Butterfly

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From Wikipedia, the free encyclopedia

For other uses, see Butterfly (disambiguation) and Butterflies (disambiguation).

Butterflies
 Temporal range: Cretaceous–
Present, 101.4–0 Ma

PreꞒ

Pg

Papilio machaon

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta
 Order: Lepidoptera

Suborder: Rhopalocera

Subgroups

 Superfamily Hedy
loidea:
 Hedyli
dae
 Superfamily Papil
ionoidea:
 Hesper
iidae
 Lycae
nidae
 Nymp
halidae
 Papilio
nidae
 Pierida
e
 Riodin
idae

Butterflies are winged insects from the lepidopteran suborder Rhopalocera,

characterized by large, often brightly coloured wings that often fold together when at
rest, and a conspicuous, fluttering flight. The group comprises
the superfamilies Hedyloidea (moth-butterflies in the Americas)
and Papilionoidea (all others). The oldest butterfly fossils have been dated to
the Paleocene, about 56 million years ago, though they likely originated in the
Late Cretaceous, about 101 million years ago.[1]

Butterflies have a four-stage life cycle, and like other holometabolous 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,
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.[citation needed]
Butterflies are often polymorphic, and many species make use
of camouflage, mimicry, and aposematism to evade their predators.[2] 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".[3]

Etymology

Possibly the original butter-fly.[4] A male brimstone

(Gonepteryx rhamni) in flight.
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 season while the
grass was growing.[4][5]

Paleontology
Further information: Prehistoric Lepidoptera
The earliest Lepidoptera fossils date to the Triassic-Jurassic boundary, around
200 million years ago.[6] 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).[7] Molecular clock estimates suggest that butterflies
originated sometime in the Late Cretaceous, but only significantly diversified during
the Cenozoic,[8][1] with one study suggesting a North American origin for the group.
[1]
The oldest American butterfly is the Late Eocene Prodryas persephone from
the Florissant Fossil Beds,[9][10] approximately 34 million years old.[11]

 Butterfly fossils
 

Prodryas persephone, a Late Eocene butterfly from the Florissant Fossil Beds,
1887 engraving

Lithopsyche antiqua, an Early Oligocene butterfly from the Bembridge Marls, Isle
of Wight, 1889 engraving
Taxonomy and phylogeny
Further information: Taxonomy of the Lepidoptera
Butterflies are scientifically classified in
the macrolepidopteran suborder clade Rhopalocera from the order Lepidoptera,
which also includes moths.[citation needed] 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.[12][13]

Butterfly families

Family Common name Characteristics Image

Small, brown, like geometrid

American moth-
Hedylidae moths; antennae not clubbed; long slim
butterflies
abdomen
 Small, darting flight; clubs on antennae
Hesperiidae Skippers
hooked backwards

Small, brightly coloured; often have

Blues, coppers,
Lycaenidae false heads with eyespots and small tails
hairstreaks
resembling antennae

Brush-footed or Usually have reduced forelegs, so

Nymphalidae four-footed appear four-legged; often brightly
butterflies coloured

Often have 'tails' on wings; caterpillar

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

Mostly white, yellow or orange; some

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

Often have metallic spots on wings;

Riodinidae Metalmarks often conspicuously coloured with
black, orange and blue

Biology
General description
Further information: Glossary of entomology terms and Comparison of butterflies and
moths

A zoomed in view of the wing scales on a Aglais io, or

peacock butterfly.
Butterfly adults are characterized by their four scale-covered wings, which give the
Lepidoptera their name (Ancient Greek λεπίς lepís, scale + πτερόν pterón, wing).
These scales give butterfly wings their colour: they are pigmented with melanins that
give them blacks and browns, as well as uric acid derivatives and flavones that give
them yellows, but many of the blues, greens, reds and iridescent colours are created
by structural coloration produced by the micro-structures of the scales and hairs.[14][15]
[16][17]

Butterfly antennal shapes, mainly clubbed, unlike

those of moths. Drawn by C. T. Bingham, 1905
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.[18]

Unlike butterflies, most moths (like Laothoe populi) fly

by night and hide by day.
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,[19] are
exceptions to these rules.[18][20]

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.[18] 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.[18]

Sexual dimorphism in Anthocharis cardamines

Male

Female

Many butterflies are sexually dimorphic. Most butterflies have the ZW sex-
determination system where females are the heterogametic sex (ZW) and males
homogametic (ZZ).[21]

Distribution and migration

See also: Lists of butterflies
of Australia (Tasmania, Victoria), Britain, India, Menorca, North America, Taiwan,
and Trinidad and Tobago
Further information: Lepidoptera migration, Insect migration, and Animal navigation
Butterflies are distributed worldwide except Antarctica, totalling some 18,500
species.[22] Of these, 775 are Nearctic; 7,700 Neotropical; 1,575 Palearctic;
3,650 Afrotropical; and 4,800 are distributed across the
combined Oriental and Australian/Oceania regions.[22] 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.[23]
 Monarch migration route

Overwintering monarchs cluster on oyamel trees

near Angangueo, Mexico.
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.[24][25] 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.[26] Spectacular large-
scale migrations associated with the monsoon are seen in peninsular India.
[27]
Migrations have been studied in more recent times using wing tags and also
using stable hydrogen isotopes.[28][29]

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.[30][31] Many migratory
butterflies live in semi-arid areas where breeding seasons are short.[32] The life
histories of their host plants also influence butterfly behaviour.[33]

Life cycle

Life cycle of the monarch 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.
 The Melissa Arctic (Oeneis melissa) overwinters twice as a caterpillar.[35] Butterflies
[34]

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.[36]

Mating pair of spotted fritillaries on greater pignut

The male small skipper (Thymelicus sylvestris)

has pheromone-releasing "sex brands" (dark line) on the upperside of its forewings.
Courtship is often aerial and often involves pheromones. Butterflies then land on the
ground or on a perch to mate.[18] 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.[37] 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.[38]

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.[39]

Eggs
 Eggs of black-veined white (Aporia crataegi)

on apple leaf A butterfly from the genus Euploea,

laying eggs underneath the leaf
Butterfly eggs are protected by a hard-ridged outer layer of shell, called the chorion.
This is lined with a thin coating of wax which prevents the egg from drying out before
the larva has had time to fully develop. Each egg contains a number of tiny funnel-
shaped openings at one end, called micropyles; the purpose of these holes is to
allow sperm to enter and fertilize the egg. Butterfly eggs vary greatly in size and
shape between species, but are usually upright and finely sculptured. Some species
lay eggs singly, others in batches. Many females produce between one hundred and
two hundred eggs.[39]

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.
[40]
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.[41]

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.[42] 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.[43]

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.[44] Some temperate region butterflies, such as
the Camberwell beauty, lay their eggs in the spring and have them hatch in the
summer.[45]
Caterpillar larva

Aposematic caterpillar of Papilio machaon, in threat

pose
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,[46] while lycaenids such
as Liphyra brassolis are myrmecophilous, eating ant larvae.[47]

Mutualism: ant tending

a lycaenid caterpillar, Catapaecilma major
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.[48][49] 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.[50]

Cryptic countershaded caterpillar of a

hawkmoth, Ceratomia amyntor
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.[51] Development of butterfly wing patterns
begins by the last larval instar.
Caterpillars have short antennae and several simple eyes. The mouthparts are
adapted for chewing with powerful mandibles and a pair of maxillae, each with a
segmented palp. Adjoining these is the labium-hypopharynx which houses a tubular
spinneret which is able to extrude silk.[14] 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.[52] 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.[53] 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.[14]

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 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.[39] 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.[14]

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.[54]

Adult

An adult Parthenos sylvia butterfly

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.[14]

Duration: 9 seconds.0:09Butterfly resting on flower at Magic Wings Conservatory

Many Heliconius butterflies also use their proboscis to feed on pollen;[55] 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.[56]

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.[14]

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.[14] 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. [57]

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.[58]

Mating
When the butterfly Bicyclus anynana is subjected to repeated inbreeding in the
laboratory, there is a dramatic decrease in egg hatching.[59] 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.[59] Although B.
anynana experiences inbreeding depression when forcibly inbred in the laboratory it
recovers within a few generation when allowed to breed freely.[60] 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.[60] 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

An Australian painted lady feeding on a flowering

shrub
Butterflies feed primarily on nectar from flowers. Some also derive nourishment
from pollen,[61] 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.[62] Flower constancy has been observed for at
least one species of butterfly.[63]

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.[64]

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.[65]

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.[66] Many butterflies use chemical
signals, pheromones; some have specialized scent scales (androconia) or other
structures (coremata or "hair pencils" in the Danaidae).[67] 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.[68] Colour vision
may be widespread but has been demonstrated in only a few species.[69][70] Some
butterflies have organs of hearing and some species make stridulatory and clicking
sounds.[71]

Heteronympha merope taking off

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.[72] 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.[73]

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.[74]

Ecology
Parasitoids, predators, and pathogens

Braconid parasitoidal wasp (Apanteles species)

cocoons attached to lime butterfly (Papilio demoleus) caterpillar
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.[75] 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.[76] 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.[77] Predators of butterflies
include ants, spiders, wasps, and birds.[78]

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.[77] 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.[79]

Endangered species
Queen Alexandra's birdwing, found in Papua New Guinea, 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.[80]

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
Further information: Defense in insects, Anti-predator adaptation, Mimicry,
and Seasonal polyphenism
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.[81] 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.
[82]
The common Mormon of India has female morphs which imitate the unpalatable
red-bodied swallowtails, the common rose and the crimson rose.[83] 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.[82]

Camouflage is found in many butterflies. Some like the oakleaf butterfly and autumn
leaf are remarkable imitations of leaves.[84] As caterpillars, many defend themselves
by freezing and appearing like sticks or branches.[85] Others
have deimatic behaviours, such as rearing up and waving their front ends which are
marked with eyespots as if they were snakes.[86] Some papilionid caterpillars such as
the giant swallowtail (Papilio cresphontes) resemble bird droppings so as to be
passed over by predators.[87] Some caterpillars have hairs and bristly structures that
provide protection while others are gregarious and form dense aggregations.[82] Some
species are myrmecophiles, forming mutualistic associations with ants and gaining
their protection.[88] 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.[89]

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.[90][91] Many butterflies have eyespots on the
wings; these too may deflect attacks, or may serve to attract mates.[54][92]

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.[93]

Many tropical butterflies have seasonal forms for dry and wet seasons.[94][95] These are
switched by the hormone ecdysone.[96] 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.[97][98][92]

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.[99]

[100]
Heliconius warns off predators with Müllerian mimicry.

Giant swallowtail caterpillar everting its osmeterium in defence; it is also mimetic,

resembling a bird dropping.

Eyespots of speckled wood (Pararge aegeria) distract predators from attacking

the head. This insect can still fly with a damaged left hindwing.
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.[101][102]

In culture
In art and literature

Ancient Egyptian relief sculpture, 26th

dynasty, Thebes. c. 664–525 BC
Butterflies have appeared in art from 3500 years ago in ancient Egypt.[103] 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.[104]

Alice meets the caterpillar. Illustration by Sir John

Tenniel in Lewis Carroll's Alice in Wonderland, c. 1865
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.[105] 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.[106]

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.[4] 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. [4]

A butterfly appeared in one of Rudyard Kipling's Just So Stories, "The Butterfly that
Stamped".[107]

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.[108]

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 John Luther Long's short story written in 1898.[109]

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 a swarm of butterflies appeared in Kyoto.
The people were frightened, thinking the apparition to be a portent of coming evil. [110]

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.[111] In line with this, the ancient Greek word for
"butterfly" is ψυχή (psȳchē), which primarily means "soul" or "mind".[112] According
to Mircea Eliade, some of the Nagas of Manipur claim ancestry from a butterfly.[113] In
some cultures, butterflies symbolise rebirth.[114] The butterfly is a symbol of
being transgender, because of the transformation from caterpillar to winged adult.
[115]
In the English county of Devon, people once hurried to kill the first butterfly of the
year, to avoid a year of bad luck.[116] 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.[117] Several American states have chosen an official state butterfly.[118]

Collecting, recording, and rearing

See also: Insect collecting
 A collection of butterflies and moths in the Manitoba
Museum, c. 2010
"Collecting" means preserving dead specimens, not keeping butterflies as pets.[119]
[120]
Collecting butterflies was once a popular hobby; it has now largely been replaced
by photography, recording, and rearing butterflies for release into the wild.[4][dubious – discuss]
[full citation needed]
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.[4]

Butterflies and moths can be reared for recreation or for release.[121]

In technology
Further information: Biomimetics
Study of the structural coloration of the wing scales of swallowtail butterflies has led
to the development of more efficient light-emitting diodes,[122] and is
inspiring nanotechnology research to produce paints that do not use toxic pigments
and the development of new display technologies.[123]

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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. PM
C 10250192. PMID 37188966.
External links

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 Papilionoidea 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 insectoid.info
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