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.

    |--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


[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.

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