Temporal range: Triassic – Recent 251–0Ma
|female Netelia producta|
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, and is derived from the Ancient Greek ὑμήν (hymen): membrane and πτερόν (pteron): wing. The hind wings are connected to the fore wings by a series of hooks called hamuli.
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.
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 closest relations.
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.
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.
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.
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.
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 do not have 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.
- Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed. Oxford University Press. p. 320. ISBN 0-19-510033-6.
- Mao M, Gibson T, Dowton M (2014) Higher-level phylogeny of the Hymenoptera inferred from mitochondrial genomes. Mol Phylogenet Evol
- Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed. Oxford University Press. pp. 570–579. ISBN 0-19-510033-6.
- David P. Cowan and Julie K. Stahlhut (July 13, 2004). "Functionally reproductive diploid and haploid males in an inbreeding hymenopteran with complementary sex determination". PNAS 101 (28): 10374–9. doi:10.1073/pnas.0402481101. PMC 478579. PMID 15232002.
- Elias, J.; Mazzi, D.; Dorn, S. (2009). Bilde, Trine, ed. "No Need to Discriminate? Reproductive Diploid Males in a Parasitoid with Complementary Sex Determination". PLoS ONE 4 (6): e6024. doi:10.1371/journal.pone.0006024. PMC 2696080. PMID 19551142.
- Davies, N.R., Krebs, J.R., and West, S.A. An Introduction to Behavioral Ecology. 4th ed. West Sussex: Wiley-Blackwell, 2012. Print. pp. 387–388
- The Evolution of Social Wasps
- 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.
- Aguiar, A.P., Deans, A.R., Engel, M.S., Forshage, M., Huber, J.T., Jennings, J.T., Johnson, N.F., Lelej, A.S., Longino, J.T., Lohrmann, V., Mikó, I., Ohl, M., Rasmussen, C., Taeger A. & Yu, D.S.K. 2013, Order Hymenoptera Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal Biodiversity: An Outline of Higher-level Classification and Survey of Taxonomic Richness (Addenda 2013). Zootaxa, 3703, 1–82.
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