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 typically have a special ovipositor for inserting eggs into hosts or places that are otherwise inaccessible. This ovipositor is often modified into a stinger. The young develop through holometabolism (complete metamorphosis)—that is, they have a wormlike larval stage and an inactive pupal stage before they reach adulthood.
Etymology The name Hymenoptera comes from Ancient Greek ὑμήν (humḗn) 'membrane' and πτερόν (pterón) 'wing'.
Evolution Molecular analysis finds that Hymenoptera is the earliest branching group of Holometabola.
Hymenoptera originated in the Triassic, with the oldest fossils belonging to the family Xyelidae. Social hymenopterans appeared during the Cretaceous. The evolution of this group has been intensively studied by Alex Rasnitsyn, Michael S. Engel, and others.
Phylogenetic relationships within the Hymenoptera, based on both morphology and molecular data, have been intensively studied since 2000. In 2023, a molecular study based on the analysis of ultra-conserved elements confirmed many previous findings and produced a relatively robust phylogeny of the whole Order, although the relationships among the Symphyta lineages are still a source of active research . Basal superfamilies are shown in the cladogram below.
Anatomy
Hymenopterans range in size from very small to large insects. 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. Hymenopterans usually have two pairs of wings, but some solitary wasps and worker ants don't. They typically have large compound eyes with 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 (hamuli wing coupling). 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 immobilize prey, but in some wasps and bees may be used in defense. Hymenopteran larvae typically have a distinct head region, three thoracic segments, and usually nine or 10 abdominal segments. In the suborder Symphyta, the eruciform larvae 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, six or eight abdominal prolegs. Unlike caterpillars, however, the prolegs have no grasping spines, and the antennae are reduced to mere stubs. Symphytan larvae that are wood borers or stem borers have no abdominal legs and the thoracic legs are smaller than those of non-borers. With rare exceptions, larvae of the suborder Apocrita have no legs and are maggotlike in form, 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. In parasitic forms, the head is often greatly reduced and partially withdrawn into the prothorax (anterior part of the thorax). Sense organs appear to be poorly developed, with no ocelli, very small or absent antennae, and toothlike, sicklelike, or spinelike mandibles. They are also unable to defecate until they reach adulthood due to having an incomplete digestive tract (a blind sac), presumably to avoid contaminating their environment. The larvae of stinging forms (Aculeata) generally have 10 pairs of spiracles, or breathing pores, whereas parasitic forms usually have nine pairs present.
Reproduction
Sex determination
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 develop into diploid females, while unfertilized eggs only contain one set (from the mother) and develop into haploid males. The act of fertilization is under the voluntary control of the egg-laying female, giving her control of the sex of her offspring. 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 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 have more genes in common with their sisters than they do with their 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. Another consequence is that hymenopterans may be more resistant to the deleterious effects of inbreeding. As males are haploid, any recessive genes will automatically be expressed, exposing them to natural selection. Thus, the genetic load of deleterious genes is purged relatively quickly.
Thelytoky
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