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Pupation - Inachis io
A Peacock Butterfly (Inachis io) larva (caterpillar) becoming a pupa. Time: Nine minutes
Inachis io-02 (xndr)
An adult Peacock Butterfly

A pupa (plural: pupae or pupas) is part of becoming an adult for some insects, like some butterflies, moths, beetles, flies, and bees. The pupal stage comes after being a larva; the next stage is adulthood.

The pupa stage of a butterfly, called a chrysalis (plural "chrysalids"), is a well-known example of a pupa because they are often found in the open.

Position in life cycle

The pupal stage follows the larval stage and precedes adulthood (imago) in insects with complete metamorphosis. The pupa is a non-feeding, usually sessile stage, or highly active as in mosquitoes. It is during pupation that the adult structures of the insect are formed while the larval structures are broken down. The adult structures grow from imaginal discs.

Duration

Pupation may last weeks, months, or even years, depending on temperature and the species of insect. For example, pupation lasts eight to fifteen days in monarch butterflies. The pupa may enter dormancy or diapause until the appropriate season to emerge as an adult insect. In temperate climates pupae usually stay dormant during winter, while in the tropics pupae usually do so during the dry season.

Hercus fontinalis eclosion
Adult Hercus fontinalis emerging from cocoon

Emergence

Insects emerge (eclose) from pupae by splitting the pupal case. Most butterflies emerge in the morning. In mosquitoes the emergence is in the evening or night. In fleas the process is triggered by vibrations that indicate the possible presence of a suitable host. Prior to emergence, the adult inside the pupal exoskeleton is termed pharate. Once the pharate adult has eclosed from the pupa, the empty pupal exoskeleton is called an exuvia; in most hymenopterans (ants, bees and wasps) the exuvia is so thin and membranous that it becomes "crumpled" as it is shed.

Types

Based on the presence or absence of articulated mandibles that are employed in emerging from a cocoon or pupal case, the pupae can be classified in to two types:

Based on whether the pupal appendages are free or attached to the body, the pupae can be classified in three types:

  • Exarate pupa – appendages are free and are not usually encapsulated within a cocoon. All decticous pupa and some adecticous pupa are always exarate. (Neuroptera, Trichoptera, Cyclorrhapha of Dipterans, Siphonaptera, most Coleoptera, Hymenoptera, and few Lepidoptera).
  • Obtect pupa – appendages are attached closely to the body and are commonly encapsulated within a cocoon. Some adecticous pupa are obtect forms. (Most Lepidoptera, Nematocera and Brachycera of Dipterans, Staphylinidae and Chrysomelidae Coleopterans, many Chalcidoidea Hymenopterans)
  • Coarctate pupa – enclosed in a hardened cuticle of the penultimate larval instar called puparium. However, the pupa itself is of exarate adecticous pupa forms. (Cyclorrhapha of Dipterans).

Chrysalis

Common crow pupa
Common crow butterfly (Euploea core) chrysalis illustrating the Greek origin of the term: χρυσός (chrysós) for gold

A chrysalis (Latin chrysallis, from Greek χρυσαλλίς = chrysallís, plural: chrysalides, also known as an aurelia) or nympha is the pupal stage of butterflies. The term is derived from the metallic gold-coloration found in the pupae of many butterflies, referred to by the Greek term χρυσός (chrysós) for gold.

When the caterpillar is fully grown, it makes a button of silk which it uses to fasten its body to a leaf or a twig. Then the caterpillar's skin comes off for the final time. Under this old skin is a hard skin called a chrysalis.

Because chrysalises are often showy and are formed in the open, they are the most familiar examples of pupae. Most chrysalides are attached to a surface by a Velcro-like arrangement of a silken pad spun by the caterpillar, usually cemented to the underside of a perch, and the cremastral hook or hooks protruding from the rear of the chrysalis or cremaster at the tip of the pupal abdomen by which the caterpillar fixes itself to the pad of silk. (Gr. kremastos 'suspended')

Like other types of pupae, the chrysalis stage in most butterflies is one in which there is little movement. However, some butterfly pupae are capable of moving the abdominal segments to produce sounds or to scare away potential predators. Within the chrysalis, growth and differentiation occur. The adult butterfly emerges (ecloses) from this and expands its wings by pumping haemolymph into the wing veins. Although this sudden and rapid change from pupa to imago is often called metamorphosis, metamorphosis is really the whole series of changes that an insect undergoes from egg to adult.

When emerging, the butterfly uses a liquid, sometimes called cocoonase, which softens the shell of the chrysalis. Additionally, it uses two sharp claws located on the thick joints at the base of the forewings to help make its way out. Having emerged from the chrysalis, the butterfly will usually sit on the empty shell in order to expand and harden its wings. However, if the chrysalis was near the ground (such as if it fell off from its silk pad), the butterfly would find another vertical surface to rest upon and harden its wings (such as a wall or fence).

Moth pupae are usually dark in color and either formed in underground cells, loose in the soil, or their pupa is contained in a protective silk case called a cocoon. The pupa of some species such as the hornet moth develop sharp ridges around the outside called adminicula that allow the pupa to move from its place of concealment inside a tree trunk when it is time for the adult to emerge.

Pupa, chrysalis, and cocoon are frequently confused, but are quite distinct from each other. The pupa is the stage between the larva and adult stages. The chrysalis generally refers to a butterfly pupa although the term may be misleading as there are some moths whose pupae resembles a chrysalis, e.g.: the plume winged moths of the family Pterophoridae and some geometrid moths. A cocoon is a silk case that the larvae of moths, and sometimes other insects, spin around the pupa.

Cocoon

Caterpillars cocoon
The tough brown cocoon of an emperor gum moth

A cocoon is a casing spun of silk by many moths and caterpillars, and numerous other holometabolous insect larvae as a protective covering for the pupa.

Cocoons may be tough or soft, opaque or translucent, solid or meshlike, of various colors, or composed of multiple layers, depending on the type of insect larva producing it. Many moth caterpillars shed the larval hairs (setae) and incorporate them into the cocoon; if these are urticating hairs then the cocoon is also irritating to the touch. Some larvae attach small twigs, fecal pellets or pieces of vegetation to the outside of their cocoon in an attempt to disguise it from predators. Others spin their cocoon in a concealed location—on the underside of a leaf, in a crevice, down near the base of a tree trunk, suspended from a twig or concealed in the leaf litter.

The silk in the cocoon of the silk moth can be unraveled to harvest silk fibre which makes this moth the most economically important of all lepidopterans. The silk moth is the only completely domesticated lepidopteran and does not exist in the wild.

Insects that pupate in a cocoon must escape from it, and they do this either by the pupa cutting its way out, or by secreting enzymes, sometimes called cocoonase, that soften the cocoon. Some cocoons are constructed with built-in lines of weakness along which they will tear easily from inside, or with exit holes that only allow a one-way passage out; such features facilitate the escape of the adult insect after it emerges from the pupal skin.

Eupeodes americanus
Puparium of Eupeodes americanus

Puparium

Some pupae remain inside the exoskeleton of the final larval instar and this last larval "shell" is called a puparium (plural, puparia). Flies of the group Muscomorpha have puparia, as do members of the order Strepsiptera, and the Hemipteran family Aleyrodidae.

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See also

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