Melinaea maenius, photographed by Rich Hoyer.

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

The Nymphalidae, brush-footed butterflies, are a diverse group of butterflies characterised by the presence of three carinae on the antennae. The first pair of legs is reduced in most nymphalids except females of the basal subfamily Libytheinae, so that only four functional walking legs are visible.

Characters (from Nielsen & Common 1991): Small to large; most antennal segments with two ventral grooves; maxillary palps one-segmented; labial palps ascending; epiphysis absent, male forelegs lack pretarsus, with fewer than five tarsomeres, lack tarsal spines, and covered with long scales; female forelegs usually reduced in size and not used for walking; spurs 0-2-2 or 0-0-0; fore wing with all branches of R present, 1A+2A simple or forming a basal fork; hind wing with humeral vein usually present, Sc+R1 diverging from Rs near base, two anal veins. Egg taller than diameter, with vertical and horizontal ribs, or nearly spherical and sometimes nearly smooth. Larva with long paired filaments, branching spines, or fine secondary setae and bifid anal segment or horned head; crochets multiordinal. Pupa suspended by cremaster or, rarely, loose on ground beneath debris.

    |--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, Ithomiinae] 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
            |         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
  Lethites reynesii GE05
  Neorinopis sepulta GE05
  Eugonia 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

*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.

[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.

[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.

Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera. In: CSIRO. The Insects of Australia 2nd ed. vol. 2 pp. 817–915. Melbourne University Press.

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

[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: 20 August 2017.

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