Butterflies

Generated on: 17 July 2025

Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, 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 the Paleocene, about 56 million
years ago, though molecular evidence suggests that they likely originated in the Cretaceous.
Butterflies have a four-stage life cycle, and like other holometabolous insects they undergo
complete metamorphosis. Winged adults lay eggs on plant foliage 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, 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. 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. The Smithsonian Institution says "butterflies are
certainly one of the most appealing creatures in nature". == Etymology == The Oxford English
Dictionary derives the word straightforwardly from Old English butorfl■oge, butter-fly; similar
names in Old Dutch and Old High German show that the name is ancient, but modern Dutch and Germ
use different words (vlinder and Schmetterling) 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. == Paleontology == The earliest
Lepidoptera fossils date to the Triassic-Jurassic boundary, around 200 million years ago.
Butterflies evolved from moths, so while the butterflies are monophyletic (forming a single clade),
the moths are not. The oldest known butterfly is Protocoeliades kristenseni from the Palaeocene aged
Fur Formation of Denmark, approximately 55 million years old, which belongs to the family
Hesperiidae (skippers). Molecular clock estimates suggest that butterflies originated sometime in
the Late Cretaceous, but only significantly diversified during the Cenozoic, with one study
suggesting a North American origin for the group. The oldest American butterfly is the Late Eocene
Prodryas persephone from the Florissant Fossil Beds, approximately 34 million years old.
Butterfly fossils
== Taxonomy and phylogeny == Butterflies are divided into seven families that contain a total of
about 20,000 species. Traditionally, butterflies have been divided into the superfamilies
Papilionoidea and the moth-like Hedyloidea. Recent work has discovered that Hedylidae, the only
family within Hedyloidea, is nested within the Papilionoidea, meaning that Papilionoidea would be
synonymous with Rhopalocera. The relationships between the rest of the 6 families are extremely well
resolved, which is summarized in the below cladogram. == Biology == === General description ===
Butterfly adults are characterized by their four scale-covered wings, which give the Lepidoptera
their name (Ancient Greek λεπ■ς lepís, scale + πτερ■ν 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. 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.
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, are exceptions to these rules. 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. 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. Many butterflies are sexually dimorphic. Most butterflies have the ZW sex-
determination system where females are the heterogametic sex (ZW) and males homogametic (ZZ). =
Distribution and migration === Butterflies are distributed worldwide except Antarctica, totalling
some 18,500 species. 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.
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. 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. 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. Spectacular large-scale migrations associated
with the monsoon are seen in peninsular India. Migrations have been studied in more recent times
using wing tags and also using stable hydrogen isotopes. 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.
Many migratory butterflies live in semi-arid areas where breeding seasons are short. The life
histories of their host plants also influence butterfly behaviour. === 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. The Melissa Arctic (Oeneis melissa) overwinters twice as a caterpillar.
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.
Courtship is often aerial and often involves pheromones. Butterflies then land on the ground or on a
perch to mate. 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. The
male passes a spermatophore 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. 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. ==== Eggs ==== 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 hundr
eggs. 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. 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. 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. 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. 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. Some temperate
region butterflies, such as the Camberwell beauty, lay their eggs in the spring and have them hatch
in the summer. ==== 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, while
lycaenids such as Liphyra brassolis are myrmecophilous, eating ant larvae. 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. 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. 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. Caterpillars have short antennae and several simple eyes. The mouthparts
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. 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. 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. 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. ==== 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. 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. 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. ==== 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 cannot view distant objects clearly.
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 an
do not feed as adults. Many Heliconius butterflies also use their proboscis to feed on pollen; 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. 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. 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. 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. ==== Pattern formation ====
The colourful patterns on many butterfly wings tell potential predators that they are toxic. Hence,
the genetic basis of wing pattern formation can illuminate both the evolution of butterflies as well
as their developmental biology. The colour of butterfly wings is derived from tiny structures called
scales, each of which have their own pigments. In Heliconius 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, a gene called cortex determines the
colour of scales: deleting cortex turned black and red scales yellow. Mutations, e.g. transposon
insertions of the non-coding DNA around the cortex gene can turn a black-winged butterfly into a
butterfly with a yellow wing band. === Mating === When the butterfly Bicyclus anynana is subjected
to repeated inbreeding in the laboratory, there is a dramatic decrease in egg hatching. This severe
inbreeding depression is considered to be likely due to a relatively high mutation rate to recessive
alleles with substantial damaging effects and infrequent episodes of inbreeding in nature that might
otherwise purge such mutations. Although B. anynana experiences inbreeding depression when forcibly
inbred in the laboratory it recovers within a few generation when allowed to breed freely. 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. Inbreeding may persist in B 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. === Behaviour === Butterflies feed primarily on nectar from flowers. Some
also derive nourishment from pollen, 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. Flower constancy has been observed for at least one species of butterfly.
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 and scavenge 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. 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. 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 organ
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. Many butterflies use chemical signals,
pheromones; some have specialized scent scales (androconia) or other structures (coremata or "hair
pencils" in the Danaidae). 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. Colour vision may be widespread but has been demonstrated in only a few species. Some
butterflies have organs of hearing and some species make stridulatory and clicking sounds. 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.
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. 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. === 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. 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. 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. Predators of butterflies include ants, spiders,
wasps, and birds. 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. 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. ==== Endangered species ==== Queen Alexandra's birdwing, found in Papua New Gu
is the largest butterfly in the world. The species is endangered, and is one of only three insects
(the other two being butterflies as well) to be listed on Appendix I of CITES, making international
trade illegal. ==== Defences ==== 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 as protection against herbivores.
Butterflies have evolved mechanisms to sequester these plant toxins and use them instead in their
own defence. 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. The common Mormon of India has female morphs
which imitate the unpalatable red-bodied swallowtails, the common rose and the crimson rose.
Müllerian mimicry occurs when aposematic species evolve to resemble each other, presumably to redu
predator sampling rates; Heliconius butterflies from the Americas are a good example. Camouflage is
found in many butterflies. Some like the oakleaf butterfly and autumn leaf are remarkable imitations
of leaves. As caterpillars, many defend themselves by freezing and appearing like sticks or
branches. Others have deimatic behaviours, such as rearing up and waving their front ends which are
marked with eyespots as if they were snakes. Some papilionid caterpillars such as the giant
swallowtail (Papilio cresphontes) resemble bird droppings so as to be passed over by predators. Some
caterpillars have hairs and bristly structures that provide protection while others are gregarious
and form dense aggregations. Some species are myrmecophiles, forming mutualistic associations with
ants and gaining their protection. Behavioural defences include perching and angling the wings to
reduce shadow and avoid being conspicuous. Some female Nymphalid butterflies guard their eggs from
parasitoidal wasps. 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.
Many butterflies have eyespots on the wings; these too may deflect attacks, or may serve to attract
mates. Auditory defences 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. Many tropical butterflies have seasonal forms for dry and wet seasons. These are switched
by the hormone ecdysone. 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. 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.
== Declining numbers == Declining butterfly populations have been noticed in many areas of the
world, and this phenomenon is consistent with the rapidly 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 by global climate change, specifically, by warmer
autumns. Butterfly populations in the United States declined by 22% between 2000 and 2020, mainly
because of habitat loss, pesticides and climate change. == In culture == === In art and
literature === Butterflies have appeared in art from 3500 years ago in ancient Egypt. 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 upholding ma'at. They also
were suggestive of regeneration or rebirth and protection. Certain butterflies, such as the tiger
butterfly, may have been associated with solar deities, particularly Ra. The tiger butterfly also
would have a particular resemblance to the ankh, 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. 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. 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. 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. The butterfly is a symbol of being transgender, because of the
transformation from caterpillar to winged adult. 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. 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. A butterfly appeared in one of
Rudyard Kipling's Just So Stories, "The Butterfly that Stamped". 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's seen in Haga), one of his Fredman's Songs. Madam Butterfly is
a 1904 opera by Giacomo Puccini about a romantic young Japanese bride who is deserted by her
American officer husband soon after they are married. It was based on an 1898 short story by John
Luther Long. === 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 his famous revolt, a vast swarm
of butterflies appeared in Kyoto. The people were frightened, thinking the apparition to be a
portent of coming evil. 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. In line with this, the ancient Greek word for "butterfly" is ψυχ■
(ps■ch■), which primarily means "soul" or "mind". According to Mircea Eliade, some of the Nagas of
Manipur claim ancestry from a butterfly. In popular Burmese culture, the butterfly (called leippya)
is symbolic of the soul or consciousness of a person. During the transitory period after death, the
Burmese believe that the "butterfly soul" becomes a wandering spirit in search of a new corporeality
medium. In some cultures, butterflies symbolise rebirth. In the English county of Devon, people once
hurried to kill the first butterfly of the year, to avoid a year of bad luck. 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. Several American states have chosen an official state butterfly. === Collecting,
recording, and rearing === "Collecting" means preserving dead specimens, not keeping butterflies as
pets. Collecting butterflies was once a popular hobby; it has now largely been replaced by
photography, recording, and rearing butterflies for release into the wild. 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.
Butterflies and moths can be reared for recreation or for release. === 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, and is inspiring nanotechnology research to produce paints
that do not use toxic pigments and the development of new display technologies. == References ==
== Further reading == Kawahara, A.Y.; Storer, C.; Carvalho, A.P.S.; et al. (15 May 2023). "A Global
Phylogeny of Butterflies Reveals Their Evolutionary History, Ancestral Hosts and Biogeographic
Origins". Nat Ecol Evol. 7 (6): 903–913. Bibcode:2023NatEE...7..903K.
doi:10.1038/s41559-023-02041-9. PMC 10250192. PMID 37188966. == External links == Papilionoid
on the Tree of Life Archived 11 December 2008 at the Wayback Machine Butterfly species and
observations on iNaturalist Lamas, Gerardo (1990). "An Annotated List of Lepidopterological
Journals" (PDF). Journal of Research on the Lepidoptera. 29 (1–2): 92–104. doi:10.5962/p.266621.
S2CID 108756448. Archived from the original (PDF) on 27 August 2016. Rhopalocera at [Link]
=== Regional lists === North America Africa: Ghana Archived 16 January 2021 at the Wayback Machin
Asia: Singapore Israel Archived 22 October 2022 at the Wayback Machine Indo-China Sulawesi
(Southeastern Sulawesi) Turkey Wilkes, Benjamin (1749). The English Moths and Butterflies.