Butterfly - Wikipedia, the free encyclopedia

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Butterfly
From Wikipedia, the free encyclopedia

Butterflies

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
very many other families within the Lepidoptera are
referred to as moths. Butterflies exhibit polymorphism,
mimicry and aposematism. Some migrate over long
distances. Some butterflies have evolved symbiotic and
parasitic relationships with social insects such as ants.
Butterflies are important economically as agents of
pollination. In addition, 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

1 Life cycle

1.1 Egg

1.2 Caterpillars

1.2.1 Wing development

1.3 Pupa
1.4 Adult or imago

2 External morphology

2.1 Scales

3 Polymorphism

3.1 Mimicry

3.2 Seasonal polyphenism

4 Habits
5 Flight

5.1 Migration

6 Defense
7 Notable species
8 In culture

8.1 Art

8.2 Symbolism

8.3 Technological inspiration

Charaxes brutus natalensis in Dar es

Salaam, Tanzania

Scientific classification
Kingdom:
Animalia
Phylum:
Arthropoda
Class:
Insecta
Order:
Lepidoptera
(unranked):
Rhopalocera
Subgroups

Superfamily Hedyloidea:

Hedylidae

Superfamily Hesperioidea:

Hesperiidae

Superfamily Papilionoidea:

Papilionidae

Pieridae

Nymphalidae

Lycaenidae

Riodinidae

Spider lily and butterfly Papilio

xuthus Linnaeus 1767

9 See also
10 Cited references
11 Other references

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12 Field guides to butterflies

13 External links

13.1 General interest

13.2 Regional lists
14 Literature

14.1 Images/Movies

Life cycle
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.

Mating Common Buckeye Butterflies

Egg
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
Egg of Ariadne merione
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
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

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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 softern epidermis beneath, and the epidermis begins
Caterpillars of Junonia coenia.
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
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.

Detail of a butterfly 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

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without damage to the wings.

Pupa

Last instar wing disk,

Junonia coenia

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
Chrysalis of Gulf Fritillary
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
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.

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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
Parts of an adult butterfly
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.
Butterflies have two antennae, two
compound eyes, and a proboscis

Scales on the wing give the

colours

Closeup of the scales of the

Inachis io

A Monarch butterfly with the

scales rubbed off a section of
the wing.

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

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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 dryseason 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
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]

Antennae shape in the lepidoptera

from C. T. Bingham (1905)

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

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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]

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The Australian painted lady feeding

on a flowering shrub

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 egglaying 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]
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
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]

Geitoneura klugii taking off

Migration

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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 (25003000 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
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.
The wings of a butterfly
(Leopard Lacewing Cethosia
cyane) become increasingly
damaged as it ages, and do
not repair

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]

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Eyespots on the wings of this

butterfly are part of the animal's
defense

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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 predators.
[43]

Notable species
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 or Yellows, Family Pieridae

Small White, Pieris rapae

Green-veined White, Pieris napi

Common Jezebel, Delias eucharis

Rusty-tipped Page (Siproeta

epaphus), Butterfly World (Florida)

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.[44] Today, butterflies are widely used in various objects of art and jewelry: mounted
in frame, embedded in resin, displayed in bottles, laminated in paper, and used in some mixed media
artworks and furnishings.[45] Butterflies have also inspired the "butterfly fairy" as an art and fictional
character, including in the Barbie Mariposa film.

Symbolism

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According to the Butterflies chapter in Kwaidan: Stories and Studies of Strange Things, by
Lafcadio Hearn, a butterfly is seen 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.[46]
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", "mind".[47]
According to Mircea Eliade's Encyclopedia of Religion, some of the Nagas of Manipur trace their
ancestry from a butterfly.[48]
In Chinese culture two butterflies flying together are a symbol of love.
Also a famous Chinese folk story called Butterfly Lovers. The Taoist
philosopher Zhuangzi once had a dream of being a butterfly flying
without care about humanity, however when he woke up and realized
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 into a new life
after being inside a cocoon for a period of time.
Jose Rizal delivered a speech in 1884 in a banquet and mentioned "the
Oriental chrysalis ... is about to leave its cocoon" comparing the
emergence of a "new Philippines" with that of butterfly
metamorphosis.[49] He has also often used the butterfly imagery in his
poems and other writings to express the Spanish Colonial Filipinos'
longing for liberty.[50] Much later, in a letter to Ferdinand Blumentritt,
Rizal compared his life in exile to a weary butterfly with sun-burnt
wings.[51]

Butterfly and Chinese

wisteriaflowers, by X Xi
(c.886c.975), painted
around 970 during the early
Song Dynasty.

Some people say that when a butterfly lands on you it means good
luck.[citation needed] However, in Devonshire, people would
traditionally rush around to kill the first butterfly of the year that they
see, or else face a year of bad luck.[52] 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.[53][54]
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
Researches on the wing structure of Palawan Birdwing butterflies led
to new wide wingspan kite and aircraft designs.[55]

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Der Schmetterlingsjger
(The butterfly hunter) by
Carl Spitzweg (1840), a
depiction from the era of
butterfly collection.

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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.[56]
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.[57]

See also

Butterfly zoo
Florida Museum of Natural History#McGuire Center for Lepidoptera and Biodiversity
List of butterflies in Taiwan
List of butterflies of Great Britain
List of butterflies of Tobago
List of butterflies of Menorca
List of butterflies of India
List of butterflies of North America
List of U.S. state butterflies
Moth

Cited references
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upon attending ants". Zoological Journal of the Linnean Society 94: 379. doi:10.1111/j.10963642.1988.tb01201.x.
3. ^ Devries, Pj (Jun 1990). "Enhancement of Symbioses Between Butterfly Caterpillars and Ants by
Vibrational Communication.". Science (New York, N.Y.) 248 (4959): 11041106.
doi:10.1126/science.248.4959.1104. PMID 17733373.
4. ^ Ehrlich, P. R., and P. H. Raven. 1964. Butterflies and plants: a study in coevolution. Evolution 18:586
608
5. ^ Mason, C. W. (1927). The Journal of Physical Chemistry 31: 321. doi:10.1021/j150273a001.
6. ^ Vukusic, P., J.R.Sambles, and H. Ghiradella (2000) Optical Classification of Microstructure in Butterfly
Wing-scales. Photonics Science News, 6, 61-66 [1]
7. ^ Prum, Ro; Quinn, T; Torres, Rh (Feb 2006). "Anatomically diverse butterfly scales all produce
structural colours by coherent scattering." (Free full text). The Journal of experimental biology 209 (Pt 4):
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9. ^ Rousset, F; Bouchon, D; Pintureau, B; Juchault, P; Solignac, M (Nov 1992). "Wolbachia endosymbionts
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Royal Society 250 (1328): 918. doi:10.1098/rspb.1992.0135. ISSN 0962-8452. PMID 1361987.
10. ^ Jiggins, Francis M (2001). "Two male-killing Wolbachia strains coexist within a population of the
butterfly Acraea encedon". Heredity 86: 161. doi:10.1046/j.1365-2540.2001.00804.x.
11. ^ Meyer, A (Oct 2006). "Repeating patterns of mimicry." (Free full text). PLoS biology 4 (10): e341.
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14. ^ Brakefield, Pm; Kesbeke, F; Koch, Pb (Dec 1998). "The regulation of phenotypic plasticity of eyespots
in the butterfly Bicyclus anynana.". The American naturalist 152 (6): 85360. doi:10.1086/286213.
ISSN 0003-0147. PMID 18811432.
15. ^ Nijhout, Hf (Jan 2003). "Development and evolution of adaptive polyphenisms.". Evolution &
development 5 (1): 918. ISSN 1520-541X. PMID 12492404.
16. ^ Brakefield, PAUL M. (1984). "The evolutionary significance of dry and wet season forms in some
tropical butterflies". Biological Journal of the Linnean Society 22: 1. doi:10.1111/j.10958312.1984.tb00795.x.
17. ^ Lyytinen, A., P. M. Brakefield, L. Lindstrm, and J. Mappes. 2004. Does predation maintain eyespot
plasticity in Bicyclus anynana. The Proceedings of the Royal Society B-Biological Sciences 271:279-283.
18. ^ Brakefield, P. M., J. Gates, D. Keys, F. Kesbeke, P. J. Wijngaarden, A. Monteiro, V. French, and S. B.
Carroll. 1996. Development, plasticity and evolution of butterfly eyespot patterns. Nature 384:236-242.
19. ^ Gilbert LE (1972). "Pollen feeding and reproductive biology of Heliconius butterflies". Proceedings of
the National Academy of Sciences 69: 14021407. doi:10.1073/pnas.69.6.1403.
http://www.pnas.org/content/69/6/1403.abstract.
20. ^ At 7 to 8 pm 27 October 2009 the BBC2 television program Ray Mears's Northern Wilderness showed a
butterfly feeding from wolf faeces in the Canadian boreal forest
21. ^ At 7.30 to 8 pm 27 October 2009 the ITV1 television program Grimefighters showed a butterfly feeding
from a decaying dead rat in a town in England
22. ^ Herrera, C.M. (1987). "Components of pollinator "quality": comparative analysis of a diverse insect
assemblage.". Oikos 50: 7990. doi:10.2307/3565403.
http://ebd06.ebd.csic.es/pdfs/Herrera.1987.Oikos.pdf.
23. ^ Inouye, David W. (2001) Role of Pollinators pp. 723-730 in Encyclopaedia of Biodiversity Volume 4.
Academic Press.
24. ^ Molleman Freerk, Grunsven Roy H. A., Liefting Maartje, Zwaan Bas J., Brakefield Paul M. (2005) Is
male puddling behaviour of tropical butterflies targeted at sodium for nuptial gifts or activity? Biol. J.
Linn. Soc. 86, (3):345-361
25. ^ "Article on San Diego Zoo website". Sandiegozoo.org. http://www.sandiegozoo.org/animalbytes/tbutterfly.html. Retrieved 2009-03-30.
26. ^ Obara Y, Hidaki T. (1968). Recognition of the female by the male, on the basis of ultra-violet reflection,
in the white cabbage butterfly Pieris rapae crucivora Boisduval. Proc. Japan Acad., 44: 829-832.
27. ^ Tadao Hirota and Yoshiomi Kato 2004 Color discrimination on orientation of female Eurema hecabe
(Lepidoptera: Pieridae) Applied Entomology and Zoology Vol. 39:229-233 [2]
28. ^ Michiyo Kinoshita, Naoko Shimada And Kentaro Arikawa (1999) Color vision of the foraging
swallowtail butterfly Papilio xuthus. The Journal of Experimental Biology 202:95 102 [3]
29. ^ Swihart, S. L (1967). Hearing in butterflies. J. Insect Physiol 13, 469
30. ^ Reppert, Steven M.; Haisun Zhu; White, Richard H. (2004) Polarized light helps monarch butterflies
navigate. Current biology 14(2):155-158
31. ^ Ellers, J. and Carol L. Boggs (2002) The evolution of wing color in Colias butterflies: Heritability, Sex
Linkage, and population divergence. Evolution, 56(4):836 840 [4]
32. ^ Srygley, R. B. and A. L. R. Thomas (2002) Aerodynamics of insect flight: flow visualisations with free
flying butterflies reveal a variety of unconventional lift-generating mechanisms. Nature 420: 660-664.
PDF
33. ^ Williams, C. B. 1927 A study of butterfly migration in south India and Ceylon, based largely on records
by Messrs. G Evershed, E.E.Green, J.C.F. Fryer and W. Ormiston. Trans. Ent. Soc. London 75:1-33
34. ^ Urquhart, F. A. & N. R. Urquhart. 1977. Overwintering areas and migratory routes of the Monarch
butterfly (Danaus p. plexippus, Lepidoptera: Danaidae) in North America, with special reference to the
western population. Can. Ent. 109: 1583-1589
35. ^ Wassenaar L.I., Hobson K.A. 1998. Natal origins of migratory monarch butterflies at wintering colonies
in Mexico: new isotopic evidence. Proc Natl Acad Sci U S A. 95(26):15436-9. Full text
36. ^ Ivo Sauman, Adriana D. Briscoe, Haisun Zhu, Dingding Shi, Oren Froy, Julia Stalleicken, Quan Yuan,
Amy Casselman, and Steven M. Reppert (2005) Connecting the Navigational Clock to Sun Compass Input
in Monarch Butterfly Brain. Neuron. 46:457-467 [5]
37. ^ Southwood, T. R. E. 1962. Migration of terrestrial arthropods in relation to habitat. Biol. Rev. 37:171214
38. ^ Dennis, R L H, Tim G. Shreeve, Henry R. Arnold and David B. Roy (2005) Does diet breadth control
herbivorous insect distribution size? Life history and resource outlets for specialist butterflies. Journal of
Insect Conservation 9(3):187-200
39. ^ Nishida, Ritsuo (2002). Sequestration of defensive substances from plants by Lepidoptera. Annu. Rev.
Entomol. 47:5792
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Lycaenid Butterflies. American Naturalist 118(5):770-775

41. ^ Nafus, D. M. and I. H. Schreiner (1988) Parental care in a tropical nymphalid butterfly Hypolimas
anomala. Anim. Behav. 36: 1425- 143
42. ^ William E. Cooper, Jr. (1998) Conditions favoring anticipatory and reactive displays deflecting
predatory attack. Behavioral Ecology
43. ^ Hind Wings Help Butterflies Make Swift Turns to Evade Predators Newswise, Retrieved on January 8,
2008.
44. ^ Larsen, Torben (1994) Butterflies of Egypt. Saudi Aramco world. 45(5):24-27 Online
45. ^ "Table complete with real butterflies embedded in resin". Mfjoe.com. http://mfjoe.com/tag/furniture/.
Retrieved 2009-03-30.
46. ^ Hearn, Lafcadio (1904). Kwaidan: Stories and Studies of Strange Thing. Dover Publications, Inc..
ISBN 0-486-21901-1.
47. ^ Hutchins, M., Arthur V. Evans, Rosser W. Garrison and Neil Schlager (Eds) (2003) Grzimek's Animal
Life Encyclopedia, 2nd edition. Volume 3, Insects, Farmington Hills, MI: Gale Group, 2003.
48. ^ Rabuzzi, M. 1997. Butterfly etymology. Cultural Entomology November 1997 Fourth issue online
49. ^ "The Best Known Speech of Jose Rizal". Joserizal.info.
http://joserizal.info/Writings/Speeches/speeches.htm. Retrieved 2009-03-30.
50. ^ "The Life and Writings of Dr. Jose Rizal". Joserizal.info.
http://joserizal.info/Biography/man_and_martyr/chapter14.htm. Retrieved 2009-03-30.
51. ^ "193. Rizal, Dapitan, 19 December 1898". Univie.ac.at.
http://www.univie.ac.at/Voelkerkunde/apsis/aufi/rizal/rbcor193.htm. Retrieved 2009-03-30.
52. ^ Dorset Chronicle, May 1825, reprinted in: "The First Butterfly", in The Every-day Book and Table
Book; or, Everlasting Calendar of Popular Amusements, Sports, Pastimes, Ceremonies, Manners,
Customs, and Events, Each of the Three Hundred and Sixty-Five Days, in Past and Present Times;
Forming a Complete History of the Year, Months, and Seasons, and a Perpetual Key to the Almanac,
Including Accounts of the Weather, Rules for Health and Conduct, Remarkable and Important Anecdotes,
Facts, and Notices, in Chronology, Antiquities, Topography, Biography, Natural History, Art, Science,
and General Literature; Derived from the Most Authentic Sources, and Valuable Original
Communication, with Poetical Elucidations, for Daily Use and Diversion. Vol III., ed. William Hone,
(London: 1838) p 678.
53. ^ "Death practices Philippine style". Sunstar.com.ph. 2005-10-30.
http://www.sunstar.com.ph/static/dav/2005/10/30/feat/death.practices.philippine.style.html. Retrieved
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54. ^ don.herrington@livinginthephilippines.com. "Superstitions and Beliefs related to Death".
Livinginthephilippines.com.
http://www.livinginthephilippines.com/philculture/superstitions_and_beliefs.html. Retrieved 2009-03-30.
55. ^ "An Introduction to The World of Birdwing Butterflies". Nagypal.net. 2000-05-28.
http://www.nagypal.net/. Retrieved 2009-03-30.
56. ^ Vukusic, Pete and Ian Hooper. 2005. Directionally Controlled Fluorescence Emission in Butterflies
Science. 310(5751):1151 DOI: 10.1126/science.1116612
57. ^ "The Butterflies that Rizal Chased, Collected, and Studied". Books.google.com.
http://books.google.com/books?id=GiUuFWeM6uQC&pg=PA13&dq=isbn:9781430323693#PPP1,M1.
Retrieved 2009-03-30.

Other references

Boggs, C., Watt, W., Ehrlich, P. 2003. Butterflies: Evolution and Ecology Taking Flight.
University of Chicago Press, Chicago, USA.
Fadul, J. A. 2008. The Butterflies that Rizal Chased, Collected, and Studied. Morrisville, NC:
Lulu Press. ISBN 978-1-430-32369-3
Heppner, J. B. 1998. Classification of Lepidoptera. Holarctic Lepidoptera, Suppl. 1.
Pyle, R. M. 1992. Handbook for Butterfly Watchers. Houghton Mifflin. First published, 1984.
ISBN 0-395-61629-8
Nemos, F. ca. 1895. Europas bekannteste Schmetterlinge. Beschreibung der wichtigsten Arten
und Anleitung zur Kenntnis und zum Sammeln der Schmetterlinge und Raupen Oestergaard
Verlag, Berlin, (pdf 77MB)
Pea, C., N. Waklberg, E. Weingartner, U. Kodandaramaiah, S. Nylin, A. V. L. Freitas, and A.
V. Z. Brower. 2006. Higher level phylogeny of Satyrinae butterflies (Lepidoptera:
Nymphalidae) based on DNA sequence data. Molecular Phylogenetics and Evolution 40:29-49.

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Stevens, M. 2005. The role of eyespots as anti-predator mechanisms, principally demonstrated

in the Lepidoptera. Biological Reviews 80:573-588.
Monteiro, A. and N. E. Pierce. 2001. Phylogeny of Bicyclus (Lepidoptera : Nymphalidae)
inferred from COI, COII, and EF-1 alpha gene sequences. Molecular Phylogenetics and
Evolution 18:264-281.

Field guides to butterflies

Butterflies of North America, Jim P. Brock and Kenn Kaufman (2003)

Butterflies through Binoculars: The East, Jeffrey Glassberg (1999)
Butterflies through Binoculars: The West, Jeffrey Glassberg (2001)
A Field Guide to Eastern Butterflies, Paul Opler (1994)
A Field Guide to Western Butterflies, Paul Opler (1999)
Peterson First Guide to Butterflies and Moths, Paul Opler (1994)
Las Mariposas de Machu Picchu by Gerardo Lamas (2003)
The Millennium Atlas of Butterflies in Britain and Ireland by Jim Asher (Editor), et al.
Pocket Guide to the Butterflies of Great Britain and Ireland by Richard Lewington
Butterflies of Britain and Europe (Collins Wildlife Trust Guides) by Michael Chinery
Butterflies of Europe by Tom Tolman and Richard Lewington (2001)
Butterflies of Europe New Field Guide and Key by Tristan Lafranchis (2004)
Butterflies of Lebanon by Torben B. Larsen. Beirut. (1974)
The butterflies of Saudi Arabia and its neighbours. by Torben B. Laren (Stacey intl.) (1984)
The butterflies of Egypt by Torben B. Larsen (Apollo Books, Denmark). (1990)
Field Guide to Butterlies of South Africa by Steve Woodhall (2005)
The butterflies of Kenya and their natural history by Torben B. Larsen (OUP) (1991)
Butterflies of Sikkim Himalaya and their Natural History by Meena Haribal (1994).
Butterflies of Peninsular India by Krushnamegh Kunte, Universities Press (2005).
Butterflies of the Indian Region by Col M. A. Wynter-Blyth, Bombay Natural History Society,
Mumbai, India (1957).
A Guide to Common Butterflies of Singapore by Steven Neo Say Hian (Singapore Science
Centre)
Butterflies of West Malaysia and Singapore by W.A.Fleming. (Longman Malaysia)
The Butterflies of the Malay Peninsula by A.S. Corbet and H. M. Pendlebury. (The Malayan
Nature Society)
Butterflies of West Africa (two vols.) by Torben B. Larsen. (Apollo Books, Denmark) (2005)
Oxford Butterflies of India by Thomas Gray, I.D.Kehimkar, J Punetha, Oxford University Press
(2008)

External links
General interest

The Royal Horticultural Society butterfly exhibition

Papilionoidea on the Tree of Life Web project
Butterflies on the UF / IFAS Featured Creatures Web site

Regional lists

Collodi Butterfly House Tuscany

Butterflies and Moths of North America
Butterflies of America
Butterflies of Canada
North American Butterfly Association (NABA)
Butterflies and Moths in the Netherlands

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Insect and butterfly diversity of Pakistan

Butterflies of the Philippines
Butterflies of Southern India
Butterflies of Sri Lanka
Butterflies of Singapore
Israel Insect World
Singapore Butterfly Checklist
Butterfly Conservation Society of Taiwan
Butterflies of Morocco
Butterflies of Indo-China Chiefly Thailand, Laos and Vietnam.
Butterflies of Southeastern Sulawesi
Naturalis Butterflies of Sulawesi (Illustrated pdf)
Butterflies of Thailand
Butterflies of Mexico
Butterflies of Ghana

Literature

Literaturatenbank Free downloads

Images/Movies

BugGuide.net Many images of North American butterflies, many licensed via Creative
Commons
Butterfly Pictures and Information
Butterfly of Brazil
Reference quality large format photographs, common butterflies of North America
Gallery of Florida Butterflies and Moths
Butterfly Picture Gallery
Photographs of most of the Butterflies in Southern California
Butterflies of Southern Illinois
Butterflies of France
Butterfly Movies (Tree of Life)
1000+ photos of Massachusetts butterflies
European butterfly pictures - common names and wildlife photography
Online videos of Skippers of the Northeast-USA

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