A butterfly is an insect of the order Lepidoptera. The scope of the term depends on how far the concept of "butterfly" is extended. Currently, most experts include the superfamilies Hedyloidea (the American moth-butterflies), Hesperioidea (the skippers) and Papilionoidea (the so-called 'true' butterflies). This concept of butterflies including the Hedyloidea is an only recently expanded one, but it makes the group a natural clade, the Rhopalocera. Like all Lepidoptera, butterflies are notable for their unusual life cycle with a larval caterpillar stage, an inactive pupal stage and a spectacular metamorphosis into a familiar and colorful winged adult form, and most species being day-flying, they regularly attract attention. The diverse patterns formed by their brightly colored wings and their erratic-yet graceful-flight have made butterfly watching a popular hobby.
![]()
|
Butterflies are nested within the evolutionary tree of moths.
Their origins may date back to the Cretaceous Period, ending 65
million years ago. Unfortunately, the fossil record is very
limited. The oldest known fossil is an unnamed possible skipper
butterfly from the Upper Palaeocene (around 57 million year old)
of Fur, Denmark . One of the most beautifully preserved is a
Metalmark butterfly (Voltinia dramba) from 25 million year old
Dominican amber.
Butterflies are today distributed throughout the world except in
the very cold and arid regions. There are an estimated 17,500
species of butterflies (Papilionoidea) out of about 180,000
species of Lepidoptera.
Presently butterflies are classified in three superfamilies,
Hedyloidea, consisting of the 'American moth-butterflies',
Hesperioidea, consisting of the 'skippers' and Papilionoidea or
'true butterflies'. The last two superfamilies are probably
sister taxa, so the butterflies collectively are thought to
constitute a natural group or clade.
A study combining morphological and molecular data concluded
that Hesperiidae, Papilionidae, Pieridae, Lycaenidae and
Riodinidae could all be strongly supported as monophyletic
groups, but the monophyletic status of Nymphalidae is uncertain.
Lycaenidae and Riodinidae were confirmed as sister taxa, and
Papilionidae as the outgroup to the rest of the true
butterflies, but the location of Pieridae within the pattern of
descent was unclear, with different lines of evidence suggesting
different conclusions. The data suggested that the Hedyloidea
are indeed more closely related to the butterflies than to other
moths.
Some older classifications recognize additional families, for
example Danaidae, Heliconiidae, Libytheidae and Satyridae, but
modern classifications treat these as subfamilies within the
Nymphalidae.
The dichotomous classification of lepidopterans into butterflies
and moths is one that is popular but not used in taxonomy. The
folk groups of butterflies and moths can be distinguished using
several features but there are exceptions to most of these
rules.
Unlike many insects, butterflies do not experience a nymph
period, but instead go through a pupal stage which lies between
the larva and the adult stage (the imago). Butterflies are
termed as holometabolous insects, and go through complete
metamorphosis.
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.
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.
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 are restricted to
just one species, others use a range of plant species, often
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 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 northernly
species (Mourning cloak, Tortiseshells)
Larvae, or caterpillars, are multi-legged eating machines. They
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. 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 mixture of chitin and
specialized proteins, is released from the epidermis 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 rapidly hardens and pigments. 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 colors. 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.
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.
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.
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 3
hours to dry while others take about 1 hour. Most butterflies
and moths will excrete excess dye after hatching. This fluid may
be white, red, orange, or in rare cases, blue.
|
|
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 microstructure of the scales.
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).
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 or the selective killing of
males in the egg stage.
Batesian and Mullerian mimicry in butterflies is common. 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.
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. Experimental
modification by ecdysone hormone treatment has demonstrated that
it is possible to control the continuum of expression of the
different forms. 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 colors in the wet-season
form which may have thermoregulatory advantages by increasing
ability to absorb solar radiation.
Butterflies feed primarily on nectar from flowers. Some also
derive nourishment from pollen, tree sap, rotting fruit, dung,
and dissolved minerals in wet sand or dirt. Butterflies play an
important ecological role as pollinators.
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
behavior 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.
Butterflies sense the air for scents, wind and nectar using
their antennae. The antennae come in various shapes and colors.
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. 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. Color
vision may be widespread but has been demonstrated in only a few
species.
Some butterflies have organs of hearing and some species are
also known to make stridulatory and clicking sounds.
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.
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.
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. (See also
Insect flight)
Many butterflies migrate over long distances. Particularly
famous migrations being those of the Monarch butterfly from
Mexico to North America, a distance of about 4,000 to 4,800
kilometres (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. Migrations have been
studied in more recent times using wing tags and also using
stable hydrogen isotopes.
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 particular
important.
It is suggested that most migratory butterflies are those that
belong to semi-arid areas where breeding seasons are short. The
life-histories of their host plants also influence the
strategies of the butterflies.
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 often based on
chemicals of plant origin. In many cases the plants themselves
have evolved these toxic substances to reduce attack to them.
These defense mechanisms are effective only if they are also
well advertised. Many unpalatable butterflies are brightly
colored. This has led to unprotected butterflies evolving forms
that appear like the unpalatable butterflies. These mimetic
forms are usually restricted to the females.
Cryptic coloration is found in many butterflies. Some like the
oakleaf butterfly are remarkable imitations of leaves. 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).
Behavioral defenses include perching and wing positions to avoid
being conspicuous. Some female Nymphalid butterflies are known
to guard their eggs from parasitoid wasps.
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.

This Butterfly Page is Copyright The Animal Web Guide © 2004 - 2009 Chuck Ayoub