A pupa (Latin

Latin

Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 pupa for doll, pl: pupae or pupas) is the life stage of some insect

Insect

Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s undergoing transformation. The pupal stage is found only in holometabolous

Holometabolism

Holometabolism, also called complete metamorphism, is a term applied to insect groups to describe the specific kind of insect development which includes four life stages - as an embryo or egg, a larva, a pupa and an imago or adult. Holometabolism is a monophyletic trait that all insects in the...

 insects, those that undergo a complete metamorphosis, going through four life stages; embryo

Embryo

An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...

, larva

Larva

A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

, pupa and imago

Imago

In biology, the imago is the last stage of development of an insect, after the last ecdysis of an incomplete metamorphosis, or after emergence from the pupa where the metamorphosis is complete...

. (For a list of such insects see Holometabolism

Holometabolism

Holometabolism, also called complete metamorphism, is a term applied to insect groups to describe the specific kind of insect development which includes four life stages - as an embryo or egg, a larva, a pupa and an imago or adult. Holometabolism is a monophyletic trait that all insects in the...

).

The pupae of different groups of insects have different names such as chrysalis in the Lepidoptera

Lepidoptera

Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...

 and tumbler in mosquitoes. Pupae may further be enclosed in other structures such as cocoons, nest

Nest

A nest is a place of refuge to hold an animal's eggs or provide a place to live or raise offspring. They are usually made of some organic material such as twigs, grass, and leaves; or may simply be a depression in the ground, or a hole in a tree, rock or building...

s or shells.

Some pupae remain inside the exoskeleton of the final larval instar and receive the name of puparium (plural, puparia); the flies of the families Stratiomyidae

Stratiomyidae

The soldier flies , are a family of flies . The family contains about 1,500 species in about 400 genera worldwide. Adults are found near larval habitats...

, Syrphidae and others have puparia.

Position in life cycle

In the life

Biological life cycle

A life cycle is a period involving all different generations of a species succeeding each other through means of reproduction, whether through asexual reproduction or sexual reproduction...

 of an insect the pupal stage follows the larva

Larva

A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

l stage and precedes adult

Adult

An adult is a human being or living organism that is of relatively mature age, typically associated with sexual maturity and the attainment of reproductive age....

hood (imago

Imago

In biology, the imago is the last stage of development of an insect, after the last ecdysis of an incomplete metamorphosis, or after emergence from the pupa where the metamorphosis is complete...

). It is during the time of pupation that the adult structures of the insect are formed while the larval structures are broken down. Pupae are inactive, and usually sessile

Sessility (zoology)

In zoology, sessility is a characteristic of animals which are not able to move about. They are usually permanently attached to a solid substrate of some kind, such as a part of a plant or dead tree trunk, a rock, or the hull of a ship in the case of barnacles. Corals lay down their own...

 (not able to move about). They have a hard protective coating and often use camouflage

Camouflage

Camouflage is a method of concealment that allows an otherwise visible animal, military vehicle, or other object to remain unnoticed, by blending with its environment. Examples include a leopard's spotted coat, the battledress of a modern soldier and a leaf-mimic butterfly...

 to evade potential predators.

Duration

Pupation may last weeks, months or even years. For example it is 2 weeks in monarch butterflies

Monarch butterfly

The Monarch butterfly is a milkweed butterfly , in the family Nymphalidae. It is perhaps the best known of all North American butterflies. Since the 19th century, it has been found in New Zealand, and in Australia since 1871 where it is called the Wanderer...

. The pupa may enter dormancy

Dormancy

Dormancy is a period in an organism's life cycle when growth, development, and physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be closely associated with environmental conditions...

 or diapause

Diapause

Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

until the appropriate season for the adult insect. In temperate

Temperate

In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

 climate pupae usually stay dormant during winter, in the tropics

Tropics

The tropics is a region of the Earth surrounding the Equator. It is limited in latitude by the Tropic of Cancer in the northern hemisphere at approximately  N and the Tropic of Capricorn in the southern hemisphere at  S; these latitudes correspond to the axial tilt of the Earth...

 pupae usually do so during the dry season

Dry season

The dry season is a term commonly used when describing the weather in the tropics. The weather in the tropics is dominated by the tropical rain belt, which oscillates from the northern to the southern tropics over the course of the year...

. Anise Swallowtail

Anise Swallowtail

The Anise Swallowtail, Papilio zelicaon, is a common swallowtail butterfly of western North America. Both the upper and lower sides of its wings are black, but the upper wing has a broad yellow stripe across it, which gives the butterfly an overall yellow appearance...

s sometimes emerge after years as a chrysalis.

Emergence

Insects emerge (eclose) from pupae by splitting the pupal case, and the whole process of pupation is controlled by the insect's hormone

Hormone

A hormone is a chemical released by a cell or a gland in one part of the body that sends out messages that affect cells in other parts of the organism. Only a small amount of hormone is required to alter cell metabolism. In essence, it is a chemical messenger that transports a signal from one...

s. 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

Exoskeleton

An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...

 is termed "pharate". Once the pharate adult has eclosed from the pupa, the empty pupal exoskeleton is called an "exuvium" (or exuvia); in most hymenopterans (ants, bees and wasps) the exuvium is so thin and membranous that it becomes "crumpled" as it is shed.

Pupal mating

In a few taxa of the Lepidoptera, especially Heliconius

Heliconius

Heliconius comprise a colorful and widespread brush-footed butterfly genus distributed throughout the tropical and subtropical regions of the New World. These butterflies utilize Passion flower plants as their larval food source and rely on bright wing color patterns to signal their distastefulness...

, pupal mating is an extreme form of reproductive strategy where adult males mate with female pupa about to emerge or with the newly moulted female; this is accompanied by other actions such as capping of the reproductive system of the female with the sphragis, denying access to other males, or by exuding an anti-aphrodisiac pheromone.

Defense

Pupae are usually immobile and are largely defenseless. To overcome this, a common feature is concealed placement. There are some species of Lycaenid butterflies

Lycaenidae

The Lycaenidae are the second-largest family of butterflies, with about 6000 species worldwide, whose members are also called gossamer-winged butterflies...

 who are protected in their pupal stage by ants. Another means of defense by pupae of other species is the capability of making sounds or vibrations to scare potential predators. A few species use chemical defenses including toxic secretions. The pupae of social hymenoptera

Hymenoptera

Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

ns are protected by adult members of the hive.

Chrysalis

A chrysalis (Latin

Latin

Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 chrysallis, from Greek

Greek language

Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 χρυσαλλίς = chrysallís, pl: chrysalides) or nympha is the pupal stage of butterflies

Butterfly

A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...

. The term is derived from the metallic gold-coloration found in the pupae of many butterflies, referred to by the Greek

Greek language

Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 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 chrysalides 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

Velcro

Velcro is the brand name of the first commercially marketed fabric hook-and-loop fastener, invented in 1948 by the Swiss electrical engineer George de Mestral...

-like arrangement of a silk

Silk

Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...

en pad spun by the caterpillar

Caterpillar

Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

, usually cemented to the underside of a perch, and the cremaster, a hook-shaped protuberance from the rear of the chrysalis at the tip of the pupal abdomen

Abdomen

In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...

 by which the caterpillar fixes itself to the pad of silk.

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

Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...

, metamorphosis is really the whole series of changes that an insect undergoes from egg to adult.

On emerging the butterfly uses a liquid 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

Moth

A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...

 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

Silk

Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...

 case called a cocoon.

It is important to differentiate between pupa, chrysalis and cocoon. The pupa is the stage between the larva and adult stages. The chrysalis is a butterfly pupa. A cocoon is a silk case that moths, and sometimes other insects, spin around the pupa.

Cocoon

A cocoon is a casing spun of silk

Silk

Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...

 by many moth

Moth

A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...

 caterpillar

Caterpillar

Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

s, and numerous other holometabolous insect larva

Larva

A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

e 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 (seta

Seta

Seta is a biological term derived from the Latin word for "bristle". It refers to a number of different bristle- or hair-like structures on living organisms.-Animal setae:In zoology, most "setae" occur in invertebrates....

e) and incorporate them into the cocoon; if these are urticating hair

Urticating hair

Urticating hairs, i.e. stinging hairs, are one of the primary defense mechanisms used by numerous plants, some New World tarantulas, and various lepidopteran caterpillars. Urtica is Latin for "nettle", and hairs that urticate are characteristic of this type of plant, and many other plants in...

s 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

Leaf

A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

, in a crevice, down near the base of a tree trunk

Tree Trunk

"Tree Trunk" is a song recorded and released in 1972 by The Doors."Tree Trunk" was recorded by the surviving members of The Doors as they assembled material for their second and final album, Full Circle, following the death of Jim Morrison...

, suspended from a twig or concealed in the leaf litter.

The silk in the cocoon of the silk moth can be unravelled to get silk fibre which makes this moth the most economically important of all Lepidopterans. The 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 fluids 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.

External links

• Cocoon Silk: A Natural Architecture
• Pupas photos plus species descriptions at Lepidoptera.pro