The Nymphalidae are the largest

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...

, 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 An emperor (from la, imperator, via fro, empereor) is a monarch, and usually the sovereign ruler of an empire or another type of imperial realm. Empress, the female equivalent, may indicate an emperor's wife ( empress consort), mother (empr ...

monarch butterfly The monarch butterfly or simply monarch (''Danaus plexippus'') is a milkweed butterfly (subfamily Danainae) in the family Nymphalidae. Other common names, depending on region, include milkweed, common tiger, wanderer, and black-veined brown. ...

admirals Admiral is one of the highest ranks in some navies. In the Commonwealth nations and the United States, a "full" admiral is equivalent to a "full" general in the army or the air force, and is above vice admiral and below admiral of the fleet, ...

, 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 Cryptic may refer to: In science: * Cryptic species complex, a group of species that are very difficult to distinguish from one another * Crypsis, the ability of animals to blend in to avoid observation * Cryptic era, earliest period of the Earth ...

effect that helps the butterflies blend into their surroundings.

Nomenclature

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 11.7.1.1). The attribution of the Nymphalidae to Rafinesque has now been widely adopted.

Classification

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

caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sy ...

s 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. 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. Cithaerias Esmeralda MHNT

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 ''Pseudergolis'' is a butterfly genus from the family Nymphalidae found in Southeast Asia. Some authorities place it in the subfamily Cyrestinae. Species Listed alphabetically.Calinaga'') in the female also, is reduced in size and functionally impotent; in some, the

atrophy Atrophy is the partial or complete wasting away of a part of the body. Causes of atrophy include mutations (which can destroy the gene to build up the organ), poor nourishment, poor circulation, loss of hormonal support, loss of nerve supply ...

of the forelegs is considerable, e.g., the Danainae and

Satyrinae The Satyrinae, the satyrines or satyrids, commonly known as the browns, are a subfamily of the Nymphalidae (brush-footed butterflies). They were formerly considered a distinct family, Satyridae. This group contains nearly half of the known diver ...

. In many of the forms of these subfamilies, the forelegs are kept pressed against the underside of the

thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the c ...

, and are in the male often very inconspicuous.

Systematics and phylogeny

The

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological s ...

of the Nymphalidae is complex. Several

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

are of unclear position, reflecting the fact that some subfamilies were formerly well-recognized as distinct families due to insufficient study. The five main

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...

s within the family are: 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 Apocynaceae (from '' Apocynum'', Greek for "dog-away") is a family of flowering plants that includes trees, shrubs, herbs, stem succulents, and vines, commonly known as the dogbane family, because some taxa were used as dog poison Members ...

, Asclepiadoideae (subfamily of

), and Moraceae. :* Ithomiini (about 300

Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...

species, sometimes considered a subfamily Ithomiinae) :: Most species have long wings, and some have transparent wings. Host plants are in the families

Gesneriaceae Gesneriaceae, the gesneriad family, is a family of flowering plants consisting of about 152 genera and ca. 3,540 species in the tropics and subtropics of the Old World (almost all Didymocarpoideae) and the New World (most Gesnerioideae), ...

, and

Solanaceae The Solanaceae , or nightshades, are a family of flowering plants that ranges from annual and perennial herbs to vines, lianas, epiphytes, shrubs, and trees, and includes a number of agricultural crops, medicinal plants, spices, weeds, and or ...

. :* Tellervini (about 6–10 species in

Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologic ...

, sometimes considered a subfamily

Tellervinae Tellervini is a tribe of danaid butterflies with only the one genus ''Tellervo'', with six widely distributed species found in the Australasian realm and the Indomalayan realm (also called the ''Oriental region''). The taxon is apparently monoph ...

) :: Caterpillars resemble those of the Danainae and feed on Apocynaceae. The satyrine clade *

Calinaginae ''Calinaga'' is the sole genus of the monotypic nymphalid butterfly subfamily Calinaginae. Its species occur in South Asia and Southeast Asia. The evolutionary history of ''Calinaga'' remains a mystery until today. Recent research indicated that ...

(about six species, restricted to the

Himalayas The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 10 ...

) : Mimics of the Danainae, they are restricted to host plants in the family Moraceae. *

Charaxinae __NOTOC__ The Charaxinae, the leafwings, are a nymphalid subfamily of butterflies that includes about 400 species, inhabiting mainly the tropics, although some species extend into temperate regions in North America, Europe, China, and southern ...

: 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 The Annonaceae are a family of flowering plants consisting of trees, shrubs, or rarely lianas commonly known as the custard apple family or soursop family. With 108 accepted genera and about 2400 known species, it is the largest family in the Ma ...

, Celastraceae,

Convolvulaceae Convolvulaceae (), commonly called the bindweeds or morning glories, is a family of about 60 genera and more than 1,650 species. These species are primarily herbaceous vines, but also include trees, shrubs and herbs. The tubers of several spe ...

Euphorbiaceae Euphorbiaceae, the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of a genus in the family. Most spurges, such as '' Euphorbia paralias'', are herbs, but some, ...

Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...

Flacourtiaceae The Flacourtiaceae is a defunct family of flowering plants whose former members have been scattered to various families, mostly to the Achariaceae and Salicaceae. It was so vaguely defined that hardly anything seemed out of place there and it beca ...

Lauraceae Lauraceae, or the laurels, is a plant family that includes the true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genera worldwide (Christenhusz & Byng 2016 ). They are dicotyledons, and occur m ...

Myrtaceae Myrtaceae, the myrtle family, is a family of dicotyledonous plants placed within the order Myrtales. Myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, and eucalyptus are some notable members of this group. All sp ...

Piperaceae The Piperaceae (), also known as the pepper family, are a large family of flowering plants. The group contains roughly 3,600 currently accepted species in 5 genera. The vast majority of species can be found within the two main genera: '' Piper' ...

Poaceae Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...

Rhamnaceae The Rhamnaceae are a large family of flowering plants, mostly trees, shrubs, and some vines, commonly called the buckthorn family. Rhamnaceae is included in the order Rosales. The family contains about 55 genera and 950 species. The Rhamnaceae ...

, Rutaceae, Santalaceae, and

Sapindaceae The Sapindaceae are a family of flowering plants in the order Sapindales known as the soapberry family. It contains 138 genera and 1858 accepted species. Examples include horse chestnut, maples, ackee and lychee. The Sapindaceae occur in temp ...

. * Morphinae (including

Amathusiini Amathusiini is a tribe of the nymphalid butterfly subfamily Morphinae. They are large butterflies. They are sometimes treated as a distinct subfamily Amathusiinae or family Amathusiidae. Genera and selected species * ''Aemona'' ** '' Aemona ama ...

, sometimes considered a subfamily Amathusiinae) : Include the spectacular neotropical '' Morpho'', its food plants include the

Arecaceae The Arecaceae is a family of perennial flowering plants in the monocot order Arecales. Their growth form can be climbers, shrubs, tree-like and stemless plants, all commonly known as palms. Those having a tree-like form are called palm trees ...

, Bignoniaceae, Fabaceae, Menispermaceae, Poaceae, and Sapindaceae. :*

Brassolini Brassolini is a tribe usually placed in the brush-footed butterfly subfamily Morphinae, which is often included in the Satyrinae as a tribe Morphini. If this is accepted, the Brassolini become the sister tribe of the Morphini among the Satyrinae ...

(owls, neotropical with 70–80 species, mostly

crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of dayli ...

, sometimes considered a subfamily Brassolinae) :: Host plants in the families Arecaceae,

Bromeliaceae The Bromeliaceae (the bromeliads) are a family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, ...

, Heliconiaceae,

Musaceae Musaceae is a family of flowering plants composed of three genera with about 91 known species, placed in the order Zingiberales. The family is native to the tropics of Africa and Asia. The plants have a large herbaceous growth habit with leaves ...

, and Poaceae. *

(satyrs and browns, earlier treated as distinct family Satyridae) : Host plants are in the families Arecaceae,

Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). A ...

Cyperaceae The Cyperaceae are a family of graminoid (grass-like), monocotyledonous flowering plants known as sedges. The family is large, with some 5,500 known species described in about 90 genera, the largest being the "true sedges" genus '' Carex'' ...

, Heliconiaceae, Poaceae, and Selaginellaceae. The heliconiine clade (

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

of the nymphaline clade, excludes former

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...

s 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 The Passifloraceae are a family of flowering plants, containing about 750 species classified in around 27 genera. They include trees, shrubs, lianas, and climbing plants, and are mostly found in tropical regions. The family takes its name fr ...

. :* Acraeini (mostly African, but some species in Asia, sometimes considered a family

Acraeinae The Acraeini are a tribe of butterflies of the subfamily Heliconiinae in the family Nymphalidae. Genera The recognized genera are: Tree of Life * ''Abananote'' Potts, 1943 * ''Acraea'' Fabricius, 1807 – acraeas * ''Actinote'' Hübner, 8 ...

) :: Host plants are in the families

Asteraceae The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae ...

, Passifloraceae, Sterculiaceae,

Tiliaceae Tiliaceae () is a family of flowering plants. It is not a part of the APG, APG II and APG III classifications, being sunk in Malvaceae mostly as the subfamilies Tilioideae, Brownlowioideae and Grewioideae, but has an extensive historical reco ...

, and

Urticaceae The Urticaceae are a family, the nettle family, of flowering plants. The family name comes from the genus ''Urtica''. The Urticaceae include a number of well-known and useful plants, including nettles in the genus ''Urtica'', ramie (''Boehmeri ...

. * Limenitidinae The nymphaline clade (

of the heliconiine clade, also includes tribes

Coeini Coeini is a tribe of brush-footed butterflies.Tribe 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 Cyrestinae is the name of a small subfamily of nymphalid brush-footed butterflies. It is considered to include only three genera – ''Marpesia'', ''Chersonesia'', and ''Cyrestis'' – distributed in the tropics.N. Wahlberg , J. Leneveu , U. Ko ...

(formerly in Limenitidinae) *

Nymphalinae The Nymphalinae are a subfamily of brush-footed butterflies (family Nymphalidae). Sometimes, the subfamilies Limenitidinae, and Biblidinae are included here as subordinate tribe(s), while the tribe Melitaeini is occasionally regarded as a distin ...

(a large subfamily that sometimes includes the Limenitidinae and Biblidinae) : Some species migrate. Caterpillars are sometimes covered in spines. Host plants include Acanthaceae,

Caprifoliaceae The Caprifoliaceae or honeysuckle family is a clade of dicotyledonous flowering plants consisting of about 860 species, in 33, to 42 genera, with a nearly cosmopolitan distribution. Centres of diversity are found in eastern North America and ...

, Fagaceae,

, Lamiaceae,

Loranthaceae Loranthaceae, commonly known as the showy mistletoes, is a family of flowering plants. It consists of about 75 genera and 1,000 species of woody plants, many of them hemiparasites. The three terrestrial species are ''Nuytsia floribunda'' (the W ...

, Moraceae, Plantaginaceae, Poaceae, Rubiaceae, Rutaceae, Salicaceae,

Scrophulariaceae The Scrophulariaceae are a family of flowering plants, commonly known as the figwort family. The plants are annual and perennial herbs, as well as shrubs. Flowers have bilateral (zygomorphic) or rarely radial (actinomorphic) symmetry. The Scr ...

, Urticaceae, and

Verbenaceae The Verbenaceae ( ), the verbena family or vervain family, is a family of mainly tropical flowering plants. It contains trees, shrubs, and herbs notable for heads, spikes, or clusters of small flowers, many of which have an aromatic smell. The ...

Example species from this family

* Archdukes, genus ''Lexias'' * California tortoiseshell, ''Nymphalis californica'' *

Comma The comma is a punctuation mark that appears in several variants in different languages. It has the same shape as an apostrophe or single closing quotation mark () in many typefaces, but it differs from them in being placed on the baseline ...

, ''Polygonia c-album'' * Common buckeye, ''Junonia coenia'' * Common snout butterfly, ''Libytheana carinenta'' * Cracker butterflies, genus '' Hamadryas'' *

Crimson patch ''Chlosyne janais'', the crimson patch or janais patch, is a common New World butterfly found from Colombia north through Central America and Mexico to southern Texas, with occasional sightings in southeastern Mexico and northern Texas. The dors ...

, ''Chlosyne janais'' * Edith's checkerspot, ''Euphydryas editha'' *

Grayling (butterfly) The grayling or rock grayling (''Hipparchia semele'') is a species in the brush-footed butterfly family Nymphalidae. Although found all over Europe, the grayling mostly inhabits coastal areas, with inland populations declining significantly in r ...

, ''Hipparchia semele'' * Hackberry emperor, ''Asterocampa celtis'' *

Lorquin's admiral Lorquin's admiral (''Limenitis lorquini'') is a butterfly in the Nymphalinae subfamily. The butterfly is named after Pierre Joseph Michel Lorquin, a French naturalist who came to California from France during the Gold Rush, and made important d ...

, ''Limenitis lorquini'' *

Marsh fritillary The marsh fritillary (''Euphydryas aurinia'') is a butterfly of the family Nymphalidae. Commonly distributed in the Palearctic region, the marsh fritillary's common name derives from one of its several habitats, marshland. The prolonged larval s ...

, ''Euphydryas aurinia'' * Meadow brown, ''Maniola jurtina'' * Mourning cloak, ''Nymphalis antiopa'' *

Monarch butterfly The monarch butterfly or simply monarch (''Danaus plexippus'') is a milkweed butterfly (subfamily Danainae) in the family Nymphalidae. Other common names, depending on region, include milkweed, common tiger, wanderer, and black-veined brown. ...

, ''Danaus plexippus'' * Blue morpho, ''Morpho menelaus'' * Painted lady, ''Vanessa cardui'' *

Peacock Peafowl is a common name for three bird species in the genera '' Pavo'' and '' Afropavo'' within the tribe Pavonini of the family Phasianidae, the pheasants and their allies. Male peafowl are referred to as peacocks, and female peafowl are r ...

, ''Aglais io'' * Plain tiger, ''Danaus chrysippus'' *

Question mark The question mark (also known as interrogation point, query, or eroteme in journalism) is a punctuation mark that indicates an interrogative clause or phrase in many languages. History In the fifth century, Syriac Bible manuscripts used ...

, ''Polygonia interrogationis'' * Red admiral, ''Vanessa atalanta'' * Small heath, ''Coenonympha pamphilus'' * Small tortoiseshell, ''Nymphalis urticae'' *

Gatekeeper A gatekeeper is a person who controls access to something, for example via a city gate or bouncer, or more abstractly, controls who is granted access to a category or status. Gatekeepers assess who is "in or out", in the classic words of manage ...

, ''Pyronia tithonus'' *

Small pearl-bordered fritillary ''Boloria selene'', known in Europe as the small pearl-bordered fritillary and in North America as the silver-bordered fritillary, is a species of butterfly of the family Nymphalidae. It is found across Europe, Asia and North America, and feeds ...

, ''Boloria selene'' *

Andromeda satyr ''Cithaerias andromeda'', the Andromeda satyr, is a species of butterfly of the family Nymphalidae. It is found from Suriname, French Guiana, Venezuela, Peru, Bolivia and Brazil. Differentiation between subspecies and their corresponding s ...

, ''Cithaerias andromeda''

Morphology

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

seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...

e, 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.

References

External links

* Canadian Biodiversity Information Facility (2003)
Family Nymphalidae

Nymphalidae
"Family Nymphalidae". ''Insecta.pro''. * Peter Chew

Brisbane butterflies web site (2005). * Tree of Life Web Project
Nymphalidae

{{Authority control Butterfly families Taxa named by Constantine Samuel Rafinesque Papilionoidea