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 causing the death of these hosts. Different species specialise in hosts from different insect orders, most often Lepidoptera, though some select beetles, flies, or bugs; the spider wasps (Pompilidae) exclusively attack spiders. Parasitoid wasp species differ in which host life-stage they attack: eggs, larvae, pupae, or adults. They mainly follow one of two major strategies within parasitism: either they are endoparasitic, developing inside the host, and koinobiont, allowing the host to continue to feed, develop, and moult; or they are ectoparasitic, developing outside the host, and idiobiont, paralysing the host immediately. Some endoparasitic wasps of the superfamily Ichneumonoidea have a mutualistic relationship with polydnaviruses, the viruses suppressing the host's immune defenses. Parasitoidism evolved only once in the Hymenoptera, during the Permian, leading to a single clade, but the parasitic lifestyle has secondarily been lost several times including among the ants, bees, and yellowjacket wasps. As a result, the order Hymenoptera contains many families of parasitoids, intermixed with non-parasitoid groups. The parasitoid wasps include some very large groups, some estimates giving the Chalcidoidea as many as 500,000 species, the Ichneumonidae 100,000 species, and the Braconidae up to 50,000 species. Host insects have evolved a range of defences against parasitoid wasps, including hiding, wriggling, and camouflage markings. Many parasitoid wasps are considered beneficial to humans because they naturally control agricultural pests. Some are applied commercially in biological pest control, starting in the 1920s with Encarsia formosa to control whitefly in greenhouses. Historically, parasitoidism in wasps influenced the thinking of Charles Darwin. Parasitoidism Parasitoid wasps range from some of the smallest species of insects to wasps about an inch long. Most females have a long, sharp ovipositor at the tip of the abdomen, sometimes lacking venom glands, and almost never modified into a sting. Parasitoids can be classified in a variety of ways. They can live within their host's body as endoparasitoids, or feed on it from outside as ectoparasitoids: both strategies are found among the wasps. Parasitoids can also be divided according to their effect on their hosts. Idiobionts prevent further development of the host after initially immobilizing it, while koinobionts allow the host to continue its development while they are feeding upon it;