IX. and Mites Acarina

1. PARASITES OF TICKS

RICKETTSIAE Fusarium (killed eggs in water) (Lom- of Argas not of public health importance have yielded intracellular -like micro-organisms, and bardini, 1950). work in progress includes a survey of representatives of Unidentified fungus other genera (Roshdy, 1961). Boophilus calcaratus (fine white filaments covered surface of ticks, which died in one week owing to obstruction of stigmata) (Oswald, 1938a, b). Bacillus cereus niverus (have many chitinized papules, mostly milky white on body; one tick had 1000 papules) americanum (isolated numerous times) (Hoogstraal, 1956). (Steinhaus, 1946). moubata (opaque white spot thought to Bacteria be fungus is probably fluid from Malpighian tubules) Ornithodorus sevignyi (enormous masses of threads of (Hoogstraal, 1956; Wellman, 1907). bacteria in Malpighian tubules) (Leishman, 1909). Ornithodoros savignyi (pathogenic); 0. moubata (Hoog- Pasteurella tularensis straal, 1956). Rhipicephalus bursa (nymphs have fungus-like disease) (Parker, 1934). (Schulze, in Olenev & Rozdestvenskaja, 1933). Dermacentor silvarum (60% of nymphs and larvae died) (Zasuhin & Tihomirova, 1936). Haemaphysalis concinna (Olsuf'ev & Petrov, 1960). PROTOZOA Salmonella enteritides MASTIGOPHORA (oral infection led to death) Coelomoplasma hyalommae (Parker & Steinhaus, 1943). Hyalomma aegyptium (= H. syriacum) (occurs in coelom Serratia marcescens of tick) (Brumpt, 1938c). Dermacentor andersoni (killed a number of ticks that C. rhipicephali had ingested infected guinea-pig blood) (Steinhaus, 1942). Rhipicephalus bursa (in coelom of tick, no mortality caused; 5.7% of ticks infected) (Brumpt, 1938d). SPIROCHAETALES Crithidia christophersi sogdianum Rhipicephalus sanguineus (in thoracic muscles, legs, and Argasid ticks (Sidorov, 1960). body cavity) (MacHattie & Chadwick, 1930; Novy, B. hispanica (= Treponema hispanicum) MacNeal & Torrey, 1907). Ornithodoros erraticus (Buen, 1937). C. haemaphysalidis Haemaphysalisflava; H. birmaniae (Patton & Strickland, B. duttonii 1908). Ornithodoros moubata (Wolbach, 1914). C. hyalommae B. kochii Hyalomma aegyptium (infects body cavity) (O'Farrell, Ornithodoros moubata (Wolbach, 1914). 1913a, b). Hyalomma excavatum (= H. anatolicum) (Arifdlanov & FUNGI Nikitina, 1961). Beauveria cinerea (=Botrytis cinerea) (Cryptoplasma rhipicephali) ricinus (considered for control of ticks in pastures) These supposed parasites proved to be sperm cells of (Boicev & Rizvanov, 1960). Rhipicephalus sanguineus) (Chatton & Blanc, 1916a, b).

- 80 TICKS AND MITES 81

Trypanosoma christophersi Dermacentor reticulatus (Brumpt, 1930). Rhipicephalus sanguineus (Brumpt, 1938a). (Cobb, 1942; Cooley, 1927; Larrousse, King & Wolbach, 1928; Smith & Cole, 1943). Trypanosoma sp. Dermacentor venustus (Brumpt, 1913). infection levels result in low Hyalomma pusillum (high Dermacentor sp. (recommended for control) (Zasuhin, egg production) (Brumpt, 1938a). 1934). SPOROZOA Haemaphysalis concinna (Cooley, 1927; Brumpt, 1913, Nosema ixodis 1930b). (in nymph) (Weiser, 1957, 1959). H. concinna (19% death of nymphs in nature) (Pervo- majskij, 1947). The literature on the role of ticks as vectors of babe- sioids is not referred to here (see Lavier, 1921). Other Haemaphysalis inermis (Brumpt, 1930; Thompson, 1943). blood protozoa transmitted by ticks include Hepatozoon Haemaphysalisjaponica douglassi (Pervomajskij, 1947). argantis (, Argas brumpti-Garnham, 1954) and Haemaphysalis leachii leachii (Cooley, 1927, 1934a). Haemogregarina mauritanica (vector, Hyalomma aegyp- Haemaphysalis leporis-palustris (Pervomajskij, 1947; tium-Laveran & Pettit, 1909). Smith & Cole, 1943). Hyalomma aegyptium impressum (Thompson, 1943). NEMATODA Hyalomma asiaticum; H. dromedarii (Bernadskaja, 1939). Filaria grassi Hyalomma dromedarii (Pervomajskij, 1947). Rhipicephalus sanguineus (enter body cavity from gut) Hyalomma (as H. aegyptium) rufipes (Cooley, 1934a; (Nob, 1901). Cooley & Kohls, 1940). F. quadrispina Hyalomma transiens; H. (as H. aegyptium) truncatum (Cooley, 1934a). Ixodes ricinus (Baldasseroni, 1909). Ixodes cookei (Cooley, 1934b). Ixodes dentatus (Smith & Cole, 1943). INSECTA Ixodes muris (Smith & Cole, 1943). (1) HYMENOPTERA (Alfeev, 1941; Blagovescenskij, 1948; Pomerancev, 1950). CHALCIDOIDEA Ixodes persulcatus (7 % mortality of nymphs in nature) ENCYTRIDAE (Pervomajskij, 1943, 1947). Hunterellus hookeril (= Hunterellus caniphila, Ixo- Ixodes ricinus (Alfeev, 1941; Du Buysson, 1912). diphagus caucurtei, Habrolepis caniphila) Ixodes ricinus (10%-17% of nymphs infected in nature; up to 95% infection level achieved experimentally) Amblyomma americanum (Bishopp, 1934). (Brumpt, 1913, 1930b). Amblyomma tholloni (Fiedler, 1953; Santos Dias, 1948). Ixodes ricinus californicus (Gregson, 1942). Dermacentor andersoni (Cooley, 1927; Cooley & Kohls, Ixodes ricinus scapularis (Bishopp, 1934; Larrousse, 1934). King & Wolbach, 1928; Smith & Cole, 1943). Dermacentor parumapertus (Cooley, 1929a). Rhipicephalus appendiculatus (Cooley, 1934a). Dermacentorparumapertus marginatus (Thompson, 1943). Rhipicephalus evertsi evertsi (Cooley, 1929a, 1934a). Dermacentor nitens (Gahan, 1934). Rhipicephalus oculatus (Cooley, 1934a). Dermacentor pictus (Alfeev, 1941). Rhipicephalus sanguineus sanguineus (Brumpt, 1930b; 1 The tick parasite Hunterellus hookeri was released in Montana Costa Lima, 1915; Smith & Cole, 1943). and adjacent area in 1927-32 to control Dermacentor andersoni. Rhipicephalus sanguineus sanguineus About 4 158 000 wasps were released but the parasite did not become (90% of nymphs established nor did it reduce the tick population (Cooley & Kohls, parasitized) (Philip, 1931a,b, 1954). 1934). This parasite, originating in France, was also released in Massachusetts and adjacent islands in 1926 in an attempt to control Rhipicephalus sanguineus sanguineus (= Habrolepis sp.) Dermacentor variabilis. It became established and persisted for at (Blanc, Goiran & Baltazard, 1937; Risbec, 1944; Cooley, least 12 years. Nymphs of Dermacentor variabilis became parasitized by the chalcids from a distance of 100 yards. Larrousse, King & 1929a; Fiedler, 1953; Steyn, 1955). Wolbach (1928) stated that it survived the winter after release and that the tick population on Naushon Island was significantly reduced Rhipicephalus sanguineus sanguineus (= R. texanus) the following season in contrast with high populations near by. In (distribution of H. hookeri is Africa, Asia, Europe, and 1937-39, 91 000 adults were released on Martha's Vineyard Island, Mass., but no recoveries were made during the following years. North and South America) (Thompson, 1943). Workers in the USSR have also studied the possibilities of using this wasp for controlling Ixodes persulcatus, but no data on releases have Rhipicephalus sp. (Blanc, Goiran & Baltazard, 1938; been seen. Howard, 1907, 1908). 6 82 TICKS AND MITES

Hunterellus theilerae 1942; Howard, 1907; Joyce & Eddy, 1943; Larson, 1937). Hyalomma marginatum rufipes (Hoogstraal & Kaiser, Haemaphysalis leporis-palustris (3.3% of nymphs and 1961; Kaiser & Hoogstraal, 1958). 7.4% of larvae were parasitized) (Larson & Green, 1938). Hyalomma truncatum (=H. transiens); Rhipicephalus Ixodes cookei ? (Cooley, 1934b). oculatus (distribution of H. theileri is South and South- Ixodes dentatus (Cobb, 1942). West Africa and Egypt) (Fiedler, 1953). Ixodes dentatus (Ixodiphagus texanus is restricted to Ixodiphagus hirtus the USA) (Smith, 1945). Ixodespersulcatus (Nikol'skaja, 1950; Pomerancev, 1950). (2) DIPTERA L mysorensis Ormthodoros sp. (Mani, 1941). PHORIDAE L. texanus Megaselia rufipes Dermacentor variabilis (Cobb, 1942; Smith & Cole, 1943). Ixodes ricinus (larvae parasitize adult ticks and eat Haemaphysalis leporis-palustris (Bishopp, 1934; Cobb, internal organs) (Olenev, 1941).

2. PREDATORS OF TICKS

ARACHNIDA TIISTERIDAE Spiders Hyalomma dromedarii (Vlasov, 1940). Argas persicus (eat eggs and larvae) (Bishopp, 1913a). Teutena triangulosa STAPHYLINIDAE Rhipicephalus sanguineus sanguineus (fed on immature and adult ticks and reduced prevalence in Corsica) Juracekia asphaltina (Sautet, 1936). Rhipicephalus sanguineus ? (Flegontova, 1938). ACARINA (3) HEMIPTERA Argas persicus Cannibalistic (Nikitina, 1959). Phonergates bicolor Ornithodoros moubata (sucks blood of tick) (Weilman, Hyalomma mauritanicum 1907). Cannibalistic (A. Sergent, 1930). INSECITA (4) HYMENOPTERA (1) NEUROPTERA Ants Ornithodoros moubata (carry off eggs and larvae) (Hoog- MYRMELIONIDAE straal, 1956). Ornithodoros moubata (on nymphs) (Ghesquiere, 1922). Monomorium minimum (2) coLEomRA Otobius megnini (adults) (Parish, 1949). CARABIDAE Hyalomma dromedarii (eat larvae and nymphs) (Vlasov, M. minutum 1940). Argas persicus (eats eggs and larvae) (Bishopp, 1913a). Calosoma auropunctata, Calosoma denticolle, Carabus Pogonomyrmex barbatus molefaciens clathratus, Carabus coriaceus, Carabus cumanus, and Otobius megnini (Parish, 1949). Sphodrus leucophthalmus Solenopsis sp. Ate average of 5 engorged nymphs and 3-4 adults daily of Ixodes ricinus, Dermacentor marginatus and Boophilus Boophilus annulatus (eats eggs and engorged females) calcaratus confined in jars (Pelipeicenko, 1957). (Bishopp, 1932). TICKS AND MITES 83

(5) LEPIDOPTERA Amblyomma variegatum (186 ticks in 16 birds); Boophilus decoloratus (51 ticks in 13 birds); Rhipicephalus appendi- Tineola biselliela culatus (500 ticks in 3 birds); R. evertsi evertsi (3 ticks Argas persicus (caterpillars eat tick larvae in indoor in 58 birds); R. simus simus (15 ticks in 4 birds) (Moreau, colonies) (Voilmer, 1931). 1933). Ornithodoros moubata (Vollmer, 1931). Boophilus decoloratus (38 ticks in 4 birds) (Van Someren, 1951). VERTEBRATA Tick bird (Buphagus sp.) BIRDS Rhipicephalus kochi ? (Moreau, 1933). Poultry Wagtail (Motacilla alba) Argas persicus (Bishopp, 1913a). Hyalomma marginatum marginatum (= H. mauritanicunt) Ornithodoros moubata (Hoogstraal, 1956). (Ed. Sergent et al., 1945). Egret (Bubulcus ibis) Hyalomma marginatum marginatum (= H. mauritanicum) MAMMALS (Ed. Sergent et at., 1945). Tick bird (Buphagus africanus subspp.) Hibernating Hyalomma marginatum marginatum (=H. mauritanicum) (Ed. Sergent et al., 1945). Amblyomma variegatum; Boophilus decoloratus (1 12 ticks in 7 birds); Rhipicephalus evertsi evertsi (3 ticks in 2 Hibernating Ornithodoros moubata (Hoogstraal, 1956). birds); Rhipicephalus simus simus (4 ticks in 2 birds) (Van Someren, 1951). Microtus arvalis Tick bird (Buphagus erythrorhynchus erythrorhyn- Dermacentor pictus (Petrov & Kuceruk, 1951). chus) Ornithodoros moubata (Hoogstraal, 1956).

3. PARASITES OF MITES

FUNGI PROTOZOA

SPOROZOA BLASTOMYCETES Blood protozoa transmitted by mites include Hepatozoon muris (vector, Laelaps echidninus-Miller, 1908) and H. Chlamydomyces akamushi balfouri (vector, Haemolaelaps aegyptius-Hoogstraal & 25% infected (Kitashima & Miyajima, 1918). Kaiser, 1961).

4. PREDATORS OF MITES

ACARINA Cunaxa sp. Bdella sp. Eutrombicula alfreddugesi (Williams, 1946). Eutrombicula alfreddugesi (Williams, 1946).