An Introduction to Medical Entomology
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Sarcophagidae. -- <Habits>; <Adults> & <Juveniles> -- The flesh flies are similar to Calliphoridae, but they are usually black or gray with stripes on their thorax. Adults feed on sweet foods such as flower nectar, fruit juice and honeydew.
Their larvae show diverse habits, but most feed on animal material, with many being scavengers. Some species are scavengers, some are parasites of other insects and a few are parasites of vertebrates that develop in skin wounds.
Description & Statistics
Early reviews of host preferences were by Aldrich (1915) and Greene (1925a). Entomophagous species are in subfamilies Sarcophaginae and Melanophorinae. Many species of Sarcophagidae are limited to carrion, others to manure; but there are both predaceous and parasitic species. Predaceous species attack egg pods of Acrididae. Well-known genera having this habit are Sarcophaga and Blaesoxipha. Oophagomyia and Wohlfahrtia are predaceous in the egg capsules of the same host group, and an occasional species of Blaesoxipha is both parasitic on the active stages and predaceous on the eggs. Mantidophaga is an internal parasitoid of the late nymphs and adults of Mantidae.
Clausen (1940) noted that an unusual degree of plasticity was revealed in the behavior of species of this family, and many were apparently only in the transitional stages to obligate parasitism. Most parasitic species are primary, solitary, endoparasitoids, though gregarious species are known. The principal host range includes Orthoptera (Acrididae, Mantispidae) and Lepidoptera, but other insect orders may be attacked. A few species are parasites of snails, while others are carrion feeders or vertebrate parasites.
A host group frequently attacked by species of the Sarcophaginae is the social wasps and bees. The relationship ins some cases is strictly parasitic, while in others it is commensal. Myiapis and Senotainia are internal parasitoids of worker honeybees, and Sphixapate develops within the larvae. Metopia and Brachicoma are external parasitoids or predators of the brood of wild bees, the latter genus attacking mainly bumblebees. Hilarella and Miltogramma develop on various insects, which are stored in cells of hunting wasps, or on the material with which the cell of bees are provisioned. Several genera have widely different host preferences. Lepidopterous larvae and pupae frequently yield sarcophagid flies, and it has been thought that there were parasitic. Several species of Sarcophaga associated with the gypsy moth were found to be scavengers only (Patterson 1911). Young larvae were unable to enter healthy larvae or pupae, and if artificially introduced into the bodies of living individuals they died. However, species of Agria are predaceous on pupae of Lepidoptera. Eleodiomyia has been reared from adult beetles of the family Tenebrionidae; Scarabaeophaga from pupae and adults of Cotinus nitida L.; and Sarcophaga spp. from adult Pentatomidae, Blattidae, etc (Clausen 1940/62). Arachnidomyia sp. has been reared from egg sacs of spiders and various genera and species from snails.
There is a wide range in host preference found among the parasitic and predaceous species of Sarcophaga. Not much is known regarding insect hosts of Melanophorinae, although species have been reared occasionally from spider egg masses and from coleopterous larvae and adults. Melanophora, Cirillia, and closely related forms are parasitic in Isopoda (Porcellio, Metaponorthus, Oniscus), and some species of this subfamily have been reared from snails.
Biology & Behavior
In Brachycoma lineata, S. lellyi and S. caridei, the maggots enter the host body through the thin membrane at the base of the wing. S. filipjevi enters through the membranes of the abdomen or through the genital opening. The latter behavior is similar to that of Eleodiomyia in attacking tenebrionid beetles. Wood (1933) noticed that the maggots of S. destructor readily enter freshly molted hosts but are not able to if the integument is fully hardened. The host dies within a short time after the larvae have entered the body. Mature larvae of S. linerata and S. caridei emerge from the host while the latter is still alive, and some parasitized individuals may recover. However, the hosts of Wohlfahrtia are usually dead before the larvae finish feeding. They usually emerge through the thin membranes of the neck, although some individuals of S. kellyi are believed to emerge through the anal opening.
Wood (1933) found that 78 % of attacked hosts of Brachycoma lineata recovered, but only 38% were able to reproduce thereafter. Relatively little growth occurs in S. destructor as long as the host remains alive. The young larvae of this species attack the wing muscles, nd death results primarily through infection. After this, development of the parasitoid is rapid. Only 16% of hosts containing one parasitoid larva died, while 92% died when two or more were present. If hosts are immature at the time of attack, they do not attain the adult stage. Larval feeding is confined mostly to the fat body. The number of individuals developing in each host varies, being usually only 2 in the case of Brachycoma lineata, a maximum of 11 in B. filipjevi and 9 in B. caridei.
There is often a high percent parasitization by Sarcophagidae, but opinions vary as to their value in natural control. Smith (1915) stated that swarms of Dissosteira longipennis Thoms. in New Mexico were almost eliminated by S. kellyi. Kunckel d'Herculais (1894) found parasitization of Schistocerca by sarcophagids in Algeria to be 69% in 1889 and 75% in 1890. The flies followed host swarms, harassing them continuously. In the case of Wohlfahrtia euvittata in South Africa, 50-90% of Locustana were found parasitized, and in some areas this attack was responsible for discontinuing poisoning programs.
Some species that were discussed as internal parasitoids of nymphs and adults of locusts are also predaceous in egg masses of the same hosts. This range in habit has been found for Sarcophaga opifera Coq. in British Columbia, and Treherne & Buckell (Clausen 1940/62) thought that the larvae, after leaving the body of the adult host, continued their development on the eggs in soil. Potgieter (1929) in South Africa observed that W. euvittata is very important in natural control of Locusta pardalina Wlk., when parasitic on the active stages. About 50% of the egg masses in one area were destroyed by this fly. The maggots are laid in groups in the openings of partly hatched egg pods or in the froth at the upper ends of those freshly laid or exposed. Larvae in various stages of development were found on the surface of the ground, and these were migrating to other egg pods for further feeding (Potgieter 1929).
Sarcophagids that are parasitic or predaceous on the brood of bees and wasps are mostly in genera Metopia, Brachicoma and Hilarella. Bougy (1935) described the attack of H. stictica Meig on Ammophila hirsuta Scop. in France. The host stores its nest with noctuid larvae, and the female fly appears while the prey is being transported to the nest. She does not attempt to larviposit on it at this time. It is only after the caterpillar has been placed in the cell and the Ammophila egg laid that she evades the host, enters the burrow and lays her own minute larva alongside the host egg. This egg is consumed within 24 hrs., and the larva then enters the body of the caterpillar to complete its development. Each individual may be regarded as a predator on the egg of Ammophila and an internal parasitoid of noctuid caterpillars.
Mature larvae and young pupae of bumblebees are parasitized by Brachycoma sarcophagina Tns. in North America. The live young are laid on or in the brood cells. They enter the body and feed until larval maturity. Pupation occurs in the nest material at the bottom of the comb. B. davidsoni Coq. is thought to lay eggs directly on the larvae; and after one is consumed, the parasitoid larva enters other cells to attack their occupants. Metopia leucocephala Rossi has been found in cells of Philanthus. Females enter the host burrow for a short distance and there lay their larvae, which have to find their own way to the cells, sometimes several feet away. Adult honeybees are found heavily parasitized by Senotainia tricuspis Meig. in some parts of Russia. The larvae feed principally in the thoracic region, the same habit being recorded for Myiapis angellozi Seguy (Seguy 1930).
Agria mamillata Pand. is predaceous on pupae of Hyponomenta in Italy, with flies appearing in the field in early June to lay their partially incubated eggs on caterpillars when they are mature but before cocoon formation. The young larva enters the body of the pupa and quickly consumes its contents. It then penetrates the adjoining cocoons and continues its feeding, destroying 50 or more pupae per single larva before maturity (Servadei 1931).
S. latisterna Perk was reared from various pupae of Lepidoptera, where it was believed to be a true facultative parasite (Hallock 1929).
Thompson (1920a, 1934) studied several sarcophagids that are parasitic in isopods of genera Oniscus, Porcellio and Metaponorthus. They differ in several ways from the general habits of the family. The adaptive characters of the 1st instar larvae, as well as the habits of the immature stages, show a closer biological affinity with Tachinidae than by any other members of Sarcophagidae. Parafeburia maculata Fall, is a solitary internal parasitoid of the first two genera. Its unincubated eggs are probably laid in the general vicinity of the hosts or where they are in the habit of congregating. They hatch in ca. one week when these membranous eggs give rise to planidium type larvae. This is the only instance in the Muscoidea in which this larval form hatches from membranous eggs that are unincubated at the time of laying. Young larvae enter the host body through the soft cuticle separating the ventral sclerites or at the bases of the appendages. Once inside the host, the larva is found with its posterior end fixed in a perforation in the integument, and a respiratory funnel is formed. The 2nd instar larva has a very thin integument, and tests have shown that an exchange of gases takes place through it; The greater part of the oxygen requirements of the larva may be secured in this way, and pupation occurs within the remains of the host.
Clausen (1940) commented on the definite effect on the reproductive system and the secondary sexual characters of the host as a result of parasitism by Parafeburia. Female ovaries are atrophied, owing to absorption of fat by the parasitoid, and such females do not develop a brood pouch. Less complete information regarding Cirillia angustifrons Rond. was presented by Thompson (1920a). General habits are similar to those for Parafeburia, with the outstanding distinction in the host relationships being the formation of the integumentary respiratory funnel by the larvae. This habit is unknown elsewhere in Sarcophagidae, although it is common in Tachinidae, indicating a higher development of the parasitic relationship than has been attained by other species.
The life cycle of parasitic Sarcophagidae, from larviposition to adult emergence, is relatively short, being completed in 16-30 days, of which the larval feeding period takes only 5-10 days. In Wohlfahrtia, larval maturity is followed by a resting period of 6-12 days, and the pupal stage then requires 16-30 days. Several generations are usually produced each year, and 5-6 are recorded for S. kellyi. Hibernation is known for Brachycoma lineata and S. kellyi, in both of which mature larvae rather than pupae persist through winter. The life cycle of P. maculata in Oniscus and Porcellio differs from the general habit of the family by having only a single generation each year. Adults appear in midsummer, and winter is passed as 2nd instar larvae within the live host (Clausen 1940/62).
For detailed descriptions of immature stages of Sarcophagidae, please see Clausen (1940/62).
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