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"पुतली" का संशोधनहरू बिचको अन्तर

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[[File:Charaxes brutus natalensis.jpg|thumb|250px|right|<center>'''पुतली''']]
 
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** [[:en:Riodinidae|रएडिनिडी]]
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'''पुतली''' [[कीरा वर्ग]]को सामान्य रूपले हरेक ठाँउमा पाइने प्राणी हो। यो धेरै सुन्दर तथा आएर्षक हुन्छ।
'''पुतली''' [[कीरा वर्ग]]को सामान्य रूपले हरेक ठाँउमा पाइने प्राणी हो। यो धेरै सुन्दर तथा आएर्षक हुन्छ।<ref>{{cite book |last=त्रिपाठी |first=नरेन्द्र नाथ |title= सरल जीवन विज्ञान, भाग-२ |year=जुलाई २००४ |publisher=शेखर प्रकाशन |location=कोलकाता |id= |page=81-82 |accessday= २३|accessmonth= जून|accessyear= २००९}}</ref> दिनमा जब यो एक [[फूल]] बाट दोस्रो फूलमा उड्छ र [[मधु|मधुपान]] गर्छ तब यसको रंग-बिरंगि पखेटा बाहिर देखिन्छन्। यसको शरीरको मुख्य तीन भाग हों सिर, वक्ष तथा उदर। यिनको दुइ जोडी पखेटा तथा तीन जोडी सन्धियुक्त खुट्टा हुन्छन् अतः यो एक कीरा हो। यसको सिरमा एक जोडी [[संयुक्त आँखे|संयुक्त आँखा]] हुन्छ तथा मुखमा घडीको स्प्रिंग जस्तै प्रोभोसिस नामक खोक्रो लामो सूँडमा जीब्रो हुन्छ जसबाट यो फूलहरूको रस (नेक्टर) चूस्छ। यो एन्टिनाको मदद बाट कुनै वस्तु एवं उनको गंधको ठेगाना पत्ता लगाउँछ।
 
पुतली एकलिंगी प्राणी हो अर्थात नर तथा पोथी अलग-अलग हुन्छन्। पोथी पुतली आफ्नो अण्डे पत्तीको तल्लो सतहमा दिन्छ। अण्डा भन्दा केही दिनहरू पछि एक सानो कीरा निस्किन्छ जसलाई क्याटरपिलर [[लार्भा]] भनिन्छ। यो बिरुवाहरूको पातहरूलाई खाएर ठूलो हुन्छ र फेरि यसको चारै तिर कडा खोल बनाउँछ। अब यसलाई [[प्यूपा]] भनिन्छ। केही समय पछि प्यूपालाई तोडेर उनमा भन्दा एक सुन्दर सानो पुतली बाहिर निस्किन्छ।<ref>{{cite book |last=शर्मा |first=कैलाश नाथ |title= आधुनिक जीव विज्ञान, भाग-२ |year=जुलाई २००४ |publisher=कमला पुस्तक भवन |location=कोलकाता |id= |page=80-81 |accessday= २३|accessmonth= जून|accessyear= २००९}}</ref> पुतलीको दिमाग़ धेरै छिटो हुन्छ। देखने, सूंघने, स्वाद चखने तथा उडनेको वाहेक ठाँउको पहिचाननको यिनीहरूमा अद्भुत क्षमता हुन्छ। वयस्क भए पछि सामान्यतया यो उनी बिरुवाहरू या रुखको तनेमा फिर्ता आउँछं, जहाँ इन्होंने आफ्नो प्रारंभिक समय बिताया हुन्छ।
[[Image:TYOU&HIGANBANA.JPG|thumbnail|left|260px|पुष्प बाट रसास्वादन गर्दै पुतली]]पुतलीको जीवनकाल धेरै सानो हुन्छ। यो ठोस भोजन खाँदैन, हुनत केही पुतलीहरू फूलहरूको रस पीउँछन्। दुनियाको सबै भन्दा छिटो उडने वाला पुतली [[मोनार्च]] छ। यो एक घन्टामा १७ मीलको दूरी तय गर्छ। कोस्टा रीकामा पुतलीहरूको १३०० भन्दा ज्यादा प्रजातिहरू पाइन्छं। दुनियाको सबै भन्दा ठूलो पुतली जायंट बर्डविंग छ, जुन सोलमन आएलैंड्समा पाइन्छ। यस पोथी पुतलीको पंखहरूको फैलाव १२ इंच भन्दा ज्यादा हुन्छ। <ref>{{cite web |url= http://www.bhaskar.com/2009/02/15/0902151845_butterfly.html|title=राम्रो मात्र हैन छिटो दिमाग़ पनि: पुतली
|accessmonthday=[[१ सितंबर]]|accessyear=[[वर्ष २००९|२००९]]|format=एचटीएमएल|publisher=दैनिक भास्कर|language=}}</ref>
 
{{translation}}
 
A '''butterfly''' is a mainly day-flying [[insect]] of the [[Order (biology)|order]] [[Lepidoptera]], the butterflies and [[moth]]s. Like other [[Holometabolism|holometabolous insects]], the butterfly's [[Biological life cycle|life cycle]] consists of four parts, [[egg (biology)|egg]], [[larva]], [[प्युपा]] and adult. Most species are [[Diurnality|diurnal]]. Butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. Butterflies comprise the ''true butterflies'' (superfamily [[Papilionoidea]]), the ''skippers'' (superfamily [[Hesperioidea]]) and the ''moth-butterflies'' (superfamily [[Hedyloidea]]). All the many other families within the Lepidoptera are referred to as [[moth]]s.
 
Butterflies exhibit [[Polymorphism (biology)|polymorphism]], [[mimicry]] and [[aposematism]]. Some, like the [[Monarch (butterfly)|Monarch]], will [[Insect migration|migrate]] over long distances. Some butterflies have evolved symbiotic and parasitic relationships with social insects such as ants. Some species are pests because in their larval stages they can damage domestic crops or trees; however, some species are agents of [[pollination]] of some plants, and caterpillars of a few butterflies (e.g., [[Miletinae|Harvesters]]) eat harmful insects. Culturally, butterflies are a popular motif in the visual and literary arts.
 
==Life cycle==
[[File:Mating Common Buckeyes.jpg|thumb|right|Mating Common Buckeye Butterflies]]
It is a popular belief that butterflies have very short life spans. However, butterflies in their adult stage can live from a week to nearly a year depending on the species. Many species have long larval life stages while others can remain [[Diapause|dormant]] in their pupal or [[egg (biology)|egg]] stages and thereby survive [[winter]]s.<ref>Powell, J. A. 1987. Records of prolonged diapause in Lepidoptera. J. Res. Lepid. 25: 83-109.</ref>
 
Butterflies may have one or more broods per year. The number of generations per year varies from temperate to tropical regions with tropical regions showing a trend towards [[voltinism|multivoltinism]].
 
===Egg===
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===Caterpillars===
[[File:Common Buckeye larva variation, Megan McCarty42.JPG|thumb|right|[[Caterpillars]] of ''[[Junonia coenia]]''.]]
Butterfly larvae, or [[caterpillar]]s, consume [[plant]] [[leaf|leaves]] and spend practically all of their time in search of food. Although most caterpillars are herbivorous, a few species such as ''[[Spalgis epius]]'' and ''[[Liphyra brassolis]]'' are [[entomophagous]] (insect eating).
 
Some larvae, especially those of the [[Lycaenidae]], form [[Mutualism (biology)|mutual associations]] with ants. They communicate with the ants using vibrations that are transmitted through the substrate as well as using chemical signals.<ref>{{cite journal |doi=10.1111/j.1096-3642.1988.tb01201.x |title=The larval ant-organs of Thisbe irenea (Lepidoptera: Riodinidae) and their effects upon attending ants |year=1988 |author=Devries, P.J. |journal=Zoological Journal of the Linnean Society |volume=94 |pages=379}}</ref><ref>{{cite journal |pmid=17733373 |year=1990 |month=Jun |author=Devries, Pj |title=Enhancement of Symbioses Between Butterfly Caterpillars and Ants by Vibrational Communication. |volume=248 |issue=4959 |pages=1104–1106 |doi=10.1126/science.248.4959.1104 |journal=Science (New York, N.Y.)}}</ref> The ants provide some degree of protection to these larvae and they in turn gather [[Honeydew (secretion)|honeydew secretions]].
 
Caterpillars mature through a series of stages called instars. Near the end of each instar, the larva undergoes a process called [[apolysis]], in which the [[cuticle]], a tough outer layer made of a mixture of [[chitin]] and specialized [[proteins]], is released from the softer [[Squamous epithelium|epidermis]] beneath, and the epidermis begins to form a new [[cuticle]] beneath. At the end of each instar, the larva [[ecdysis|moults]] the old [[cuticle]], and the new [[cuticle]] expands, before rapidly hardening and developing pigment. Development of butterfly wing patterns begins by the last larval instar.
 
Butterfly caterpillars have three pairs of true legs from the thoracic segments and up to 6 pairs of [[proleg]]s arising from the abdominal segments. These prolegs have rings of tiny hooks called crochets that help them grip the substrate.
 
Some caterpillars have the ability to inflate parts of their head to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called [[osmeterium|osmeteria]] which are everted to produce smelly chemicals. These are used in defense.
 
Host plants often have toxic substances in them and caterpillars are able to [[Sequestering#Biology|sequester]] these substances and retain them into the adult stage. This helps making them unpalatable to birds and other predators. Such unpalatibility is advertised using bright red, orange, black or white warning colours. The toxic chemicals in plants are often evolved specifically to prevent them from being eaten by insects. Insects in turn develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants.<ref>Ehrlich, P. R., and P. H. Raven. 1964. Butterflies and plants: a study in coevolution. Evolution 18:586 – 608</ref>
 
====Wing development====
[[File:Fifthinstarwingdisk.png|thumb|upright|Last instar wing disk, ''Junonia coenia'']]
[[File:Butterfly Wing close-up.jpg|thumb|Detail of a butterfly wing]]
Wings or wing pads are not visible on the outside of the larva, but when larvae are dissected, tiny developing ''wing disks'' can be found on the second and third thoracic segments, in place of the spiracles that are apparent on abdominal segments.
Wing disks develop in association with a trachea that runs along the base of the wing, and are surrounded by a thin ''peripodial membrane'', which is linked to the outer epidermis of the larva by a tiny duct.
 
Wing disks are very small until the last larval instar, when they increase dramatically in size, are invaded by branching [[Invertebrate trachea|tracheae]] from the wing base that precede the formation of the wing veins, and begin to develop patterns associated with several landmarks of the wing.
 
Near pupation, the wings are forced outside the epidermis under pressure from the [[hemolymph]], and although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval cuticle they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled without damage to the wings.
 
===Pupa===
[[File:Chrysalis5504.jpg|thumb|upright|Chrysalis of [[Gulf Fritillary]]]]
 
When the larva is fully grown, hormones such as prothoracicotropic [[hormone]] (PTTH) are produced. At this point the larva stops feeding and begins "wandering" in the quest of a suitable pupation site, often the underside of a leaf.
 
The larva transforms into a pupa (or [[Pupa#Chrysalis|chrysalis]]) by anchoring itself to a substrate and moulting for the last time. The chrysalis is usually incapable of movement, although some species can rapidly move the abdominal segments or produce sounds to scare potential predators.
 
The pupal transformation into a butterfly through [[metamorphosis (biology)|metamorphosis]] has held great appeal to mankind. To transform from the miniature wings visible on the outside of the pupa into large structures usable for flight, the pupal wings undergo rapid mitosis and absorb a great deal of nutrients. If one wing is surgically removed early on, the other three will grow to a larger size. In the pupa, the wing forms a structure that becomes compressed from top to bottom and pleated from proximal to distal ends as it grows, so that it can rapidly be unfolded to its full adult size. Several boundaries seen in the adult color pattern are marked by changes in the expression of particular transcription factors in the early pupa.
 
===Adult or imago===
The adult, sexually mature, stage of the insect is known as the [[imago]]. As Lepidoptera, butterflies have four wings that are covered with tiny scales (see photo). The fore and hindwings are not hooked together, permitting a more graceful flight. An adult butterfly has six legs, but in the nymphalids, the first pair is reduced. After it emerges from its pupal stage, a butterfly cannot fly until the wings are unfolded. A newly emerged butterfly needs to spend some time inflating its wings with blood and letting them dry, during which time it is extremely vulnerable to [[predator]]s.
Some butterflies' wings may take up to three hours to dry while others take about one hour. Most butterflies and moths will excrete excess dye after hatching. This fluid may be white, red, orange, or in rare cases, blue.
 
==External morphology==
{{Main|Glossary of Lepidopteran terms}}
<gallery>
File:Butterfly parts.svg|Parts of an adult butterfly
File:Butterfly portrait.jpg|Butterflies have two [[Antenna (biology)|antennae]], two compound eyes, and a [[proboscis]]
</gallery>
Adult butterflies have four wings: a forewing and hindwing on both the left and the right side of the body. The body is divided into three segments: the [[head]], [[thorax]], and the [[abdomen]]. They have two [[Antenna (biology)|antennae]], two [[compound eye]]s, and a [[proboscis]].
 
===Scales===
Butterflies are characterized by their scale-covered wings.
The coloration of butterfly wings is created by minute scales. These scales are pigmented with [[melanin]]s that give them blacks and browns, but blues, greens, reds and iridescence are usually created not by pigments but the microstructure of the scales. This structural coloration is the result of [[coherent scattering]] of light by the [[photonic crystal]] nature of the scales.<ref>{{cite journal |doi=10.1021/j150273a001 |year=1927 |author=Mason, C. W. |journal=The Journal of Physical Chemistry |volume=31 |pages=321}}</ref><ref>Vukusic, P., J. R. Sambles, and H. Ghiradella (2000) Optical Classification of Microstructure in Butterfly Wing-scales. Photonics Science News, 6, 61-66, [http://newton.ex.ac.uk/research/emag/butterflies/classification_page.htm EX.ac.uk]</ref><ref>{{cite journal |doi=10.1242/jeb.02051 |year=2006 |month=Feb |author=Prum, Ro; Quinn, T; Torres, Rh |title=Anatomically diverse butterfly scales all produce structural colours by coherent scattering. |volume=209 |issue=Pt 4 |pages=748–65 |issn=0022-0949 |pmid=16449568 |journal=The Journal of experimental biology |url=http://jeb.biologists.org/cgi/pmidlookup?view=long&pmid=16449568 |format=Free full text}}</ref> The scales cling somewhat loosely to the wing and come off easily without harming the butterfly.
 
{{ButterflyScaleMicroscopy}}
 
==Polymorphism==
Many adult butterflies exhibit polymorphism, showing differences in appearance. These variations include geographic variants and seasonal forms. In addition many species have females in multiple forms, often with mimetic forms. Sexual dimorphism in coloration and appearance is widespread in butterflies. In addition many species show [[sexual dimorphism]] in the patterns of ultraviolet reflectivity, while otherwise appearing identical to the unaided human eye. Most of the butterflies have a [[sex-determination system]] that is represented as [[ZW sex-determination system|ZW]] with females being the heterogametic sex (ZW) and males homogametic (ZZ).<ref>{{cite journal |pmid=9292232 |year=1997 |month=Aug |author=Traut, W; Marec, F |title=Sex chromosome differentiation in some species of Lepidoptera (Insecta). |volume=5 |issue=5 |pages=283–91 |issn=0967-3849 |doi=10.1023/B:CHRO.0000038758.08263.c3 |journal=Chromosome research : an international journal on the molecular, supramolecular and evolutionary aspects of chromosome biology}}</ref>
 
Genetic abnormalities such as [[gynandromorph]]y also occur from time to time. In addition many butterflies are infected by ''[[Wolbachia]]'' and infection by the bacteria can lead to the conversion of males into females<ref>{{cite journal |pmid=1361987 |year=1992 |month=Nov |author=Rousset, F; Bouchon, D; Pintureau, B; Juchault, P; Solignac, M |title=Wolbachia endosymbionts responsible for various alterations of sexuality in arthropods. |volume=250 |issue=1328 |pages=91–8 |issn=0962-8452 |doi=10.1098/rspb.1992.0135 |journal=Proceedings. Biological sciences / the Royal Society}}</ref> or the selective killing of males in the egg stage.<ref>{{cite journal |doi=10.1046/j.1365-2540.2001.00804.x |title=Two male-killing Wolbachia strains coexist within a population of the butterfly Acraea encedon |year=2001 |author=Jiggins, Francis M |journal=Heredity |volume=86 |pages=161 |pmid=11380661 |last2=Hurst |first2=GD |last3=Schulenburg |first3=JH |last4=Majerus |first4=ME |issue=Pt 2}}</ref>
 
===Mimicry===
[[File:Heliconius mimicry.png|right|thumb|The [[Heliconius]] butterflies from the tropics of the Western Hemisphere are the classical model for Müllerian mimicry.<ref>{{cite journal |doi=10.1371/journal.pbio.0040341 |year=2006 |month=Oct |author=Meyer, A |title=Repeating patterns of mimicry. |volume=4 |issue=10 |pages=e341 |issn=1544-9173 |pmid=17048984 |pmc=1617347 |journal=PLoS biology |url=http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0040341 |format=Free full text}}</ref>]]
 
[[Batesian mimicry|Batesian]] and [[Mullerian mimicry]] in butterflies is common. Batesian mimics imitate other species to enjoy the protection of an attribute they do not share, [[aposematism]] in this case. The [[Papilio polytes|Common Mormon]] of India has female morphs which imitate the unpalatable red-bodied swallowtails, the [[Pachliopta aristolochiae|Common Rose]] and the [[Pachliopta hector|Crimson Rose]]. Mullerian mimicry occurs when aposematic species evolve to resemble each other, presumably to reduce predator sampling rates, the [[Heliconius]] butterflies from the Americas being a good example.
 
Wing markings called [[Eyespot (mimicry)|eyespots]] are present in some species; these may have an [[automimicry]] role for some species. In others, the function may be intraspecies communication, such as mate attraction. In several cases, however, the function of butterfly eyespots is not clear, and may be an evolutionary anomaly related to the relative elasticity of the genes that encode the spots.<ref>
{{cite journal | author = Brunetti CR et al. | year = 2001 | month = 10 | title = The generation and diversification of butterfly eyespot color patterns. | journal = J. of Cell Biology | volume = 11 | issue = 20 | pages = 1578–85 | pmid = : 11676917 | accessdate = 2006-08-22}}
</ref><ref>{{cite journal | author = Brakefield, PM et al. | year = 1996 | title = Development, plasticity and evolution of butterfly eyespot patterns. | journal = Nature | issue = 384 | pages = 236–242 | accessdate = 2006-08-22 | volume = 384 | doi = 10.1038/384236a0 | pmid = 12809139}}</ref>
 
===Seasonal polyphenism===
<div name="wet-dry forms"/>Many of the tropical butterflies have distinctive seasonal forms. This phenomenon is termed [[seasonal polyphenism]] and the seasonal forms of the butterflies are called the dry-season and wet-season forms. How the season affects the genetic expression of patterns is still a subject of research.<ref>{{cite journal |pmid=18811432 |year=1998 |month=Dec |author=Brakefield, Pm; Kesbeke, F; Koch, Pb |title=The regulation of phenotypic plasticity of eyespots in the butterfly Bicyclus anynana. |volume=152 |issue=6 |pages=853–60 |issn=0003-0147 |doi=10.1086/286213 |journal=The American naturalist}}</ref> Experimental modification by ecdysone hormone treatment has demonstrated that it is possible to control the continuum of expression of variation between the wet and dry-season forms.<ref>{{cite journal |pmid=12492404 |year=2003 |month=Jan |author=Nijhout, Hf |title=Development and evolution of adaptive polyphenisms. |volume=5 |issue=1 |pages=9–18 |issn=1520-541X |journal=Evolution & development |doi=10.1046/j.1525-142X.2003.03003.x}}</ref> The dry-season forms are usually more cryptic and it has been suggested that the protection offered may be an adaptation. Some also show greater dark colours in the wet-season form which may have thermoregulatory advantages by increasing ability to absorb solar radiation.<ref>{{cite journal |doi=10.1111/j.1095-8312.1984.tb00795.x |title=The evolutionary significance of dry and wet season forms in some tropical butterflies |year=1984 |author=Brakefield, PAUL M. |journal=Biological Journal of the Linnean Society |volume=22 |pages=1 |last2=Larsen |first2=Torben B.}}</ref>
 
''Bicyclus anynana'' is a species of butterfly that exhibits a clear example of seasonal [[polyphenism]]. These butterflies, endemic to Africa, have two distinct phenotypic forms that alternate according to the season. The wet-season forms have large, very apparent ventral eyespots whereas the dry-season forms have very reduced, oftentimes nonexistent, ventral eyespots. Larvae that develop in hot, wet conditions develop into wet-season adults where as those growing in the transition from the wet to the dry season, when the temperature is declining, develop into dry-season adults.<ref>Lyytinen, A., P. M. Brakefield, L. Lindström, and J. Mappes. 2004. Does predation maintain eyespot plasticity in Bicyclus anynana. The Proceedings of the Royal Society B-Biological Sciences 271:279-283.</ref> This polyphenism has an adaptive role in'' B. anynana''. In the dry-season it is disadvantageous to have conspicuous eyespots because '' B. anynana'' blend in with the brown vegetation better without eyespots. By not developing eyespots in the dry-season they can more easily camouflage themselves in the brown brush. This minimizes the risk of visually mediated predation. In the wet-season, these brown butterflies cannot as easily rely on cryptic coloration for protection because the background vegetation is green. Thus, eyespots, which may function to decrease predation, are beneficial for ''B. anynana'' to express.<ref>Brakefield, P. M., J. Gates, D. Keys, F. Kesbeke, P. J. Wijngaarden, A. Monteiro, V. French, and S. B. Carroll. 1996. Development, plasticity and evolution of butterfly eyespot patterns. Nature 384:236-242.</ref>
 
==Habits==
[[File:antennae ctb.png|thumb|Antennal shape in the [[Lepidoptera]] from C. T. Bingham (1905)]]
[[File:Australian painted lady feeding.jpg|thumb|The [[Australian painted lady]] feeding on a flowering shrub]]
Butterflies feed primarily on [[nectar]] from flowers. Some also derive nourishment from [[pollen]],<ref>{{cite journal|author=Gilbert LE|year=1972|title= Pollen feeding and reproductive biology of ''Heliconius'' butterflies|journal=Proceedings of the National Academy of Sciences|volume=69|pages=1402–1407|url=http://www.pnas.org/content/69/6/1403.abstract|doi=10.1073/pnas.69.6.1403}}</ref> tree sap, rotting fruit, dung<ref>At 7 to 8 pm 27 October 2009 the [[BBC2]] television program ''Ray Mears's Northern Wilderness'' showed a butterfly feeding from wolf faeces in the Canadian [[boreal forest]]</ref>, decaying flesh<ref>At 7.30 to 8 pm 27 October 2009 the [[ITV1]] television program ''[[Grimefighters]]'' showed a butterfly feeding from a decaying dead rat in a town in England</ref>, and dissolved minerals in wet sand or dirt. Butterflies are important as pollinators for some species of plants although in general they do not carry as much pollen load as [[bee]]s. They are however capable of moving pollen over greater distances.<ref>{{cite journal|author=Herrera, C.M.|year= 1987 |title=Components of pollinator "quality": comparative analysis of a diverse insect assemblage.|journal= Oikos|volume=50|pages=79–90 |url=http://ebd06.ebd.csic.es/pdfs/Herrera.1987.Oikos.pdf|doi=10.2307/3565403|jstor=3565403|issue=1|publisher=Oikos, Vol. 50, No. 1}}</ref>
 
As adults, butterflies consume only liquids and these are sucked by means of their [[proboscis]]. They feed on [[nectar]] from flowers and also sip water from damp patches. This they do for water, for energy from sugars in nectar and for sodium and other minerals which are vital for their reproduction. Several species of butterflies need more [[सोडियम|sodium]] than provided by [[nectar (plant)|nectar]]. They are attracted to sodium in [[salt]] and they sometimes land on people, attracted by human sweat. Besides damp patches, some butterflies also visit dung, rotting fruit or carcasses to obtain minerals and nutrients. In many species, this [[mud-puddling]] behaviour is restricted to the males, and studies have suggested that the nutrients collected are provided as a nuptial gift along with the spermatophore during mating.<ref>Molleman Freerk, Grunsven Roy H. A., Liefting Maartje, Zwaan Bas J., Brakefield Paul M. (2005) Is male puddling behaviour of tropical butterflies targeted at sodium for nuptial gifts or activity? Biol. J. Linn. Soc. 86, (3):345-361</ref>
 
Butterflies sense the air for scents, wind and nectar using their antennae. The antennae come in various shapes and colours. The hesperids have a pointed angle or hook to the antennae, while most other families show knobbed antennae. The antennae are richly covered with sensillae. A butterfly's sense of [[taste]] is coordinated by chemoreceptors on the [[Arthropod leg#Insects|tarsi]], which work only on contact, and are used to determine whether an egg-laying insect's offspring will be able to feed on a leaf before eggs are laid on it.<ref>{{cite web|url=http://www.sandiegozoo.org/animalbytes/t-butterfly.html |title=Article on San Diego Zoo website |publisher=Sandiegozoo.org |date= |accessdate=2009-03-30}}</ref> Many butterflies use chemical signals, [[pheromones]], and specialized scent scales (androconia) and other structures (coremata or 'Hair pencils' in the Danaidae) are developed in some species.
 
Vision is well developed in butterflies and most species are sensitive to the ultraviolet spectrum. Many species show sexual dimorphism in the patterns of UV reflective patches.<ref>Obara Y, Hidaki T. (1968). Recognition of the female by the male, on the basis of ultra-<script type="text/javascript" src="http://ne.wikipedia.org/w/index.php?title=MediaWiki:translit.js&action=raw&ctype=text/javascript&dontcountme=s"&action="raw"></script>violet reflection, in the white cabbage butterfly ''Pieris rapae crucivora'' Boisduval. Proc. Japan Acad., 44: 829-832.</ref> Color vision may be widespread but has been demonstrated in only a few species.<ref>Tadao Hirota and Yoshiomi Kato 2004 Color discrimination on orientation of female ''Eurema hecabe'' (Lepidoptera: Pieridae) Applied Entomology and Zoology Vol. 39:229-233 [http://www.jstage.jst.go.jp/article/aez/39/2/229/_pdf JST.go.jp]</ref><ref>Michiyo Kinoshita, Naoko Shimada And Kentaro Arikawa (1999) Color vision of the foraging swallowtail butterfly ''Papilio
xuthus''. The Journal of Experimental Biology 202:95 – 102 [http://jeb.biologists.org/cgi/reprint/202/2/95.pdf]</ref>
 
Some butterflies have organs of hearing and some species are also known to make stridulatory and clicking sounds.<ref>Swihart, S. L (1967). Hearing in butterflies. J. Insect Physiol 13, 469</ref>
[[File:Monarch butterflies.jpg|thumb|left|Monarch butterflies]]
Many butterflies, such as the [[Monarch butterfly]], are [[Insect migration|migratory]] and capable of long distance flights. They migrate during the day and use the sun to orient themselves. They also perceive polarized light and use it for orientation when the sun is hidden.<ref>Reppert, Steven M.; Haisun Zhu; White, Richard H. (2004) Polarized light helps monarch butterflies navigate. Current biology 14(2):155-158</ref>
 
Many species of butterfly maintain territories and actively chase other species or individuals that may stray into them. Some species will bask or perch on chosen perches. The flight styles of butterflies are often characteristic and some species have courtship flight displays. Basking is an activity which is more common in the cooler hours of the morning. Many species will orient themselves to gather heat from the sun. Some species have evolved dark wingbases to help in gathering more heat and this is especially evident in alpine forms.<ref>Ellers, J. and Carol L. Boggs (2002) The evolution of wing color in ''Colias'' butterflies: Heritability, Sex Linkage,
and population divergence. Evolution, 56(4):836 – 840 [http://www.stanford.edu/group/CCB/Pubs/Boggs_pdfs/2002_Ellers_Boggs_Coliaswingcolor.pdf Stanford.edu]</ref>
 
==Flight==
[[File:Butterfly midflight.jpg|thumb|''Geitoneura klugii'' taking off]]
:''See also [[Insect flight]]''
Like many other members of the [[insect]] world, the lift generated by butterflies is more than what can be accounted for by steady-state, non-transitory [[aerodynamics]]. Studies using ''[[Vanessa atalanta]]'' in a windtunnel show that they use a wide variety of aerodynamic mechanisms to generate force. These include wake capture, vortices at the wing edge, rotational mechanisms and Weis-Fogh 'clap-and-fling' mechanisms. The butterflies were also able to change from one mode to another rapidly.<ref>Srygley, R. B. and A. L. R. Thomas (2002) Aerodynamics of insect flight: flow visualisations with free flying butterflies reveal a variety of unconventional lift-generating mechanisms. Nature 420: 660-664. [http://users.ox.ac.uk/~zool0206/s&t02.pdf OX.ac.uk]</ref>
 
===Migration===
[[File:Monarch In May.jpg|thumb|The [[Monarch butterfly]] migrates large distances]]
{{Main|Lepidoptera migration}}
:''See also [[Insect migration]]''
Many butterflies migrate over long distances. Particularly famous migrations are those of the [[Monarch butterfly]] from [[Mexico]] to northern [[संयुक्त राज्य अमेरिका|USA]] and southern [[Canada]], a distance of about 4000 to 4800&nbsp;km (2500–3000 miles). Other well known migratory species include the [[Vanessa cardui|Painted Lady]] and several of the [[Danainae|Danaine]] butterflies. Spectacular and large scale migrations associated with the [[Monsoons]] are seen in peninsular [[भारत|India]].<ref>Williams, C. B. 1927 A study of butterfly migration in south India and Ceylon, based largely on records by Messrs. G Evershed, E.E.Green, J.C.F. Fryer and W. Ormiston. Trans. Ent. Soc. London 75:1-33</ref> Migrations have been studied in more recent times using wing tags and also using stable hydrogen isotopes.<ref>Urquhart, F. A. & N. R. Urquhart. 1977. Overwintering areas and migratory routes of the Monarch butterfly (''Danaus p. plexippus'', Lepidoptera: Danaidae) in North America, with special reference to the western population. Can. Ent. 109: 1583-1589</ref><ref>Wassenaar L.I., Hobson K.A. 1998. Natal origins of migratory monarch butterflies at wintering colonies in Mexico: new isotopic evidence. Proc Natl Acad Sci U S A. 95(26):15436-9. [http://www.pnas.org/cgi/content/full/95/26/15436 Full text PNAS.org]</ref>
 
Butterflies have been shown to navigate using time compensated sun compasses. They can see polarized light and therefore orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is suggested to be particularly important.<ref>Ivo Sauman, Adriana D. Briscoe, Haisun Zhu, Dingding Shi, Oren Froy, Julia Stalleicken, Quan Yuan, Amy Casselman, and Steven M. Reppert (2005) Connecting the Navigational Clock to Sun Compass Input in Monarch Butterfly Brain. Neuron. 46:457-467 [http://www.neuron.org/content/article/abstract?uid=PIIS0896627305002369 Neuron.org]</ref>
 
It is suggested that most migratory butterflies are those that belong to semi-arid areas where breeding seasons are short.<ref>Southwood, T. R. E. 1962. Migration of terrestrial arthropods in relation to habitat. Biol. Rev. 37:171-214</ref> The life-histories of their host plants also influence the strategies of the butterflies.<ref>Dennis, R L H, Tim G. Shreeve, Henry R. Arnold and David B. Roy (2005) Does diet breadth control herbivorous insect distribution size? Life history and resource outlets for specialist butterflies. Journal of Insect Conservation 9(3):187-200</ref>
 
==Defense==
[[File:Leopard Lacewing Cethosia cyane 2 Richard Bartz .jpg|thumb|upright|The wings of a butterfly (Leopard Lacewing ''[[Cethosia cyane]]'') become increasingly damaged as it ages, and do not repair]]
Butterflies are threatened in their early stages by [[parasitoid]]s and in all stages by [[predator]]s, diseases and environmental factors. They protect themselves by a variety of means.
 
Chemical defenses are widespread and are mostly based on chemicals of plant origin. In many cases the plants themselves evolved these toxic substances as [[plant defense against herbivory|protection]] against herbivores. Butterflies have evolved mechanisms to [[sequester]] these plant toxins and use them instead in their own defense.<ref>Nishida, Ritsuo (2002). Sequestration of defensive substances from plants by Lepidoptera. Annu. Rev. Entomol. 47:57–92</ref> These defense mechanisms are effective only if they are also well advertised and this has led to the evolution of bright colours in unpalatable butterflies. This signal may be [[mimicry|mimicked]] by other butterflies. These mimetic forms are usually restricted to the females.
[[File:Citrus Swallowtail Papilio demodocus.jpg|right|thumb|Eyespots on the wings of this butterfly are part of the animal's defense]]
Cryptic coloration is found in many butterflies. Some like the oakleaf butterfly are remarkable imitations of leaves.<ref>Robbins, Robert K. (1981) The "False Head" Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies. American Naturalist 118(5):770-775</ref> As caterpillars, many defend themselves by freezing and appearing like sticks or branches. Some papilionid caterpillars resemble bird dropping in their early instars. Some caterpillars have hairs and bristly structures that provide protection while others are gregarious and form dense aggregations. Some species also form associations with ants and gain their protection (See [[Myrmecophile]]).
 
Behavioural defenses include perching and wing positions to avoid being conspicuous. Some female Nymphalid butterflies are known to guard their eggs from parasitoid wasps.<ref>Nafus, D. M. and I. H. Schreiner (1988) Parental care in a tropical nymphalid butterfly ''Hypolimas anomala''. Anim. Behav. 36: 1425- 143</ref>
 
Eyespots and tails are found in many lycaenid butterflies and these divert the attention of predators from the more vital head region. An alternative theory is that these cause ambush predators such as [[spider]]s to approach from the wrong end and allow for early visual detection.<ref>William E. Cooper, Jr. (1998) Conditions favoring anticipatory and reactive displays deflecting predatory attack. Behavioral Ecology</ref>
 
A butterfly's hind wings are thought to allow the butterfly to take swift, tight turns to evade predators.<ref>[http://newswise.com/articles/view/547899/ Hind Wings Help Butterflies Make Swift Turns to Evade Predators] Newswise, Retrieved on January 8, 2008.</ref>
 
==Notable species==
[[File:Siproeta epaphus Galawebdesign.jpg|thumb|right|Rusty-tipped Page (''[[Siproeta epaphus]]''), Butterfly World (Florida)]]
There are between 15,000 and 20,000 [[species]] of butterflies worldwide. Some well-known species from around the world include:
* [[Swallowtail butterfly|Swallowtail]]s and [[Birdwing]]s, Family '''[[Papilionidae]]'''
** [[Swallowtail butterfly|Common Yellow Swallowtail]], ''Papilio machaon''
** [[Spicebush Swallowtail]], ''Papilio troilus''
** [[Papilio demoleus|Lime Butterfly]], ''Papilio demoleus''
** [[Birdwing|''Ornithoptera'']] genus (Birdwings; the largest butterflies)
* [[Pieridae|White]]s and Yellows, Family '''[[Pieridae]]'''
** [[Small White]], ''Pieris rapae''
** [[Green-veined White]], ''Pieris napi''
** [[Delias eucharis|Common Jezebel]], ''Delias eucharis''
* [[Lycaenidae|Blues and Coppers]] or Gossamer-Winged Butterflies, Family '''[[Lycaenidae]]'''
** [[Xerces Blue]], ''Glaucopsyche xerces'' (extinct)
** [[Karner Blue]], ''Lycaeides melissa samuelis'' (endangered)
** [[Talicada nyseus|Red Pierrot]], ''Talicada nyseus''
* [[Metalmark butterfly|Metalmark butterflies]], Family '''[[Riodinidae]]'''
** [[Hamearis lucina|Duke of Burgundy]], ''Hamearis lucina''
** [[Abisara echerius|Plum Judy]], ''Abisara echerius''
* [[Brush-footed butterfly|Brush-footed butterflies]], Family '''[[Nymphalidae]]'''
** [[Vanessa cardui|Painted Lady]], or Cosmopolitan, ''Vanessa cardui''
** [[Monarch butterfly]], ''Danaus plexippus''
** [[Morpho (butterfly)|''Morpho'']] genus
** [[Speckled Wood]], ''Pararge aegeria''
* [[Skipper butterfly|Skippers]], Family '''[[Hesperiidae]]'''
** [[Mallow Skipper]], ''Carcharodus alceae''
** [[Zabulon Skipper]], ''Poanes zabulon''
 
==In culture==
===Art===
Artistic depictions of butterflies have been used in many cultures including Egyptian hieroglyphs 3500 years ago.<ref>Larsen, Torben (1994) Butterflies of Egypt. Saudi Aramco world. 45(5):24-27 [http://www.saudiaramcoworld.com/issue/199405/butterflies.of.egypt.htm Online]</ref> Today, butterflies are widely used in various objects of art and jewelry: mounted in frame, embedded in resin, displayed in bottles, laminated in paper, and used in some mixed media artworks and furnishings.<ref>{{cite web|url=http://mfjoe.com/tag/furniture/ |title=Table complete with real butterflies embedded in resin |publisher=Mfjoe.com |date= |accessdate=2009-03-30}}</ref> Butterflies have also inspired the "butterfly [[fairy]]" as an art and fictional character, including in the ''[[Barbie Mariposa]]'' film.
 
===Symbolism===
According to the “Butterflies” chapter in ''[[Kwaidan: Stories and Studies of Strange Things]]'', by [[Lafcadio Hearn]], a butterfly is seen as the [[personification]] of a person's soul; whether they be living, dying, or already dead. One Japanese superstition says that if a butterfly enters your guestroom and perches behind the [[bamboo]] screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad [[omen]]s. When [[Taira no Masakado]] was secretly preparing for his famous revolt, there appeared in [[Kyoto]] so vast a swarm of butterflies that the people were frightened — thinking the apparition to be a portent of coming evil.<ref>{{cite book | last = Hearn | first = Lafcadio | authorlink = Lafcadio Hearn | year = 1904 | title = Kwaidan: Stories and Studies of Strange Thing | publisher = Dover Publications, Inc. | isbn = 0-486-21901-1}}</ref>
 
The Russian word for "butterfly", бабочка (''bábochka''), also means "[[bow tie]]". It is a diminutive of "baba" or "babka" (= "woman, grandmother, [[cake]]"), whence also "babushka" = "[[grandmother]]".
 
The Ancient Greek word for "butterfly" is ψυχή (''psȳchē''), which primarily means "[[soul]]", "[[mind]]".<ref>Hutchins, M., Arthur V. Evans, Rosser W. Garrison and Neil Schlager (Eds) (2003) Grzimek's Animal Life Encyclopedia, 2nd edition. Volume 3, Insects, Farmington Hills, MI: Gale Group, 2003.</ref>
 
According to Mircea Eliade's ''Encyclopedia of Religion'', some of the [[Naga people|Nagas]] of [[Manipur]] trace their ancestry from a butterfly.<ref name="Rabuzzi">Rabuzzi, M. 1997. Butterfly etymology. Cultural Entomology November 1997 Fourth issue [http://www.insects.org/ced4/etymology.html online]</ref>
 
[[File:Xvxi1.jpg|thumb|upright|Butterfly and Chinese [[wisteria]]flowers, by Xü Xi (c.886&ndash;c.975), painted around 970 during the early [[Song Dynasty]].]]
In [[Chinese culture]] two butterflies flying together are a symbol of love. Also a famous Chinese folk story called [[Butterfly Lovers]]. The [[Taoist]] philosopher [[Zhuangzi]] once had a dream of being a butterfly flying without care about humanity, however when he woke up and realized it was just a dream, he thought to himself "Was I before a man who dreamt about being a butterfly, or am I now a butterfly who dreams about being a man?"
 
In some old cultures, butterflies also symbolize [[Rebirthing|rebirth]] into a new life after being inside a cocoon for a period of time.
 
[[Jose Rizal]] delivered a speech in [[1884]] in a banquet and mentioned "the Oriental chrysalis ... is about to leave its cocoon" comparing the emergence of a "new Philippines" with that of butterfly metamorphosis.<ref>{{cite web|url=http://joserizal.info/Writings/Speeches/speeches.htm |title=The Best Known Speech of Jose Rizal |publisher=Joserizal.info |date= |accessdate=2009-03-30}}</ref> He has also often used the butterfly imagery in his poems and other writings to express the Spanish Colonial Filipinos' longing for liberty.<ref>{{cite web|url=http://joserizal.info/Biography/man_and_martyr/chapter14.htm |title=The Life and Writings of Dr. Jose Rizal |publisher=Joserizal.info |date= |accessdate=2009-03-30}}</ref> Much later, in a letter to [[Ferdinand Blumentritt]], Rizal compared his life in exile to a weary butterfly with sun-burnt wings.<ref>{{cite web|url=http://www.univie.ac.at/Voelkerkunde/apsis/aufi/rizal/rbcor193.htm |title=193. Rizal, Dapitan, 19 December 1898 |publisher=Univie.ac.at |date= |accessdate=2009-03-30}}</ref>
 
[[File:Carl Spitzweg 033.jpg|thumb|upright|Der Schmetterlingsjäger (The butterfly hunter) by [[Carl Spitzweg]] (1840), a depiction from the era of butterfly collection.]]
 
Some people say that when a butterfly lands on you it means good luck.{{Citation needed|date=March 2009}} However, in [[Devon]]shire, people would traditionally rush around to kill the first butterfly of the year that they see, or else face a year of bad luck.<ref>[[Dorset Chronicle]], May 1825, reprinted in: [http://books.google.co.uk/books?ct=result&id=6-E8AAAAIAAJ&jtp=678 "The First Butterfly"], in ''The Every-day Book and Table Book; or, Everlasting Calendar of Popular Amusements, Sports, Pastimes, Ceremonies, Manners, Customs, and Events, Each of the Three Hundred and Sixty-Five Days, in Past and Present Times; Forming a Complete History of the Year, Months, and Seasons, and a Perpetual Key to the Almanac, Including Accounts of the Weather, Rules for Health and Conduct, Remarkable and Important Anecdotes, Facts, and Notices, in Chronology, Antiquities, Topography, Biography, Natural History, Art, Science, and General Literature; Derived from the Most Authentic Sources, and Valuable Original Communication, with Poetical Elucidations, for Daily Use and Diversion. Vol III.'', ed. William Hone, (London: 1838) p 678.</ref> Also, in the [[Philippines]], a lingering black butterfly or moth in the house is taken to mean that someone in the family has died or will soon die.<ref>{{cite web|url=http://www.sunstar.com.ph/static/dav/2005/10/30/feat/death.practices.philippine.style.html |title=Death practices Philippine style |publisher=Sunstar.com.ph |date=2005-10-30 |accessdate=2009-03-30}}</ref><ref>{{cite web|author=don.herrington@livinginthephilippines.com |url=http://www.livinginthephilippines.com/philculture/superstitions_and_beliefs.html |title=Superstitions and Beliefs related to Death |publisher=Livinginthephilippines.com |date= |accessdate=2009-03-30}}</ref>
 
The idiom "[[butterflies in the stomach]]" is used to describe a state of nervousness.
 
In the [[NBC]] television show [[Kings (American TV series)|''Kings'']], butterflies are the national symbol of the fictional nation of Gilboa and a sign of God's favor.
 
===Technological inspiration===
Researches on the wing structure of [[Birdwing|Palawan Birdwing]] butterflies led to new wide wingspan kite and aircraft designs.<ref>{{cite web|url=http://www.nagypal.net/ |title=An Introduction to The World of Birdwing Butterflies |publisher=Nagypal.net |date=2000-05-28 |accessdate=2009-03-30}}</ref>
 
Studies on the reflection and scattering of light by the scales on wings of swallowtail butterflies led to the innovation of more efficient [[light-emitting diode]]s.<ref>Vukusic, Pete and Ian Hooper. 2005. Directionally Controlled Fluorescence Emission in Butterflies Science. 310(5751):1151 DOI: 10.1126/science.1116612</ref>
 
The structural coloration of butterflies is inspiring nanotechnology research to produce paints that do not use toxic pigments and in the development of new display technologies.
 
The discoloration and health of butterflies in butterfly farms, is now being studied for use as indicators of air quality in several cities.
 
==See also==
*[[Butterfly Alphabet]]
*[[Butterfly zoo]]
*[[Differences between butterflies and moths]]
*[[Florida Museum of Natural History#McGuire Center for Lepidoptera and Biodiversity]]
*[[List of butterflies of Great Britain]]
*[[List of butterflies of India]]
*[[List of butterflies of Menorca]]
*[[List of butterflies of North America]]
*[[List of butterflies in Taiwan]]
*[[List of butterflies of Tobago]]
*[[List of U.S. state butterflies]]
*[[List of Australian butterflies]]
*[[Moth]]
 
==Field guides to butterflies==
Some [[field guides]] to butterfly species include:<ref>For a more comprehensive list, see the [http://media.library.uiuc.edu/cgi/b/bib/bix-idx?type=simple&c=bix&sid=54d8a20e4f1eb5f2de074bad4caba7ae&Submit=search&sort=title&q1=butterflies&rgn1=Entire+record International Field Guides] database</ref>
* ''Butterflies of North America'', Jim P. Brock and Kenn Kaufman (2003)
* ''Butterflies through Binoculars: The East'', Jeffrey Glassberg (1999)
* ''Butterflies through Binoculars: The West'', Jeffrey Glassberg (2001)
* ''A Field Guide to Eastern Butterflies'', Paul Opler (1994)
* ''A Field Guide to Western Butterflies'', Paul Opler (1999)
* ''Peterson First Guide to Butterflies and Moths'', Paul Opler (1994)
* ''Las Mariposas de Machu Picchu'' by Gerardo Lamas (2003)
* ''The Millennium Atlas of Butterflies in Britain and Ireland'' by Jim Asher (Editor), et al.
* ''Pocket Guide to the Butterflies of Great Britain and Ireland'' by Richard Lewington
* ''Butterflies of Britain and Europe'' (Collins Wildlife Trust Guides) by Michael Chinery
* ''Butterflies of Europe'' by Tom Tolman and Richard Lewington (2001)
* ''Butterflies of Europe New Field Guide and Key'' by Tristan Lafranchis (2004)
* ''Butterflies of Lebanon'' by Torben B. Larsen. Beirut. (1974)
* ''The butterflies of Saudi Arabia and its neighbours''. by Torben B. Laren (Stacey intl.) (1984)
* ''The butterflies of Egypt'' by Torben B. Larsen (Apollo Books, Denmark). (1990)
* ''Field Guide to Butterlies of South Africa'' by Steve Woodhall (2005)
* ''The butterflies of Kenya and their natural history'' by Torben B. Larsen (OUP) (1991)
* ''Butterflies of Sikkim Himalaya and their Natural History'' by Meena Haribal (1994).
* ''Butterflies of Peninsular India'' by Krushnamegh Kunte, Universities Press (2005).
* ''Butterflies of the Indian Region'' by Col M. A. Wynter-Blyth, Bombay Natural History Society, Mumbai, India (1957).
* ''A Guide to Common Butterflies of Singapore'' by Steven Neo Say Hian (Singapore Science Centre)
* ''Butterflies of West Malaysia and Singapore'' by W.A.Fleming. (Longman Malaysia)
* ''The Butterflies of the Malay Peninsula'' by A.S. Corbet and H. M. Pendlebury. (The Malayan Nature Society)
* ''Butterflies of West Africa'' (two vols.) by Torben B. Larsen. (Apollo Books, Denmark) (2005)
* ''Oxford Butterflies of India'' by Thomas Gray, I.D.Kehimkar, J Punetha, [[Oxford University Press]] (2008)
 
==Cited references==
{{Reflist|2}}
 
==Other references==
* Boggs, C., Watt, W., Ehrlich, P. 2003. Butterflies: Evolution and Ecology Taking Flight. [[University of Chicago Press]], Chicago, USA.
* Darby, Gene, 1958. What Is A Butterfly. Chicago, [[Benefic Press]]. pp.&nbsp;5 – 48.
* Heppner, J. B. 1998. Classification of Lepidoptera. ''Holarctic Lepidoptera'', Suppl. 1.
* Monteiro, A. and N. E. Pierce. 2001. Phylogeny of Bicyclus (Lepidoptera : Nymphalidae) inferred from COI, COII, and EF-1 alpha gene sequences. Molecular Phylogenetics and Evolution 18:264-281.
* Nemos, F. ca. 1895. Europas bekannteste Schmetterlinge. Beschreibung der wichtigsten Arten und Anleitung zur Kenntnis und zum Sammeln der Schmetterlinge und Raupen Oestergaard Verlag, Berlin, [http://hdl.handle.net/10013/epic.28790.d001 (pdf 77MB)]
* Peña, C., N. Waklberg, E. Weingartner, U. Kodandaramaiah, S. Nylin, A. V. L. Freitas, and A. V. Z. Brower. 2006. Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Molecular Phylogenetics and Evolution 40:29-49.
* Pyle, R. M. 1992. Handbook for Butterfly Watchers. [[Houghton Mifflin]]. First published, 1984. ISBN 0-395-61629-8
* Stevens, M. 2005. The role of eyespots as anti-predator mechanisms, principally demonstrated in the Lepidoptera. Biological Reviews 80:573-588.
 
==External links==
===General interest===
{{commons+cat|Butterfly|Butterflies}}
{{Wiktionary|Butterfly}}
*[http://www.itvlocal.com/meridian/news/?void=279599 The Royal Horticultural Society butterfly exhibition]
*[http://tolweb.org/Papilionoidea/12027 Papilionoidea on the Tree of Life Web project]
*[http://entomology.ifas.ufl.edu/creatures/main/search_common.htm#bfly Butterflies] on the [[University of Florida|UF]] / [[Institute of Food and Agricultural Sciences|IFAS]] Featured Creatures Web site
 
===Regional lists===
* [http://www.pinocchio.it/eng/butterflyhouse/ Collodi Butterfly House] Tuscany
* [http://www.butterfliesandmoths.org/ Butterflies and Moths of North America]
* [http://butterfliesofamerica.com/intro.htm Butterflies of America]
* [http://www.cbif.gc.ca/spp_pages/butterflies/speciesindex_e.php Butterflies of Canada]
* [http://www.naba.org North American Butterfly Association (NABA)]
* [http://www.leps.nl/ Butterflies and Moths in the Netherlands]
* [http://perso.orange.es/meul_12345/indgalb.htm Butterflies of Spain and Portugal]
* [http://www.wwfpak.org/nap/dnap_wildlife_wildfauna_mammals_insect.php Insect and butterfly diversity of Pakistan]
* [http://www.philippinebutterfly.com/gallery.php?category=BUTTERFLIES Butterflies of the Philippines]
* [http://www.naturemagics.com/butterfly.shtm Butterflies of Southern India]
* [http://www.srilankaninsects.net/Butterflies/MainPage/ButterfliesMain.htm Butterflies of Sri Lanka]
* [http://www.oocities.com/RainForest/Vines/2382/sgchecklist.htm Butterflies of Singapore]
* [http://www.nature-of-oz.com/israelsspecieslist1.htm Israel Insect World]
* [http://www.oocities.com/RainForest/Vines/2382/sgchecklist.htm Singapore Butterfly Checklist]
* [http://www.butterfly.org.tw Butterfly Conservation Society of Taiwan]
* [http://www.ctv.es/USERS/tarrier/tarrier_M/index.htm Butterflies of Morocco]
* [http://yutaka.it-n.jp/ Butterflies of Indo-China] Chiefly Thailand, Laos and Vietnam.
* [http://www.flmnh.ufl.edu/butterflies/neotropica/sulawesi/checklist.html Butterflies of Southeastern Sulawesi]
* [http://www.repository.naturalis.nl/document/46743 Naturalis] Butterflies of Sulawesi (Illustrated pdf)
* [http://www.malaeng.com/blog/?page_id=1293 Butterflies of Thailand]
* [http://www.mariposasmexicanas.com/ Butterflies of Mexico]
* [http://ghanabutterflies.uw.hu/ Butterflies of Ghana]
 
==Literature==
*[http://www.entomologische-literatur.de/startseite.html Literaturatenbank] Free downloads
 
===Images/Movies===
<!-- Wikipedia is not a sales tool. Only include images that are free for non-commercial use. That license is not appropriate for the wiki, but does provide value to the readers. -->
* [http://bugguide.net/node/view/81/bgimage BugGuide.net Many images of North American butterflies, many licensed via Creative Commons]
* [http://greennature.com/article1273.html Butterfly Pictures and Information]
* [http://marlonstein.com/fotos/album/Insetos/Borboletas/ Butterfly of Brazil]
* [http://cirrusimage.com/butterfly_photos.htm Reference quality large format photographs, common butterflies of North America]
* [http://harperkay.homestead.com/ButterfliesAndMothsIndex.html Gallery of Florida Butterflies and Moths]
* [http://www.floridanaturepictures.com/butterflies/butter.html Butterfly Picture Gallery]
* [http://socalbutterflies.com Photographs of most of the Butterflies in Southern California]
* [http://www.oocities.com/haarfager9/ Butterflies of Southern Illinois]
* [http://www.butterfliesoffrance.com/ Butterflies of France]
* [http://perso.orange.es/meul_12345/indgalb.htm Butterflies of Spain and Portugal]
* [http://tolweb.org/movies/Papilionoidea/12027 Butterfly Movies (Tree of Life)]
* [http://www.massbutterflies.org/construct-group-page.asp?gr=All 1000+ photos of Massachusetts butterflies]
* [http://www.nature-pictures.org/en/Butterfly-photos/ European butterfly pictures - common names and wildlife photography]
* [http://www.rkwalton.com/skip.html Online videos of Skippers of the Northeast-USA]
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[[श्रेणी:Butterflies| ]]
[[श्रेणी:Lepidoptera]]
[[श्रेणी:Pollinators]]
 
[[श्रेणी:पुतली]]
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