Nymphalidae

Melinaea maenius, photographed by Rich Hoyer.

Belongs within: Obtectomera.
Contains: Catagramma, Adelpha, Satyrinae, Morphinae, Charaxinae, Heliconiinae, Limenitidinae, Biblidinae, Nymphalinae, Apaturinae, Danaini.

The Nymphalidae, brush-footed butterflies, are a cosmopolitan group of butterflies in which only four normal walking legs are usually present. The first pair of legs is reduced and pressed against the thorax except in females of the basal subfamily Libytheinae. The Libytheinae are a small but widely distributed subfamily whose larvae feed on plants of the genus Celtis (Braby 2000b).

Characters (from Braby 2000b): Adults with antennae relatively close together at base; fore wing with five branches in radial vein, some staked, and one anal vein; hind wing with humeral vein usually present (sometimes vestigial or absent), two anal veins, and tornus usually rounded, rarely tailed; both sexes with only four normal functional legs, fore legs substantially smaller in size, pressed against thorax and non-functional except in female Libytheinae, male with fore tarsus often reduced to a single elongate segment that may be clubbed or brush-like without an apical claw, female with fore tarsus usually five-segmented and clawed apically; fore tibia lacking epiphysis. Egg very variable in shape, may be spherical, almost spherical with apex flattened, dome-shaped, barrel-shaped, or elongate and subconical with apex slightly truncate; base always flattened; chorion often with strong longitudinal ribs and fine transverse ribs, sometimes smooth. Final instar larva cylindrical with fine hairs, may also have numerous bristles, branched spines or several long paired filaments; head sometimes bearing horns; osmeterium usually absent; crochets of prolegs in uni-, bi- or triordinal mesoseries. Pupa normally suspended by cremaster, occasionally lying unattached on ground, never provided with central silken girdle.

<==Nymphalidae
    |--Libytheinae [Libytheidae] WWN03
    |    |--Prolibythea vagabunda GE05
    |    |--Barbarothea florissanti GE05
    |    |--Libytheana Michener 1943 B00b
    |    `--Libythea Fabricius 1807 B00b
    |         |--L. carinenta (Cramer 1779) S55
    |         |--L. celtis WWN03
    |         `--L. geoffroy Godart 1824 B00b
    |              |--L. g. geoffroy B12
    |              |--L. g. genia Waterhouse 1938 B12
    |              `--L. g. nicevillei Olliff 1891 B00b
    `--+--+--+--Satyrinae WWN03
       |  |  `--+--Morphinae WWN03
       |  |     `--+--Charaxinae WWN03
       |  |        `--Calinaga Moore 1857 WWN03, B00b [Calinaginae]
       |  |             `--C. buddha WWN03
       |  `--+--+--Heliconiinae WWN03
       |     |  `--Limenitidinae WWN03
       |     `--+--+--Biblidinae WWN03
       |        |  `--+--Nymphalinae WWN03
       |        |     `--Apaturinae WWN03
       |        `--Cyrestinae WWN03
       |             |--Cyrestini WWN03
       |             |    |--Cyrestis thyodamas WWN03
       |             |    `--Dicorrhagia WWN03
       |             `--Pseudergolini WWN03
       |                  |--Amnosia decora WWN03
       |                  |--Stibochiona nicea WWN03
       |                  `--Pseudergolis WWN03
       `--Danainae [Danaidae, Euploeiinae, Ithomiinae, Limnadinae] WWN03
            |  i. s.: Hirsutis salvadoris (Staudinger 1885) FS54
            |         Mechanitis FS54
            |           |--M. doryssus Bates 1864 FS54
            |           |--M. istmia FS54
            |           |    |--M. i. istmia FS54
            |           |    `--M. i. lycidice Bates 1863 FS54
            |           `--M. polymnia [=Heliconius (Mechanitis) polymnia] G20
            |         Napeogenes tolosa (Hewitson 1856) FS54
            |         Dircenna S55
            |           |--D. dero (Hübner 1823) S55
            |           |--D. klugii (Geyer 1837) FS54
            |           `--D. loreta Haensch 1903 S55
            |         Leucothyris victorina FS54
            |           |--L. v. victorina FS54
            |           `--L. v. paula Weymer 1884 FS54
            |         Hymenitis FS54
            |           |--H. nero (Hewitson 1854) FS54
            |           `--H. oto (Hewitson 1854) FS54
            |         Thyridia confusa S55
            |           |--T. c. confusa S55
            |           `--T. c. psamathe (Godman & Salvin 1898) S55
            |         Xanthocleis psidii (Linnaeus 1758) S55
            |         Ceratinia S55
            |           |--C. nise S55
            |           |    |--C. n. nise S55
            |           |    `--C. n. peruensis (Haensch 1905) S55
            |           `--C. tutia S55
            |                |--C. t. tutia S55
            |                `--C. t. chanchamaya (Haensch 1905) S55
            |         Corbulis cassotis S55
            |           |--C. c. cassotis S55
            |           `--C. c. cajona (Haensch 1905) S55
            |         Heterosais edessa S55
            |           |--H. e. edessa S55
            |           `--H. e. nephele (Bates 1862) S55
            |         Archaeolycorea ferreirai GE05
            |         Parantica Moore 1880 NS07
            |--Danaini WWN03
            |--Tellervo Kirby 1894 B00b [Tellervinae, Tellervini WWN03]
            |    `--T. zoilus (Fabricius 1775) B00b
            |         |--T. z. zoilus B00a
            |         |--T. z. digulica B00a
            |         `--T. z. gelo Waterhouse & Lyell 1914 B00b
            `--Ithomiini WWN03
                 |--Greta oto WWN03
                 |--Ithomia patilla Hewitson 1852 FS54
                 `--Melinaea maenius GE05, S55
                      |--M. m. maenius S55
                      |--M. m chincha Druce 1876 S55
                      `--M. m. mothone (Hewitson 1866) S55

Nymphalidae incertae sedis:
  Aglais KIR02
    |--A. karaganica KIR02
    `--A. urticae A71
  Percnodaimon pluto (Fereday 1872) H82
  Philaethria dido S55
    |--P. d. dido S55
    `--P. d. ostara (Röber 1906) S55
  Eueides eanes S55
    |--E. e. eanes S55
    `--E. e. eanides Stichel 1903 S55
  Historis orion (Fabricius 1775) S55
  Gynaecia dirce (Linnaeus 1764) S55
  Pyrrhogyra S55
    |--P. edocla S55
    |    |--P. e. edocla S55
    |    `--P. e. maculata Staudinger 1888 S55
    `--P. otolais Bates 1864 S55
  Temenis laothoe S55
    |--T. l. laothoe S55
    |--T. l. ottonis Fruhstorfer 1907 S55
    `--T. l. violetta Fruhstorfer 1907 S55
  Nessaea obrinus (Linnaeus 1764) S55
  Myscelia sophronia (Godtart 1823) S55
  Eunica excelsa Godman & Salvin 1877 S55
  Catagramma S55
  Adelpha S55
  Doxocopa S55
    |--D. agathina (Cramer 1777) S55
    |--D. cherubina S55
    |    |--D. c. cherubina S55
    |    `--D. c. parva (Röber 1916) S55
    `--D. lavinia (Butler 1866) S55
  Polygrapha cyanea (Godman & Salvin 1868) S55
  Zaretes isidora S55
    |--Z. i. isidora S55
    `--Z. i. cacica Staudinger 1888 S55
  Paramo Adams & Bernard 1977 O08
    `--*P. aculata (Krüger 1924) [=Pedaliodes aculata] O08
  Blepolensis catherinae GE05
  Neorinopis sepulta GE05
  Eugonia GE05
    |--E. angularia R13
    `--E. atava GE05
  Batesia hypochlora GE05
  Hypanartia lethe (Fabricius 1793) S55
  Archimestra L88
  Issoria sinha TYM08
    |--I. s. sinha TYM08
    `--I. s. pelewensis Nakamura 1932 TYM08
  Callerebia cyclopia TYM08
    |--C. c. cyclopia TYM08
    `--C. c. yoshikurana Kishida & Nakamura 1941 TYM08
  Neorinopsis sepulta RJ93
  Limnas chrysippus B01a (see below for synonymy)
  Samanta B01a
    |--S. perspicua B01a
    `--S. simonsi B01a
  Monotrichtis B01b
    |--M. auricruda B01b
    |--M. kenia B01b
    `--M. sufitza B01a
  Neocoenyra B01b
    |--N. duplex B01a
    `--N. gregorii B01b
  Physcoenura leda B01a
  Panopea expansa B01a
  Euralia B01a
    |--E. deceptor B01a
    `--E. wahlbergi B01a
  Hamanumida daedalus B01a
  Euphaedra B01a
    |--E. neophron B01a
    `--E. violacea B01a
  Euryphene senegalensis B01a
  Hypanis ilithyia B01a
  Pardopsis punctatissima B01a
  Crenis boisduvali B01b
  Atella B01b
    |--A. columbina B01c
    `--A. phalantha B01b
  Neptidopsis ophione [incl. N. ophione var. velleda] B01b
  Byblia B01b
    |--B. ilithyia [incl. B. ilithyia var. simplex] B01b
    `--B. vulgaris B01b
  Planema B01b
    |--P. aganice B01b
    |--P. gea B01b
    `--P. montana [incl. P. bertha, P. meruana] B01b

Limnas chrysippus B01a [incl. L. alcippoides B01c, L. alcippus B01c, L. chrysippus var. dorippus B01a, L. chrysippus var. klugii B01b]

*Type species of generic name indicated

REFERENCES

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

[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).

[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.

[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.

[B01c] Butler, A. G. 1901c. 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.

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

[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.

[H82] Helmore, D. W. 1982. Drawings of New Zealand insects. Bulletin of the Entomological Society of New Zealand 8: 1–52.

[KIR02] Kozlov, M. V., V. D. Ivanov & A. P. Rasnitsyn. 2002. Order Lepidoptera Linné, 1758. The butterflies and moths (=Papilionida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 220–227. Kluwer Academic Publishers: Dordrecht.

[L88] Liebherr, J. K. 1988. General patterns in West Indian insects, and graphical biogeographic analysis of some circum-Caribbean Platynus beetles (Carabidae). Systematic Zoology 37 (4): 385–409.

[NS07] Nässig, W. A., & C. H. Schulze. 2007. A second species with diurnal males of the genus Eupterote from Indonesia: Eupterote (Eupterote) splendens sp. n. from Sulawesi (Insecta, Lepidoptera, Bombycoidea, Eupterotidae). Senckenbergiana Biologica 87 (2): 189–194.

[O08] Özdikmen, H. 2008. Nomenclatural changes for some preoccupied harvestman genus group names (Arachnida: Opiliones). Turkish Journal of Arachnology 1 (1): 37–43.

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

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

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

[TYM08] Tennent, W. J., M. Yasuda & K. Morimoto. 2008. Lansania Journal of arachnology and zoology—a rare and obscure Japanese natural history journal. Archives of Natural History 35 (2): 252–280.

[WWN03] Wahlberg, N., E. Weingartner & S. Nylin. 2003. Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution 28: 473–484.

Last updated: 27 February 2021.

Satyrinae

Common evening brown Melanitis leda, photographed by Ravi Vaidyanathan.

Belongs within: Nymphalidae.
Contains: Calisto, Oreixenica, Elymniini, Satyrini.

The Satyrinae are a cosmopolitan group of butterflies with the greater diversity in the tropics. Most species are relatively drab in coloration, with browns and greys predominating, though there are some more colourful tropical species. Larvae have bifid tails and mostly feed on monocots.

<==Satyrinae [Satyridae]
    |--+--Elymniini WWN03
    |  `--Satyrini WWN03
    `--+--Manataria maculata WWN03
       `--Melanitis Fabricius 1807 WWN03, B00 [Melanitini]
            |--M. amabilis (Boisduval 1832) B00
            |    |--M. a. amabilis B00
            |    `--M. a. valentina Fruhstorfer 1908 B00
            |--M. ariadne [=Biblis (Melanitis) ariadne] G20
            |--M. constantia (Cramer 1777) B00
            |--M. ismene RD77
            |--M. leda (Linnaeus 1758) B00
            |    |--M. l. leda B12
            |    |--M. l. bankia (Fabricius 1775) B12
            |    |--M. l. levuna PP02
            |    `--M. l. saipanensis Nakamura 1929 TYM08
            `--M. undularis WM66

Satyrinae incertae sedis:
  Haetera GE05 [Haeterini WWN03]
    |--H. diaphana [=Satyrus (Haetera) diaphanus] G20
    `--H. macleannania GE05
  Eritini WWN03
  Ragadiini WWN03
  Taygetis S55
    |--T. andromeda (Cramer 1779) S55
    |--T. kerea Butler 1869 FS54
    `--T. uncinata Weymer 1907 FS54
  Pierella lena (Linnaeus 1767) S55
  Bia [Biini] S55
    `--B. actorion (Linnaeus 1767) S55
  Dodonidia helmsi T27
  Neorinella garciae GE05
  Lethe GE05
    |--L. corbieri GE05
    `--L. diana I92
         |--L. d. diana I92
         `--L. d. mikuraensis Shirôzu 1975 I92
  Pseudoneorina coulleti GE05
  Tisiphone Hübner 1819 B00
    |--T. abeona (Donovan 1805) B00
    |    |--T. a. abeona B00
    |    |--T. a. albifascia Waterhouse 1904 [incl. T. abeona antoni Tindale 1947] B00
    |    |--T. a. aurelia Waterhouse 1915 B00
    |    |--T. a. morrisi Waterhouse 1914 B00
    |    |--T. a. rawnsleyi (Miskin 1876) B00
    |    `--T. a. regalis Waterhouse 1928 B00
    `--T. helena (Olliff 1888) [=T. abeona helena] B00
  Calisto L88
  Eretris L88
  Pararge P01
    |--P. achine P01
    |--P. aegeria KP19
    `--P. megaera P01
  Epinephele P01
    |--E. hyperanthus P01
    |--E. janira F92
    `--E. lycaon P01
  Oreixenica B00
  Nesoxenica Waterhouse & Lyell 1914 B00
    `--N. leprea (Hewitson 1864) B00
         |--N. l. leprea B00
         `--N. l. elia Waterhouse & Lyell 1914 B00
  Argynnina Butler 1867 B00
    |--A. cyrila Waterhouse & Lyell 1914 [=A. hobartia cyrila] B00
    `--A. hobartia (Westwood 1851) B00
         |--A. h. hobartia B00
         |--A. h. montana Couchman & Couchman 1977 B00
         `--A. h. tasmanica (Lyell 1900) B00
  Geitoneura Butler 1867 B00
    |--G. acantha (Donovan 1805) B00
    |--G. klugii (Guérin-Méneville 1830) [incl. G. k. insula Burns 1951, G. k. mulesi (Burns 1948)] B00
    `--G. minyas (Waterhouse & Lyell 1914) [incl. G. minyas mjobergi (Aurivillius 1920)] B00
  Lethites reynesii RJ93
  Cithaerias RD77

*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).

[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.

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

[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.

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

[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.

[L88] Liebherr, J. K. 1988. General patterns in West Indian insects, and graphical biogeographic analysis of some circum-Caribbean Platynus beetles (Carabidae). Systematic Zoology 37 (4): 385–409.

[PP02] Patrick, B., & H. Patrick. 2002. Butterflies of Fiji. Weta 24: 5–12.

[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.

[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.

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

[TYM08] Tennent, W. J., M. Yasuda & K. Morimoto. 2008. Lansania Journal of arachnology and zoology—a rare and obscure Japanese natural history journal. Archives of Natural History 35 (2): 252–280.

[T27] Thomson, G. M. 1927. The pollination of New Zealand flowers by birds and insects. Transactions and Proceedings of the New Zealand Institute 57: 106–125.

[WWN03] Wahlberg, N., E. Weingartner & S. Nylin. 2003. Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution 28: 473–484.

[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: 27 February 2021.

Charaxinae

Charaxes jasius, photographed by Siga.

Belongs within: Nymphalidae.

The Charaxinae are a group of fast-flying butterflies with thick bodies and sharp wings that live in the canopy of trees (Wahlberg et al. 2003). Larvae mostly lack noticeable hairs or spines and usually bear two pairs of prominent tapering horns on the head (Braby 2000b).

<==Charaxinae
    |--Euxanthini WWN03
    |--Prothoini WWN03
    |--Pallini WWN03
    |--Anaea S55 [Anaeini WWN03]
    |    |--A. ates (Druce 1877) S55
    |    |--A. glauce (Felder 1862) S55
    |    |--A. lineata (Salvin 1869) S55
    |    |--A. onophis (Felder 1861) S55
    |    |--A. perenna Godman & Salvin 1884 FS54
    |    |--A. xenocles (Westwood 1850) S55
    |    `--A. xenocrates (Westwood 1850) S55
    |--Preponini WWN03
    |    |--Archeoprepona demophon WWN03
    |    `--Prepona S55
    |         |--P. demophon (Linnaeus 1764) S55
    |         |--P. laertes (Hübner 1806) S55
    |         |--P. meander S55
    |         |    |--P. m. meander S55
    |         |    `--P. m. magos Fruhstorfer 1916 S55
    |         `--P. omphale S55
    |              |--P. o. omphale S55
    |              `--P. o. amesia Fruhstorfer 1904 S55
    `--Charaxes Ochsenheimer 1816 B00b [Charaxini WWN03]
         |--C. affinis B66
         |--C. argynnides B66
         |--C. brutus B01a
         |--C. castor WWN03
         |--C. cithaeron B01b
         |--C. euryalus B66
         |--C. fabius [incl. Papilio euphanes Esper 1801] B66
         |--C. galaxia B66
         |--C. horatius B66
         |--C. jahlusa B66
         |--C. jasius AN-A04
         |--C. latona Butler 1865 B00b
         |    |--C. l. latona B00b
         |    `--C. l. meridionalis B00b
         |--C. manica B01a
         |--C. neanthes B01a
         |--C. pleione (Godart 1819) [=Nymphalis pleione, Paphia pleione] B66
         |--C. pollux B01b
         |--C. pyrrhus B66
         |--C. rosae B01a
         |--C. sempronius (Fabricius 1793) B12
         `--C. varanes B01a

Charaxinae incertae sedis:
  Polyura Billberg 1820 B00b
    |--P. andrewsi (Butler 1900) B00b
    |--P. caphontis PP02
    |--P. eudamippus I92
    |    |--P. e. eudamippus I92
    |    `--P. e. weismanni (Fritze 1894) I92
    |--P. jupiter (Butler 1869) B00b
    |--P. pyrrhus B00b
    `--P. sempronius (Fabricius 1793) [=P. pyrrhus sempronius] B00b
         |--P. s. sempronius B00a
         `--P. s. tiberius B00a
  Agrias beatifica GE05

*Type species of generic name indicated

REFERENCES

[AN-A04] Askew, R. R. & J. L. Nieves-Aldrey. 2004. Further observations on Eupelminae (Hymenoptera, Chalcidoidea, Eupelmidae) in the Iberian Peninsula and Canary Islands, including descriptions of new species. Graellsia 60 (1): 27–39.

[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).

[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.

[B66] Butler, A. G. 1866. Corrections and addenda to certain papers on Lepidoptera published during the years 1865–66; with additional notes on some of the species described. Proceedings of the Zoological Society of London 1866: 451–458.

[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.

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

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

[PP02] Patrick, B., & H. Patrick. 2002. Butterflies of Fiji. Weta 24: 5–12.

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

[WWN03] Wahlberg, N., E. Weingartner & S. Nylin. 2003. Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution 28: 473–484.

Last updated: 27 February 2021.

Nymphalinae

Orange oakleaf Kallima inachus, copyright Suppu.

Belongs within: Nymphalidae.
Contains: Precis, Nymphalini, Melitaeini, Hypolimnas, Junonia.

The Nymphalinae is a cosmopolitan clade of butterflies supported in its current circumscription by molecular analysis (Wahlberg et al. 2003). Members include the oakleaf butterflies of the genus Kallima, found in southern and eastern Asia, which have the undersides of the wings marked to resemble a dead leaf when closed.

<==Nymphalinae
    |--+--Nymphalini WWN03
    |  `--Colobura [Coeini] WWN03
    |       `--C. dirce WWN03
    `--Kallimini WWN03
         |--Kallima WWN03
         |    |--K. inachus I92
         |    |    |--K. i. inachus I92
         |    |    `--K. i. eucerca Fruhstorfer 1898 I92
         |    `--K. paralekta WWN03
         `--+--+--Siproeta stelenes WWN03
            |  `--Melitaeini WWN03
            `--+--Hypolimnas WWN03
               `--+--Junonia WWN03
                  `--Protogoniomorpha WWN03
                       |--P. anacardii WWN03
                       `--P. nebulosa B01a

Nymphalinae incertae sedis:
  Doleschallia Felder & Felder 1860 B00
    `--D. bisaltide (Cramer 1777) B00
         |--D. b. bisaltide PP02
         |--D. b. australis Felder & Felder 1867 B00
         `--D. b. vomana PP02
  Euptoieta hegesia (Cramer 1779) FS54
  Anthanassa FS54
    |--A. frisia FS54
    |    |--A. f. frisia FS54
    |    `--A. f. tulcis (Bates 1864) FS54
    `--A. ptolyca (Bates 1864) FS54
  Microtia elva Bates 1864 FS54
  Pyrameis FS54
    |--P. abyssinica B01b
    |--P. atalanta F92
    |--P. cardui P27
    |--P. gonerilla P27
    `--P. huntera (Fabricius 1775) FS54
  Precis FS54
  Anartia FS54
    |--A. amathea (Linnaeus 1758) S55
    |--A. fatima (Fabricius 1793) FS54
    |    |--A. f. fatima FS54
    |    `--A. f. venusta FS54
    `--A. jatrophae (Linnaeus 1764) S55
  Victorina FS54
    |--V. epaphus Latreille 1811 S55
    |--V. steneles (Linnaeus 1764) S55
    |    |--V. s. steneles S55
    |    |--V. s. bipunctata Fruhstorfer 1907 S55
    |    `--V. s. pallida Fruhstorfer 1907 FS54
    |--V. sulpitia (Cramer 1782) S55
    `--V. superba (Bates 1864) [=Amphirene superba; incl. V. aphrodite Butler 1865] B66b
  Megalura FS54
    |--M. berania Hewitson 1852 S55
    |--M. coresia (Godart 1819) FS54
    |--M. livius Kirby 1852 S55
    |--M. marcella Felder 1861 S55
    `--M. peleus Sulzer 1776 S55
  Smyrna blomfildia FS54
    |--S. b. blomfildia FS54
    `--S. b. datis Fruhstorfer 1908 FS54
  Cyclogramma pandama Doubleday & Hewitson 1850 FS54
  Heterochroa FS54
    |--H. basiloides (Bates 1864) FS54
    `--H. celerio FS54
         |--H. c. celerio FS54
         `--H. c. diademata Fruhstorfer 1915 FS54
  Mynes Boisduval 1832 B00
    `--M. geoffroyi (Guérin-Méneville 1830) B00
         |--M. g. geoffroyi B00
         `--M. g. guerini Wallace 1869 B00
  Symbrenthia J03
  Prodryas persephone GE05
  Lithodryas styx GE05
  Chlorippe GE05
    |--C. laurentia B66a
    |--C. lucasii B66a
    `--C. wilmattae GE05
  Jupitellia charon GE05
  Nymphalites GE05
    |--N. obscurum GE05
    `--N. scudderi GE05
  Apanthesis leuce GE05
  Yoma Doherty 1886 B00
    `--Y. sabina (Cramer 1780) [incl. Y. algina] B00
         |--Y. s. sabina B00
         `--Y. s. parva (Butler 1876) B00
  Brenthis RD77
  Araschnia levana [incl. A. prorsa] RD77

*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).

[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. Corrections and addenda to certain papers on Lepidoptera published during the years 1865-66; with additional notes on some of the species described. Proceedings of the Zoological Society of London 1866: 451–458.

[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.

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

[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.

[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.

[PP02] Patrick, B., & H. Patrick. 2002. Butterflies of Fiji. Weta 24: 5–12.

[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.

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

[WWN03] Wahlberg, N., E. Weingartner & S. Nylin. 2003. Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution 28: 473–484.

Last updated: 27 February 2021.