Butterflies are characterized by their scale-covered wings.The coloration of butterfly wings is created by minute scales. These scales are pigmented with melanins 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 crystalnature of the scales. [Mason C. W., 1927 Structural colors in insects. II. Iridescent colors. J. Phys. Chem., 31, 321-354] [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] ] [Prum, Richard O., Tim Quinn and Rodolfo H. Torres 2006. Anatomically diverse butterfly scales all produce structural colours by coherent scattering. Journal of Experimental Biology 209, 748-765 doi: 10.1242/jeb.02051 [http://jeb.biologists.org/cgi/content/full/209/4/748 Full text] ]
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 dimorphismin the patterns of ultraviolet reflectivity, while otherwise appearing identical to the unaided human eye. Most of the butterflies have a sex-determination systemthat is represented as ZW with females being the heterogametic sex (ZW) and males homogametic (ZZ). [Traut W, Marec F (1997) Sex chromosome differentiation in some species of Lepidoptera (Insecta). Chromosome Research 5: 283-291. ]
Genetic abnormalities such as
gynandromorphy 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 [Rousset, F., D. Bouchon, B. Pintureau, P. Juchault, and M. Solignac. 1992. Wolbachia endosymbionts responsible for various alterations of sexuality in arthropods. Proceedings of the Royal Society of London B, Biological Sciences 250:91 – 98.] or the selective killing of males in the egg stage. [Jiggins, Francis M, Gregory D D Hurst, J Hinrich G V D Schulenburg and Michael E N Majerus (2001) Two male-killing Wolbachia strains coexist within a population of the butterfly "Acraea encedon". Heredity. 86(2):161-166]
Mullerian mimicryin butterflies is common. Batesian mimics imitate other species to enjoy the protection of an attribute they do not share, aposematismin this case. The Common Mormon of India has female morphs which imitate the unpalatable red-bodied swallowtails, the Common Rose and the Crimson Rose. Mullerian mimicry occurs when aposematic species evolve to resemble each other, presumably to reduce predator sampling rates, the Heliconiusbutterflies from the Americas being a good example.
Wing markings called eyespots are present in some species; these may have an
automimicryrole 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.  [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 ]
easonal polyphenismMany 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. [Brakefield, Paul M., Fanja Kesbeke, P. Bernhardt Koch (1998) The Regulation of Phenotypic Plasticity of Eyespots in the Butterfly "Bicyclus anynana". The American Naturalist. 152(6):853-860] 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. [Nijhout, H. F. (2003) Development and evolution of adaptive polyphenisms. Evolution and Development 5(1):9-18.] 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. [Brakefield, P. M. and Torben Larsen (1984) The evolutionary significance of dry and wet season forms in some tropical butterflies. Biological Journal of the Linnean Society 22: 1-12 [https://openaccess.leidenuniv.nl/bitstream/1887/11011/1/029_009.pdf Full text] ]
Butterflies feed primarily on
nectarfrom flowers. Some also derive nourishment from pollen, [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] tree sap, rotting fruit, dung, and dissolved minerals in wet sand or dirt. Butterflies play an important ecological role as pollinators.Fact|date=October 2008
As adults, butterflies consume only liquids and these are sucked by means of their
proboscis. They feed on nectarfrom 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 sodiumthan provided by nectar. They are attracted to sodium in saltand 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-puddlingbehaviour 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. [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]
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
tasteis coordinated by chemoreceptors on the 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. [ [http://www.sandiegozoo.org/animalbytes/t-butterfly.html Article on San Diego Zoo website] ] 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. [Obara Y, Hidaki T. (1968). Recognition of the female by the male, on the basis of ultra-violet reflection, in the white cabbage butterfly "Pieris rapae crucivora" Boisduval. Proc. Japan Acad., 44: 829-832.] Color vision may be widespread but has been demonstrated in only a few species. [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] ] [Michiyo Kinoshita, Naoko Shimada And Kentaro Arikawa (1999) Color vision of the foraging swallowtail butterfly "Papilioxuthus". The Journal of Experimental Biology 202:95 – 102 [http://jeb.biologists.org/cgi/reprint/202/2/95.pdf] ]
Some butterflies have organs of hearing and some species are also known to make stridulatory and clicking sounds. [Swihart, S. L (1967). Hearing in butterflies. J. Insect Physiol 13, 469]
Many butterflies, such as the
Monarch butterfly, are 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. [Reppert, Steven M.; Haisun Zhu; White, Richard H. (2004) Polarized light helps monarch butterflies navigate. Current biology 14(2):155-158]
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. [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] ]
Like many other members of the
insectworld, 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. [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 PDF] ] (See also Insect flight)
Insect migration"Many butterflies migrate over long distances. Particularly famous migrations being those of the Monarch butterflyfrom Mexicoto North America, a distance of about 4,000 to 4,800 kilometres (2500-3000 miles). Other well known migratory species include the Painted Lady and several of the Danaine butterflies. Spectacular and large scale migrations associated with the Monsoonsare seen in peninsular India. [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] Migrations have been studied in more recent times using wing tags and also using stable hydrogen isotopes. [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] [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] ]
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 particular important. [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] ]
It is suggested that most migratory butterflies are those that belong to semi-arid areas where breeding seasons are short. [Southwood, T. R. E. 1962. Migration of terrestrial arthropods in relation to habitat. Biol. Rev. 37:171-214] The life-histories of their host plants also influence the strategies of the butterflies. [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]
Butterflies are threatened in their early stages by
parasitoids and in all stages by predators, 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 protection against herbivores. Butterflies have evolved mechanisms to
sequesterthese plant toxins and use them instead in their own defense. [Nishida, Ritsuo (2002). Sequestration of defensive substances from plants by Lepidoptera. Annu. Rev. Entomol. 47:57–92] 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 mimicked by other butterflies. These mimetic forms are usually restricted to the females.Cryptic coloration is found in many butterflies. Some like the oakleaf butterfly are remarkable imitations of leaves. [Robbins, Robert K. (1981) The "False Head" Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies. American Naturalist 118(5):770-775] 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. [Nafus, D. M. and I. H. Schreiner (1988) Parental care in a tropical nymphalid butterfly "Hypolimas anomala". Anim. Behav. 36: 1425- 143]
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 spiders to approach from the wrong end and allow for early visual detection. [William E. Cooper, Jr. (1998) Conditions favoring anticipatory and reactive displays deflecting predatory attack. Behavioral Ecology]
There are between 15,000 and 20,000
speciesof butterflies worldwide. Some well known species from around the world include:
* Swallowtails and
Birdwings, Family Papilionidae
** Common Yellow Swallowtail, "Papilio machaon"
Spicebush Swallowtail, "Papilio troilus"
** Lime Butterfly, "Papilio demoleus"
** "Ornithoptera" genus (Birdwings; the largest butterflies)
* Whites or Yellows, Family Pieridae
Small White, "Pieris rapae"
Green-veined White, "Pieris napi"
** Common Jezebel, "Delias eucharis"
* Blues and Coppers or Gossamer-Winged Butterflies, Family Lycaenidae
Xerces Blue, "Glaucopsyche xerces" (extinct)
Karner Blue, "Lycaeides melissa samuelis" (endangered)
** Red Pierrot, "Talicada nyseus"
* Metalmark butterflies, Family Riodinidae
** Lange's Metalmark Butterfly
** Plum Judy, "Abisara echerius"
* Brush-footed butterflies, Family Nymphalidae
** Painted Lady, or Cosmopolite, "Vanessa cardui"
Monarch butterfly, "Danaus plexippus"
** "Morpho" genus
Speckled Wood, "Pararge aegeria"
Artistic depictions of butterflies have been used in many cultures including Egyptian hieroglyphics 3500 years ago. [Larsen, Torben (1994) Butterflies of Egypt. Saudi Aramco world. 45(5):24-27 [http://www.saudiaramcoworld.com/issue/199405/butterflies.of.egypt.htm Online] ] Today, butterflies are widely used in various objects of art, and have inspired the "butterfly
fairy" as an art and fictional character.
According to the “Butterflies” chapter in "", by
Lafcadio Hearn, a butterfly is seen as the personificationof 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 bambooscreen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad omens. When Taira no Masakadowas secretly preparing for his famous revolt, there appeared in Kyotoso vast a swarm of butterflies that the people were frightened — -thinking the apparition to be a portent of coming evil. [cite book | last = Hearn | first = Lafcadio | authorlink = Lafcadio Hearn | year = 1904 | title = Kwaidan: Stories and Studies of Strange Thing | publisher = Dover Publications, Inc. | id = ISBN 0-486-21901-1]
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 ψύχη ("psȳchē"), which primarily means "
soul", " mind". [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.]
According to Mircea Eliade's "Encyclopedia of Religion", some of the Nagas of
Manipurtrace their ancestry from a butterfly.Rabuzzi, M. 1997. Butterfly etymology. Cultural Entomology November 1997 Fourth issue [http://www.insects.org/ced4/etymology.html online] ]
Chinese culturetwo butterflies flying together are a symbol of love. Also a famous Chinese folk story called Butterfly Lovers. The Taoistphilosopher Zhuangzionce had a dream of being a butterfly flying without care about humanity, however when he woke up and realised 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 rebirth into a new life after being inside a cocoon for a period of time.
Some people say that when a butterfly lands on you it means good luck.
Devonshire, people would traditionally rush around to kill the first butterfly of the year that they see, or else face a year of bad luck. [ 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.]
The idiom "
butterflies in the stomach" is used to describe a state of nervousness.
Studies on the reflection of light by the scales on wings of swallowtail butterflies have to led to the innovation of more efficient
light-emitting diodes. [Vukusic, Pete and Ian Hooper. 2005. Directionally Controlled Fluorescence Emission in Butterflies Science. 310(5751):1151 DOI: 10.1126/science.1116612]
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. [ [http://www.qualcomm.com/technology/imod/index.html Biomimetics at Qualcomm] ]
Papilionidae- The Swallowtails
Pieridae- The Whites and Yellows
Riodinidae- The Metalmarks, Punches and Judies
Nymphalidae- The Brush-footed Butterflies
Lycaenidae- The Blues
Hesperiidae- The Skippers
List of British butterflies
* List of official state butterflies for each state of the U.S.A.
List of butterflies of India
List of Butterflies of North America
Butterflies of Taiwan
* McGuire Center for Lepidoptera and Biodiversity,
University of Florida
* Boggs, C., Watt, W., Ehrlich, P. 2003. Butterflies: Evolution and Ecology Taking Flight. University of Chicago Press, Chicago, USA.
* Heppner, J. B. 1998. Classification of Lepidoptera. "Holarctic Lepidoptera", Suppl. 1.
* Pyle, R. M. 1992. Handbook for Butterfly Watchers. Houghton Mifflin. First published, 1984. ISBN 0-395-61629-8
* 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)]
Field guides to butterflies
* "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)
* "Field Guide to Butterlies of South Africa" by Steve Woodhall (2005)
* "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)
* [http://tolweb.org/Papilionoidea/12027 Papilionoidea on the Tree of Life Web project]
* [http://creatures.ifas.ufl.edu/main/search_common.htm#bfly Butterflies] on the
UF/ IFAS Featured Creatures Web site
* [http://www.pinocchio.it/eng/butterflyhouse/ Collodi Butterfly House] Tuscany
* [http://www.butterfliesandmoths.org/ Butterflies and Moths of North America]
* [http://www.naba.org North American Butterfly Association (NABA)]
* [http://www.leps.nl/ Butterflies and Moths in the Netherlands]
* [http://www.wwfpak.org/nap/dnap_wildlife_wildfauna_mammals_insect.php Insect and butterfly diversity of Pakistan]
* [http://www.naturemagics.com/butterfly.shtm Butterflies of Southern India]
* [http://www.srilankaninsects.net/Butterflies/MainPage/ButterfliesMain.htm Butterflies of Sri Lanka]
* [http://www.geocities.com/RainForest/Vines/2382/sgchecklist.htm Butterflies of Singapore]
* [http://www.nature-of-oz.com/israelsspecieslist1.htm Israel Insect World]
* [http://www.geocities.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.malaeng.com/blog/?page_id=1293 Butterflies of Thailand]
* [http://www.mariposasmexicanas.com/ Butterflies of Mexico]
* [http://bugguide.net/node/view/81/bgimage BugGuide.net Many images of North American butterflies, many licensed via Creative Commons]
* [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.butterfliesoffrance.com/ Butterflies of France]
* [http://tolweb.org/movies/Papilionoidea/12027 Butterfly Movies (Tree of Life)]
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