Nymphalidae is the largest family of butterflies with about 6,000 species distributed throughout most of the world. 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. Many species are brightly colored and include popular species such as the emperors, admirals, tortoiseshells, and fritillaries. However, the underwings 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 butterfly disappear into its surroundings.
Rafinesque introduced the name Nymphalia as a subfamily name in diurnal Lepidoptera. Rafinesque did not include Nymphalis among the listed genera, but Nymphalis is unequivocally implied in the formation of the name (Code Article 126.96.36.199). The attribution of the Nymphalidae to Rafinesque has now been widely adopted (e.g., Vane-Wright & de Jong, 2003: 167; Pelham, 2008; Wahlberg, 2010).
In adult butterflies the first pair of legs are 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 forewing has the submedial vein (vein 1) unbranched and in one subfamily forked near base; medial vein with three branches, veins 2, 3, and 4; veins 5 and 6 arising from the points of junction of the discocellulars; subcostal vein and its continuation beyond apex of cell, vein 7, with never more than four branches, veins 8–11; 8 and 9 always arising from vein 7, 10, and also 11 sometimes from vein 7 but more often free, i.e., given off by the subcostal vein before apex of cell.
The hindwing has internal (1a) and precostal veins. The cell in both wings closed or open, often closed in the fore, open in the hindwing. Dorsal margin of hind wing channelled to receive the abdomen in many of the forms.
Antennae always with two grooves on the underside; club 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 fore legs 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 study.
The libytheine clade (basal)
The danaine clade (basal)
- Host plant families include Apocynaceae, Asclepiadoideae (subfamily of Apocynaceae), and Moraceae.
- Most species with long wings, some having transparent wings. Host plants in the families Apocynaceae, Gesneriaceae, and Solanaceae.
The satyrine clade
- Tropical canopy butterflies. Caterpillars often with head spines or projections. Mostly edible species with some Batesian mimics. Host plants in the families Annonaceae, Celastraceae, Convolvulaceae, Euphorbiaceae, Fabaceae, Flacourtiaceae, Lauraceae, Myrtaceae, Piperaceae, Poaceae, Rhamnaceae, Rutaceae, Santalaceae, and Sapindaceae.
- Include the spectacular neotropical Morphos. 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, Araceae, Cyperaceae, Heliconiaceae, Poaceae, and Selaginellaceae.
- Colourful tropical butterflies noted for Müllerian mimicry. All species use host plants in the family Passifloraceae.
- Apaturinae (Mostly tropical)
- Biblidinae (formerly in Limenitidinae)
- Cyrestinae (formerly in Limenitidinae)
- Nymphalinae (A large subfamily that sometimes includes the Limenitidinae and Biblidinae.)
- Some species migratory. Caterpillars 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
- 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
- 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, Inachis io
- Question Mark, Polygonia interrogationis
- Red Admiral, Vanessa atalanta
- Small Heath, Coenonympha pamphilus
- Small Tortoiseshell, Nymphalis urticae
- Rafinesque, C.S. (1815) Analyse de la Nature, ou Tableau de l'Univers et des Corps Organisés. Jean Barravecceia: Palermo. 224 pages, p 127.
- Charles Thomas Bingham (1905). Butterflies, Volume 1. The Fauna of British India, Including Ceylon and Burma. London: Taylor and Francis. http://www.archive.org/details/TheFunaOfBritishIndiaButterfliesVolI.
- Niklas Wahlberg, Elisabet Weingartner & Sören Nylin (2003). Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). In Gisella Caccone & Giacomo Bernardi. "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" (PDF). Molecular Phylogenetics and Evolution 28 (3): 473–484. doi:10.1016/S1055-7903(03)00052-6. PMID 12927132. http://nymphalidae.utu.fi/Wahlbergetal2003b.pdf.
- 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. ISBN 978-0-12-226865-6.