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8/13/2021 Butterfly - Wikipedia

Butterfly
Butterflies are insects in the macrolepidopteran clade
Rhopalocera from the order Lepidoptera, which also includes Butterflies

moths. Adult butterflies have large, often brightly coloured Temporal range: Palaeocene-
wings, and conspicuous, fluttering flight. The group comprises Present, 56–0 Ma
the large superfamily Papilionoidea, which contains at least one
former group, the skippers (formerly the superfamily PreꞒ Ꞓ O S D C P T J K PgN
"Hesperioidea"), and the most recent analyses suggest it also
contains the moth-butterflies (formerly the superfamily
"Hedyloidea"). Butterfly fossils date to the Paleocene, about 56
million years ago.

Butterflies have a four-stage life cycle, as like most insects they


undergo complete metamorphosis. 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 Papilio machaon
a year, while others have a single generation, and a few in cold
Scientific classification
locations may take several years to pass through their entire life
cycle. Kingdom: Animalia

Butterflies are often polymorphic, and many species make use Phylum: Arthropoda
of camouflage, mimicry and aposematism to evade their Class: Insecta
predators.[1] Some, like the monarch and the painted lady,
migrate over long distances. Many butterflies are attacked by Order: Lepidoptera
parasites or parasitoids, including wasps, protozoans, flies, and Suborder: Rhopalocera
other invertebrates, or are preyed upon by other organisms.
Some species are pests because in their larval stages they can Subgroups
damage domestic crops or trees; other species are agents of
pollination of some plants. Larvae of a few butterflies (e.g., Superfamily Hedyloidea:
harvesters) eat harmful insects, and a few are predators of ants,
while others live as mutualists in association with ants. Hedylidae
Culturally, butterflies are a popular motif in the visual and
Superfamily Papilionoidea:
literary arts.
Hesperiidae
Lycaenidae
Contents Nymphalidae
Etymology Papilionidae
Taxonomy and phylogeny
Pieridae
Biology
Riodinidae
General description
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Distribution and migration


Life cycle
Egg
Caterpillar larva
Pupa
Adult
Pattern formation
Behaviour
Ecology
Parasitoids, predators, and pathogens
Endangered species
Defences
Declining numbers
In culture
In art and literature
In mythology and folklore
Collecting, recording, and rearing
In technology
References
External links
Regional lists

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, but modern Dutch and German 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 The original "butter-fly"?[2] A male
season while the grass was growing.[2][3] brimstone (Gonepteryx rhamni) in
flight

Taxonomy and phylogeny


The earliest Lepidoptera fossils date to the Triassic-Jurassic boundary, around 200 million years
ago.[4] 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).[5] Molecular clock estimates suggest that butterflies originated
sometime in the mid-Cretaceous, but only signficantly diversified during the Cenozoic.[6] The oldest
American butterfly is the Late Eocene Prodryas persephone from the Florissant Fossil Beds,[7][8]
approximately 34 million years old.[9]
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Traditionally, 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.[10][11]

Butterfly families
Prodryas persephone, a Late
Common Eocene butterfly from the Florissant
Family Characteristics Image
name
Fossil Beds, 1887 engraving
Small, brown, like
American
geometrid moths;
Hedylidae moth-
antennae not clubbed;
butterflies
long slim abdomen

Small, darting flight;


Hesperiidae Skippers clubs on antennae
hooked backwards

Small, brightly coloured;


Blues, often have false heads
Lycaenidae coppers, with eyespots and small
hairstreaks tails resembling
antennae Lithopsyche antiqua, an Early
Oligocene butterfly from the
Brush- Usually have reduced Bembridge Marls, Isle of Wight,
footed or forelegs, so appear 1889 engraving
Nymphalidae
four-footed four-legged; often
butterflies brightly coloured

Often have 'tails' on


wings; caterpillar
Papilionidae Swallowtails generates foul taste with
osmeterium organ; pupa
supported by silk girdle

Mostly white, yellow or


Whites and orange; some serious
Pieridae
allies pests of Brassica; pupa
supported by silk girdle

Often have metallic


spots on wings; often
Riodinidae Metalmarks conspicuously coloured
with black, orange and
blue

Biology

General description

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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
Butterfly antennal shapes, mainly
with melanins that give them
clubbed, unlike those of moths.
blacks and browns, as well as uric
Drawn by C. T. Bingham, 1905
acid derivatives and flavones that The wings of butterflies, here
give them yellows, but many of the Aglais io, are covered with
blues, greens, reds and iridescent coloured scales.
colours are created by structural coloration produced by the micro-
structures of the scales and hairs.[12][13][14][15]

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.[16]

Nearly all butterflies are diurnal, have relatively bright colours, and Unlike butterflies, most moths
hold their wings vertically above their bodies when at rest, unlike the (like Laothoe populi) fly by
majority of moths which fly by night, are often cryptically coloured night and hide by day.
(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,[17] are exceptions to these
rules.[16][18]

Sexual dimorphism in Butterfly larvae, caterpillars, have a hard (sclerotised) head with strong
Anthocharis cardamines 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.[16] Many are well camouflaged;
Female
others are aposematic with bright colours and bristly projections
Male
containing toxic chemicals obtained from their food plants. The pupa
or chrysalis, unlike that of moths, is not wrapped in a cocoon.[16]

Many butterflies are sexually dimorphic. Most butterflies have the ZW sex-determination system
where females are the heterogametic sex (ZW) and males homogametic (ZZ).[19]

Distribution and migration

Butterflies are distributed worldwide except Antarctica, totalling some 18,500 species.[20] 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.[20] 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

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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.[21]

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
Monarch migration route completes the whole trip. The
eastern North American
population of monarchs can Overwintering monarchs cluster on
travel thousands of miles south-west to overwintering sites in oyamel trees near Angangueo,
Mexico. There is a reverse migration in the spring.[22][23] It has Mexico.
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.[24] Spectacular large-scale migrations associated with the
monsoon are seen in peninsular India.[25] Migrations have been studied in more recent times using
wing tags and also using stable hydrogen isotopes.[26][27]

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.[28][29] Many migratory butterflies live in semi-arid areas where breeding
seasons are short.[30] The life histories of their host plants also influence butterfly behaviour.[31]

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.[32]
The Melissa Arctic (Oeneis
Mating pair of spotted fritillaries on
melissa) overwinters twice as
greater pignut
a caterpillar.[33] Butterflies Life cycle of the monarch butterfly
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.[34]

Courtship is often aerial and often involves pheromones. Butterflies then land on the ground or on a
perch to mate.[16] 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.[35] The

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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.[36]

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;
The male small skipper (Thymelicus
when the female dies, a partially developed larva emerges from
sylvestris) has pheromone-releasing
her abdomen.[37] "sex brands" (dark line) on the
upperside of its forewings.
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
Eggs of black-veined white (Aporia A butterfly laying eggs underneath
funnel-shaped openings at
crataegi) on apple leaf the leaf
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.[37]

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.[38]
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.[39]

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.[40] 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.[41]

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.[42] Some temperate region butterflies, such as the Camberwell beauty, lay their eggs in the
spring and have them hatch in the summer.[43]

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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,[44] while lycaenids
such as Liphyra brassolis are myrmecophilous, eating ant
larvae.[45]

Some larvae, especially those


of the Lycaenidae, form
Aposematic caterpillar of Papilio
mutual associations with
machaon, in threat pose
ants. They communicate with
the ants using vibrations that
are transmitted through the
substrate as well as using chemical signals.[46][47] The ants
provide some degree of protection to these larvae and they in turn
Mutualism: ant tending a lycaenid
gather honeydew secretions. Large blue (Phengaris arion)
caterpillar, Catapaecilma major
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.[48]

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.[49] Development of butterfly Cryptic countershaded caterpillar of
wing patterns begins by the last larval instar. a hawkmoth, Ceratomia amyntor

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.[12] 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.[50] 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.[51] 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.[12]

Pupa

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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.[37] 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.[12]
Chrysalis of gulf fritillary
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.[52]

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 and do not feed as adults.[12]

Many Heliconius butterflies also use their proboscis to feed on pollen;[53] 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.[54]

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.[12]

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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.[12] 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.[55]

Pattern formation

The colorful 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 color 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 color 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.[56]

Behaviour

Butterflies feed primarily on nectar from flowers. Some also


derive nourishment from pollen,[57] 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.[58] Flower
constancy has been observed for at least one species of
butterfly.[59]
An Australian painted lady feeding
on a flowering shrub
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.[60]

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.[61]

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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.[62] Many butterflies use chemical signals, pheromones;
some have specialized scent scales (androconia) or other structures (coremata or "hair pencils" in the
Danaidae).[63] 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.[64] Colour
vision may be widespread but has been demonstrated in only a few species.[65][66] Some butterflies
have organs of hearing and some species make stridulatory and clicking sounds.[67]

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
Heteronympha merope taking off
wings edgewise to the sun.[68] 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.[69]

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.[70]

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 Braconid parasitoidal wasp
been used as biological controls of pest butterflies like the large (Apanteles species) cocoons
attached to lime butterfly (Papilio
white butterfly.[71] When the small cabbage white was
demoleus) caterpillar
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

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was thus regained.[72] 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.[73]
Predators of butterflies include ants, spiders, wasps, and birds.[74]

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.[73] 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.[75]

Endangered species

Queen Alexandra's birdwing 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.[76]

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

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
Heliconius warns off these plant toxins and use them
predators with Müllerian instead in their own defence.[78]
[77] Giant swallowtail caterpillar everting
mimicry. These defence mechanisms are its osmeterium in defence; it is also
effective only if they are well mimetic, resembling a bird dropping.
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.[79] The
common Mormon of India has female morphs which imitate the unpalatable red-bodied swallowtails,
the common rose and the crimson rose.[80] 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.[79]

Camouflage is found in many butterflies. Some like the oakleaf butterfly and autumn leaf are
remarkable imitations of leaves.[81] As caterpillars, many defend themselves by freezing and
appearing like sticks or branches.[82] Others have deimatic behaviours, such as rearing up and waving
their front ends which are marked with eyespots as if they were snakes.[83] Some papilionid
caterpillars such as the giant swallowtail (Papilio cresphontes) resemble bird droppings so as to be
passed over by predators.[84] Some caterpillars have hairs and bristly structures that provide
protection while others are gregarious and form dense aggregations.[79] Some species are
myrmecophiles, forming mutualistic associations with ants and gaining their protection.[85]
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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.[86]

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.[87][88]
Many butterflies have eyespots on the wings; these too may deflect
attacks, or may serve to attract mates.[52][89] Eyespots of speckled wood
(Pararge aegeria) distract
Auditory defences can also be used, which in the case of the grizzled predators from attacking the
skipper refers to vibrations generated by the butterfly upon expanding head. This insect can still fly
its wings in an attempt to communicate with ant predators.[90] with a damaged left
hindwing.
Many tropical butterflies have seasonal forms for dry and wet
seasons.[91][92] These are switched by the hormone ecdysone.[93] 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.[94][95][89]

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.[96]

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.[97][98]

In culture

In art and literature

Butterflies have appeared in art from 3500 years ago in ancient


Egypt.[99] 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.
Ancient Egyptian relief
The close association of butterflies with fire and warfare persisted into
sculpture, 26th dynasty,
the Aztec civilisation; evidence of similar jaguar-butterfly images has
Thebes. c. 664–525 BC
been found among the Zapotec and Maya civilisations.[100]

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.[101] The Norwegian naturalist Kjell Sandved compiled a photographic
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Butterfly Alphabet containing all 26 letters and


the numerals 0 to 9 from the wings of
butterflies.[102]

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
A butterfly in the coat
forelegs of the larva, or as suggesting a face with
of arms of Perho
protruding nose and chin.[2] Eric Carle's
children's book The Very Hungry Caterpillar
Butterfly and Chinese portrays the larva as an extraordinarily
wisteria, by Xü Xi. Early hungry animal, while also teaching children
Song Dynasty, c. 970 how to count (to five) and the days of the
week.[2]

A butterfly appeared in one of Rudyard Kipling's Just So Stories, "The


Butterfly that Stamped".[103]

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.[104]

Madam Butterfly is a 1904 opera by Giacomo Puccini about a romantic Alice meets the caterpillar.
young Japanese bride who is deserted by her American officer husband Illustration by Sir John
soon after they are married. It was based on John Luther Long's short Tenniel in Lewis Carroll's
story written in 1898.[105] Alice in Wonderland, c.
1865

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.[106]

Diderot's Encyclopédie cites butterflies as a symbol for the soul. A


Roman sculpture depicts a butterfly exiting the mouth of a dead man,
Der Schmetterlingsjäger
(The butterfly hunter)
representing the Roman belief that the soul leaves through the
painting by Carl Spitzweg, mouth.[107] In line with this, the ancient Greek word for "butterfly" is
1840 ψυχή (psȳchē), which primarily means "soul" or "mind".[108] According
to Mircea Eliade, some of the Nagas of Manipur claim ancestry from a
butterfly.[109] In some cultures, butterflies symbolise rebirth.[110] The
butterfly is a symbol of being transgender, because of the transformation from caterpillar to winged
adult.[111] In the English county of Devon, people once hurried to kill the first butterfly of the year, to
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avoid a year of bad luck.[112] In the Philippines, a lingering black


butterfly or moth in the house is taken to mean a death in the
family.[113] Several American states have chosen an official state
butterfly.[114]

Collecting, recording, and rearing

"Collecting" means preserving dead specimens, not keeping A serving tray decorated with
[115][116] butterfly wings
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.[2] 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.[2]

Butterflies and moths can be reared for recreation or for release.[117]

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,[118] and is inspiring nanotechnology research to
produce paints that do not use toxic pigments and the development of new display technologies.[119]

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External links
Papilionoidea on the Tree of Life (http://tolweb.org/Papilionoidea/12027)
Lamas, Gerardo (1990). "An Annotated List of Lepidopterological Journals" (https://web.archive.or
g/web/20160827214204/http://lepidopteraresearchfoundation.org/journals/29/PDF29/29-092.pdf)
(PDF). Journal of Research on the Lepidoptera. 29 (1–2): 92–104. Archived from the original (htt
p://lepidopteraresearchfoundation.org/journals/29/PDF29/29-092.pdf) (PDF) on 27 August 2016.
Retrieved 22 November 2012.
Rhopalocera (https://web.archive.org/web/20170211080551/http://insectoid.info/insecta/lepidopter
a/rhopalocera-sub/) at insectoid.info

Regional lists
North America (http://www.butterfliesandmoths.org/)
Africa (http://abdb-africa.org/): Ghana (http://ghanabutterflies.uw.hu/)
Asia: Singapore (https://web.archive.org/web/20091028083659/http://www.geocities.com/RainFor
est/Vines/2382/sgchecklist.htm) Israel (http://www.nature-of-oz.com/israelsspecieslist1.htm) Indo-
China (http://yutaka.it-n.jp/) Sulawesi (http://www.repository.naturalis.nl/document/46743)
(Southeastern Sulawesi (http://www.flmnh.ufl.edu/butterflies/neotropica/sulawesi/checklist.html))
Turkey (http://www.adamerkelebek.org/)

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