II. VIRUS PATHOGENS OF CULICIDAE (MOSQUITOS) a Brian A. Federici Department of Entomology Division of Biological Control University of California Riverside, CA 92502, USA

a The following table and bibliography are mainly based on literature published up to and including 1973, although a few references to work published in 1974 are inicluded.

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ABSTRACTS Mary Ann Strand An iridescent virus infecting the mosquito Aedes stimulans. Anderson, J. F. (1970). J. Invertebr. Pathol., 15: 219-224.

An iridescent virus with a particle size of 135-140 nm was isolated from larvae of The virus appears to be Aedes stimulans collected from woodland pools in Connecticut. a different strain than those isolated from other insect species and the name Aedes Infected larvae were recognized by their stimulans iridescent virus is proposed. The incidence in the field was considerably less than 1%. opalescent turquoise colour. The presence of the virus in ovarian tissue suggests transovarial passage.

Electron microscope studies of the "R" and "T" strains Anthony, D. W. & Hall, D. W. (1970). Proc. 4th Int. Colloq. of mosquito iridescent virus in Aedes taeniorhynchus (Wied.) larvae. Insect Pathol., pp. 386-395.

Electron microscope studies of the "R" (regular brown-orange iridescence) and "T" (turquoise iridescence) strains of mosquito iridescent virus in 4th-instar A. taeniorhynchus larvae showed the fat body to be the primary site of replication for In tissue sections, both strains appear hexagonal or pentagonal in shape, both strains. "T" strain is approximately which suggests that each particle is an icosahedron. 30-40 nm smaller than the "R" strain.

Anthony, D. W. et al. (1973). Pathol., 22: 1-5.

A virus disease in Anopheles quadrimaculatus.

J. Invertebr.

Free and occluded virus particles were found in the cytoplasm of the midgut epithelial cells of 2nd instar larvae of A. quadrimaculatus infected with Thelohania legeri. Spherical crystals, which contained these particles, showed a macromolecular paraIn a few instances, the cuboidal crystalline lattice typical of polyhedral protein. The observations suggest crystals appeared to have coalesced to form larger crystals. that the free particles and their occluded forms may represent stages of a cytoplasmic polyhedrosis virus.

Double infection of mosquitoes with a virus and a malarial parasite. Bertram, D. S. (1965). Proc. 12th Int. Cong. Entomol. (London), pp. 766-767.

Malarial transmission by Aedes aegypti can be suppressed by concurrent infection of the The interaction may be fatal to the mosquito. vector with an arbovirus. Evidence of insect viruses in colonies of Anopheles stephensi. Bird, R. G. et al. (1970). Trans. Roy. Soc. Trop. Med. Hyg., 64(1): 28-29 (abstract).

Three different virus-like particles were found in gut lesions of adult A. stephensi and One was identified as a cytoplasmic polyhedrosis virus a cell line derived from them. Another from the midgut and was associated with difficulty in malarial transmission. The was found in the gut of individuals of the same strain but different colonies. third type was observed in cell cultures and appeared to be TMV type rods.

Chapman, H. C. (1972).

Personal communication.

38

Chapman, H. C. (1974).

Biological control of mosquito larvae.

Ann. Rev. Entomol., 19:

33-59. Review article.

Chapman, H. C. et al. (1970). Publ. Entomol. Soc. Amer., 7:

Protozoans, nematodes, and viruses of anophelines.

Misc.

134-139.

Viruses such as the mosquito iridescent virus, cytoplasmic polyhedrosis viruses of the hypodermal and gut cells, and a nuclear polyhedrosis virus are reported only from culicines, but 2 probable cytoplasmic polyhedrosis viruses, one in the hypodermal cells and the other in the gut cells are reported from larvae of Anopheles crucians. A two-year survey of pathogens and parasites of Culicidae, Chapman, H. C. et al. (1969). New Jersey Mosquito Extermin. Assoc. Proc., Chaoboridae, and Ceratopogonidae in Louisiana. 56: 203-212.

Adult and larval populations of mosquitos were sampled from 1967 to 1969. Forty-four species of mosquitos were captured and 35 were hosts to one or more pathogens. Inclusion and noninclusion viruses were recorded.

Chapman, H. C. et al. (1966). Additional hosts of the mosquito iridescent virus. J. Invertebr. Pathol., 8: 545-546. Mosquito iridescent virus has been verified in Aedes taeniorhynchus, A. fulvus pallens, Infected larvae had an external A. vexans, and Psorophora ferox from Louisiana. iridescent hue that was usually first visible in the thorax of late 3rd- or early 4thinstar larvae. Most patently infected larvae die before pupation.

Chapman, H. C. et al. (1972). Med. Hyg., 21(5): 777-781.

Predators and pathogens for mosquito control.

Amer. J. Trop. ,

The pathogens that can be or show promise of being manipulated and mass-reared in the laboratory are presented.

Chapman, H. C. et al. (1967). Pathogens and parasites in Louisiana Culicidae and New Jersey Mosquito Exterm. Assoc. Proc., 54: 54-60.

Chaoboridae.

Twenty-seven species of mosquitos were found infected with internal parasites or pathogens in southwestern Louisiana. Mosquito iridescent virus was reported in some

samples.

Clark, T. B. (1972).

Personal communication.

Clark, T. B. & Chapman, H. C. (1969). J. Invertebr. Pathol., 13: 312.

A polyhedrosis in Culex salinarius of Louisiana.

A viral disease characterized by the presence of teragonal inclusion bodies in limb anlage and hypodermal cells was found in the larvae of Culex tarsalis in California. A similar virus has been found in C. salinarius in Louisiana.

Not a true cytoplasmic polyhedrosis virus in that the inclusions studied in this paper are made up entirely of virus particles. This virus is most commonly referred to as a "tetragonal virus" because of the tetragonal nature of the inclusions. It is possibly a parvovirus.

39 Nuclear polyhedrosis and cytoplasmic polyhedrosis virus J. Invertebr. Pathol., 14: 284-286. infections in Louisiana mosquitoes.

Clark, T. B. et al. (1969).

A nuclear polyhedrosis virus infecting the gastric caeca and midgut of Aedes sollicitans and a cytoplasmic polyhedrosis virus infecting the gastric caeca and midgut of The most obvious sign of Culex salinarius were found in larvae collected in Louisiana. the disease was the whiteness of the gut wall due to the large number of inclusion Attempts at laboratory transmission of the nuclear polyhedrosis virus have bodies. yielded infection in only about 5% of the exposed larvae, but those infected died before The cytoplasmic polyhedrosis virus was easily transmitted in the laboratory pupation. but even heavily infected individuals pupated and emerged as apparently healthy adults.

Field and laboratory observations of two viral diseases in Clark, T. B. & Fukuda, T. (1971). Mosquito News, 31: 193-199. Aedes sollicitans (Walker) in southwestern Louisiana. A viral epizootic involving both a cytoplasmic polyhedrosis and a nuclear polyhedrosis Experimental results suggest occurred in populations of Aedes sollicitans in Louisiana. that a series of overlapping broods of Aedes sollicitans may lead to a buildup of infective viral material in the habitat, but a period of drying between broods appears to Transovum transmission and lateral transmission could reduce it very significantly. The introduction of explain the levels of infection found after the dry period. infective material into an area previously almost free of disease resulted in a significant rise in the rate of infection.

Clark, T. B. et al. (1965). A mosquito iridescent virus (MIV) from Aedes taeniorhynchus J. Invertebr. Pathol., 7: 519-521. (Wiedemann). Examination Diseased 4th-instar larvae of A. taeniorhynchus were collected in Florida. revealed a noninclusion virus with a dense central core surrounded by a 6-sided capsule The principal site of infection is the cytowhich is composed of at least 2 layers. Infected larvae became iridescent orange during the 4th-instar and plasm of fat cells. Per os transmission of MIV was sometimes appeared milky white late in the disease. achieved in only a small percentage of trials.

Cunningham, J. C. & Tinsley, T. W. (1968). A serological comparison of some iridescent J. Gen. Virol., 3: 1-8. nonoccluded insect viruses. Tipula iridescent virus and Sericesthis iridescent virus were found to be serologically They related by complement-fixation, tube-precipitation and agar-gel diffusion tests. were unrelated to mosquito iridescent virus when compared by complement fixation.

Dasgupta, B. (1968). Med. Hyg., 62: 730.

A possible virus disease of the malaria parasite.

Inclusion bodies have been observed in oocysts of Plasmodium. showed partial to total loss of nuclei.

Trans. Roy. Soc.

The abnormal oocysts

The intranuclear inclusions in the mid-gut of the larva Dasgupta, B. & Ray, H. N. (1957). Parasitology 47: 194-195. of Anopheles subpictus.

Feulgen-positive intranuclear inclusions were observed in the secretory cells of the midA mature inclusion body appeared as a lump of gut of larvae collected near Calcutta. Staining reactions suggested that the amount of DNA in DNA formed of globular bodies. the nucleus of other cells in the neighbourhood of the affected nucleus was reduced considerably.

40 Microbial infections associated with plasmodial development in Davies, E. E. et al. (1971). Anopheles stephensi. Ann. Trop. Med. Parasitol., 63: 403-408. Two forms of virus-like particles and a rickettsia-like organism have been observed in the midgut epithelial cells of a laboratory colony of A. stephensi. One of the viral particles does not appear to affect either the mosquito or the plasmodial infection. The other occurs in oocysts which show evidence of degenerative changes.

Day, M. F. & Mercer, E. H. (1964). Properties of an iridescent virus from the beetle, Sericesthis purinosa. Aust. J. Biol. Sci., 17: 892-902.

Sericesthis iridescent virus (SIV) was added to the medium in which second instar larvae of Aedes aegypti were developing. After two weeks the fat body in two living larvae became iridescent blue indicating infection by SIV. Nucleic acids in the blue-green and orange mosquito iridescent Faust, R. M. et al. (1968). viruses (MIV) isolated from larvae of Aedes taeniorhynchus. J. Invertebr. Pathol., 10: 160. Analysis of the nucleic acids revealed that DNA was present in both blue-green and orange MIV and that RNA was absent. DNA constituted 10.5% of the blue-green form and 11.7% of the orange.

Federici, B. A. (1970).

Unpublished observations.

Federici, B. A. (1973). Preliminary studies on a cytoplasmic polyhedrosis virus of Aedes Abstracts of papers, 5th Int. Collog. Insect Pathol. Microbial Cont. taeniorhynchus. (Oxford), p. 34. The 4th-instar larva of A. taeniorhynchus from Louisiana was found infected with a cytoInfection plasmic polyhedrosis virus (CPV) in the posterior portion of the stomach. trials gave an average rate of patent infection of 4.3% and mortality rate less than 1%. Occlusion bodies were present in the posterior stomach and occasionally in the gastric The coalescing process during occlusion formation is similar to that observed caeca. in a baculovirus in larvae of A. triseriatus. This similarity suggests that the mosquito host plays a role in the formation of occlusions. Virus pathogens of mosquitoes and their potential use in mosquito Federici, B. A. (1974). In: Aubin, A. et al., ed. Le controle des moustiques/Mosquito control. control. Quebec, Univ. Quebec Press, pp. 93-135. Review article with 56 references. Formation of virion-occluding proteinic Federici, B. A. & Anthony, D. W. (1972). a baculovirus of Aedes triseriatus. J. Invertebr. Pathol., 20: 129-138.

spindles in

An unusual process of inclusion formation was studied in A. triseriatus larvae infected with a Baculovirus (BV) similar to the nuclear polyhedrosis virus (NPV) type. In this disease virion-occluding proteinic inclusions initially developed individually. However, as the disease progressed the proteinic inclusions gradually coalesced eventually forming large rugose ellipsoids and finally, large smooth-surfaced spindles. Nuclei in late stages of infection usually contained two to five rugose ellipsoidal inclusions, frequently measuring 5 pm to 7 pm in diameter by 10 pm to 15 pm in length. The ellipsoidal forms exhibited different chemical behaviour from the spindles.

41 Studies on the pathology of a baculovirus in Federici, B. A. & Lowe, R. E. (1972). Aedes triseriatus. J. Invertebr. Pathol., 20: 14-21. The virus infected The pathology of a Baculovirus (BV) in A. triseriatus was studied. The the cardia, gastric caeca, and the entire stomach of larval midgut epithelium. progress of the disease was similar to that of other Baculoviruses of the nuclear polyThe disease differed from other BVs of the NPV type in that hedrosis virus (NPV) type. small proteinic inclusions gradually coalesced as they grew, forming large fusiform inclusions. Per os transmission of Chilo iridescent virus to mosquitoes. Fukuda, T. (1971). J. Invertebr. Pathol., 18: 152-153.

Chilo iridescent virus (CIV) was successfully transmitted to 13 species of mosquitos, The CIV in mosquito larvae was however the percentage contracting the disease was low. first detected at the time of 3rd-instar when the blue-violet iridescence began to show. The virus had a wide host range in mosquitos Death usually occurred in the 4th-instar. and can be more easily mass-produced than MIV. Pathology of a mosquito iridescent virus (MIV) infecting Hall, D. W. & Anthony, D. W. (1971). Aedes taeniorhynchus. J. Invertebr. Pathol., 18: 61-69.

Cells of the RMIV was capable of infecting a variety of tissues within its host. body, tracheal epithelium, imaginal discs, and epidermis were the primary sites of Extensive destruction of the fat body by this virus resulted in the replication. of most infected mosquitos before they reached the adult stage. The transovarial mission of RMIV was confirmed, and when transovarial transmission occurred, either or none of the progeny of a given female were infected.

fat viral death transall

A Baculovirus from the mosquito Wyeomyia smithii. Hall, D. W. & Fish, D. D. (1974). J. Invertebr. Pathol., 23: 383-388. A Baculovirus was found in larvae of W. smithii collected from a sphagnum bog in This virus is similar in size and appearance to Baculoviruses from Massachusetts. A unique feature of the virus is the formation of polymorphic other mosquitos. The prepatent period for this virus is 3-5 days. inclusions.

Hall, D. W. & Lowe, R. E. (1971). virus (MIV). Mosquito News, 31:

A new distribution record for the mosquito iridescent

448-449.

Aedes taeniorhynchus larvae collected off the west coast of Florida were found to be The location About 0.12% of the specimens examined were infected. infected with RMIV. of these collections, in addition to those previously reported, suggests that RMIV is TMIV has not been found in Florida. present throughout the range of A. taeniorhynchus.

Hall, D. W. & Lowe, R. E. (1972).

Physical and serological comparisons of "R" and "T" strains of mosquito iridescent virus from Aedes taeniorhynchus. J. Invertebr. Pathol., 19:

317-324. Gel diffusion studies with alkaline degraded virus preparations exhibited four antigens Electron common to both viruses but no unique antigens were detected for either virus. micrographs of infected tissue sections showed tubular structures associated with both RMIV and TMIV. RMIV and TMIV sedimented at different rates in sucrose density gradients and RMIV was found to be slightly more dense than TMIV by equilibrium ultracentrifugation in cesium chloride. Mixtures of the two viruses can be separated by sucrose density gradient centrifugation.

42 Hasan, S. et al. (1970). Infection a virus irisant dans une population naturelle d'Aedes detritus Haliday en France. Ann. Zool. Ecol. anim., 2: 295-299 (English summary).

An iridescent virus has been isolated from a naturally infected population of A. detritus in southern France. Morphologically and serologically the virus is identical to the iridescent virus described from this mosquito in Tunisia. (See Vago, Rioux, Duthoit & Dedet, 1969.) Infection of Aedes detritus Hal. with mosquito iridescent virus. Hasan, S. et al. (1971). Bull. World Health Organ., 45(2): 268-269. Attempts were made to infect larvae of A. detritus collected in France with MIV. Some larvae were kept for 24 hrs in water in which infected larvae had been macerated; others were infected with a virus suspension. In both cases, only 4th-instars showed disease symptoms, which appeared after 10 days for those injected and 20 days for those in the suspension. Infected larvae died within 2 weeks of appearance of symptoms.

Hazard, E. I. (1972).

Personal communication.

Kellen, W. R. et al. (1963). A possible polyhedrosis in Culex tarsalis Coquillet (Diptera: Culicidae). J. Insect Pathol., 5: 98-103. Tetragonal intranuclear inclusion bodies have been observed in larvae collected from stagnant ponds in California. Infected larvae succumb in the 4th-instar, and successful transmissions of the disease agent have been obtained in the laboratory. It was concluded that an infectious agent is definitely involved, and that it might be a virus.

Kellen, W. R. et al. (1966). A cytoplasmic-polyhedrosis virus of Culex tarsalis (Diptera: J. Invertebr. Pathol., 8: 390-394. Culicidae). The tetragonal crystals described by Kellen, Clark & Lindegren (1963) have been confirmed as inclusion bodies of a cytoplasmic-polyhedrosis virus* upon further examination. Polyhedra were present in hypodermal cells, in the developing leg, wing, and antennal This is the first cytoplasmic polyhedrosis reported which is not restricted to buds. gut tissue.

Kelly, D. C. & Robertson, J. S. (1973). J. Gen. Virol., 20(Suppl.): 17-41.

Icosahedral cytoplasmic deoxyriboviruses.

A comprehensive catalogue of icosahedral cytoplasmic deoxyriboviruses (ICDV) isolated from animals and plants is given. The list includes 7 from mosquitos.

Lebedeva, 0. P.

et al.

(1973).

Investigation of

in a laboratory culture of Aedes aegypti.

a virus disease of the densonucleosis type Acta Virol., 17: 253-256.

Histological changes in infected larvae were observed by light and electron microscopy. The virus The most obvious pathological changes were in the cells of the fat body. particles were about 200 A in size, having a paraspherical -shape and polygonal outlines. Observations revealed a similarity between the viral infection found in Aedes aegypti larvae and densonucleosis of Galleria mellonella.

See footnote to Chapman et al., 1970.

43 Virus-like formations in larvae of Aedes and Lebedeva, 0. P. & Zelenko, A. P. (1972). Culex mosquitoes. Med. Parazitol. Parazit. Bolenzi., 41: 490-492 (Russian, with English summary). in the cytoplasm of fat body cells were found in 4th-instar larvae and and Culex pipiens molestus from laboratory colonies and in some Culex sp. from natural water bodies. The inclusions resembled in size, shape, and staining properties those of Entomopoxvirus described from other insects. Also, the diseased mosquitos exhibited the hypertrophied nuclei structurally similar to those described in densonucleosis of Galleria mellonella. (See Vago, 1963, and Vago et al., Inclusion bodies

aegypti

pupae of Aedes

1964.) Linley, J. R. & Nielsen, H. Aedes taeniorhynchus. I.

T. (1968a). Transmission of a mosquito iridescent virus in 7-16. J. Invertebr. Pathol., 12: Laboratory experiments.

can be effectively

The virus

transmitted to the larvae

per os;

transovarial transmission

female to her eggs, and evidence was obtained that all, or a very high proportion, of the eggs are infected. Larvae infected per os in their early instars may develop signs and symptoms of disease, and die before pupation. Exposure to infection progressively later in their development results in later development of signs and symptoms, and a reduction in the total number of larvae that show disease before pupation. Rearing larvae under different conditions of temperature, diet, and crowding produced no detectable differences in infection rates among either groups of larvae exposed from the time of hatching to the same dosage of virus, or groups of larvae hatched from a large batch of eggs in which a proportion carrying infection had been from an infected

occurs

homogeneously distributed. Linley,

R.

J.

Nielsen, H. T. (1968b). II. Experiments

&

taeniorhynchus.

Aedes

Pathol., 12:

Transmission of a mosquito iridescent virus in J. Invertebr. related to transmission in nature.

17-24.

showed that infection could be acquired per os by larvae exposed under and that when healthy 4th-instar larvae were given access to intact diseased cadavers for a short time before pupation, they became infected and produced adults that laid infected eggs. Transovarial transmission of virus gives rise to diseased larvae, which die in the 4th-instar. The cadavers so formed provide the source of new infection, which is acquired per os by healthy larvae just before pupation. Infected adults from these larvae complete the cycle by depositing infected eggs.

A

field experiment

natural conditions,

Lowe,

R.

E.

et al.

(1970).

iridescent viruses.

Proc.

Comparison of the mosquito iridescent virus (MIV) with other Collog. Insect Pathol., pp. 163-170.

4th Int.

R isolate of the mosquito iridescent virus, although closely related to other insect iridescent viruses, possesses distinct physico-chemical characteristics. The diameter of RMIV (195 nm), molecular weight (2.486 x daltons), sedimentation coefficient (4.458), and DNA content (15.977%) are greater than the respective values for similar

The

109

Although biologically

viruses.

the two colour Matta,

J.

F.

(1970).

similar, and showing

no

unique antigens serologically,

isolates of MIV differ in size, sedimentation rate, and density. The characterization of a mosquito

chemical characterization.

J. Invertebr. Pathol., 16:

iridescent virus. 157-164.

II.

Physico-

for the purification of the R type of MIV is presented. The RMIV was distinctly different from the other iridescent viruses in terms of several physical parameters, and it would be difficult to justify considering it as a strain of these viruses. There was, however, a similarity in amino acid composition, indicating that

A procedure

they have

a common phylogeny.

44 A differential staining technique for a mosquito Matta, J. F. & Lowe, R. E. (1969). J. Invertebr. Pathol., 13: 457-458. iridescent virus. A staining technique, that allows routine screening for MIV infections using a light microscope, was devised.

The characterization of a mosquito iridescent virus (MIV). Matta, J. F. & Lowe, R. E. (1970). J. Invertebr. Pathol., 16: I. Biological characteristics, infectivity, and pathology. 38-41.

Infected Aedes taeniorhynchus were mass-produced by rearing them in dishes containing The average mortality was dependent on crowding homogenized infected 4th-instar larvae. The increase in virus over the inoculum varied between of the larvae during infection. The RMIV infects only the fat body and imaginal discs in A. taeniorhynchus 70 and 310%. and destruction of these tissues is usually complete. Evidence of two inapparent nonoccluded viral infections of Richardson, J. et al. (1974). Culex tarsalis. J. Invertebr. Pathol., 23: 213-224. C. tarsalis from a laboratory colony in California were found to be infected with 2 Both viruses were seen in thin sectioned material noninclusion cytoplasmic viruses. C. tarsalis from salivary glands, fat body, and nervous tissue of infected mosquitos. virus 1 (CTV 1) was retained through 6 serial passages by needle inoculations of infected Both CTV1 and CTV2 appeared to multiply in mosquito suspensions into virus-free adults. the hosts. Infection may cause degeneration of salivary glands in adult females.

The ecology of the immature stages of Aedes detritus (Diptera: J. Appl. Ecol., 5: 613-630.

Service, M. W. (1968).

Culicidae).

Large numbers of larvae collected on Brownsea Island were infected by various pathogens including non-inclusion iridescent viruses.

The structure of icosahedral Stoltz, D. B. (1971). J. Ultrastruct. Res., 37: 219-239.

cytoplasmic deoxyriboviruses.

The presence of two unit membranes in mosquito iridescent virus particles is clearly The unit membranes in this study are defined on the basis of morphology illustrated. (See Stoltz, 1973.) alone. The structure of icosahedral cytoplasmic deoxyriboviruses. Stoltz, D. B. (1973). J. Ultrastruct. Res., 43: 58-74. alternative model.

II.

An

The available evidence seems to indicate that ICDV particles contain only a single structural unit membrane, associated with the viral nucleoid.

Stoltz, D. B. et al. (1974). J. Microscopie, 19: 109-112.

Virus-like particles in the mosquito Culex salinarius.

Larvae of C. salinarius collected in Louisiana were found to have identical symptoms as The causative those described in Culex tarsalis by Kellen, Clark & Lindegren (1966). Although electron microscopic examinaagent is readily transmissible to C. tarsalis. tion revealed virus-like particles, the symptoms are not those expected from a typical

cytoplasmic polyhedrosis virus.

45

Stoltz, D. B. & Summer, M. D. (1971). Pathway of infection of mosquito iridescent virus. I. Preliminary observations on the fate of ingested virus. J. Virology, 8: 900-909. MIV is ingested in large amounts by lst- and 2nd-instar Aedes taeniorhynchus larvae without causing a high rate of infection. Preliminary observations suggest that most, MIV and if not all, ingested particles are degraded shortly after entering the midgut. other virus particles were apparently unable to penetrate the peritrophic membrane; consequently none was observed inside, or in contact with, midgut epithelial cells.

Tinsley, T. W. & Kelly, D. C. (1970). An interim nomenclature system for the iridescent group of insect viruses. J. Invertebr. Pathol., 16: 470-472. In the new system each iridescent virus is given a type number so additional host citing would pose no problem. This system is suggested to be employed until a system of more permanent names can be assigned based on comparisons of physical, chemical, and biologica properties of the various isolates. An iridescent virus of Aedes cantans in Great Britain. Tinsley, T. W. et al. (1971). J. Invertebr. Pathol., 18: 427-428.

Larvae of A. cantans were collected from a pond in Kent, England, and found to be infected with a virus of the iridescent group. They developed a lime-green colour at an advanced stage of infection.

Infection spontanee a virus irisant dans une population d'Aedes Vago, C. et al. (1969). detritus (Hal., 1883) des environs de Tunis. Ann. Parasit. Hum. Comp., 44: 667-676

(English summary). A virus disease was observed in natural populations of A. detritus in Tunisia. Diseased larvae could be identified by their slower movements and milky white coloration. The purified virus has a cubical Extensive damage to the fat bodies was observed. It appears to be related to the iridescent symmetry and measured 180 nm in diameter. viruses by its morphology, its location, and the iridescence of affected tissue.

Biochemical and biophysical properties of two strains of Wagner, G. W. et al. (1973). mosquito iridescent virus. Virology, 52: 72-80.

Several differing biophysical properties of the two strains (RMIV and TMIV) were observed due to size differences. The two strains appeared to be antigenically identical but the percent protein, DNA, and lipid of the two were dissimilar. Other than size variation, no morphological differences between RMIV and TMIV could be detected.

Weiser, J. (1965). 33(2): 586-588.

A new virus infection of mosquito larvae.

Bull. World Health Organ.,

Infected Diseased Aedes annulipes and A. cantans were found in South-western Bohemia. In their normal individuals are opaque and opalescent, and are usually less motile. habitat, they appear to die before pupation. Nearly every tissue seems to be affected. In general Infection is not common, not more than 1% of the larvae are infected. appearance, the virus is similar to the Tipula iridescent virus.

Wildy, P. (1971).

Classification and nomenclature of viruses.

Monogr. Virol., 5:

1-81.

46 Laboratory studies with mosquito iridescent virus Woodard, D. B. & Chapman, H. C. (1968). J. Invertebr. Pathol., 11: 296-301. (MIV). RMIV and a blue MIV of Aedes taeniorhynchus were serially passed through 68 and 30 The average rate of infection generation, respectively, of larvae of A. taeniorhynchus. The maximum rate of infection was usually for RMIV was 167% and 21% for the blue MIV. reached when large numbers of early-instar larvae were exposed for 48 hours to substantial amounts of MIV material and the larvae were reared to the 4th stadium at 25°C. Attempts to transmit MIV to larvae of 9 species of mosquitos not known to be hosts were About 20%/ patent infection occurred in the larval successful only with A. sollicitans. progeny of adult A. taeniorhynchus derived from early 4th-instar larvae exposed to MIV. Such transovarial transmission was observed in the MIV of Psorophora ferox.