“Butterfly”的意思、由来-开放百科全书

Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises the large superfamily Papilionoidea, which contains at least one former group, the skippers (formerly the superfamily "Hesperioidea"), and the most recent analyses suggest it also contains the moth-butterflies (formerly the superfamily "Hedyloidea"). Butterfly fossils date to the Paleocene, which was about 56 million years ago.

Butterflies have the typical four-stage insect life cycle. Winged adults lay eggs on the food plant on which their larvae, known as caterpillars, will feed. The caterpillars grow, sometimes very rapidly, and when fully developed, pupate in a chrysalis. When metamorphosis is complete, the pupal skin splits, the adult insect climbs out, and after its wings have expanded and dried, it 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.

Butterflies are often polymorphic, and many species make use of camouflage, mimicry and aposematism to evade their predators. Some, like the monarch and the painted lady, migrate over long distances. Many butterflies are attacked by parasites or parasitoids, including wasps, protozoans, flies, and other invertebrates, or are preyed upon by other organisms. Some species are pests because in their larval stages they can damage domestic crops or trees; other species are agents of pollination of some plants. Larvae of a few butterflies (e.g., harvesters) eat harmful insects, and a few are predators of ants, while others live as mutualists in association with ants. Culturally, butterflies are a popular motif in the visual and literary arts.

Etymology

The Oxford English Dictionary derives the word straightforwardly from Old English butorflēoge, butter-fly; similar names in Old Dutch and Old High German show that the name is ancient. 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.^[1]^[2]

Taxonomy and phylogeny

The earliest Lepidoptera fossils are of a small moth, Archaeolepis mane, of Jurassic age, around 190 million years ago (mya).^[3]^[4] Butterflies evolved from moths, so while the butterflies are monophyletic (forming a single clade), the moths are not. The oldest butterflies are from the Palaeocene MoClay or Fur Formation of Denmark, approximately 55 million years old. The oldest American butterfly is the Late Eocene Prodryas persephone from the Florissant Fossil Beds,^[5]^[6] approximately 34 million years old.^[7]

Traditionally, the butterflies have been divided into the superfamily Papilionoidea excluding the smaller groups of the Hesperiidae (skippers) and the more moth-like Hedylidae of America. Phylogenetic analysis suggests that the traditional Papilionoidea is paraphyletic with respect to the other two groups, so they should both be included within Papilionoidea, to form a single butterfly group, thereby synonymous with the clade Rhopalocera.^[8]^[9]

Biology

General description

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 with melanins that give them blacks and browns, as well as uric acid derivatives and flavones that give them yellows, but many of the blues, greens, reds and iridescent colours are created by structural coloration produced by the micro-structures of the scales and hairs.^[11]^[10]^[11]^[12]

As in all insects, the body is divided into three sections: the head, thorax, and abdomen. The thorax is composed of three segments, each with a pair of legs. In most families of butterfly the antennae are clubbed, unlike those of moths which may be threadlike or feathery. The long proboscis can be coiled when not in use for sipping nectar from flowers.^[13]

Nearly all butterflies are diurnal, 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 often cryptically 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 the hummingbird hawk-moth,^[14] are exceptions to these rules.^[13]^[15]

Butterfly larvae, 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.^[13] Many are well camouflaged; others are aposematic with bright colours and bristly projections containing toxic chemicals obtained from their food plants. The pupa or chrysalis, unlike that of moths, is not wrapped in a cocoon.^[13]

Many butterflies are sexually dimorphic. Most butterflies have the ZW sex-determination system where females are the heterogametic sex (ZW) and males homogametic (ZZ).^[16]

Distribution and migration

Butterflies are distributed worldwide except Antarctica, totalling some 18,500 species.^[22] Of these, 775 are Nearctic; 7,700 Neotropical; 1,575 Palearctic; 3,650 Afrotropical; and 4,800 are distributed across the combined Oriental and Australian/Oceania regions.^[17] The monarch 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 the Iberian 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.^[18]

Many butterflies, such as the painted lady, monarch, and several danaine 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 to overwintering sites in Mexico. There is a reverse migration in the spring.^[19]^[20] 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.^[21] Spectacular large-scale migrations associated with the monsoon are seen in peninsular India.^[22] Migrations have been studied in more recent times using wing tags and also using stable hydrogen isotopes.^[23]^[24]

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.^[25]^[26] Many migratory butterflies live in semi-arid areas where breeding seasons are short.^[27] The life histories of their host plants also influence butterfly behaviour.^[28]

Life cycle

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 dormant in their pupal or egg stages and thereby survive winters.^[29] The Melissa Arctic (Oeneis melissa) overwinters twice as a caterpillar.^[30] 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 multivoltinism.^[31]

The vast majority of butterflies have a four-stage life cycle; egg, larva (caterpillar), pupa (chrysalis) and imago (adult). In the genera Colias, Erebia, Euchloe, and Parnassius, a small number of species are known that reproduce semi-parthenogenetically; when the female dies, a partially developed larva emerges from her abdomen.^[41]

Egg

Butterfly eggs are protected by a hard-ridged outer layer of shell, called the chorion. 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, called micropyles; 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.^[34]

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 of Pieris 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.^[35] Butterflies in the genus Agathymus do not fix their eggs to a leaf, instead the newly laid eggs fall to the base of the plant.^[36]

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.^[37] In some species, such as the great 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 do violets in this example.^[38]

The egg stage lasts a few weeks in most butterflies, but eggs laid close to winter, especially in temperate regions, go through a diapause (resting) stage, and the hatching may take place only in spring.^[39] Some temperate region butterflies, such as the Camberwell beauty, lay their eggs in the spring and have them hatch in the summer.^[40]

Caterpillar larva

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 are predators: Spalgis epius eats scale insects,^[41] while lycaenids such as Liphyra brassolis are myrmecophilous, eating ant larvae.^[42]

Some larvae, especially those of the Lycaenidae, form mutual associations with ants. They communicate with the ants using vibrations that are transmitted through the substrate as well as using chemical signals.^[43]^[44] The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions. Large blue (Phengaris arion) caterpillars trick Myrmica ants into taking them back to the ant colony where they feed on the ant eggs and larvae in a parasitic relationship.^[45]

Caterpillars mature through a series of developmental stages known as instars. Near the end of each stage, the larva undergoes a process called apolysis, mediated by the release of a series of neurohormones. During this phase, the cuticle, a tough outer layer made of a mixture of chitin and specialized proteins, is released from the softer epidermis beneath, and the epidermis begins to form a new cuticle. At the end of each instar, the larva moults, the old cuticle splits and the new cuticle expands, rapidly hardening and developing pigment.^[46] Development of butterfly wing patterns begins by the last larval instar.

Caterpillars have short antennae and several simple eyes. The mouthparts are adapted for chewing with powerful mandibles and a pair maxillae, each with a segmented palp. Adjoining these is the labium-hypopharynx which houses a tubular spinneret which is able to extrude silk.^[11] Caterpillars such as those in the genus Calpodes (family Hesperiidae) have a specialized tracheal system on the 8th segment that function as a primitive lung.^[47] Butterfly caterpillars have three pairs of true legs on the thoracic segments and up to six pairs of prolegs 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.^[48] The epidermis bears tufts of setae, 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 the gonads start development in the egg stage.^[11]

Pupa

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 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 a cocoon 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.^[34] 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.^[49]

The pupal transformation into a butterfly through 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 colour pattern are marked by changes in the expression of particular transcription factors in the early pupa.^[50]

Adult

The reproductive stage of the insect is the winged adult or imago. The surface of both butterflies and moths is covered by scales, each of which is an outgrowth from a single epidermal cell. The head is small and dominated by the two large compound eyes. These are capable of distinguishing flower shapes or motion but not for clearly viewing distant objects. Colour perception is good, especially in some species in the blue/violet range. The antennae 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 the mandibles are usually reduced in size or absent. The first maxillae are elongated into a tubular proboscis 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.^[49]

Many Heliconius butterflies also use their proboscis to feed on pollen;^[51] 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.^[52]

The thorax of the butterfly is devoted to locomotion. Each of the three thoracic segments has two legs (among nymphalids, 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 of spiracles which are used in respiration.^[49]

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. A spermatophore 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.^[49] 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 hemolymph and letting them dry, during which time it is extremely vulnerable to predators.^[53]

Behaviour

Butterflies feed primarily on nectar from flowers. Some also derive nourishment from pollen,^[54] 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 as bees, but they are capable of moving pollen over greater distances.^[55] Flower constancy has been observed for at least one species of butterfly.^[56]

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, and sodium 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, 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 may be provided as a nuptial gift, along with the spermatophore, during mating.^[57]

In hilltopping, 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.^[58]

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 as sensillae. A butterfly's sense of taste is coordinated by chemoreceptors on the tarsi, 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.^[59] Many butterflies use chemical signals, pheromones; some have specialized scent scales (androconia) or other structures (coremata or "hair pencils" in the Danaidae).^[60] 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.^[61] Colour vision may be widespread but has been demonstrated in only a few species.^[62]^[63] Some butterflies have organs of hearing and some species make stridulatory and clicking sounds.^[64]

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 {{convert|27|°C}}; 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 {{convert|40|°C}}, they can orientate themselves with the folded wings edgewise to the sun.^[65] 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.^[66]

As in many other insects, the lift generated by butterflies is more than can be accounted for by steady-state, non-transitory aerodynamics. Studies using Vanessa atalanta in a wind tunnel 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 the Weis-Fogh 'clap-and-fling' mechanism. Butterflies are able to change from one mode to another rapidly.^[67]

Ecology

Parasitoids, predators, and pathogens

Butterflies are threatened in their early stages by parasitoids 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 the large white butterfly.^[68] When the small 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.^[69] 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.^[70] Predators of butterflies include ants, spiders, wasps, and birds.^[71]

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.^[70] The bacterium Bacillus thuringiensis has been used in sprays to reduce damage to crops by the caterpillars of the large white butterfly, and the entomopathogenic fungus Beauveria bassiana has proved effective for the same purpose.^[72]

Endangered Species

Black grass-dart butterfly (Ocybadistes knightorum) is a butterfly of the family Hesperiidae. It is endemic to New South Wales. It has a very limited distribution in the Boambee area.

Defences

Chemical defences 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 sequester these plant toxins and use them instead in their own defence.^[75] 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 commonly mimicked by other butterflies, usually only females. A Batesian mimic imitates another species to enjoy the protection of that species' aposematism.^[91] The common Mormon of India has female morphs which imitate the unpalatable red-bodied swallowtails, the common rose and the crimson rose.^[76] 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.^[77]

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.^[84]^[85] Many butterflies have eyespots on the wings; these too may deflect attacks, or may serve to attract mates.^[50]^[86]

Auditory defenses can also be used, which in the case of the Grizzled skipper refers to vibrations generated by the butterfly upon expanding its wings in an attempt to communicate with ant predators.^[87]

Many tropical butterflies have seasonal forms for dry and wet seasons.^[88]^[89] These are switched by the hormone ecdysone.^[90] 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.^[91]^[92]^[93]

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 the turbulence created by the small whirlpools formed by the wings during flight.^[94]

In culture

In art and literature

Butterflies have appeared in art from 3500 years ago in ancient Egypt.^[95] In the ancient Mesoamerican city of Teotihuacan, the brilliantly coloured image of the butterfly was carved into many temples, buildings, jewellery, and emblazoned on incense burners. The butterfly was sometimes depicted with the maw of a jaguar, 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 the Aztec civilisation; evidence of similar jaguar-butterfly images has been found among the Zapotec and Maya civilisations.^[96]

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.^[97] The Norwegian naturalist Kjell Sandved compiled a photographic Butterfly Alphabet containing all 26 letters and the numerals 0 to 9 from the wings of butterflies.^[98]

Sir John Tenniel drew a famous illustration of Alice meeting a caterpillar for Lewis Carroll's Alice in Wonderland, c. 1865. The caterpillar is seated on a toadstool and is smoking a hookah; the image can be read as showing either the forelegs of the larva, or as suggesting a face with protruding nose and chin.^[1] Eric Carle's children's book The Very Hungry Caterpillar portrays the larva as an extraordinarily hungry animal, while also teaching children how to count (to five) and the days of the week.^[1]

One of the most popular, and most often recorded, songs by Sweden's eighteenth-century bard, Carl Michael Bellman, is "Fjäriln vingad syns på Haga" (The butterfly wingèd is seen in Haga), one of his Fredman's Songs.^[99]

In mythology and folklore

According to Lafcadio 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 bad omens. 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.^[101]

Diderot's Encyclopédie 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.^[102] In line with this, the ancient Greek word for "butterfly" is ψυχή (psȳchē), which primarily means "soul" or "mind".^[103] According to Mircea Eliade, some of the Nagas of Manipur claim ancestry from a butterfly.^[104] In some cultures, butterflies symbolise rebirth.^[105] The butterfly is a symbol of being transgender, because of the transformation from caterpillar to winged adult.^[106] In the English county of Devon, people once hurried to kill the first butterfly of the year, to avoid a year of bad luck.^[107] In the Philippines, a lingering black butterfly or moth in the house is taken to mean a death in the family.^[108] Several American states have chosen an official state butterfly.^[109]

Collecting, recording, and rearing

Collecting butterflies was once a popular hobby; it has now largely been replaced by photography, recording, and rearing butterflies for release into the wild.^[1] The zoological illustrator Frederick 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 handbook The Natural History of British Butterflies in 1924.^[1]

In technology

Study of the structural coloration of the wing scales of swallowtail butterflies has led to the development of more efficient light-emitting diodes,^[110] and is inspiring nanotechnology research to produce paints that do not use toxic pigments and the development of new display technologies.^[111]

References

1. ^¹²³⁴⁵{{cite book |last1=Marren |first1=Peter |last2=Mabey |first2=Richard |title=Bugs Britannica |date=2010 |publisher=Chatto and Windus |isbn=978-0-7011-8180-2 |pages=196–205}}
2. ^Donald A. Ringe, A Linguistic History of English: From Proto-Indo-European to Proto-Germanic (Oxford: Oxford, 2003), 232.
3. ^{{cite book |last1=Grimaldi |first1=David A. |author2=Engel, Michael S. |title=Evolution of the Insects |url=https://books.google.com/books?id=Ql6Jl6wKb88C&pg=PA561 |year=2005 |publisher=Cambridge University Press |isbn=978-0-521-82149-0 |page=561}}
4. ^{{cite book |last1=Davies |first1=Hazel |last2=Butler |first2=Carol A. |title=Do Butterflies Bite? |url=https://books.google.com/books?id=6bOI4-tfxJkC&pg=PA48 |year=2008 |publisher=Rutgers University Press |isbn=978-0-8135-4268-3 |page=48}}
5. ^{{cite book |last1=Meyer |first1=Herbert William |last2=Smith |first2=Dena M .|title=Paleontology of the Upper Eocene Florissant Formation, Colorado |url=https://books.google.com/books?id=-OcQecTR5_QC&pg=PA6 |year=2008 |publisher=Geological Society of America |isbn=978-0-8137-2435-5 |page=6}}
6. ^{{cite web |url=http://www.faculty.ucr.edu/~legneref/identify/lepidopt.htm |title=Lepidoptera – Latest Classification |work=Discoveries in Natural History & Exploration |publisher=University of California |accessdate=15 July 2011}}
7. ^{{cite journal | last1 = McIntosh | first1 = W. C. |display-authors=et al | year = 1992 | title = Calibration of the Latest Eocene-Oligocene geomagnetic Polarity Time Scale Using 40Ar/39Ar Dated Ignimbrites | url = | journal = Geology | volume = 20 | issue = 5| pages = 459–463 | doi=10.1130/0091-7613(1992)020<0459:cotleo>2.3.co;2| bibcode = 1992Geo....20..459M }}
8. ^{{cite journal | last1 = Heikkilä | first1 = M. | last2 = Kaila | first2 = L. | last3 = Mutanen | first3 = M. | last4 = Peña | first4 = C. | last5 = Wahlberg | first5 = N. | year = 2012 | title = Cretaceous Origin and Repeated Tertiary Diversification of the Redefined Butterflies | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 279 | issue = 1731| pages = 1093–1099 | doi=10.1098/rspb.2011.1430 | pmid=21920981 | pmc=3267136}}
9. ^{{cite journal | last1 = Kawahara | first1 = A. Y. | last2 = Breinholt | first2 = J. W. | year = 2014 | title = Phylogenomics Provides Strong Evidence for Relationships of Butterflies And Moths | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 281 | issue = 1788| page = 20140970 | doi=10.1098/rspb.2014.0970 | pmid=24966318 | pmc=4083801}}
10. ^{{cite journal |last=Mason |first=C. W. |title=Structural Colors in Insects. II |year=1927 |journal=The Journal of Physical Chemistry |volume=31 |pages=321–354 |issue=3 |doi=10.1021/j150273a001}}
11. ^{{cite journal |author1=Vukusic, P. |author2=J. R. Sambles |author3=H. Ghiradella |last-author-amp=yes |date=2000 |title=Optical Classification of Microstructure in Butterfly Wing-scales |journal=Photonics Science News |volume=6 |pages=61–66}}
12. ^{{cite journal |doi=10.1242/jeb.02051 |date=February 2006 |author1=Prum, R. |author2=Quinn, T. |author3=Torres, R. |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 }}
13. ^¹²³⁴{{cite book |author1=Gullan, P. J. |author2=Cranston, P. S. |title=The Insects: An Outline of Entomology | edition=5 |url=https://books.google.com/books?id=lF5hBAAAQBAJ&pg=PT790 |year=2014 |publisher=Wiley |isbn=978-1-118-84616-2 |pages=523–524}}
14. ^{{cite journal | last1 = Herrera | first1=Carlos M. | title=Activity Pattern and Thermal Biology of a Day-Flying Hawkmoth (Macroglossum stellatarum) under Mediterranean summer conditions | journal=Ecological Entomology | volume=17 | pages=52–56 | year=1992 | doi=10.1111/j.1365-2311.1992.tb01038.x | hdl=10261/44693 }}
15. ^{{cite web |title=Butterflies and Moths (Order Lepidoptera) |url=http://www.amentsoc.org/insects/fact-files/orders/lepidoptera.html |publisher=Amateur Entomologists' Society |accessdate=13 September 2015}}
16. ^{{cite journal |pmid=9292232 |date=August 1997 |last1=Traut |first1=W. |last2=Marec |first2=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}}
17. ^¹{{cite web |last1=Williams |first1=Ernest |last2=Adams |first2=James |last3=Snyder |first3=John |title=Frequently Asked Questions |url=http://www.lepsoc.org/frequently_asked_questions.php |publisher=The Lepidopterists' Society |accessdate=9 September 2015 |deadurl=yes |archiveurl=https://web.archive.org/web/20150513015255/http://www.lepsoc.org/frequently_asked_questions.php |archivedate=13 May 2015 |df=dmy-all }}
18. ^{{cite web |title=Global Distribution |url=http://monarchlab.org/biology-and-research/biology-and-natural-history/global-distribution |publisher=Monarch Lab |accessdate=9 September 2015}}
19. ^{{cite journal |url=http://www.sciencenews.org/view/generic/id/348485/description/Chill_turns_monarchs_north |title=Chill Turns Monarchs North; Cold Weather Flips Butterflies' Migratory Path |journal=Science News |date=23 March 2013 |volume=183 |issue=6}}
20. ^{{cite book |author=Pyle, Robert Michael |title=National Audubon Society Field Guide to North American Butterflies |pages=712–713 |publisher=Alfred A. Knopf |isbn=978-0-394-51914-2|year=1981 }}
21. ^{{cite web|title=Butterfly Conservation: Secrets of Painted Lady migration unveiled|url=http://www.birdguides.com/webzine/article.asp?a=3493|publisher=BirdGuides Ltd|accessdate=22 October 2012|date=2012-10-22}}
22. ^{{cite journal | last1=Williams | first1=C. B. | year=1927 | title=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 | journal=Transactions of the Entomological Society of London | volume=75 | issue=| pages=1–33 | doi=10.1111/j.1365-2311.1927.tb00054.x}}
23. ^{{cite journal | doi=10.4039/Ent1091583-12 | last1=Urquhart | first1=F. A. | last2=Urquhart | first2=N. R. | year=1977 | title=Overwintering Areas and Migratory Routes of the Monarch butterfly (Danaus p. plexippus, Lepidoptera: Danaidae) in North America, with Special Reference to the Western Population | journal=Can. Ent. | volume=109 | issue=12| pages=1583–1589}}
24. ^{{cite journal | doi=10.1073/pnas.95.26.15436 | last1=Wassenaar | first1=L.I. | last2=Hobson | first2=K.A. | year=1998 | title=Natal Origins of Migratory Monarch Butterflies at Wintering Colonies in Mexico: New Isotopic Evidence | journal=Proc. Natl. Acad. Sci. U.S.A. | volume=95 | issue=26| pages=15436–9 | pmid=9860986 | pmc=28060| bibcode=1998PNAS...9515436W }}
25. ^{{cite journal | doi=10.1016/j.cub.2003.12.034 | last1=Reppert | first1=Steven M. | last2=Zhu | first2=Haisun | last3=White | first3=Richard H. | year=2004 | title=Polarized light Helps Monarch Butterflies Navigate | journal=Current Biology | volume=14 | issue=2| pages=155–158 | pmid=14738739}}
26. ^{{cite journal | doi=10.1016/j.neuron.2005.03.014 | last1=Sauman | first1=Ivo | last2=Briscoe | first2=Adriana D. | last3=Zhu | first3=Haisun | last4=Shi | first4=Dingding | last5=Froy | first5=Oren | last6=Stalleicken | first6=Julia | last7=Yuan | first7=Quan | last8=Casselman | first8=Amy | last9=Reppert | first9=Steven M. | year=2005 | title=Connecting the Navigational Clock to Sun Compass Input in Monarch Butterfly Brain | journal=Neuron | volume=46 | issue=3| pages=457–467 | pmid=15882645 |display-authors=etal}}
27. ^{{cite journal | doi=10.1111/j.1469-185X.1962.tb01609.x | last1=Southwood | first1=T. R. E. | year=1962 | title=Migration of terrestrial arthropods in relation to habitat | journal=Biol. Rev. | volume=37 | issue=2| pages=171–214}}
28. ^{{cite journal | doi=10.1007/s10841-005-5660-x | last1=Dennis | first1=R L H | last2=Shreeve | first2=Tim G. | last3=Arnold | first3=Henry R. | last4=Roy | first4=David B. | year=2005 | title=Does Diet Breadth Control Herbivorous Insect Distribution Size? Life History and Resource Outlets for Specialist Butterflies | journal=Journal of Insect Conservation | volume=9 | issue=3| pages=187–200}}
29. ^{{cite journal | last1=Powell | first1=J. A. | year=1987 | title=Records of Prolonged Diapause in Lepidoptera | journal=J. Res. Lepid. | volume=25 | pages=83–109}}
30. ^{{cite web |url=http://www.butterfliesandmoths.org/species/Oeneis-melissa |title=Melissa Arctic |work=Butterflies and Moths of North America |accessdate=15 September 2015}}
31. ^{{cite book |author=Timothy Duane Schowalter |title=Insect Ecology: An Ecosystem Approach |url=https://books.google.com/books?id=2KzokTLIysQC&pg=PA159 |year=2011 |publisher=Academic Press |isbn=978-0-12-381351-0 |page=159}}
32. ^{{cite journal |last1=Arikawa |first1=Kentaro |title=Hindsight of Butterflies: The Papilio butterfly has light sensitivity in the genitalia, which appears to be crucial for reproductive behavior |journal=BioScience |volume=51|issue=3 |pages=219–225 |doi=10.1641/0006-3568(2001)051[0219:HOB]2.0.CO;2 |year=2001 }}
33. ^{{cite book |last=Schlaepfer |first=Gloria G. |title=Butterflies |url=https://books.google.com/books?id=HZKIf_QBbfgC&pg=PA52 |year=2006 |publisher=Marshall Cavendish |isbn=978-0-7614-1745-3 |page=52}}
34. ^¹²{{cite book |author=Capinera, John L. |title=Encyclopedia of Entomology |url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA640 |year=2008 |publisher=Springer Science & Business Media |isbn=978-1-4020-6242-1 |page=640}}
35. ^{{cite journal |author1=Beament, J.W.L. |author2=Lal, R. |year=1957 |title=Penetration Through the Egg-shell of Pieris brassicae |journal=Bulletin of Entomological Research |volume=48 |issue=1 |pages=109–125 |doi=10.1017/S0007485300054134}}
36. ^{{cite book|author=Scott, James A. |title=The Butterflies of North America: A Natural History and Field Guide |url=https://books.google.com/books?id=Oa5m8gZcGjMC&pg=PA121 |year=1992 |publisher=Stanford University Press |isbn=978-0-8047-2013-7 |page=121}}
37. ^{{cite book|author=Capinera, John L. |title=Encyclopedia of Entomology|url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA676 |year=2008 |publisher=Springer Science & Business Media |isbn=978-1-4020-6242-1 |page=676}}
38. ^{{cite book |author1=Shepard, Jon |author2=Guppy, Crispin |title=Butterflies of British Columbia: Including Western Alberta, Southern Yukon, the Alaska Panhandle, Washington, Northern Oregon, Northern Idaho, and Northwestern Montana |url=https://books.google.com/books?id=il6rJ7glHNQC&pg=PA55 |year=2011 |publisher=UBC Press |isbn=978-0-7748-4437-6 |page=55}}
39. ^{{cite web |title=British Butterflies: Education: Butterflies in Winter |url=http://www.britishbutterflies.co.uk/winter.asp |accessdate=12 September 2015}}
40. ^{{cite web |title=Camberwell Beauty |url=http://www.luontoportti.com/suomi/en/perhoset/camberwell-beauty |publisher=NatureGate |accessdate=12 September 2015}}
41. ^{{cite journal |author1=Venkatesha, M. G. |author2=Shashikumar, L. |author3=Gayathri Devi, S.S. |year=2004 |title=Protective Devices of the Carnivorous Butterfly, Spalgis epius (Westwood) (Lepidoptera: Lycaenidae) |journal=Current Science |volume=87 |issue=5 |pages=571–572}}
42. ^{{cite book |last1=Bingham |first1=C.T. |authorlink=Charles Thomas Bingham |title=The Fauna of British India, Including Ceylon and Burma |url=https://archive.org/details/butterflies02bingiala |volume=II |edition=1st |publisher= Taylor and Francis, Ltd. |location=London |year=1907 }}
43. ^{{cite journal |last=Devries |first=P. J. |title=The larval Ant-organs of Thisbe irenea (Lepidoptera: Riodinidae) and Their Effects Upon Attending Ants |year=1988 |journal=Zoological Journal of the Linnean Society |volume=94 |issue=4 |doi=10.1111/j.1096-3642.1988.tb01201.x |pages=379–393}}
44. ^{{cite journal |last=Devries |first=P. J. |title=Enhancement of Symbioses Between Butterfly Caterpillars and Ants by Vibrational Communication |date=June 1990 |journal=Science |volume=248 |issue=4959 |doi=10.1126/science.248.4959.1104 |pmid=17733373 |pages=1104–1106|bibcode=1990Sci...248.1104D }}
45. ^{{cite journal |last1=Thomas |first1=Jeremy |last2=Schönrogge |first2=Karsten |last3=Bonelli |first3=Simona |last4=Barbero |first4=Francesca |last5=Balletto |first5=Emilio |title=Corruption of Ant Acoustical Signals by Mimetic Social Parasites |journal=Communicative and Integrative Biology |year=2010 |volume=3 |issue=2 |pages=169–171 |pmid=20585513 |pmc=2889977 |doi=10.4161/cib.3.2.10603}}
46. ^{{cite book |author=Klowden, Marc J. |title=Physiological Systems in Insects |url=https://books.google.com/books?id=CABp1YL0F8gC&pg=PA114 |year=2013 |publisher=Academic Press |isbn=978-0-12-415970-9 |page=114}}
47. ^{{cite journal | last=Locke | first=Michael | title=Caterpillars have evolved lungs for hemocyte gas exchange | journal=Journal of Insect Physiology | volume=44 | issue=1 | year=1997 | doi=10.1016/s0022-1910(97)00088-7 | pmid=12770439 | pages=1–20 |url=https://www.academia.edu/33633960/Caterpillars_have_evolved_lungs_for_hemocyte_gas_exchange}}
48. ^{{cite web | url=http://www.naturemuseum.org/online/thebutterflylab/anatomy/larva.html | archiveurl=https://web.archive.org/web/20120319072020/http://www.naturemuseum.org/online/thebutterflylab/anatomy/larva.html |archivedate=19 March 2012 | title=Peggy Notebaert Nature Museum | publisher=Chicago Academy of Sciences | work=Larva Legs | accessdate=7 June 2012}}
49. ^¹²³⁴⁵⁶{{cite web |url=http://www.britannica.com/animal/lepidopteran/Form-and-function |title=Lepidopteran: Form and function |author=Culin, Joseph |publisher=Encyclopædia Britannica |accessdate=8 September 2015}}
50. ^¹{{cite journal |author1=Brunetti, Craig R. |author2=Selegue, Jayne E. |author3=Monteiro, Antonia |author4=French, Vernon |author5=Brakefield, Paul M. |author6=Carroll, Sean B. |year=2001 |title=The Generation and Diversification of Butterfly Eyespot Color Patterns |journal=Current Biology |volume=11 |issue=20 |pages=1578–1585 |doi=10.1016/S0960-9822(01)00502-4 |pmid=11676917}}
51. ^{{Cite journal |pmid = 25958827|year = 2015|last1 = Harpel|first1 = D.|title = Pollen feeding proteomics: Salivary proteins of the passion flower butterfly, Heliconius melpomene|journal = Insect Biochemistry and Molecular Biology|volume = 63|pages = 7–13|last2 = Cullen|first2 = D. A.|last3 = Ott|first3 = S. R.|last4 = Jiggins|first4 = C. D.|last5 = Walters|first5 = J. R.|doi = 10.1016/j.ibmb.2015.04.004|url=https://lirias.kuleuven.be/handle/123456789/509592}}
52. ^{{cite web |url=https://sta.uwi.edu/fst/lifesciences/sites/default/files/lifesciences/documents/ogatt/Heliconius_ethilla%20-%20Ethilia%20Longwing%20Butterfly.pdf |title=The Online Guide to the Animals of Trinidad and Tobago {{!}} Heliconius ethilla (Ethilia Longwing Butterfly) |publisher=UWI St. Augustine |accessdate=28 May 2018}}
53. ^{{cite book |author=Woodbury, Elton N. |title=Butterflies of Delmarva |url=https://books.google.com/books?id=cDNHAAAAYAAJ |year=1994 |publisher=Delaware Nature Society; Tidewater Publishers |isbn=978-0-87033-453-5 |page=22}}
54. ^{{cite journal |last=Gilbert |first=L. E. |year=1972 |title=Pollen Feeding and Reproductive Biology of Heliconius Butterflies |journal=Proceedings of the National Academy of Sciences |volume=69 |issue=6 |pages=1402–1407 |doi=10.1073/pnas.69.6.1403|pmid=16591992 |pmc=426712 |bibcode=1972PNAS...69.1403G }}
55. ^{{cite journal|last=Herrera |first=C. M. |year=1987 |title=Components of Pollinator 'Quality': Comparative Analysis of a Diverse Insect Assemblage |journal=Oikos |volume=50 |number=1 |pages=79–90 |url=http://ebd06.ebd.csic.es/pdfs/Herrera.1987.Oikos.pdf |doi=10.2307/3565403 |jstor=3565403 |deadurl=yes |archiveurl=https://web.archive.org/web/20090225100931/http://ebd06.ebd.csic.es/pdfs/Herrera.1987.Oikos.pdf |archivedate=2009-02-25 |df= }}
56. ^{{cite journal | doi=10.1006/anbe.1996.0390 | last1=Goulson | first1=D. | last2=Ollerton | first2=J. | last3=Sluman | first3=C. | year=1997 | title=Foraging strategies in the small skipper butterfly, Thymelicus flavus: when to switch? | journal=Animal Behaviour | volume=53 | issue=5| pages=1009–1016}}
57. ^{{cite journal | doi=10.1111/j.1095-8312.2005.00539.x | last1=Molleman |first1=Freerk | last2=Grunsven |first2=Roy H. A. | last3=Liefting |first3=Maartje | last4=Zwaan | year=2005 | first4=Bas J. | last5=Brakefield | first5=Paul M. | title=Is Male Puddling Behaviour of Tropical Butterflies Targeted at Sodium for Nuptial Gifts or Activity? | journal=Biological Journal of the Linnean Society | volume=86 | issue=3 | pages=345–361}}
58. ^{{cite book |last1=Gochfeld |first1=Michael |last2=Burger |first2=Joanna |title=Butterflies of New Jersey: A Guide to Their Status, Distribution, Conservation, and Appreciation |url=https://books.google.com/books?id=8SzNCNPkgQcC&pg=PA55 |year=1997 |publisher=Rutgers University Press |isbn=978-0-8135-2355-2 |page=55}}
59. ^{{cite web |url=http://www.sandiegozoo.org/animalbytes/t-butterfly.html |title=Article on San Diego Zoo website |publisher=Sandiegozoo.org |accessdate=2009-03-30}}
60. ^{{cite journal |last1=Birch |first1=M. C. |last2=Poppy |first2=G. M. |title=Scents and Eversible Scent Structures of Male Moths |journal=Annual Review of Entomology |date=1990 |volume=35 |pages=25–58 |url=http://ento.psu.edu/publications/512BirchEtAl1990.pdf |doi=10.1146/annurev.ento.35.1.25}}
61. ^{{cite journal | last1=Obara | first1=Y. | last2=Hidaki | first2=T. | year=1968 | title=Recognition of the Female by the Male, on the Basis of Ultra-Violet Reflection, in the White Cabbage Butterfly Pieris rapae crucivora | journal=Boisduval. Proc. Japan Acad. | volume=44 | issue=8| pages=829–832| doi=10.2183/pjab1945.44.829 }}
62. ^{{cite journal | last1=Hirota | first1=Tadao | last2=Yoshiomi | first2=Yoshiomi| year=2004| title=Color Discrimination on Orientation of Female Eurema hecabe (Lepidoptera: Pieridae) | journal=Applied Entomology and Zoology | volume=39 | issue=2| pages=229–233 | doi=10.1303/aez.2004.229}}
63. ^{{cite journal | last1=Kinoshita | first1=Michiyo | last2=Shimada | first2=Naoko | last3=Arikawa | first3=Kentaro | year=1999 | title=Color Vision of the Foraging Swallowtail Butterfly Papilio xuthus | journal=The Journal of Experimental Biology | volume=202 | issue=2| pages=95–102 | pmid=9851899}}
64. ^{{cite journal | last1=Swihart | first1=S. L | year=1967 | title=Hearing in Butterflies | journal=Journal of Insect Physiology | volume=13 | issue=3| pages=469–472 | doi=10.1016/0022-1910(67)90085-6}}
65. ^{{cite journal |title=Butterflies Make Best Use of the Sunshine |journal=New Scientist |url=https://books.google.com/books?id=PfHEmTgmHacC&pg=PA13 |date=17 December 1988 |page=13 |issn=0262-4079|author1=Information |first1=Reed Business }}
66. ^{{cite journal | last1=Ellers | first1=J. | last2=Boggs | first2=Carol L. | year=2002 | title=The Evolution of Wing Color in Colias Butterflies: Heritability, Sex Linkage, and population divergence | url=http://www.stanford.edu/group/CCB/Pubs/Boggs_pdfs/2002_Ellers_Boggs_Coliaswingcolor.pdf | journal=Evolution | volume=56 | issue=4| pages=836–840 | pmid=12038541 | doi=10.1554/0014-3820(2002)056[0836:teowci]2.0.co;2}}
67. ^{{cite journal | last1=Srygley | first1=R. B. | last2=Thomas | first2=A. L. R. | year=2002 | title=Aerodynamics of Insect Flight: Flow Visualisations with Free Flying Butterflies Reveal a Variety of Unconventional Lift-Generating Mechanisms | journal=Nature | volume=420 | issue=6916| pages=660–664 | pmid=12478291 | doi=10.1038/nature01223| bibcode=2002Natur.420..660S }}
68. ^{{cite book |author=Feltwell, J. |title=Large White Butterfly: The Biology, Biochemistry and Physiology of Pieris brassicae (Linnaeus) |url=https://books.google.com/books?id=8XnrCAAAQBAJ&pg=PA401 |year=2012 |publisher=Springer |isbn=978-94-009-8638-1 |pages=401–}}
69. ^{{cite book |author1=Burton, Maurice |author2=Burton, Robert |title=International Wildlife Encyclopedia: Brown bear - Cheetah |url=https://books.google.com/books?id=WLIOwLemzyQC&pg=PA416 |year=2002 |publisher=Marshall Cavendish |isbn=978-0-7614-7269-8 |page=416}}
70. ^¹{{cite book |author=Allen, Thomas J. |title=A Field Guide to Caterpillars |url=https://books.google.com/books?id=SS6s7tck0fIC&pg=PA15 |year=2005 |publisher=Oxford University Press |isbn=978-0-19-803413-1 |page=15}}
71. ^{{cite web |title=Parasites and Natural Enemies |url=http://monarchlab.org/biology-and-research/biology-and-natural-history/parasites-natural-enemies |publisher=University of Minnesota |accessdate=16 October 2015}}
72. ^{{cite book |author=Feltwell, J. |title=Large White Butterfly: The Biology, Biochemistry and Physiology of Pieris Brassicae (Linnaeus) |url=https://books.google.com/books?id=8XnrCAAAQBAJ&pg=PA429 |year=2012 |publisher=Springer |isbn=978-94-009-8638-1 |page=429}}
73. ^CITES appendices I, II and III, official website
74. ^{{cite journal |doi=10.1371/journal.pbio.0040341 |date=October 2006 |last=Meyer |first=A. |title=Repeating Patterns of Mimicry |volume=4 |issue=10 |pages=e341 |issn=1544-9173 |pmid=17048984 |pmc=1617347 |journal=PLoS Biology}}
75. ^{{cite journal | doi=10.1146/annurev.ento.47.091201.145121 | last1=Nishida | first1=Ritsuo | year=2002 | title=Sequestration of Defensive Substances from Plants by Lepidoptera | journal=Annual Review of Entomology | volume=47 | pages=57–92 | pmid=11729069}}
76. ^{{cite web |last1=Halloran |first1=Kathryn |last2=Wason |first2=Elizabeth |title=Papilio polytes |url=http://animaldiversity.org/accounts/Papilio_polytes/|publisher=University of Michigan Museum of Zoology |accessdate=12 September 2015 |date=2013}}
77. ^¹²{{cite book | author=Edmunds, M. | year=1974 | title=Defence in Animals | publisher=Longman | pages=74–78, 100–113}}
78. ^{{cite journal | doi=10.1086/283868 | last1=Robbins | first1=Robert K. | year=1981 | title=The "False Head" Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies | journal=American Naturalist | volume=118 | issue=5| pages=770–775}}
79. ^{{cite book| author=Forbes, Peter |date=2009 |title=Dazzled and Deceived: Mimicry and Camouflage |publisher=Yale University Press |isbn=978-0-300-17896-8|title-link=Dazzled and Deceived: Mimicry and Camouflage }}
80. ^{{cite web | url=http://www.springerreference.com/docs/html/chapterdbid/86053.html | title=Deimatic Behavior | publisher=Springer | year=2012 | accessdate=31 December 2012 | author=Edmunds, Malcolm}}
81. ^{{cite web|title=Featured Creatures: Giant Swallowtail |url=http://entnemdept.ifas.ufl.edu/creatures/citrus/giantswallowtail.htm |publisher=University of Florida |accessdate=12 September 2015}}
82. ^{{cite journal|author1=Fiedler, K. |author2=Holldobler, B. |author3=Seufert, P. |title=Butterflies and Ants: The Communicative Domain |journal=Cellular and Molecular Life Sciences |volume=52 |date=1996 |pages=14–24 |doi=10.1007/bf01922410}}
83. ^{{cite journal | last1 = Nafus | first1 = D. M. | last2 = Schreiner | first2 = I. H. | year = 1988 | title = Parental Care in a Tropical Nymphalid Butterfly Hypolimas anomala | url = | journal = Animal Behaviour | volume = 36 | issue = 5| pages = 1425–1443 | doi=10.1016/s0003-3472(88)80213-6}}
84. ^{{cite journal | author=Cooper, William E. Jr. |year=1998 |title=Conditions Favoring Anticipatory and Reactive Displays Deflecting Predatory Attack |journal=Behavioral Ecology |volume=9 |issue=6 |pages=598–604 |doi=10.1093/beheco/9.6.598}}
85. ^{{cite journal | doi=10.1017/S1464793105006810 | last1=Stevens | first1=M. | year=2005 | title=The Role of Eyespots as Anti-Predator Mechanisms, Principally Demonstrated in the Lepidoptera | journal=Biological Reviews | volume=80 | issue=4| pages=573–588 | pmid=16221330}}
86. ^{{cite journal | author=Brakefield, PM | year=1996 | title=Development, Plasticity and Evolution of Butterfly Eyespot Patterns | journal=Nature | issue=6606| pages=236–242 | volume=384 | doi=10.1038/384236a0 | pmid=12809139| last2=Gates | first2=Julie | last3=Keys | first3=Dave | last4=Kesbeke | first4=Fanja | last5=Wijngaarden | first5=Pieter J. | last6=Montelro | first6=Antónia | last7=French | first7=Vernon | last8=Carroll | first8=Sean B. |display-authors=etal| bibcode=1996Natur.384..236B }}
87. ^{{Cite journal|last=Elfferich|first=Nico W.|date=1998|title=Is the larval and imaginal signalling of Lycaenidae and other Lepidoptera related to communication with ants|url=http://natuurtijdschriften.nl/search?identifier=538588|journal=Deinsea|volume=4|issue=1}}
88. ^{{cite journal |pmid=18811432 |date=December 1998 |author1=Brakefield, Pm |author2=Kesbeke, F |author3=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}}
89. ^{{cite journal | doi=10.1006/mpev.2000.0872 | last1=Monteiro | first1=A. | last2=Pierce | first2=N. E. | year=2001 | title=Phylogeny of Bicyclus (Lepidoptera: Nymphalidae) Inferred from COI, COII, and EF-1 Alpha Gene Sequences | journal=Molecular Phylogenetics and Evolution | volume=18 | issue=2 | pages=264–281 | pmid=11161761}}
90. ^{{cite journal |pmid=12492404 |date=Jan 2003 |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}}
91. ^{{cite journal |last1=Brakefield |first1=Paul M. |last2=Larsen |first2=Torben B. |title=The Evolutionary Significance of Dry and Wet Season Forms in some Tropical Butterflies |year=1984 |journal=Biological Journal of the Linnean Society |volume=22 |doi=10.1111/j.1095-8312.1984.tb00795.x |pages=1–12|hdl=1887/11011 }}
92. ^{{cite journal | doi=10.1098/rspb.2003.2571 | last1=Lyytinen | first1=A. | last2=Brakefield | first2=P. M. | last3=Lindström | first3=L. | last4=Mappes | first4=J. | year=2004 | title=Does Predation Maintain Eyespot Plasticity in Bicyclus anynana | journal=Proceedings of the Royal Society B | volume=271 | issue=1536| pages=279–283| pmc=1691594 | pmid=15058439}}
93. ^{{cite journal | doi=10.1038/384236a0 | last1=Brakefield | first1=P. M. | last2=Gates | first2=J. | last3=Keys | first3=D. | last4=Kesbeke | first4=F. | last5=Wijngaarden | first5=P. J. | last6=Monteiro | first6=A. | last7=French | first7=V. | last8=Carroll | first8=S. B. | year=1996 | title=Development, Plasticity and Evolution of Butterfly Eyespot Patterns | journal=Nature | volume=384 | issue=6606 | pages=236–242 | pmid=12809139| bibcode=1996Natur.384..236B }}
94. ^[https://www.insidescience.org/news/mathematical-butterfly-simulations-provide-new-insights-flight The Mathematical Butterfly: Simulations Provide New Insights On Flight]
95. ^{{cite journal |author=Larsen, Torben |year=1994 |title=Butterflies of Egypt |journal=Saudi Aramco World |volume=45 |issue=5 |pages=24–27 |url=http://www.saudiaramcoworld.com/issue/199405/butterflies.of.egypt.htm}}
96. ^{{cite book | title=The Gods and Symbols of Ancient Mexico and the Maya |author=Miller, Mary |year=1993 |publisher=Thames & Hudson |isbn=978-0-500-27928-1}}
97. ^{{cite web|url=http://mfjoe.com/tag/furniture/ |title=Table Complete with Real Butterflies Embedded in Resin |accessdate=30 March 2009 |deadurl=yes |archiveurl=https://web.archive.org/web/20100506094545/http://mfjoe.com/tag/furniture/ |archivedate=6 May 2010 |df= }}
98. ^{{cite news |last=Pinar |title=Entire Alphabet Found on the Wing Patterns of Butterflies |url=http://www.mymodernmet.com/profiles/blogs/kjell-bloch-sandved-butterfly-alphabet |date=13 November 2013}}
99. ^{{cite web |last1=Nilsson |first1=Hans |title=Bellman på Spåren |url=http://www.bellman.net/artiklar/sparen.html |publisher=Bellman.net |accessdate=13 September 2015 |language=Swedish}}
100. ^{{cite book |author=Van Rij, Jan |url=https://books.google.com/books?id=rV3VeiACJuYC |title=Madame Butterfly: Japonisme, Puccini, and the Search for the Real Cho-Cho-San |publisher=Stone Bridge Press |year=2001|isbn=9781880656525 }}
101. ^{{cite book | last=Hearn | first=Lafcadio | authorlink=Lafcadio Hearn | year=1904 | title=Kwaidan: Stories and Studies of Strange Things | titlelink=Kwaidan: Stories and Studies of Strange Things | publisher=Dover | isbn=978-0-486-21901-1}}
102. ^{{Cite journal | url=http://quod.lib.umich.edu/cgi/t/text/text-idx?c=did;cc=did;rgn=main;view=text;idno=did2222.0001.694 | title=Butterfly |journal=Encyclopedia of Diderot and d'Alembert | accessdate=1 April 2015| date=January 2011 | last1=Louis | first1=Chevalier de Jaucourt (Biography) }}
103. ^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.
104. ^Rabuzzi, M. 1997. Butterfly etymology. Cultural Entomology November 1997 Fourth issue online
105. ^{{cite web |title=Church Releases Butterflies as Symbol of Rebirth |url=http://staugustine.com/living/religion/2015-04-16/church-releases-butterflies-symbol-rebirth#.Ve77UZdUWHg |publisher=The St. Augustine Record |accessdate=8 September 2015}}
106. ^{{cite web |title=I'm Scared to Be a Woman |url=https://www.hrw.org/report/2014/09/24/im-scared-be-woman/human-rights-abuses-against-transgender-people-malaysia |publisher=Human Rights Watch |accessdate=8 September 2015 |date=24 September 2014 |quote=a 22-year-old transgender woman sports a tattoo of a butterfly—a transgender symbol signifying transformation}}
107. ^Dorset Chronicle, May 1825, reprinted in: [https://books.google.com/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, etc. Vol III., ed. William Hone, (London: 1838) p 678.
108. ^{{cite web|title=Superstitions and Beliefs Related to Death|url=http://www.livinginthephilippines.com/culture-and-people/philippine-culture/superstitions-and-beliefs/419-superstitions-and-beliefs|website=Living in the Philippines|accessdate=9 October 2015}}
109. ^{{cite web |url=http://www.netstate.com/states/tables/state_butterflies.htm |title=Official State Butterflies |publisher=NetState.com |accessdate=9 September 2015}}
110. ^{{cite journal |author1=Vukusic, Pete |author2=Hooper, Ian |year=2005 |title=Directionally Controlled Fluorescence Emission in Butterflies |journal=Science |volume=310 |issue=5751 |page=1151 |doi=10.1126/science.1116612 |pmid=16293753}}
111. ^{{cite web |last1=Vanderbilt |first1=Tom |title=How Biomimicry is Inspiring Human Innovation |url=http://www.smithsonianmag.com/science-nature/how-biomimicry-is-inspiring-human-innovation-17924040/?no-ist |publisher=Smithsonian |accessdate=8 September 2015}}