I. -Musca domestica and Related Species

1. PARASITES

BACTERIA Aspergillus sp. Bacillus cereus Usui (1960). A filterable principle interferes with development (Briggs, Beauveria bassiana 1960). Has knock-down effect on ; 100% kill in three hours B. lutzae (Dresner, 1949). Pathogenic for adults (Brown, 1927). Entomophthora 1 americana B. thuringiensis 50% kill in 24 hours (Dresner, 1949). Can be used against larvae, but not highly effective E. muscae (Hall & Arakawa, 1959). Causes death of many adults especially in autumn; A filterable principle interferes with development (Briggs, larvae fed on a culture died as pupae (Graham-Smith, 1960). 1916; Hesse, 1913; Lakon, 1919a). When fed to caged chickens, achieved a variable measure Experimental infection killed 100% (Bernstein, 1914). of control in their droppings (Bums, Wilson & Tower, Mucor heimalus and M. racemosus 1961). When fed Mucor-containing syrup flies died with infection Bacterium delendae-muscae resembling that due to Entomophthora (Bernstein, 1914). Larvae ingest, and die at end of pupal stage (Roubaud & Mucor sp. Descazeaux, 1923). Usui (1960). Bact. mathisi Stigmatomyces baeri Kills in three hours or more after ingestion (Roubaud & Treillard, 1935). Musca planiceps (infected cut leg) (Senior-White, 1945). Micrococcus rushmorei Pathogenic for adults (Brown, 1927). MASTIGOPHORA Neisseria luciliarum Crithidia luciliae (=Herpetomonas luciliae) Pathogenic (Brown, 1927). Found in 14% of flies collected; experimentally infected Serratia kielensis 3 of 20 flies (Strickland, 1911; Wallace & Clark, 1959). Adults died earlier when fed (Cao, 1906a). Herpetomonas muscarum (= H. muscidarum, H. S. marcescens luciliae, H. muscae-domesticae) Pathogenic for adults (Steinhaus, 1951). 50% of flies infected; intestinal tissue said to be invaded (Macfie, 1917a; Ross & Hussain, 1924; Wallace & Clark, Staphylococcus flaccicifex 1959). Pathogenic for adults (Sweetman, 1936). In haemocoele of larval M. domestica (Kramer, 1961a). Staph. muscae Effects unknown (Noguchi, 1926). Caused fatal infection in adults (Glaser, 1924a). Infected larval M. domestica; no pathology (Becker, 1923, Not pathogenic and passage in fly did not increase 1924; Bellosillo, 1937; Fantham & Robertson, 1927). virulence (Glaser 1926a). In gut of adult (Glaser, 1926b). Is tissue parasite (Ross & Hussain, 1924). FUNGI 42.5 % infected (Macfie, 1917a).

Aspergillus parasiticus 1 I have followed other contemporary workers in regarding Entomophthora Fresenius as the valid name for this , with Pathology uncertain (Marchionatto, 1945). Empusa Cohn as a synonym (cf. MacLeod, 1963).

- 11 - 12 HOUSEFLIES

SPOROZOA H. muscae (=Filaria muscae) Larvae Octosporea muscae domesticae ingest parasite, which causes necrosis of fat body; it encysts in pupae, and migrates to head of fly (Mello & In midgut, hind gut, Malpighian tubules and larval yolk Cuocolo, 1943). (Chatton & Krempf, 1911; Fantham & Porter, 1958; Flu, 1911; Kudo, 1924c; Steinhaus, 1946). ACARINA Spiroglugea sp. In Malpighian tubules, fat body and gut cells (Chatham & (1) MESOSTIGMATA Krempf, 1911; Fantham & Porter, 1958; Flu, 1911; Kudo, 1924c; Steinhaus, 1946). Gamasid mites Telomyxa sp. Immature forms swarm on flies emerging from rubbish heaps (Banks, 1905). In Malpighian tubules, haemocoele, and larval fat body (Chatham & Krempf, 1911; Fantham & Porter, 1958; Flu, 1911; Kudo, 1924c; Steinhaus, 1946). MACROCHELIDAE Thelohania ovata Glyptholaspis confusa (= Macrocheles plumiventis) In rectum of scalaris (Dunkerly, 1912). Evans & Browning (1956) T. thomsoni Average 9.9, range 7.4-13.2, fly eggs and larvae eaten per day by adult female mites, which were not found attached In epithelium of chyle, stomach and proximal intestine of to adult flies (Axtell, 1961a, b). Muscina assimilis (Kramer, 1961b). Gave encouraging results in field experiments, in com- bination with Macrocheles muscae domesticae, and shared with this species credit for up to 96% control PLATYHELMINTHES when added to cow manure seeded with fly eggs (Axtell, Choanotaenia infundibulum 1961a). In body of larvae (Grassi & Rovelli, 1892). G. fimicola (=Holotaspis marginatus) Berlese (1882, 1912). Davainea cesticillus Filipponi & Pegazzano (1960). Not known whether this fowl cestode harms its fly (Ackert, 1918). Holotaspis sp. D. tetragona Thought to be parasitic; occurred singly or in groups on any part of the fly for up to 192 hours (Graham-Smith, Not known whether this avian cestode harms its fly host 1916). (Bushnell & Hinshaw, 1924). Macrocheles glaber Syngamus trachea Attached to houseflies (Filipponi, 1960). Infection of adults causes sluggishness, rendering them easily caught by birds (Clapham, 1939). M. insignitus Reared on fly eggs and horse manure; not known to be phoretic on adult flies (Berlese, 1918; Filipponi, 1960). NEMATODA M. medarius Allantonema muscae Found in Canada (Chant, 1960). Musca vicina (Roy & Mukherjee, 1937b). Found in USA (Axtell, 1961a). A. stricklandi M. muscaedomesticae (=Acarus muscae, Macrocheles Musca vicina (Roy & Mukherjee, 1937a). muscae, Acarus marginatus, Holotaspis badius; world-wide distribution) Habronema megastoma Houseflies are heavily infested in autumn in (Ewing, Larvae causes infected, adults die early; tumours in 1913; Zanini, 1930); see also Filipponi (1955a; 1955b). Malpighian tubules (Johnson, 1913; Roubaud & Des- cazeaux, 1921, 1922a, b). Found commonly on adult houseflies in , the mites feeding on eggs and lst- larvae, especially in dung H. microstoma where the flies breed (Pereira & de Castro, 1945, 1947). Larvae infected, adults die early; parasitizes Female mites feed from ventral surface of base of (Johnston, 1913; Roubaud & Descazeaux (1922a, b). of adult fly (Ewing, 1913). HOUSEFLIES 13

Macrocheles muscaedomesticae (continued) (3) SARCOPTIFORMES Average 2.7, range 2.1-4.3, fly eggs and larvae eaten per ACARIDAE day by adult female mites (Axtell, 1961a, b). Acarus muscarum Gave encouraging results in field experiments, in com- On bination with Glyptholaspis confusa, and shared with head and neck of fly (Linnaeus, 1758). this species credit for up to 96% control when added to A. reflexus cow manure seeded with fly eggs (Axtell, 1961b). Larval form on body of fly (Attimonelli, 1940). Mites were reared in the laboratory, predation being demonstrated in the laboratory and to some extent in Tyroglyphus siro the field (Rodriguez & Wade, 1961; Wade & Rodriguez, Nymphs and adults found on adult flies (Graham-Smith, 1961). 1916). M. robustulus (=Macrocheles rothamstedensis) Berlese (1904); Evans & Browning (1956). PSEUDOSCORPIONIDA Average 4.6, range 3.3-5.6, fly eggs and larvae eaten per Chelifer panzeri day by female mites (Axtell, 1961a, b). Attached as parasites (Preudhomme de Borre, 1873). M. subbadius Chernes nodosus Berlese (1904). Chelifer stage dispersed (Godfrey, 1909). Average 1.2, range 0.9-1.5, fly eggs and larvae eaten per day by female mites (Axtell, 1961a, b). Pseudoscorpions Life of fly may be shortened by the burden, up to 10 per M. subglaber fly (Hewitt, 1914a). Attached to houseflies (Filipponi, 1960). INSECTA PARASrTIDAE Gamasus coleoptratorum Attached to flies especially in autumn, feeding on weak BRACONIDAE ones (Graham-Smith, 1916). Alysia manducator (2) TROMBIDIFORMES Attacks full-grown (Froggatt, 1921; James, 1928). TROMBIDIIDAE Opius nitidulator Atomus parasiticus (=Acarus parasiticus) Thompson (1943). On maggots; since the Trombidiidae parasitize , this group deserves greater attention; the nymphs and adults are known to eat eggs and early larval of pachycerus the host (Latreille, 1795). Effectively attacks pupae; in experiments parasitized 92 % Metathrombidium poriceps (Roy & Siddons, 1939; Roy, Siddons & Mukherjee, On adults (Ewing & Hartzell, 1918). 1940). " " Red mite CYNIPIDAE Frequent in China (Feng, 1933). Eucoila impatiens Trombidium muscae Parasitizes (Howard, 1911a). Larvae on hou%effies (Howard, 191 la). Eucoila sp. T. muscarum Parasitizes in Fiji (Simmonds, 1940). Larvae at wing bases of adult ffies (Howard, 191 la). Very common; up to 100% infestation (Ewing & Hartzell, Kleidotoma marshalli 1918). Attacks maggots (James, 1928). T. parasiticum Kleidotoma sp. Red larvae on flies (Murray, 1877). Attacks maggots (James, 1928). 14 HOUSEFLIES

DIAPRHIDAE Pachycrepoideus vindemmiae (=P. dubius) True parasite (Girault & Sanders, 1910; Hewitt, 1914a; Ashmeadopria spp. Johnston & Tiegs, 1921). Up to 10% parasitized (Puerto Rico Agricultural Experi- ment Station, 1938). Prospalangia platensis Parasitized 60 %-74Y% of pupae (Brethes, 1915). Hemilexomyia abrupta Scarce parasite (Johnston & Tiegs, 1921). Spalangia cameroni Introduced into Hawaii and Fiji for fly control (Sim- Muscidifurax raptor monds, 1929a). From pupae (Girault & Sanders, 1910). S. hirta Trichopria commoda Parasitizes pupae (Richardson, 1913a). Being mass-produced at Kyoto Municipal Institute of for liberation in a control campaign S. muscae (Muesebeck, 1961). Parasitizes pupae (Howard, 1911a). S. muscidarum ENCYRTIDAE Parasitizes pupae (Johnston & Tiegs, 1921; Pinkus, 1913; Tachinaephagus giraulti Richardson, 1913b). Reared from Musca (Ferriere, 1933). S. nigra Parasitizes pupae (Howard, 1911a; Richardson, 1913a). FIGITIDAE S. philippinensis Figites anthomyiarum Introduced into Hawaii and Puerto Rico for fly control Reared from maggots (Hewitt, 1914a; Howard, 191 la). (Fullaway, 1917; Puerto Rico Agricultural Experiment Station, 1938). F. scutellaris Reared from maggots (Howard, 191 la). S. stomoxysiae Parasitizes pupae (Lindquist, 1936; Simmonds, 1940; F. striolatus Thompson, 1943). Reared from adult housefly (Hewitt, 1914a). Spalangia spp. Tried for control in Guam and Fiji (Simmonds, 1922). PTEROMALIDAE Reared in laboratory in 15-17 days, caused 75 %-85 % Nasonia vitripennis (= Mormoniella vittripennis, parasitization in Guam (Vandenberg, 1930, 1931a). Nasonia brevicornis) In (Parker & Thompson, 1928). Parasitizes pupae (Fujita, 1932; Girault & Sanders, 1910; Hewitt, 1914a; Moursi, 1946; Parker & Thomson, 1928; Stenomalus muscarum Smirnov & Kuzina, 1933; Smirnov & Vladimirov, 1934; Parasitizes pupae (Howard, 191 la). Vladimirov & Smirnov, 1934). Found on hibernating houseflies (Waterston, 1916c).

2. PREDATORS

CHILOPODA ARANEIDA Scutigera coleoptrata (= S. forceps) Among the numerous records of predation by spiders the Destroys flies at night (Hewitt, 1914a). following may be mentioned: Destroys large numbers of adult flies (Hewitt, 1914a; Agelena naevia Howard & Hutchison, 1917). Howard (191 la). S. smithii Araneus sericatus (=Epeira sericata) Feeds on adult houseflies at night (West, 1951). West (1951). HOUSEFLIES 15

Coenotele gregalis Mesembrina meridiana Diguet (1915a, b). Predacious (Simmonds, 1940). Plexippus paykulli Mathis (1938). Larvae eat housefly larvae (Por6inskij, 1913b; Schoof & Siverly, 1954). Poecilochroa convictrix Diguet (1915a, b). Ophyra anthrax Preys on larvae (Lejkina, 1942). Salticus scenicus Hewitt (1914a). 0. leucostoma Preys on larvae (Lejkina, 1942). Steatoda borealis Howard (191 la). STRATIOMYIIDAE Teutana triangulosa Derbeneva-Uhova (1940). Howard (191la). illucens Theridion tepidariorum Not predacious, but larvae in privies keep medium too Howard (191 la). moist and " soupy " for production of housefly larvae; insecticide treatment against resistant houseflies kills the Hermetia larvae and results in increase of housefly pro- PSEUDOSCORPIONIDA duction (Furman, Young & Catts, 1959; Kilpatrick & Schoof, 1959). Chermes nodocus Hewitt (1914a). (4) HYMENOPTERA INSECTA FORMICIDAE Iridomyrmex humilis (1) ORTHOPTERA Destroys eggs, larvae, and pupae (Howard, 191 la). MANTIDAE Monomorium minimum Eat many adults (Hewitt, 1914a). Destroys eggs, larvae and pupae (Howard, 191 la).

(2) DERMAPTERA Paratrechina longicornis Destroys eggs, larvae and pupae (Pimentel, 1955). Destroy larvae (Compere, 1912). Pheidole megacephala (3) DIPTERA Destroys eggs, larvae and pupae and newly emerged adults (Bridwell, 1918; Illingworth, 1915). ANTHOMYIIDAE Gave 50% control (Phillips, 1934; Simmonds, 1928, Scatophaga stercoraria 1940, 1958). Adults eat housefly adults (Hewitt, 1914b). Ants ate eggs and larvae, and in an experiment brought about 99% control (Phillips, 1934). ASILIDAE P. subarmata borinquensis Eats larvae (Pimentel, 1955). Laphria canis Eats adult flies (Howard, 191 la). Solenopsis corticalis Eats eggs and pupae (Pimentel, 1955). S. gemminata rufa (=S. g. diabola) Hydrotaea dentipes Destroys eggs, larvae and pupae (Howard, 191 Ia). Larvae eat housefly larvae; very important (Lejkina, Killed 50 % of larvae and nearly emerged adults (Pimentel, 1942; Porcinskij, 1913a). 1955). 16 HOUSEFLIES

Tapinoma melanocephalum HISTERIDAE Eats eggs and larvae (Pimentel, 1955). Placodes caffer Ants, 5 species Simmonds (1940). Gave 72%-91 % control in test cages (Pimentel, 1955). Platylister chinensis (=Hister chinensis) Fed on larvae, eating 10-20 per 24 hours; 1400 introduced OTHER HYMENOPTERA into Fiji for fly control (Lever, 1938). Bembix spinolae Helped to control houseflies following introduction into and Samoa Predatory on adult flies (Parker, 1917). Fiji (Simmonds, 1958). Larvae ate 8 fly larvae per day (Peckham & Peckham, 1905). SCARABAEIDAE Copris incertus prociduus Bembix sp. Introduced into Fiji; not effective in fly control (Sim- Predatory on flies (Vaillard, 1914). monds, 1929b). Mellinus arvensis Adults ate housefly adults (Herold, 1922). STAPHYLINIDAE Nysson sp. Eat larvae (Hewitt, 1914a; Howard, 191 la). Adults ate housefly adults (Froggatt, 1917). Baryodma ontarionis (=Aleochara bilineata) Eats larvae, as parasite (Colhoun, 1953; Senior-White, Polistes olivaceus (=P. hebraeus) 1945; Sweetman, 1958). Predatory on larvae and adults; reduced fly incidence in Fiji (Jepson, 1915). VERTEBRATA Stizus spp. Adults ate housefly adults (Froggatt, 1917). AMPHIBIA Frogs Tenthredo variegatus Eat adults (Hewitt, 1914a). Adults ate housefly adults (Venables, 1914). Toads Vespa orientalis Eat adults (Hewitt, 1914a). Adults ate housefly adults (Zmeev, 1941). Vespa spp. REPTILES Adults ate housefly adults (Hewitt, 1914a; West, 1951). Lizards Eat adults (Hewitt, 1914a). Vespula germanica (= Vespa germanica) Adults ate housefly adults (Herold, 1922). BIRDS Poultry (5) COLEOPTERA Eat larvae and adults (Hewitt, 1914a). CARABIDAE Wild birds Agonoderus spp. Eat adults (Hewitt, 1914a). Eat larvae (Howard, 191 la). MAMMALS Harpalus spp. Cats Eat larvae (Howard, 191 la). Eat adults (Hewitt, 1914a). Platyhus spp. Rats Eat larvae (Howard, 1911 a). Eat adults (Hewitt, 1914a).