Pterygota

Holasicia rasnitsyni, from Lutz Koch.

Belongs within: Insecta.
Contains: Palaeodictyoptera, Diaphanopterodea, Megasecoptera, Ephemeroptera, Odonatoptera, Polyneoptera, Paraneoptera, Holometabola.

The Pterygota are the winged insects, the first group of animals to have evolved the power of flight. Ancestrally, two pairs of wings are present, though one pair of wings has been lost or modified in a number of sublineages.

Synapomorphies (from Grimaldi & Engel 2005): Eversible vesicles absent; transverse stipital muscle present; pleural apophyses fused with sternal apophyses; pterothoracic walls strengthened by pleural sulcus; two coxal proprioreceptor organs present; corporotentorium present; two pairs of wings present; sperm transfer through copulation.

<==Pterygota (see below for synonymy)
    |--Megalometer lata R02b
    |--Heterologellus R02b
    |--Schuchertiella R02a
    |--Merlebachia R02a
    |--Anthracotremma [Anthracotremmatidae] R02b
    |    `--A. robusta R02b
    |--Evenka [Evenkidae] R02b
    |    `--E. archaica R02a
    `--+--+--Palaeodictyopteroidea [Dictyoneuridea, Palaeodictyopterida] GC12
       |  |    |--Palaeodictyoptera GE05
       |  |    `--+--Diaphanopterodea GE05
       |  |       `--Megasecoptera GE05
       |  `--+--Ephemeroptera GE05
       |     `--Syntonopterodea [Syntonopterida] GE05
       |          |  i. s.: Aedoeophasma anglica R02c
       |          |         Miracopteron mirabile R02c
       |          |--Bojophlebia [Bojophlebiidae] R02c
       |          |    `--B. procopi R02c
       |          `--Syntonopteridae GE05
       |               |--Syntonoptera R02c
       |               `--Lithoneura GE05
       |                    |--L. lameerei GE05
       |                    `--L. mirifica R02c
       `--+--Triplosoba [Protephemerida, Protephemeroptera, Triplosobida, Triplosobidae] GE05
          |    `--*T. pulchella R02c
          `--Metapterygota GE05
               |--Odonatoptera GE05
               `--Neoptera [Caloneuridea, Planoneoptera] GE05
                    |  i. s.: Cymbopsis R02d [Cymbopsidae]
                    |           `--C. excelsa Kukalová 1965 BN02
                    |         Glaphyrokoris R02d
                    |         Cymenophlebia R02d
                    |         Kliveria R02d
                    |         Endoiasmus R02d
                    |--+--Polyneoptera GE05
                    |  `--Eumetabola GE05
                    |       |--Paraneoptera GE05
                    |       `--Holometabola GE05
                    `--+--Blattinopseida [Blattinopseidae] BN02
                       |    |--Blattinopsis R02d
                       |    `--Glaphyrophlebia R02d
                       |         |--G. subcostalis R02d
                       |         `--G. uralensis R02d
                       `--Paoliida [Protoptera] R02b
                            |--Stygne [Stygneidae] RZ02
                            |    `--S. roemeri R02b
                            `--Paoliidae BN02
                                 |--Zdenekia grandis Kukalová 1958 K-P83
                                 |--Olinka R02b
                                 |--Kemperala hagenensis GE05, R02b
                                 |--Paolia R02b
                                 |--Pseudofouquea R02b
                                 |--Sustaia R02b
                                 `--Holasicia R02b
                                      |--H. rasnitsyni R02b
                                      `--H. vetula R02b

Pterygota incertae sedis:
  Kelleropteron R02a
  Propachytylopsis R02b
  Kochopteron R02a
  Herbstiala R02a
  Hadentomum Handlirsch 1906 [Hadentomidae] BN05
  Bardapteron Si02
  Mesoforficula Sh02
  Forficularia Sh02
  Permoneuridae Si02
  Mesogrammatidae R02d
  Rhyniognatha hirsti Tillyard 1928 FT05
  Stereopteron Carpenter 1950 [Stereopteridae] BN02
  Archaeologus falcatus Handlirsch 1906 BN02
  Proedischia [Proedischiidae] BN02
    `--P. mezzalarai Pinto & Pinto de Ornellas 1978 BN02
  Rigattoptera [Rigattopteridae] BN02
    `--R. ornellasae Pinto 1996 BN02
  Platyphlebopteron Germer 1971 BN02
    `--P. jakobyi Germer 1971 BN02
  Ferganiinae BN02
    |--Fergania Sharov 1968 BN02
    `--Parafergania Gorokhov 1987 BN02
         `--P. sharovi Gorokhov 1987 BN02
  Vitimia Sharov 1968 [Vitimiidae] BN02
  Strudiella Garrouste, Clément et al. 2012 GC12
    `--*S. devonica Garrouste, Clément et al. 2012 [=Strudiella antennata (l. c.)] GC12

Pterygota [Ephemeridea, Exopterygota, Hydropalaeoptera, Libelluliformes, Palaeoptera, Panephemeroptera, Protorthoptera, Scarabaeona, Scarabaeones, Subulicornes]

*Type species of generic name indicated

REFERENCES

[BN02] Béthoux, O., & A. Nel. 2002. Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa 96: 1-88.

[BN05] Béthoux, O., A. Nel, J. Lapeyrie & G. Gand. 2005. New data on Paleozoic grylloblattid insects (Neoptera). Journal of Paleontology 79 (1): 125-138.

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

[GC12] Garrouste, R., G. Clément, P. Nel, M. S. Engel, P. Grandcolas, C. D'Haese, L. Lagebro, J. Denayer, P. Gueriau, P. Lafaite, S. Olive, C. Prestianni & A. Nel. 2012. A complete insect from the Late Devonian period. Nature 488: 82-85.

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

[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.

[R02a] Rasnitsyn, A. P. 2002a. Subclass Scarabaeona Laicharting, 1781. The winged insects (=Pterygota Lang, 1888). In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 75-83. Kluwer Academic Publishers: Dordrecht.

[R02b] Rasnitsyn, A. P. 2002b. Infraclass Scarabaeones Laicharting, 1781. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 84-85. Kluwer Academic Publishers: Dordrecht.

[R02c] Rasnitsyn, A. P. 2002c. Cohors Libelluliformes Laicharting, 1781 (=Subulicornes Latreille, 1807, =Hydropalaeoptera Rohdendorf, 1968). In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 85-89. Kluwer Academic Publishers: Dordrecht.

[R02d] Rasnitsyn, A. P. 2002d. Cohors Cimiciformes Laicharting, 1781. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 104-115. Kluwer Academic Publishers: Dordrecht.

[RZ02] Rasnitsyn, A. P., & V. V. Zherikhin. 2002. Impression fossils. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 437-444. Kluwer Academic Publishers: Dordrecht.

[Sh02] Shcherbakov, D. E. 2002. Order Forficulida Latreille, 1810. The earwigs and protelytropterans (=Dermaptera DeGeer, 1773 +Protelytroptera Tillyard, 1931). In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 288-291. Kluwer Academic Publishers: Dordrecht.

[Si02] Sinitshenkova, N. D. 2002. Superorder Dictyoneuridea Handlirsch, 1906 (=Palaeodictyopteroidea). In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 115-124. Kluwer Academic Publishers: Dordrecht.