Adeloidea

Green longhorns Adela reaumurella, copyright Nick Wood.

Belongs within: Heteroneura.

The Adeloidea are a group of small moths of which larvae include leaf-miners, gall-makers or internal borers of plants. Members include the Adelidae which have the longest antennae relative to body size of any Lepidoptera (Nielsen & Common 1991). The Cecidosidae, known from South Africa, South America and New Zealand, mostly develop as gall-formers on Anacardiaceae though larvae of the basal New Zealand Xanadoses nielseni are miners under smooth tree bark (Hoare & Dugdale 2003).

Characters (from Nielsen & Common 1991, as Incurvarioidea): Small to very small; ocelli and chaetosemata absent; antennae short to markedly longer than fore wing, scape without eye-cap, often with pecten; proboscis long, short or absent, scaled near base; maxillary palps 5-segmented to reduced; labial palps 3-segmented (rarely absent), second segment with lateral bristles; epiphysis usually present; tibial spurs 0-2-4 (rarely 0-2-2); wings usually with microtrichia, hind wing usually with frenulum and pseudofrenular bristles; male often with rows of blunt spines (pectinifers) on valva and with arrow-shaped juxta; female with long vestibulum supported by strong muscle tendons ('guy wires') and piercing ovipositor for inserting eggs in plant tissues. Larva apodous or with short prolegs, crochets in transverse rows, anal prolegs and crochets usually absent; leaf-miners later cutting out or forming flattened cases used as portable shelters while leaf-skeletonising or feeding on flowers or for pupation, or gall-makers of internal borers in stems and twigs. Pupa protruded from case at ecdysis.

<==Adeloidea [Incurvarioidea] KP19
    |--Heliozelidae NC91
    |    |  i. s.: Hoplophanes chlorochrysa NC91
    |    |--Plesiozela Karsholt & Kristensen 2003 KK03
    |    |    |--*P. nielseni Karsholt & Kristensen 2003 (see below for synonymy) KK03
    |    |    `--P. patagonica Karsholt & Kristensen 2003 KK03
    |    `--+--Antispila cornifoliella KK03, WRM02
    |       |--Heliozela KK03
    |       |    |--H. prodela NC91
    |       |    `--H. resplendella (Stainton 1851) KK03
    |       `--Coptodisca KK03
    |            |--C. kalmiella WRM02
    |            `--C. splendiferella G17
    `--+--Adelidae NC91
       |    |  i. s.: Chalceopla WRM02
       |    |           |--C. cyanella P27
       |    |           `--C. simpliciella WRM02
       |    |         Adelites P92
       |    |--Ceromitia [Nematopogoninae] NC91
       |    |    `--C. iolampra NC91
       |    `--Adelinae NC91
       |         |--Nemophora C70
       |         |    |--N. degeerella KP19
       |         |    |--N. sparsella C91
       |         |    `--N. topazias C70
       |         `--Adela Latreille 1802 L02
       |              |--*A. reaumurella [=Alucita reaumurella] L02
       |              |--A. bella P27
       |              |--A. degeerella G20
       |              |--A. kuznetzovi RJ93
       |              |--A. pantherella Guénée 1849 E12
       |              |--A. septentrionalis WRM02
       |              |--A. trigrapha WRM02
       |              `--A. viridella R13
       `--+--Crinopterygidae NC91
          `--+--Incurvariidae [Incurvariinae] NC91
             |    |  i. s.: ‘Tinea’ monophthalma NC91
             |    |         ‘Tinea’ nectaria C70
             |    |         ‘Tinea’ phauloptera C70
             |    |--Perthida glyphopa KK03, SC04
             |    |--Incurvaria masculella DG03
             |    |--Incurvarites alienella P92
             |    `--Prophalonia P92
             `--+--Prodoxidae NC91
                |    |  i. s.: Greya politella GE05, WRM02
                |    `--Prodoxinae GE05
                |         |--Parategeticula GE05
                |         |--Prodoxus GE05
                |         |    |--P. decipiens EN20
                |         |    `--P. quinquepunctella WRM02
                |         `--Tegeticula GE05
                |              |--T. alba P27
                |              |--T. cassandra EN20
                |              |--T. corruptrix GE05
                |              |--T. filamentosa RD77
                |              |--T. intermedia GE05
                |              `--T. yuccasella WRM02
                `--Cecidosidae NC91
                     |--Xanadoses Hoare & Dugdale 2003 HD03
                     |    `--*X. nielseni Hoare & Dugdale 2003 HD03
                     `--+--+--Ptisanora Meyrick 1913 [incl. Liopseustis Meyrick 1928] HD03
                        |  |    `--*P. trivialis Meyrick 1913 [incl. P. planicola Meyrick 1928] HD03
                        |  `--Scyrotis Meyrick 1909 HD03
                        |       |--*S. athleta Meyrick 1909 HD03
                        |       |--S. diplopsamma Meyrick 1918 HD03
                        |       `--S. granosa Meyrick 1912 HD03
                        `--+--Cecidoses Curtis 1835 HD03
                           |    `--*C. eremita Curtis 1835 HD03
                           |--Eucecidoses Brèthes 1916 HD03
                           |    `--*E. minutanus Brèthes 1916 HD03
                           |--Oliera Brèthes 1916 HD03
                           |    `--*O. argentinana Brèthes 1916 HD03
                           `--Dicranoses Kieffer & Jörgensen 1910 [incl. Ridiaschina Brèthes 1916] HD03
                                |--*D. capsulifex Kieffer & Jörgensen 1910 HD03
                                `--D. congregatella Brèthes 1916 HD03

*Plesiozela nielseni Karsholt & Kristensen 2003 [=P. obliqua Nielsen ex Karsholt & Kristensen 2003 (n. n.)] KK03

*Type species of generic name indicated

REFERENCES

[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

[C91] CSIRO. 1991. The Insects of Australia: A textbook for students and research workers vol. 1. Melbourne University Press: Carlton (Victoria).

[DG03] Davis, D. R., & P. Gentili. 2003. Andesianidae, a new family of monotrysian moths (Lepidoptera: Andesianoidea) from South America. Invertebrate Systematics 17: 15–26.

[EN20] Eggli, U., & R. Nyffeler (eds) 2020. Illustrated Handbook of Succulent Plants: Monocotyledons 2^nd ed. Springer.

[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.

[G17] Girault, A. A. 1917. Descriptiones stellarum novarum. 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: 80–101).

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

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

[HD03] Hoare, R. J. B., & J. S. Dugdale. 2003. Description of the New Zealand incurvarioid Xanadoses nielseni, gen. nov., sp. nov. and placement in Cecidosidae (Lepidoptera). Invertebrate Systematics 17: 47–57.

[KK03] Karsholt, O., & N. P. Kristensen. 2003. Plesiozela, gen. nov. from temperate South America: apparent sister-group of the previously known Heliozelidae (Lepidoptera: Incurvarioidea: Heliozelidae). Invertebrate Systematics 17: 39–46.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

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

[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.

[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.

[SC04] Stone, C., & N. C. Coops. 2004. Assessment and monitoring of damage from insects in Australian eucalypt forests and commercial plantations. Australian Journal of Entomology 43 (3): 283–292.

[WRM02] Wiegmann, B. M., J. C. Regier & C. Mitter. 2002. Combined molecular and morphological evidence on the phylogeny of the earliest lepidopteran lineages. Zoologica Scripta 31 (1): 67–81.

Last updated: 25 April 2022.

Ditrysia

Bucculatrix cristatella, copyright Ilia Ustyantsev.

Belongs within: Heteroneura.
Contains: Tineidae, Apoditrysia, Yponomeutoidea, Gracillariidae, Lyonetiidae, Psychinae.

The Ditrysia are a major clade of Lepidoptera accounting for some 98% of the species of the order. They are characterised by female genitalia with an internal connection between the copulatory opening and ovipore, as well as by large apodemes on the second abdominal sternite and short bands rather than long fibres in the proboscis muscles (Grimaldi & Engel 2005). Basal members of the clade, historically gathered into the superfamily Tineoidea (now recognised as non-monophyletic) are small to medium-sized species with narrow, typically grey fore wings that often retain the M vein in the discal cell. Most lack ocelli though prominent ocelli are retained in the Douglasiidae (Nielsen & Common 1991). The Simaethistiidae are a small, poorly known group of basal ditrysians found in eastern Asia and Australia

<==Ditrysia [Gracillarioidea, Papilionina, Tineina, Tineites, Tineoidea, Tortrices]
    |--Meessiinae [Meessiidae] KP19
    |    |--Meessia leopoldella YS10
    |    |--Demobrotis NC91
    |    `--Eudarcia simulatricella KP19
    `--+--+--+--Tineidae KP19
       |  |  `--Dryadaula [Dryadaulidae] KP19
       |  |       `--D. visaliella KP19
       |  `--+--Apoditrysia GE05
       |     `--+--+--Yponomeutoidea KP19
       |        |  `--Roeslerstammiidae [Amphitheridae, Amphitherinae] NC91
       |        |       |--Amphithera heteroleuca KP19, NC91
       |        |       |--Enchoptila idiopis P27
       |        |       |--Thereutis NC91
       |        |       |--Macarangela leucochrysa NC91, C91
       |        |       |--Nematobola NC91
       |        |       |--Chalcoteuches phlogera NC91
       |        |       `--Roeslerstammia NC91
       |        `--+--Gracillariidae KP19
       |           `--+--Bucculatricidae NC91
       |              |    |--Ogmograptis scribula NC91
       |              |    `--Bucculatrix KP19
       |              |         |--B. alaternella Constant 1890 C90
       |              |         |--B. cristatella KP19
       |              |         |--B. gossypii NC91
       |              |         |--B. helichrysella Constant 1890 C90
       |              |         `--B. platani RJ93
       |              `--+--Lyonetiidae KP19
       |                 |--Bedellia NC91 [Bedelliidae M03, Bedelliinae]
       |                 |    |--B. psamminella L27
       |                 |    `--B. somnulentella P27
       |                 `--Heliodinidae M03
       |                      |--Thylacosceles acridomima P27
       |                      |--Heliodines princeps NC91
       |                      |--Epicroesa ambrosia NC91
       |                      |--Baltonides roeselliformis RJ93
       |                      `--Vanicela P27
       |                           |--V. disjunctella P27
       |                           `--V. xenadelpha P27
       `--Psychidae B37
            |  i. s.: Liothula omnivora WFS04
            |         Solenobia triquetrella D51
            |         Iphierga macarista NC91
            |         Epichnopteryx plumella (Denis & Schiffermüller 1775) JP05
            |         Canephora JP05
            |           |--C. hirsuta Poda 1761 JP05
            |           `--C. unicolor RD77
            |         Paramictoides febretta YS10
            |         Pachythelia unicolor YS10
            |         Apterona YS10
            |         Plutorectis G84
            |         Paracharactis MC13
            |         Cryptothelea GE05
            |           |--C. cardiophaga IR05
            |           |--C. gloverii VC07
            |           `--C. junodi F92
            |         Luffia YS10
            |           |--L. ferchaultella BC91
            |           `--L. lapidella YS10
            |         Animula huebneri H01
            |         Cochliotheca RD77
            |           |--C. crenulella RD77
            |           `--C. helix RD77
            |         Acanthopsyche opacella [incl. A. atra] RD77
            |         Amicta RD77
            |         Sterrhopteryx pristinella Rebel 1934 P92
            |--Psychinae NC91
            |--Oiketicinae KP19
            |    |--Thyridopteryx ephemeraeformis KP19
            |    `--Oiketicus G84
            |         |--O. elongatus G84
            |         `--O. kirbyi L81
            `--Taleporiinae NC91
                 |--Ctenocompa C70
                 |--Narycia saxosa C70, P27
                 |--Trigonocyttara clandestina C70
                 |--Lepidoscia arctiella C70
                 |--Cebysa leucotelus C70, NC91
                 `--Taleporia JP05
                      |--T. cawthronella P27
                      `--T. tubulosa (Retzius 1783) JP05

Ditrysia incertae sedis:
  Azinis hilarella WM66
  Apoprogonidae NC91
  Cerostoma Latreille 1802 L02
    |--*C. dorsatum [=Ypsolophus dorsatus] L02
    `--C. persicella F92
  Simaethistiidae GE05
  Pseudarbelidae NC91
  Arrhenophanidae NC91
  Acrolophus D01 [Acrolophidae GE05]
    |--A. boucardi Druce 1901 D01
    |--A. linus Druce 1901 D01
    `--A. underwoodi Druce 1901 D01
  Eucryptogona [Eriocottidae] NC91
    `--E. trichobathra NC91
  Douglasiidae NC91
    |--Douglasia Stainton 1854 [=Douglassia (l. c.) non Bartsch 1934] P66
    |--Tinagma leucanthes NC91
    `--Klimeschia NC91
  Melusina [Compsoctenidae] RD77
    `--M. energa RD77
  Euplocamus guttella G20

*Type species of generic name indicated

REFERENCES

[BC91] Baker, G. T., & A. Chandrapatya. 1991. Fine structure of the chorion of the broak mite Polyphagotarsonemus latus (Prostigmata: Tarsonemidae). 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. 2 pp. 325–328. SPB Academic Publishing: The Hague.

[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.

[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

[C90] Constant, A. 1890. Description de dix espèces nouvelles de microlépidoptères de la faune française. Annales de la Société Entomologique de France, 6^e série 9: cxxiv–cxxvi.

[C91] CSIRO. 1991. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1. Melbourne University Press: Carlton (Victoria).

[D51] Dobzhansky, T. 1951. Genetics and the Origin of Species 3^rd ed. Columbia University Press: New York.

[D01] Druce, H. 1901. Descriptions of some new species of Lepidoptera from east Africa and tropical America. Annals and Magazine of Natural History, series 7, 7: 432–444.

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

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

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

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

[H01] Hampson, G. F. 1901. The Lepidoptera-Phalaenae of the Bahamas. Annals and Magazine of Natural History, series 7, 7: 246–261.

[IR05] Idris, A. B., & A. Rizki. 2005. Notes on the tribe Ephialtini (Hymenoptera: Ichneumonidae: Pimplinae) of Malaysia. Serangga 10 (1–2): 111–126.

[JP05] Jungmann, E., & U. Poller. 2005. Neunachweise und Wiederfunde für die Schmetterlingsfauna (Insecta; Lepidoptera) des Altenburger Landes. Mauritiana 19 (2): 317–326.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L27] Lindsay, S. 1927. A list of the Lepidoptera of Deans Bush, Riccarton, Canterbury. Transactions and Proceedings of the New Zealand Institute 57: 693–696.

[L81] Long, J. L. 1981. Introduced Birds of the World: The worldwide history, distribution and influence of birds introduced to new environments. Reed: Sydney.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[M03] McQuillan, P. B. 2003. The giant Tasmanian ‘pandani’ moth Proditrix nielseni, sp. nov. (Lepidoptera: Yponomeutoidea: Plutellidae s. l.) Invertebrate Systematics 17: 59–66.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[P66] Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae: an evaluation of the valid taxa, both Recent and fossil, with lists of characteristic species. Bulletin of the Auckland Institute and Museum 5: 1–184, pls 1–23.

[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.

[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.

[VC07] Villanueva, R. T., & C. C. Childers. 2007. Insect predators of eriophyids on Florida citrus including a new predacious Lepidoptera and two cecidomyiids. In: Morales-Malacara, J. B., V. M. Behan-Pelletier, E. Ueckermann, T. M. Pérez, E. G. Estrada-Venegas & M. Badii (eds) Acarology XI: Proceedings of the International Congress pp. 391–395. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[WM66] Wallace, A. R., & F. Moore. 1866. List of lepidopterous insects collected at Takow, Formosa, by Mr. R. Swinhoe. Proceedings of the Zoological Society of London 1866: 355–365.

[WFS04] Winks, C. J., S. V. Fowler & L. A. Smith. 2004. Invertebrate fauna of boneseed, Chrysanthemoides monilifera ssp. monilifera (L.) T. Norl. (Asteraceae: Calenduleae), an invasive weed in New Zealand. New Zealand Entomologist 27: 61–72.

[YS10] Yefremova, Z. A., & I. S. Strakhova. 2010. A review of the species of the genus Elasmus Westwood (Hymenoptera, Eulophidae) from Russia and neighboring countries. Entomologicheskoe Obozrenie 89 (3): 634–661.

Last updated: 25 April 2022.

Gracillariidae

Cocoa pod borer Acrocercops cramerella, from the Papua Insects Foundation.

Belongs within: Ditrysia.

The Gracillariidae are very small moths with long antennae and long fringes on the hind wings (Nielsen & Common 1991).

Characters (from Nielsen & Common 1991): Very small, slender; head usually smooth scaled; ocelli and chaetosemata absent; antennae nearly as long as, or longer than, fore wing, filiform, scape slender, usually without pecten, sometimes with small eye-cap; proboscis present, unsealed; maxillary palps slender, porrect or ascending, four-segmented, or reduced; labial palps slender, porrect or ascending; epiphysis present, spurs 0-2-4, hind tibiae smooth or with dorsal row of stiff bristles, never hairy, fore and mid tibiae sometimes thickened with scales; fore wing lanceolate or linear-lanceolate with long fringing piliform scales, R₅ to costa or apex, some veins often lost, 1A+2A without basal fork; hind wing lanceolate or linear, venation often reduced; male abdomen often with coremata on segments 7 and 8. Larva leaf- or gall-mining, at least in early stages, with heteromorphosis; early instars flattened, with large blade-like mandibles, sap-feeding, thoracic legs and prolegs absent; after second or third ecdysis larvae often feed on parenchyma, body cylindrical, thoracic legs present, prolegs on abdominal segments 3–5, crochets usually in lateral penellipse. Pupa with antennae and proboscis often extending beyond wing tips; abdomen with fine scattered dorsal spines, segments 5–7 movable in male, 5–6 in female; usually in flattened cocoon.

<==Gracillariidae [Lithocolletidae]
    |  i. s.: Parectopa formosa P27
    |         Epicephala P27
    |           |--E. chalybacma VHK02
    |           `--E. frugicola B88
    |         Geometridites larentiiformis RJ93
    |         Ornativalva plutelliformis SKR96
    |         Gracillariites P92
    |--Gracillariinae NC91
    |    |--Macarostola formosa NC91
    |    |--Caloptilia KP19
    |    |    |--C. alchimiella [=Gracilaria alchimiella] V09
    |    |    |--C. azaleella NC91
    |    |    |--C. elaeas M83
    |    |    `--C. triadicae KP19
    |    `--Gracilaria GE05
    |         |--G. chrysitis L27
    |         |--G. elaeas L27
    |         |--G. linearis P27
    |         |--G. ononidis R13
    |         |--G. selenitis P27
    |         `--G. xylophanes P27
    `--+--+--Philodoria [Ornixolinae] KP19
       |  `--Lithocolletinae KP19
       |       |--Cameraria ohridella KP19
       |       |--Lithocolletis B88
       |       |    |--L. aglaozona C70
       |       |    |--L. hamadryadella G17c
       |       |    `--L. populifolia [=Gracilaria populifolia] H04
       |       `--Phyllonorycter NC91
       |            |--P. aglaozona NC91
       |            |--P. blancardella MC02
       |            |--P. crataegella G17b
       |            |--P. hamadryadella RD77
       |            `--P. messaniella NC91
       `--+--Phyllocnistis [Phyllocnistidae, Phyllocnistinae] KP19
          |    |--P. citrella NC91
          |    |--P. diaugella NC91
          |    `--P. suffusella B88
          `--Acrocercops KP19
               |--A. antimima NC91
               |--A. calicella NC91
               |--A. chionosoma B88
               |--A. cramerella G17a
               |--A. diffluella B88
               |--A. ordinatella P27
               |--A. plebeia NC91
               |--A. transecta KP19
               `--A. tricuneatella NC91

*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).

[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

[G17a] Girault, A. A. 1917a. New Javanese Hymenoptera. 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: 59–71).

[G17b] Girault, A. A. 1917b. New chalcid flies. 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: 72–77).

[G17c] Girault, A. A. 1917c. 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).

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

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[L27] Lindsay, S. 1927. A list of the Lepidoptera of Deans Bush, Riccarton, Canterbury. Transactions and Proceedings of the New Zealand Institute 57: 693–696.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: An annotated species list. DSIR Entomology Division Report 3: 1–98.

[MC02] Miliczky, E. R., & C. O. Calkins. 2002. Spiders (Araneae) as potential predators of leafroller larvae and egg masses (Lepidoptera: Tortricidae) in central Washington apple and pear orchards. Pan-Pacific Entomologist 78 (2): 140–150.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

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

[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.

[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.

[SKR96] Sobhian, R., L. Knutson & J. S. Rodier. 1996. Biology and host specificity notes on Styphlus penicillus Gyllenhal (Coleoptera: Curculionidae), examined as a biological control agent for Crupina vulgaris in the United States with remarks on its host plant. Proceedings of the Entomological Society of Washington 98 (2): 317–323.

[V09] Verdcourt, B. (ed.) 2009. Additions to the Wild Fauna and Flora of the Royal Botanic Gardens, Kew. XXVI. Miscellaneous records. Kew Bulletin 64 (1): 183–194.

[VHK02] Verma, M., M. Hayat & S. I. Kazmi. 2002. The species of Elasmus from India (Hymenoptera: Chalcidoidea: Eulophidae). Oriental Insects 36: 245–306.

Eucosmini

Macadamia nut-borer Cryptophlebia ombrodelta, copyright Victor Fazio.

Belongs within: Tortricidae.

The Eucosmini are a group of tortricid moths characterised by male genitalia with a one‐branched protuberance on the valve, and reduced gnathos and uncus.

<==Eucosmini NC91
    |--Bathrotoma constrictana NC91
    |--Crocidosema plebejana NC91
    |--Epinotia NC91
    |    |--E. aporema J-PWM04
    |    `--E. lantana NC91
    |--Eucosma KN03
    |    |--E. coniogramma Clarke 1976 [incl. E. okubiensis Kawabe 1987] KN03
    |    |--E. critica VHK02
    |    |--E. querula L27
    |    |--E. sordidana F92
    |    `--E. triangulana C70
    `--Cryptophlebia NC91
         |--C. amamiana Komai & Nasu 2003 KN03
         |--C. horii Kawabe 1987 KN03
         |--C. lasiandra KN03
         |--C. leucotreta B88
         |--C. ombrodelta KN03
         |--C. palustris Komai & Nasu 2003 KN03
         `--C. rhizophorae Vári 1981 KN03

*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).

[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

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

[J-PWM04] Jiménez-Pérez, A., Q. Wang & N. Markwick. 2004. Intrinsic rate of increase in ‘Cnephasia’ jactatana Walker (Lepidoptera: Tortricidae), a pest of kiwifruit. New Zealand Entomologist 27: 109–112.

[KN03] Komai, F., & Y. Nasu. 2003. Four species of Olethreutinae (Lepidoptera: Tortricidae) associated with viviparous seedlings of the mangrove Rhizophoraceae in the Ryukyu Islands, Japan. Invertebrate Systematics 17: 75–87.

[L27] Lindsay, S. 1927. A list of the Lepidoptera of Deans Bush, Riccarton, Canterbury. Transactions and Proceedings of the New Zealand Institute 57: 693–696.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[VHK02] Verma, M., M. Hayat & S. I. Kazmi. 2002. The species of Elasmus from India (Hymenoptera: Chalcidoidea: Eulophidae). Oriental Insects 36: 245–306.

Archipini

Tea tortrix Homona coffearia, copyright Challiyil Eswaramangalath Pavithran Vipin.

Belongs within: Tortricidae.

The Archipini are a group of tortricid moths whose larvae feed between joined, living leaves (Nielsen & Common 1991). Many species have wings marked with a distinct basal and median fascia and a dark costal spot.

<==Archipini NC91
    |--Epiphyas postvittana NC91
    |--Archips G05
    |    |--A. argyrospila G05
    |    `--A. eucrora F92
    |--Cryptoptila australana NC91, C70
    |--Arotrophora NC91
    |    |--A. arcuatalis NC91 [=Scopula arcuatalis M86; incl. Crambus submarginellus M86, Eromene transcissella M86]
    |    `--A. siniocosma C70
    |--Merophyas NC91
    |    |--M. divulsana NC91
    |    `--M. leucariana M83
    `--Homona NC91
         |--H. coffearia VHK02
         |--H. similans P27
         `--H. spargotis NC91

*Type species of generic name indicated

REFERENCES

[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

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

[G05] Grissell, E. E. 2005. A review of North American species of Microdontomerus Crawford (Torymidae: Hymenoptera). Journal of Hymenoptera Research 14 (1): 22–65.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

[M86] Meyrick, E. 1886. Notes on synonymy of Australian micro-Lepidoptera. Proceedings of the Linnean Society of New South Wales, series 2, 1 (3): 803–806.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[VHK02] Verma, M., M. Hayat & S. I. Kazmi. 2002. The species of Elasmus from India (Hymenoptera: Chalcidoidea: Eulophidae). Oriental Insects 36: 245–306.

Obtectomera

Twenty-plume moth Orneodes hexadactyla, copyright Entomart.

Belongs within: Apoditrysia.
Contains: Copromorphoidea, Pyraloidea, Macroheterocera, Gelechioidea, Thyrididae, Alucitoidea, Pterophoridae, Papilionidae, Hesperiidae, Pieridae, Nymphalidae, Riodinidae, Lycaenidae.

The Obtectomera are a clade of Lepidoptera united in its circumscription below by molecular phylogenetic analysis (Kawahara et al. 2019). Historically, the clade has been characterised by pupae with abdominal segments 1–4 immobile and without a dorsal row of spines on the tergites (Grimaldi & Engel 2005); however, Obtectomera in this sense may not be monophyletic. Representatives of the Obtectomera include the Calliduloidea, a small group of diurnal moths with simple antennae and wings held back to back at rest, resembling small butterflies (Common 1970). Members of the superfamilies Alucitoidea and Pterophoroidea, plume moths, often have the fore and hind wings deeply cleft into multiple plumes. Pterophoroidea usually possess vein CuP in the fore wing whereas this vein is absent in Alucitoidea (Nielsen & Common 1991). The Papilionoidea, butterflies and skippers, are diurnal Lepidoptera with a pouch on abdominal tergite 1 that lay eggs upright and have a girdle around the pupa (Grimaldi & Engel 2005).

Obtectomera [Bombycina, Bombycites, Macrolepidoptera, Pyralidina] KP19
    |  i. s.: Hyblaea IR05 [Hyblaeidae, Hyblaeoidea NC91]
    |           |--H. ibidias NC91
    |           `--H. pleura IR05
    |         Copromorphoidea GE05
    |         Whalleyaniidae GE05
    |         Cypra delicatula [incl. Cozistra membranacea] M86
    |         Artaxa M86
    |           |--A. lutea M86
    |           `--A. paraneura Meyrick 1886 M86
    |--+--+--Pyraloidea KP19
    |  |  `--+--Macroheterocera KP19
    |  |     `--Mimallonidae [Lacosomidae, Mimallonoidea] KP19
    |  |          |--Bedosia strigifera (Felder & Felder 1874) [=Perophora strigifera] NSp07
    |  |          |--Lacosoma chiridota KP19
    |  |          |--Menevia lantona KP19
    |  |          `--Cicinnus NSp07
    |  |               |--C. despecta (Walker 1855) [incl. Perophora sanguinolenta Felder & Felder 1874] NSp07
    |  |               `--C. melsheimeri P27
    |  `--+--Gelechioidea KP19
    |     `--+--+--Thyrididae KP19
    |        |  `--Calliduloidea KP19
    |        |       |--Pterothysanus [Pterothysanidae] RD77
    |        |       `--Callidulidae KP19
    |        |            |--Cleis scotti C70
    |        |            `--Pterodecta [Callidulinae] KP19
    |        |                 `--P. felderi KP19
    |        `--+--Alucitoidea KP19
    |           `--Pterophoroidea [Pterophorii] GE05
    |                |--Pterophoridae GE05
    |                `--Orneodes Latreille 1802 L02 [Orneodidae P27]
    |                     |--*O. hexadactyla L02, RD77 [=Pterophorus hexadactylus L02, Alucita hexadactyla H04]
    |                     |--O. objurgatella B88
    |                     |--O. phricodes P27
    |                     `--O. pygmaea RD77
    `--Papilionoidea [Papilionides, Rhopalocera] KP19
         |  i. s.: Thanatites vetula GE05
         |         Corades B66a
         |           |--C. cybele Butler 1866 B66a
         |           `--C. sareba B66a
         |         Daedelma B66a
         |           |--D. doraete B66a
         |           `--D. emilia Butler 1866 B66a
         |         Debis B66a
         |           |--D. chandica B66a
         |           |--D. europa WM66
         |           |--‘Papilio’ halyma Fabricius 1793 [=Neonympha halyma, Satyrus halyma] B66b
         |           |--D. isabella Butler 1866 B66a
         |           `--D. portlandia B66b
         |         Hetaera B66a
         |           |--H. aurora [incl. H. andromeda] B66a
         |           |--H. esmeralda B66a
         |           |--H. harpalyce Butler 1866 B66a
         |           |--H. pellucida Butler 1866 B6a
         |           |--H. philis B66a
         |           `--H. pireta B66a
         |         Callidryas WM66
         |           |--C. alcmeone WM66
         |           |--C. pyranthe WM66
         |           `--C. scylla R13
         |         Messaris erymanthus WM66
         |         Diadema WM66
         |           |--D. auge WM66
         |           |--D. bolina WM66
         |           `--D. dexithea O90
         |         Hestina assimilis WM66
         |           |--H. a. assimilis Iw92
         |           `--H. a. shirakii Shirôzu 1955 Iw92
         |         Athyma leucothoe WM66
         |         Ergolis WM66
         |           |--E. coryta WM66
         |           `--E. merione A73
         |         Cyllo leda WM66
         |         Pterygospidea folus [incl. P. cicero] WM66
         |         Chionobas bore [incl. Papilio polixenes Fabricius 1793, Neonympha polixenes] B66b
         |         Enodia creola C37
         |         Amblyscirtes carolina C37
         |         Sarangesa eliminata B01
         |         Argyrophenga antipodum P27
         |         Archaeprepona GE05
         |         Heraclides cresphontes Im92
         |         Ericina G20
         |           |--E. cupido G20
         |           `--E. lysippus G20
         |         Telchinia violae [=Acraea violae] A73
         |         Chilasa clytia [incl. C. clytia f. dissimilis] A73
         |         Ixias pyrene A73
         |         Stallingsia maculosa EN20
         |         Aegiale hesperiaris EN20
         |         Meneris tulbaghia [=Aeropetes tulbaghia] EN20
         |--Papilionidae KP19
         `--+--+--Hesperiidae KP19
            |  `--Hedylidae [Hedyloidea] GE05
            |       |--Hedyle Guenée 1857 BR05
            |       `--Macrosoma hedylaria KP19, GE05
            `--+--Pieridae KP19
               `--+--Nymphalidae GE05
                  `--+--Riodinidae J03
                     `--Lycaenidae GE05

*Type species of generic name indicated

REFERENCES

[A73] Ashton, R. 1973. Butterflies of New Delhi (Papilionoidea). Journal of the Bombay Natural History Society 69 (3): 502–509.

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

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

[B66a] Butler, A. G. 1866a. Descriptions of some new exotic butterflies in the National Collection. Proceedings of the Zoological Society of London 1866: 39–42.

[B66b] Butler, A. G. 1866b. A monograph of the genus Euptychia, a numerous race of butterflies belonging to the family Satyridae; with descriptions of sixty species new to science, and notes on their affinities &c. Proceedings of the Zoological Society of London 1866: 458–504.

[B01] Butler, A. G. 1901. On a collection of butterflies from the Uganda Protectorate, forwarded by C. Steuart Betton, Esq., in 1900. Annals and Magazine of Natural History, series 7, 7: 562.

[C37] Clark, A. H. 1937. The butterflies of Virginia. Proceedings of the Entomological Society of Washington 39 (5): 116.

[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

[EN20] Eggli, U., & R. Nyffeler (eds) 2020. Illustrated Handbook of Succulent Plants: Monocotyledons 2^nd ed. Springer.

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

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

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[IR05] Idris, A. B., & A. Rizki. 2005. Notes on the tribe Ephialtini (Hymenoptera: Ichneumonidae: Pimplinae) of Malaysia. Serangga 10 (1–2): 111–126.

[Im92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[Iw92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[J03] Jong, R. de. 2003. Are there butterflies with Gondwanan ancestry in the Australian region? Invertebrate Systematics 17: 143–156.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[M86] Meyrick, E. 1886. On some Lepidoptera from the Fly River. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 241–258.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[O90] Oberthür, C. 1890. Note suivante. Annales de la Société Entomologique de France, 6^e série 9: ccxli–ccxlii.

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

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

[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.

[WM66] Wallace, A. R., & F. Moore. 1866. List of lepidopterous insects collected at Takow, Formosa, by Mr. R. Swinhoe. Proceedings of the Zoological Society of London 1866: 355–365.

Last updated: 24 April 2022.

Heliothinae

Heliocheilus cramboides, copyright Donald Hobern.

Belongs within: Noctuidae.
Contains: Heliothis.

The Heliothinae, bollworms and related forms, are a group of moths whose larvae feed mostly on the reproductive organs (flowers, fruits and seeds) of plants, with some species being significant horticultural pests.

Characters (from Matthews 1999): Small to medium-sized, robust moths. Frons generally rounded, bulging with well-developed ventral lip. Labial palps moderately long, weakly upcurved. Antennal filiform or minimally biserrate in male, filiform in female. Compound eye smooth, unlashed. Head and thorax covered with mixed hairlike and spathulate scales. Hind wing venation trifine; vein M₂, if present, reduced and lying in median fold of hind wing. Male genitalia with valve usually simple and strap-like, quite long and narrow, rather flat, often curved distally, usually lacking processes except sometimes with a small ampulla; apical corona usually consisting of one or more rows of spine-like setae, occasionally absent; sacculus generally not well distinguished. Aedeagus short or long, straight or slightly curved, often bearing a denticulate 'scobinate' patch at apex; scobinate patch may occur at base of vesica; vesica varying from simple and sac-like to long and coiled, usually a number of diverticula present from a 'basal pouch'; vesica may bear cornuti. Female genitalia with papillae anales (ovipositor) varying from soft and pad-like with may setae to sclerotised and knife-like with fewer setae; ductus bursae varying from short to long, opening posteriorly through ostium bursae, variably ribbed and sclerotised; appendix bursae usually present, with ductus seminalis lying at apex; fundus bursae usually bearing from one to four signa. Larvae with spiny skin; prothoracic lateral setae with transverse position in later instars.

Heliothinae [Heliothidae] M99
    |  i. s.: Eutricopis nexilis M99
    |         Heliothodes diminutiva M99
    |         Periphanes ZK11
    |--Pyrrhia umbra (Hufnagel 1766) M99, JP05
    |--Melaporphyria M99
    |--+--Protadisura M99
    |  `--+--+--Heliolonche M99
    |     |  `--Schinia M99
    |     |       |--S. dobla M99
    |     |       `--S. florida M99
    |     `--Adisura Moore 1881 [incl. Astonycha Turner 1920] M99
    |          |--*A. atkinsoni Moore 1881 M99
    |          |--A. cana M99
    |          |--A. litarga (Turner 1920) [=*Astonycha litarga] M99
    |          |--A. marginalis (Walker 1858) (see below for synonymy) M99
    |          |--A. parva M99
    |          `--A. stigmatica M99
    `--+--Heliothis M99
       |--+--Australothis Matthews 1991 [=Australoverpa Matthews 1991] M99
       |  |    |--*A. rubrescens (Walker 1858) (see below for synonymy) M99
       |  |    |--A. exopisso Matthews 1999 M99
       |  |    |--A. tertia (Roepke 1941) [=Heliothis tertia] M99
       |  |    `--A. volatilis M99
       |  `--Helicoverpa Hardwick 1965 M99
       |       |  i. s.: H. assulta (Guenée 1852) (see below for synonymy) M99
       |       |         H. fletcheri M99
       |       |         H. hardwicki Matthews 1999 M99
       |       |         H. prepodes (Common 1985) [=Heliothis prepodes] M99
       |       |--H. punctigera (Wallengren 1860) [=Heliothis punctigera; incl. Chloridea marmada Swinhoe 1918] M99
       |       `--+--H. gelotopoeon M99
       |          `--+--H. hawaiiensis M99
       |             `--+--*H. armigera (Hübner 1803–1808) M99 (see below for synonymy)
       |                `--+--H. confusa M99
       |                   `--H. zea M99
       `--Heliocheilus Grote 1865 [incl. Canthylidia Butler 1886, Raghuva Moore 1881, Rhodosea Grote 1883] M99
            |--*H. paradoxus Grote 1865 M99
            |--H. abaccheutus Matthews 1999 M99
            |--H. aberrans (Butler 1886) [=Heliothis aberrans] M99
            |--H. albipunctella M99
            |--H. albivenata (Montague 1914) (see below for synonymy) M99
            |--H. aleurota (Lower 1902) (see below for synonymy) M99
            |--H. atrilinea (Turner 1943) [=Canthylidia atrilinea] M99
            |--H. canusina (Swinhoe 1901) (see below for synonymy) M99
            |--H. cistella (Swinhoe 1901) [=Canthylidia cistella] M99
            |--H. cladotus Swinhoe 1901 (see below for synonymy) M99
            |--H. confertissima (Walker 1865) [=Leucania confertissima, *Raghuva confertissima] M99
            |--H. confundens (Warren 1913) [=Canthylidia confundens] M99
            |--H. cramboides (Guenée 1852) (see below for synonymy) M99
            |--H. eodora (Meyrick 1902) [=Heliothis eodora; incl. Canthylidia tenuistria Turner 1902] M99
            |--H. ferruginosa (Turner 1911) [=Canthylidia ferruginosa; incl. C. epigrapha Turner 1920] M99
            |--H. flavitincta (Lower 1908) [=Melicleptria flavitincta] M99
            |--H. halimolimnus Matthews 1999 M99
            |--H. ionola (Swinhoe 1901) [=Adisura ionola, Melicleptria ionola] M99
            |--‘*Rhodosea’ julia Grote 1883 M99
            |--H. melibaphes (Hampson 1903) [=Melicleptria melibaphes] M99
            |--H. mesoleuca (Lower 1902) [=Canthylidia mesoleuca, Melicleptria mesoleuca] M99
            |--H. moribunda (Guenée 1852) M99 (see below for synonymy)
            |--H. neurota (Lower 1903) (see below for synonymy) M99
            |--H. pallida (Butler 1886) (see below for synonymy) M99
            |--H. puncticulata (Warren 1913) [=Canthylidia puncticulata] M99
            |--H. ranalaetensis Matthews 1999 M99
            |--H. rhodopolia (Turner 1911) [=Canthylidia rhodopolia] M99
            |--H. thelycritus Matthews 1999 M99
            `--H. vulpinotatus Matthews 1999 M99

Adisura marginalis (Walker 1858) [=Anthophila marginalis; incl. Heliothis delicia Felder & Rogenhofer 1874, A. dulcis Moore 1881, A. purgata Warren 1913, A. similis Moore 1881] M99

*Australothis rubrescens (Walker 1858) [=Thalpophila rubrescens; incl. Chloridea aresca Turner 1911, C. caesia Warren 1913, Heliothis hyperchroa Turner 1920, C. rufa Warren 1913] M99

*Helicoverpa armigera (Hübner 1803–1808) M99 [=Noctua armigera M99, Heliothis (Helicoverpa) armigera M83; incl. N. barbara Fabricius 1794 (nom. rej.) M99, Helicoverpa armigera commoni Hardwick 1965, Heliothis conferta Walker 1857 M99, Helic. armigera conferta M99, Heliothis armigera ab. fusca Cockerell 1889 M99, Heliothis guidelii Costantino 1922 M99, Heliothis obsoleta (Fabricius 1793) M99, GM79, Heliothis pulverosa Walker 1857 M99, Heliothis rama Bhattacharjee & Gupta 1972 M99, Chloridea obsoleta ab. rufa Warren 1911 non C. rufa Warren 1913 M99, Heliothis uniformis Wallengren 1860 M99]

Helicoverpa assulta (Guenée 1852) [=Heliothis assulta; incl. Helic. assulta afra Hardwick 1965, Heliothis separata Walker 1857, Heliothis succinea Moore 1881, Heliothis temperata Walker 1857] M99

Heliocheilus albivenata (Montague 1914) [=Melicleptria albivenata; incl. M. bipartita Strand 1924, M. stramineipicta Strand 1924, Canthylidia zorophanes Turner 1929, C. zurophanes (l. c.)] M99

Heliocheilus aleurota (Lower 1902) [=Melicleptria aleurota, Canthylidia alleurota (l. c.); incl. C. cana Turner 1943, C. aleurota ab. spissata Warren 1913] M99

Heliocheilus canusina (Swinhoe 1901) [=Canthylidia canusina; incl. C. anemodes Lower 1902, C. canusina ab. intacta Warren 1913] M99

Heliocheilus cladotus Swinhoe 1901 [incl. Canthylidia cladotus ab. densata Warren 1913, C. cladotus ab. hyalina Warren 1913, C. cladotus ab. hyalosticta Warren 1913, C. cramboides ab. semigrisea Warren 1913] M99

Heliocheilus cramboides (Guenée 1852) [=Leucania cramboides; incl. Canthylidia capnoneura Turner 1932, C. discolor Warren 1913, Heliocheilus discolor, C. cramboides ab. intensa Warren 1913, C. nervosa Turner 1943, Heliothis neurias Meyrick 1902, Heliocheilus neurias, C. cramboides ab. semiochrea Warren 1913, C. sericea Warren 1913, C. sulphurea Warren 1913, C. sumbensis Warren 1913, C. venata Warren 1913, Heliocheilus venata] M99

Heliocheilus moribunda (Guenée 1852) M99 [=Leucania moribunda M99, Canthylidia moribunda P27; incl. C. arenosa Turner 1943 M99, Leucania invaria Walker 1856 M99, C. invaria pallescens Warren 1913 M99]

Heliocheilus neurota (Lower 1903) [=Tarache neurota; incl. Canthylidia clathrata Warren 1913, C. crocopepla Turner 1925, Oruza crocotoschema Turner 1936, C. eurhythma Turner 1915, C. clathrata ab. exesa Warren 1913, C. clathrata ab. notata Warren 1913, C. clathrata ab. pallida Warren 1913 non C. pallida Butler 1886] M99

Heliocheilus pallida (Butler 1886) [=*Canthylidia pallida; incl. C. albida Warren 1913, H. osmidus Swinhoe 1901, C. rosea Warren 1913] M99

*Type species of generic name indicated

REFERENCES

[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.

[JP05] Jungmann, E., & U. Poller. 2005. Neunachweise und Wiederfunde für die Schmetterlingsfauna (Insecta; Lepidoptera) des Altenburger Landes. Mauritiana 19 (2): 317–326.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

[M99] Matthews, M. 1999. Heliothine Moths of Australia: A guide to bollworms and related noctuid groups. CSIRO Publishing.

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[ZK11] Zahiri, R., I. J. Kitching, J. D. Lafontaine, M. Mutanen, L. Kaila, J. D. Holloway & N. Wahlberg. 2011. A new molecular phylogeny offers hope for a stable family level classification of the Noctuoidea (Lepidoptera). Zoologica Scripta 40 (2): 158–173.