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HYMENOPTERA, Spalangiinae (Pteromalidae) (Chalcidoidea) -- <Images> & <Juveniles>

This subfamily was originally a separate family Spalangiidae. Pteromalidae now also includes the former separate families, Cleonymidae, Miscogasteridae and Spalangiidae, which have been designated subfamilies Cleoneminae, Miscogasterinae and Spalangiinae, respectively. For the present, discussions of these various subfamilies will be separate because of considerable distinctness among them.

Legner (unpub. data) working with parasitoids of synanthropic Diptera has observed what he regards as a higher degree of sophistication among species of Spalangia than of various other pteromalid parasitoids, as manifested by more intense searching of the host's environment and examination of the host prior to parasitization. Also, during numerous laboratory experiments with various species of this genus, they have shown a degree of learning toward the ultimate goal of escape from the experimental environment. After repeated handling their ability to feign death, for example, is remarkable, which frequently results in their escape from the observer.

There are only a few genera, of which Spalangia is the most commonly encountered. Richardson (1913) listed six species from dipterous hosts, one from Lepidoptera and two with myrmecophilous habits. The recording from the lepidopterous host is probably in error, for the species concerned, S. nigra Latr, has been frequently recorded from housefly puparia (Bou…ek 1963). Silvestri (1914) recorded several species from Trypetidae in West Africa. Those which attack Diptera utilize puparia and are solitary and external in habit. Clausen (1940) noted that these are parasitoids of dipterous pupae and consequently are considered hyperparasitoids only when the particular host species that they attack are themselves of parasitic habit, although S drosophilae Ashm., attacking dung-infesting Diptera, is recorded as attacking Alysia and Psilodora, which are primary parasitoids of the same hosts. Entomophagous Diptera which form their puparia on or near the soil surface are frequently attacked by Spalangiidae. Cerocephala develops on larvae and pupae of the coleopterous families Scolytidae, Curculionidae, etc. (Clausen 1940/1962).

Spalangiidae are valuable in the natural control of synanthropic Diptera both naturally and through artificial field inundation (please refer to section on medical/veterinary entomology <bc-37.htm>). Lindquist (1936) recorded up to 64% parasitization of dung-infesting Diptera, mainly by Spalangia muscidarum var. stomoxysiae Gir.

Biology and Behavior

Richardson (1913) did an extensive study on S. nigroaenea Curtis (= S. muscidarum) as a solitary parasitoid of the pupae of housefly, Musca domestica L. In oviposition, the female crawls over the host puparium, examines it carefully with the antennae, and then inserts the ovipositor through the puparial wall, usually in the posterior half. The body of the larva or pupa within the puparium is not penetrated, and the egg is placed externally.

The 1st instar larva is very active and capable of extended movement over the surface of the host, which is for the purpose of finding a suitable feeding point, which usually proves to be the dorsum or dorsolateral areas of the abdomen (Gerling & Legner 1968). The pupal integument is much thinner at these points and is more easily punctured than elsewhere. The 2nd and 3rd instar larvae are relatively more fixed in their feeding positions in S. cameroni Perkins (Gerling & Legner 1968). When feeding is complete, the mature larva moves toward the anterior end of the puparium, casts its meconium and then pupates. The adult emerges through a hole cut in the anterior end of the puparium (Clausen 1940/1962).

An account of S. nigroaenea as a parasitoid of stablefly, Stomoxys calcitrans L. was given by Pinkus (1913), which differs in several respects from that by Richardson. Oviposition took place in the anterior portion of the puparium, usually through a suture. The female was able to detect prior parasitization and would not deposit a second egg on a host already bearing one. Many of the pupae attacked died, even though no egg was deposited, which would indicate that the body had been penetrated by the ovipositor. The adults mate soon after emergence, and females are able to deposit eggs the same day..

A minimum life cycle under cool laboratory conditions was found to be ca. 88 days, but under warmer summer conditions outdoors the cycle is less than half this duration. Hibernation was thought to take place in any immature stage, and development progressed at any time that the temperature became higher again.

Spalangia parasitizing the pupae of Lyperosia were studied by Handschen (1932, 1934). These were S. sundaica Graham of Java and S. orientalis Graham from Australia. S. sundaica deposited an average of 160-170 eggs during a 4-weeks period. The cycle from egg to adult was completed in 18-21 days, and males emerged two days earlier than females. Adults were attracted to dung in which the hosts develop. S. orientalis had the same general habits and life cycle but produced an average of only 85 eggs during 15 days. These two "species" were hybridized in an effort to produce a more effective "race" that was better adapted to Australian conditions. Female S. orientalis when mated with male S. sundaica produced progeny more prolific than either parent form, the average egg deposition being 240 in 32 days. The reverse cross produced 100 eggs during 10 days. The fact that the hybrids were fertile and highly fecund, it is indicative that races instead of species had been studied. Yet Bou…ek (1963) considered S. orientalis a synonym of S. endius Walk. and S. sundaica a synonym of S. nigroaenea Curtis. During three decades of research on Spalangia and related genera, Legner (unpub. data) has never obtained hybrids between S. cameroni and S. nigroaenea. Studies with Australian Spalangia nevertheless have raised some interesting questions, as for example the reproductive isolation of a race of S. endius secured from the southeastern portion of the continent from that of morphologically indistinguishable isolates from North America. A morphologically and biologically distinct race from New Zealand, however, although isolated from the Australian isolate fully interbred with North American S. endius (Legner 1983).

In Cerocephala cornigera Westw., parasitoid of scolytid larvae and pupae, the female first paralyzes the host and then places an egg either directly on it or in its immediate vicinity. The ovipositor is usually inserted into the entrance of the host's oviposition tunnel.

Adult feeding habits in the family were studied by Lindquist (1936) who found that Spalangia muscidarum var. stomoxysiae paid little attention to artificial foods, and the length of life when confined with puparia suggested host feeding. Parker & Thompson (1925) found such feeding in S. nigra Latr, where they noted the construction of a feeding tube (see Host‑feeding>.

Life cycles in Spalangiidae are short, ranging from 17 days in S. muscidarum var. stomoxysiae and S. drosophilae to 25-30 days in C. cornigera. More than one generation is produced each year, and winter is passed in the mature larval stage, although Richardson found that S. muscidarum is in the pupal stage during winter.

The developmental life history of Spalangia cameroni Perkins was presented by Gerling & Legner (1968), with observations on physiology of ovum formation, and sperm translocation through the male reproductive system. Pertinent aspects treated in detail were host-feeding and selection, oviposition, superparasitization, length of developmental stages, oocyte development, ovisorption and sperm activation. Particularly interesting were the interinvolvement of host-feeding and oviposition, the high moisture requirement for embryonic development and the 100+ feeding punctures made by a larva while feeding ectophagously on the host pupa encased in the puparium. There was a prolongation of female pupal development with respect to the male, a deposition of partially resorbed eggs, two chambers in the seminal vesicle, and a short duration of testes function.

The sex ratio varies with environmental conditions (Legner 1967a,b; 1969, 1979a,b; Legner & Gerling 1967), but females tend to predominate in a ratio of 2:1 in S. muscidarum var. stomoxysiae. There has been no thelytoky discovered in this family.

For detailed descriptions of immature stages please see (Clausen 1940/1962).

References: Please refer to <biology.ref.htm>, [Additional references may be found at: MELVYL Library]

Ables, J. R. & M. Shepard. 1974b. Responses and competition of the parasitoids Spalangia endius and Muscidifurax raptor (Hymenoptera: Pteromalidae) at different densities of house fly pupae. Canad. Ent. 106: 825-30.

Ables, J. R. & M. Shepard. 1976a. Seasonal abundance and activity of indigenous hymenopterous parasitoids attacking the house fly (Diptera: Muscidae). Canad. Ent. 108: 841-44.

Ables, J. R. & M. Shepard. 1976b. Influence of temperature on oviposition by the parasitoids Spalangia endius and Muscidifurax raptor. Environ. Ent. 5: 511-13.

Ables, J. R., R. P. West & M. Shepard. 1975. Response of the house fly and its parasitoids to Dimilin (Th-6040). J. Econ. Ent. 68: 622-24.

Ables, J. R., M. Shepard and J. R. Holman. 1976. Development of the parasitoids Spalangia endius and Muscidifurax raptor in relation to constant and variable temperature: simulation and validation. Environ. Ent. 5: 329-32.

Abraham, R. & H. König. Der Einfluss der Temperatur auf die Anstichaktivität bei Nasonia vitripennis und Spalangia nigra (Chalcidoidea: Pteromalidae). Entomophaga 22: 299-308.

Azizov, N. 1972. Some details on the biology of Spalangia nigroaenea (Hymenoptera, Spalangiidae). Zool. Zurn. 51(6): 925-26. [in Russian].

Bathon, H., & K. Fabritius. 1985. Der Fliegenpuppenparasit Spalangia endius Walker (Hymenoptera, Pteromalidae), ein Neufund für die Bundesrepublik Deutschland. Z. angew. Zool. 72(3): 343-47.

Bloomcamp, C. L. 1985. Development of resistance to cyromazine in the house fly and its affect on the hymenopteran pupal parasite, Spalangia endius. M.S. Thesis, Univ. Florida, Gainesville. 75 p.

Blume, R. R. 1987. Bionomics of some species of the genus Spalangia Latreille. Southwest. Ent. 12: 191-95.

Blume, R. R., S. E. Kunz, B. F. Hogan & J. J. Matter. 1970. Biological and ecological investigations of horn flies in central Texas: influence of other insects in cattle manure. J. Econ. Ent. 63: 1121-23.

Boucek, Z. 1963. A taxonomic study in Spalangia Latr. (Hym., Chalcidoidea). Acta Ent. Musei Nation. Pragae, Praha 35: 429-512.

Boucek, Z. 1965. Descriptions of new species of Spalangia Latr. (Hym., Chalcidoidea) from Africa and South America with notes on some known species. Acta Ent. Musei Nation. Pragae, Praha 36: 593-602.

Boucek, Z. 1965. A review of the chalcidoid fauna of the Moldavian S.S.R., with descriptions of new species (Hymenoptera: Chalcidoidea). Acta faun. ent. Musei Nation. Pragae, Praha 11: 5-38.

Boucek, Z. 1965. Contributions to the Czechoslovak fauna of Chalcidoidea (Hym.). Acta faun. ent. Musei Nation. Pragae, Praha 12: 231-60.

Boucek, Z. 1970. Contribution to the knowledge of Italian Chalcidoidea, based mainly on a study at the Institute of Entomology in Turin, with descriptions fo some new European species (Hymenoptera). Mem. Soc. Ent. Ital. 59: 35-102.

Boucek, Z. 1977. A faunistic review of the Yugoslavian Chalcidoidea (parasitic Hymenoptera). Acta ent. Jugoslavica, Zagreb 13: 1-145.

Burks, B. D. 1969. Species of 'Spalangia' Latreille in the United States National Museum collection. Smithson. Contrib. Zool., Washington 2: 1-7.

Cabrales, G., R. Figueroa, F. Uribe & C. I. Trochez. 1985. Evaluación del efecto del parasitismo por Spalangia endius W. (Hym. Pteromalidae) sobre la dinámica de población de Musca domestica L. en galpones para aves (La Florida-Risaralda). Acta Agronómica, Palmira 35: 93-105.

Capehart, J. S., R. L. Harris & D. E. Bay. 1981. The effect of host species on developmental time of Muscidifurax raptor and Spalangia drosophilae. Southwest. Ent. 6: 136-9.

Chu, F. Y. 1984. Discovery of Spalangia sp.-- a pupal parasite of Musca domestica vicina Macquart (M. domestica domestica). Natural Enemies of Insects, Kunchong Tiandi 6: 149 p. [in Chinese].

Donaldson, J. S. & G. H. Walter. 1984. Sex ratios of Spalangia endius (Hymenoptera; Pteromalidae), in relation to current theory. Ecol. Ent., London 9: 395-402.

Fullaway, D. T. 1915. Description of a new species of Spalangia. Proc. Hawaii. Ent. Soc. 3: 292-94.

<SPALANGI.G>

Gerling, D. & E. F. Legner. 1968. Developmental history and reproduction of Spalangia cameroni, parasite of synanthropic flies. Ann. Ent. Soc. Amer. 61: 1436-43.

Gersdorf, E. 1964. Neues über Spalangia cameroni Grav. (Pter., Chalc., Hym.). Ber. Naturhist. Ges. Hannover 108: 47-8.

Girault, A. A. 1921. New serphoid, cynipoid, and chalcidoid Hymenoptera. Proc. U. S. Natl. Mus., Washington, D.C. 58: 177-216. [Spalangia p. 213].

Graham, L. F. 1932. Descriptions of two nes species of the genus Spalangia Latreille (Pteromalidae) from northern Australia and the Sunda Islands. In: E. A. Handschin, A preliminary report on investigations on teh buffalo fly (Lyperosia exigua De Meij.) and its parasites in Javan and northern Australia. Comm. Australia Counc. Sci. Res. Pamph. 31: 24 p.

Handschin, E. 1934b. Studien an Lyperosia exigua Meijere und ihren Parasiten. III. Teil. Die Anziehung von Spalangia zu ihrem Wirte. Rev. Suisse Zool., Genéve 41: 267-97.

Hoelscher, C. E. & R. L. Combs, Jr. 1969. Laboratory rearing of Spalangia nigroaenea (Hymenoptera; Pteromalidae). Florida Ent. 52: 103-5.

Ienistea, M. A. & K. Fabritius. 1978. Emplenota albopila Muls. et Rey und Spalangia endius Walk., Parasiten von Eucellia tergina Zett. an der Schwarzmeerküste. Stud. Com. Muz. Brukenthal, St. Nat., Sibiu 22: 339-41.

Kochetova, N. I. & N. Azizov. 1972. A contribution to the biology of Spalangia nigroaenea (Hymenoptera, Spalangiidae), a parasite of synanthropic flies in the Samarkand area of the Uzbek SSR. Zool. Zhurnal., Moskva 51: 665-70. [in Russian].

Koval…uk, L. A. 1967. Biological peculiarities of Spalangia drosophilae Ashm., a parasite of the frit fly and a primary assessment in the north-west of the USSR, p. 42-7. In: Mater. nau…noj konf. molodyh u…enyh, Leningrad. [in Russian].

Koval…uk, L. A. 1970. Communication on research of Spalangia drosophilae, a parasite of frit flies in the Lettish SSR. Mater. 7. Pribalt. Soveshch. Zashch. Rast., Pt. 1: 40-41. [in Russian].

Koval…uk, L. A. 1971. Methodical recommendations for the rearing and use of Spalangia drosophilae Ashm., a parasite of frit fly and other flies of the genus Oscinella. Rept. Leningrad. p. 1-19. [in Russian].

Koval…uk, L. A. 1972. Spalangia drosophilae, a parasite of frit flies. Zashch. Rast., Moskva (1972): 23-5. [in Russian].

Legner, E. F. 1967a. Two exotic strains of Spalangia drosophilae merit consideration in biological control of Hippelates collusor (Diptera: Chloropidae). Ann. Entomol. Soc. Amer. 60(2): 458-462.

Legner, E. F. 1967b. Behavior changes the reproduction of Spalangia cameroni, S. endius, Muscidifurax raptor and Nasonia vitripennis [Hymenoptera: Pteromalidae] at increasing fly host densities. Ann. Ent. Soc. Amer. 60: 819-26.

Legner, E. F. 1969. Distribution pattern of hosts and parasitization by Spalangia drosophilae (Hymenoptera: Pteromalidae). Canad. Ent. 101: 551-57.

Legner, E. F. 1979a. Reproduction of Spalangia endius, Muscidifurax raptor and M. zaraptor on fresh vs. refrigerated fly hosts. Ann. Entomol. Soc. Amer. 72(1): 155-157.

Legner, E. F. 1979b. The relationship between host destruction and parasite reproductive potential in Muscidifurax raptor, M. zaraptor, and Spalangia endius [Chalcidoidea: Pteromalidae]. Entomophaga 24(2): 145-152.

Legner, E. F. 1983. Broadened view of Muscidifurax parasites associated with endophilous synanthropic flies and sibling species in the Spalangia endius complex. Proc. Calif. Mosq. & Vect. Contr. Assoc., Inc. 51: 47-48.

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Madeira, N. G. 1986. Pupation behaviour of Calliphoridae (Diptera) in nature and the discovery of the parasite Spalangia endius (Hymenoptera: Pteromalidae). Rev. Brasil. Biol. 45: 481-84. [in Portuguese].

Mandeville, J. D. 1988. Host preference, fecundity, and longevity of Muscidifurax zaraptor Kogan and Legner (Hymenoptera: Pteromalidae) influenced by Musca domestica L. and Fannia canicularis (L.) (Diptera: Muscidae), with notes on parasitism by Spalangia spp. (Hymenoptera: Pteromalidae) in the field. Ph.D. Thesis, Univ. of California, Riverside.

Mandeville, J. D. & B. A. Mullens. 1990b. Host species and size as factors in parasitism by Muscidifurax spp. and Spalangia spp. (Hymenoptera: Pteromalidae) in the field. Ann. Ent. Soc. Amer. 83: 1074-83.

Marshakov, V. G. 1983. The introduced parasite Spalangia drosophilae Ashm. (Hymenoptera, Pteromalidae) and its effect in the control of the fritflies. critical analysis. Ent. Obozrenie, Leningrad 52: 245-51. [in Russian].

McCoy, C. W. 1963. Mass liberation of laboratory reared parasites, Spalangia muscidarum (Richardson) for control of Stomoxys calcitrans (L.) and Musca domestica (L.) in Lancaster County, Nebraska. M.S. Thesis, Univ. of Nebraska, Lincoln.

McCoy, C. W. 1965. Biological control studies of Musca domestica and Fannia sp. on southern California poultry ranches. Proc. Calif. Mosq. Contr. Assoc., Inc. 33: 40-2.

Merritt, R. W., E. F. Gersabeck & M. K. Kennedy. 1981. The contribution of Spalangia endius and Muscidifurax raptor to a stable fly management program on Mackinac Island, Michigan: a question of effort, p. 44-51. In: Patterson, R. S., P. G. Koehler, P. B. Morgan & R. L. Harris (eds.), Status of Biological Control of Filth Flies. Proc. Workshop, Feb. 4-5, 1981, Gainesville, Florida. Sci. Educ. Admin., USDA, Washington, D.C. 212 p.

Moon, R. D., I. L. Berry & J. J. Petersen. 1982. Reproduction of Spalangia cameroni Perkins (Hymenoptera: Pteromalidae) on table fly (Diptera; Muscidae) in the laboratory. J. Kansas Ent. Soc. 55: 77-85.

Morgan, P. B. 1980. Sustained releases of Spalangia endius Walker (Hymenoptera: Pteromalidae) for the control of Musca domestica L. and Stomoxys calcitrans (L.) (Diptera: Muscidae). J. Kansas Ent. Soc. 53: 367-72.

Morgan, P. B. 1980. Mass culturing three species of microhymenopteran pupal parasites, Spalangia endius Walker, Muscidifurax raptor Girault and Sanders, and Pachycrepoideus vindemiae (Rondani) (Hymenoptera: Pteromalidae). Proceedings VIII National Meeting on Biological Control, Tecoman, Colima, Mexico, April 22‑25, 1980.

Morgan, P. B. 1981a. The potential use of parasites to control Musca domestica L. and other filth breeding flies at agricultural installations in the southern United States, p. 11-25. In: Patterson, R. S., P. G. Koehler, P. B. Morgan & R. L. Harris (eds.), Status of Biological Control of Filth Flies. Proc. Workshop, Feb. 4-5, 1981, Gainesville, Florida. Sci. Educ. Admin., USDA, Washington, D.C. 212 p.

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Morgan, P. B. & R. S. Patterson. 1975a. Field parasitization of house flies by natural populations of Pachycrepoideus vindemiae (Rondani), Muscidifurax raptor Girault and Sanders, and Spalangia nigroaenes Curtis. Florida Ent. 58: 202.

Morgan, P. B. & R. S. Patterson. 1975b. Possibilities of controlling stable flies and house flies with protelean parasitoids. Proc. 46th Ann. Mtg. Florida Anti-Mosquito Contr. Assoc., Ft. Meyers. p. 29-35.

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Morgan, P. B. & R. S. Patterson. 1989. Efficiency of target formulations of pesticides plus augmentative releases of Spalangia endius Walker (Hymenoptera: Pteromalidae) to suppress populations of Musca domestica L. (Diptera: Muscidae) at poultry installations in the southeastern United States. In: R. S. Patterson & D. A. Rutz (eds.), Biocontrol of Arthropods Affecting Livestock and Poultry. Ent. Soc. of Amer. Misc. Pub. (Accepted by R. S. Patterson, February 13, 1989).

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Propp, G. D. & P. B. Morgan. 1984. Multiparasitism of house fly, Musca domestica L., pupae by Spalangia endius Walker and Muscidifurax raptor Girault and Sanders (Hymenoptera: Pteromalidae). Environ. Ent. 12: 1232-38.

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