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260 salt, and in many species the larvae are truly aquatic. Larvae of some species feed on algae, but generally they and the adults are carnivorous or carrion feeders, sometimes almost to the point of being parasitic. Two examples, illustrating the extreme habitats in which Ephydridae are found, are Ephydra riparia, which is found in the Great Salt Lake of Utah (see also Chapter 18, Section 4.3), and Psilopa (Helaeomyia) petrolei, whose larvae live in pools of crude petroleum in California. The closely related DROSOPHILIDAE (pomace or fruit flies) are small flies generally seen in the vicinity of decaying vegetation or fruit, or near breweries and vinegar factories. The larvae are mostly fungivorous, though a few are leaf miners or prey on other insects. Various species of Drosophila have, of course, been extensively used for a wide range of biological research.

Subdivision Calyptratae

How the Calyptratae should be subdivided remains debatable; some authors lump all families in the Muscoidea while others, including McAlpine et al. (1981-1989), believe that there are three monophyletic subgroups in the subdivision, Muscoidea sensu stricto, Oestroidea, and Hippoboscoidea (= Pupiparia of earlier authors).

Superfamily Muscoidea

In the cosmopolitan family MUSCIDAE (3000 species) are many common pests, for example, the Australian bush fly (Musca vetustissima), house flies [Musca domestica (Figure 9.17A) and Fannia canicularis], and bloodsucking species such as the stable fly (Stomoxys calcitrans) and face fly (Musca autumnalis). However, the great majority of species are non-pestiferous. Adults are predaceous, saprophagous, pollenophagous, hematophagous, or feed on exudates of mammals. Females are typically oviparous, though a few are ovoviviparous or larviparous. Larvae are mostly saprophagous or dung feeders, though some are predaceous and a few are ectoparasites on birds. Most of the 1000 species of the primarily holarctic group ANTHOMYIIDAE (root maggot flies) are phytophagous in the larval stage and, as a result, many are economically important, for example, the cabbage

FIGURE 9.17. Muscoidea. (A) The house fly, Musca domestica (Muscidae); (B) the wheat bulb fly, Hylemya coarctata (Anthomyiidae); and (C) a dung fly, Cordilura criddlei (Scathophagidae). (A, from V. B. Wigglesworth, 1959, Metamorphosis, polymorphism, differentiation, Scientific American, February 1959. By permission of Mr. Eric Mose, Jr. B, C, from F. R. Cole and E. I. Schlinger, 1969, The Flies of Western North America. By permission of the University of California Press.]

FIGURE 9.17. Muscoidea. (A) The house fly, Musca domestica (Muscidae); (B) the wheat bulb fly, Hylemya coarctata (Anthomyiidae); and (C) a dung fly, Cordilura criddlei (Scathophagidae). (A, from V. B. Wigglesworth, 1959, Metamorphosis, polymorphism, differentiation, Scientific American, February 1959. By permission of Mr. Eric Mose, Jr. B, C, from F. R. Cole and E. I. Schlinger, 1969, The Flies of Western North America. By permission of the University of California Press.]

root fly, Hylemya brassicae, and the wheat bulb fly, H. coarctata (Figure 9.17B). Others are saprophagous, dung feeders, or are inquilines in the burrows of solitary Hymenoptera or rodents. The small, primarily holarctic family SCATHOPHAGIDAE (dung flies) (500 species) is considered the most primitive of the Calyptratae. Adults (Figure 9.17C) are mostly predaceous on other insects, though some feed on the juice of dung. Larvae have varied habits; many are leaf and stem miners, others predaceous, and some feed on dung.

Superfamily Oestroidea

Five families are included in this very large group of Diptera. CALLIPHORIDAE (1000 species, 80% of which are restricted to the Old World) is a cosmopolitan group that includes blow flies, green- and bluebottles, and screwworm flies. Among members of the family, which may be paraphyletic (Rognes, 1997), a complete spectrum of larval feeding habits can be seen, ranging from true carrion feeders, through species that feed on exudates or open wounds of living animals, to truly parasitic forms. Calliphoridae of medical or veterinary importance include the sheep blow flies (Lucilia spp.), the screwworms [Cochliomyia (Callitroga) spp.] (Figure 9.18A), and bluebottles (Calliphora spp.), which are vectors of human diseases. Closely related to the calliphorids are the SARCOPHAGIDAE (flesh flies) (Figure 9.18B), whose larvae feed on decaying animal tissue or are true parasites of arthropods, mollusks, or annelids. Most of the 2000 species in this cosmopolitan group are viviparous, depositing first-instar larvae directly into the food source. The TACHINIDAE (Figure 9.18C,D), with some 8000 species worldwide, form the second largest family of Diptera. Without exception, the larvae are parasitic on other arthropods, mainly insects. An egg, or in the many viviparous species, a larva, is frequently deposited directly on the body of the host. Alternatively, the egg is laid on the host's food plant. The host usually dies as a result of the parasitism, and there is little doubt that tachinids play a role equal to that of many parasitic Hymenoptera in controlling the population level of certain species. Not surprisingly, some have been employed as biological control agents against pests (for examples, see Table 24.6). Included in the OESTRIDAE (bot flies and warble flies) (Figure 9.18E-G), a group of about 150 species, are four well-defined subfamilies that are often given family rank. OESTRINAE are holarctic and African flies that larviposit in the nasal and pharyngeal cavities of large herbivores; HYPODERMATINAE have a similar distribution, their hosts include rodents as well as herbivores, females oviposit on the host's skin, and the larvae develop subcutaneously; CUTEREBRINAE are restricted to the New World where they oviposit in places frequented by the host, the larvae hatching in response to radiant heat from an adjacent host and developing subcutaneously on primates, rodents, and lagomorphs; and the cosmopolitan GASTEROPHILINAE mostly oviposit on the legs or near the mouth of horses, zebras, and elephants, the larvae eventually making their way to the host's stomach or intestine where they complete development.

Superfamily Hippoboscoidea

Four families are included in this group, GLOSSINIDAE, HIPPOBOSCIDAE, STREBLIDAE, and NYCTERIBIIDAE, the last three formerly being considered to constitute the Pupiparia. In all of the families the adults are bloodsucking parasites of birds or mammals, and the larvae mature, one at a time, entirely within the genital tract of the female,

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