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

Suborder †Frankenholziina Guthörl 1962 Family †Hanidae Kukalová-Peck 1975 Genus †Forcynthia Sinitshenkova 2004 Genus †Hana Kukalová-Peck 1975

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

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

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

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

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

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.