Holometabola

Fossil wing of stem-hymenopteran Avioxyela gallica, from Nel et al. (2013). Scale bar = 1 mm.

Belongs within: Pterygota.
Contains: Neuropterida, Strepsiptera, Coleoptera, Panorpoidea, Eusymphyta, Unicalcarida.

The Holometabola are the major clade of insects, accounting for nearly 85% of all insect species, characterised by the evolution of a complete metamorphosis with distinct larval, pupal and adult stages. The larval stage is more or less soft-bodied and lacks external wing buds, with the larval cuticle being more or less entirely replaced during the quiescent pupal stage by adult cuticle derived specialised pockets of epidermal cells called imaginal discs or anlage (Grimaldi & Engel 2005).

<==Holometabola [Endopterygota, Halteria, Meronida, Oligoneoptera, Panorpatae, Scarabaeiformes]
    |  i. s.: Mormolucoides P02
    |         Baryshnyala Ilger & Brauckmann 2011 [Baryshnyalidae] IB11
    |           `--*B. occulta Ilger & Brauckmann 2011 IB11
    |--Metabolarva Kirejtshuk, Prokin et al. in Nel, Roques et al. 2013 NR13
    |    `--*M. bella Kirejtshuk, Prokin et al. in Nel, Roques et al. 2013 NR13
    |--Neuropteriformia A99
    |    |--Neuropterida GE05
    |    `--Coleopterida [Coleopteroidea] KN13
    |         |  i. s.: Coleopsis archaica Kirejtshuk et al. 2013 NR13
    |         |--Stephanastus Kirejtshuk & Nel in Nel, Roques et al. 2013 [Skleroptera, Stephanastidae] NR13
    |         |    `--*S. polinae Kirejtshuk & Nel. in Nel, Roques et al. 2013 NR13
    |         |--+--Strepsiptera KN13
    |         |  `--Coleoptera GE05
    |         `--Umenocoleidae KN13
    |              |--Umenocoleus Chen & T’an 1973 NR13
    |              `--Jantarimantis G03 [incl. Archimantis Vršanský 2002 V02]
    |                   `--J. zherikhini (Vršanský 2002) G03 [=*Archimantis zherikhini V02]
    `--Mecopteriformia A99
         |--Panorpoidea GE05
         `--+--Avioxyela Nel, Engel et al. in Nel, Roques et al. 2013 [Avioxyelidae] NR13
            |    `--*A. gallica Nel, Engel et al. in Nel, Roques et al. 2013 NR13
            `--Hymenoptera (see below for synonymy) GE05
                 |  i. s.: Stenopolybia Z02
                 |         Orisema uchancoi H03
                 |         Ophrynopus schauinslandi G27
                 |         Amalthea Rafinesque 1815 BR05
                 |         Giraudia Foerster 1868 BR05
                 |         Dineura virididorsata (Retzius 1783) HP-W05
                 |         Schizonotus Ratzeburg 1852 H11
                 |         Myzina E12
                 |           |--M. guerinii Lucas 1848 E12
                 |           `--M. oraniensis Lucas 1848 E12
                 |         Tarpa levaillantii Lucas 1849 E12
                 |         Scleroderma ruficornis Lucas 1849 E12
                 |         ‘Promachus’ Cresson 1887 nec Loew 1848 nec Stål 1875 Z01
                 |         Triassoxyela foveolata PK17
                 |         Gigantoxyela quadrifurcata NR13
                 |         Baeosomus Thomson 1891 Z93
                 |         Diastrophus Hartig 1840 Z94
                 |         Patasson nitens M94
                 |         Lyda A83
                 |           |--L. campestris R13
                 |           |--L. hypotrophica R13
                 |           `--L. sylvatica A83
                 |         Cephaleia F92
                 |           |--C. abietis F92
                 |           `--C. alashanica F92
                 |         Lygaeonematus laricis F92
                 |         Hoplocampa Hartig 1837 M99
                 |           |--H. chrysorrhaea (Klug 1816) M99
                 |           `--H. fulvicornis (Panzer 1801) [incl. H. rutilicornis (Klug 1816)] M99
                 |         Aprosthema Konow 1899 M99
                 |           `--A. melanura (Klug 1814) [incl. A. tarda (Klug 1814)] M99
                 |         Heptamelus Haliday 1855 M99
                 |           `--H. ochroleucus (Stephens 1835) M99
                 |         Strongylogaster Dahlbom 1835 M99
                 |           `--S. multifasciata (Geoffroy 1785) [incl. S. lineata (Christ 1791)] M99
                 |         Dulophanes Konow 1907 [incl. Nesoselandria Rohwer 1910] M99
                 |         Ardis Konow 1886 M99
                 |           `--A. pallipes (Serville 1823) [incl. A. brunniventris (Hartig 1837)] M99
                 |         Claremontia Rohwer 1909 M99
                 |           |--C. alternipes [=Monophadnoides alternipes] M99
                 |           |--C. confusa [=Monophadnoides confusa] M99
                 |           |--C. tenuicornis [=Monophadnoides tenuicornis] M99
                 |           `--C. waldheimii [=Monophadnoides waldheimii] M99
                 |         Fenusa Leach 1817 M99
                 |           `--F. pumila Leach 1817 [incl. F. pusilla (Lepeletier 1823)] M99
                 |         Fenusella Enslin 1912 [incl. Messa Leach 1817 (n. n.)] M99
                 |         Scolioneura Konow 1890 M99
                 |           |--S. betuleti (Klug 1916) M99
                 |           `--S. vicina Konow 1894 M99
                 |         Empria Lepeletier & Serville 1828 M99
                 |           |--E. abdominalis F92
                 |           |--E. candidata F92
                 |           |--E. klugii (Stephens 1835) [incl. E. waldstaetterense Liston 1980] M99
                 |           `--E. pulverata F92
                 |         Harpiphorus Hartig 1837 M99
                 |           `--H. lepidus (Klug 1818) M99
                 |         Apethymus Benson 1939 M99
                 |           |--A. braccatus (Gmelin 1790) M99
                 |           `--A. serotinus (Müller 1776) M99
                 |         Siobla Cameron 1877 M99
                 |           `--S. sturmii (Klug 1817) M99
                 |         Aglaostigma Kirby 1882 M99
                 |           |--A. fulvipes F92
                 |           `--A. nebulosum (André 1881) M99
                 |         Elinora Benson 1946 [incl. Cuneala Zirngiebl 1956] M99
                 |           `--E. longipes (Konow 1886) (see below for synonymy) M99
                 |         Blankia Lacourt 1997 M99
                 |           `--*B. koehleri (Klug 1817) [=Allantus koehleri] M99
                 |         Sharliphora Wong 1969 M99
                 |           `--S. nigella (Förster 1854) [incl. S. ambigua (Fallén 1808) (preoc.)] M99
                 |         Konowia Brauns 1884 M99
                 |           `--K. betulae (Enslin 1911) [incl. K. guntionensis Zombori 1969] M99
                 |         Trachelus Jurine 1807 M99
                 |           `--T. troglodytus (Fabricius 1787) [=T. troglodyta] M99
                 |         Iota Saussure 1855 S00
                 |         Coeloides dendroctoni S00
                 |         Cephenopsis mirabilis Hong 1985 RJ93
                 |         Pararchexyelidae RJ93
                 |         Spategaster R13
                 |           |--S. blattarum R13
                 |           `--S. tricolor R13
                 |         Teras terminalis R13
                 |         Poecilosoma pulveratum R13
                 |         Agenia R13
                 |           |--A. intermedia R13
                 |           |--A. punctata R13
                 |           `--A. variegata R13
                 |         Pogonius variegatus R13
                 |         Tachus Jurine 1807 KM20
                 |--Eusymphyta PK17
                 `--Unicalcarida PK17

Elinora longipes (Konow 1886) [=Allantus longipes; incl. *Cuneala tricolor Zirngiebl 1956] M99

Hymenoptera [Chalastrogastra, Neohymenoptera, Orthandria, Piezata, Siricina, Siricomorpha, Symphyta, Tenthredines, Tenthredinetae, Urocerata, Vespida, Vespidea] GE05

*Type species of generic name indicated

REFERENCES

[A83] Adolph, E. 1883. Zur Morphologie der Hymenopterenflügel. Zugleich ein Beitrag zu den Fragen der Speciesbildung und des Atavismus. Verhandlungen der Kaiserlichen Leopoldinisch-Carolinischen Deutschen Akademie der Naturforscher [Nova Acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum] 46 (2): 41–132, pls 1–6.

[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).

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[F92] Fan Z. 1992. Key to the Common Flies of China 2^nd ed. Science Press: Beijing.

[G27] Gourlay, E. S. 1927. Notes on the New Zealand wood-wasp Ophrynopus schauinslandi Ashmead. Transactions and Proceedings of the New Zealand Institute 57: 691–693.

[G03] Grimaldi, D. 2003. A revision of Cretaceous mantises and their relationships, including new taxa (Insecta: Dictyoptera: Mantodea). American Museum Novitates 3412: 1–47.

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

[H11] Harvey, M. S. 2011. Notes on some Old World schizomids of the genera Ovozomus and Schizomus (Schizomida: Hubbardiidae). Records of the Western Australian Museum 26 (2): 202–208.

[H03] Heads, M. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311–449.

[HP-W05] Heitland, W., & H. Pschorn-Walcher. 2005. Biology and parasitoids of the peculiar alder sawfly, Platycampus luridiventris (Fallen) (Insecta, Hymenoptera, Tenthredinidae). Senckenbergiana Biologica 85 (2): 215–231.

[IB11] Ilger, J.-M., & C. Brauckmann. 2011. The smallest Neoptera (Baryshnyalidae fam. n.) from Hagen-Vorhalle (early Late Cretaceous: Namurian B; Germany). ZooKeys 130: 91–102.

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

[KM20] Kury, A. B., A. C. Mendes, L. Cardoso, M. S. Kury & A. A. Granado. 2020. WCO-Lite: Online world catalogue of harvestmen (Arachnida, Opiliones). Version 1.0—Checklist of all valid nomina in Opiliones with authors and dates of publication up to 2018. Published by the author: Rio de Janeiro.

[M99] Magis, N. 1999. Répertoire des mouches à scie reconnues en Belgique et au Grand-Duché de Luxembourg (Hymenoptera: Symphyta). Additions et corrections. Notes Fauniques de Gembloux 36: 85–93.

[M94] May, B. M. 1994. An introduction to the immature stages of Australian Curculionoidea. In: Zimmerman, E. C. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May pp. 365–728. CSIRO Australia.

[NR13] Nel, A., P. Roques, P. Nel, A. A. Prokin, T. Bourgoin, J. Prokop, J. Szwedo, D. Azar, L. Desutter-Grandcolas, T. Wappler, R. Garrouste, D. Coty, D. Huang, M. S. Engel & A. G. Kirejtshuk. 2013. The earliest known holometabolous insects. Nature 503: 257–261.

[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.

[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.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[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.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2^nd ed. CABI Publishing: Wallingford (UK).

[V02] Vršanský, P. 2002. Origin and the early evolution of mantises. AMBA Projekty 6 (1): 1–16.

[Z02] Zherikhin, V. V. 2002. Insect trace fossils. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 303–324. Kluwer Academic Publishers: Dordrecht.

[Z93] Zimmerman, E. C. 1993. Australian Weevils (Coleoptera: Curculionoidea) vol. 3. Nanophyidae, Rhynchophoridae, Erirhinidae, Curculionidae: Amycterinae, literature consulted. CSIRO Australia.

[Z94] Zimmerman, E. C. 1994. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May. CSIRO Australia.

[Z01] Zompro, O. 2001. A review of Eurycanthinae: Eurycanthini, with a key to genera, notes on the subfamily and designation of type species. Phasmid Studies 10 (1): 19–23.