Hymenoptera

Larva of horned hickory sawfly Macroxyela, photographed by Charley Eiseman.

Belongs within: Holometabola.
Contains: Tenthredinoidea, Siricoidea, Vespina.

The Hymenoptera include the wasps and their derivative taxa such as bees and ants. Members of the group are primitively characterised by the presence of hamuli, small hooks on the anterior margin of the hind wing that hook onto the fore wing and hold the wings locked together. The earliest known members of the Hymenoptera in the Late Triassic are assigned to the Xyelidae, though it is uncertain whether all members of the fossil subfamily Archexyelinae form a clade with modern xyelids (Grimaldi & Engel 2005). Larvae of the Xyelidae are herbivorous, with Xyelinae feeding on developing cones or shoots of conifers, and Macroxyela on leaves of elms or walnuts. The Xyelidae have been regarded in the past as the sister group of all other Hymenoptera but a molecular phylogenetic analysis of the order by Peters et al. (2017) united them in a clade Eusymphyta with the other ectophytophagous wasp groups Pamphilioidea and Tenthredinoidea. The Pamphilioidea are a relatively small Holarctic group of sawflies with an enlarged, almost prognathous, head that is widest near the clypeus.

The remaining Hymenoptera formed a clade Unicalcarida combining the carnivorous wasps of the Vespina with the endophytophagous groups Xiphydriidae, Siricoidea and Cephoidea, whose larvae develop in burrows within their plant hosts.

Synapomorphies (from Grimaldi & Engel 2005): Hamuli present; protibial spur with velum; haplodiploid sex determination.

Hymenoptera (see below for synonymy)
    |--Eusymphyta PK17
    |    |--Pamphilioidea [Megalodontoidea] PK17
    |    |    |  i. s.: Parapamphiliidae GE05
    |    |    |         Acantholyda hieroglyphica PK17
    |    |    |--Pamphiliidae R02b
    |    |    |    |--Atocus Z02b
    |    |    |    |--Pamphilius Latreille 1802 L02
    |    |    |    |    `--*P. sylvaticus (Linnaeus 1758) L02, L58 [=Tenthredo sylvatica L02]
    |    |    |    |--Cephalcia abietis HR11
    |    |    |    `--Onycholyda amplecta HR11
    |    |    `--+--Xyelydidae R02b
    |    |       `--+--Praesiricidae R02b
    |    |          `--Megalodontes Latreille 1802 L02 [Megalodontesidae R02b]
    |    |               |--*M. cephalotes [=Tenthredo cephalotes] L02
    |    |               `--M. spireae MS01
    |    `--+--Tenthredinoidea PK17
    |       `--Xyelidae [Archihymenoptera, Xyeloidea] PK17
    |            |  i. s.: Spathoxyela pinicola R02b
    |            |         Angaridyela vitimica GE05
    |            |         Eoxyela atra R02a
    |            |         Megaxyela gigantea Mocsary 1909 Z02b, I92
    |            |         Angarixyela NR13
    |            |--Xyela [Xyelinae, Xyelini] R02a
    |            |    |--X. alpigena PK17
    |            |    `--X. julii HR11
    |            |--Macroxyela [Macroxyelinae] NPH04
    |            |    `--M. ferruginea HR11
    |            `--Archexyelinae GE05
    |                 |--Archexyela crosbyi R70
    |                 `--Madygenius GE05
    `--Unicalcarida PK17
         |  i. s.: Daohugoa Rasnitsyn & Zhang 2004 [Daohugoidae] RZ04
         |           `--*D. tobiasi Rasnitsyn & Zhang 2004 RZ04
         |--+--Xiphydriidae [Xiphydrioidea] HR11
         |  |    |--Derycyrtinae N91
         |  |    |    |--Austrocyrta australiensis N91
         |  |    |    `--Derecyrta circularis HR11
         |  |    `--Xiphydriinae N91
         |  |         |--Rhysacephala N91
         |  |         `--Xiphydria Latreille 1802 L02
         |  |              |--*X. camelus [=Sirex camelus] L02
         |  |              `--X. prolongata HR11
         |  `--Siricoidea HR11
         `--+--Cephoidea GE05
            |    |--Sepulcidae GE05
            |    `--Cephidae HR11
            |         |--Mesocephus sibiricus Z02b
            |         |--Cephus Latreille 1802 L02
            |         |    |--*C. pygmaeus [=Sirex pygmaeus] L02
            |         |    |--C. cinctus PK17
            |         |    |--C. flavicornis Lucas 1849 E12
            |         |    `--C. spinipes PK17
            |         `--Hartigia Schioedte 1838 HR11, L59
            |              `--H. trimaculata HR11
            `--Vespina HR11

Hymenoptera incertae sedis:
  Stenopolybia Z02a
  Orisema uchancoi H03
  Rielia manticida A71
  Euryxyela Hong 1984 [=Xaxexis Pagliano & Scaramozzino 1990] NPH04
  Ophrynopus schauinslandi G27
  Amalthea Rafinesque 1815 BR05
  Giraudia Foerster 1868 BR05
  Janus Stephens 1835 BR05
  Dineura virididorsata (Retzius 1783) HP-W05
  Lionotus delphinalis A-B91
  Schizonotus Ratzeburg 1852 H11
  Myzina E12
    |--M. guerinii Lucas 1849 E12
    `--M. oraniensis Lucas 1849 E12
  Tarpa levaillantii Lucas 1849 E12
  Scleroderma ruficornis Lucas 1849 E12
  ‘Promachus’ Cresson 1887 nec Loew 1848 nec Stål 1875 Z01
  Myzinum quinquecinctum (Fabricius 1775) [=Tiphia quinquecincta, Sphex quinquecinctus] BM76
  Triassoxyela foveolata PK17
  Gigantoxyela quadrifurcata NR13
  Baeosomus Thomson 1891 Z93
  Diastrophus Hartig 1840 Z94
  Patasson nitens M94
  Lophyrus Latreille 1802 L02
    `--*L. pini (Linnaeus 1758) L02, L58 [=Tenthredo pini L02]

Hymenoptera [Chalastrogastra, Neohymenoptera, Siricina, Siricomorpha, Symphyta, Tenthredines, Urocerata, Vespida, Vespidea]

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[A-B91] Athias-Binche, F. 1991. Ecology and evolution of phoresy in mites. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 27–41. SPB Academic Publishing: The Hague.

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

[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.

[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.

[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.

[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.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[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.

[L59] Lopes, H. de S. 1959. A revision of Australian Sarcophagidae (Diptera). Studia Ent. 2 (1–4): 33–67.

[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.

[MS01] Mocsáry, A., & V. Szépligeti. 1901. Hymenopterák [Hymenopteren]. 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. 121–169. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

[NPH04] Nel, A., J. F. Petrulevicius & M. Henrotay. 2004. New Early Jurassic sawflies from Luxembourg: the oldest record of Tenthredinoidea (Hymenoptera: “Symphyta”). Acta Palaeontologica Polonica 49 (2): 283–288.

[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.

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

[R02b] Rasnitsyn, A. P. 2002b. Superorder Vespidea Laicharting, 1781. Order Hymenoptera Linné, 1758 (=Vespida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 242–254. Kluwer Academic Publishers: Dordrecht.

[RZ04] Rasnitsyn, A. P., & H. Zhang. 2004. A new family, Daohugoidae fam. n., of siricomorph hymenopteran (Hymenoptera=Vespida) from the Middle Jurassic of Daohugou in Inner Mongolia (China). Proceedings of the Russian Entomological Society, St. Petersburg 75 (1): 12–16.

[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.

[Z02a] Zherikhin, V. V. 2002a. Insect trace fossils. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 303–324. Kluwer Academic Publishers: Dordrecht.

[Z02b] Zherikhin, V. V. 2002b. Ecological history of the terrestrial insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 331–388. 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.

Last updated: 3 October 2017.