Butterflies arewinged insects from thelepidopteransuperfamilyPapilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossils have been dated to thePaleocene, about 56 million years ago, though molecular evidence suggests that they likely originated in theCretaceous.[1]
Butterflies have a four-stagelife cycle, and like otherholometabolous insects they undergocomplete metamorphosis.[2] Winged adults lay eggs on the food plant on which theirlarvae, known ascaterpillars, will feed. The caterpillars grow, sometimes very rapidly, and when fully developed,pupate in achrysalis. Whenmetamorphosis is complete, the pupal skin splits, the adult insect climbs out, expands its wings to dry, and flies off.
Some butterflies, especially in the tropics, have several generations in a year, while others have a single generation, and a few in cold locations may take several years to pass through their entire life cycle.[3]
Butterflies are oftenpolymorphic, and many species make use ofcamouflage,mimicry, andaposematism to evade their predators.[4] Some, like themonarch and thepainted lady,migrate over long distances. Many butterflies are attacked byparasites orparasitoids, includingwasps,protozoans,flies, and other invertebrates, or arepreyed upon by other organisms. Some species are pests because in their larval stages they can damage domestic crops or trees; other species are agents ofpollination of some plants. Larvae of a few butterflies (e.g.,harvesters) eat harmful insects, and a few are predators ofants, while others live asmutualists in association with ants. Culturally, butterflies are a popular motif in the visual and literary arts. TheSmithsonian Institution says "butterflies are certainly one of the most appealing creatures in nature".[5]
Etymology
Possibly the original butter-fly.[6] A male brimstone (Gonepteryx rhamni) in flight.
TheOxford English Dictionary derives the word straightforwardly fromOld Englishbutorflēoge, butter-fly; similar names inOld Dutch andOld High German show that the name is ancient, but modern Dutch and German use different words (vlinder andSchmetterling) and the common name often varies substantially between otherwise closely related languages. A possible source of the name is the bright yellow male of the brimstone (Gonepteryx rhamni); another is that butterflies were on the wing in meadows during the spring and summer butter season while the grass was growing.[6][7]
Often have metallic spots on wings; often conspicuously coloured with black, orange and blue
Traditionally, butterflies have been divided into the superfamiliesPapilionoidea and the moth-likeHedyloidea. Recent work has discovered that Hedylidae, the only family within Hedyloidea, is nested within the Papilionoidea, meaning that Papilionoidea would besynonymous with Rhopalocera. The relationships between the rest of the 6 families are extremely well resolved, which is summarized in the below cladogram.[1][14][15]
A zoomed in view of the wing scales on aAglais io, or peacock butterfly.
Butterfly adults are characterized by their four scale-covered wings, which give the Lepidoptera their name (Ancient Greek λεπίς lepís, scale + πτερόν pterón, wing). These scales give butterfly wings their colour: they are pigmented withmelanins that give them blacks and browns, as well asuric acid derivatives andflavones that give them yellows, but many of the blues, greens, reds andiridescent colours are created bystructural coloration produced by the micro-structures of the scales and hairs.[16][17][18][19]
Butterfly antennal shapes, mainly clubbed, unlike those of moths. Drawn by C. T. Bingham, 1905
As in all insects, the body is divided into three sections: the head,thorax, andabdomen. The thorax is composed of three segments, each with a pair of legs. In most families of butterfly the antennae are clubbed, unlike those ofmoths which may be threadlike or feathery. The long proboscis can be coiled when not in use for sipping nectar from flowers.[20]
Unlike butterflies, most moths (likeLaothoe populi) fly by night and hide by day.
Nearly all butterflies arediurnal, have relatively bright colours, and hold their wings vertically above their bodies when at rest, unlike the majority of moths which fly by night, are oftencryptically coloured (well camouflaged), and either hold their wings flat (touching the surface on which the moth is standing) or fold them closely over their bodies. Some day-flying moths, such as thehummingbird hawk-moth,[21] are exceptions to these rules.[20][22]
Butterflylarvae,caterpillars, have a hard (sclerotised) head with strong mandibles used for cutting their food, most often leaves. They have cylindrical bodies, with ten segments to the abdomen, generally with short prolegs on segments 3–6 and 10; the three pairs of true legs on the thorax have five segments each.[20] Many are well camouflaged; others are aposematic with bright colours and bristly projections containing toxic chemicals obtained from their food plants. Thepupa or chrysalis, unlike that of moths, is not wrapped in a cocoon.[20]
Butterflies are distributed worldwide except Antarctica, totalling some 18,500 species.[24] Of these, 775 areNearctic; 7,700Neotropical; 1,575Palearctic; 3,650Afrotropical; and 4,800 are distributed across the combinedOriental andAustralian/Oceania regions.[24] Themonarch butterfly is native to the Americas, but in the nineteenth century or before, spread across the world, and is now found in Australia, New Zealand, other parts of Oceania, and theIberian Peninsula. It is not clear how it dispersed; adults may have been blown by the wind or larvae or pupae may have been accidentally transported by humans, but the presence of suitable host plants in their new environment was a necessity for their successful establishment.[25]
Monarch migration routeOverwintering monarchs cluster onoyamel trees nearAngangueo, Mexico.
Many butterflies, such as thepainted lady, monarch, and severaldanaine migrate for long distances. These migrations take place over a number of generations and no single individual completes the whole trip. The eastern North American population of monarchs can travel thousands of miles south-west tooverwintering sites in Mexico. There is a reverse migration in the spring.[26][27] It has recently been shown that the British painted lady undertakes a 9,000-mile round trip in a series of steps by up to six successive generations, from tropical Africa to the Arctic Circle — almost double the length of the famous migrations undertaken by monarch.[28] Spectacular large-scale migrations associated with themonsoon are seen in peninsular India.[29] Migrations have been studied in more recent times using wing tags and also usingstable hydrogen isotopes.[30][31]
Butterflies navigate using a time-compensated sun compass. They can see polarized light and therefore orient even in cloudy conditions. The polarized light near the ultraviolet spectrum appears to be particularly important.[32][33] Many migratory butterflies live in semi-arid areas where breeding seasons are short.[34] The life histories of their host plants also influence butterfly behaviour.[35]
Life cycle
Life cycle of the monarch butterfly
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 remaindormant in their pupal or egg stages and thereby survive winters.[36] TheMelissa Arctic (Oeneis melissa) overwinters twice as a caterpillar.[37] Butterflies may have one or more broods per year. The number of generations per year varies fromtemperate totropical regions with tropical regions showing a trend towardsmultivoltinism.[38]
Courtship is often aerial and often involvespheromones. Butterflies then land on the ground or on a perch to mate.[20] Copulation takes place tail-to-tail and may last from minutes to hours. Simple photoreceptor cells located at the genitals are important for this and other adult behaviours.[39] The male passes aspermatophore to the female; to reduce sperm competition, he may cover her with his scent, or in some species such as the Apollos (Parnassius)plugs her genital opening to prevent her from mating again.[40]
The vast majority of butterflies have a four-stage life cycle:egg,larva (caterpillar),pupa (chrysalis) andimago (adult). In the generaColias,Erebia,Euchloe, andParnassius, a small number of species are known that reproducesemi-parthenogenetically; when the female dies, a partially developed larva emerges from her abdomen.[41]
Eggs
Eggs of black-veined white (Aporia crataegi) onapple leafA butterfly from the genusEuploea, laying eggs underneath the leaf
Butterfly eggs are protected by a hard-ridged outer layer of shell, called thechorion. This is lined with a thin coating of wax which prevents the egg from drying out before the larva has had time to fully develop. Each egg contains a number of tiny funnel-shaped openings at one end, calledmicropyles; the purpose of these holes is to allow sperm to enter and fertilize the egg. Butterfly eggs vary greatly in size and shape between species, but are usually upright and finely sculptured. Some species lay eggs singly, others in batches. Many females produce between one hundred and two hundred eggs.[41]
Butterfly eggs are fixed to a leaf with a special glue which hardens rapidly. As it hardens it contracts, deforming the shape of the egg. This glue is easily seen surrounding the base of every egg forming a meniscus. The nature of the glue has been little researched but in the case ofPieris brassicae, it begins as a pale yellow granular secretion containing acidophilic proteins. This is viscous and darkens when exposed to air, becoming a water-insoluble, rubbery material which soon sets solid.[42] Butterflies in the genusAgathymus do not fix their eggs to a leaf; instead, the newly laid eggs fall to the base of the plant.[43]
Eggs are almost invariably laid on plants. Each species of butterfly has its own host plant range and while some species of butterfly are restricted to just one species of plant, others use a range of plant species, often including members of a common family.[44] In some species, such as thegreat spangled fritillary, the eggs are deposited close to but not on the food plant. This most likely happens when the egg overwinters before hatching and where the host plant loses its leaves in winter, as doviolets in this example.[45]
The egg stage lasts a few weeks in most butterflies, but eggs laid close to winter, especially in temperate regions, go through adiapause (resting) stage, and the hatching may take place only in spring.[46] Some temperate region butterflies, such as theCamberwell beauty, lay their eggs in the spring and have them hatch in the summer.[47]
Butterfly larvae, or caterpillars, consume plant leaves and spend practically all of their time searching for and eating food. Although most caterpillars are herbivorous, a few species arepredators:Spalgis epius eatsscale insects,[48] while lycaenids such asLiphyra brassolis aremyrmecophilous, eating ant larvae.[49]
Some larvae, especially those of theLycaenidae, formmutual associations with ants. They communicate with the ants using vibrations that are transmitted through thesubstrate as well as using chemical signals.[50][51] The ants provide some degree of protection to these larvae and they in turn gatherhoneydew secretions.Large blue (Phengaris arion) caterpillars trickMyrmica ants into taking them back to theant colony where they feed on the ant eggs and larvae in a parasitic relationship.[52]
Caterpillars mature through a series of developmental stages known asinstars. Near the end of each stage, the larva undergoes a process calledapolysis, mediated by the release of a series ofneurohormones. During this phase, thecuticle, a tough outer layer made of a mixture ofchitin and specializedproteins, is released from the softerepidermis beneath, and the epidermis begins to form a new cuticle. At the end of each instar, the larvamoults, the old cuticle splits and the new cuticle expands, rapidly hardening and developing pigment.[53]
Caterpillars have short antennae and severalsimple eyes. Themouthparts are adapted for chewing with powerful mandibles and a pair of maxillae, each with a segmented palp. Adjoining these is the labium-hypopharynx which houses a tubular spinneret which is able to extrude silk.[16] Caterpillars such as those in the genusCalpodes (family Hesperiidae) have a specialized tracheal system on the 8th segment that function as a primitive lung.[54] Butterfly caterpillars have three pairs of true legs on the thoracic segments and up to six pairs ofprolegs arising from the abdominal segments. These prolegs have rings of tiny hooks called crochets that are engaged hydrostatically and help the caterpillar grip the substrate.[55] The epidermis bears tufts ofsetae, the position and number of which help in identifying the species. There is also decoration in the form of hairs, wart-like protuberances, horn-like protuberances and spines. Internally, most of the body cavity is taken up by the gut, but there may also be large silk glands, and special glands which secrete distasteful or toxic substances. The developing wings are present in later stage instars and thegonads start development in the egg stage.[16]
When the larva is fully grown, hormones such asprothoracicotropic hormone (PTTH) are produced. At this point the larva stops feeding, and begins "wandering" in the quest for a suitable pupation site, often the underside of a leaf or other concealed location. There it spins a button of silk which it uses to fasten its body to the surface and moults for a final time. While some caterpillars spin acocoon to protect the pupa, most species do not. The naked pupa, often known as a chrysalis, usually hangs head down from the cremaster, a spiny pad at the posterior end, but in some species a silken girdle may be spun to keep the pupa in a head-up position.[41] Most of the tissues and cells of the larva are broken down inside the pupa, as the constituent material is rebuilt into the imago. The structure of the transforming insect is visible from the exterior, with the wings folded flat on the ventral surface and the two halves of the proboscis, with the antennae and the legs between them.[16]
The pupal transformation into a butterfly throughmetamorphosis 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 colour pattern are marked by changes in the expression of particular transcription factors in the early pupa.[56]
The reproductive stage of the insect is the winged adult orimago. The surface of both butterflies and moths is covered by scales, each of which is an outgrowth from a singleepidermal cell. The head is small and dominated by the two largecompound eyes. These are capable of distinguishing flower shapes or motion but cannot view distant objects clearly. Colour perception is good, especially in some species in the blue/violet range. Theantennae are composed of many segments and have clubbed tips (unlike moths that have tapering or feathery antennae). The sensory receptors are concentrated in the tips and can detect odours. Taste receptors are located on the palps and on the feet. The mouthparts are adapted to sucking and themandibles are usually reduced in size or absent. The first maxillae are elongated into a tubularproboscis which is curled up at rest and expanded when needed to feed. The first and second maxillae bear palps which function as sensory organs. Some species have a reduced proboscis or maxillary palps and do not feed as adults.[16]
Butterfly resting on flower at Magic Wings Conservatory
ManyHeliconius butterflies also use their proboscis to feed on pollen;[57] in these species only 20% of the amino acids used in reproduction come from larval feeding, which allow them to develop more quickly as caterpillars, and gives them a longer lifespan of several months as adults.[58]
The thorax of the butterfly is devoted to locomotion. Each of the three thoracic segments has two legs (amongnymphalids, the first pair is reduced and the insects walk on four legs). The second and third segments of the thorax bear the wings. The leading edges of the forewings have thick veins to strengthen them, and the hindwings are smaller and more rounded and have fewer stiffening veins. The forewings and hindwings are not hooked together (as they are in moths) but are coordinated by the friction of their overlapping parts. The front two segments have a pair ofspiracles which are used in respiration.[16]
The abdomen consists of ten segments and contains the gut and genital organs. The front eight segments have spiracles and the terminal segment is modified for reproduction. The male has a pair of clasping organs attached to a ring structure, and during copulation, a tubular structure is extruded and inserted into the female's vagina. Aspermatophore is deposited in the female, following which the sperm make their way to a seminal receptacle where they are stored for later use. In both sexes, the genitalia are adorned with various spines, teeth, scales and bristles, which act to prevent the butterfly from mating with an insect of another species.[16] 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 withhemolymph and letting them dry, during which time it is extremely vulnerable to predators.[59]
Pattern formation
The colourful patterns on many butterfly wings tell potential predators that they are toxic. Hence, the genetic basis of wingpattern formation can illuminate both theevolution of butterflies as well as theirdevelopmental biology. The colour of butterfly wings is derived from tiny structures called scales, each of which have their ownpigments. InHeliconius butterflies, there are three types of scales: yellow/white, black, and red/orange/brown scales. Some mechanism of wing pattern formation are now being solved using genetic techniques. For instance, agene calledcortex determines the colour of scales: deletingcortex turned black and red scales yellow. Mutations, e.g.transposon insertions of thenon-coding DNA around thecortex gene can turn a black-winged butterfly into a butterfly with a yellow wing band.[60]
Mating
When the butterflyBicyclus anynana is subjected to repeated inbreeding in the laboratory, there is a dramatic decrease in egg hatching.[61] This severeinbreeding depression is considered to be likely due to a relatively highmutation rate to recessivealleles with substantial damaging effects and infrequent episodes ofinbreeding in nature that might otherwise purge such mutations.[61] AlthoughB. anynana experiences inbreeding depression when forcibly inbred in the laboratory it recovers within a few generation when allowed to breed freely.[62] During mate selection, adult females do not innately avoid or learn to avoid siblings, implying that such detection may not be critical to reproductive fitness.[62] Inbreeding may persist inB anynana because the probability of encountering close relatives is rare in nature; that is, movement ecology may mask the deleterious effect of inbreeding resulting in relaxation of selection for active inbreeding avoidance behaviors.
Butterflies feed primarily onnectar from flowers. Some also derive nourishment frompollen,[63] tree sap, rotting fruit, dung, decaying flesh, and dissolved minerals in wet sand or dirt. Butterflies are important as pollinators for some species of plants. In general, they do not carry as much pollen load asbees, but they are capable of moving pollen over greater distances.[64]Flower constancy has been observed for at least one species of butterfly.[65]
Adult butterflies consume only liquids, ingested through the proboscis. They sip water from damp patches for hydration and feed on nectar from flowers, from which they obtain sugars for energy, andsodium and other minerals vital for reproduction. Several species of butterflies need more sodium than that provided by nectar and are attracted by sodium in salt; they sometimes land on people, attracted by the salt in human sweat. Some butterflies also visit dung and scavenge rotting fruit or carcasses to obtain minerals and nutrients. In many species, thismud-puddling behaviour is restricted to the males, and studies have suggested that the nutrients collected may be provided as anuptial gift, along with the spermatophore, during mating.[66]
Inhilltopping, males of some species seek hilltops and ridge tops, which they patrol in search for females. Since it usually occurs in species with low population density, it is assumed these landscape points are used as meeting places to find mates.[67]
Butterflies use their antennae to sense the air for wind and scents. The antennae come in various shapes and colours; the hesperiids have a pointed angle or hook to the antennae, while most other families show knobbed antennae. The antennae are richly covered with sensory organs known assensillae. A butterfly's sense of taste is coordinated by chemoreceptors on thetarsi, or feet, 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.[68] Many butterflies use chemical signals,pheromones; some have specialized scent scales (androconia) or other structures (coremata or "hair pencils" in the Danaidae).[69] 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.[70] Colour vision may be widespread but has been demonstrated in only a few species.[71][72] Some butterflies have organs of hearing and some species makestridulatory and clicking sounds.[73]
Heteronympha merope taking off
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. Butterflies can only fly when their temperature is above 27 °C (81 °F); when it is cool, they can position themselves to expose the underside of the wings to the sunlight to heat themselves up. If their body temperature reaches 40 °C (104 °F), they can orientate themselves with the folded wings edgewise to the sun.[74] Basking is an activity which is more common in the cooler hours of the morning. Some species have evolved dark wingbases to help in gathering more heat and this is especially evident in alpine forms.[75]
As in many other insects, thelift generated by butterflies is more than can be accounted for by steady-state, non-transitoryaerodynamics. Studies usingVanessa atalanta in awind tunnel show that they use a wide variety of aerodynamic mechanisms to generate force. These includewake capture,vortices at the wing edge, rotational mechanisms and theWeis-Fogh 'clap-and-fling' mechanism. Butterflies are able to change from one mode to another rapidly.[76]
Butterflies are threatened in their early stages byparasitoids and in all stages by predators, diseases and environmental factors.Braconid and other parasitic wasps lay their eggs in lepidopteran eggs or larvae and the wasps' parasitoid larvae devour their hosts, usually pupating inside or outside the desiccated husk. Most wasps are very specific about their host species and some have been used as biological controls of pest butterflies like thelarge white butterfly.[77] When thesmall cabbage white was accidentally introduced to New Zealand, it had no natural enemies. In order to control it, some pupae that had been parasitised by a chalcid wasp were imported, and natural control was thus regained.[78] Some flies lay their eggs on the outside of caterpillars and the newly hatched fly larvae bore their way through the skin and feed in a similar way to the parasitoid wasp larvae.[79] Predators of butterflies include ants, spiders, wasps, and birds.[80]
Caterpillars are also affected by a range of bacterial, viral and fungal diseases, and only a small percentage of the butterfly eggs laid ever reach adulthood.[79] The bacteriumBacillus thuringiensis has been used in sprays to reduce damage to crops by the caterpillars of the large white butterfly, and theentomopathogenic fungusBeauveria bassiana has proved effective for the same purpose.[81]
Butterflies protect themselves from predators by a variety of means.
Chemical defences are widespread and are mostly based on chemicals of plant origin. In many cases the plants themselves evolved these toxic substances asprotection against herbivores. Butterflies have evolved mechanisms to sequester these plant toxins and use them instead in their own defence.[83] These defence mechanisms are effective only if they are well advertised; this has led to the evolution of bright colours in unpalatable butterflies (aposematism). This signal is commonlymimicked by other butterflies, usually only females. ABatesian mimic imitates another species to enjoy the protection of that species' aposematism.[84] Thecommon Mormon of India has female morphs which imitate the unpalatable red-bodied swallowtails, thecommon rose and thecrimson rose.[85]Müllerian mimicry occurs when aposematic species evolve to resemble each other, presumably to reduce predator sampling rates;Heliconius butterflies from the Americas are a good example.[84]
Camouflage is found in many butterflies. Some like the oakleaf butterfly andautumn leaf are remarkable imitations of leaves.[86] As caterpillars, many defend themselves by freezing and appearing like sticks or branches.[87] Others havedeimatic behaviours, such as rearing up and waving their front ends which are marked with eyespots as if they were snakes.[88] Some papilionid caterpillars such as the giant swallowtail (Papilio cresphontes) resemble bird droppings so as to be passed over by predators.[89] Some caterpillars have hairs and bristly structures that provide protection while others are gregarious and form dense aggregations.[84] Some species aremyrmecophiles, formingmutualistic associations withants and gaining their protection.[90] Behavioural defences include perching and angling the wings to reduce shadow and avoid being conspicuous. Some femaleNymphalid butterflies guard their eggs from parasitoidalwasps.[91]
The Lycaenidae have a false head consisting of eyespots and small tails (false antennae) to deflect attack from the more vital head region. These may also cause ambush predators such as spiders to approach from the wrong end, enabling the butterflies to detect attacks promptly.[92][93] Many butterflies haveeyespots on the wings; these too may deflect attacks, or may serve to attract mates.[56][94]
Auditory defences can also be used, which in the case of thegrizzled skipper refers to vibrations generated by the butterfly upon expanding its wings in an attempt to communicate with ant predators.[95]
Many tropical butterflies haveseasonal forms for dry and wet seasons.[96][97] These are switched by the hormoneecdysone.[98] The dry-season forms are usually more cryptic, perhaps offering better camouflage when vegetation is scarce. Dark colours in wet-season forms may help to absorb solar radiation.[99][100][94]
Butterflies without defences such as toxins or mimicry protect themselves through a flight that is more bumpy and unpredictable than in other species. It is assumed this behavior makes it more difficult for predators to catch them, and is caused by theturbulence created by the small whirlpools formed by the wings during flight.[101]
Declining butterfly populations have been noticed in many areas of the world, and this phenomenon is consistent with therapidly decreasing insect populations around the world. At least in the Western United States, this collapse in the number of most species of butterflies has been determined to be driven byglobal climate change, specifically, by warmer autumns.[103][104] Butterfly populations in the United States declined by 22% between 2000 and 2020, mainly because of habitat loss, pesticides and climate change.[105]
Butterflies have appeared in art from 3500 years ago inancient Egypt.[106] In hunting scenes, butterflies were sometimes included in a way that suggested life, freedom, and the strength to escape capture, creating a balance to scenes concerned with death and upholdingma'at. They also were suggestive of regeneration or rebirth and protection. Certain butterflies, such as thetiger butterfly, may have been associated with solar deities, particularlyRa. The tiger butterfly also would have a particular resemblance to theankh, due to its black body and wingtips, that was likely noted by the Ancient Egyptians. Butterflies may also have been understood as one of the deceased's guides in the afterlife.[107]
In the ancientMesoamerican city ofTeotihuacan, the brilliantly coloured image of the butterfly was carved into many temples, buildings, jewellery, and emblazoned onincense burners. The butterfly was sometimes depicted with the maw of ajaguar, and some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies with fire and warfare persisted into theAztec civilisation; evidence of similar jaguar-butterfly images has been found among theZapotec andMaya civilisations.[108]
Butterflies are widely used in objects of art and jewellery: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and used in some mixed media artworks and furnishings.[109] TheNorwegiannaturalistKjell Sandved compiled a photographicButterfly Alphabet containing all 26 letters and the numerals 0 to 9 from the wings of butterflies.[110] The butterfly is a symbol of beingtransgender, because of the transformation from caterpillar to winged adult.[111]
According toLafcadio Hearn, a butterfly was seen in Japan 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 guest room and perches behind the bamboo screen, the person whom you most love is coming to see you. Large numbers of butterflies are viewed as badomens. WhenTaira no Masakado was secretly preparing his famous revolt, a vast a swarm of butterflies appeared inKyoto. The people were frightened, thinking the apparition to be a portent of coming evil.[115]
Diderot'sEncyclopédie cites butterflies as a symbol for the soul. A Roman sculpture depicts a butterfly exiting the mouth of a dead man, representing the Roman belief that the soul leaves through the mouth.[116] In line with this, the ancient Greek word for "butterfly" is ψυχή (psȳchē), which primarily means "soul" or "mind".[117] According toMircea Eliade, some of theNagas ofManipur claim ancestry from a butterfly.[118] In some cultures, butterflies symboliserebirth.[119] In the English county ofDevon, people once hurried to kill the first butterfly of the year, to avoid a year of bad luck.[120] In the Philippines, a lingering black or dark butterfly or moth in the house is taken to mean an impending or recent death in the family.[121] Several American states have chosen anofficial state butterfly.[122]
"Collecting" means preserving dead specimens, not keeping butterflies as pets.[123][124] Collecting butterflies was once a popular hobby; it has now largely been replaced by photography, recording, and rearing butterflies for release into the wild.[6][dubious –discuss][full citation needed] The zoological illustratorFrederick William Frohawk succeeded in rearing all the butterfly species found in Britain, at a rate of four per year, to enable him to draw every stage of each species. He published the results in the folio sized handbookThe Natural History of British Butterflies in 1924.[6]
Butterflies andmoths can be reared for recreation or for release.[125]
Study of thestructural coloration of the wing scales of swallowtail butterflies has led to the development of more efficientlight-emitting diodes,[126] and is inspiringnanotechnology research to produce paints that do not use toxic pigments and the development of new display technologies.[127]
^Mason, C. W. (1927). "Structural Colors in Insects. II".The Journal of Physical Chemistry.31 (3):321–354.doi:10.1021/j150273a001.
^Vukusic, P.; J. R. Sambles & H. Ghiradella (2000). "Optical Classification of Microstructure in Butterfly Wing-scales".Photonics Science News.6:61–66.
^abWilliams, Ernest; Adams, James; Snyder, John."Frequently Asked Questions". The Lepidopterists' Society. Archived fromthe original on 13 May 2015. Retrieved9 September 2015.
^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".Transactions of the Entomological Society of London.75:1–33.doi:10.1111/j.1365-2311.1927.tb00054.x.
^Urquhart, F. A.; Urquhart, N. R. (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. Entomol.109 (12):1583–1589.doi:10.4039/Ent1091583-12.S2CID86198255.
^Dennis, R L H; Shreeve, Tim G.; Arnold, Henry R.; Roy, David B. (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.Bibcode:2005JICon...9..187D.doi:10.1007/s10841-005-5660-x.S2CID20605146.
^Beament, J.W.L.; Lal, R. (1957). "Penetration Through the Egg-shell ofPieris brassicae".Bulletin of Entomological Research.48 (1):109–125.doi:10.1017/S0007485300054134.
^Devries, P. J. (1988). "The larval Ant-organs of Thisbe irenea (Lepidoptera: Riodinidae) and Their Effects Upon Attending Ants".Zoological Journal of the Linnean Society.94 (4):379–393.doi:10.1111/j.1096-3642.1988.tb01201.x.
^abSaccheri IJ, Brakefield PM, Nichols RA. "Severe Inbreeding Depression and Rapid Fitness Rebound in the ButterflyBicyclus anynana (Satyridae)".Evolution. 1996 Oct; 50 (5): 2000-2013. doi: 10.1111/j.1558-5646.1996.tb03587.x. PMID 28565613
^abRobertson DN, Sullivan TJ, Westerman EL. Lack of sibling avoidance during mate selection in the butterflyBicyclus anynana.Behavioural Processes. 2020 Apr; 173: 104062. doi: 10.1016/j.beproc.2020.104062. Epub 2020 Jan 22. PMID 31981681
^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?".Biological Journal of the Linnean Society.86 (3):345–361.doi:10.1111/j.1095-8312.2005.00539.x.
^Halloran, Kathryn; Wason, Elizabeth (2013)."Papilio polytes".Animal Diversity Web. University of Michigan Museum of Zoology.Archived from the original on 7 October 2015. Retrieved12 September 2015.
^Fiedler, K.; Holldobler, B.; Seufert, P. (1996). "Butterflies and Ants: The Communicative Domain".Cellular and Molecular Life Sciences.52:14–24.doi:10.1007/bf01922410.S2CID33081655.
^Nafus, D. M.; Schreiner, I. H. (1988). "Parental Care in a Tropical Nymphalid ButterflyHypolimas anomala".Animal Behaviour.36 (5):1425–1443.doi:10.1016/s0003-3472(88)80213-6.S2CID53183529.
^Cooper, William E. Jr. (1998). "Conditions Favoring Anticipatory and Reactive Displays Deflecting Predatory Attack".Behavioral Ecology.9 (6):598–604.CiteSeerX10.1.1.928.6688.doi:10.1093/beheco/9.6.598.
^Larsen, Torben (1994)."Butterflies of Egypt".Saudi Aramco World.45 (5):24–27.Archived from the original on 13 January 2010. Retrieved15 November 2006.
^"I'm Scared to Be a Woman". Human Rights Watch. 24 September 2014.Archived from the original on 6 September 2015. Retrieved8 September 2015.a 22-year-old transgender woman sports a tattoo of a butterfly—a transgender symbol signifying transformation
^Louis, Chevalier de Jaucourt (Biography) (January 2011)."Butterfly".Encyclopedia of Diderot and d'Alembert.Archived from the original on 11 August 2016. Retrieved1 April 2015.
^Hutchins, M., Arthur V. Evans, Rosser W. Garrison and Neil Schlager (Eds) (2003) Grzimek's Animal Life Encyclopedia, 2nd edition. Volume 3, Insects. Gale, 2003.
^Rabuzzi, M. 1997. Butterfly etymology. Cultural Entomology November 1997 Fourth issueonlineArchived 3 December 1998 at theWayback Machine
^Dorset Chronicle, May 1825, reprinted in:"The First Butterfly", inThe Every-day Book and Table Book; or, Everlasting Calendar of Popular Amusements, Etc. Vol III, ed. William Hone, (London: 1838) p 678.
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