Defence

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. 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.

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 .

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

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.

Migration

Many butterflies migrate over long distances. Particularly famous migrations are those of the Monarch butterfly from Mexico to northern USA and southern Canada, a distance of about 4000 to 4800 km (2500–3000 miles). Other well known migratory species include the Painted Lady and several of the Danaine butterflies. Spectacular and large scale migrations associated with the Monsoons are seen in peninsular India.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 particularly 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.

Habits

Butterflies feed primarily on nectar from flowers. Some also derive nourishment from pollen,tree sap, rotting fruit, dung, decaying flesh, and dissolved minerals in wet sand or dirt. Butterflies are important as pollinators for some species of plants although in general they do not carry as much pollen load as bees. They are however capable of moving pollen over greater distances.Flower constancy has been observed for at least one species of butterfly.
As adults, butterflies consume only liquids and these are sucked by means of their proboscis. They feed on nectar from flowers and also sip water from damp patches. This they do for water, for energy from sugars in nectar and for sodium and other minerals which are vital for their reproduction. Several species of butterflies need more sodium than provided by nectar. They are attracted to sodium in salt and they sometimes land on people, attracted by human sweat. Besides damp patches, some butterflies also visit dung, rotting fruit or carcasses to obtain minerals and nutrients. In many species, this mud-puddling behaviour is restricted to the males, and studies have suggested that the nutrients collected are provided as a nuptial gift along with the spermatophore during mating.

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, 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. 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

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.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.The dry-season forms are usually more cryptic and it has been suggested that the protection offered may be an adaptation. Some also show greater dark colours in the wet-season form which may have thermoregulatory advantages by increasing ability to absorb solar radiation.

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.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 .

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.

Pupa

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.

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.

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.

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 entomophagous (insect eating).
Some larvae, especially those of the Lycaenidae, form mutual associations with ants. They communicate with the ants using vibrations that are transmitted through the substrate as well as using chemical signals.^[2][3] The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions.
Caterpillars mature through a series of stages called instars. Near the end of each instar, the larva undergoes a process called apolysis, in which the cuticle, a tough outer layer made of a mixture of chitin and specialized proteins, is released from the softer epidermis beneath, and the epidermis begins to form a new cuticle beneath. At the end of each instar, the larva moults the old cuticle, and the new cuticle expands, before rapidly hardening and developing pigment. Development of butterfly wing patterns begins by the last larval instar.

Butterfly caterpillars have three pairs of true legs from the thoracic segments and up to 6 pairs of prolegs arising from the abdominal segments. These prolegs have rings of tiny hooks called crochets that help them grip the substrate.
Some caterpillars have the ability to inflate parts of their head to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called osmeteria which are everted to produce smelly chemicals. These are used in defense.
Host plants often have toxic substances in them and caterpillars are able to sequester these substances and retain them into the adult stage. This helps making them unpalatable to birds and other predators. Such unpalatibility is advertised using bright red, orange, black or white warning colours. The toxic chemicals in plants are often evolved specifically to prevent them from being eaten by insects. Insects in turn develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants.

Egg

Butterfly eggs are protected by a hard-ridged outer layer of shell, called the chorion. This is lined with a thin coating of wax which prevents the egg from drying out before the larva has had time to fully develop. Each egg contains a number of tiny funnel-shaped openings at one end, called micropyles; the purpose of these holes is to allow sperm to enter and fertilize the egg. Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate.
Butterfly eggs are fixed to a leaf with a special glue which hardens rapidly. As it hardens it contracts, deforming the shape of the egg. This glue is easily seen surrounding the base of every egg forming a meniscus. The nature of the glue is unknown and is a suitable subject for research. The same glue is produced by a pupa to secure the setae of the cremaster. This glue is so hard that the silk pad, to which the setae are glued, cannot be separated.

Eggs are usually laid on plants. Each species of butterfly has its own hostplant range and while some species of butterfly are restricted to just one species of plant, others use a range of plant species, often including members of a common family.
The egg stage lasts a few weeks in most butterflies but eggs laid close to winter, especially in temperate regions, go through a diapause (resting) stage, and the hatching may take place only in spring. Other butterflies may lay their eggs in the spring and have them hatch in the summer. These butterflies are usually northern species, such as the Mourning Cloak (Camberwell Beauty) and the Large and Small Tortoiseshell butterflies.