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DIPTERA, Sarcophagidae (Brauer 1889) -- <Images> & <Juveniles> Please refer also to the following links for details on this
group: Sarcophagidae = Link 1 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
are 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 B. lineata, S. lellyi and S. caridei,
the maggots enter the host body through the thin membrane at the base of the
wing. B. 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 B. 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 B. 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 B. lineata, a maximum
of 11 in B. filipjevi and 9 in S. 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 B. 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. Life
Cycle
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 B.
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). References: Please refer to <biology.ref.htm>, [Additional references
may be found at: MELVYL
Library] Aldrich, J.
M. 1916.
Vol. 1, Thomas Say Found., Ent. Soc. Amer. 302 p. Allen, H.
W. 1926. Proc.
U. S. Natl. Mus. 68: 1-106. Clausen, C. P.
1940/1962. Entomophagous
Insects. McGraw-Hill Book Co., Inc.,
NY. & London. 688 p. [reprinted 1962 by Hafner Publ. Co.]. Cole, F.
R. 1969.
The Flies of Western North America.
Univ. Calif. Press, Berkeley & Los Angeles. 693 p. James, M. T. & R. F. Harwood. 1969.
Herm's Medical Entomology, 6th ed.
MacMillan Co. 484 p. Kunckel D'Herculais, J. 1894. Les Dipteres parasites des Acridiens: les
Muscidae vivipares a larves Sarcophages.
Aptenie et castration parasitaire. Acad. Sci. Compt. Rend., Paris 118: 1106- Potgieter, J.
T. 1929. A
contribution to the biology of the brown swarm locust Locustana pardalina
(Wlk.) and its natural enemies. Proc.
Agr., Union So. Afr., Dept. Agr. & For., Pan-Afr. Agr. Vet. Conf.,
Pretoria. p. 265-308. Roback, S. S.
1954. Illinois Biol. Mon.
23: 1-181. Wood, O. H.
1933. Notes on some dipterous
parasites of Schistocerca
and Locusta in the
Sudan. Bull. Ent. Res. 24: 521-30. |