Palaeodictyoptera

The Palaeodictyoptera are an extinct order of medium-sized to very large, primitive Palaeozoic paleopterous insects. They are informative about the evolution of wings in insects.

Overview

They were characterized by beak-like mouthparts, used to pierce plant tissues for feeding. There is a similarity between their fore- and hindwings, and an additional pair of winglets on the prothorax, in front of the first pair of wings. They are known as "six-winged insects" because of the presence of a pair of wings on each of the thoracic segments. Their winglets provide clues to the origins of the first insect wings.

The mouthparts were elongated, and included sharp piercing stylets, and possibly a sucking pump-like organ. Unlike modern sucking insects, such as the Hemipterans, the mouthparts were held vertically below the head, or projected forwards. They probably used these organs to suck juices from plants, although some may have been ectoparasites, or predators.

Some types attained huge size. For example, Mazothairos is estimated to have a wingspan of about . Another distinctive feature was the presence of unusually long cerci, about twice the length of the abdomen.

The Palaeodictyoptera are a paraphyletic assemblage of basal palaeodictyopteroidean insects, rather than a clade, because they gave rise to other insect orders. They range in time from the Middle Carboniferous (late Serpukhovian or early Bashkirian in age) to the late Permian.

Taxonomy

The following taxa are recognized as being included in the order Palaeodictyoptera:

• Suborder †Dictyoneurina <small>Handlirsch 1906</small>
• Superfamily †Calvertielloidea <small>Martynov 1932</small>
• Family †Calvertiellidae <small>Martynov 1931</small>
• Genus †Calvertiella <small>Tillyard 1925</small>
• Genus †Carrizopteryx <small>Kukalová-Peck 1976</small>
• Genus †Lodetiella <small>Béthoux et al. 2007</small>
• Genus †Moravia <small>Kukalová 1964</small>
• Genus †Moraviptera <small>Kukalová 1955</small>
• Genus †Sharovia <small>Sinitshenkova 1977</small>
• Genus †Xiaheyanella <small>Fu et al. 2015</small>
• Superfamily †Dictyoneuroidea <small>Handlirsch 1906</small>
• Family †Dictyoneuridae <small>Handlirsch 1906</small>
• Genus †Arltia <small>Guthörl 1934</small>
• Genus †Cleffia <small>Guthörl 1931</small>
• Genus †Dictyoneura <small>Goldenberg 1854</small>
• Genus †Dictyoneurula <small>Handlirsch 1906</small>
• Genus †Goldenbergia <small>Scudder 1885</small>
• Genus †Hongius <small>Özdikmen 2008</small>
• Genus †Kallenbergia <small>Guthörl 1930</small>
• Genus †Macrodictya <small>Guthörl 1940</small>
• Genus †Microdictya <small>Brongniart 1894</small>
• Genus †Polioptenus <small>Scudder 1885</small>
• Genus †Rotundopteris <small>Guthörl 1940</small>
• Genus †Saarodictyum <small>Guthörl 1939</small>
• Genus †Sagenoptera <small>Handlirsch 1906</small>
• Genus †Schmidtopteron <small>Brauckmann and Hahn 1978</small>
• Genus †Siberiodictya <small>Sharov and Sinitshenkova 1977</small>
• Genus †Stenodictya <small>Brongniart 1894</small>
• Genus †Stenodictyoneura <small>Leriche 1911</small>
• Genus †Stilbocrocis <small>Handlirsch 1906</small>
• Genus †Titanodictya <small>Handlirsch 1906</small>

• Suborder †Frankenholziina <small>Guthörl 1962</small>
• Family †Hanidae <small>Kukalová-Peck 1975</small>
• Genus †Forcynthia <small>Sinitshenkova 2004</small>
• Genus †Hana <small>Kukalová-Peck 1975</small>

Superfamilies incertae sedis

• Superfamily †Breyerioidea <small>Handlirsch 1906</small>
• Family †Breyeriidae <small>Handlirsch 1906</small>
• Genus †Aviobreyeria <small>Prokop et al. 2013</small>
• Genus †Breyeria <small>de Borre 1875</small>
• Genus †Hasala <small>Brauckmann 1995</small>
• Genus †Jugobreyeria <small>Brauckmann et al. 1985</small>
• Genus †Megaptiloides <small>Handlirsch 1906</small>
• Genus †Stobbsia <small>Handlirsch 1908</small>
• Genus †Vernooijia <small>Prokop et al. 2018</small>
• Family †Cryptoveniidae <small>Bolton 1912</small>
• Genus †Cryptovenia <small>Bolton 1912</small>

• Superfamily †Eugereonoidea <small>Sinitshenkova 2002</small>
• Family †Archaemegaptilidae <small>Handlirsch 1919</small>
• Genus †Archaemegaptilus <small>Meunier 1908</small>
• Genus †Haseneura <small>Rosová et al. 2024</small>
• Genus †Osnabrugapteron <small>Rosová et al. 2022</small>
• Family †Eugereonidae <small>Handlirsch 1906</small>
• Genus †Dictyoptilus <small>Brongniart 1894</small>
• Genus †Eugereon <small>Dohrn 1866</small>
• Genus †Peromaptera <small>Brongniart 1894</small>
• Genus †Sandiella <small>Carpenter 1970</small>
• Genus †Valdeania <small>Teixeira 1941</small>
• Family †Graphiptilidae <small>Handlirsch 1906</small>
• Genus †Graphiptilus <small>Brongniart 1894</small>
• Genus †Patteiskya <small>Laurentiaux 1958</small>
• Genus †Rhabdoptilus <small>Brongniart 1894</small>
• Family †Jongmansiidae <small>Laurentiaux 1950</small>
• Genus †Jongmansia <small>Laurentiaux 1950</small>
• Genus †Katosaxoniapteron <small>Prokop et al. 2023</small>
• Family †Lithomanteidae <small>Handlirsch 1906</small>
• Genus †Archaeophasma <small>Matthew 1910</small>
• Genus †Catadyesthus <small>Handlirsch 1906</small>
• Genus †Dyscritus <small>Scudder 1868</small>
• Genus †Lithomantis <small>Woodward 1876</small>
• Genus †Lusiella <small>Laurentiaux and Teixeira 1958</small>
• Family †Lycocercidae <small>Handlirsch 1906</small>
• Genus †Apopappus <small>Handlirsch 1906</small>
• Genus †Lycocercus <small>Handlirsch 1906</small>
• Genus †Lycodemas <small>Carpenter and Richardson 1971</small>
• Genus †Lycodus <small>Carpenter 1992</small>
• Genus †Madera <small>Carpenter 1970</small>
• Genus †Notorachis <small>Carpenter and Richardson 1971</small>
• Family †Namuroningxiidae <small>Prokop and Ren 2007</small>
• Genus †Namuroningxia <small>Prokop and Ren 2007</small>
• Family †Polycreagridae <small>Handlirsch 1906</small>
• Genus †Polycreagra <small>Handlirsch 1906</small>
• Family †Protagriidae <small>Handlirsch 1906</small>
• Genus †Protagrion <small>Brongniart 1894</small>
• Family †Synarmogidae <small>Handlirsch 1910</small>
• Genus †Synarmoge <small>Handlirsch 1910</small>
• Family †Tchirkovaeidae <small>Sinitshenkova 1979</small>
• Genus †Paimbia <small>Sinitshenkova 1979</small>
• Genus †Tchirkovaea <small>Zalessky 1931</small>

• Superfamily †Spilapteroidea <small>Brongniart 1894</small>
• Family †Fouqueidae <small>Handlirsch 1906</small>
• Genus †Fouquea <small>Brongniart 1894</small>
• Genus †Neofouquea <small>Carpenter 1967</small>
• Family †Mecynostomatidae <small>Kukalová 1969</small>
• Genus †Mecynostomata <small>Metcalf 1952</small>
• Family †Spilapteridae <small>Brongniart 1894</small>
• Genus †Abaptilon <small>Zalessky 1946</small>
• Genus †Baeoneura <small>Sinitshenkova 1977</small>
• Genus †Becquerelia <small>Brongniart 1894</small>
• Genus †Bizarrea <small>Prokop et al. 2015</small>
• Genus †Delitzschala <small>Brauckmann and Schneider 1996</small>
• Genus †Dunbaria <small>Tillyard 1924</small>
• Genus †Epitethe <small>Handlirsch 1906</small>
• Genus †Epithele <small>Handlirsch 1906</small>
• Genus †Homaloneura <small>Brongniart 1885</small>
• Genus †Lamproptilia <small>Brongniart 1885</small>
• Genus †Mcluckiepteron <small>Richardson 1956</small>
• Genus †Neuburgia <small>Martynov 1930</small>
• Genus †Palaeoptilus <small>Brongniart 1894</small>
• Genus †Paradunbaria <small>Sharov and Sinitshenkova 1977</small>
• Genus †Permiakovia <small>Martynov 1940</small>
• Genus †Saxonyptilus <small>Rosová et al. 2022</small>
• Genus †Sinodunbaria <small>Li et al. 2013</small>
• Genus †Spilaptera <small>Brongniart 1885</small>
• Genus †Spiloptilus <small>Handlirsch 1906</small>
• Genus †Tectoptilus <small>Kukalová 1969</small>
• Genus †Tytthospilaptera <small>Liu et al. 2015</small>
• Genus †Vorkutoneura <small>Sinitshenkova 1977</small>

Families incertae sedis

• Family †Elmoboriidae <small>Carpenter 1976</small>
• Genus †Elmoboria <small>Carpenter 1976</small>
• Genus †Oboria <small>Kukalová 1960</small>
• Family †Eubleptidae <small>Handlirsch 1906</small>
• Genus †Eubleptus <small>Handlirsch 1906</small>
• Family †Homoiopteridae <small>Handlirsch 1906</small>
• Genus †Adolarryia <small>Kukalová-Peck and Richardson 1983</small>
• Genus †Ametretus <small>Handlirsch 1911</small>
• Genus †Amousus <small>Handlirsch 1911</small>
• Genus †Anglopterum <small>Prokop et al. 2006</small>
• Genus †Boltonia <small>Pruvost 1919</small>
• Genus †Homoioptera <small>Brongniart 1894</small>
• Genus †Larryia <small>Kukalová-Peck and Richardson 1983</small>
• Genus †Mazonopterum <small>Kukalová-Peck and Richardson 1983</small>
• Genus †Mazothairos <small>Kukalová-Peck and Richardson 1983</small>
• Genus †Ostrava <small>Kukalová 1960</small>
• Genus †Paraostrava <small>Prokop and Nel 2004</small>
• Genus †Parathesoneura <small>Sinitshenkova 1977</small>
• Genus †Pharciphyzelus <small>Beckemeyer and Engel 2011</small>
• Genus †Scepasma <small>Handlirsch 1911</small>
• Genus †Thesoneura <small>Carpenter 1943</small>
• Genus †Turneropterum <small>Kukalová-Peck and Richardson 1983</small>
• Family †Homothetidae <small>Scudder 1885</small>
• Genus †Homothetus <small>Scudder 1867</small>
• Family †Megaptilidae <small>Handlirsch 1906</small>
• Genus †Lithoptilus <small>Lameere 1917</small>
• Genus †Megaptilus <small>Brongniart 1885</small>
• Family †Psychroptilidae <small>Riek 1976</small>
• Genus †Psychroptilus <small>Riek 1976</small>
• Family †Straeleniellidae <small>Laurentiaux-Vieira and Laurentiaux 1986</small>
• Genus †Straeleniella <small>Laurentiaux-Vieira and Laurentiaux 1986</small>

Genera incertae sedis

• Genus †Achaeoptilus <small>Scudder 1881</small>
• Genus †Aenigmatopsis <small>Handlirsch 1937</small>
• Genus †Althansia <small>Guthörl 1934</small>
• Genus †Anagesthes <small>Handlirsch 1906</small>
• Genus †Anthracosta <small>Pruvost 1930</small>
• Genus †Asiodictya <small>Rohdendorf 1961</small>
• Genus †Bathytaptus <small>Handlirsch 1906</small>
• Genus †Bojoptera <small>Kukalová 1958</small>
• Genus †Brongniartites <small>Handlirsch 1919</small>
• Genus †Brongniartula <small>Handlirsch 1919</small>
• Genus †Commentryites <small>Handlirsch 1922</small>
• Genus †Compsoneura <small>Brongniart 1894</small>
• Genus †Delafondia <small>Langiaux and Parriat 1975</small>
• Genus †Diexodus <small>Handlirsch 1911</small>
• Genus †Eumecoptera <small>Handlirsch 1906</small>
• Genus †Eurydictyella <small>Carpenter 1986</small>
• Genus †Eurythmopteryx <small>Handlirsch 1906</small>
• Genus †Gegenemene <small>Handlirsch 1906</small>
• Genus †Haplophlebium <small>Scudder 1867</small>
• Genus †Heolus <small>Handlirsch 1906</small>
• Genus †Idoptilus <small>Wootton 1972</small>
• Genus †Kansasia <small>Tillyard 1937</small>
• Genus †Mammia <small>Handlirsch 1906</small>
• Genus †Mecynoptera <small>Handlirsch 1904</small>
• Genus †Mecynostomites <small>Handlirsch 1919</small>
• Genus †Monsteropterum <small>Kukalová-Peck 1972</small>
• Genus †Palaiotaptus <small>Handlirsch 1906</small>
• Genus †Palapteris <small>Guthörl 1940</small>
• Genus †Paraboltonia <small>Handlirsch 1922</small>
• Genus †Paramecynostoma <small>Handlirsch 1919</small>
• Genus †Paramegaptilus <small>Handlirsch 1906</small>
• Genus †Platephemera <small>Scudder 1867</small>
• Genus †Propalingenia <small>Handlirsch 1906</small>
• Genus †Pseudomecynostoma <small>Handlirsch 1919</small>
• Genus †Pteronidia <small>Bolton 1912</small>
• Genus †Pyebrodia <small>Brauckmann and Herd 2002</small>
• Genus †Rochdalia <small>Woodward 1913</small>
• Genus †Saarlandia <small>Guthörl 1930</small>
• Genus †Sabitaptus <small>Pruvost 1930</small>
• Genus †Schedoneura <small>Carpenter 1963</small>
• Genus †Severinopsis <small>Kukalová 1958</small>
• Genus †Synaptoneura <small>Pruvost 1919</small>
• Genus †Titanophasma <small>Brongniart 1882</small>
• Genus †Turnbullia <small>Richardson 1956</small>

References

External links

• Paleodictyoptera at the Tree of Life project (list of taxa)

Further reading

• Carpenter, F. M. 1992. Superclass Hexapoda. Volume 3 of Part R, Arthropoda 4; Treatise on Invertebrate Paleontology, Boulder, Colorado, Geological Society of America.