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Over 600 genera
About 5,700 species
Nymphalidae are the largest family of butterflies with more than
6,000 species distributed throughout most of the world, belonging to
the superfamily Papilionoidea. These 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.
2.1 Systematics and phylogeny
2.2 Example species from this family
4 See also
6 Further reading
7 External links
Rafinesque 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 188.8.131.52). The attribution of the
Nymphalidae to Rafinesque has now been widely adopted.
For terms see External morphology of Lepidoptera.
In the adult butterflies, the first pair of legs is small or
reduced, 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 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.
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.,
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.
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
The five main clades within the family are:
The libytheine clade (basal)
Libytheinae (snout butterflies, earlier treated as the distinct family
The danaine clade (basal)
Danainae (milkweed butterflies, earlier treated as the distinct family
Host plant families include Apocynaceae,
Asclepiadoideae (subfamily of
Apocynaceae), and Moraceae.
Ithomiini (about 300 neotropical species, sometimes considered a
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
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
Morphinae (including Amathusiini, sometimes considered a subfamily
Include the spectacular neotropical Morpho, its food plants include
the Arecaceae, Bignoniaceae, Fabaceae, Menispermaceae, Poaceae, and
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.
Satyrinae (satyrs and browns, earlier treated as distinct family
Host plants are in the families Arecaceae, Araceae, Cyperaceae,
Heliconiaceae, Poaceae, and Selaginellaceae.
The heliconiine clade (sister group of the nymphaline clade, excludes
Biblidini and Cyrestini, and tribes Pseudergolini and
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
Coeini and Pseudergolini)
Apaturinae (mostly tropical)
Host plants are in the family Ulmaceae. Caterpillars are smooth with
bifid tails and horns on the head.
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.
Example species from this family
Archdukes, genus Lexias
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
Monarch butterfly, Danaus plexippus
Painted lady, Vanessa cardui
Plain tiger, Danaus chrysippus
Question mark, Polygonia interrogationis
Red admiral, Vanessa atalanta
Small heath, Coenonympha pamphilus
Gatekeeper, Pyronia tithonus
Small pearl-bordered fritillary, "Boloria Selene"
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.
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 .
^ a b c Bingham, C.T. (1905). The Fauna of British India, Including
Ceylon and Burma Butterflies. 1 (1st ed.). London: Taylor and Francis,
^ Niklas Wahlberg, Elisabet Weingartner & Sören Nylin (2003).
Gisella Caccone & Giacomo Bernardi, ed. "Papers presented at the
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
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.
^ a b c d e Philip J. DeVries (2001). "Nymphalidae". In Simon A.
Levin. Encyclopedia of Biodiversity. Academic Press.
pp. 559–573. doi:10.1016/B0-12-226865-2/00039-0.
^ Gould, S.E. "
Butterfly watch: four legs vs. six legs". Scientific
American. Scientific American. Retrieved 7 Sep 2013.
Glassberg, Jeffrey Butterflies through Binoculars, The West (2001)
Guppy, Crispin S. and Shepard, Jon H. Butterflies of British Columbia
James, David G. and Nunnallee, David Life Histories of Cascadia
Pelham, Jonathan Catalogue of the Butterflies of the United States and
Pyle, Robert Michael The Butterflies of Cascadia (2002)
Wikimedia Commons has media related to Nymphalidae.
Wikispecies has information related to Nymphalidae
Biodiversity Information Facility (2003): Family Nymphalidae
Nymphalidae at Lepidoptera.pro
Peter Chew: Danaids and Browns - Family Nymphalidae, Brisbane
butterflies web site (2005).
Tree of Life Web Project: Nymphalidae
Nymphalidae from all over the world
Butterflies and Moths of North America
Butterflies of America
Micropterigidae (mandibulate archaic moths)
Agathiphagidae (kauri moths)
Acanthopteroctetidae (archaic sun moths)
Neopseustidae (archaic bell moths)
Hepialidae (swift moths, ghost moths)
Neotheoridae (Amazonian primitive ghost moths)
Palaeosetidae (miniature ghost moths)
Prototheoridae (African primitive ghost moths)
Mnesarchaeidae (New Zealand primitive moths)
Adelidae (fairy longhorn moths)
Prodoxidae (yucca moths)
Andesianidae (Andean endemic moths)
Nepticulidae (pigmy, or midget moths)
Opostegidae (white eyecap moths)
Palaephatidae (Gondwanaland moths)
Tischeriidae (trumpet leaf miner moths)
Acrolophidae (burrowing webworm moths)
Eriocottidae (Old World spiny-winged moths)
Psychidae (bagworm moths)
Tineidae (fungus moths)
Bucculatricidae (ribbed cocoon makers)
Douglasiidae (Douglas moths)
Acrolepiidae (false diamondback moths)
Glyphipterigidae (sedge moths)
Yponomeutidae (ermine moths)
Coleophoridae (case-bearers, case moths)
Cosmopterigidae (cosmet moths)
Elachistidae (grass-miner moths)
Gelechiidae (twirler moths)
Lecithoceridae (long-horned moths)
Momphidae (mompha moths)
Oecophoridae (concealer moths)
Scythrididae (flower moths)
Xyloryctidae (timber moths)
Zygaenidae (burnet, forester, or smoky moths)
Megalopygidae (flannel moths)
Epipyropidae (planthopper parasite moths)
Dalceridae (slug caterpillars)
Limacodidae (slug, or cup moths)
Cossidae (carpenter millers, or goat moths)
Dudgeoneidae (dudgeon carpenter moths)
Brachodidae (little bear moths)
Castniidae (castniid moths: giant butterfly-moths, sun moths)
Sesiidae (clearwing moths)
Choreutidae (metalmark moths)
Tortricidae (tortrix moths)
Urodidae (false burnet moths)
Schreckensteiniidae (bristle-legged moths)
Epermeniidae (fringe-tufted moths)
Alucitidae (many-plumed moths)
Tineodidae (false plume moths)
Pterophoridae (plume moths)
Copromorphidae (tropical fruitworm moths)
Carposinidae (fruitworm moths)
Hyblaeidae (teak moths)
Pyralidae (snout moths)
Crambidae (grass moth)
Thyrididae (picture-winged leaf moths)
Mimallonidae (sack bearer moths)
Lasiocampidae (eggars, snout moths, or lappet moths)
Anthelidae (Australian lappet moth)
Bombycidae (silk moths)
Brahmaeidae (Brahmin moths)
Carthaeidae (Dryandra moth)
Endromidae (Kentish glory and relatives)
Sphingidae (hawk moths, sphinx moths and hornworms)
Erebidae (underwing, tiger, tussock, litter, snout, owlet moths)
Noctuidae (daggers, sallows, owlet moths, quakers, cutworms, darts)
Nolidae (tuft moths)
Notodontidae (prominents, kittens)
Epicopeiidae (oriental swallowtail moths)
Geometridae (geometer moths)
Cimeliidae (gold moths)
Callidulidae (Old World butterfly-moths)
Hedylidae (American moth-butterflies)
Lycaenidae (gossamer-winged butterflies: blues, coppers and relatives)
Nymphalidae (brush-footed, or four-footed butterflies)
Papilionidae (swallowtail butterflies)
Pieridae (whites, yellows, orangetips, sulphurs)
Monotrysia is not a clade.
Taxonomy of the Lepidoptera
Lists by region
Fauna Europaea: 7202