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Description & Behavior
Hymenoptera is one of the largest orders of insects that includes the sawflies, wasps, bees, and ants. There are over 130,056 identified species. The name relates to the heavy wings of the insects, and comes from the Greek for hymen membrane and pteron for wing. The hindwings are connected to the forewings by a series of hooks called hamuli.
Females usually have a special ovipositor for inserting eggs into hosts or otherwise remote places. The ovipositor also is usually modified into a stinger. The immatures develop through complete metamorphosis with a worm-like larval stage and an inactive pupal stage before they mature.
The order dates from the Triassic, the oldest fossils belonging to the family Xyelidae. Social hymenopterans first appeared during the Cretaceous. The evolution of this group has been intensively studied by A. Rasnitsyn, M. S. Engel, G. Dlussky (see References).
The size ranges from medium to large, usually with two pairs of wings. The mouthparts are for chewing, with well-developed mandibles. Many species have further developed the mouthparts into a lengthy proboscis, with which they can imbibe liquids, such as nectar. They have large compound eyes, and usually three ocelli.
The forward margin of the hindwing bears a number of hooked bristles, or "hamuli", which lock onto the forewing, holding them 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 very tightly. The wings have few veins compared with many other insects, especially in the smaller species.
In the more primitive species, the ovipositor takes the shape of a blade, and has adapted to slicing plant tissues. However most species have it modified for piercing, and, in some cases, is several times the length of the body. The ovipositor may also 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 for boring and to inject venom. The stinger is used to immobilise prey, but in some wasps and bees may be used for defence.
The larvae of the primitive species resemble caterpillars, and like them, they feed on leaves. They have large chewing mandibles, three thoracic limbs, and usually a number of abdominal prolegs. But unlike caterpillars, the prolegs have no grasping spines, and the antennae are greatly reduced.
The larvae of other Hymenoptera more closely resemble maggots, and are adapted to life in a protected environment. This may be the body of a host, or a cell in a nest, where the adults will care for the larva (e.g., Bethylidae). Such larvae have soft bodies with no limbs. Defecation occurs when they reach adulthood due to having an incomplete digestive tract, probably to avoid contaminating their surroundings.
Sex is determined by the number of chromosomes that an insect has. Fertilized eggs receive two sets. 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, a process that is known as haplodiploidy.
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 into a male. This is common in an individual whose parents were siblings or other close relatives. Diploid males are known to be produced by inbreeding in ants, bees and wasps.
One result 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 may contribute to the multiple origins of eusociality within this order. Different species show a wide range of feeding habits. The most primitive forms are usually herbivorous, feeding on leaves or pine needles. Stinging wasps are predators, and will provide their larvae with immobilised prey, while bees feed on nectar and pollen
Quite a few species are parasitoids as larvae. The adults inject the eggs into a paralysed host, which the larvae begin to consume after hatching. Some species are even hyperparasitoids, with the host itself being another parasitoid insect. Habits intermediate between those of the herbivorous and parasitoid forms occur in some species that inhabit the galls or nests of other insects, utilizing their food, and even killing and devouring the occupant.
The suborder Symphyta includes the sawflies, horntails, and parasitic wood wasps. The group may be paraphyletic, as it has been thought that the family Orussidae may be the ancestral group of the Apocrita. They have an unconstricted junction between the thorax and abdomen, and the larvae of free-living forms are herbivorous, have legs, prolegs (on every segment, unlike Lepidoptera and ocelli. The wasps, bees, and ants together make up the suborder Apocrita, characterized by a constriction between the first and second abdominal segments called a petiole, that also involves the fusion of the firs abdominal segment to the thorax. Also, the larvae of all Apocrita do not have legs, prolegs, or ocelli
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