For other uses, see Butterfly (disambiguation).

Butterflies are part of the class of Insects in the Moth order Lepidoptera. Adult butterflies have
large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises the true
butterflies (superfamilyPapilionoidea), the skippers (superfamily Hesperioidea) and the moth-
butterflies (superfamily Hedyloidea). Butterfly fossils date to the mid Eoceneepoch, 4050 million
years ago.
[1]

Butterflies exhibit polymorphism, mimicry and aposematism. Some, like theMonarch,
will migrate over long distances. Some butterflies have parasitic relationships with organisms
including protozoans, flies, ants, other invertebrates, and vertebrates.
[2]

[3]
Some species are pests
because in their larval stages they can damage domestic crops or trees; however, some species are
agents of pollination of some plants, and caterpillars of a few butterflies (e.g., Harvesters) eat
harmful insects. Culturally, butterflies are a popular motif in the visual and literary arts.
Etymology
The Oxford English Dictionary derives the word from a combination of butter and fly. It adds: "The
reason of the name is unknown", and refers to Hensleigh Wedgwood, who "points out a Dutch
synonym boterschijte in Kilian, which suggests that the insect was so called from the appearance of
its excrement".
[4]

Donald Ringe writes that the name is derived from Middle English buterflie, butturflye, boterflye,
from Old English butorfloge, buttorfloge, buterfloge, perhaps a compound of butor (beater),
mutation of batan (to beat), and floge (fly).
[5]

Life cycle
The face of a Dryas iulia, more commonly known as a Julia butterfly.Butterflies in their adult stage can live
from a week to nearly a year depending on the species. Many species have long larval life stages
while others can remain dormant in their pupal or egg stages and thereby survive winters.
[6]

The number of generations per year varies fromtemperate to tropical regions with tropical regions
showing a trend towards multivoltinism.
Eggs

Egg of Ariadne merione
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 and moth eggs vary greatly in size between species, but they are all either spherical or
ovate.
[citation needed]

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 is unknown and is a suitable subject for research. The same glue
is produced by a pupa to secure the setae of the cremaster. This glue is so hard that the silk pad, to
which the setae are glued, cannot be separated.
[citation needed]

Eggs are almost invariably laid on plants. Each species of butterfly has its own hostplant 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.
[citation needed]

The egg stage lasts a few weeks in most butterflies but eggs laid close to winter, especially in
temperate regions, go through adiapause (resting) stage, and the hatching may take place only in
spring. Other butterflies may lay their eggs in the spring and have them hatch in the summer. These
butterflies are usually northern species, such as the Mourning Cloak (Camberwell Beauty) and
the Large and Small Tortoiseshell butterflies.
[

Caterpillars

Caterpillars of Junonia coenia.
Butterfly larvae, or caterpillars, consume plant leaves and spend practically all of their time in search
of food. Although most caterpillars are herbivorous, a few species such asSpalgis epius and Liphyra
brassolis are entomophagous (insect eating).
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.
[7][8]
The ants provide some degree of protection to these larvae and they in
turn gather honeydew secretions. Others such as Phengaris arioncommunicate with ants to form a
parasitic relationship.
[9]

Caterpillars mature through a series of stages called instars. Near the end of each instar, the larva
undergoes a process called apolysis, in which the cuticle, a tough outer layer made of a mixture
of chitin and specialized proteins, is released from the softer epidermisbeneath, and the epidermis
begins to form a new cuticle beneath. At the end of each instar, the larva moults the old cuticle, and
the new cuticle expands, before rapidly hardening and developing pigment. Development of butterfly
wing patterns begins by the last larval instar.
Butterfly caterpillars have three pairs of true legs from the thoracic segments and up to 6 pairs
of prolegs arising from the abdominal segments. These prolegs have rings of tiny hooks called
crochets that help them grip the substrate.
[10]

Some caterpillars have the ability to inflate parts of their head to appear snake-like. Many have false
eye-spots to enhance this effect. Some caterpillars have special structures called osmeteria which
are everted to produce foul-smelling chemicals. These are used in defense.
Host plants often have toxic substances in them and caterpillars are able to sequester these
substances and retain them into the adult stage. This makes them unpalatable to birds and other
predators. Such unpalatibility is advertised using bright red, orange, black or white warning colours,
a practice known as aposematism. The toxic chemicals in plants are often evolved specifically to
prevent them from being eaten by insects. Insects in turn develop countermeasures or make use of
these toxins for their own survival. This "arms race" has led to the coevolution of insects and their
host plants.
[11]

Wing development

Last instar wing disk,Junonia coenia

Detail of a butterfly wing
Wings or wing pads are not visible on the outside of the larva, but when larvae are dissected, tiny
developing wing disks can be found on the second and third thoracic segments, in place of the
spiracles that are apparent on abdominal segments. Wing disks develop in association with a
trachea that runs along the base of the wing, and are surrounded by a thin peripodial membrane,
which is linked to the outer epidermis of the larva by a tiny duct.
Wing disks are very small until the last larval instar, when they increase dramatically in size, are
invaded by branching tracheae from the wing base that precede the formation of the wing veins, and
begin to develop patterns associated with several landmarks of the wing.
Near pupation, the wings are forced outside the epidermis under pressure from the hemolymph, and
although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval
cuticle they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the
wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled
without damage to the wings.
[citation needed]

Pupa

Chrysalis of Gulf Fritillary
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 of a suitable pupation site,
often the underside of a leaf.
The larva transforms into a pupa (or chrysalis) by anchoring itself to a substrate and moulting for the
last time. The chrysalis is usually incapable of movement, although some species can rapidly move
the abdominal segments or produce sounds to scare potential predators.
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
color pattern are marked by changes in the expression of particular transcription factors in the early
pupa.
[citation needed]

Adult or imago
The adult, sexually mature, stage of the insect is known as the imago. As Lepidoptera, butterflies
have four wings that are covered with tiny scales (see photo). The fore and hindwings are not
hooked together, permitting a more graceful flight. An adult butterfly has six legs, but in
the nymphalids, the first pair is reduced. 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 let them dry, during which time it is extremely vulnerable to predators. Some
butterflies' wings may take up to three hours to dry while others take about one hour. Most butterflies
and moths will excrete excess dye after hatching. This fluid may be white, red, orange, or in rare
cases, blue.
[citation needed]

External morphology
Main article: Glossary of Lepidopteran terms

Parts of an adult butterfly

Butterflies have twoantennae, two compound eyes, and a proboscis
Adult butterflies have four wings: a forewing and hindwing on both the left and the right side of the
body. The body is divided into three segments: the head, thorax, and the abdomen. They have
two antennae, two compound eyes, and a proboscis.
Scales
Butterflies are characterized by their scale-covered wings. The coloration of butterfly wings is
created by minute scales. These scales are pigmented with melanins that give them blacks and
browns, but blues, greens, reds and iridescence are usually created not by pigments but the
microstructure of the scales. This structural coloration is the result of coherent scattering of light by
the scales.
[12][13][14]
The scales cling somewhat loosely to the wing and come off easily without harming
the butterfly.
Photographi
c and light
microscopic
images




Zoomed-out view of anInachis
io.
Closeup of the scales of the same
specimen.
High magnification of the
coloured scales (probably a
different species).

Electron
microscopic
images



A patch of wing Scales close up A single scale Microstructure of a scale

Magnificatio
n Approx. 50 Approx. 200 1000 5000



Polymorphism
Main article: Polymorphism (biology)
Many adult butterflies exhibit polymorphism, showing differences in appearance. These variations
include geographic variants and seasonal forms. In addition many species have females in multiple
forms, often with mimetic forms. Sexual dimorphism in coloration and appearance is widespread in
butterflies. In addition many species show sexual dimorphism in the patterns of ultraviolet reflectivity,
while otherwise appearing identical to the unaided human eye. Most of the butterflies have a sex-
determination system that is represented as ZW with females being the heterogametic sex (ZW) and
males homogametic (ZZ).
[15]

Genetic abnormalities such as gynandromorphy also occur from time to time. In addition many
butterflies are infected byWolbachia and infection by the bacteria can lead to the conversion of
males into females
[16]
or the selective killing of males in the egg stage.
[17]

Mimicry

The Heliconius butterflies from the tropics of the Western Hemisphere are the classical model for Mllerian
mimicry.
[18]

Batesian and Mullerian mimicry in butterflies is common. Batesian mimics imitate other species to
enjoy the protection of an attribute they do not share, aposematism in this case. The Common
Mormon of India has female morphs which imitate the unpalatable red-bodied swallowtails,
the Common Rose and the Crimson Rose. Mullerian mimicry occurs when aposematic species
evolve to resemble each other, presumably to reduce predator sampling rates,
the Heliconius butterflies from the Americas being a good example.
Wing markings called eyespots are present in some species; these may have anautomimicry role for
some species. In others, the function may be intraspecies communication, such as mate attraction.
In several cases, however, the function of butterfly eyespots is not clear, and may be an evolutionary
anomaly related to the relative elasticity of the genes that encode the spots.
[19][20]

Seasonal polyphenism
Many of the tropical butterflies have distinctive seasonal forms. This phenomenon is
termed seasonal polyphenism and the seasonal forms of the butterflies are called the dry-season
and wet-season forms. How the season affects the genetic expression of patterns is still a subject of
research.
[21]
Experimental modification by ecdysone hormone treatment has demonstrated that it is
possible to control the continuum of expression of variation between the wet and dry-season
forms.
[22]
The dry-season forms are usually more cryptic and it has been suggested that the
protection offered may be an adaptation. Some also show greater dark colours in the wet-season
form which may have thermoregulatory advantages by increasing ability to absorb solar radiation.
[23]

Bicyclus anynana is a species of butterfly that exhibits a clear example of seasonal polyphenism.
These butterflies, endemic to Africa, have two distinct phenotypic forms that alternate according to
the season. The wet-season forms have large, very apparent ventral eyespots whereas the dry-
season forms have very reduced, often nonexistent, ventral eyespots. Larvae that develop in hot,
wet conditions develop into wet-season adults whereas those growing in the transition from the wet
to the dry season, when the temperature is declining, develop into dry-season adults.
[24]
This
polyphenism has an adaptive role in B. anynana. In the dry-season it is disadvantageous to have
conspicuous eyespots because B. anynana blend in with the brown vegetation better without
eyespots. By not developing eyespots in the dry-season they can more easily camouflage
themselves in the brown brush. This minimizes the risk of visually mediated predation. In the wet-
season, these brown butterflies cannot as easily rely on cryptic coloration for protection because the
background vegetation is green. Thus, eyespots, which may function to decrease predation, are
beneficial for B. anynana to express.
[25]

Habits

Antennal shape in the Lepidopterafrom C. T. Bingham (1905)

The Australian painted lady feeding on a flowering shrub
Butterflies feed primarily on nectar from flowers. Some also derive nourishment frompollen,
[26]
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 although in general they do not carry as much
pollen load as bees. They are however capable of moving pollen over greater distances.
[27]
Flower
constancy has been observed for at least one species of butterfly.
[28]

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 thespermatophore, during mating.
[29]

Butterflies use their antennae to sense the air for wind and scents. The antennae come in various
shapes and colours; the hesperids have a pointed angle or hook to the antennae, while most other
families show knobbed antennae. The antennae are richly covered with sensory organs known
as sensillae. A butterfly's sense of taste, 200 times stronger than humans,
[30]
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.
[31]
Many
butterflies use chemical signals, pheromones, and specialized scent scales (androconia) and other
structures (coremata or "hair pencils" in the Danaidae) are developed in some species.
Vision is well developed in butterflies and most species are sensitive to the ultraviolet spectrum.
Many species show sexual dimorphism in the patterns of UV reflective patches.
[32]
Color vision may
be widespread but has been demonstrated in only a few species.
[33][34]

Some butterflies have organs of hearing and some species are also known to make stridulatory and
clicking sounds.
[35]


Monarch butterflies
Many butterflies, such as the Monarch butterfly, are migratory and capable of long distance flights.
They migrate during the day and use the sun to orient themselves. They also perceive polarized light
and use it for orientation when the sun is hidden.
[36]

Many species of butterfly maintain territories and actively chase other species or individuals that may
stray into them. Some species will bask or perch on chosen perches. The flight styles of butterflies
are often characteristic and some species have courtship flight displays. Basking is an activity which
is more common in the cooler hours of the morning. Many species will orient themselves to gather
heat from the sun. Some species have evolved dark wingbases to help in gathering more heat and
this is especially evident in alpine forms.
[37]

Flight

Heteronympha merope taking off
See also Insect flight
Like many other members of the insect world, the lift generated by butterflies is more than what
can be accounted for by steady-state, non-transitory aerodynamics. Studies usingVanessa
atalanta in a windtunnel show that they use a wide variety of aerodynamic mechanisms to
generate force. These include wake capture, vortices at the wing edge, rotational mechanisms
and Weis-Fogh 'clap-and-fling' mechanisms. The butterflies were also able to change from one
mode to another rapidly.
[38]

Migration
Main article: Lepidoptera migration
See also Insect migration
Many butterflies migrate over long distances.

Monarch migration route

Overwintering monarchs cluster onoyamel trees in a preserve outside ofAngangueo, Michoacan,
Mexico; one tree is completely covered in butterflies.

Monarch Butterflies in Pacific Grove, California
The eastern population of monarchs migrates hundreds to thousands of miles to
overwintering sites in Mexico. There is a northward migration in the spring.
[39][40]

Other well known migratory species include thePainted Lady and several of
the Danainebutterflies. Spectacular and large scale migrations associated with
the Monsoons are seen in peninsular India.
[41]
Migrations have been studied in more recent
times using wing tags and also using stable hydrogen isotopes.
[42][43]

Butterflies have been shown to navigate using time compensated sun compasses. They can
see polarized light and therefore orient even in cloudy conditions. The polarized light in the
region close to the ultraviolet spectrum is suggested to be particularly important.
[44]

It is suggested that most migratory butterflies are those that belong to semi-arid areas where
breeding seasons are short.
[45]
The life-histories of their host plants also influence the
strategies of the butterflies.
[46]

Defense

Wings of a butterfly (Leopard Lacewing Cethosia cyane) become increasingly damaged as they age,
and do not repair
See also Defense in insects
Butterflies are threatened in their early stages by parasitoids and in all stages
bypredators, diseases and environmental factors. They protect themselves by a variety
of means.
Chemical defenses are widespread and are mostly based on chemicals of plant origin.
In many cases the plants themselves evolved these toxic substances
as protection against herbivores. Butterflies have evolved mechanisms to sequester
these plant toxins and use them instead in their own defense.
[47]
These defense
mechanisms are effective only if they are also well advertised and this has led to the
evolution of bright colours in unpalatable butterflies. This signal may bemimicked by
other butterflies. These mimetic forms are usually restricted to the females.
Cryptic coloration is found in many butterflies. Some like the Oakleaf butterfly
and Autumn Leaf are remarkable imitations of leaves.
[48]
As caterpillars, many defend
themselves by freezing and appearing like sticks or branches. Some papilionid
caterpillars resemble bird dropping in their early instars. Some caterpillars have hairs
and bristly structures that provide protection while others are gregarious and form dense
aggregations. Some species also form associations with ants and gain their protection
(See Myrmecophile).
Behavioural defenses include perching and wing positions to avoid being conspicuous.
Some female Nymphalid butterflies are known to guard their eggs from parasitoid
wasps.
[49]


Eyespots on wings of Speckled Wood, Pararge aegeria, appear to distract predators from
attacking the head. The left hind wing has been badly damaged by birds, but the insect is alive and
able to fly.
Eyespots and tails are found in many lycaenid butterflies. It is thought that their function
is to divert the attention of predators from the more vital head region. An alternative
theory is that these cause ambush predators such as spiders to approach from the
wrong end and allow for early visual detection.
[50]

A butterfly's hind wings are thought to allow them to take swift, tight turns to evade
predators.
[51]

Notable species

Rusty-tipped Page (Siproeta epaphus), Butterfly World (Florida)
There are between 15,000 and 20,000 species of butterflies worldwide. Some well-
known species from around the world include:
Swallowtails and Birdwings, Family Papilionidae
Common Yellow Swallowtail, Papilio machaon
Spicebush Swallowtail, Papilio troilus
Lime Butterfly, Papilio demoleus
Ornithoptera genus (Birdwings; the largest butterflies)
Whites and Yellows, Family Pieridae
Small White, Pieris rapae
Large White, Pieris brassicae
Green-veined White, Pieris napi
Common Jezebel, Delias eucharis
Blues and Coppers or Gossamer-Winged Butterflies, Family Lycaenidae
Xerces Blue, Glaucopsyche xerces (extinct)
Karner Blue, Lycaeides melissa samuelis (endangered)
Red Pierrot, Talicada nyseus
Metalmark butterflies, Family Riodinidae
Duke of Burgundy, Hamearis lucina
Plum Judy, Abisara echerius
Brush-footed butterflies, Family Nymphalidae
Painted Lady, or Cosmopolitan, Vanessa cardui
Monarch butterfly, Danaus plexippus
Morpho genus
Speckled Wood, Pararge aegeria
Skippers, Family Hesperiidae
Mallow Skipper, Carcharodus alceae
Zabulon Skipper, Poanes zabulon
In culture
Art
Artistic depictions of butterflies have been used in many cultures including Egyptian
hieroglyphs 3500 years ago.
[52]

In the ancient Mesoamerican city of Teotihuacan, the brilliantly colored image of the
butterfly was carved into many temples, buildings, jewelry, and emblazoned on incense
burners in particular. The butterfly was sometimes depicted with the maw of ajaguar and
some species were considered to be the reincarnations of the souls of dead warriors.
The close association of butterflies to fire and warfare persisted through to the Aztec
civilization and evidence of similar jaguar-butterfly images has been found among
the Zapotec, and Mayan civilizations.
[53]


A serving tray decorated with butterfly wings
Today, butterflies are widely used in various objects of art and jewelry: mounted in
frames, embedded in resin, displayed in bottles, laminated in paper, and used in some
mixed media artworks and furnishings.
[54]
Butterflies have also inspired the "butterfly
fairy" as an art and fictional character, including in the Barbie Mariposa film.
Symbolism
According to Kwaidan: Stories and Studies of Strange Things, by 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 guestroom and perches behind the bamboo screen, the person whom you most
love is coming to see you. However, 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.
[55]

The Russian word for "butterfly", (bbochka), also means "bow tie". It is a
diminutive of "baba" or "babka" (= "woman, grandmother, cake"), whence also
"babushka" = "grandmother".
The ancient Greek word for "butterfly" is (psch), which primarily means "soul" or
"mind".
[56]

According to Mircea Eliade's Encyclopedia of Religion, some of
the Nagas of Manipur trace their ancestry from a butterfly.
[57]


Butterfly and Chinesewisteriaflowers, by X Xi (c.886c.975), painted around 970 during the
early Song Dynasty.
In Chinese culture, two butterflies flying together symbolize love. Also, Butterfly
Lovers is a famous Chinese folktale. The Taoist philosopher, Zhuangzi, once had a
dream about being a butterfly that flew without care about humanity; however; when he
awoke and realized that it was just a dream, he thought to himself, "Was I before a man
who dreamt about being a butterfly, or am I now a butterfly who dreams about being a
man?"
In some old cultures, butterflies also symbolize rebirth after being inside a cocoon for a
period of time.

Der Schmetterlingsjger(The butterfly hunter) by Carl Spitzweg (1840), a depiction from the era of
butterfly collection.
Some people say that when a butterfly lands on you it means good luck.
[citation
needed]
However, inDevonshire, people would traditionally rush around to kill the first
butterfly of the year that they see, or else face a year of bad luck.
[58]
Also, in the
Philippines, a lingering black butterfly or moth in the house is taken to mean that
someone in the family has died or will soon die.
[59]

The idiom "butterflies in the stomach" is used to describe a state of nervousness.
In the NBC television show Kings, butterflies are the national symbol of the fictional
nation of Gilboa and a sign of God's favor.
Technological inspiration
Studies on the reflection and scattering of light by the scales on wings of swallowtail
butterflies led to the innovation of more efficient light-emitting diodes.
[60]

The structural coloration of butterflies is inspiring nanotechnology research to produce
paints that do not use toxic pigments and in the development of new display
technologies.
The discoloration and health of butterflies in butterfly farms, is now being studied for use
as indicators of air quality in several cities.
Citizen/Scientists
Butterfly counts are organized to assess the numbers and species of butterflies in a
given locale. Much of this work is organized and recorded by volunteers who share their
information with researchers.