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A planidium is a specialized form of
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
larva A larva (; : larvae ) 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 ...
seen in the first-
instar An instar (, from the Latin '' īnstar'' 'form, likeness') 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 ...
of a few families of insects that have
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
al 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
pupa A pupa (; : pupae) 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 th ...
tes.


Etymology

The term "planidium" is derived from the
Greek language Greek (, ; , ) is an Indo-European languages, Indo-European language, constituting an independent Hellenic languages, Hellenic branch within the Indo-European language family. It is native to Greece, Cyprus, Italy (in Calabria and Salento), south ...
πλανής (planis) meaning "wanderer", the same origin of the word ''
planet A planet is a large, Hydrostatic equilibrium, rounded Astronomical object, astronomical body that is generally required to be in orbit around a star, stellar remnant, or brown dwarf, and is not one itself. The Solar System has eight planets b ...
.'' The term
planula A planula is the free-swimming, flattened, ciliated, bilaterally symmetric larval form of various cnidarian species and also in some species of Ctenophores, which are not related to cnidarians at all. Some groups of Nemerteans also produce larva ...
was similarly derived in reference to the wandering larvae of certain
Cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
. Accordingly, "planidium" is the general term for such an adaptation, and it is not limited to any particular species or
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
. Planidia of different species differ variously from each other in form. The first-instar larva in the beetle family Meloidae has three claws on each foot, and is therefore called a triungulin (plural ''triungula''). The term is derived from the Latin ''tri'' meaning "three", and ''ungula'' meaning "a claw". An
obsolescent Obsolescence is the process of becoming antiquated, out of date, old-fashioned, no longer in general use, or no longer useful, or the condition of being in such a state. When used in a biological sense, it means imperfect or rudimentary when comp ...
variant ''triungulus'' (plural ''triunguli'') may still be encountered. For practical purposes of uniform terminology, except where there is some special reason for the use of the more narrowly specific term "triungulin", it is best to use only the more general term "planidium".


Function and occurrence

Planidia occur among subsets of the members of several orders, including:
Neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
,
Hymenoptera Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typi ...
,
Coleoptera Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
,
Strepsiptera 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, Zygentoma, silverfish, and cockroaches. Females of most s ...
, and
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advance ...
. Examples include: the neuropteran family Mantispidae; the
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
families Meloidae and Ripiphoridae; and the fly families
Acroceridae 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 ...
,
Bombyliidae The Bombyliidae are a family of flies, commonly known as bee flies. Some are colloquially known as bomber flies. Adults generally feed on nectar and pollen, some being important pollinators. Larvae are mostly parasitoids of other insects. Over ...
, Nemestrinidae, and
Tachinidae The Tachinidae are a large and variable family of true fly, flies within the insect order Fly, Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in t ...
. Among the Hymenoptera examples include the
parasitic wasp Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps ( Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causi ...
families Eucharitidae and Perilampidae. All
Strepsiptera 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, Zygentoma, silverfish, and cockroaches. Females of most s ...
have planidial larvae. The term "triungulin", originally coined in reference to the planidia of the beetle family Meloidae, is commonly applied to similar-looking planidial larvae of other families of beetles or of Strepsiptera. It is purely descriptive, and of no theoretical importance; without implying any conceptual difference from other planidia.


Planidial behaviour

Depending on their species, planidial larvae either wait for a passing host, or actively seek one out. In many species, the planidia depend on
phoresy Phoresis or phoresy is a temporary commensalistic relationship when an organism (a phoront or phoretic) attaches itself to a host organism solely for travel. It has been seen in ticks and mites since the 18th century, and in fossils 320  ...
to gain access to the actual host life stage. For instance, they may ride on the adult form of the host, or on an intermediate vector that might carry them to where their later instars might feed till they are ready for pupation. Typically, such a planidium then enters the body of the host larva, but some of the species attack host eggs; for example, some Meloidae feed on the subterranean egg pods of grasshoppers and locusts, and Mantispidae feed on egg purses of spiders. A striking example of phoresy is that planidia of beetles of the genus '' Meloe'' will form a group and produce a
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
that mimics the sex attractant of its host bee species; when the male bee arrives and attempts to mate with the mass of larvae, they climb onto his abdomen, and from there, they transfer in turn to a female bee, and finally to the bee nest, where they attack the bee larvae as their hosts. It is common for planidia to molt shortly after entering the host body or nest, but some species postpone further development while the host larva grows. Whether after a delay or not, the first
ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remnant ...
changes the planidial form into an extra larval form that is remarkably different from the planidium; this reflects the lapsed need for the larva to wander any further, together with an increased need for efficiency in feeding. The changes in
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
usually include de-
sclerotization Sclerotization is a biochemical process that produces the rigid shell of sclerotin that comprises an insect's chitinous exoskeleton. It is prominent in the thicker, armored parts of insects and arachnid Arachnids are arthropods in the Class (b ...
, and loss of the legs and eyes of the larvae. Inclusion of the extra, functionally distinct form of larva into the life history is an example of hypermetamorphosis


See also

* * * * * *


References


External links

{{Commons category-inline Parasitic insects Insect physiology Larvae