Lepidoptera (Butterflies & Moths)

**Butterflies** are part of the class of Insects in the order Lepidoptera. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises the *true butterflies* (superfamily Papilionoidea), the *skippers*(superfamily Hesperioidea) and the *moth-butterflies* (superfamily Hedyloidea). Butterfly fossils date to the mid Eocene epoch, 40–50 million years ago.

Butterflies exhibit polymorphism, mimicry and aposematism. Some, like the Monarch, will migrate over long distances. Some butterflies have parasitic relationships with organisms including protozoans, flies, ants, other invertebrates, and vertebrates. Some species are pests because in their larval stages they can damage domestic crops or trees; however, some species are agents of pollination of some plants, and caterpillars of a few butterflies (e.g., Harvesters) eat harmful insects. Culturally, butterflies are a popular motif in the visual and literary arts.

## Lifecycle

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.

### Eggs

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 almost invariably 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 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. Others such as *Phengaris arion* communicate with ants to form a parasitic relationship.

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 foul-smelling 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 makes them unpalatable to birds and other predators. Such unpalatibility is advertised using bright red, orange, black or white warning colours, a practice known as aposematism. 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.

#### 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.

When the larva is fully grown, hormones such as prothoracicotropic hormone (PTTH) are produced. At this point the larva stops feeding and begins "wandering" in the quest of a suitable pupation site, often the underside of a leaf.

The larva transforms into a pupa (or chrysalis) by anchoring itself to a substrate and moulting for the last time. The chrysalis is usually incapable of movement, although some species can rapidly move the abdominal segments or produce sounds to scare potential predators.

The pupal transformation into a butterfly through metamorphosis has held great appeal to mankind. To transform from the miniature wings visible on the outside of the pupa into large structures usable for flight, the pupal wings undergo rapid mitosis and absorb a great deal of nutrients. If one wing is surgically removed early on, the other three will grow to a larger size. In the pupa, the wing forms a structure that becomes compressed from top to bottom and pleated from proximal to distal ends as it grows, so that it can rapidly be unfolded to its full adult size. Several boundaries seen in the adult color pattern are marked by changes in the expression of particular transcription factors in the early pupa.

### Adult or imago

The adult, sexually mature, stage of the insect is known as the imago. As Lepidoptera, butterflies have four wings that are covered with tiny scales. The fore and hindwings are not hooked together, permitting a more graceful flight. An adult butterfly has six legs, but in thenymphalids, 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 hemolymph and let 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.

## 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.

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, 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 giftalong with the spermatophore, during mating.

Butterflies use their antennae to sense the air for wind and scents. 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 sensory organs known as sensillae. A butterfly's sense of taste, 200 times stronger than humans, 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.

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.

## 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 and Yellows, Family **Pieridae**
	+ Small White, *Pieris rapae*
	+ Large White, *Pieris brassicae*
	+ Green-veined White, *Pieris napi*
	+ Common Jezebel, *Delias eucharis*
* 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*