Variety of Life

Blue underwing Catocala fraxini, copyright Harald Süpfle.

Belongs within: Erebidae.
Contains: Melipotini, Euclidiini, Ophiusini, Poaphilini.

The Erebinae are a clade of noctuoid moths identified by molecular data, members of which have a smooth apex on the proboscis with all the styloconic sensilla placed dorsally. Females have the seventh sternite reduced and divided into two lobes with with ostium bursae situated in the cleft between the lobes. Larvae are slender semi-loopers with a pair of dorsolateral tubercles on the eighth abdominal segment.

Erebinae [Catocalinae]
    |--+--Ugia Walker 1858 ZK11, Z21
    |  |    `--U. insuspecta ZK11
    |  `--Acantholipes Lederer 1857 ZK11, FL05 [Acantholipini]
    |       |--A. circumdata ZK11
    |       `--A. regularis ZK11
    `--+--Sypnini ZK11
       |    |--Sypnoides fumosa ZK11
       |    `--Sypna ZK11
       `--+--Thysania Dalman 1824 FL05 [Thysaniini ZK11]
          |    |--T. agrippina DS73
          |    `--T. zenobia DS73
          |--+--Pandesma [Pandesmini] ZK11
          |  |    |--P. costalis S01
          |  |    |--P. pagana Swinhoe 1901 S01
          |  |    |--P. robusta ZK11
          |  |    `--P. submurina MC13
          |  `--+--Heteropalpia acrosticta ZK11
          |     `--+--Pericyma Herrich-Schäffer 1851 FL05 [Pericymini ZK11]
          |        |    `--P. cruegeri ZK11
          |        `--+--Catephiini [Catephidae] ZK11
          |           |    |--Anophia acronyctoides WM66
          |           |    `--Catephia Ochsenheimer 1816 FL05
          |           |         `--C. alchymista ZK11
          |           `--Sphingomorpha chlorea ZK11
          `--+--+--+--+--Serrodes campana ZK11
             |  |  |  `--Melipotini ZK11
             |  |  `--+--Erebus Latreille 1910 FL05 [Erebini ZK11]
             |  |     |    |--E. ephesperis ZK11
             |  |     |    `--E. odora H01
             |  |     `--Erygia apicalis ZK11
             |  `--+--Sympis rufibasis ZK11
             |     `--+--Anisoneura salebrosa ZK11
             |        `--Hulodini [Hulodides] ZK11
             |             |--Ericeia subcinerea ZK11
             |             `--Hulodes Guenée 1852 FL05
             |                  `--H. caranea ZK11
             `--+--+--Hypopyrini [Hypopyridae] ZK11
                |  |    |--Hypopyra Guenée 1852 FL05
                |  |    |    `--H. capensis ZK11
                |  |    `--Spirama ZK11
                |  `--+--Ercheia Walker 1858 FL05 [Ercheini ZK11]
                |     |    `--E. cyllaria ZK11
                |     `--+--Pantydia ZK11
                |        |    |--P. capistrata MC13
                |        |    |--P. diemeni ZK11
                |        |    `--P. sparsa G84
                |        `--Euclidiini ZK11
                |--+--Catocalini [Catocalidi] ZK11
                |  |    |--Catocala Schrank 1802 FL05
                |  |    |    |--C. fraxini DS73
                |  |    |    |--C. koreana Staudinger 1892 I92
                |  |    |    |--C. nupta DS73
                |  |    |    |--C. promissa DS73
                |  |    |    `--C. sponsa (Linnaeus 1767) JP05
                |  |    `--Ulotrichopus Wallengren 1860 Z21
                |  |         |--U. macula ZK11
                |  |         `--U. variegata Hampson 1902 Z21
                |  `--Audeini ZK11
                |       |--Audea Walker 1858 Z21
                |       |    `--A. bipunctata ZK11
                |       `--Crypsotidia Rothschild 1901 Z21
                `--+--Ophiusini ZK11
                   `--Poaphilini LS13

Erebinae incertae sedis:
  Dasypodia selenophora NC91, MC13
  Diatenes MC13
    |--D. aglossoides MC13
    `--D. marmarinopa MC13
  Donuca memorabilis MC13
  Eudesmeola lawsoni MC13
  Prorocopis melanochorda MC13
  Lyncestis acontioides [=Melipotis acontioides] PP72
  Psorya hadesia PP72
  Leistera Swinhoe 1909 Z21
    `--*L. pulchristrigata (Bethune-Baker 1906) [=Catephia pulchristrigata] Z21
  Cosmophila Boisduval 1833 Z21
    `--C. flava (Fabr. 1775) P27
  Homodes Guenée 1852 Z21
  Hypocala Guenée 1852 [Hypocalini] Z21
  Tamba Hübner 1823 Z21
  Arytrura John 1912 [Arytrurini] FL05
  Brevipecten FL05
  Ascalapha Hübner 1809 FL05, Z21
  Hemeroblemma FL05
  Latebraria FL05
  Antiblemma FL05
  Athyrma FL05
  Omopterini Z21
    |--Zale Hübner 1818 FL05, Z21 [incl. Omoptera Guérin-Méneville 1832 FL05]
    `--Matigramma Grote 1872 Z21
  Amphigonia Guenée 1852 [Amphigoniidae] FL05
  Anydrophila John 1909 [Anydrophilini] FL05
  Ctenusa Hampson 1910 [Ctenusini] FL05
  Hypogramma Guenée 1852 [Hypogrammidae] FL05
  Ommatophora Guenée 1852 [Ommatophoridae] FL05
  Speiredonia Hübner 1823 [Speiredoniinae] FL05
    |--S. mutabilis ZS10
    `--S. zamis ZS10
  Tachosa Walker 1869 [Tachosini] FL05
  Tinolius Walker 1855 [Tinoliidae] FL05
  Yrias Guenée 1852 [Yriasidae] FL05
  Thermesia Hübner 1823 [Thermesidae] FL05

*Type species of generic name indicated

REFERENCES

[DS73] Dickens, M., & E. Storey. 1973. The World of Moths. Macmillan Publishing Co., Inc.: New York.

[FL05] Fibiger, M., & J. D. Lafontaine. 2005. A review of the higher classification of the Noctuoidea (Lepidoptera) with special reference to the Holarctic fauna. Esperiana 11: 7–92.

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

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

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

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

[LS13] Lafontaine, J. D., & B. C. Schmidt. 2013. Comments on differences in classification of the superfamily Noctuoidea (Insecta, Lepidoptera) between Eurasia and North America. ZooKeys 264: 209–217.

[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.

[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).

[PP72] Parkin, P., D. T. Parkin, A. W. Ewing & H. A. Ford. 1972. A report on the arthropods collected by the Edinburgh University Galapagos Islands expedition, 1968. Pan-Pacific Entomologist 48 (2): 100–107.

[P27] Philpott, A. 1927. New Zealand Lepidoptera: notes and descriptions. Transactions and Proceedings of the New Zealand Institute 57: 703–709.

[S01] Swinhoe, C. 1901. New genera and species of eastern and Australian moths (continued). Annals and Magazine of Natural History, series 7, 7: 489–501.

[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.

[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.

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

[Z21] Zilli, A. 2021. Tabwecala robinsoni gen. nov., sp. nov., from Vanuatu and its systematic postion in the 'Ophiusini-Poaphilini' clade (Lepidoptera, Erebidae). Nota Lepidopterologica 44: 193–211.

Barea consignatella, copyright Donald Hobern.

Belongs within: Gelechioidea.

The Oecophorinae are a group of moths in which Rs and M₁ of the hind wing are nearly parallel, the abdominal terga are densely spinose, and the male usually has a beak-like gnathos (Nielsen & Common 1991).

<==Oecophorinae NC91
    |--Garrha carnea NC91
    |--Thema chlorochyta NC91
    |--Chrysonoma fascialis NC91
    |--Hippomacha callista NC91
    |--Aristeis hepialella NC91
    |--Heliocausta hemiteles NC91
    |--Hofmannophila pseudospretella NC91
    |--Endrosis sarcitrella NC91
    |--Oecophora Latreille 1802 L02
    |    |--*O. sulphurella [=Tinea sulphurella] L02
    |    `--O. linneella G20
    |--Philobota NC91
    |    |--P. disjunctella P27
    |    |--P. productella NC91
    |    `--P. xiphostola P27
    |--Wingia NC91
    |    |--W. aurata ZS10
    |    |--W. lambertella P27
    |    `--W. rectiorella NC91
    `--Barea NC91
         |--B. ambigua L27
         |--B. confusella L27
         |--B. consignatella NC91
         `--B. leucocephala P27

*Type species of generic name indicated

REFERENCES

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

[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.

[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.

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

Stathmopoda melanochra, copyright Donald Hobern.

Belongs within: Gelechioidea.

The Stathmopodinae are a group of usually very small moths that commonly rest with the hind wings raised (Nielsen & Common 1991).

Characters (from Nielsen & Common 1991): Usually very small; head smooth-scaled; wings very narrow; fore wing lacking pterostigma, R₂ well before upper angle of discal cell, female with subcubital retinaculum only; hind tibiae and tarsi usually bearing whorls of stiff bristles.

<==Stathmopodinae [Stathmopodidae, Tinaegeriidae] NC91
    |--Snellenia lineata NC91
    |--Pseudaegeria phlogina NC91
    |--Coracistis erythrocosma NC91
    |--Hieromantis NC91
    |--Calicotis NC91
    |--Vanacela C70
    `--Stathmopoda NC91
         |--S. arachnophthora NC91
         |--S. callichrysa NC91
         |--S. cephalaea NC91
         |--S. chalcotypa NC91
         |--S. coracodes P27
         |--S. crocophanes NC91
         |--S. melanochra NC91
         |--S. phlegyra L27
         `--S. skelloni L27

*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.

[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).

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

Cryptophasa rubescens, copyright Donald Hobern.

Belongs within: Gelechioidea.

The Xyloryctinae are a group of small to medium-sized moths in which the hind wings have veins Rs and M₁ stalked or arising from the same point and divergent, and Sc+R₁ diverging from Rs well before the end of the cell. The gnathos in the male is not spinose (Nielsen & Common 1991).

<==Xyloryctinae [Xyloryctidae] NC91
    |--Telecrates laetiorella NC91
    |--Odites pudica P27
    |--Chalarotona intabescens NC91
    |--Eschatura lemurias P27
    |--Uzucha humeralis NC91
    |--Catoryctis eugramma NC91, P27
    |--Neodrepta luteotactella G84
    |--Oegocniites borisjaki RJ93
    |--Lichenaula NC91
    |    |--L. choriodes NC91
    |    `--L. lichenea NC91
    |--Scieropepla NC91
    |    |--S. reversella P27
    |    `--S. typhicola NC91
    |--Maroga NC91
    |    |--M. melanostigma NC91
    |    `--M. unipunctana P27
    |--Xylorycta NC91
    |    |--X. luteotactella NC91
    |    |--X. melaleuca P27
    |    `--X. strigata NC91
    `--Cryptophasa NC91
         |--C. ecclesiastis Meyrick 1887 M87
         |--C. irrorata M87
         |--C. leucadelpha Meyrick 1887 M87
         |--C. melanostigma C70
         |--C. nubila P27
         `--C. rubescens 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.

[M87] Meyrick, E. 1887. Descriptions of new Lepidoptera. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1037–1048.

[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.

[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.

Triangle plume Platyptilia gonodactyla, copyright Ilia Ustyantsev.

Belongs within: Obtectomera.

The Pterophoridae, plume moths, are a group of small moths in which the hind wings are usually deeply cleft into three plumes.

Characters (from Nielsen & Common 1991): Small; head usually smooth-scaled; ocelli and chaetosemata absent; antennae filiform; proboscis unsealed; maxillary palps minute, one-segmented; labial palps slender or stout, porrect or upturned; legs long, slender; epiphysis present; spurs 0-2-4, prominent; hind tibiae sometimes with specialised scaling; fore wing divided by terminal clefts into two or three plumes, rarely four plumes, or entire; with complex wing-folding mechanism, pterostigma sometimes present, wing-locking microtrichia absent, chorda and M absent in cell, one branch of R often absent, M₁ and M₂ weak or absent, CuP usually present, 1A+2A without fork; hind wing deeply cleft into three plumes, rarely entire, underside with double series of specialised scales along M3 and CuA₂ or CuA₁, CuP absent, female frenulum of one or two bristles; abdomen long and slender, tympanal organs absent. Eggs of flat type, oval, smooth. Larva usually with numerous secondary, often modified setae, sometimes on verrucae, prothorax with 3 L setae, ventral prolegs long, crochets uniordinal in mesoseries or mesal penellipse; larvae usually leaf-mining at first, later exposed, eroding surface of leaves or burrowing in flower buds or in stems. Pupa slender, smooth or spined, often with numerous setae on head, thorax and abdomen, pilifers present; fully exposed, attached by hooked setae on underside of abdominal segments 9 and 10.

<==Pterophoridae
    |  i. s.: Exelastis atomosa B88
    |         Buckleria paludum RD77
    |--Agdistinae NC91
    |    |--Ochyrotica kurandica NC91
    |    |--Agdistopis halieutica NC91
    |    `--Agdistis staticis RD77
    |--Pterophorinae KP19
    |    |--Lantanophaga pusillidactyla KP19
    |    `--+--Emmelina monodactyla KP19
    |       `--Pterophorus KP19
    |            |--P. aptalis NC91
    |            |--P. didactylus L02
    |            |--P. oligocenicus Bigot et al. 1986 RJ93
    |            |--P. pentadactyla KP19 [=Aciptilia pentadactyla H04]
    |            `--P. rhododactylus [=Cnaemidophorus rhododactylus] H04
    `--Platyptilinae NC91
         |--Cosmoclostis aglaodesma NC91, ZS10
         |--Stangeia xerodes NC91
         |--Stenoptilia NC91
         |    |--S. lithoxesta P27
         |    `--S. zophodactyla NC91
         |--Sphenarches C70
         |    |--S. anisodactylus NC91
         |    `--S. caffer P27
         |--Trichoptilus C70
         |    |--T. adelphodes P27
         |    `--T. wahlbergi C70
         `--Platyptilia NC91
              |--P. acanthodactyla M38
              |--P. aeolodes L27
              |--P. falcatalis P27
              |--P. gonodactyla P01
              `--P. omissalis 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.

[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.

[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.

[M38] Muesebeck, C. F. W. 1938. Three new reared species of Apanteles from California (Hymenoptera: Braconidae). Proceedings of the Entomological Society of Washington 40 (7): 201–204.

[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).

[P01] Pável, J. 1901. Lepkék [Lepidopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 171–177. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

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

[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.

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

Alucita xanthodes, copyright Alison Pearson.

Belongs within: Obtectomera.

The Alucitoidea are a group of moths in which the hind wing is deeply divided into six or seven plumes.

Characters (from Nielsen & Common 1991): Small, head smooth scaled, ocelli present or absent; chaetosemata absent; scape without pecten; proboscis unsealed; epiphysis present; spurs 0-2-4 or 0-0-2; wings often divided into plumes, CuP absent in fore wing; tympanal organs absent; S2 with V-shaped sclerotisation, T3–6 and sometimes T2–7 with anterior, transverse, sclerotised band set with spines. Larva densely spinulose, without secondary setae, L group on prothorax bisetose on one pinaculum, ventral prolegs with uniordinal crochets. Pupa without dorsal abdominal spines, in cocoon, not protruded at ecdysis.

<==Alucitoidea NC91
    |--Tineodidae [Oxychirotidae] NC91
    |    |--Tineodes adactylalis NC91
    |    |--Tanycnema anomala NC91
    |    |--Cenoloba obliteralis NC91
    |    |--Oxychirota paradoxa NC91
    |    |--Palaeodes samealis NC91
    |    `--Euthrausta NC91
    |         |--E. holophaea C70
    |         `--E. oxyprora NC91
    `--Alucita [Alucitidae] GE05
         |--A. cereella Bosc 1800 B00
         |--A. desmodactyla KP19
         |--A. monospilalis P27
         |--A. phricodes NC91
         |--A. pygmaea NC91
         |--A. vittata G20
         |--A. vitella L02
         |--A. xanthodes C70
         `--A. xylostella L02

*Type species of generic name indicated

REFERENCES

[B00] Bosc, C. 1800. Description de trois espèces de lepidoptères de la Caroline. Bulletin des Sciences, par la Societé Philomathique de Paris 2 (39): 114–115.

[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.

[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.

[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 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.

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.

Grizzled skipper Pyrgus malvae, copyright Gail Hampshire.

Belongs within: Hesperiidae.

The Pyrginae, flats, are a cosmopolitan group of relatively large, robust and fast-flying skippers whose larvae feed on a wide range of dicotyledonous plants (Braby 2000a).

Characters (from Braby 2000a): Antenna more than half length of costa of fore wing; third segment of labial palp erect or porrect, short to medium in length, stout; fore wing with vein M₂ straight near origin, arising midway between M₁ and M₃ or closer to M₁ than M₃; hind wing with cell normal, termen rounded; male may have costal gold or linear scent-pouch, containing numerous sex-scales, on costa near base of fore wing.

<==Pyrginae [Pyrgidae] J03
    |--Capila J03
    |--Telemiades J03
    |--Eagris J03
    |--Celaenorrhinus Hübner 1919 B00a
    |    `--C. fritzgaertneri (Balley 1880) FS54
    |--Exometoeca Meyrick 1888 B00a
    |    `--E. nycteris Meyrick 1888 B00a
    |--Euschemon Doubleday 1846 [Euschemoninae] B00a
    |    `--E. rafflesia (Macleay 1826) B00a
    |         |--E. r. rafflesia B00a
    |         `--E. r. alba Mabille 1903 B00a
    |--Netrocoryne Felder & Felder 1867 B00a
    |    |--N. repanda Felder & Felder 1867 B00a
    |    |    |--N. r. repanda B00a
    |    |    `--N. r. expansa Waterhouse 1932 B00a
    |    `--N. thaddaeus (Hewitson 1876) B00a
    |--Chaetocneme Felder 1860 B00a
    |    |--C. beata (Hewitson 1867) B00a
    |    |--C. critomedia (Guérin-Méneville 1831) B00a
    |    |    |--C. c. critomedia B00a
    |    |    `--C. c. sphinterifera (Fruhstorfer 1910) B00a
    |    |--C. denitza (Hewitson 1867) B00a
    |    `--C. porphyropsis (Meyrick & Lower 1902) B00a
    |--Tagiades Hübner 1819 B00a
    |    |--T. flesus B01a
    |    |--T. japetus (Stoll 1781) B00a
    |    |    |--T. j. japetus B00a
    |    |    `--T. j. janetta Butler 1870 B00a
    |    `--T. nestus (Felder 1860) B00b
    |         |--T. n. nestus B00b
    |         `--T. n. korela Mabille 1891 B00b
    `--Pyrgus Hübner 1819 BR05
         |--P. dromus B01a
         |--P. machacosa B01b
         |--P. maculatus (Bremer & Grey 1852) I92
         |--P. malvae (Linnaeus 1758) I92
         |--P. notata (Blanchard 1852) FS54
         |--P. orcus (Cramer 1782) S55
         `--P. syrichtus (Fabricius 1775) FS54
              |--P. s. syrichtus FS54
              `--P. s. montivagus (Reakirt 1866) FS54

*Type species of generic name indicated

REFERENCES

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

[B00a] Braby, M. F. 2000a. Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).

[B00b] Braby, M. F. 2000b. Butterflies of Australia: their identification, biology and distribution vol. 2. CSIRO Publishing: Collingwood (Victoria).

[B01a] Butler, A. G. 1901a. An account of a collection of butterflies made by the Rev. K. St. Aubyn Rogers between Mombasa and the forests of Taveta. Annals and Magazine of Natural History, series 7, 7: 22–35.

[B01b] Butler, A. G. 1901b. An account of a collection of butterflies obtained by Lord Delamere, chiefly at Munisu, near Mount Kenya. Annals and Magazine of Natural History, series 7, 7: 197–203.

[FS54] Franz, E., & H. Schröder. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador. Senckenbergiana Biologica 35: 75–87.

[I92] 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.

[S55] Schröder, H. 1955. Eine Falter-Ausbeute aus dem westlichen Bolivien. (Ins. Lepid. Rhopal.) Senckenbergiana Biologica 36: 329–338.

Silver sedge-skipper Hesperilla crypsargyra, copyright Donald Hobern.

Belongs within: Hesperiidae.

Hesperilla, sedge-skippers, is a genus of skippers found in coastal and subcoastal regions of Australia, with larvae feeding on species of Cyperaceae (Braby 2000).

Characters (from Braby 2000): Club of antenna bent before middle, apiculus long and pointed; third segment of labial palp of moderate length; fore wing with vein CuA₂ arising midway between end and base of cell, vein 1A+2A in male relatively straight; hind wing with vein CuA₁ arising closer to M₃ than CuA₂; hind tibia with two pairs of spurs; male with linear patch of sex-scales. Pupal cap strongly produced and heavily sclerotised; abdominal segments of pupa containing long stiff setae.

<==Hesperilla Hewitson 1868 B00
    |--H. chrysostricha (Meyrick & Lower 1902) B00
    |    |--H. c. chrysotricha (see below for synonymy) B00
    |    `--H. c. cyclospila (Meyrick & Lower 1902) B00
    |--H. crypsargyra (Meyrick 1888) B00
    |    |--H. c. crypsargyra [incl. H. crypsargyra lesouefi Tindale 1953] B00
    |    `--H. c. hopsoni Waterhouse 1927 B00
    |--H. crypsigramma (Meyrick & Lower 1902) B00
    |--H. donnysa Hewitson 1868 B00
    |    |--H. d. donnysa (see below for synonymy) B00
    |    |--H. d. albina Waterhouse 1932 B00
    |    |--H. d. aurantia Waterhouse 1927 B00
    |    `--H. d. galena Waterhouse 1927 B00
    |--H. flavescens Waterhouse 1941 (see below for synonymy) B00
    |--H. furva Sands & Kerr 1973 B00
    |--H. idothea (Miskin 1889) B00
    |    |--H. i. idothea B00
    |    `--H. i. clara Waterhouse 1932 B00
    |--H. malindeva Lower 1911 [incl. H. malindeva dagoomba Johnson & Valentine 1994] B00
    |--H. mastersi Waterhouse 1900 B00
    |--H. ornata (Leach 1814) B00
    |    |--H. o. ornata B00
    |    `--H. o. monotherm Lower 1907 B00
    |--H. picta (Leach 1814) B00
    |--H. sarnia Atkins 1978 B00
    `--H. sexguttata Herrich-Schäffer 1869 B00

Hesperilla chrysotricha chrysotricha (Meyrick & Lower 1902) [incl. H. chrysotricha leucosia Waterhouse 1938, H. chrysotricha lunawanna Couchman 1949, H. chrysotricha naua Couchman 1949, H. chrysotricha plebeia Waterhouse 1927] B00

Hesperilla donnysa donnysa Hewitson 1868 [incl. H. donnysa delos Waterhouse 1941, H. donnysa diluta Waterhouse 1932, H. donnysa icaria Waterhouse 1941, H. donnysa patmos Waterhouse 1941, H. donnysa samos Waterhouse 1941] B00

Hesperilla flavescens Waterhouse 1941 [=H. donnysa flavescens; incl. H. donnysa flavia Waterhouse 1941, H. flavescens flavia] B00

*Type species of generic name indicated

REFERENCES

[B00] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).

Western grass dart Taractrocera papyria agraulia, copyright Fred Hort.

Belongs within: Hesperiidae.
Contains: Telicota, Ocybadistes, Suniana.

The Hesperiinae are a cosmopolitan group of skippers whose larvae feed on species of monocotyledons, particularly on grasses (Braby 2000).

Characters (from Braby 2000): Length of antenna variable, less or greater than half length of costa of fore wing; labial palp with third segment porrect or erect, short and stout, or long and slender; fore wing with vein M₂ downwardly curved near origin, arising closer to M₃ than M₁; hind wing with cell normal with discocellular vein between M₂ and M₃ angled or inclined towards tornus or termen, termen rounded or distinctly lobed and produced at tornus.

<==Hesperiinae J03
    |--+--Plastingia J03
    |  |--Prada J03
    |  `--Tiacellia J03
    |--Hesperia J03
    |    |--H. agna Moore 1865 WM66
    |    |--H. comma P01
    |    |--H. conjuncta R13
    |    |--H. divodasa Moore 1865 WM66
    |    |--H. evalthe [=Papilio evalthe] L02
    |    |--H. licas [=Papilio licas] L02
    |    |--H. lineola P01
    |    |--H. malvae L02
    |    `--H. proteus L02
    `--+--+--Pastria J03
       |  `--+--Oriens angustula (Herrich-Schäffer 1869) J03, B00
       |     `--Cephrenes Waterhouse & Lyell 1914 J03, B00
       |          |--C. augiades (Felder 1860) B00
       |          |    |--C. a. augiades B00
       |          |    `--C. a. sperthias (Felder 1862) B00
       |          `--C. trichopepla (Lower 1908) B12
       `--+--+--Banta J03
          |  |--Kobrona J03
          |  |--Telicota J03
          |  `--+--Mimene Joicey & Talbot 1917 J03, B00
          |     |    `--M. atropatene (Fruhstorfer 1911) B00
          |     `--Sabera Swinhoe 1908 J03, B00
          |          |--S. caesina (Hewitson 1866) B00
          |          |    |--S. c. caesina B00
          |          |    `--S. c. albifascia (Miskin 1889) B00
          |          |--S. dobboe (Plötz 1885) B00
          |          |    |--S. d. dobboe B00
          |          |    `--S. d. autoleon (Miskin 1889) B00
          |          `--S. fuliginosa (Miskin 1889) B00
          `--+--Arrhenes Mabille 1904 J03
             |    |--A. dschilus (Plötz 1885) B00
             |    |    |--A. d. dschilus B00
             |    |    `--A. d. iris (Waterhouse 1932) B00
             |    `--A. marnas (Felder 1860) B00
             |         |--A. m. marnas B00
             |         `--A. m. affinis (Waterhouse & Lyell 1912) B00
             `--+--Potanthus J03
                `--+--Ocybadistes J03
                   `--+--Suniana J03
                      `--Taractrocera Butler 1870 J03, B00
                           |--T. anisomorpha (Lower 1911) B12
                           |--T. dolon (Plötz 1884) B00
                           |    |--T. d. dolon B00
                           |    `--T. d. diomedes Waterhouse 1933 B00
                           |--T. ilia Waterhouse 1932 B00
                           |--T. ina Waterhouse 1932 B00
                           `--T. papyria (Boisduval 1832) B00
                                |--T. p. papyria B00
                                `--T. p. agraulia (Hewitson 1868) B00

Hesperiinae incertae sedis:
  Pelopidas Walker 1870 B00
    |--P. agna (Moore 1866) B12
    |    |--P. a. agna B00
    |    `--P. a. dingo Evans 1949 B00
    |--P. lyelli (Rothschild 1915) B12
    `--P. mathias VHK02
         |--P. m. mathias F92
         `--P. m. oberthueri F92
  Notocrypta de Niceville 1889 B00
    `--N. waigensis (Plötz 1882) B00
         |--N. w. waigensis B00
         `--N. proserpina (Butler 1883) B00
  Parnara Moore 1881 B00
    |--P. amalia (Semper 1879) B00
    |--P. bada (Moore 1878) [=P. naso bada] B00
    |    |--P. b. bada B00
    |    `--P. b. sida (Waterhouse 1934) B00
    |--P. guttata NI11
    |--P. mathias P01
    `--P. ogasawaraensis Matsumura 1906 I92
  Borbo Evans 1949 B00
    |--B. cinnara (Wallace 1866) B00
    `--B. impar (Mabille 1883) B00
         |--B. i. impar B00
         |--B. i. lavinia (Waterhouse 1932) B00
         `--B. i. tetragraphus (Mabille 1891) B00
  Pseudoborbo Lee 1966 B00
    `--P. bevani (Moore 1878) [=Borbo bevani] B00

Nomen nudum: Hesperia cinnara Moore in Wallace & Moore 1866 WM66

*Type species of generic name indicated

REFERENCES

[B00] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).

[B12] Braby, M. F. 2012. The butterflies of El Questro Wilderness Park, with remarks on the taxonomy of the Kimberley fauna, Australia. Records of the Western Australian Museum 27 (2): 161–175.

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

[I92] 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.

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

[NI11] Norhafiza, A. F., & A. B. Idris. 2011. A catalogue of Ichneumoninae (Hymenoptera: Ichneumonidae) from Malaysia. Serangga 16 (1): 29–50.

[P01] Pável, J. 1901. Lepkék [Lepidopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 171–177. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

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

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

[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.

Duke of Burgundy Hamearis lucina, copyright Gilles San Martin.

Belongs within: Obtectomera.
Contains: Nymphidiini.

The Riodinidae, metalmarks, are a group of butterflies that are most diverse in the Neotropical region.

Characters (from Braby 2000, as Riodininae): Male fore leg non-functional, less than half length of mid and hind legs, tarsus fused into single segment, rarely with claws; hind wing usually with basal vein along costal margin, tornus may be produced, lobed or dentate but rarely tailed; males rarely with sex-scales. Larva with prothorax bearing pair of vibratory papillae, metathorax bearing pair of dorsolateral eversible organs, pair of eversible nectary organs present on segment 8.

<==Riodinidae [Nemeobiidae]
    |--Hamearinae [Nemeobiinae] J03
    |    |--Abisara J03
    |    |--Zemeros J03
    |    `--Hamearis lucina [=Nemeobius lucina] RD77
    `--Riodininae HH02
         |  i. s.: Praetaxila Fruhstorfer 1914 B00
         |           `--P. segecia (Hewitson 1861) B00
         |                |--P. s. segecia B00
         |                |--P. s. punctaria (Fruhstorfer 1904) B00
         |                `--P. s. yaniya Fruhsorfer 1914 B00
         |         Styx [Styginae] B00
         |           `--S. infernalis Staudinger 1875 B00
         |         Semomesia capanea KP19
         `--Nymphidiini HH02

Riodinidae incertae sedis:
  Petrocerus catiena HH02
  Callistium HH02
  Emesis HH02
    |--E. guttata [=Audre guttata] HH02
    |--E. mandana (Cramer 1780) S55
    `--E. xanthosa HH02
  Calydna caieta HH02
  Stalachtis HH02
  Apodemia mormo HH02
    |--A. m. mormo USDI77
    `--A. m. langei USDI77
  Polystichtis S55
    |--P. emylius S55
    |    |--P. e. emylius S55
    |    `--P. e. crispinella Stichel 1911 S55
    `--P. lasthenes FS54
         |--P. l. lasthenes FS54
         `--P. l. zeurippa Boisduval 1836 FS54
  Calephelis virginiensis FS54
    |--C. v. virginiensis FS54
    `--C. v. laverna (Godman & Salvin 1886) FS54
  Lymnas FS54
    |--L. cephise (Ménétriés 1855) FS54
    `--L. pixe (Boisduval 1910) FS54
  Esthemopsis caeruleata (Godman & Salvin 1878) FS54
  Mesosemia S55
    |--M. jeziella S55
    |    |--M. j. jeziella S55
    |    `--M. j. thyestes Druce 1878 S55
    `--M. judicialis Butler 1874 S55
  Lyropteryx apollonia Westwood 1851 S55
  Cyrenia martia Westwood 1851 S55
  Ancyluris S55
    |--A. aulestes S55
    |    |--A. a. aulestes S55
    |    `--A. a. eryxo (Saunders 1858) S55
    `--A. inca GE05
  Rhetus S55
    |--R. arcius GE05
    `--R. periander (Cramer 1777) S55
  Caria S55
    |--C. mantinea (Felder 1861) S55
    `--C. trochilus S55
         |--C. t. trochilus S55
         `--C. t. arete (Felder 1861) S55
  Baeotis nesaea S55
    |--B. n. nesaea S55
    `--B. n. bacaenita Schaus 1902 S55
  Amarynthis meneria (see below for synonymy) S55
  Peplia lamis [incl. P. lamis f. molpe (Hübner 1816)] S55
  Helicopis GE05
    |--H. acis R13
    `--H. cupido GE05
  Riodinella nympha GE05
  Voltinia GE05
    |--V. danforthi V-W04
    |--V. dramba GE05
    `--V. umbra V-W04
  Euselasia GE05
  Mesenopsis melanochlora GE05
  Eurybiini HH02
    |--Alesa amesis HH02
    `--Eurybia S55
         |--E. nicaea (Fabricius 1775) S55
         `--E. patrona GE05

Amarynthis meneria [incl. A. meneria f. maecenas (Fabricius 1793), A. meneria f. micalia (Cramer 1776)] S55

*Type species of generic name indicated

REFERENCES

[B00] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 2. CSIRO Publishing: Collingwood (Victoria).

[FS54] Franz, E., & H. Schröder. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador. Senckenbergiana Biologica 35: 75–87.

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

[HH02] Hall, J. P. W., & D. J. Harvey. 2002. Basal subtribes of the Nymphidiini (Lepidoptera: Riodinidae): phylogeny and myrmecophily. Cladistics 18: 539–569.

[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.

[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.

[S55] Schröder, H. 1955. Eine Falter-Ausbeute aus dem westlichen Bolivien. (Ins. Lepid. Rhopal.) Senckenbergiana Biologica 36: 329–338.

[USDI77] United States Department of the Interior. 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

[V-W04] Vane-Wright, D. 2004. Butterflies at that awkward age. Nature 428: 477–480.