Editing Nymphalidae

{{Short description|Largest butterfly family}}
{{Automatic taxobox
| image = NovaraExpZoologischeTheilLepidopteraAtlasTaf53.jpg
| image_caption = 
| taxon = Nymphalidae
| authority = [[Constantine Samuel Rafinesque|Rafinesque]], 1815
| diversity = Over 600 genera<br/>About 5,700 species
| subdivision_ranks = Subfamilies
| subdivision = 
* [[Apaturinae]]
* [[Biblidinae]]
* [[Calinaginae]]
* [[Charaxinae]]
* [[Cyrestinae]]
* [[Danainae]]
* [[Heliconiinae]]
* [[Libytheinae]]
* [[Limenitidinae]]
* [[Morphinae]]
* [[Nymphalinae]]
* [[Satyrinae]]
and see [[#Systematics and phylogeny|text]]
}}
[[File:Peacock butterfly (Aglais io) 2.jpg|thumb|Peacock (''[[Aglais io]])'']]
[[File:Crimson-spotted forester (Euphaedra themis) male.jpg|thumb|Crimson-spotted forester (''[[Euphaedra themis]]'')]]
[[File:Monarch butterflies (Danaus plexippus plexippus) Piedra Herrada 2.jpg|thumb|Monarch butterflies (''[[Danaus plexippus]]'')]]
[[File:Dark blue tiger (Tirumala septentrionis septentrionis) male underside Phi Phi.jpg|thumb|Dark blue tiger (''[[Tirumala septentrionis]]'')]]
[[File:Small pearl-bordered fritillary (Boloria selene).jpg|thumb|Small pearl-bordered fritillary (''[[Boloria selene]]'')]]
[[File:Crescent butterfly.webm|thumb|thumbtime=8|[[Phyciodes|Crescent]] butterfly [[nectar]]ing on [[Verbesina alternifolia|yellow ironweed]]]]

The '''Nymphalidae''' are the largest [[family (biology)|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 [[Apaturinae|emperors]], [[monarch butterfly]], [[Nymphalini|admirals]], [[Aglais|tortoiseshells]], and [[Heliconiini|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 [[crypsis|cryptic]] effect that helps the butterflies blend into their surroundings.

== Nomenclature ==

Rafinesque<ref>Rafinesque, C.S. (1815). ''Analyse de la Nature, ou Tableau de l'Univers et des Corps Organisés.'' Jean Barravecceia: Palermo. 224 pages, p 127.</ref> 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.<ref>Vane-Wright & de Jong, 2003: 167; Pelham, 2008; Wahlberg, 2010</ref>
<!-- not a sentence
and contradictory = Nymphalidae Swainson, 1827, Phil. Mag. (n. Ser.), 1: 187] is an invalid name and a junior homonym of Nymphalidae Rafinesque, 1815. -->

== Morphology ==

{{For|an explanation of technical terms|External morphology of Lepidoptera}}

In the adult butterflies, the first pair of legs is small or reduced,<ref>{{Cite journal|last1=Wolfe|first1=Joanna M.|last2=Oliver|first2=Jeffrey C.|last3=Monteiro|first3=Antónia|date=2011-01-01 |title=Evolutionary reduction of the first thoracic limb in butterflies|url= |journal=Journal of Insect Science|language=en|volume=11|issue=1|pages=66|doi=10.1673/031.011.6601|issn=1536-2442|pmc=3281478|pmid=21867433}}</ref> giving the family the other names of four-footed or brush-footed butterflies. The [[caterpillar]]s are hairy or spiky with projections on the head, and the [[Pupa|chrysalids]] have shiny spots.

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]]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.<ref>{{cite web|last1=Gould|first1=S. E |title=Butterfly watch: four legs vs. six legs|url=http://blogs.scientificamerican.com/lab-rat/butterfly-watch-four-legs-vs-six-legs/|website=Scientific American |access-date=7 Sep 2013}}</ref>

The [[forewing]]s 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.<ref name="ctb">{{cite book |last1=Bingham |first1=C. T. |author-link=Charles Thomas Bingham |title=[[The Fauna of British India, Including Ceylon and Burma]]: Butterflies Volume I |publisher= [[Taylor & Francis|Taylor and Francis, Ltd.]] |location=London |year=1905}}</ref>

The [[hindwing]]s 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.<ref name="ctb"/>

The [[Antenna (biology)|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 (insect anatomy)|thorax]], and are in the male often very inconspicuous.<ref name="ctb"/>

=== Systematics and phylogeny ===

The [[phylogeny]] of the Nymphalidae is complex. Several [[Taxon|taxa]] are of unclear position, reflecting the fact that some [[Family (biology)|subfamilies]] were formerly well-recognized as distinct families due to insufficient study.

The five main [[clade]]s within the family are:<ref name="wahlberg">{{cite journal |author=Niklas Wahlberg, Elisabet Weingartner & Sören Nylin |year=2003 |journal=[[Molecular Phylogenetics and Evolution]] |volume=28 |issue=3 |pages=473–484 |doi=10.1016/S1055-7903(03)00052-6 |pmid=12927132 |url=http://nymphalidae.utu.fi/Wahlbergetal2003b.pdf |title=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)) |editor=Gisella Caccone & Giacomo Bernardi |url-status=dead |archive-url=https://web.archive.org/web/20071025220449/http://nymphalidae.utu.fi/Wahlbergetal2003b.pdf |archive-date=2007-10-25}}</ref>

The '''libytheine clade''' ([[basal (biology)|basal]])

* [[Libytheinae]] ([[Snout butterfly|snout butterflies]], earlier treated as the distinct family Libytheidae)

The '''danaine clade''' ([[basal (biology)|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 realm|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]].<ref name="eob">{{cite book |author=Philip J. DeVries |author-link=Philip James DeVries |year=2001 |chapter=Nymphalidae |pages=[https://archive.org/details/encyclopediabiod00levi/page/n567 559]–573 |editor=Simon A. Levin |editor-link=Simon A. Levin |title=Encyclopedia of Biodiversity |url=https://archive.org/details/encyclopediabiod00levi|url-access=limited|publisher=[[Academic Press]] |isbn=978-0-12-226865-6 |doi=10.1016/B0-12-226865-2/00039-0}}</ref>
* [[Charaxinae]]
: Tropical [[Canopy (biology)|canopy]] butterflies, the caterpillars often have head spines or projections. Mostly edible species, have some [[Batesian mimicry|Batesian mimics]]. Host plants are in the families [[Annonaceae]], [[Celastraceae]], [[Convolvulaceae]], [[Euphorbiaceae]], [[Fabaceae]], [[Flacourtiaceae]], [[Lauraceae]], [[Myrtaceae]], [[Piperaceae]], [[Poaceae]], [[Rhamnaceae]], [[Rutaceae]], [[Santalaceae]], and [[Sapindaceae]].<ref name="eob"/>
* [[Morphinae]] (including [[Amathusiini]], sometimes considered a subfamily [[Amathusiinae]])
: Include the spectacular neotropical ''[[Morpho (genus)|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.<ref name="eob"/>
* [[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 [[tribe (biology)|tribe]]s [[Biblidini]] and [[Cyrestini]], and tribes Pseudergolini and Coeini)

* [[Heliconiinae]] (earlier treated as distinct family Heliconiidae)
: Colourful tropical butterflies, they are noted for [[mimicry|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]].
* [[Limenitidinae]]

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.<ref name="eob"/>
* [[Biblidinae]] (formerly in [[Limenitidinae]])
* [[Cyrestinae]] (formerly in Limenitidinae)
* [[Nymphalinae]] (a large subfamily that sometimes includes the Limenitidinae and [[Biblidinae]])
: Some species [[Lepidoptera migration|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]].<ref name="eob"/>

=== Example species from this family ===
{{div col|colwidth=23em}}
* ''[[Actinote zikani]]'', genus ''Actinote''
* [[Archduke (butterfly)|Archdukes]], genus ''Lexias''
* [[California tortoiseshell]], ''Nymphalis californica''
* [[Polygonia c-album|Comma]], ''Polygonia c-album''
* [[Junonia coenia|Common buckeye]], ''Junonia coenia''
* [[Common snout butterfly]], ''Libytheana carinenta''
* Cracker butterflies, genus ''[[Hamadryas (butterfly)|Hamadryas]]''
* [[Crimson patch]], ''Chlosyne janais''
* [[Edith's checkerspot]], ''Euphydryas editha''
* [[Grayling (butterfly)]], ''Hipparchia semele''
* [[Asterocampa celtis|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''
* [[Morpho menelaus|Blue morpho]], ''Morpho menelaus''
* [[Vanessa cardui|Painted lady]], ''Vanessa cardui''
* [[Aglais io|Peacock]], ''Aglais io''
* [[Danaus chrysippus|Plain tiger]], ''Danaus chrysippus''
* [[Polygonia interrogationis|Question mark]], ''Polygonia interrogationis''
* [[Vanessa atalanta|Red admiral]], ''Vanessa atalanta''
* [[Small heath (butterfly)|Small heath]], ''Coenonympha pamphilus''
* [[Small tortoiseshell]], ''Nymphalis urticae''
* [[Gatekeeper (butterfly)|Gatekeeper]], ''Pyronia tithonus''
* [[Small pearl-bordered fritillary]], ''Boloria selene''
* [[Andromeda satyr]], ''Cithaerias andromeda''
* [[Anthanassa texana|Texan crescentspot butterfly]], ''Anthanassa texana texana''
<!-- needs verification * Westin indigo, ''Westin siblis'' -->
* [[Speyeria zerene|Zerene fritillary]], ''Speyeria zerene'' (includes several subspecies such as [[Speyeria zerene hippolyta|Oregon silverspot]], ''Speyeria zerene hippolyta''){{div col end}}

==See also==
{{Portal|Insects|Arthropods}}
*[[List of fritillaries (butterflies)]]

== References ==
{{Reflist}}

== 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)

== External links ==
{{Commons category}}
{{Wikispecies}}

* Canadian Biodiversity Information Facility (2003): [https://web.archive.org/web/20030210152905/http://www.cbif.gc.ca/spp_pages/butterflies/families/nymphalidae_e.php Family Nymphalidae]
* [https://insecta.pro/taxonomy/93 Nymphalidae] "Family Nymphalidae". ''Insecta.pro''.
* Peter Chew: [https://web.archive.org/web/20091027084931/http://www.geocities.com/brisbane_nymphs/index.html Danaids and Browns - Family Nymphalidae], Brisbane butterflies web site (2005).
* [[Tree of Life Web Project]]: [http://tolweb.org/Nymphalidae/12172 Nymphalidae]
* [https://web.archive.org/web/20170312064020/http://en.butterflycorner.net/Family-Nymphalidae.327.0.html Nymphalidae from all over the world]

{{Lepidoptera}}
{{Taxonbar |from=Q156449}}
{{Authority control}}

[[Category:Nymphalidae| ]]
[[Category:Butterfly families]]
[[Category:Taxa named by Constantine Samuel Rafinesque]]
[[Category:Papilionoidea]]