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.

Pergidae

Spitfire Perga affinis larvae, copyright Gordon Smith.

Belongs within: Tenthredinoidea.

The Pergidae are a group of sawflies most diverse in Australia and South America with fewer species in North America and Asia. They are divided between numerous subfamilies with the largest species belonging to the Australasian Perginae. Larvae of this subfamily are often notable defoliators of Eucalyptus (Naumann 1991).

Characters (from Naumann 1991): Adult 2.8-30 mm; antennae 5-24-segmented; head sometimes with complete hypostomal bridge; neck short; fore wing without crossvein 2r, fore tibia with one or two apical spurs. Larva with 3-5-segmented legs with claws; abdominal prolegs on segments 2-6, 2-8 or 3-8 and 10, sometimes only on 10 or completely absent; stemmata present; antennae variable from disc-like to multisegmented.

<==Pergidae
    |  i. s.: Platypsectra analis ZS10
    |         Pergagrapta G84
    |           |--P. gravenhorstii G84
    |           `--P. polita PK17
    |         Acanthoperga R70
    |         Decameria similis HR11
    |         Heteroperreyia hubrichi HR11
    |--Philomastix [Philomastiginae] Na91
    |--Syzygoniinae Na91
    |--Acordulecera BD17 [Acordulecerinae Na91]
    |    `--A. pellucida BD17
    |--Loboceratinae Na91
    |--Paralypiinae Na91
    |--Conocoxinae Na91
    |--Cladomacra [Perreyiinae] Na91
    |--Styracotechys [Styracotechyinae] Na91
    |--Pergula [Pergulinae] Na91
    |--Pteryperga [Pteryperginae] Na91
    |    `--P. galla R70
    |--Pterygophorinae Na91
    |    |--Pterygophorus cinctus Boisduval 1835 Na91, B35
    |    `--Lophyrotoma interrupta Na91
    |--Phylacteophaginae Na91
    |    |--Leptoperga Riek 1970 RC74
    |    `--Phylacteophaga Na91
    |         |--P. eucalypti B88
    |         `--P. froggatti Na91
    |--Euryinae Na91
    |    |--Clarissa Na91
    |    |--Eurys Na91
    |    |--Polyclonus atratus RC74
    |    |--Warra Na91
    |    `--Ancyloneura Na91
    `--Perginae Na91
         |--Pseudoperga Na91
         |    |--P. belinda G84
         |    `--P. lewisii No91
         `--Perga Na91
              |--P. affinis Na91
              |--P. dorsalis R35
              |--P. leaski G84
              `--P. lewisi N70

*Type species of generic name indicated

REFERENCES

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[BD17] Branstetter, M. G., B. N. Danforth, J. P. Pitts, B. C. Faircloth, P. S. Ward, M. L. Buffington, M. W. Gates, R. R. Kula & S. G. Brady. 2017. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27: 1019–1025.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[G84] Gauld, I. D. 1984. An Introduction to the Ichneumonidae of Australia. British Museum (Natural History).

[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.

[Na91] 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).

[N70] Norris, K. R. 1970. General biology. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 107–140. Melbourne University Press.

[No91] Norris, K. R. 1991. General biology. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 68–108. Melbourne University Press: Carlton (Victoria).

[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.

[R35] Rayment, T. 1935. A Cluster of Bees: Sixty essays on the life-histories of Australian bees, with specific descriptions of over 100 new species. Endeavour Press: Sydney.

[R70] Riek, E. F. 1970. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 867–959. Melbourne University Press.

[RC74] Riek, E. F., & J. C. Cardale. 1974. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 107–111. Melbourne University Press.

[ZS10] Zborowski, P., & R. Storey. 2010. A Field Guide to Insects in Australia 3^rd ed. Reed New Holland: Sydney.

Last updated: 25 April 2022.

Anagrini

Anagrus epos, from the California Department of Food and Agriculture.

Belongs within: Mymaridae.

The Anagrini are a group of fairy wasps characterised by four-segmented tarsi and a truncately sessile metasoma held in place by a large thoracic muscle, the phragma (Girault 1929).

<==Anagrini G29
    |--Pteratomus Packard 1866 G29, GM79 [incl. Packardiella Ashmead 1904 G29, GM79]
    |--Metanthemus Girault 1928 G28
    |    `--*M. aureus Girault 1928 G28
    |--Parallelaptera Enock 1909 G29, GM79 [incl. Anthemiella Girault 1911 G29, GM79]
    |    `--P. rex (Girault 1911) GM79
    `--Anagrus Haliday 1833 GM79
         |--A. armatus (Ashmead 1887) GM79 [incl. A. spiritus G29]
         |    |--A. a. armatus GM79
         |    |--A. a. giraulti Crawford 1913 GM79
         |    `--A. a. nigriceps Girault 1915 GM79
         |--A. atomus TT02
         |--A. avalae TT02
         |--A. epos Girault 1911 GM79
         |--A. frequens N91
         |--A. nigriventris Girault 1911 TT02, GM79 [=A. armatus var. nigriventris TT02]
         |--A. puella Girault 1911 GM79
         `--A. ustulatus TT02

*Type species of generic name indicated

REFERENCES

[G28] Girault, A. A. 1928. Some new hexapods stolen from authority. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 221–224).

[G29] Girault, A. A. 1929. North American Hymenoptera Mymaridae. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 233–261).

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1–400.

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

[TT02] Triapitsyn, S. V., & D. A. J. Teulon. 2002. On the identity of Anagrus (Hymenoptera: Mymaridae) egg parasitoids of Froggatt’s apple leafhopper, Edwardsiana crataegi (Douglas) (Homoptera: Cicadellidae), in Christchurch, New Zealand. New Zealand Entomologist 25: 91–92.

Anaphes

Female Anaphes iole, photographed by Max E. Badgley.

Belongs within: Mymaridae.

Anaphes is a cosmopolitan genus of fairy wasps with a subsessile metasoma and subdivided scutellum (Girault 1929).

Characters (from Girault 1929): Antenna of female nine-segmented; antennal club solid. Scutellum preceded by large frustrum-shaped sclerite separating large triangular axillae and by a second, linear-transverse one; scutellum large, oval. Tarsi four-segmented. Metasoma subsessile, with no distinct petiole. Ovipositor free.

<==Anaphes Haliday 1833 GM79
    |--A. behmani Girault 1929 GM79
    |--A. conotracheli RD77
    |--A. diana N91
    |--A. hercules Girault 1911 GM79
    |--A. iole Girault 1911 GM79 (see below for synonymy)
    |    |--A. i. iole GM79
    |    `--A. i. anomocerus Girault 1929 GM79
    |--A. nigrellus Girault 1911 GM79
    |--A. pallipes Ashmead 1887 GM79
    |--A. pratensis Foerster 1847 GM79
    |--A. sinipennis Girault 1911 GM79
    `--A. victus MH11

Anaphes iole Girault 1911 GM79 [incl. A. ovijentatus Crosby & Leonard 1914 G29, GM79, A. perdubius Girault 1916 G29, GM79]

*Type species of generic name indicated

REFERENCES

[G29] Girault, A. A. 1929. North American Hymenoptera Mymaridae. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 233–261).

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1–400.

[MH11] Munro, J. B., J. M. Heraty, R. A. Burks, D. Hawks, J. Mottern, A. Cruaud, J.-Y. Rasplus & P. Jansta. 2011. A molecular phylogeny of the Chalcidoidea (Hymenoptera). PLoS One 6 (11): e27023.

[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).

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

Chalcididae

Trigonura sp., copyright Lynn Bergen.

Belongs within: Chalcidoidea.
Contains: Cratocentrini, Haltichellinae, Epitranus, Dirhininae, Phasgonophorini, Conura, Brachymeria (Brachymeria).

The Chalcididae are a group of chalcidoid wasps characterised by large, ventrally dentate hind femora that are adapted for jumping.

Characters (from Bouček 1988): Body often large and non-metallic, gaster convex, not collapsing; gena posteriorly with strong carina; thorax strongly punctate, prepectus very small, tegula only slightly longer than broad; hind femur strongly enlarged with teeth on ventral edge.

<==Chalcididae [Chalcidae, Chalcidinae] HB13
    |  i. s.: Spilochalcis mariae B37
    |--+--Cratocentrini HB13
    |  `--Xenanastatus Bouček 1988 MH11, B88
    |       `--*X. partisanguineus (Girault 1923) [=Eupelmus partisanguineus] B88
    `--+--+--Haltichellinae HB13
       |  `--+--+--Epitranus HB13
       |     |  `--Dirhininae HB13
       |     `--Smicromorpha Girault 1913 B88 [incl. Smicromorphella Girault 1930 B88; Smicromorphinae HB13]
       |          |--*S. doddi Girault 1913 [incl. S. cadaverosa Girault 1914] B88
       |          |--S. banksi Naumann 1986 B88
       |          |--S. eudela Naumann 1986 B88
       |          |--S. lagynos Naumann 1986 B88
       |          `--S. minera Girault 1926 [=*Smicromorphella minerva] B88
       `--+--+--Phasgonophorini HB13
          |  `--Conura HB13
          `--+--Trigonura Sichel 1865 HB13, B88 (see below for synonymy)
             |    |--*T. crassicauda [=Phasgonophora (*Trigonura) crassicauda] B88
             |    |--T. algerti HB13
             |    |--T. californica HB13
             |    |--T. euthyrrhinni (Dodd 1921) [=Chalcis euthyrrhini, *Chalcidellia euthyrrhini] B88
             |    |--T. guttatipennis (Girault 1927) [=Chalcidellia guttatipennis] B88
             |    `--‘Chalcidellia’ ucalegon R70
             `--Brachymeria Westwood in Stephens 1829 B88 (see below for synonymy)
                  |  i. s.: B. coloradensis (Cresson 1872) GM79 [incl. Chalcis tachinae Howard 1885 G17, GM79]
                  |         B. femorata (Panzer 1801) G38 [=Chalcis femorata G38; incl. C. ornatipes Cameron 1906 B88]
                  |         B. ovata N91 [=Chalcis ovata B37
                  |           |--B. o. ovata G17
                  |           `--‘Chalcis’ o. abiesae Girault 1917 G17
                  |         B. pedalis (Cresson 1872) GM79
                  |         B. pulchripes Holmgren 1868 B88
                  |         B. tibialis HB13
                  |--B. (Brachymeria) B88
                  |--B. (Australochalcis Girault 1939) B88
                  |    `--B. (*A.) humilicrus (Girault 1939) [=*Australochalcis humilicrus] B88
                  |--B. (Dirrhinomorpha Girault & Dodd in Girault 1915) B88
                  |    `--B. (*D.) angusta (Girault & Dodd in Girault 1915) [=*Dirrhinomorpha angusta] B88
                  `--B. (Mirochalcis Girault 1915) B88
                       `--B. (*M.) atricorpus (Girault 1915) [=*Mirochalcis atricorpus] B88

Brachymeria Westwood in Stephens 1829 B88 [incl. Ceyxia Cameron 1911 B88, Gahanula Burks 1960 B88, Meyeriella Krausse 1916 B88, Oncochalcis Cameron 1904 B88, Pseudobrachymeria Burks 1960 B88; Brachymerariae, Brachymeriini HB13]

Trigonura Sichel 1865 HB13, B88 [incl. Bactrochalcis Kieffer 1912 B88, Centrochalcidea Gahan & Fagan 1923 B88, Centrochalcis Cameron 1913 non Cameron 1905 B88, Chalcidellia Girault 1924 B88]

*Type species of generic name indicated

REFERENCES

[B37] Balduf, W. V. 1937. Bionomic notes on the common bagworm, Thyridopteryx ephemeraeformis Haw., (Lepid., Psychidae) and its insect enemies (Hym., Lepid.). Proceedings of the Entomological Society of Washington 39 (7): 169–184.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[G38] Gahan, A. B. 1938. Notes on some genera and species of Chalcidoidea (Hymenoptera). Proceedings of the Entomological Society of Washington 40 (8): 209–227.

[G17] Girault, A. A. 1917. Descriptiones stellarum novarum. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 80–101).

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1–400.

[HB13] Heraty, J. M., R. A. Burks, A. Cruaud, G. A. P. Gibson, J. Liljeblad, J. Munro, J.-Y. Rasplus, G. Delvare, P. Janšta, A. Gumovsky, J. Huber, J. B. Woolley, L. Krogmann, S. Heydon, A. Polaszek, S. Schmidt, D. C. Darling, M. W. Gates, J. Mottern, E. Murray, A. D. Molin, S. Triapitsyn, H. Baur, J. D. Pinto, S. van Noort, J. George & M. Yoder. 2013. A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics 29: 466–542.

[MH11] Munro, J. B., J. M. Heraty, R. A. Burks, D. Hawks, J. Mottern, A. Cruaud, J.-Y. Rasplus & P. Jansta. 2011. A molecular phylogeny of the Chalcidoidea (Hymenoptera). PloS One 6 (11): e27023.

[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).

[R70] Riek, E. F. 1970. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 867–959. Melbourne University Press.

Encyrtinae

Copidosoma floridanum, copyright Scott Justis.

Belongs within: Encyrtidae.
Contains: Mirini.

The Encyrtinae are encyrtid wasps in which paratergites (lateral subdivisions of the metasomal tergites) are absent (Naumann 1991).

<==Encyrtinae [Encyrtini] HB13
    |--Neocladia MH11
    `--+--+--Acerophagus flavus HB13
       |  `--Metaphycus HB13
       |       |--M. helvolus A71
       |       |--M. lounsburyi A71
       |       `--M. luteolus HB13
       `--+--+--+--Mirini MH11
          |  |  `--Comperiella bifasciata HB13
          |  `--Cheiloneurus HB13
          |       |--C. cervantesi Girault 1933 G33
          |       |--C. dumasi Girault 1932 G32a
          |       |--C. flaccus HB13
          |       |--C. fulvescens HB13
          |       `--C. javensis Girault 1917 G17a
          `--+--Trechnites fuscitarsis HB13
             `--Copidosoma Ratzeburg 1844 HB13, GM79
                  |--C. charon Guerrieri & Noyes 2001 F05
                  |--C. floridanum HB13
                  |--C. ortyx Guerrieri & Noyes 2001 F05
                  |--C. shakespearei Girault 1923 G23
                  |--C. truncatellum MH11
                  |--C. zdeneki Guerrieri & Noyes 2001 F05
                  `--C. zolta Guerrieri & Noyes 2001 F05

Encyrtinae incertae sedis:
  Tassonia Girault 1921 G21
    `--*T. gloriae Girault 1921 G21
  Chinchilla Girault 1928 G28
    `--*C. keatsi Girault 1928 G28
  Australia Girault 1928 G28 [=Australica (l. c.) GM79]
    `--*A. minuta Girault 1928 G28
  Encyrtus Latreille 1809 GM79 [incl. Comys RD77]
    |--E. arsanes Walker 1839 W39
    |--E. aurantifasciatus G32a [incl. E. barbatus Timberlake 1919 G32a, GM79]
    |--E. cheles Walker 1839 W39
    |--E. epytus Walker 1839 W39
    |--E. fasciatus (Fonscolombe 1832) GM79
    |--E. garibaldia Girault 1933 G33
    |--E. infidus [=Scelio (Encyrtus) infidus] G20
    |--E. longifuniculus Girault 1932 G32b
    |--E. lucetius Walker 1839 W39
    |--E. pacorus Walker 1839 W39
    |--E. purpureiventris Girault 1915 GM79
    |--E. salacon Walker 1839 W39
    |--E. xuthus Walker 1839 W39
    |--E. zameis Walker 1839 W39
    `--E. zebina Walker 1839 W39
  Psyllaephagus umbro (Walker 1839) N91, B88 [=Miscogaster umbro B88]
  Microteryina [Microteryini] S96
    |--Allencyrtus S96
    `--Microterys Thomson 1876 N91, GM79
         |--M. cincticornis Ashmead 1899 GM79
         |--M. clauseni A71
         |--M. flavus B88
         |--M. lunatus (Dalman 1820) GM79
         |--M. mazzinini Girault 1917 G17b
         |--M. speciosissimus Girault 1911 GM79
         |--M. sylvius (Dalman 1820) GM79
         |--M. tesselatus (Dalman 1820) GM79
         `--M. titiani Girault 1917 G17b
  Saprencyrtus N91
  Aenasiella N91
    |--A. africa Girault 1917 G17b
    `--A. ovi Girault 1925 G25
  Eugahania N91
  Tachinaephagus Ashmead 1904 N91, GM79 [incl. Australencyrtus Johnson & Tiegs 1921 G24, GM79]
    `--T. zealandicus Ashmead 1904 (see below for synonymy) G38
  Syrphophagus aphidivorus N91, B88
  Ooencyrtus N91
    |--O. bicolor Girault 1915 GM79
    |--O. californicus Girault 1917 [=Oenocyrtus (l. c.) californicus] G17b
    |--O. johnsoni (Howard 1898) GM79
    |--O. lacteiclavus Girault 1932 G32b
    |--O. pacificus Waterston 1915 GM79
    `--O. pityocampae (Mercet 1921) L-SSY04
  Fulgoridicida N91
  Xenoencyrtus niger N91, C81
  Amira N91
  Comperia N91
  Paralitomastix kohleri R70
  Coelopencyrtus N91
  Ixodiphagus Howard 1907 N91, ST08
  Hunterellus R70
  Quaylea whittieri R70, A71
  Litomastix R70
    |--L. auricollis Thompson 1875 GM79
    `--L. truncatellus A71

Tachinaephagus zealandicus Ashmead 1904 [incl. T. australiensis Girault 1917, Stenoterys fulvoventralis Dodd in Groggatt 1921, Australencyrtus giraulti Johnston & Tiegs 1931] G38

*Type species of generic name indicated

REFERENCES

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

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[C81] Caltagirone, L. E. 1981. Landmark examples in classical biological control. Annual Review of Entomology 26: 213–232.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

[G38] Gahan, A. B. 1938. Notes on some genera and species of Chalcidoidea (Hymenoptera). Proceedings of the Entomological Society of Washington 40 (8): 209–227.

[G17a] Girault, A. A. 1917a. New Javanese Hymenoptera. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 59–71).

[G17b] Girault, A. A. 1917b. Descriptiones stellarum novarum. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 80–101).

[G21] Girault, A. A. 1921. New animals of Australia and old men of the earth. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 161–163).

[G23] Girault, A. A. 1923. Loves wooed and won in Australia. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 167–169).

[G24] Girault, A. A. 1924. Homo perniciosus and new Hymenoptera. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 178–181).

[G25] Girault, A. A. 1925. An essay on when a fly is lovable, the ceremony of baptizing some and unlovely hate. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 187–190).

[G28] Girault, A. A. 1928. Some new hexapods stolen from authority. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 221–224).

[G32a] Girault, A. A. 1932a. New pests from Australia, X. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 286–291).

[G32b] Girault, A. A. 1932b. Hymenoptera, Thysanoptera nova Australiensis. II. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 292).

[G33] Girault, A. A. 1933a. Some beauties inhabitant not of commercial boudoirs but of Nature’s bosom, notably new insects. Privately published (reprinted in: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 299–303).

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1–400.

[HB13] Heraty, J. M., R. A. Burks, A. Cruaud, G. A. P. Gibson, J. Liljeblad, J. Munro, J.-Y. Rasplus, G. Delvare, P. Janšta, A. Gumovsky, J. Huber, J. B. Woolley, L. Krogmann, S. Heydon, A. Polaszek, S. Schmidt, D. C. Darling, M. W. Gates, J. Mottern, E. Murray, A. D. Molin, S. Triapitsyn, H. Baur, J. D. Pinto, S. van Noort, J. George & M. Yoder. 2013. A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics 29: 466–542.

[L-SSY04] López-Sebastián, E., J. Selfa & J. Ylla. 2004. Primeros datos del parasitismo de Ooencyrtus pityocampae (Mercet, 1921) sobre Graellsia isabelae (Graells, 1849) en condiciones de laboratorio. Graellsia 60 (1): 121–123.

[MH11] Munro, J. B., J. M. Heraty, R. A. Burks, D. Hawks, J. Mottern, A. Cruaud, J.-Y. Rasplus & P. Jansta. 2011. A molecular phylogeny of the Chalcidoidea (Hymenoptera). PloS One 6 (11): e27023.

[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).

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[R70] Riek, E. F. 1970. Hymenoptera (wasps, bees, ants). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 867–959. Melbourne University Press.

[S96] Sharkov, A. 1996. A review of the species of the genus Mucrencyrtus Noyes (Hymenoptera: Encyrtidae). Proceedings of the Entomological Society of Washington 98 (2): 350–368.

[ST08] Simutnik, S. A., & V. A. Trjapitzin. 2008. Description of a new species of the genus Globulencyrtus (Hymenoptera, Chalcidoidea, Encyrtidae) from Australia with information of G. politus, the type species, a parasitoid of mites of the family Caeculidae (Acarina). Vestnik Zoologii 42 (6): 559–562.

[W39] Walker, F. 1839. Monographia Chalciditum vol. 2. Species collected by C. Darwin, Esq. Hyppolite Baillière: London.

Asaphinae

Asaphes cf. vulgaris, copyright Pierre Bornand.

Belongs within: Chalcidoidea.

The Asaphinae are a group of chalcidoid wasps mostly developing as parasites or hyperparasites on Sternorrhyncha, particularly aphids and scales (Bouček 1988).

Characters (from Bouček 1988): Occipital carina distinctly developed, distinct carina present posteriorly on gena; scutellum anteriorly acuminate but axillar grooves interrupted in the middle and their descending part very deep; propodeum medially with rather coarse irregular sculpture often suggesting areolation; petiole anteriorly with transverse raised edge or carina, its body often parallel-sided and with longitudinal ribs; basal tergites more or less greatly enlarged, not collapsing.

<==Asaphinae [Asaphini]
    |--Ausasaphes Bouček 1988 B88
    |    |--*A. pallipes Bouček 1988 B88
    |    `--A. atripes Bouček 1988 B88
    |--Enoggera Girault 1926 [incl. Enoggerella Girault 1926] B88
    |    |--*E. polita Girault 1926 B88
    |    `--E. nassaui (Girault 1926) [=*Enoggerella nassaui] B88
    `--Asaphes Walker 1834 [incl. Isocratus Förster 1856, Parectroma Brethes 1913] B88
         |--*A. vulgaris Walker 1834 B88
         |--A. americanus Girault 1914 GM79
         |--A. californicus Girault 1917 G17
         |--A. rufipes Brues 1908 G17, GM79
         `--A. suspensus HB13

*Type species of generic name indicated

REFERENCES

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[G17] Girault, A. A. 1917. Descriptiones hymenopterorum chalcidoidicarum variorum cum observationibus. V. Privately published (reprinted Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 116–131).

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1–400.

[HB13] Heraty, J. M., R. A. Burks, A. Cruaud, G. A. P. Gibson, J. Liljeblad, J. Munro, J.-Y. Rasplus, G. Delvare, P. Janšta, A. Gumovsky, J. Huber, J. B. Woolley, L. Krogmann, S. Heydon, A. Polaszek, S. Schmidt, D. C. Darling, M. W. Gates, J. Mottern, E. Murray, A. D. Molin, S. Triapitsyn, H. Baur, J. D. Pinto, S. van Noort, J. George & M. Yoder. 2013. A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics 29: 466–542.