Hypermetamorphosis

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Hypermetamorphosis, or heteromorphosis, [1] is a term used mainly in entomology; it refers to a class of variants of holometabolism, that is to say, complete insect metamorphosis. Hypermetamorphosis is exceptional in that some instars, usually larval instars, are functionally and visibly distinct from the rest. The differences between such instars usually reflect transient stages in the life cycle; for instance, one instar might be mobile while it searches for its food supply, while the following instar immediately sheds its locomotory organs and settles down to feed until it is fully grown and ready to change into the reproductive stage, which in turn, does not have the same nutritional requirements as the larvae.

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Description

These planidia are fairly typical of certain species of parasitoid wasps, in this case the genus Perilampus. Hypermetamorphosis2.JPG
These planidia are fairly typical of certain species of parasitoid wasps, in this case the genus Perilampus .
Hypermetamorphosis in Meloidae. This form of planidium is a typical triungulin and feeds in its first instar. In its second instar it turns into a less triungulin-like form, and feeds again. It then turns into the Scarabaeoid form for two or more instars, depending on species. After that it adopts pre-pupal forms, pupates, and finally emerges as an adult beetle. Hypermetamorphosis in Meloidae.JPG
Hypermetamorphosis in Meloidae. This form of planidium is a typical triungulin and feeds in its first instar. In its second instar it turns into a less triungulin-like form, and feeds again. It then turns into the Scarabaeoid form for two or more instars, depending on species. After that it adopts pre-pupal forms, pupates, and finally emerges as an adult beetle.
Triungulin on a butterfly. This probably is an example of phoresy rather than parasitism. Triungulin on a butterfly (5739202307).jpg
Triungulin on a butterfly. This probably is an example of phoresy rather than parasitism.

Hypermetamorphosis, as the term normally is used in entomology, refers to a class of variants of holometabolism. In hypermetamorphosis some larval instars are functionally and morphologically distinct from each other.

The general case in holometabolous insects such as flies, moths, or wasps, is that all larval stages look similar, growing larger as the insect matures. In hypermetamorphic insects however, at least one instar, usually the first, differs markedly from the rest. In many hypermetamorphic species, the first instars are numerous, tiny, very mobile larvae that must find their way to a food source. The general term for a mobile first instar is a planidium, from the Greek language πλάνος (planos) meaning "roaming".

In typical examples the first-instar larval morphology is campodeiform (meaning: elongated, flattened, and active, more or less resembling the morphology of hexapods in the genus Campodea ). There is however, considerable variety in the forms of planidia that occur in various families and orders; in the beetle family Meloidae, the three-clawed planidium originally was called a triungulin, and similar planidia for example, those of the Strepsiptera, may also be called triungula.

In their planidial form, many species do not feed; they first molt into a form suited to eating rather than seeking out food. The second instar is completely different in appearance and behavior, often becoming grub- or maggot-like in the instars before pupation. As a rule, the instars after the first ecdysis are of more or less constant form and not highly mobile, being specialised for feeding and growth until the final larval instar metamorphoses into the pupal form. [2]

The moth family Gracillariidae is an unusual case of hypermetamorphosis in that its first few larval instars do feed, but differently to later instars. Specifically, early instars feed on plant sap (and for this reason have modified mandibles) while later instars feed on plant tissue. Furthermore, instead of becoming less mobile during the larval stage, larvae become more mobile, with early instars lacking legs while later instars possess legs. [3]

There are examples of holometabolic species in which there are certain striking differences between the earliest instars and the later instars, though without their generally being regarded as hypermetamorphic. For example, early instars of many Papilionidae are of a colour, shape and texture that suggest bird droppings; later instars that are larger and would simply stand out in such camouflage, typically become green. [4] The prepupa or last larval instar of insects ceases to feed and (in some cases) searches for a place to pupate, [5] [6] but this also is not considered hypermetamorphosis.

Vv cm early instar.jpg
Papilio Polyetes 08062012 (1).jpg
Early instars of Papilio polytes (left) resemble a bird dropping, while the late ones (right) are too big to do so, and are simply camouflaged as a leaf. This striking coloration difference between the instars is generally not regarded as hypermetamorphosis.

Various forms of hypermetamorphosis

Hypermetamorphosis usually occurs as an adaptation of the ontogeny of certain parasitoid insects, notably:

Technically, the subimago of the Ephemeroptera might be described as a stage in a form of hypermetamorphosis, but that is not common practice. [7]

Examples of hypermetamorphosis in any given insect order are analogous and not homologous to those in any other order; for example, hypermetamorphosis in the Acroceridae was not derived from the Strepsiptera.

Related Research Articles

<span class="mw-page-title-main">Metamorphosis</span> Profound change in body structure during the postembryonic development of an organism

Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some insects, jellyfish, fish, amphibians, mollusks, crustaceans, cnidarians, echinoderms, and tunicates undergo metamorphosis, which is often accompanied by a change of nutrition source or behavior. Animals can be divided into species that undergo complete metamorphosis ("holometaboly"), incomplete metamorphosis ("hemimetaboly"), or no metamorphosis ("ametaboly").

<span class="mw-page-title-main">Strepsiptera</span> Order of insects

The Strepsiptera are an order of insects with eleven extant families that include about 600 described species. They are endoparasites of other insects, such as bees, wasps, leafhoppers, silverfish, and cockroaches. Females of most species never emerge from the host after entering its body, finally dying inside it. The early-stage larvae do emerge because they must find an unoccupied living host, and the short-lived males must emerge to seek a receptive female in her host. They are believed to be most closely related to beetles, from which they diverged 300–350 million years ago, but do not appear in the fossil record until the mid-Cretaceous around 100 million years ago.

<span class="mw-page-title-main">Larva</span> Juvenile form of distinct animals before metamorphosis

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

<span class="mw-page-title-main">Pupa</span> Life stage of some insects undergoing transformation

A pupa is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages thereof being egg, larva, pupa, and imago. The processes of entering and completing the pupal stage are controlled by the insect's hormones, especially juvenile hormone, prothoracicotropic hormone, and ecdysone. The act of becoming a pupa is called pupation, and the act of emerging from the pupal case is called eclosion or emergence.

<span class="mw-page-title-main">Instar</span> Developmental stage of arthropods between moults

An instar is a developmental stage of arthropods, such as insects, which occurs between each moult (ecdysis) until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, changes in the number of body segments or head width. After shedding their exoskeleton (moulting), the juvenile arthropods continue in their life cycle until they either pupate or moult again. The instar period of growth is fixed; however, in some insects, like the salvinia stem-borer moth, the number of instars depends on early larval nutrition. Some arthropods can continue to moult after sexual maturity, but the stages between these subsequent moults are generally not called instars.

<span class="mw-page-title-main">Holometabola</span> Superorder of insects

Holometabola, also known as Endopterygota, is a supra-ordinal clade of insects within the infraclass Neoptera that go through distinctive larval, pupal, and adult stages. They undergo a radical metamorphosis, with the larval and adult stages differing considerably in their structure and behaviour. This is called holometabolism, or complete metamorphism.

<i>Manduca sexta</i> Species of moth

Manduca sexta is a moth of the family Sphingidae present through much of the Americas. The species was first described by Carl Linnaeus in his 1763 Centuria Insectorum.

<span class="mw-page-title-main">Saturniidae</span> Family of moths

Saturniidae, members of which are commonly named the saturniids, is a family of Lepidoptera with an estimated 2,300 described species. The family contains some of the largest species of moths in the world. Notable members include the emperor moths, royal moths, and giant silk moths.

Larviform female is a biological phenomenon occurring in some insect species, where the females in the adult stage of metamorphosis resemble the larvae to various degrees, while the male appears more morphologically adult. The resemblance may mean the larviform female has the same coloring as the larvae and/or similar body plans, and may be the result of the female arresting development at earlier stages of ecdysis than males. The female may not pupate at all, as in Xenos vesparum. Typically, the female is wingless and generally larger than the male. Larviform females still reach sexual maturity. Larviform females occur in several insect groups, including most Strepsiptera and Bagworm moths, many elateroid beetles, and some gall midges.

Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago. Holometabolism is a synapomorphic trait of all insects in the superorder Holometabola. Immature stages of holometabolous insects are very different from the mature stage. In some species the holometabolous life cycle prevents larvae from competing with adults because they inhabit different ecological niches. The morphology and behavior of each stage are adapted for different activities. For example, larval traits maximize feeding, growth, and development, while adult traits enable dispersal, mating, and egg laying. Some species of holometabolous insects protect and feed their offspring. Other insect developmental strategies include ametabolism and hemimetabolism.

<span class="mw-page-title-main">Varied carpet beetle</span> Species of beetle

The varied carpet beetle is a 3 mm-long beetle belonging to the family Dermestidae, positioned in subgenus Nathrenus. They are a common species, often considered a pest of domestic houses and, particularly, natural history museums, where the larvae may damage natural fibers and can damage carpets, furniture, clothing, and insect collections. A. verbasci was also the first insect to be shown to have an annual behavioral rhythm and to date remains a classic example of circannual cycles in animals.

<i>Manduca quinquemaculata</i> Species of moth

Manduca quinquemaculata, the five-spotted hawkmoth, is a brown and gray hawk moth of the family Sphingidae. The caterpillar, often referred to as the tomato hornworm, can be a major pest in gardens; they get their name from a dark projection on their posterior end and their use of tomatoes as host plants. Tomato hornworms are closely related to the tobacco hornworm Manduca sexta. This confusion arises because caterpillars of both species have similar morphologies and feed on the foliage of various plants from the family Solanaceae, so either species can be found on tobacco or tomato leaves. Because of this, the plant on which the caterpillar is found does not indicate its species.

<span class="mw-page-title-main">Acroceridae</span> Family of flies

The Acroceridae are a small family of odd-looking flies. They have a hump-backed appearance with a strikingly small head, generally with a long proboscis for accessing nectar. They are rare and not widely known. The most frequently applied common names are small-headed flies or hunch-back flies. Many are bee or wasp mimics. Because they are parasitoids of spiders, they also are sometimes known as spider flies.

<span class="mw-page-title-main">Planidium</span> Form of insect larva

A planidium is a specialized form of insect larva seen in the first-instar of a few families of insects that have parasitoidal ways of life. They are usually flattened, highly sclerotized (hardened), and quite mobile. The function of the planidial stage is to find a host on which the later larval instars may feed, generally until the insect pupates.

<i>Acronicta rumicis</i> Species of moth

Acronicta rumicis, the knot grass moth, is a species of moth which is part of the genus Acronicta and family Noctuidae. It was first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae. It is found in the Palearctic region. A. rumicis lives and feeds on plants located in wide-open areas. At its larval stage, as a caterpillar, it causes such a large impact as a crop pest that it has received much attention and research. A. rumicis feeds on maize, strawberries and other herbaceous plants.

<i>Synthesiomyia nudiseta</i> Species of fly

Synthesiomyia nudiseta is one of the largest flies in the family Muscidae. The fly has a pair of forewings; the paired hind wings have been reduced to halteres that help with stability and movement during flight. Key characteristics of this species include plumose segmented aristae, well-developed calypters, and sternopleural bristles. Synthesiomyia nudiseta is a forensically important species because it is necrophilous and can therefore help determine the time of colonization for the post mortem interval with its known life cycle.

<i>Spodoptera eridania</i> Species of moth

Spodoptera eridania is a moth that is known to be a pest. They are one of the most important defoliators in the tropical and subtropical regions of the western hemisphere that feed heavily on plants while they are young, often resulting in skeleton leaves on their food plants. They are also heavy feeders on tomato in Florida. There is a lot of development in producing pesticides against the S. eridania, specifically a neem-based pesticide that can result in smaller and prolonged development. The wingspan is 33–38 mm. Adults are on wing year-round. The larvae feed on various weeds but prefer Amaranthus species and Phytolacca americana.

<i>Samea multiplicalis</i> Species of moth

Samea multiplicalis, the salvinia stem-borer moth, is an aquatic moth commonly found in freshwater habitats from the southern United States to Argentina, as well as in Australia where it was introduced in 1981. Salvinia stem-borer moths lay their eggs on water plants like Azolla caroliniana, Pistia stratiotes, and Salvinia rotundifolia. Larval feeding on host plants causes plant death, which makes S. multiplicalis a good candidate for biological control of weedy water plants like Salvinia molesta, an invasive water fern in Australia. However, high rates of parasitism in the moth compromise its ability to effectively control water weeds. S. multiplicalis larvae are a pale yellow to green color, and adults develop tan coloration with darker patterning. The lifespan, from egg to the end of adulthood is typically three to four weeks. The species was first described by Achille Guenée in 1854.

<i>Hemileuca lucina</i> Species of moth

Hemileuca lucina, the New England buck moth, is a species of moth in the family Saturniidae. This moth species is only found in the New England region of the United States. Larvae in early stages mainly feed on broadleaf meadowsweet whereas larvae in later stages show variation in food sources such as blackberry and black cherry leaves. Larvae have a black body with orange/black spines on their back that are used to deter predators. Pupation occurs during the summer and adult moths come out around September.

<i>Meloe americanus</i> Species of beetle

Meloe americanus is a type of blister beetle (Meloidae) found in North America. It is most relevant to the fields of agriculture and veterinary medicine. Adult beetles feed on different types of plants, which cause crop damage. They also release a fluid containing a chemical that is toxic, and at high concentrations lethal, to mammals. The first instar larvae are uniquely active and mobile, utilizing phoresy and parasitism to feed and mature through their developmental stages.

References

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