IV. Sandflies and Midges - Psychodidae and Ceratopogonidae

1. PARASITES

RICKETTSIAE Grubyella ochoterenai Culicoides phlebotomus (exposed adults died, exhibiting Ricksettia sp. fungal outgrowths) (Ciferri, 1929). Phlebotomus vexator (in gonads) (Hertig, 1936). Penicillium glaucum P. papatasii (killed larvae in the laboratory) (Zotov, 1930). BACTERIA PROTOZOA Bacteria Ceratopogonidae (larvae) 1 (Mayer, 1934). (1) MASTIGOPHORA Culicoides nubeculosus (in fat body of larvae) (Lawson, 1951). Crithidia sp. C. nubeculosus (Steinhaus, 1946). P. baghdadis (Adler & Theodor, 1929). Pseudomonas sp. Herpetomonas phlebotomi Culicoides salinarius (Becker, 1958). P. minutus (10% incidence in India) (Mackie, 1914; Patton, 1919). Spirochaeta phlebotomi (= Treponema phlebotomi) P. minutus (in gut) (Shortt, 1925). P. perniciosus (in gut) (Pringault, 1921a). P. papatasii (in gut) (Mackie, 1914).

(2) SPOROZOA FUNGI (a) GREGARINIDA Aspergillus sp. Lankesteria ? Phlebotomus spp. (young larvae may become entangled P. papatasii (no pathological damage) (Missiroli, 1932). in mycelium; spores germinate in larval intestine, the mycelium invading muscles of thoracic area and causing Monocystis mackiei death; this fungus is highly pathogenic in laboratory P. argentipes (25 % in nature) (Shortt & Swaminath, cultures) (Hertig & Johnson, 1961). 1927). P. papatasii (Missiroli, 1929b). Entomophthora papatasii P. papatasii (Marett, 1915). (b) HAEMOSPORIDIIDA E. phlebotomnus Haemoproteus canachites P. papatasii (Adler & Theodor, 1929). Culicoides sphagnumensis (Fallis & Bennett, 1961a). P. longipalpis (Adler, personal communication, 1961). H. nettionis Fungus Culicoides nr. piliferus (Fallis & Bennett, 1961a). P. papatasii; P. sergenti (destroyed eggs) (Adler & Haemoproteus spp. Theodor, 1929). Culicoides crepuscularis (oocysts found in stomach walls); 1 Identification of parasite uncertain. C. stilobezzioides (Fallis & Bennett, 1961a).

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Hepatozoon (oocysts) Culicoides peregrinus (in haemocoele) (Abe, 1927; Ghosh, 1925; Grasse & Boissezon, 1929). P. papatasii (Adler & Theodor, 1929). Leucocytozoon caulleryi Culicoides arakawae (Akiba, 1960). NEMATODA

Schizocystis gregarinoides Filaria perstans Bezzia solstitialis ( Ceratopogon solstitialis) (in gut) Culicoides (Sharp, 1928). (Leger, 1900). Wuchereria bancrofti S. gregarinoides is ingested and the oocysts release sporozoites; these attach to the gut epithelium, which P. sergenti var. mongolensis (Yao, Wu & Sun, 1938). elongates up to 150 /s. After reproduction by the game- tocytes, oocysts are eliminated in the faeces and the cycle Unidentified nematodes is repeated. There is little pathological damage to the P. papatasii; P. sergenti (in haemocoele and ovaries) ceratopogonids. (Adler & Theodor, 1929).

(3) MICROSPORIDIA/MONOCNIDEA ACARINA COUGOURDELLIDAE Allothrombium sp. Spiroglugea octospora (= Spironemna octospora) Culicoides heliophilus (attached to abdomen of female) Ceratopogon sp. (in larval fat body) (Jirovec, 1936; Kudo, (Becker, 1958). 1924c; Leger & Hesse, 1922; Weiser, 1961). Evansiella culicoides Toxoglugea vibrio (= Toxonema vibrio) Culicoides circumscriptus; C. maritimus; C. pulicaris; Ceratopogon sp. (larval fat body) (Jirovec, 1936; Kudo, C. punctatus (may interfere with flight of adults) (Becker, 1924c; Leger & Hesse, 1922; Weiser, 1961). 1958). " Hydrachnid mites" NOSEMATIDAE P. algeriensis (on thorax) (Parrot, 1919). Phlebotomus spp. (on thorax) (Marett, 1915). Nosema sphaeromiadis Raphignathus youngi Sphaeromias sp. (in larval fat body) (Weiser, 1957, 1961). P. papatasii (adults) (Hirst, 1926). P. papatasii; Phlebotomus spp. (Burakova, 1930). (4) CILIOPHORA/CILIATA Trombidium hindustanicum " Ectoparasitic cysts " P. papatasii (larvae) (Hirst, 1926). Culicoides cubitalis (larvae); C. pallidicornis (larvae) " Mites" (Kettle & Lawson, 1952). Culicoides austeni (attached to legs and body) (Sharp, " Ciliates " 1928). Culicoides austeni (Becker, 1958; Sharp, 1928).

Perezella sp. INSECTA Culicoides odibilis; C. pulicaris; C. riethi; C. salinarius (probably contributes towards larval mortality) (Becker, HYMENOPTERA 1958). Planidium (type) Tetrahymena pyriformis (=Probalantidium knowlesi, P. freetownensis var. sudanicus (adults) (Lariviere & Balantidium knowlesii?, Leptoglena knowlesi). Abbonenc, 1958). 32 SANDFLIES AND MIDGES

2. PREDATORS

WORMS Scatopse fuscipes Nereis diversicolor Phlebotomus sp. (larvae) (Burakova, 1930). Culicoides circumscriptus; C. salinarius (under experi- " Tanypinen-larva " mental conditions worms ate no larvae in mud, but 4 worms ate 57 of 60 larvae in water in 18 hours; may Culicoides (larvae) (Mayer, 1934). be of some importance in nature) (Becker, 1958). Tachydromia minuta INSECTA Culicoides circumscriptus (Becker, 1958). " Tipulid fly " (1) ODONATA Culicoidesfurens (undoubtedly feeds on larvae and pupae) Dragonflies (Painter, 1926). Probably feed on sandflies (Painter, 1926). "Empid fly" (2) HEMIPTERA Culicoides maritimus (Becker, 1958). REDUVIIDAE Ploiaria domestica (4) HYMENOPTERA P. papatasii (feeds on engorged adults) (Schulze, 1919). Ants (3) DIPTERA Culicoides (adults) (Adler, personal communication, 1961). Bezzia sp. Culicoides sp. (ate larvae under laboratory conditions) VERTEBRATA (Thomsen, 1937). Probezzia sp. Gasterosteus aculeatus Culicoides sp. (ate larvae under laboratory conditions) Culicoides spp. (voracious predator under laboratory (Thomsen, 1937). conditions, but not in nature) (Becker, 1958).