
                         |         Sininocellia Yang 1985 AA04
                         |--Negha Navás 1916 VM20, AA04
                         |    `--N. meridionalis TW05
                         `--+--Inocellia Schneider 1843 VM20, AA04
                            |    |--I. crassicornis VM20
                            |    `--I. japonica VM20
                            `--Fibla Navás 1915 [incl. Burcha Navás 1915, Estoca Navás 1919] E02
                                 |  i. s.: F. erigena (Menge 1856) AA04 [=Raphidia erigena AA04, R. (Inocellia) erigena E02]
                                 |         F. exusta (Cockerell & Custer 1925) AA04 [=Inocellia exusta E02]
                                 |         F. maclachlani (Albarda 1891) VM20, E02 (see below for synonymy)
                                 |         ‘*Estoca’ peyerimhoffi Navás 1919 E02
                                 |--F. (Fibla) AA04
                                 |    |--*F. (F.) hesperica Navás 1915 E02
                                 |    `--F. (F.) carpenteri Engel 1995 AA04
                                 `--F. (Reisserella) Aspöck & Aspöck 1971 [incl. Miofibla Nel 1993] E02
                                      |--F. (*R.) pasiphae Aspöck & Aspöck 1971 [=Inocellia (R.) pasiphae] E02
                                      `--F. (R.) cerdanica (Nel 1993) [=Miofibla cerdanica] E02

Fibla maclachlani (Albarda 1891) VM20, E02 [=Inocellia maclachlani E02, *Burcha maclachlani E02]

*Type species of generic name indicated

REFERENCES

[AA91] Aspöck, H., & U. Aspöck. 1991. Raphidioptera (snake-flies, camelneck-flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 521–524. Melbourne University Press: Carlton (Victoria).

[AA04] Aspöck, U., & H. Aspöck. 2004. Two significant new snakeflies from Baltic amber, with discussion on autapomorphies of the order and its included taxa (Raphidioptera). Systematic Entomology 29: 11–19.

[E02] Engel, M. S. 2002. The smallest snakefly (Raphidioptera: Mesoraphidiidae): a new species in Cretaceous amber from Myanmar, with a catalogue of fossil snakeflies. American Museum Novitates 3363: 1–22.

[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.

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

[S63] Steinmann, H. 1963. Raphidiopterological studies I. Navasana gen. n. from Hungary, Harraphidia gen. n. from Morocco, new Lesna Nav. and Subilla Nav. species from Europe. Acta Zoologica Academiae Scientiarum Hungaricae 9 (1–2): 183–198.

[TW05] Terry, M. D., & M. F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240–257.

[VM20] Vasilikopoulos, A., B. Misof, K. Meusemann, D. Lieberz, T. Flouri, R. G. Beutel, O. Niehuis, T. Wappler, J. Rust, R. S. Peters, A. Donath, L. Podsiadlowski, C. Mayer, D. Bartel, A. Böhm, S. Liu, P. Kapli, C. Greve, J. E. Jepson, X. Liu, X. Zhou, H. Aspöck & U. Aspöck. 2020. An integrative phylogenomic approach to elucidate the evolutionary history and divergence times of Neuropterida (Insecta: Holometabola). BMC Evolutionary Biology 20: 64.

[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.

Neuropterida

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

REFERENCES

[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).

Osmylidae

Porismus strigatus, copyright Donald Hobern.

Belongs within: Euneuroptera.

The Osmylidae are a diverse group of lacewings distinguished by the presence of more or less distinct ocelli (New 1991).

Characters (from New 1991): Generally medium to large (fore wing length 12-30 mm or more), with wings often patterned with grey or brown and rather broad; antennae usually less than half as long as fore wing; three ocelli present, at least represented by raised tubercles on vertex; female genitalia with enlarged gonocoxite 9, and 2 spermathecae. Larvae long and fusiform, with long tapering straight jaws.

Osmylidae
    |--Spilosmylomorpha EWB18
    |    |--Gumilla A69 [Gumillinae EWB18]
    |    |--Spilosmylinae RY03
    |    |    |--Spilosmylus flavicornis RD77
    |    |    `--Conchylosmylus triseriatus R70b
    |    `--Protosmylinae EWB18
    |         |--Paryphosmylus ornatus A69
    |         `--Protosmylus P92
    `--Osmylomorpha EWB18
         |  i. s.: Eidoporismus N91 [Eidoporisminae EWB18]
         |           `--E. pulchellus N91
         |--Osmylinae RY03
         |    |--Plethosmylus A69
         |    `--Osmylus Latreille 1802 WHW10, L02
         |         |--*O. maculatus [=Hemerobius maculatus] L02
         |         |--O. chrysops A99
         |         |--O. fulvicephalus GE05
         |         `--O. strigosus G27
         `--Kempyninae EWB18
              |  i. s.: Clydosmylus N91
              |--Australysmus lacustris WHW10
              `--+--Kempynus WHW10
                 |    |--K. falcatus A69
                 |    `--K. kimminsi WHW10
                 `--+--Porismus [Porisminae] WHW10
                    |    `--P. strigatus N91
                    `--Stenosmylinae EWB18
                         |--Euporismus albatrox N91
                         |--Oedosmylus brevis WHW10
                         |--Stenosmylus N91
                         |--Carinosmylus N91
                         |--Stenolysmus N91
                         |--Isostenosmylus A69
                         `--Phymatosmylus Adams 1969 A69
                              `--*P. caprorum Adams 1969 A69

Osmylidae incertae sedis:
  Mesosmylus atalantus P02
  Kazakhstania fasciata P02
  Epiosmylus Panfilov 1980 RY02
    |--E. longicornis RY02
    `--E. panfilovi Ren & Yin 2002 RY02
  Euporismites R70a
  Sogjuta speciosa RJ93
  Kalosmylinae R70b
  Archeosmylus RJ93 [Archeosmylidae, Archeosmylinae EWB18]
    `--A. complexus Whalley 1988 RJ93

*Type species of generic name indicated

REFERENCES

[A69] Adams, P. A. 1969. A new genus and species of Osmylidae (Neuroptera), from Chile and Argentina, with a discussion of planipennian genitalic homologies. Postilla 141: 1–11.

[A99] Ax, P. 1999. Das System der Metazoa II. Ein Lehrbuch der phylogenetischen Systematik. Gustav Fisher Verlag: Stuttgart (translated: 2000. Multicellular Animals: The phylogenetic system of the Metazoa vol. 2. Springer).

[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.

[G27] Girault, A. A. 1927. Thysanoptera nova Australiensis. II. Privately published (reprinted: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 214–215).

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

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[N91] New, T. R. 1991. Neuroptera (lacewings). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 525–542. Melbourne University Press: Carlton (Victoria).

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[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.

[RY02] Ren D. & Yin J.-C. 2002. [A new Middle Jurassic species of Epiosmylus from Inner Mongolia, China (Neuroptera: Osmylidae)]. Acta Zootaxonomica Sinica 27 (2): 274–277.

[RY03] Ren, D., & J. Yin. 2003. New ‘osmylid-like’ fossil Neuroptera from the Middle Jurassic of Inner Mongolia, China. Journal of the New York Entomological Society 111 (1): 1–11.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[R70a] Riek, E. F. 1970a. Fossil history. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 168–186. Melbourne University Press.

[R70b] Riek, E. F. 1970b. Neuroptera (lacewings). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 472–494. 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.

[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.

Megaloptera

New Zealand dobsonfly Archichauliodes diversus, copyright Steve Reekie.

Belongs within: Neuropterida.

The Megaloptera, alderflies and dobsonflies, are a group of long-winged insects with aquatic predatory larvae.

Characters (from Theischinger 1991): Mandibulate, endopterygote Neoptera, with two pairs of subequal, functional membranous wings and very soft abdomen. Larvae aquatic; mandibulate, with large prothorax, functional legs and lateral abdominal gills. Pupae decticous and exarate.

<==Megaloptera [Corydalida, Sialoidea]
    |  i. s.: Chauliosialis sukatshevae K-P91, S02
    |         Tychtodelopterum [Tychtodelopteridae] MW15
    |--Euchauliodes [Euchauliodidae] GE05
    |    `--E. distinctus K-P91
    |--Sialidae GE05
    |    |--Indosialis P02
    |    |--Austrosialis ignicollis T91, ZS10
    |    |--Stenosialis T91
    |    |--Protosialis R70
    |    |--Dobbertinia [Dobbertiniidae] GE05
    |    |    `--D. reticulata GE05
    |    `--Sialis Latreille 1802 L02
    |         |--*S. lutaria L02, VM20 [=Semblis lutaria L02]
    |         |--S. californica WHW10
    |         |--S. fuliginosa RD77
    |         |--S. hamata TW05
    |         |--S. mohri WHW10
    |         `--S. strausi S02
    `--Corydalidae GE05
         |  i. s.: Parachauliodes YS01
         |--+--Corydalus Latreille 1802 L02 [Corydalinae T91]
         |  |    |--*C. cornutus (Linnaeus 1758) L02, L58 [=Hemerobius cornutus L02]
         |  |    `--C. armatus WHW10
         |  `--Protohermes VM20
         |       |--P. grandis K-P83
         |       `--P. xanthodes VM20
         `--Chauliodinae [Chauliodidae] T91
              |  i. s.: Cretochaulus lacustris P02
              |         Neohermes californicus WHW10
              |         Nigronia WHW10
              |           |--N. fasciatus HR11
              |           `--N. serricornis WHW10
              `--Chauliodini R70
                   |--Chauliodes Latreille 1802 L02
                   |    |--*C. pectinicornis [=Hemerobius pectinicornis] L02
                   |    `--C. prisca Pictet 1854 P92
                   |--Protochauliodes T91
                   |    |--P. biconicus T91
                   |    `--P. kirramae T91
                   `--Archichauliodes T91
                        |  i. s.: A. dubitatis H74
                        |--A. (Archichauliodes) T91
                        |    |--A. (A.) anagaurus T91
                        |    |--A. (A.) deceptor T91
                        |    |--A. (A.) diversus T91
                        |    |--A. (A.) guttiferus T91
                        |    |--A. (A.) polypastus T91
                        |    `--A. (A.) uncinatus T91
                        `--A. (Apochauliodes) cervulus T91

*Type species of generic name indicated

REFERENCES

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

[H74] Helson, G. A. H. 1974. Insect Pests: Identification, life history, and control of pests of farms, horticulture, gardens, and public health. A. R. Shearer, Government Printer: Wellington (New Zealand).

[HR11] Heraty, J., F. Ronquist, J. M. Carpenter, D. Hawks, S. Schulmeister, A. P. Dowling, D. Murray, J. Munro, W. C. Wheeler, N. Schiff & M. Sharkey. 2011. Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution 60: 73–88.

[K-P83] Kukalová-Peck, J. 1983. Origin of the insect wing and wing articulation from the arthropodan leg. Canadian Journal of Zoology 61: 1618–1669.

[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).

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[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.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[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.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[R70] Riek, E. F. 1970. Megaloptera (alderflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 465–471. Melbourne University Press.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

[TW05] Terry, M. D., & M. F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240–257.

[T91] Theischinger, G. 1991. Megaloptera (alderflies, dobsonflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 516–520. Melbourne University Press: Carlton (Victoria).

[VM20] Vasilikopoulos, A., B. Misof, K. Meusemann, D. Lieberz, T. Flouri, R. G. Beutel, O. Niehuis, T. Wappler, J. Rust, R. S. Peters, A. Donath, L. Podsiadlowski, C. Mayer, D. Bartel, A. Böhm, S. Liu, P. Kapli, C. Greve, J. E. Jepson, X. Liu, X. Zhou, H. Aspöck & U. Aspöck. 2020. An integrative phylogenomic approach to elucidate the evolutionary history and divergence times of Neuropterida (Insecta: Holometabola). BMC Evolutionary Biology 20: 64.

[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.

[YS01] Yoshizawa, K., & T. Saigusa. 2001. Phylogenetic analysis of paraneopteran orders (Insecta: Neoptera) based on forewing base structure with comments on monophyly of Auchenorrhycha (Hemiptera). Systematic Entomology 26: 1–13.

[ZS10] Zborowski, P., & R. Storey. 2010. A Field Guide to Insects in Australia 3^rd ed. Reed New Holland: Sydney.

Archaeognatha

Sea bristletails Petrobius maritimus, photographed by Jymm.

Belongs within: Panhexapoda.
Contains: Meinertellidae.

The Archaeognatha, bristletails, are a group of wingless insects with an arched thorax modified for jumping.

Characters (from Ferguson 1990): Body form subcylindrical, thorax strongly arched; head hypognathous; eyes large, contiguous, with hundreds of ommatidia; vertex greatly reduced; frons large, with median ocellus; lateral ocelli large; mandibles primitive, consisting of a basal part with a single point of articulation to the cranium and a distal part with separate molar and incisor processes; palpus of maxillae with seven segments; labium with palpus of three unequal segments, the distal not greatly expanded apically; thoracic terga with very large paranotal lobes against sides; all leg segments essentially cylindrical; meso— and metathoracic coxae typically with styli (may be absent on mesothoracic coxa or both); femur and tibia large; tarsus with three tarsomeres; abdominal segments II—VII covered ventrally by three distinct sclerites; an anteromedial sternite and two lateral coxopodites; coxopodites of abdominal segments II—IX each with a posterolateral stylus; posterior edge of segments I—VII typically with one or two pairs of exsertile vesicles mesad of the styli.

<==Archaeognatha [Machilida, Machiloidea, Microcoryphia, Monocondylia]
    |  i. s.: Triassomachilis Sharov 1948 KS02 [Triassomachilidae GE05]
    |           `--T. uralensis Sharov 1948 K03
    |         Ditrigoniophthalmus oreophilus SBG11
    |--Dasyleptidae [Cercopodata, Monura] R02
    |    |--Leoidodasypus sharoui RJ93
    |    `--Dasyleptus Brongniart 1885 FT05
    |         |--D. brongniarti Sharov 1957 K-P83
    |         `--D. sharovi GE05
    `--+--Meinertellidae R02
       `--Machilidae R02
            |--Dilta littoralis GEW01, MG06
            |--Petridiobus arcticus F90
            |--Mesomachilis nearcticus F90
            |--Mixomachilis remingtoni F90
            |--Neomachilis halophilus F90
            |--Trigoniophthalmus Verhoeff 1910 F90
            |    `--T. alternatus (Silvestri 1904) F90
            |--Pedetontus Silvestri 1911 F90
            |    |--P. saltator Wygodzinsky & Schmidt 1980 F90
            |    `--P. unimaculatus R02
            |--Machilis Latreille 1802 L02
            |    |--*M. polypoda [=Lepisma polypoda] L02
            |    |--M. strenuva F90
            |    `--M. variabilis LNM91
            `--Petrobius Leach 1908 F90
                 |--P. brevistylis Carpenter 1913 [incl. P. canadensis Paclt 1969] F90
                 |--P. calcaratus Silvestri 1911 F90
                 |--P. californicus Silvestri 1911 F90
                 |--P. longicaudatus K-P83
                 |--P. maritimus Leach 1809 F90
                 |--P. persquamosus Silvestri 1911 F90
                 |--P. submutans Silvestri 1911 F90
                 `--P. superior Silvestri 1911 F90

*Type species of generic name indicated

REFERENCES

[FT05] Fayers, S. R., & N. H. Trewin. 2005. A hexapod from the Early Devonian Windyfield Chert, Rhynie, Scotland. Palaeontology 48 (5): 1117–1130.

[F90] Ferguson, L. M. 1990. Insecta: Microcoryphia and Thysanura. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 935-949. John Wiley & Sones: New York.

[GEW01] Giribet, G., G. D. Edgecombe & W. C. Wheeler. 2001. Arthropod phylogeny based on eight molecular loci and morphology. Nature 413: 157-161.

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

[K03] Kluge, N. J. 2003. About evolution and homology of genital appendages of insects. Trudy Russkogo Entomologicheskogo Obshestva [Proceedings of the Russian Entomological Society] 74: 3–16.

[KS02] Kluge, N. Yu., & N. D. Sinitshenkova. 2002. Order Ephemerida Latreille, 1810. The true mayflies (=Ephemeroptera Hyatt et Arms, 1891 (s. l.); =Euephemeroptera Kluge, 2000. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 89–97. Kluwer Academic Publishers: Dordrecht.

[K-P83] Kukalová-Peck, J. 1983. Origin of the insect wing and wing articulation from the arthropodan leg. Canadian Journal of Zoology 61: 1618–1669.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[LNM91] Lawrence, J. F., E. S. Nielsen & I. M. Mackerras. 1991. Skeletal anatomy and key to orders. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 3–32. Melbourne University Press: Carlton (Victoria).

[MG06] Mallatt, J., & G. Giribet. 2006. Further use of nearly complete 28S and 18S rRNA genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Molecular Phylogenetics and Evolution 40: 772–794.

[R02] Rasnitsyn, A. P. 2002. Subclass Lepismatona Latreille, 1804. The wingless insects (=Thysanura Latreille 1796, s. l.) In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 69–74. Kluwer Academic Publishers: Dordrecht.

[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.

[SBG11] Staniczek, A. H., G. Bechly & R. J. Godunko. 2011. Coxoplectoptera, a new fossil order of Palaeoptera (Arthropoda: Insecta), with comments on the phylogeny of the stem group of mayflies (Ephemeroptera). Insect Systematics and Evolution 42: 101–138.

Meinertellidae

Rock bristletail Machiloides banksi, copyright Katja Schulz.

Belongs within: Archaeognatha.

The Meinertellidae are a group of bristletails found primarily in the Southern Hemisphere, characterised by very small abdominal sterna which only protrude slightly, if at all, between the coxites, no scales on the legs, scape and pedicel, and male terminalia with a very short penis and lacking parameres (Watson & Smith 1991).

<==Meinertellidae [Meinertellinae] R02
    |--Neomachilellus F90
    |--Megalopsobius W39
    |--Machilontus W39
    |--Meinertellus cundinamarcensis W39, SC17
    |--Meinertelloides W39
    |--Macropsontus W39
    |--Machilellus W39
    |--Allomachilis Silvestri 1904 W39
    |    `--A. froggatti Silvestri 1904 W39
    |--Hypomachilodes Silvestri 1911 F90
    |    `--H. texanus F90
    |--Machilinus Silvestri 1904 F90
    |    |--M. aurantiacus F90
    |    `--M. nevadensis Sweetman 1937 S37
    |--Nesomachilis WS91
    |    |--N. australica WS91
    |    |--N. maoricus A99 [=Machiloides maoricus W39]
    |    |--N. queenslandica WS91
    |    `--N. tamborina WS91
    `--Machiloides Silvestri 1904 F90
         |--M. australicus Wom. 1938 W39
         |--M. banksi Silvestri 1911 F90
         |--M. hickmani Womersley 1939 W39
         |--M. petauristes Wygodzinsky & Schmidt 1980 F90
         `--M. sarasini [incl. M. s. var. montivaga, M. s. var. profuga] W39

*Type species of generic name indicated

REFERENCES

[A99] Ax, P. 1999. Das System der Metazoa II. Ein Lehrbuch der phylogenetischen Systematik. Gustav Fisher Verlag: Stuttgart (transl. 2000. Multicellular Animals: The phylogenetic system of the Metazoa vol. 2. Springer).

[F90] Ferguson, L. M. 1990. Insecta: Microcoryphia and Thysanura. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 935–949. John Wiley & Sones: New York.

[SC17] Schwentner, M., D. J. Combosch, J. P. Nelson & G. Giribet. 2017. A phylogenomic solution to the origin of insects by resolving crustacean-hexapod relationships. Current Biology 27: 1818–1824.

[S37] Sweetman, H. L. 1937. A new species of Machilinus (Thysanura: Machilidae). Proceedings of the Entomological Society of Washington 39 (8): 214–216.

[WS91] Watson, J. A. L., & G. B. Smith. 1991. Archaeognatha. Microcoryphia (bristletails). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 272–274. Melbourne University Press: Carlton (Victoria).

[W39] Womersley, H. 1939. Primitive Insects of South Australia: Silverfish, springtails, and their allies. South Australian Branch of the British Science Guild: Adelaide.