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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Panorpida
Cladus: Amphiesmenoptera
Ordo: Lepidoptera
Subordo: Glossata
Cladus: Coelolepida
Cladus: Myoglossata
Cladus: Neolepidoptera
Infraordo: Heteroneura
Cladus: Eulepidoptera
Cladus: Ditrysia
Cladus: Apoditrysia
Cladus: Obtectomera
Superfamilia: Papilionoidea

Familia: Nymphalidae
Subfamiliae (12): Apaturinae - Biblidinae - Calinaginae - Charaxinae - Cyrestinae - Danainae - Heliconiinae - Libytheinae - Limenitidinae - Morphinae - Nymphalinae - Satyrinae


Nymphalidae Rafinesque, 1815.
Primary references

Rafinesque, C.S. 1815. Analyse de la nature, ou tableau de l'univers et des corps organisés. Palerme: L'Imprimerie de Jean Barravecchia. 224 pp. BHL Reference page. 

Additional references

Espeland, M., Breinholt, J.W., Willmott, K.R., Warren, A.D., Vila, R., Toussaint, E.F.A., Maunsell, S.C., Aduse-Poku, K., Talavera, G., Eastwood, R., Jarzyna, M.A., Guralnick, R., Lohman, D.J., Pierce, N.E. & Kawahara, A.Y. 2018. A Comprehensive and Dated Phylogenomic Analysis of Butterflies. Current Biology 28(5): 770–778. DOI: 10.1016/j.cub.2018.01.061 Reference page. 
Freitas, A.V.L. & Brown Jr., K.S. 2004. Phylogeny of the Nymphalidae (Lepidoptera). Systematic Biology 53(3): 363–383. DOI: 10.1080/10635150490445670 Open access. Reference page. 
Heikkilä, M., Kaila, L., Mutanen, M., Peña, C. & Wahlberg, N. 2011. Cretaceous origin and repeated tertiary diversification of the redefined butterflies. Proceedings of the Royal Society B 279(1731): 1093–1099. DOI: 10.1098/rspb.2011.1430 Open access. Reference page. 
Wahlberg, N., Leneveu, J., Kodandaramaiah, U., Peña, C., Nylin, S., Freitas, A.V.L. & Brower, A.V.Z. 2009. Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary. Proceedings of the Royal Society B 276(1677): 4295–4302. DOI: 10.1098/rspb.2009.1303 Open access. Reference page. 
Wahlberg, N., Weingartner, E. & Nylin, S. 2003. Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). Molecular phylogenetics and evolution 28(3): 473–484. DOI: 10.1016/S1055-7903(03)00052-6 PDF. Reference page. 


Nymphalidae Systematics Group, Department of Zoology, Stockholm University, Sweden. [1]

Vernacular names
беларуская: Німфаліды
čeština: Babočkovití
dansk: Takvingefamilien
Deutsch: Edelfalter
English: Brush-footed butterfly
español: Ninfálidos
eesti: Koerlibliklased
français: nymphalidés
magyar: Tarkalepkefélék, főlepkék
日本語: タテハチョウ科
한국어: 네발나비과
lietuvių: Pleštekės
македонски: Шарени пеперутки
polski: Rusałkowate
svenska: Praktfjärilar
中文: 蛺蝶科

The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called brush-footed butterflies or four-footed butterflies, because they are known to stand on only four legs while the other two are curled up; in some species, these forelegs have a brush-like set of hairs, which gives this family its other common name. Many species are brightly coloured and include popular species such as the emperors, monarch butterfly, admirals, tortoiseshells, and fritillaries. However, the under wings are, in contrast, often dull and in some species look remarkably like dead leaves, or are much paler, producing a cryptic effect that helps the butterflies blend into their surroundings.


Rafinesque[1] introduced the name Nymphalia as a subfamily name in diurnal Lepidoptera. Rafinesque did not include Nymphalis among the listed genera, but Nymphalis was unequivocally implied in the formation of the name (Code Article The attribution of the Nymphalidae to Rafinesque has now been widely adopted.[2]
For an explanation of technical terms, see External morphology of Lepidoptera.

In the adult butterflies, the first pair of legs is small or reduced,[3] giving the family the other names of four-footed or brush-footed butterflies. The caterpillars are hairy or spiky with projections on the head, and the chrysalids have shiny spots.

The forewings have the submedial vein (vein 1) unbranched and in one subfamily forked near the base; the medial vein has three branches, veins 2, 3, and 4; veins 5 and 6 arise from the points of junction of the discocellulars; the subcostal vein and its continuation beyond the apex of cell, vein 7, has never more than four branches, veins 8–11; 8 and 9 always arise from vein 7, 10, and 11 sometimes from vein 7 but more often free, i.e., given off by the subcostal vein before apex of the cell.[4]

The hindwings have internal (1a) and precostal veins. The cell in both wings is closed or open, often closed in the fore, open in the hindwing. The dorsal margin of the hindwing is channelled to receive the abdomen in many of the forms.[4]
A specimen of Andromeda satyr (Cithaerias andromeda).

The antennae always have two grooves on the underside; the club is variable in shape. Throughout the family, the front pair of legs in the male, and with three exceptions (Libythea, Pseudergolis, and Calinaga) in the female also, is reduced in size and functionally impotent; in some, the atrophy of the forelegs is considerable, e.g., the Danainae and Satyrinae. In many of the forms of these subfamilies, the forelegs are kept pressed against the underside of the thorax, and are in the male often very inconspicuous.[4]
Systematics and phylogeny

The phylogeny of the Nymphalidae is complex. Several taxa are of unclear position, reflecting the fact that some subfamilies were formerly well-recognized as distinct families due to insufficient study.

The five main clades within the family are:[5]

The libytheine clade (basal)

Libytheinae (snout butterflies, earlier treated as the distinct family Libytheidae)

The danaine clade (basal)

Danainae (milkweed butterflies, earlier treated as the distinct family Danaidae)

Host plant families include Apocynaceae, Asclepiadoideae (subfamily of Apocynaceae), and Moraceae.

Ithomiini (about 300 Neotropical species, sometimes considered a subfamily Ithomiinae)

Most species have long wings, and some have transparent wings. Host plants are in the families Apocynaceae, Gesneriaceae, and Solanaceae.

Tellervini (about 6–10 species in Australasia, sometimes considered a subfamily Tellervinae)

Caterpillars resemble those of the Danainae and feed on Apocynaceae.

The satyrine clade

Calinaginae (about six species, restricted to the Himalayas)

Mimics of the Danainae, they are restricted to host plants in the family Moraceae.[6]


Tropical canopy butterflies, the caterpillars often have head spines or projections. Mostly edible species, have some Batesian mimics. Host plants are in the families Annonaceae, Celastraceae, Convolvulaceae, Euphorbiaceae, Fabaceae, Flacourtiaceae, Lauraceae, Myrtaceae, Piperaceae, Poaceae, Rhamnaceae, Rutaceae, Santalaceae, and Sapindaceae.[6]

Morphinae (including Amathusiini, sometimes considered a subfamily Amathusiinae)

Include the spectacular neotropical Morpho, its food plants include the Arecaceae, Bignoniaceae, Fabaceae, Menispermaceae, Poaceae, and Sapindaceae.

Brassolini (owls, neotropical with 70–80 species, mostly crepuscular, sometimes considered a subfamily Brassolinae)

Host plants in the families Arecaceae, Bromeliaceae, Heliconiaceae, Musaceae, and Poaceae.[6]

Satyrinae (satyrs and browns, earlier treated as distinct family Satyridae)

Host plants are in the families Arecaceae, Araceae, Cyperaceae, Heliconiaceae, Poaceae, and Selaginellaceae.

The heliconiine clade (sister group of the nymphaline clade, excludes former tribes Biblidini and Cyrestini, and tribes Pseudergolini and Coeini)

Heliconiinae (earlier treated as distinct family Heliconiidae)

Colourful tropical butterflies, they are noted for Müllerian mimicry. All species use host plants in the family Passifloraceae.

Acraeini (mostly African, but some species in Asia, sometimes considered a family Acraeinae)

Host plants are in the families Asteraceae, Passifloraceae, Sterculiaceae, Tiliaceae, and Urticaceae.


The nymphaline clade (sister group of the heliconiine clade, also includes tribes Coeini and Pseudergolini)

Apaturinae (mostly tropical)

Host plants are in the family Ulmaceae. Caterpillars are smooth with bifid tails and horns on the head.[6]

Biblidinae (formerly in Limenitidinae)
Cyrestinae (formerly in Limenitidinae)
Nymphalinae (a large subfamily that sometimes includes the Limenitidinae and Biblidinae)

Some species migrate. Caterpillars are sometimes covered in spines. Host plants include Acanthaceae, Caprifoliaceae, Convolvulaceae, Euphorbiaceae, Fagaceae, Flacourtiaceae, Lamiaceae, Loranthaceae, Moraceae, Plantaginaceae, Poaceae, Rubiaceae, Rutaceae, Salicaceae, Sapindaceae, Scrophulariaceae, Urticaceae, and Verbenaceae.[6]

Example species from this family

Archdukes, genus Lexias
California tortoiseshell, Nymphalis californica
Comma, Polygonia c-album
Common buckeye, Junonia coenia
Common snout butterfly, Libytheana carinenta
Cracker butterflies, genus Hamadryas
Crimson patch, Chlosyne janais
Edith's checkerspot, Euphydryas editha
Grayling (butterfly), Hipparchia semele
Hackberry emperor, Asterocampa celtis
Lorquin's admiral, Limenitis lorquini
Marsh fritillary, Euphydryas aurinia
Meadow brown, Maniola jurtina
Mourning cloak, Nymphalis antiopa
Monarch butterfly, Danaus plexippus
Blue morpho, Morpho menelaus
Painted lady, Vanessa cardui
Peacock, Aglais io
Plain tiger, Danaus chrysippus
Question mark, Polygonia interrogationis
Red admiral, Vanessa atalanta
Small heath, Coenonympha pamphilus
Small tortoiseshell, Nymphalis urticae
Gatekeeper, Pyronia tithonus
Small pearl-bordered fritillary, Boloria selene
Andromeda satyr, Cithaerias andromeda


The trait for which these butterflies are most known is the use of only four legs; the reason their forelegs have become vestigial is not yet completely clear. Some suggest the forelegs are used to amplify the sense of smell, because some species possess a brush-like set of soft hair called setae, which has led researchers to believe the forelegs are used to improve signaling and communication between the species, while standing in the other four. This ability proves useful in terms of reproduction and the overall health of the species, and it is the leading theory so far.[7]
See also

iconInsects portal iconArthropods portal

List of fritillaries (butterflies)


Rafinesque, C.S. (1815). Analyse de la Nature, ou Tableau de l'Univers et des Corps Organisés. Jean Barravecceia: Palermo. 224 pages, p 127.
Vane-Wright & de Jong, 2003: 167; Pelham, 2008; Wahlberg, 2010
Wolfe, Joanna M.; Oliver, Jeffrey C.; Monteiro, Antónia (2011-01-01). "Evolutionary reduction of the first thoracic limb in butterflies". Journal of Insect Science. 11 (1): 66. doi:10.1673/031.011.6601. ISSN 1536-2442. PMC 3281478. PMID 21867433.
Bingham, C. T. (1905). The Fauna of British India, Including Ceylon and Burma: Butterflies Volume I. London: Taylor and Francis, Ltd.
Niklas Wahlberg, Elisabet Weingartner & Sören Nylin (2003). Gisella Caccone & Giacomo Bernardi (ed.). "Papers presented at the Mammalian Phylogeny symposium during the 2002 Annual Meeting of the Society for Molecular Biology and Evolution, Sorrento, Italy, June 13–16, 2002 (Chapter: Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea))" (PDF). Molecular Phylogenetics and Evolution. 28 (3): 473–484. doi:10.1016/S1055-7903(03)00052-6. PMID 12927132. Archived from the original (PDF) on 2007-10-25.
Philip J. DeVries (2001). "Nymphalidae". In Simon A. Levin (ed.). Encyclopedia of Biodiversity. Academic Press. pp. 559–573. doi:10.1016/B0-12-226865-2/00039-0. ISBN 978-0-12-226865-6.

Gould, S. E. "Butterfly watch: four legs vs. six legs". Scientific American. Retrieved 7 Sep 2013.

Further reading

Glassberg, Jeffrey Butterflies through Binoculars, The West (2001)
Guppy, Crispin S. and Shepard, Jon H. Butterflies of British Columbia (2001)
James, David G. and Nunnallee, David Life Histories of Cascadia Butterflies (2011)
Pelham, Jonathan Catalogue of the Butterflies of the United States and Canada (2008)
Pyle, Robert Michael The Butterflies of Cascadia (2002)

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