Calliphoridae

Blow fly adults are 8-10 mm long, and generally have a metallic blue, green, coppery green, or greenish-black sheen to the thorax and abdomen. An exception is the cluster fly, Pollenia rudis,which is grayish black. In general, the flies that are metallic blue are Calliphora, and those that are metallic green are Lucilia. Full-grown larvae are 10-18 mm long and yellowish white; there are complete bands of small spines around the body on most segments. Larval anterior spiracles have 6-12 lobes; posterior spiracles are located dorsally, but in some species the spiracles are in a slight caudal depression. The posterior segment of the larva is usually surrounded with cone-

shaped tubercles, and the posterior spiracles have a complete peritreme, and three slits. Larvae usually feed on dead animals or garbage containing meat; some are scavengers on excrement and plant material, while others are parasites. Female calliphorids can locate a suitable substrate for oviposition in almost any habitat.

Oviposition begins 3-7 days after emergence. At 24 °C the preoviposition period ranges from 4-5 days for Calliphora to 6-7 days for Phormia. Eggs are usually deposited in large batches with as many as 200 eggs. Females are very efficient in locating suitable breeding sites, and if many adults are present there may be an excessive number of eggs deposited in one location. Adults rarely enter buildings to oviposit, but they will enter through a door or window when attracted to a breeding substrate; they will oviposit at night. Hatching occurs in 12-16 h. Females retain the eggs for long periods when suitable breeding sites are not available; when laid, these eggs hatch in a short time. C. vicina and C. vomitoria will deposit living, newly hatched larvae. Maggots generally avoid light and burrow into the substrate to feed. Full-grown larvae cease feeding and migrate to find a site to pupariate; they prefer loose soil and will travel 25-30 m to find a suitable location. The pupal period is 5-9 days.

Pest status is based primarily on their presence around people and food. They are strong flyers, and will travel far from their breeding site; these flies are strongly attracted to organic odors. Adults are mostactive during warm and sunny weather. Some species freely enter buildings and become a nuisance by flying atlights and windows. Adults indoors are a threat to food sanitation. Lucilia species are common in and around houses, and adults can enter openings as small as 3.2 mm. Blow fly larvae will develop in carcasses ofdead birds or rodents in attics and wall voids. Small animals can support a large number of larvae, and within a few weeks a large number of adults will emerge indoors. There are usually several generations per year.

Calliphorids are commonly used in maggot therapy, which is the treatment of wounds with live fly larvae. Blow fly species used for this type of medical treatment have some important characteristics. Their larvae feed only on necrotic or damaged tissue and not on sound, healthy tissue; larvae usually remain at the wound site and do not crawl to search for other sites; they develop rapidly; they are relatively easy to rear in vitro, and the eggs are easily sterilized. The most widely used cal-liphorid in wound therapy is Lucilia (= Phaenicia) sericata. This species feeds only on necrotic tissue and is ideally suited for use in removing dead tissue from wounds. Other calliphorids used by physicians for treating wounds include Calliphora vicina, Chrysomya rufifacies, Lucilia cuprina, L. illustris, Phormia regina, and Protophormia terraenovae. Early physicians learned that the larvae often found in wounds improved healing and reduced scarring; this type of therapy has been used for centuries. Placing maggots in wounds has been used in societies around the world, including aboriginal tribes in Australia, Hill Peoples of Northern Burma, and probably the Mayans of Central America. The founder ofmodern maggottherapy is William Baer (1872-1931), professor of orthopedic surgery at Johns Hopkins School of Medicine, Baltimore, Maryland, USA. Following his experience with maggots in the wounds of soldiers, he designed a therapy of using sterilized maggots to treata variety ofwounds. This method is successful in treating chronic or acutely infected wounds, including bone infections, abscesses, carbuncles, and ulcers.

Congo floor-maggot, Auchmeromyia senegalensis Adults are 9-13 mm long and the mesonotum has two longitudinal black lines. The abdomen is primarily black apically; abdominal segment 2 in the male is 1.5 times longer than segment 3, and segment 2 in the female is twice as long as segment 3. Full-grown larvae are about 14 mm long and yellowish white in the first stage, but red after feeding; the body has many folds in each of the segments. The posterior segment ends in a sharp angle, and dorsally on this segmentare the posterior spiracles. Adultflies remain in shaded areas near human habitations, and they feed on discarded fruits and vegetables. Eggs are laid in batches of about 50 in dry soil in shade, in the nests of burrowing animals, or indoors in the soil floor of dwellings. Larvae remain in cracks and crevices in the soil, but move out at night to suck blood from people sleeping directly on the floor. The bite from the maggots is not painful and larvae are able to feed without being disturbed. Maggots are not capable of climbing to reach individuals sleeping in cots above the floor. Development takes about 14 days under suitable conditions, but can be extended for several months if the larvae do not find food. This species occurs in Africa.

Calliphora uralensis Adults are 10-14 mm long. They are metallic blue with pale setae forming stripes on the thorax, and with yellowish-white spots on the abdomen. Eggs are usually laid on excrement, but also on meat and fish. More than 200 eggs may be deposited at one time; hatching occurs in about 20 h. Development is completed in 10-14 days, and the pupal period lasts 7-10 days. Adults live for about 3 months, and overwintering is in the pupal stage. This is a common fly in houses and food-handling establishments in the eastern regions of Europe and Russia during summer; they are uncommon in winter.

Bluebottle fly, Calliphora vicina (= C. erythrocephala) Adults are about 10 mm long and have a bluish-black thorax, and a metallic-blue abdomen. The region of the head below the eyes is orange; in the closely related species, C. vomitoria, it is black. Eggs are laid in batches of up to 180 directly on the larval substrate; fecundity is 500-700. Hatching is in about 88 h at 10°C, 38 h at 12.5 °C, 19 h at 19 °C, and 14 h at 25 0C. First-instar larvae develop in about 49 h at 12.5 °C, 22 h at 19 °C, and 18 h at 25 °C; second-instar larvae develop in about 58 h at 12.5 °C, 23 h at 19 °C, and 19 h at 25 °C; third-instar larvae develop in about 65 h at 12.5-19 °C, and 26 h at 25 °C. There may be 5-14 days postfeeding in search of a pupari-ation site. The pupal period lasts about 41 days at 10 °C, 28 days at 12.5 °C, 14 days at 19 °C, and 11 days at 25 °C. Puparia are formed in the soil and adults can emerge from puparia buried 40 cm in sandy soil. Overwintering is by adults, larvae, and pupae. Adults live about 30 days, but 188 days has been reported. Females oviposit on fresh, decaying, or cooked meat, and on human excrement. Adults appear in early spring, but they are uncommon in summer; there may be a peak of adults in the fall. These are slow-flying and loud-buzzing flies, and they often enter houses. This species is widely distributed in North America, Mexico, Europe, and in Asia, and Australia.

Blow fly, Calliphora vomitoria Adults are about 10 mm long and they have a bluish-black thorax and a metallic-blue abdomen. The region of the head below the eyes is black, but in C. vicina itis orange. Eggs are laid on the larval food and they hatch in 6-48 h; larval development is complete in about 15 days. Third-stage larvae move to a dry substrate to pupate; the pupal period is about 11 days. Adults live about 35 days. This species is widely distributed in North America and Europe.

Oriental latrinefly,Chrysomyamegacephyla Adults are about 11 mm long, and the thorax and abdomen are greenish blue with purple reflections. The first abdominal segment is black. Full-grown larvae are about 12 mm long, smooth, and yellowish white. Larvae feed on decomposing animal matter, and excrement, including human urine. Eggs are laid in a mass on or in a suitable substrate; fecundity is about 400 eggs; hatching occurs in about 10 h. Development takes about 4 days, and the pupal period lasts about 4 days. Adults live 54-90 days at 25-29 °C and 75% relative humidity (RH). This calliphorid occurs from the Middle East to southern and eastern Asia,

Australia, and New Zealand. It is a common pest in houses and outdoor markets, where adults come to meats and various household foods.

Hairy maggot blow fly, Chrysomya rufifacies Adults are 10-12 mm long and the thorax is uniformly green or violet blue; the parafacial and facial bristles are reddish brown. Full-grown larvae are about 14 mm long and brownish yellowish; the body has a median row of fleshy tubercles on each segment, which gives the maggot a somewhat hairy appearance. Small spines are present on some of these projections, especially those dorsally. In C. albiceps, a related species, the stalks of the projections have no spines. The peritreme of the posterior spiracle is very wide and the edges of the gap are forked; the slits are short and nearly fill the spiracular plate. This cal-liphorid produces unisexual progeny, which is an unusual feature among higher flies. C. rufifacies has a temperature tolerance that corresponds with its tropical origin and general distribution. Eggs are laid singly or in batches; fecundity is 210-368 eggs. Eggs fail to hatch at9 °C,butati5 °Ceggshatchand larvae develop but fail to pupate. Larvae grow and develop normally at40 °C. Adults live 23-30 days. The temperature threshold for adult flight activity is i3 °C; this species has a flight range of 0.5-1.6 km/day (Australia). C. rufifacies is nota primary invader oftissue, and feeds as a scavenger or secondary carrion fly. It is native to the tropics but has spread around the world with commerce. It is known from India, Japan, Hawaii, Australia, Central and South America (Argentina), and North America (Arizona, Texas, Florida). It is common in some urban areas in India, where it occasionally enters houses; it also enters houses in Australia, but less so in Japan. In some regions, it causes primary and secondary myiasis in animals. Ithas a role in forensic entomology. Ithas been recovered from human cadavers in Costa Rica, California, Arizona, Texas, and Florida. This species is closely related to C. albiceps, which is distributed in Africa, and they may be conspecific.

Tumbu fly, Cordylobia anthropophaga Adults are 9-12 mm long and dull yellowish white to light brown, and with two dark gray dorsal longitudinal thoracic stripes. Wings are light brown. The four visible abdominal segments are about equal-sized. Full-grown larvae are ii-i5 mm long, yellowish white, and covered with dark spines. Eggs are laid in batches on dry soil in shady sites, especially sites contaminated with urine or feces of humans and other animals (rodent, dog, monkey); oviposition sites are usually indoors, in the earthen floor ofhuts and other dwellings. Eggs are also laid on drying laundry hung out of direct sunlight; infestations develop if these eggs are not killed. Females lay eggs in batches of 200-300, and fecundity is about 500 eggs. Hatching occurs in 1-3 days, and first-stage larvae can remain alive for 9-15 days without food. Larvae usually remain concealed just below the soil surface during the day. Movement or body heat on the soil activates the larvae. They move at night and attack people or small animals sleeping on the earthen floor. They use their mouthhooks to attach directly to a host, or to clothing first then move to the host; larvae usually penetrate the skin to the depth of the posterior spiracles. Developmentis completed in 8-9 days, and the full-grown larva leaves the boil-like swelling on the skin and drops to the soil to form a puparium. Adult flies emerge in 8-15 days. Rats may be the normal host and probably serve as the natural reservoir for this fly. Dogs are also infested, and severe infestations cause death of the animal. This species occurs in Africa. The related species, Cordylobia rohaini (Lund's fly), closely resembles the Tumbu fly in appearance and life history, but attacks humans less frequently. It occurs in tropical Africa, especially in areas ofrain forest.

Cynomyopis cadaverina Adults are 9-14 mm long, the thorax is metallic bluish black, and the abdomen is shiny bluish green. The seasonal peaks in populations of this fly are in early spring and late fall, but adults occasionally appear during warm spells in winter. Adults enter houses readily. Larvae breed in carrion, excrement, and other decaying organic material. Eggs are deposited in batches of 25-50, and hatching is in about 24 h. Developmentis complete in about 6 days, and the pupal period lasts about 4 days. Overwintering is in the pupal or adult stage. Itis found throughoutNorth American, Mexico, and northern Europe.

Lucilia illustris Adults are 5-10 mm long, and the thorax is bluish green with bronze and purple reflections. Full-grown larvae are 11-13 mm long and yellowish white; the body is somewhat peg-shaped and with dark bands of spines encircling segments 2-9, but the bands may be incomplete dorsally. Prothoracic spiracles have 6-8 lobes; the posterior spiracles have a complete peritreme. It generally occurs outdoors, but will come indoors under adverse weather conditions. Females oviposit on dead animals, and feces (animals and human). Larvae feed on these substrates, and on the wool and flesh of sheep. This species is often found with L. sericata on carcasses of animals. Larvae have been used in wound therapy. This species occurs in North America and Europe.

Greenbottlefly, Luciliasericata (= Phoenicia) Adults are about 12 mm long and metallic green, with yellowish or coppery reflections. Full-grown larvae are about 14 mm long and may be colored slightly purple. Eggs are often deposited in large numbers in one location; fecundity is about 250 eggs. Hatching occurs in about 24 h; it is about 23 h at 22 °C and 18 h at 29 °C. Larval development is completed in 4-5 days at 24 °C. First instar develops in about 27 h at 22 °C and 16 h at 29 °C; second instar develops in about 22 h at 22 °C and i6hat29 °C; the third instar takes about22 hat22-29 °C. There may be 4.5-9 days postfeeding in search of a pupari-ation site. The pupal period is about 143 h at 22 °C and 130 h at 29 °C. The common breeding medium is carrion, but also includes decaying garbage and manure. Larvae generally confine their feeding to dead tissue; this species is commonly used in modern wound therapy. This species is widely distributed, and probably cosmopolitan.

Phaenicia cuprina Adults are 6-8 mm long and the thorax is metallic green. Larvae are primarily scavengers and they are usually associated with wet garbage and decaying organic matter. Eggs hatch in about 24 h, and larvae feed for about 6 days before moving away and entering the ground to form the puparium. The pupal period lasts about 7 days. Adults often come indoors and they are strongly attracted to lights; they will fly to ultraviolet lights. This species is widely distributed.

Black blow fly, Phormia regina Adults are 6-11 mm long and the thorax is black with a bluish-green luster, and it has black, longitudinal stripes. The abdomen is shiny, and bluish green. Full-grown larvae are i2-i7 mm long and yellowish white. Bands of small spines encircle segments 1-8, and segments 9 and 10 have a band ofventral spines. Prothoracic spiracles have i0 or ii short lobes; the posterior spiracles are in a depression surrounded by six tubercles. The posterior spiracle peritreme appears incomplete. Eggs are often deposited in large numbers in one location; fecundity is about 250 eggs. Hatching occurs in about 24 h; it is about 20 h at 22 °C and 18 h at 29 °C. Larval development is completed in 4-5 days at 24 °C. First instar develops in about 25 hat 22 °Cand i2hat29 °C; second instar develops in about 25 h at 22 °C and 15 h at 29 °C; the third instar takes about 25 h at 22-29 °C. There may be 5.29 days postfeeding in search of a pupariation site. The pupal period is about 116 hat 22 ° Cand 99hat29 ° C. Eversion of the pupal respiratory horns occurs by 26.5 hat 22 °C and by 22 hat 29 °C. Third-stage larvae often pupariate on the surface of the breeding substrate, unless it is very wet or exposed to bright light. Adults often overwinter in protected locations. P. regina is common in early spring when temperatures are cool and is less abundant in the summer. Larvae feed on carrion and decaying plant material, and on the wool and flesh of sheep. Larvae have been used in wound therapy. This species is widely distributed in North America, Mexico, and Europe.

Cluster fly, attic fly, Pollenia rudis Adults are 4-8 mm long. They have a broad thorax covered with golden-yellowish setae; the wings overlap when at rest. This calliphorid lacks the shiny blue or green thorax, which characterizes most of the flies in this family. Large numbers of adults enter buildings in the fall, and remain there, relatively inactive, throughout the winter. In early spring adults leave overwintering harborages during warm and sunny days; mating occurs in spring. Eggs are laid singly in the soil, and hatching occurs in about 3 days. Larval stages are predaceous on earthworms. First-stage larvae seek out earthworms (Allolobophora spp.); larvae enter through the spermiducal opening and other pores. There is usually one fly larva per earthworm. Development is completed in 27-39 days, and the puparium is formed in the soil. There may be four generations per year in the USA. In late summer and fall, large numbers of adults gather on the sun-warmed sides of buildings, and then move through cracks and crevices to enter attic space and wall voids. Natural hibernation sites for this fly probably include animal burrows, under bark, and in cavities in trees and down logs. Earthworm populations, which may be as many as 420-500 per square meter in urban and suburban soils, ensure the larval stage of this fly an abun-dantfood resource. Urban and suburban buildings provide the adult cluster fly suitable overwintering harborages throughout the distribution of the species. It occurs throughoutNorth America, and from Ireland to Europe and North Africa, to Siberia and China.

Other Pollenia Adults of P. atramentaria, P. dasypoda, P. intermedia, P. varia, and P. vespillo are found at animal feces, but are not known to overwinter in large numbers. The Japanese cluster fly, P.japonica, is widely distributed in Japan. Adults of this species lay eggs in soil and larvae feed on earthworms.

Protophormia terraenovae Adults are 8-12 mm long and the thorax is dark blue, the legs are black, and the abdomen is greenish blue. The front in the male is less than one-fifth the head width. Full-grown larvae are 10-16 mm long and yellowish white. This species has a northern distribution; it is usually confined to areas north of the Tropic of Cancer. It breeds in carrion and decaying organic matter, and it has been used in wound therapy. It is distributed in North America and Europe.

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