hymenopteran

• n.

• 膜翅目の昆虫の総称
• 膜状の4枚の翅をもつハチ・アリなど

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Hymenoptera Temporal range: Triassic – Recent 251–0 Ma PreЄ Є O S D C P T J K Pg N
female Netelia producta
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
(unranked):	Endopterygota
Superorder:	Hymenopterida
Order:	Hymenoptera Linnaeus, 1758
Suborders
Apocrita Symphyta

The Hymenoptera are the third largest orders of insects, comprising the sawflies, wasps, bees and ants. Over 150,000 species are recognized, with many more remaining to be described. The name refers to the wings of the insects, but the original derivation is ambiguous. All references agree that the derivation involves the Ancient Greek πτερόν (pteron) for wing. The Ancient Greek ὑμήν (hymen) for membrane provides a plausible etymology for the term because these insects have membranous wings. However, a key characteristic of this order is that the hind wings are connected to the fore wings by a series of hooks. Thus, another plausible etymology involves Hymen, the Ancient Greek god of marriage, as these insects have "married wings" in flight.

Females typically have a special ovipositor for inserting eggs into hosts or otherwise inaccessible places. The ovipositor is often modified into a stinger. The young develop through holometabolism (complete metamorphosis)—that is, they have a worm-like larval stage and an inactive pupal stage before they mature.

Evolution

Hymenoptera originated in the Triassic, the oldest fossils belonging to the family Xyelidae. Social hymenopterans appeared during the Cretaceous. The evolution of this group has been intensively studied by A. Rasnitsyn, M. S. Engel, G. Dlussky, and others.

This clade has been studied by examining the mitochondrial DNA. Although this study was unable to resolve all the ambiguities in this clade, some relationships could be established. Aculeata, Ichneumonomorpha and Proctotrupomorpha were monophyletic. The Megalyroidea andTrigonalyoidea are sister clades as are the Chalcidoidea+Diaprioidea. The Cynipoidea was generally recovered as the sister group to Chalcidoidea and Diaprioidea which are each other's closest relations.

Anatomy

Hymenopterans range in size from very small to large insects, and usually have two pairs of wings. Their mouthparts are adapted for chewing, with well-developed mandibles (ectognathous mouthparts). Many species have further developed the mouthparts into a lengthy proboscis, with which they can drink liquids, such as nectar. They have large compound eyes, and typically three simple eyes, (ocelli).

The forward margin of the hind wing bears a number of hooked bristles, or "hamuli", which lock onto the fore wing, keeping them held together. The smaller species may have only two or three hamuli on each side, but the largest wasps may have a considerable number, keeping the wings gripped together especially tightly. Hymenopteran wings have relatively few veins compared with many other insects, especially in the smaller species.

In the more ancestral hymenopterans, the ovipositor is blade-like, and has evolved for slicing plant tissues. In the majority, however, it is modified for piercing, and, in some cases, is several times the length of the body. In some species, the ovipositor has become modified as a stinger, and the eggs are laid from the base of the structure, rather than from the tip, which is used only to inject venom. The sting is typically used to immobilise prey, but in some wasps and bees may be used in defense.

The larvae of the more ancestral hymenopterans resemble caterpillars in appearance, and like them, typically feed on leaves. They have large chewing mandibles, three pairs of thoracic limbs, and, in most cases, a number of abdominal prolegs. Unlike caterpillars, however, the prolegs have no grasping spines, and the antennae are reduced to mere stubs.

The larvae of other hymenopterans, however, more closely resemble maggots, and are adapted to life in a protected environment. This may be the body of a host organism, or a cell in a nest, where the adults will care for the larva. Such larvae have soft bodies with no limbs. They are also unable to defecate until they reach adulthood due to having an incomplete digestive tract, presumably to avoid contaminating their environment.

Sex determination

Main article: Haplodiploid sex-determination system

Among most or all hymenopterans, sex is determined by the number of chromosomes an individual possesses. Fertilized eggs get two sets of chromosomes (one from each parent's respective gametes), and so develop into diploid females, while unfertilized eggs only contain one set (from the mother), and so develop into haploid males; the act of fertilization is under the voluntary control of the egg-laying female. This phenomenon is called haplodiploidy.

However, the actual genetic mechanisms of haplodiploid sex determination may be more complex than simple chromosome number. In many Hymenoptera, sex is actually determined by a single gene locus with many alleles. In these species, haploids are male and diploids heterozygous at the sex locus are female, but occasionally a diploid will be homozygous at the sex locus and develop as a male instead. This is especially likely to occur in an individual whose parents were siblings or other close relatives. Diploid males are known to be produced by inbreeding in many ant, bee and wasp species. Diploid biparental males are usually sterile but a few species that have fertile diploid males are known.

One consequence of haplodiploidy is that females on average actually have more genes in common with their sisters than they do with their own daughters. Because of this, cooperation among kindred females may be unusually advantageous, and has been hypothesized to contribute to the multiple origins of eusociality within this order. In many colonies of bees, ants, and wasps, worker females will remove eggs laid by other workers due to increased relatedness to direct siblings, a phenomenon known as worker policing.

Diet

Different species of Hymenoptera show a wide range of feeding habits. The most primitive forms are typically herbivorous, feeding on leaves or pine needles. Stinging wasps are predators, and will provision their larvae with immobilised prey, while bees feed on nectar and pollen.

A number of species are parasitoid as larvae. The adults inject the eggs into a paralysed host, which they begin to consume after hatching. Some species are even hyperparasitoid, with the host itself being another parasitoid insect. Habits intermediate between those of the herbivorous and parasitoid forms are shown in some hymenopterans, which inhabit the galls or nests of other insects, stealing their food, and eventually killing and eating the occupant.

Classification

Giant honey bee Apis dorsata on Tribulus terrestris in Hyderabad, India

The Hymenoptera are divided into two groups, the Symphyta which have no waist, and the Apocrita which have a narrow waist.

Symphyta

The suborder Symphyta includes the sawflies, horntails, and parasitic wood wasps. The group may be paraphyletic, as it has been suggested that the family Orussidae may be the group from which the Apocrita arose. They have an unconstricted junction between the thorax and abdomen. The larvae are herbivorous free-living eruciforms, with three pairs of true legs, prolegs (on every segment, unlike Lepidoptera) and ocelli. The prolegs do not have crochet hooks at the ends unlike the larvae of the Lepidoptera.

Apocrita

The wasps, bees, and ants together make up the suborder Apocrita, characterized by a constriction between the first and second abdominal segments called a wasp-waist (petiole), also involving the fusion of the first abdominal segment to the thorax. Also, the larvae of all Apocrita lack legs, prolegs, or ocelli. The hindgut of the larvae also remains closed during development, with feces being stored inside the body, with the exception of some bee larvae where the larval anus through developmental reversion has reappeared again. In general, the anus only opens at the completion of larval growth.

See also

• List of bees, wasps and ants of Great Britain
• Hymenoptera genome database
• Worker policing

References

Further reading

• Gullan, P. J.; Cranston, P. S. (2014). The Insects: An Outline of Entomology (Fifth ed.). Wiley Blackwell. 
• Grimaldi, D. and Engel, M.S. (2005). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5. 
• Rasnitsyn, A.P. and Quicke, D.L.J. (2002). History of Insects. Kluwer Academic Publishers. pp. 242–254. ISBN 1-4020-0026-X. 

External links

General

• Hymenoptera Anatomy Ontology project
• Hymenoptera Anatomy Glossary
• Hymenoptera Forum German and International
• Hymenoptera Information System (German)
• Hymenoptera of North America – large format reference photographs, descriptions, taxonomy
• International Society of Hymenopterists
• Bees, Wasps and Ants Recording Society (UK)
• Ants Photo Gallery (RU)
• International Palaeoentomological Society
• Sphecos Forum for Aculeate Hymenopterra
• Hymenoptera images on MorphBank, a biological image database
• Order Hymenoptera Insect Life Forms

Systematics

• Hymenopteran Systematics
• Hymenoptera Online 1000+ images

Regional Lists

• Insetos do Brasil
• New Zealand Hymenoptera
• Waspweb Afrotropical Hymenoptera Excellent images
• checklist of Australian Hymenoptera

Books

• Bees and Wasps and Ants, Oh My!

UpToDate Contents

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• 1. Allergen extracts: Composition, manufacture, and labeling

English Journal

• Aggregation and a strong Allee effect in a cooperative outbreak insect.

• Goodsman DW1, Koch D2, Whitehouse C3, Evenden ML1, Cooke BJ4, Lewis MA1,2.
• Ecological applications : a publication of the Ecological Society of America.Ecol Appl.2016 Dec;26(8):2621-2634. doi: 10.1002/eap.1404. Epub 2016 Nov 8.
• Most species that are negatively impacted when their densities are low aggregate to minimize this effect. Aggregation has the potential to change how Allee effects are expressed at the population level. We studied the interplay between aggregation and Allee effects in the mountain pine beetle (Dendr
• PMID 27862568

• The caste- and sex-specific DNA methylome of the termite Zootermopsis nevadensis.

• Glastad KM1,2, Gokhale K3, Liebig J4, Goodisman MA2.
• Scientific reports.Sci Rep.2016 Nov 16;6:37110. doi: 10.1038/srep37110.
• Epigenetic inheritance plays an important role in mediating alternative phenotype in highly social species. In order to gain a greater understanding of epigenetic effects in societies, we investigated DNA methylation in the termite Zootermopsis nevadensis. Termites are the most ancient social insect
• PMID 27848993

• Developmental DNA methyltransferase expression in the fire ant Solenopsis invicta.

• Kay S1, Skowronski D1, Hunt BG2.
• Insect science.Insect Sci.2016 Oct 24. doi: 10.1111/1744-7917.12413. [Epub ahead of print]
• DNA methylation is accomplished in animals by two classes of enzymes known as DNA methyltransferases, DNMT3 and DNMT1, which perform de novo methylation and maintenance methylation, respectively. Several studies of hymenopteran eusocial insects suggest that DNA methylation is capable of influencing
• PMID 27774769

Japanese Journal

• A new antigenic marker specifically labels a subpopulation of the class II Kenyon cells in the brain of the European honeybee Apis mellifera

• ,
• BIOPHYSICS 11(0), 73-77, 2015
• … In the social hymenopteran insects such as honeybees, the mushroom bodies are the prominent brain structures. …
• NAID 130004940816

• Sexually Dimorphic Body Color Is Regulated by Sex-Specific Expression of Yellow Gene in Ponerine Ant, Diacamma Sp.

• Miyazaki Satoshi,Okada Yasukazu,Miyakawa Hitoshi,Tokuda Gaku,Cornette Richard,Koshikawa Shigeyuki,Maekawa Kiyoto,Miura Toru
• PLOS ONE 9(3), e92875, 2014-03-25
• … Most hymenopteran species exhibit conspicuous sexual dimorphism due to ecological differences between the sexes. … As hymenopteran genomes, under the haplodiploid genetic system, exhibit quantitative differences between sexes while remaining qualitatively identical, sexual phenotypes are assumed to be expressed through sex-specific gene usage. …
• NAID 120005429614

• Insecticide susceptibility and carboxylesterase activity in leafminers (Diptera: Agromyzidae) and their associated hymenopteran parasitoids

• Matsuda Kentaro,Saito Tsutomu
• Crop Protection 55, 50-54, 2013-10-31
• NAID 120005348753

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