Butterfly

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For other uses, see Butterfly (disambiguation).

Butterflies

Charaxes brutus natalensis in Dar

es Salaam, Tanzania
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
(unranked): Rhopalocera
Subgroups
 Superfamily
Hedyloidea:
o Hedylidae
 Superfamily
Hesperioidea:
o Hesperiidae
 Superfamily
Papilionoidea:
o Papilionidae
o Pieridae
o Nymphalidae
o Lycaenidae
o Riodinidae
Spider lily and butterfly Papilio xuthus Linnaeus 1767

A butterfly is any of several groups of mainly day-flying insects of the order Lepidoptera, the
butterflies and moths. Like other holometabolous insects, butterflies' life cycle consists of four
parts, egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly
coloured wings, and conspicuous, fluttering flight. Butterflies comprise the true butterflies
(superfamily Papilionoidea), the skippers (superfamily Hesperioidea) and the moth-butterflies
(superfamily Hedyloidea). All the many other families within the Lepidoptera are referred to as
moths.

Butterflies exhibit polymorphism, mimicry and aposematism. Some, like the Monarch, will
migrate over long distances. Some butterflies have evolved symbiotic and parasitic relationships
with social insects such as ants. Butterflies are important ecologically as agents of pollination.
The caterpillars of some butterflies eat harmful insects. A few species are pests because in their
larval stages they can damage domestic crops or trees. Culturally, butterflies are a popular motif
in the visual and literary arts.

Contents
[hide]

 1 Life cycle
o 1.1 Egg
o 1.2 Caterpillars
 1.2.1 Wing development
o 1.3 Pupa
o 1.4 Adult or imago
 2 External morphology
o 2.1 Scales
 3 Polymorphism
o 3.1 Mimicry
o 3.2 Seasonal polyphenism
 4 Habits
 5 Flight
o 5.1 Migration
  6 Defense
 7 Notable species
 8 In culture
o 8.1 Art
o 8.2 Symbolism
o 8.3 Technological inspiration
 9 See also
 10 Field guides to butterflies
 11 Cited references
 12 Other references
 13 External links
o 13.1 General interest
o 13.2 Regional lists
 14 Literature
o 14.1 Images/Movies

Life cycle

Mating Common Buckeye Butterflies

It is a popular belief that butterflies have very short life spans. However, 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.[1]

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.

Egg
Egg of Ariadne merione

Butterfly eggs consist of 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.

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.

Eggs are usually 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.

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. 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 as Spalgis 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.[2][3] The ants provide some degree of protection to these larvae and they
in turn gather honeydew secretions.

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 epidermis beneath, 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.

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 smelly 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 helps making them unpalatable to birds and
other predators. Such unpalatibility is advertised using bright red, orange, black or white warning
colours. 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.[4]
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.

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.

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 blood and letting 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.
External morphology
Main article: Glossary of Lepidopteran terms

Parts of an adult
butterfly Butterflies have two antennae, 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 photonic crystal nature of the scales.[5][6][7] The scales cling somewhat loosely to the wing
and come off easily without harming the butterfly.

Photograp
hic and
light
microscop
ic images

Zoomed-out view of Closeup of the High magnification

an Inachis io. scales of the same of the coloured
specimen. scales (probably a
different species).

Electron
microscop
ic images

Microstructure of a
A patch of wing Scales close up A single scale
scale
Magnifica
tion Approx. x50 Approx. x200 x1000 x5000

Polymorphism
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).[8]

Genetic abnormalities such as gynandromorphy also occur from time to time. In addition many
butterflies are infected by Wolbachia and infection by the bacteria can lead to the conversion of
males into females[9] or the selective killing of males in the egg stage.[10]

Mimicry

The Heliconius butterflies from the tropics of the Western Hemisphere are the classical model
for Müllerian mimicry.[11]

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 an automimicry 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.[12][13]

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.[14] 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.[15] 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.[16]

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, oftentimes nonexistent, ventral eyespots. Larvae that develop in
hot, wet conditions develop into wet-season adults where as those growing in the transition from
the wet to the dry season, when the temperature is declining, develop into dry-season adults.[17]
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.[18]

Habits

Antennae shape in the lepidoptera from C. T. Bingham (1905)

The Australian painted lady feeding on a flowering shrub

Butterflies feed primarily on nectar from flowers. Some also derive nourishment from pollen,[19]
tree sap, rotting fruit, dung[20], decaying flesh[21], 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 the Hymenoptera. They are however capable of moving pollen over
greater distances.[22] Within the Lepidoptera, the Hawkmoths and the Noctuidae are dominant as
pollinators.[23]

As adults, butterflies consume only liquids and these are sucked by means of their proboscis.
They feed on nectar from flowers and also sip water from damp patches. This they do for water,
for energy from sugars in nectar and for sodium and other minerals which are vital for their
reproduction. Several species of butterflies need more sodium than provided by nectar. They are
attracted to sodium in salt and they sometimes land on people, attracted by human sweat. Besides
damp patches, 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 are provided as a nuptial gift along with the
spermatophore during mating.[24]

Butterflies sense the air for scents, wind and nectar using their antennae. 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 sensillae. A
butterfly's sense of taste is coordinated by chemoreceptors on the tarsi, 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.[25] 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.[26] Color vision
may be widespread but has been demonstrated in only a few species.[27][28]

Some butterflies have organs of hearing and some species are also known to make stridulatory
and clicking sounds.[29]
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.[30]

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.[31]

Flight

Geitoneura klugii 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 using Vanessa
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.[32]

Migration
The Monarch butterfly migrates large distances
Main article: Lepidoptera migration
See also Insect migration

Many butterflies migrate over long distances. Particularly famous migrations are those of the
Monarch butterfly from Mexico to northern USA and southern Canada, a distance of about 4000
to 4800 km (2500–3000 miles). Other well known migratory species include the Painted Lady
and several of the Danaine butterflies. Spectacular and large scale migrations associated with the
Monsoons are seen in peninsular India.[33] Migrations have been studied in more recent times
using wing tags and also using stable hydrogen isotopes.[34][35]

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.[36]

It is suggested that most migratory butterflies are those that belong to semi-arid areas where
breeding seasons are short.[37] The life-histories of their host plants also influence the strategies of
the butterflies.[38]

Defense

The wings of a butterfly (Leopard Lacewing Cethosia cyane) become increasingly damaged as it
ages, and do not repair
Butterflies are threatened in their early stages by parasitoids and in all stages by predators,
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.[39] 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 be
mimicked by other butterflies. These mimetic forms are usually restricted to the females.

Eyespots on the wings of this butterfly are part of the animal's defense

Cryptic coloration is found in many butterflies. Some like the oakleaf butterfly are remarkable
imitations of leaves.[40] 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.[41]

Eyespots and tails are found in many lycaenid butterflies and these 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.[42]

A butterfly's hind wings are thought to allow the butterfly to take swift, tight turns to evade