Genetics, Cytogenetics, and Evolution of Mosquitoes

GENETICS. CYTOGENETICS. AND EVOLUTION OF MOSQUITOES

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James B Kitzmiller Florida Medical Entomology laboratory. Vero Beach. Florida

I . Introduction . . . . . . . . . . I1. Formal Genetics . . . . . . . . A . Morphological Mutants . . . . . B. Enzymes . . . . . . . . . . C . Linkage and Crossing-Over . . . . D . Linkage- Group-Chromosome Correlations E . Sex Genetics . . . . . . . . . F. Susceptible and Refractory Strains . . G . Genetics of Autogeny . . . . . . H . Genetics of Insecticide Resistance . . I11. Cytogenetics . . . . . . . . . . A . Salivary Chromosomes . . . . . . B. Karyotype Studies . . . . . . . C . Gametogenesis . . . . . . . . D . Cytology . . . . . . . . . . IV . Evolution and Speciation . . . . . . A . Sibling Species and Gene Flow . . . B . Hybridization . . . . . . . . V . Cytoplasmic Incompatibility . . . . . VI . Genetic Manipulation . . . . . . . VII . Behavior Genetics . . . . . . . . VIII . Genetic Control . . . . . . . . . A . General . . . . . . . . . . B . International Symposia . . . . . C . Control Schemes . . . . . . . D . Laboratory Tesbs . . . . . . . E . Computer Models . . . . . . . F. Field Trials . . . . . . . . . I X . Biochemical Genetics . . . . . . . A . Pathways . . . . . . . . . B . Molecular Genetics . . . . . . . C . Biochemical Taxonomy . . . . . References . . . . . . . . . .

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I. Introduction

The starting point for this review is “Genetics of Insect Vectors of Disease” (Wright and Pal, 1967). This work, known in the trade as “THE” book, summarizes very well the work in mosquito genetics until that time. Because of the publication lag of that volume I have included in this review some papers with a publication date of 1965, and havc arbitrarily stopped with papers published through 1973. Undoubtedly, many papers which have a 1973 date are still in press as I write this. Some 1974 and 1975 papers are included. A nagging doubt is the percentage of papers covered or, better said, how many missed. I have tried several different methods of finding the pertinent literature: the usual bibliographic search, the Literature Citation Section in Mosquito News (of tremendous value), and a personal request to most of my colleagues in the field t o send bibliographies and reprints. The response to the last item has been particularly gratifying, and I hope that this review will be of some help to those who have so generously cooperated. I am sure that I have missed some papers-I hope not too many. I have read, in the original language, each paper cited except that I have had to rely upon abstracts, summaries, or partial translations for the Japanese papers. Since this subject was first reviewed (Kitzmiller, 1953) the number of papers has greatly increased although still relatively few laboratories are working in the field. Complete reviews of various aspects of mosquito genetics are given in the publication “Genetics of Insect Vectors of Disease,” Wright and Pal (1967), editors. Treated in detail are formal genetics (Kitzmiller and Mason, 1967; Laven, 1967c; Craig and Hickey, 1967b) ; cytogenetics (Kitzmiller, 1967; Rai, 1967b) ; evolution and speciation (Kitzmiller et al., 1967; Davidson et al., 1967; Laven, 1967d; McClelland, 1 9 6 7 ~ ) ; population genetics (Spielman and Kitzmiller, 1967) ; physiological genetics (Bender and Gaensslen, 1967) ; resistance (Brown, 1967c) ; behavior (Mattingly, 196713) ; susceptibility (Macdonald, 1967b), and genetic control (Knipling, 1967; Lachance, 1967). Since the publication of this book, some of these areas have been reviewed and brought up to date. The genetics of resistance (Georghiou, 1969; Brown and Pal, 1971) is most thoroughly treated, and the latter work gives a most helpful list (Brown and Pal, 1971, p. 446) of published reviews of different phases of insecticide resistance. Two synoptic reviews, one in Russian (Kiknadze, 1967) and one in Japanese (Kanda, 1971), serve as introductions

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to the cytogenetic aspects of mosquitoes, and a shorter review (Chow-

daiah et al., 1971) covers the cytogenetics of the oriental species. A comprehensive review of medical entomology by two of the giants in the field (Philip and Rozeboom, 1973) contains a brief but accurate synopsis of mosquito genetics. Condensed accounts of anopheline (Coluzzi and Kitzmiller, 1975), aedine (Rai and Hartberg, 1975), and Culez (Barr, 1975) genetics are contained in Volume 3 of King (1975), “A Survey of Genetics.”

II. Formal Genetics

True to past performance, most of the advances in formal genetics have been made in Culex and Aedes. Relatively few morphological mutants have been discovered in Anopheles, although irradiated laboratory populations and extensive field collections have been carefully screened. The analyses of the isozymes in Anopheles constitute an important exception to the general lack of anopheline markers; they will be treated in a separate section, below. A, MORPHOLOGICAL MUTANTS 1. Culex

Culex tritaeniorhynchus, thanks to the work of Baker and his colleagues, is probably, after Aedes aegypti, the best-known mosquito genetically (Baker and Aslamkhan, 1968; Baker et al., 1970; Baker and Sakai, 1972b, 1974). Their work is a remarkable example of what can be done in taming a wild species in a short time. One sex-linked recessive, golden (Baker, 1968), expresses itself in egg, larval, pupal, and adult stages, with uniform penetrance and complete expression. Another sexlinked recessive, white eye (Baker, 1969), also has uniform expression and complete penetrance, but females are sterile. Against the white background, Rabbani and Baker (1970) found an autosomal codominant mutant, red-spotted, phenotypically expressed as a red and white variegated eye. Rs/+ individuals may be distinguished from rs/rs but only in a w / w background. Also recessive and sex-linked (Baker and Sakai, 1973133, rose eye is an allele a t the white locus. Of considerable importance was the isolation of a sex-linked dominant, Delta wing (Baker and Sakai, 1972a), which, combined with the dominant marker, M , for sex and an extensive inversion (Baker et al., 1971a,b), permitted the establishment of a ClB type system (Sakai and Baker, 1972). Delta affects

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the angle of the wings and the shape of the eye. A melanotic mutant, ebony (Sakai et al., 1972a), is nonlethal (many melanotic mutants previously described in other species are lethal), autosomal, and recessive. Two additional autosomal recessives curved leg and straw (Baker and Sakai, 1973a) do not show linkage to markers of either group 1 or 2 and have therefore been placed in linkage group 3. Twenty-three mutants are described in Baker and Sakai (1974). Heat sensitive mutants were induced using ethyl methane sulfonate (Sakai and Baker, 1974). 2. Culex pipiens Complex

Barr and Narang (1972) have described a pupal mutant, enlarged tergum, autosomal, recessive, and with good penetrance. A sex-linked recessive, divided eye, is expressed as a clear band of variable width, devoid of facets, which separates an upper and lower portion of the adult eye (Barr, 1969). Two lethal characters, curved larval antenna and female lethal, have been studied by Barr and Myers (196613, 1971). The first is autosomal and recessive; the second appears to be complex and interesting, but a satisfactory genetic basis has not yet been worked out. The lethal factor can be transmitted by males by outcrossing and kills almost all females before hatching, producing “all-male” rafts. The occasional females produced from lethal rafts appear not to be contaminants (markers), and nine produced lethal rafts when bred with sibs. VandeHey (1967) described Black larva in Culex pipiens, which is inherited as a sex-linked dominant in males but shows incomplete dominance in females. Penetrance is good and, as a pleiotropic effect, developmental rate is decreased. VandeHey (1969) also lists a number of morphological variants obtained by inbreeding progeny of wild-caught females. Of 20 different phenotypes obtained, 3 were analyzed t o some degree. Shetty and Chowdaiah (1973) list 46 genetical and morphological variations, 100 gynandromorphs, 150 intersexes, and 2 mosaics from a 2-year observation of 6 field-collected strains of Culex fatiguns. One adult and five larval variants are stated to have been analyzed, although no data are given in this preliminary publication. In a long series of taxonomic and morphological papers (1961-19711, Ishii has described many features of Culex larvae. Several of these papers (Ishii, 1966, 1967a,b,c, 1969a,b, 1970a,b,c, 1971) have dealt with selection for siphonal hair tuft numbers. An index of 4-4, four tufts on each side of the siphon, is most frequent in natural populations. Selection for 4-4 phenotypes produced positive results up to about 85% by the F, generation, but other phenotypes also appeared. Similarly, selection for lower numbers of tufts, or higher numbers, never resulted in complete homozygosity, although selection was effective in both directions. Similar results

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were obtained in both pallens and molestus. Continued inbreeding resulted in poor viability and eventual loss of the population. The green larval and pupal color in C . fatigans was reinvestigated by Consoli (1972) with inconclusive results. A dominant mutant, Gold (McClelland, 1967b), has potential use in marker stocks. Gold shows recessive lethality, is not sex-linked. White eye was recovered from a C . fatigans population in Brazil (Rai, 1969a). A dark mutant in C . pipiens, “dunkel” (Dennhofer, 1973), affects especially the head capsule and siphon of the larvae. Autosomal and linked with ruby in group 2 (but see below, linkage), dunkel is not allelic with either me1 or Bl. Another dark mutant, black (Rahman et al., 1971), is an autosomal recessive in C . fatigans. In Culez tarsalis Barr and Myers (1966a) describe white eye and yellow larva. Recessive and fully penetrant, white is unusual in that it is evidently autosomal, not allelic with white of Culex pipiens. The mutant yellow is sex-linked and probably recessive, poorly expressed in males. Haeger and O’Meara (1970) describe methods for rapid incorporation of wild genotypes into laboratory populations. See also Umino (196513, 1965c, 1966). 3. Anopheles The description and isolation of morphological mutants in Anopheles continues to lag behind such studies in Aedes and Culex. There are abundant morphological variations, to be sure, in many species, but inbreeding or other genetic analyses seldom produce repeatable results. Aslamkhan et al. (1972a,b) and Aslamkhan (1973a,b,c) list about fifty morphological variants in Anopheles stephensi but give no data on crosses. Ten of these variants are reported as having been isolated in pure stocks, with experiments planned to study the modes of inheritance. A sex-linked recessive, white eye (Aslamkhan, 1973c) should be a valuable marker. A lethal, autosomal, partially dominant gene, black (Mason and Davidson, 1966), kills all preadult stages when homozygous. I n Anopheles atroparvus a red-eyed mutant or was obtained by Laudani et al. (1970) which proved to be autosomal and recessive. Larvae with a broad dorsal light stripe are often seen in laboratory and field populations of Anopheles. A few of these have been isolated and followed genetically. In A. albimanus Georghiou et al. (1967) selected white-striped individuals from the Haiti strain and concluded that the character is controlled by a single semidominant autosomal gene with variable expression in the heterozygote. Another common phenotype, black larval color is usually lethal as fourth-instar larvae or as pupae. Seawright and Anthony (1972) analyzed such a case in A . quadrimaculatus and concluded from offspring

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of normal sibs of black that it was recessive and autosomal. Brightfield and electron microscopy showed that the fat body and cuticular tissues contained spherical dark granules of variable size. More information is available on mutations in species A and B of the A. gambiae complex (Mason and Davidson, 1966; Service, 1970a; Mason, 1967). Wild-type larvae have a whitish patch of scales on the anterior part of the thorax, the collarless mutant, which is autosomal and recessive and uniformly colored. Another pigment mutant, Black Diamond, appears to be a single autosomal dominant, and red stripe is possibly a polygenic system. Two eye color mutants, pink and white, have been isolated, both being sex-linked recessives; white appears to affect the entire pigment system of the larva and is epistatic to both collarless and red stripe. Other mutants described by Mason and Davidson (1966) include white eye, a sex-linked recessive, green larva, an autosomal recessive in A . pharoensis, and nonstripe, an autosomal recessive in A . quadrimaculatus. Two albinoid females from Nigeria (Service, 1964) and four from Kenya van Someren, 1969) are worthy of note, although no eggs were obtained, and melanic variants (White and Davidson, 1972) have been reported.

4. Aedes To date about 100 mutants have been isolated and studicd in Aedes aegypti (Coker, 1967a; Craig and Hickey, 1967a,b). Most of these are morphological mutants, and most affect either color patterns or appendages. Many are useful as genetic markers and have been located on linkage maps. An excellent summary of work up to 1966 may be found in Craig and Hickey (1967b). Two genes affecting eye color, red and rust (McClelland, 1966), are recessive and partially sex-linked, constant in expression, and show complete penetrance. An especially interesting mutant, bronze (Bhalla and Craig, 1967), controls a pale tan or bronze color in the dark parts of the adult, pupa, larva, and egg shell. This sex-linked recessive has complete penetrance and uniform expressivity, fair viability. The males are fully fertile, but the females are sterile and the stock is maintained by bz/ X bz/bz crosses. Another sex-linked recessive, white (Bhalla, 1968b), with complete penetrance and good expressivity, blocks the ommochrome synthesis in compound eyes and ocelli. Dunn and Craig (1968) report still another sex-linked mutant, small antenna, which is recessive but shows partial penetrance and a wide range of expressivity, mostly dependent upon genetic background. Of considerable interest is a sex-linked, incompletely dominant gene Terminalia (Hartberg and Craig, 1973), which controls morphological features of the telomeres and

+

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basal lobes of the male genitalia. This gene is allelic in A . aegypti and in the closely related species A . mascarensis, between which crosses were made to study the inheritance of the morphological and taxonomic features of the male genitalia. Translocation experiments (Bhalla, 1973b) have localized black tarsi on the small terminal segment of chromosome 3, distal to the secondary constriction. I n A . albopictus a sex-linked, recessive, homeotic mutant proboscipediu (Quinn and Craig, 1971; Bat-Miriam and Craig, 1966) affects the labella and the maxillary palps. Ingenious feeding and rearing techniques permit the maintenance of a homozygous stock. Other mutants in A . albopictus include yellow, black palps, bulb, white proboscis, wart, and dark scutum (Bat-Miriam and Craig, 1966). An aberrant A . albopictus is described by Rudnick et al. (1971), and certain chemicals (Quraishi, 1967) will produce phenocopy-like variations. I n the salt-marsh mosquito A . taeniorhynchus, a mutant bleached-eye (O’Meara, 1975) is inherited as an autosomal recessive, but it also shows a pleiotropic lethal effect dependent upon the genotype of the mother. The pale scaling used to separate A . forrnosus, A . queenslandensis, and the type form A . aegypti (Hartberg, 1969) is probably a polymorphism. Craig and Hickey (1966) did a quantitative analysis of field populations in Africa and found a load of 0.52, 0.72, and 0.84 mutations per mosquito in three feral African populations, but loads of 2.72 and 2.96 in two domestic populations. The presence of suitable markers in A . aegypti has made possible field studies on dispersion, field mating, movement, and recovery of released individuals (Bond et al., 1970; Fay and Craig, 1969; Hausermann et al., 1971).

B. ENZYMES 1. Anopheles

One of the most significant advances in mosquito genetics in recent years has been in the electrophoretic studies of enzymes. Starch gel and acrylamide gel techniques have permitted the study of biochemical mutants in individuals, in species, and in populations. As in Drosophila, these studies have uncovered a large amount of polymorphism. Perhaps the most welcome result has been in the anophelines, where the presence of abundant biochemical alleles has provided for the first time the markers necessary for linkage group and mapping studies. Morphological markers are still rare in the anophelines. An excellent review (Bullini and Coluzzi, 1973, 1974) includes much original work especially on the Pgm alleles. Pioneering the enzyme work was Bianchi in Cagliari, who was quick to adapt the techniques developed in Drosophila to the anophelines. The

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first work with Anopheles atroparvus (Bianchi, 1965, 1966a) indicated that zymograms for alkaline phosphatase activity showed clear differences in numbers, intensities, and relative mobilities in eggs, different larval instars, pupae, and adults. This work was quickly extended (Bianchi, 1966b) to the other Palearctic members of the Anopheles maculipennis complex-labranchiae, subalpinus, maculipennis, and messeae-and it was demonstrated that all members of the complex had unique enzymatic patterns and that each species could be distinguished from most (but not all) of the others. Especially important was the fact that A. atroparvus and A. labranchiae were clearly separable, although the morphology of the larvae and adults, and even the banding patterns of the salivary gland chromosomes, are identical. This difference was exploited (Bianchi, 1968a,b,d) to demonstrate clearly that an allele for a fast moving alkaline phosphatase band was homozygous in A . Zabranchiae, A. atroparvus was homozygous for the slow moving allele, and hybrids in both directions showed bands of intermediate mobility. The data further indicate that the structural genes in the two species are homologous but not identical, differing in polypeptide subunits, and that the intermediate band of the hybrids is probably a dimer. Extending the work to A. stephensi, Bianchi (1968a, 1969) showed that in a cross between two strains that had been selected for susceptibility and resistance to infection by Plasmodium gallinaceum, each strain was homozygous for an esterase. Each strain showed a different level of electrophoretic mobility for this esterase, but F, hybrids showed two bands, one from each of the parents. F, X F, crosses produced three kinds of individuals in a 1:2:1 ratio, indicating that the Est-1 locus in these crosses in A. stephensi is under the control of a single pair of alleles. Comparing alkaline phosphatase patterns in a European species, A . labranchiae, and a North American species, A. freeborni, Bianchi and Pirodda (1968) found that these two species are clearly separable and futhermore that F, hybrid individuals show two distinct bands, indicating that molecular hybridization does not take place between these species, although it does between A. labranchiae and A. atroparvzls. Isoelectric focusing (Bianchi, 1970) did not give better results than a continuous pH system. A second system, Esterase-6, was investigated in A. atroparvus (Bianchi and Rinaldi, 1969, 1970). Four alleles, one fast, one slow, and two intermediate, produced the 10 expected phenotypes. A random sample of 134 individuals from the population gave reasonably good Rardy-Weinberg frequencies. Tissue-specific glucose-6-phosphate dehydrogenases (Bianchi and Rinaldi, 1974) are present. Also in A. atroparvus the three electrophoretic variants of xanthine dehydrogenase (Bianchi and Chessa, 1970a,b) form six phenotypes. Random samples from the stock cage were in reasonable

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accord with Hardy-Weinberg expectations. Some mechanism, as yet undetected, interferes with the expression of these alleles in adult males. Several enzymes in Anopheles punctipennis (Narang and Kitzmiller 1971a,b,c, 1972, 1973a,b) have been studied by acrylamide gel electrophoresis. Six esterases, A, B, C, D, E, and F, vary in substrate specificity, in mobility, and in staining intensity. In the A-B system (Narang and Kitzmiller, 1971a) extensive crosses and analysis showed that the eight dark bands of the A-B system are the result of seven alleles at the A locus and seven alleles at the B locus. Both loci are autosomal. The data indicate that null alleles are common in the population and that a large amount of enzyme polymorphism may be present in a relatively small population. The C system (Narang and Kitzmiller, 1971b) is specific for beta substrate, easily recognizable, autosomal and should be an excellent marker. There are three alleles a t a single locus, distinguished by mobility. There appear to be no null alleles in this system, in contrast to the A and B systems. The E system (Narang and Kitzmiller, 1973a) is also highly polymorphic with seven different alleles at a single autosomal locus. The F system (Narang and Kitzmiller, 1973b) contains both alpha and beta substrate esterase bands, is also autosomal, also has seven alleles a t a single locus. These bands were studied with a thick gel technique which gave better resolution and less interference. The D system has not yet been analyzed genetically. The dehydrogenase systems (Narang and Kitzmiller, 1972) indicate two autosomal loci for xanthine dehydrogenase, one with three alleles, the other with two. Octanol dehydrogenase is probably under the control of a single locus although the tetrazolium oxidase activity makes definitive analysis uncertain. Adults and larvae differ in xanthine dehydrogenase activity sites. Booth et al. (1971, 1973) studied the effects of selected inhibitors on the A, B, C, E, and F loci. The A and B isozymes are probably cholinesterases, since they are inhibited by carbarnates and organophosphates. The C locus is probably a carboxylesterase; and the E and F loci, aromatic esterases. Bullini, Coluzzi, and co-workers studied phosphoglucomutase activity and polymorphism in 21 species of Anopheles; they have summarized this work in an excellent review of gene-enzyme systems in mosquitoes (Bullini and Coluzzi, 1973, 1974). I n anophelines each phosphoglucomutase allele appears to determine a one-band electrophoretic pattern, while in culicines each phosphoglucomutase allele appears to determine a twoband phenotype. The more significant data are summarized here, but consult Bullini and Coluzzi’s paper for details and for additional species. Anopheles plumbeus: four codominant alleles, in spite of low population densities of this tree-hole breeding species. Anopheles claviger and petra-

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gnanii: three codominant alleles, with PgmB most frequent in both populations. Anopheles maculipennis complex : five species (maculipennis, labranchiae, atroparvus, melanoon, and sacharoui) and eleven different geographical locations yielded six different alleles. One allele PgmAl is the most frequent or the only one present in all samples except in A . sacharoui, where the PgmAz allele is most common. Anopheles gambiae: sixteen samples including all six species showed four codominant alleles with P g d most frequent except in A . merus. Anopheles stephensi, superpictus, and rujipes: I n this closely related group of species, six codominant alleles, five in A . stephensi and a different one in A . superpictus. I n A . rufipes, the two alleles are the same as two in A . stephensi. An alcohol dehydrogenase in Pakistani strains of stephensi (Iqbal et al., 1973b,d) is expressed as three phenotypes, resulting from a single autosomal locus with codominant alleles. The same authors (Iqbal et al., 1973c,d) report two additional enzymes in A. stephensi; an esterase and an acid phosphatase. All three are controlled by autosomal codominant alleles. The data indicate linkage between Adh and Acph (21.8%0), the long awaited beginning of mapping studies using enzyme markers. Other than the Pgm analyses, surprisingly little work has been done on isozyme analysis in Anopheles gambiae. In species A (Coker, 1973) a beginning has been made with the demonstration of esterases, malate dehydrogenase, lactate dehydrogenase, alkaline phosphatase, and leucine aminopeptidase. Alcohol dehydrogenase was not present. Of special promise is the initiation of the isozyme approach in Anopheles albimanus (Ved Brat and Whitt, 1974a,b). L-Lactate dehydrogenase and glycerol-3-phosphate dehydrogenase specific activities were determined during larval, pupal, and adult stages. Lactate dehydrogenase activity is initially about 100 times greater than that of glycerol-3-phosphate dehydrogenase, reaches a peak in fourth instar, undergoes a sharp drop during pupation but an increase in the adult. Glycerol-3-phosphate dehydrogenase stays low during larval stages but rises sharply in the adult. Four electrophoretically distinct esterase systems in Anopheles albimanw have been assigned to separate loci (Ved Brat and Whitt, 1974a). Test crosses reveal that all four loci are on the same chromosome with the tentative sequence and map distances as follows: A-14-B-22-C-ll-D. 2. Culicines

Trebatoski and Haynes (1969) initiated the work in the culicines. Ten species of Aedes, one Culex, and one Anopheles were analyzed for esterases, phosphatases, and dehydrogenases. All species examined showed specific enzyme patterns. No genetic analyses are given, but some inter-

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esting taxonomic and evolutionary speculations based upon the data. are presented. Not unexpectedly, further investigations focused on Aedes aegypti with the description of the first gene-enzyme in that species (Trebatoski and Craig, 1969). An esterase designated Esterase 6 differed in two inbred lines. Hybrids between these strains showed both bands, and the results are consistent with the hypothesis that the two Est 6 bands are controlled by a single pair of codominant alleles a t a single autosomal locus on chromosome 2. Crosses with another inbred line showed a third allele with intermediate mobility. The strains used had been inbred for 9 and 21 generations, which may explain the lack of polymorphism. Nonspecific esterase activity was also demonstrated in the Liverpool strain of A . aegypti (Townson, 1969a,b, 1971a, 1972) in which slow, fast, and heterozygous phenotypes were found, as well as evidently a null phenotype. Freyvogel et al. (1968) screened fourteen strains and species of three genera for nonspecific esterases, finding that most of the esterases were species and strain specific, that there was sexual dimorphism in Anopheles stephensi, and that there was differential distribution in different organs. Species specificity of protein bands was also demonstrated by Warren and Breland (1969). This work was extended in Aedes vittatus (Freyvogel and McClelland, 1969) to confirm the previously reported strain differences in esterases, but gave different results in the alkaline phosphatase zymograms from those of Bianchi (1968d) , in that the intensities of the zymogram bands in the hybrids resemble their mothers and are not intermediate, as shown by Bianchi. Briegel and Freyvogel (1971) and Freyvogel and Briegel (1971) compare nonspecific esterases during all developmental stages of Aedes aegypti, Culex pipiens fatigans, and Culex pipiens pipiens. The distribution and activity levels of the esterases is different for all three species as well as for different developmental stages. Zymogram patterns of the 4 larval instars were more uniform, although differences in intensities may be seen in larvae of different ages. Similar preliminary data are reported for AmLGeres subalbatus (Desowitz, 1969). The pattern of enzymatic activity changes during the life of female Aedes aegypti (Briegel, 1972). Females of two strains were compared for up to 50 days of age, and were also examined before, during, and after blood meals. The enzyme activity between strains was relatively uniform, but notable changes before, during, and after blood meals were evident. This work was continued (Briegel and Freyvogel, 1973) to demonstrate specific patterns of esterase isozymes in several organs of female A . aegypti. A thorough review of most of the isozyme work on the culicines (Bul-

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lini and Coluzzi, 1973, 1974) indicates that polymorphism is common and widespread. Most of the Pgm alleles in the culicines determine twoband phenotypic patterns (in contrast to the one allele-one band pattern in the anophelines). Some 22 species and 77 populations have been analyzed for Pgm alleles; an extensive review of other isozymes is included in this valuable summary. I n Aedes aegypti a t least 7 codominant Pgm alleles have been found in 19 different populations. All are in linkage group 2. Although the strains are of widely diverse geographic origin, the PgmAl allele is most frequent in 17 of the 19 populations tested. Some populations are monomorphic, several have 3 alleles, and 2 populations show 5 alleles (Bullini et al., 1970a,b, 1971e, 1972a,c, 1973a,b). Other species of Aedes also exhibit widespread polymorphism of Pgm alleles. Aedes caspius has five different Pgm alleles, A . pulchritarsis has three, and A . berlandi, two. Similarly the related species Aedes cataphylla, pulZatus, and d e t d m show 4, 2, and 5 alleles, respectively. Pgm samples have also been taken in Aedes rusticus, refiki, and excrucians. (For details see Bullini and Coluzzi, 1974.) Widespread polymorphism for Pgm alleles is also found in Culex pipiens. Best studied are eight Italian populations which had from 3 t o 6 Pgm alleles; two populations had all six (Bullini et al., 1971b; Bullini and Coluzzi, 1974). The Pgm alleles are remarkably stable in frequencies even in populations that differ considerably ecologically (Bullini et al., 1972b), a property suggesting that some selective force, not related t o the ecological conditions studied, is responsible. Data are also available (Bullini and Coluzzi, 1974) for Culex hortensis, Culex mimeticus, Culiseta Zongiareolata, Culiseta annulata, and Orthopodomyia pulchripalpis. 3. Culex

Surprisingly few papers have been published on the enzymes in Culex. Several of the papers quoted above have included reports of nonspecific esterases in Culex, but only the Pgm alleles (Bullini and Coluzzi, 1973, 1974) have received detailed genetic analyses. A developmental and genetic study of C . fatigans (Simon, 1969) confirmed the developmental picture hnd sharp differences in esterases between stages, but little variation within stages. At one of four esterase zones designated esterase E, the electrophoretic activity is determined by a pair of codominant autosoma1 alleles. Homozygotes had one band only, but heterozygotes had two bands, suggesting a dimer. Partial characterization indicated that this is probably an aliesterase. Similar studies in C. fatigans (Garnett and French, 1971) show 13 different zones of esterase activity in adult

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populations. Two of these were analyzed genetically and again found t o be the result of a pair of codominant autosomal alleles, and characterization again indicates that these isozymes are aliesterases. Zymograms of C. p. molestus (Odintsov et aZ., 1971) showed 4 esterases in nerve tissue and 9 in whole-body homogenates. Characterizations suggested one arylesterase, the others being carboxylesterases and cholinesterases. I n C. fatigam, malate dehydrogenase (Narang and Narang, 1973) shows from 9 bands in pupae to 14 in adults. Again, uniformity within developmental stages is observed. The high number of malate dehydrogenase bands is in contrast to other loci studied in Culex, in which 3 or 4 isozymes only have been reported. (See also Schumann, 1974a.) I n Culex tritaeniorhynchw five esterase alleles (Iqbal et al., 1973a) a t the E-4 locus are codominant autosomal alleles, as are two alkaline phosphatase alleles (Sakai et al., 1973). The esterase and alkaline phosphatase loci are on different autosomes. A series of five alcohol dehydrogenase variants (Baker and Sakai, 1974; Sakai et aZ., 1973b; Iqbal et al., 1973b,d) behave as codominant alleles a t a single locus and map between cZ and s on linkage group 3. Four of the five alleles have been found in natural populations in the vicinity of Lahore. The active enzyme is probably a dimer (Iqbal et aZ., 1973b). Two amylase alleles, fast and slow (Baker and Sakai, 1974), are sex-linked and do not form a hybrid band in amy-4/amy-6 combinations. A potentially powerful tool for analysis of isozymes has been reported by Narang and Narang (1974). This method involves isoelectric focusing a t neutral isoelectric points and permits separation of the multiple components of the heavily stained “blobs” often seen on acrylamide gels. Mosquitoes used were Culex fatigans and Anopheles evansae.

4. Use of Enzyme Markers Isozyme markers have been skillfully utilized by Mario Coluzzi and co-workers. In the Aedes mariae complex, the three sibling species m a k e , zammitii, and phoeniciae are kept apart by sterility barriers in both sexes. F, individuals are produced in all crosses. The M x Z or Z x M crosses give sterile males and fertile females; P X M, M X P, P X Z, or Z X P crosses produce sterile males and a very few fertile females (Coluzzi and Sabatini, 1968e; Coluzzi et aZ., 1970b). Phosphoglucomutase electrophoretic patterns were found to be monomorphic in laboratory colonies of both A . mariae and A . zammitii. Crosses between the two species (Coluzzi et al., 1971a,b) show that Pgm is determined by a pair of codominant alleles PgmA and PgmB. The heterozygotes show both bands. These data were elegantly used (Coluzzi and Bullini, 1971) t o argue for the existence of precopulatory isolating mechanisms. I n the lab-

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oratory these species will hybridize, the hybrids are heterozygous for the Pgm alleles, and the F1 males are sterile. Is this postcopulatory hybrid sterility the principal isolating mechanism, or is some precopulatory mechanism operating as well? About 25,000 A . zammitii were released in an A . mariae zone in which the density of A . mariae had previously been lowered by collecting and destroying A. mariae larvae from the isolated rock pools in which they breed. Samples from July 15 through October 11 show many A and B homozygotes (ca. 1400 A, 1000 B ) but only 10 heterozygotes, strong evidence for the existence of an efficient precopulatory mechanism. In the third member of the complex, A . phoeniciae, populations are polymorphic for two alleles PgmR and PgmC, the latter existing with a gene frequency of about 92%. Crosses among all three species produced the expected kinds of hybrid allozyme patterns. I n population surveys (Bullini and Coluzzi, 1972b) five alleles have been found in the complex: A,, A?, Aa, B, and C. I n A . mariae populations the B allele is always the most prevalent, in A . zammitii the A, allele predominates, and in A . phoeniciae, C. This argues for some sort of balanced selection rather than random drift of neutral isoalleles (Kimura, 1968). In the same paper Bullini and Coluzzi (197213) present similar evidence from A . aegypti in which the PgmAI allele is always the most frequent of the seven known alleles a t the Pgm locus. The enzyme markers will probably be of considerable value in studies aimed a t genetic control (Bullini and Colluzi, 1972a). C. LINKAGEAND CROSSING-OVER 1 . Aedes aegypti

Excellent linkage maps have already been published for Aedes aegypti (Craig and Hickey 1967a,b). Work reported here has been published since 1965, if not already included in those maps. a. Sex-Linked Loci. The position of loci on the sex chromosome began with the work of McClelland (1966), who showed that two eye color mutants, rust and red, both showed linkage with the factor for sex. Both genes were recessive and showed good penetrance. Crossover values (although variable) averaged : red-sex, 6.4% ; rust-sex, 26.4% ; w t - r e d , 22.176. The order was therefore rust, red, sex. Bronze (Bhalla and Craig, 1967) is about 3 crossover units from sex. White eye (Bhalla, 1968b) is easily recognizable in all stages, even in late embryos. Extensive crosses indicated sex-linkage with about 17.4 crossover units between white and sex. A sex-linked recessive, sma22 antenna (Dunn and Craig, 1968), with

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incomplete penetrance and reduced expressivity is about 10 crossover units from bronze in the direction of white. The evidence indicates that sma is probably very close to white. The available evidence for analysis of linkage data for chromosome I is well summarized by Bhalla and Craig (1970) and the order rust, red, sez, bronze, and white is confirmed. Between bronze and white, lie gray (Craig and Hickey, 19678) and sma, in that order. The exact location of other sex-linked genes is in doubt. Perhaps the greatest value of this paper is the generalization, previously reported by several authors and since confirmed many times in Aedes, Culez, and Anopheles, that crossover values are extremely variable, even with supposedly identical genetic backgrounds and controlled environmental conditions. Sex, age, and temperature all affect crossing over. The discrepancies may often be considerable. Some extremes are as follows:

rust-red red-sex sex-bronze bronze-while red-bronze red-filarial susceptibility

Maximum

Minimum

22 8.5 5 13.5 20 34

10 1 2 10 10 16

It is suggested that wherever possible only heterozygous males of known age should be used for crossover experiments. b. Linkage Groups S and S. Linkage group 2 is well known (Craig and Hickey, 1967b). The principal markers are DDT and Dieldrin (resistance), Gold, yellow, and spot. Studies with insecticide resistance often give puzzling results, and probably there will turn out to be several resistance loci of varying importance. A comparative study of DDT resistance (Wood, 1967a) indicates that a major gene for DDT resistance in larvae, R DDTl is on linkage group 2 about 12 units from yellow, with the tentative order as DDT-spot-yellow, an order that differs from previous studies of larval resistance (Klassen and Brown, 1964) but agrees with the order for adult resistance (Coker, 1966). Adult resistance is due, according to Wood, to another major gene, R DDTd on linkage group 3, but Coker (1966) suggests two resistance genes on chromosome 2, with linkage group 3 having no influence on adult resistance. The linkage relationships of dieldrin resistance and DDT resistance were reviewed by Lockhart et al. (1970). Using silver thorm, a recessive mutant introduced into the MYS strain of Aedes aegypti by introgressive hybridization from

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Aedes nmscarensis, and strains carrying the markers yellow, silver, spot, Gold, black palpi, and black tarsus, it was concluded that the genes for dieldrin and D D T resistance are some 45 units apart, rather than about 5 units as calculated previously (Klassen and Brown, 1964). The order in linkage group 2 was found to be (probably) Dieldrin-Silver-black pedicel-yellow-spot-DDT. The question of the Dieldrin-DDT linkage remains unresolved. Again different strains and different experiments showed variable crossover distances between the loci involved. The Esterase 6 locus (Trebatoski and Craig, 1969) is also on linkage group 2 and in crosses with spot and yellow shows linkage value of spot-Est 6,26.4% ; yellow-Est 6, 17.4%, with the order therefore spot-yellow-Est 6. The phosphoglucomutase locus (Bullini et al., 1972c) also is located on linkage group 2 about 18 units from yellow. Also in linkage group 2 is a locus for susceptibility to Plasmodium gal1inaceu.m (Kilama and Craig, 1969). This locus, pls, is between silver mesonoturn and Dieldrin resistance with the approximate distances Dl-17-pls-8-Si. The proboscipedia mutant in Aedes albopictus (Quinn and Craig, 1971) is linked with sex with about 20% crossing-over. Very little has been done with linkage group 3. Two genes, black tarsi and miniature, are on chromosome 3 (Ved Brat and Rail 1973a). Using translocations and pseudolinkage with sex, the data indicate location on opposite arms of this autosome, about 28 units apart. 2. Anopheles

In Anopheles the data remain scanty. The locus for the familiar stripe phenotype (Georghiou et al., 1967) was shown to be on the same linkage group as the locus for dieldrin resistance. Mason (1967) demonstrated sexlinkage of both white and pink, but gave no map distances. The most encouraging results are those of Iqbal et al. (1973d) working with three enzyme systems, all autosomal in Anopheles stephensi. The loci Acid phosphatase and Alcohol dehydrogenase are linked with about 14% recombination in females and 22% in males. Neither is linked to an esterase locus. A d h and Acph have tentatively been assigned to linkage group 2 and Est to linkage group 3. A preliminary report (Aslamkhan, 1973a) suggests in A . stephensi a white locus on the X chromosome, brown palpi, beaked proboscis, wart, and reduced antennae on linkage group 2, and 4th costal spot on linkage group 3. If detailed data support these observations, valuable information will have been obtained. Another preliminary report (Haridi, 1973) suggests that D D T , Dieldrin, and Diamond are on the same linkage group in A . gambiae. I n A. albimanus (Ved Brat and Whitt, 1974a) four esterase loci are on the same autosome. See Section 11,B, 1.

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3. Culex pipiens Complex Most of the data since 1967 are fragmentary and occur in C . p . fatigans, pipiens, or molestus, often in strains that have received genetic contributions from two or all three members of the complex. I n crosses with ruby and yellow marker strains, Barr and Myers (1966b) showed that the lethal mutant curved larval antenna was on linkage group 2. Yellow and ca are closely linked, both are about 20 units from ruby. The order is not yet known. Several investigations of insecticide resistance loci gave variable results. In Culex p . fatigans adults (Tadano and Brown, 1967) the dominant D D T resistance gene was found to be linked with linkage group 2 loci yellow and ruby, a t crossover distances of about 20 and 46 units, respectively. Dieldrin resistance showed linkage with the group 3 marker kps with about 3 5 4 0 % crossing-over. The relatively high crossover distances suggest verification with intermediate loci when and if available. The y-DDT values varied from 17.5% to 22.7% in different experiments, confirming the heterogeneity reported by many investigators (Sanders and Barr, 1966; Baker and Sakai, 1972a; McClelland, 1966). Tadano (1969a,b) claimed linkage with the group 2 mutant ruby for the loci for malathion resistance and fenthion resistance in C. p. pallens. These two genes are linked with about 12-15% crossing-over in the order ma-fe-ru. However, each shows about 48-50% crossing-over with ruby. Obviously more precise experiments are needed. Also in C. p . pallens, Tadano (1970b) showed that, in larvae, resistance t o Abate, Fenitrothion and Malathion were linked in group 2 and independent of white in group 1. He gives (Tadano, 1970b) recombination values as follows: AbateFenthion, 5.8-6.9% ; Fenthion and Fenitrothion, 0 . 2 4 4 % ; DieldrinAbate, 22-26%. Tadano and Sat0 (1970) give recombination values between 17 and 20% for Parathion and Malathion. Dorval and Brown (1970) analyze Fenthion resistance in C. p. fatigans. This single recessive factor is probably in linkage group 2, but with large recombination values, about 42% fe-y and also about 42% fe-ru. Although not certain, the probable order of the genes is fe-ru-y. The relatively long distance, 42 units, compares with 48 determined by Tadano (1969a,b). The C. p. fatigans order was determined to be f e - w y and the ma-fe-ru-y order in C . p . pallens. The w y crossover values agree well in both experiments. Some morphological mutants have been studied. Gold (McClelland and Smithson, 1968) was clearly shown to be in linkage group 2. Almost 10,000 backcross progeny gave ru-y recombination values of about 24% in females and 17% in males. Gold-y values are about 22%, with no significant difference in sexes. The sequence is Gold-yellow-ruby, and Gold may serve as a valuable marker because of its evidently close link-

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J A M E S B. KITZMILLER

age to the dominant resistant gene for DDT. Tadano (1969~)found that in two strains of C . p . molestus the white locus was located very close to the sex locus, less than one crossover unit distance in each case. One sex-linked recessive, divided eye (Barr, 1969) and one autosomal recessive, enlarged tergum (Barr and Narang, 1972) , have been investigated in Culex pipiens. Since divided eye showed about 30% recombination with sex (range 16-34%) and about 30% (range 22-35%) with red eye, the order is probably de-sex-r. The autosomal linkage of enlarged tergum with yellow and rust, group 2 markers was as follows: et-y 18% (13.6-21.8%) ; y-ru 20% (15.1-24.6%). The order appears to be et-y-ru. In Culex tarsalis the locus for white eye is autosomal, yellow is linked with sex with about 6% recombination (Barr and Myers, 1966a) and is independent of the locus for malathion resistance (Calman and Georghiou, 1970).

4. Culex tritaeniorhynchus The extensive studies of Baker and his group have permitted organization of preliminary linkage groups in Culex tritaeniorhynchus (Baker and Sakai, 1974). Assigned to linkage group 1, in order, are: golden, white, sex, Delta. In linkage group 2 are Red spotted, ebony, and Alkaline phosphatase and in linkage group 3, curved leg and straw. Recombination values vary, as usual (Baker and Sakai, 1972a) but are in essential agreement among the results of different experiments. Some average recombination values for the sex chromosome: go--20, 10% ; w-sex, 8% ; sex-D, 2%; go-sex, 25%; w-D, 10%. These averages (calculated by the present reviewer) represent pooled data from different experiments, and are entirely from males, since there is complete linkage in females (Baker and Rabbani, 1970; Sakai et al., 1973a,b). On chromosome 2, preliminary experiments indicate about 38 units between Rs and Aph and about 2 units between Aph and e. Likewise about 20 units separate cl and s on chromosome 3. Linkage data for chromosome 1 may be found in Baker and Rabbani (1970) , Baker and Sakai (1972a, 1973b), and Sakai et al. (1972b) ; for chromosome 2, Rabbani and Baker (1970) and Sakai et al. (1973a) ; and for chromosome 3, Baker and Sakai (1973a).

D. LINKAGE GROUP-CHROMOSOME CORRELATIONS The description of loci and linkage studies has permitted the gradual formulation of linkage groups in those species that have been sufficiently studied. Simultaneously the descriptions of karyotypes and salivary chro-. mosomes in those same species clarified the cytological picture. It re-

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mained for the production of inversions and translocations to provide the necessary cytogenetic correlations between linkage groups and specific chromosomes. No definite correlations have as yet been made in Anopheles. Translocations (Rabbani and Kitzmiller, 1972a,b) and inversions (Rabbani and Kitzmiller, 1974) are now available in Anopheles albimanus and are being combined with isozyme markers. Translocations and inversions are available in other species (Krafsur, 1972; Krafsur and Davidson, 1973) and when sufficient markers are available progress will be rapid. Definite correlations have been made between linkage groups in Aedes aegypti, Culex tritaeniorhynchus, and Culex pipiens. The first demonstration was in A . aegypti (McDonald and Rai, 1970a,c), in which two translocation stocks with suitable markers were studied in both males and females. The genetic markers in linkage group 1 were associated with the smallest chromosome. The genetic markers in linkage group 2 were associated with the longest chromosome, the markers in linkage group 3 were associated with the chromosome of intermediate size with the secondary constriction. After this positive identification, the chromosomes in A . aegypti were renumbered to correspond with the numbered linkage groups-i.e., chromosome 2 is now the longest and chromosome 3, intermediate. The same conclusions were reached independently by Bhalla ( 1971a, 1973a,b) with different inversions and translocations. Using translocations, Ved Brat and Rai (1973a) have located blt and min on chromosome 3 in A . aegypti. Following shortly was confirmation in Culex tritaeniorhynchus. As in Aedes, there is no recognizable heteromorphic sex chromosome. It had been known (Baker and Rabbani, 1970) that linkage was complete for sex-linked markers in females, that crossing-over took place only in males. Radiation-induced pericentric inversions were incorporated into stocks marked with the sex-linked recessive golden, then crosses were made involving heterozygous inversion males. Crossing-over in males was almost completely suppressed, suggesting that the inversion involved the sex chromosome. The cytogenetic studies unquestionably confirmed the genetic data. I n beautiful photographs, the shortest pair of chromosomes is shown to be heterozygous for a pericentric inversion, resembling very much a subtelocentric Anopheles X chromosome. Equally dramatic was the demonstration of the location of linkage group 2 markers with the short arm of submetacentric chromosome 2, which is easily distinguished cytologically from metacentric chromosome. 3. Of 62 radiation-induced translocations (reduced fertility screening), 46 were extensively investigated. The sex-linked marker golden and the linkage group 2 marker Red-spotted were followed. I n stocks that showed new pseudolinkage be-

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tween go2den and Red-spotted, no crossing-over took place in females; cytological examination indicated a translocation of the short arm of chromosome 2 to the sex chromosome. All data, genetic, cytogenetic, sterility, and recombination, agree and support this conclusion (Baker et al., 1971a,b; Sakai et al., 1971a,b; Baker and Sakai, 1974). Two sets of experiments have been run in Culex pipiens, with conflicting results. The first of these (Dennhofer, 1972) examined the cytogcnetic associations of eight translocations and one inversion, together with the linkage group 2 marker ruby. Seven translocations showed genetic relationships with sex; they were always also associated with the shortest pair of chromosomes. A pericentric inversion (with associated semisterility) was associated with the intermediate-sized chromosome and also showed linkage with ruby. Thus Dennhofer concludes that the shortest pair of chromosomes carries linkage group 1; the intermediate chromosome, the group 2 markers; and the longest pair of chromosomes, thc group 3 markers. The other investigation (Bhalla et al., 1975) utilized the three standard methods for linkage group chromosomal correlations, semisterility, linkage alterations, and direct observation of chromosomes. Bhalla’s evidence indicates that a translocation involving the shortest and longest chromosomes also shows close linkage betwecn sex and ruby, a group 2 marker. Other genetic and cytogenetic data bear out his conclusion that ruby and other group 2 markers are associated with the longest pair of chromosomes. By elimination, the intermediate length chromosome would contain the group 3 markers. It seems clear that further work must be done to resolve this contradiction. One of the problems is that it is not always easy to distinguish chromosomes 2 and 3, since they differ by only about 1 pm in length. Another possibility, not very likely, is that ruby and other group 2 markers are in fact on the longest chromosome in C . fatiguns (Bhalla) and on the intermediate one in C . pipiens (Dennhofer) .

E. SEX GENETICS There is a voluminous literature on hormones, reproductive physiology, and similar subjects, and in many of these investigations mosquitoes have been used as experimental animals. Consideration of this literature is outside the scope of this present review. I have attempted to limit discussion here to a few areas, clearly genetic, but other areas might well have been included. One of the few studies in sex determination (Aslamkhan, 1973a) indicates that in Anopheles stepheles sex determination is of the XY type,

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confirming the XY system proposed (Mason, 1967) for Anopheles gambiae. Isozyme evidence (Narang and Kitamiller, 1971a) also supports the XY mechanism. 1 . Monogamy

Whether a female mosquito effectively mates more than once influences genetic results. Most evidence (Craig, 1967a) indicates that multiple insemination is infrequent although it does occur. Transplantation and injection experiments (Craig, 1967a) and mating tests (Spielman et al., 1967; Jones, 1970) showed that in Aedes aegypti, although copulation may take place several times, effective insemination occurs only once. This monogamy (Anonymous, 1972, but see Jones, 1973c) is induced by a material from the male accessory glands (Craig, 1967a; Hinton, 1974) which is transferred a t the time of copulation. The active substance designated “matrone” (Fuchs et al., 1968) is a peptide (Fuchs et al., 1969; Fuchs and Hiss, 1970). Females of aegypti are refractory to insemination for from 30 to 60 hours after emergence (Lea, 1967, 1968; Gwadz, 1967; Gwadz and Craig, 1968; Akey and ,Jones, 1968), apparently owing to juvenile hormone (Lea, 1968; Gwadz et al., 1971b). The mechanisms that permit copulation but not insemination are unclear. Failure to achieve complete copulation (Gwadz and Craig, 1970), expulsion of the seminal mass (Spielman et al., 1967, 1969), and behavioral traits (Gwade et al., 1971a) have been suggested. The activity of both junvenile hormone and matrone evidently control receptivity of the female by acting upon the terminal abdominal ganglion (Gwadz, 1972). Treatment of young larvae with Apholate (Powell and Craig, 1970) reduced the volume of the accessory gland, and such males were less efficient in making females monogamous, probably owing to sperm depletion (Sharma and Rai, 1967). Matrone also has an effect (Hiss and Fuchs, 1972) on oviposition. Multiple inseminations occur rarely in the laboratory and may take place if multiple copulations occur within P 5 hours of each other (Craig, 1967a; Spielman et al., 1967). Inadequate sperm transfer may also be involved (Gwadz and Craig, 1970). The results of George (1967) with Aedes aegypti and of Bryan (1968, 1972b) with Anopheles gambiae generally are in agreement with the matrone hypothesis, and the latter experiments (Bryan, 1972b) indicate that only certain types of F, hybrid sterile males may be effective in field trials. In Anopheles farauti, sterility is not induced in females when mated to F, hybrid sterile males with normal-sized accessory glands (Bryan, 1973a,b,c). The available evidence indicates that the accessory glands of the hybrids lack secretory function.

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JAMES B. KITZMILLER

The results of Gubler (1970a) in crossing Aedes polynesiensis with Aedes albopictus are also consistent with the matrone hypothesis, as are the results of Grover and Pillai (1970b). Male Anopheles gambiae (Cuellar et aZ., 1970) may mate up to 4 times and still produce offspring. In Aedes aegypti the aggressive males may inseminate about 5 females, but then become sexually depleted and do not inseminate females for the rest of their lives (Jones, 1973a) although they do renew the supply of spermatozoa and the accessory gland secretion. Modified behavior is evidently responsible. Fecundity was lowered in several strains of A . aegypti (Hacker, 1971), but the results intriguingly suggest a possible balanced polymorphism mechanism. In Aedes atropalpus early sexual receptivity (Gwadz, 1970) appears to be under the control of a single semidominant autosomal gene. Spontaneous parthenogenesis has been reported in Culex fatigans (Das et al., 1968), but the description of the experimental techniques does not rule out possible contamination by an incompatible male. Barr (1974b) has summarized reproductive processes in the Diptera, with special reference to mosquitoes. (see also Jones, 1967, 197313; Jones and Pilitt, 1973). A male factor enhanced egg deposition in Aedes aegypti (Leahy, 1967, 1970) and nonspecific functions when transplants of Culex pipiens and Drosophila melanogaster were placed into female A . aegypti. 6. Sex Ratio

Details of the Distorter mechanism in Aedes aegypti (Craig and Hickey, 1967b) indicate (Hickey and Craig, 1966a,b) that high male ratios are not due to postfertilization mortality but to a dominant Distorter factor close to or identical with the dominant male factor M. Distorter functions only when heterozygous and only when associated with the M factor (MD md), never when associated with m. Strains differ in the proportions of MD, many strains having about 40% females (Hickey and Craig, 1966a). A Distorter-balancer mechanism is probably involved. Analysis of differential mortality, larval density, temperature, and parental age (Hickey, 1970) did not influence the action of Distorter. A presumably different mechanism involving an inversion and a translocation (McGivern and Rai, 1974) produces gametes strongly in favor of females. A high frequency (84%) of alternate segregation a t metaphase I is involved. A distorter system has also been studied in Culex pipiens (Sweeney, 1972), in which it appears to be a recessive, sex-limited, sex-linked trait with variable expression. As in A . aegypti, the d factor is close to the sex locus. Postzygotic effects were ruled out, but cytological examination of meiosis showed that in the distorter stock, chromosome 1 was broken

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in about 40% of all anaphase figures. The data are consistent with breakage, near the centromere, in the m dyads, with assumed lethal effects so that M sperms are produced in excess. The actual effective ma1e:female ratio a t any one time may be quite different from the generally accepted 1 : l (Cuellar, 1 9 7 3 ~ ) The . daily emergence of males, their sexual maturity, and probability of survival are important. 3. Sexual Aberrancies

The literature on mosquito gynandromorphs and intersexes has suffered from superficial treatment, desultory bibliographic search, and careless terminology. Brust (1966b) quite properly points out the differences between intersexes and gynandromorphs and describes in detail gynandromorphs of Aedes abserratus and Aedes excrucians as well as intersexes of A . abserratus, A . communis, and A . excrucians. He also reviews the mechanisms and theories of gynandromorph and intersex formation and sorts out the literature that deals with these two types. Rai (196810) reports preliminary data on the genetic mechanism of gynandromorph formation. It would seem that the time is ripe for a critical review of the fragmentary and superficial literature on this subject, but that will not be undertaken here. Earlier lists (Roth, 1948; Bates, 1949; Kitzmiller, 1953) clearly included both gynandromorphs and intersexes. I shall attempt to list the cases described since 1966. This listing is undoubtedly incomplete. GYNANDROMORPHS

Aedes aegypti albopictus cinereus craggi dentatus dorsalis excrucians hendersoni nigrornaculis togoi triseriatus vexans Culex cinereits erythrothorax

Craig and Hickey (1967b) Craig and Hickey (1967b) Brust (1966b) Huang (1974) van Someren (1969) Blakeslee et al. (1966) Brust (1966b) Skierska (1969) Grimstad and Defoliart (1974) Miura (1969) Chellappah (1965a,b) Ezenwa and Venard (1973) Minson (1969) van Someren (1969) Blakeslee and Rigby (1965)

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JAMES B. KITZMILLER

fatigans

f uscocephalus nigripalpus pipiens (molestus) salinarius tarsalis

tritaeniorhynchua Culiseta inornata novazealandiae Mansonia perturbans Trichosporon digitatum

Seal (1966) Meadows (1966) Shetty and Chowdaiah (1973) Aslamkhan (1970) Meadows (1966) Taylor et al. (1966) Laven (1967~) Meadows (1966) Taylor et al. (1966) Rigby (1966) Taylor et al. (1966) Rosay (1968) Harmston (1965, 1971) Mitchell and Hughes (1969) Aslamkhan and Baker (1969a) Benge (1970) Dobrotworsky (1972) Pinger (1972) Lee (1967)

INTERSEXES Aedes abserra t us communk excrucians hexodontus nigripes

Brust Brust Brust Brust Brust Brust

(1966b) (1966b) (1966b) and Smith (1972) (1966a) and Smith (1972)

F. SUSCEPTIBLE AND REFRACTORY STRAINS 1. Filariae

The literature on the genetic aspects of vector susceptibility to parasites (Macdonald, 1967a,b) has been thoroughly and ably reviewed. Since then several selection experiments have determined strain differences and the genetic basis of susceptibility to various parasites. The f" gene controls susceptibility to other Rrugia and Wuchereria, but not to Dirofilaria (Macdonald and Ramachandran, 1965), or FoleyelZa (Terwedow, 1973). Using Brugia pahangi in Aedes aegypti Townson (1971b) clearly showed that susceptible mosquitoes have higher mortality than refractory ones and that mortality is relative to the microfilarial density. T o account for the survival of the obviously disadvantageous susceptibility, Townson suggested a balanced polymorphism with selection for heterozygotes. Different strains of A . aegypti contain allelic genes for susceptibility,

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and different strains differ markedly for susceptibility to B. pahangi (Rodriguez, 1973; Rodriguez and Craig, 1973). The larval stages also develop in male susceptible mosquitoes (Terwedow and Rodriguez, 1973 ; Townson, 1974), as does the Foleyella species found in the fat body (Terwedow, 1973). Susceptibility to B. pahangi in Culez pipiens is probably controlled by a single sex-linked gene, sb, close to the sex locus (Obiamiwe and Macdonald, 1973). The Delhi strain of C. fatigans and four strains incompatible with it were tested for susceptibility to W . bancrofti. All five strains were highly susceptible and there were no evident differences among the strains (Thomas and Singh, 1974; Singh and Curtis, 1974). The D, strain with Paris cytoplasm, Freetown genome is also incompatible with local strains, but also highly susceptible to infection with W. bancrofti (Thomas, 1974) and therefore not suitable for use in the field against local populations. Some problems of formal genetics and genetic control proposals involving susceptibility are treated by McClelland and McGreevy (1973a,b). Selection for resistance and susceptibility t o infection with Dirofilark immitis (Raghavan et al., 1967) produced refractory and susceptible strains in Aedes aegypti. A single sex-linked recessive gene f t , controls susceptibility to D. immitis (McGreevy, 1971; Zielke, 1972, 1973; McGreevy et al., 1974). This is a different locus (Zielke, 1973; McGreevy et al., 1974) than f". Selection was also effective in producing resistant and susceptible strains for D. repens in A . aegypti (Coluzzi and Gironi, 1971). Selection in Culez fatigans (Partono and Oemijati, 1970, 1972) did not differentiate strains resistant or susceptible to Wuchereria bancrofti, and these authors concluded that no major gene could be hypothesized. However, Thomas and Ramachandran (1970) were able to increase susceptibility in C. fatigans to the rural strain of W. bancrofti, and Zielke (1973) also was able to improve susceptibility by selection. The role of genetics in the control of filariasis is briefly touched by Thomas (1968). The filarial worms also may undergo adaptation to a new mosquito host (Laurence and Pester, 1967) presumably by recombination of different genetic strains. 2. Malaria After demonstrating that Aedes aegypti strains differ widely in susceptibility to Plasmodium gallinaceum, Kilama and Craig (1969) showed that the refractory condition is determined by a single autosomal recessive gene, pls, located on linkage group 2 between Silver-mesonotum and Dieldrin resistance, 8 crossover units from the former and 17 from the latter. The pls gene was found in 8 African strains but not in strains from other continents (Kilama, 1973). A puzzling fact is that although

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most African strains are refractory to P . gallinaceum, this species of malaria does not occur in Africa. Similarly, a recessive gene on linkage group 3 controls resistance to Plasmodium cathemerium in Culex pipiens (Dennhofer, 1971b). Selection for and against susceptibility to P . gallinaceum is also possible in Anopheles stephensi (Corradetti et al., 1969). 3. Physiological Factors Strains of Aedes aegypti differ markedly (Machado-Allison and Craig, 1972) in their abilities to withstand desiccation, and a beginning has been made in the investigation of adaptations, life tables, and ecological genetics of this species (Machado-Allison, 1972a,b; Flores and MachadoAllison, 1972; Fergusson and Machado-Allison, 1972; Machado-Allison et al., 1972).

G. GENETICSOF AUTOGENY Autogeny has been reported in about 50 mosquito species belonging to several genera. Autogeny has a basis that is partly environmental, diet playing a major role. The subject has been ably reviewed by Vinogradova (1965) and Spielman (1971). Although most of the early work with autogeny was done with Culex, relatively little has appeared of late. In a complicated analysis using genetic markers, Aslamkhan and Laven (1970) conclude that the genetic basis of autogeny is due to a sex-linked dominant gene D, and a series of multiple alleles on chromosome 3. There are some difficulties with this hypothesis, and the authors suggest that it be taken as a working hypothesis to encourage further investigation. The genetic basis for autogeny in Aedes atropalpus (O’Meara and Craig, 1969) appears simple. Crosses among eleven widely separated field populations indicate that autogeny is controlled by a single dominant autosomal gene. Tentative evidence also indicates that blood-feeding is genetically controlled and may be closely linked to autogeny. Genetic and dietary factors (O’Meara and Krasnick, 1970) interact. Changes in larval diet affect both penetrance and expression. Autogenous fecundity in A. atropalpus (O’Meara, 1972) is controlled by a series of modifiers. Three of the four subspecies lack both the autogeny gene A and the modifiers. The monofactorial basis of autogeny has also been demonstrated in Aedes detritus (Rioux et al., 1973), but autogeny appears to be polyfactorially determined in Aedes togoi (Thomas and Leng, 1972). Local populations of Aedes taeniorhynchus (O’Meara and Evans, 1973) are polymorphic for autogeny, which increases clinally on a north to south basis.

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H. GENETICSOF INSECTICIDE RESISTANCE This important area has not lacked workers, and much information is available. Fortunately two superb reviews have appeared since 1967, and therefore only papers not covered by these reviews will be treated here. The first of these reviews (Georghiou, 1969) deals exclusively with the genetics of resistance in house flies and mosquitoes and provides a most thorough and detailed coverage of genetics. Good discussions of single gene vs polygenic inheritance, dominance and recessiveness, techniques, and detailed genetic test procedures are included. The resistance literature is carefully reviewed from a genetic point of view, by species and by insecticide. An extensive bibliography is included. The second review (Brown and Pal, 1971) is actually a 491-page monograph that covers the entire field of resistance and susceptibility in arthropods. Introductory chapters on the nature, characterization, detection, and measurement of resistance are followed by chapters dealing with groups of insects, three valuable annexes (chemical names, a list of published reviews, and the WHO computer program), and an exhaustive 36-page list of references. Two specific chapters on resistance in anopheline mosquitoes and in culicine mosquitoes are included. Genetic data are included in the discussions of each species. A shorter but up-to-date review (Pal, 1973) summarizes the data for resistance in the anophelines, especially since 1970 (see also Busvine, 1971). Strain differences in Aedes aegypti with respect to the genetic components of resistance were shown for organophosphorus compounds (Madhukar and Pillai, 1970) and for D D T (Inwang et al., 1967; Wood, 1968). Wood (1970) also showed that the y locus in group 2 reduces the penetrance of the gene for DDT resistance. In Culex fatigans dieldrin resistance in an upper Volta population (Hamon et al., 1967) was found to be monofactorial, incompletely dominant, with dominance stronger in the larvae, and males of all genotypes more susceptible than females. Field observations, however, were difficult to explain on a one-gene hypothesis. The authors suggest that modifiers are operative in nature, but that perhaps a major gene had been selected in the laboratory strain. Suzuki and Umino (1969) also suggest monofactorial inheritance of diazinone resistance in C. molestus. Also in C.fatigans, a t least two different genes are postulated (Kalrs, 1970) to account for DDT resistance. Nine generations of selection pressure with a carbamate (propoxur) failed to develop resistance in C. tarsalis (Georghiou et al., 19741, and 31 generations of selection were negative in C.nigripalpus against Paris Green (Rathburn and Boike, 1973a,b). Thirty and 25 generations, respec-

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tively, of selection for fenitrothion resistance in Culex pipiens fatigans and Anopheles albimanus were also negative (Georghiou and Calman, 1969). Similar negative results in C. fatigans (Micks and Berlin, 1970; Micks and Gaddy, 1973) were obtained after 98 generations of selection against diesel oil and 35 generations against an aromatic hydrocarbon. A D D T resistance gene in C . tarsalis may be pleiotropic for resistance to pyrethrins (Plapp and Hoyer, 1968), but close linkage or allelism have not been ruled out. Resistance to D D T is present in both species A and species B of the Anopheles gambiae complex. Crossing studies between resistant and susceptible strains (Haridi, 1970, 1972) show that there possibly are two different resistance genes, a dominant in the Togo strain of A and the Sudan strain of B, but an incompletely dominant gene in the Upper Volta strain of A. Further studies (Haridi, 1974) indicate two types of dieldrin resistance, a dominant gene in A and an incompletely dominant one in both species. Linkage tests with collarless and Diamond indicate that both dieldrin genes were closely linked, perhaps even a t the same locus, with incomplete dominance due to modifiers. Evidently Diamond, D D T , and Dieldrin are on the same linkage group, but the preliminary data given in this abstract do not permit definitive analysis. In Anopheles pharoensis (Kame1 et al., 1970; Hamed et al., 1973a) DDT resistance has been reported as due to a single autosomal recessive gene. Evidently strain differences exist in Anopheles stephensi (Pervez and Aslamkhan, 1973). Crosses among five populations of Anopheles pseudopunctipennis from Mexico (Martinez-Palacios and Davidson, 1967) produced no evidence of hybrid sterility, testes of F, males were normal, and DDT resistance was shown to be due to a single semidominant gene. Relatively little is known about the genetics of carbamate resistance in Anopheles albimanus (Georghiou et al., 1972; Zahar and Davidson, 1973a,b). The studies thus far indicate (Georghiou, 1972; Ariaratnam and Georghiou, 1971, 1973) that carbamate resistance is controlled by one major gene and several modifiers and that segregation is independent of the Stripe marker (see also Coz et al., 1968). Seasonal increases in carbamate and OP resistance (Georghiou et al., 1973) may be due to higher fitness levels of OP-resistant genotypes and poses the problem of indirect selection for resistance in agricultural areas. Genetic resistance to radiation is a little-recognized but important fact. Stahler (1971) demonstrated that Aedes aegypti could be selected for resistance to 2500 R. After 90 generations, radiation-induced mortality was 30% compared to 93% in the control. Resistance to chemosterilants may also develop (Patterson et al., 1967).

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Resistance to Malathion develops quickly in Aedes taeniorhynchus (Mount et al., 1974), but cross resistance to other adulticides did not appear. A formula (Stone, 1968) for determining the degree of dominance of resistance to chemicals, suggests revision of the standard terms for dominance. See also Suzuki ( 1968, 1969), Tadano (1970a), Thomas (1966, 1973), Umino (1965a), Umino and Suzuki (1963, 1964, 1966, 1968, 1969), Wilson et al. (1972), Wood (1967b), and Zaghloul and Brown, (1968). Ill. Cytogenetics

A. SALIVARY CHROMOSOMES Without question the most important development in mosquito genetics in the last 7 years has been the demonstration that the sibling species of the Anopheles gambiae complex, formerly inseparable on morphological grounds, could be unequivocally separated using the polytene chromosomes (Coluzzi and Sabatini, 1967, 1968a,b,c,d, 1969a,b; see also Coluzzi, 1964, 1966; Davidson and Chalkley 1970; Davidson and Hunt, 1973). The cytotaxonomic identification permitted studies on the distribution of the cryptic species with concomitant economy of time, facilities, and money in control operations. Quickly the cytogenetic method was extended to adult females (Coluzzi, 1968; Green, 1972a; Hunt, 1972a,b) with the demonstration that ovarian nurse cells also had readable polytene chromosomes, an observation that has permitted for the first time adequate field surveys of species of the complex. The ramifications and possibilities for other sibling species or complexes are important and obvious. Later a new member of the complex, species D, was discovered cytologically (Hunt, 197213; Davidson and Hunt, 1973; Davidson, 1973c; White, 1973a,b). The possibility of accurately determining distributions and proportions of the species has far-reaching implications in control operations, finances, and national and international health programs. Several studies using cytotaxonomy have clarified the geographic (Service, 1970a,b,c, 1972; Carnevale, 1972; Green, 1970; Davidson, 1967b; Chauvet et al., 1968, 1969; White, 1967b, 1968, 1969, 1971a; Coz, 1973a,b,d), seasonal (Shidrawi, 1972; White, 1972; Service, 1969; White and Maguyuka, 1972), and ecological (Service, 1970b; Coz, 1972; Shelley, 1973; Green, 1971; White and Rosen, 1972, 1973; White et al., 1972; White, 1972, 1973a,b, 1974c) distributions of the members of the Anopheles gambiae complex.

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1. Maps

Detailed salivary gland chromosome maps have recently been published for the following anophelines: Anophe lea albimanus aquasalis argyritarsis atropos atroparvus barbirostris barbirostris ahomi bradle yi crucians darlin& farauti franciscanus hectoris maculatus maculipennis messeae neomaculipalpus nigerrimus nuneztovari perplexens plumbeus pseudopunctipennia pulcherrimus punctimacula stephensi subpictus superpictus tessellatus vestitipennis walkeri

Keppler e t al. (1973) Kitzmiller and Chow (1971) Kreutzer e t al. (1975) Kreutzer e t al. (1969a) Farci e t al. (1973) (an important, more detailed revision of the original map) Chowdaiah e t al. (1970) Paaahan (1974) Kreutzer e t al. (1970) Kreutzer e t al. (1970) Kreutzer e t al. (1972a) Bryan and Coluzzi (1971) Smithson and McClelland (1972) Baker e t al. (1966) Narang e t al. (197313) Stegniy e t al. (1974) Kabanova e t al. (1972a) Kitzmiller et al. (1966) Seetharam and Chowdaiah (1971) Kitzmiller et al. (1978) Kreutzer and Kitzmiller (1971b) Coluzzi and Cancrini (1971) Baker e t al. (1965) Baker e t al. (1968) Kreutzer e t al. (1969b) Sharma e t al. (1969b) Narang e t al. (1973a) Seetharam and Chowdaiah (1974) Coluzzi e t al. (1970a, 1973a) Narang e t al. (1974) Chowdniah e t al. (1966) Kitzmiller et al. (1974a)

Preliminary descriptions or abstracts dealing with anopheline polytene chromosomes have been published as follows : Anopheles albimanus albitarsis argyritarsis annulipes apicimacula aquasalis barbirostris

Kitzmiller and Baker (1965) Kitzmiller (1973) Kitzmiller e t al. (1973a) Kitzmiller and Baker (1965) Kreutzer (1972a,b) Kitzmiller (1973) Green (1972b) Kitzmiller and Kreutzer (1967) Kitzmiller and Kreutzer (1967) Chowdaiah et al. (1967)

GENETICS, CYTOGENETICS, AND EVOLUTION OF MOSQUITOES

bengalensis braziliensis darlingi eiseni evansae fluuiatilia hectoris hyrcanus nigerrimus koreicus lindesayi japonicus maculatus willmori neomaculipalpus noroestensis pharoensis pseudopunctipennis punctimacula rangeli sinensis sineroides splendidus stephensi subpictus triannulatus uagus uaruna uestitipennis wa Lkeri Chagasia bathanus

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Kanda (1969) Kitzmiller (1973) Kreutzer (1972a,b) Kitzmiller (1973) Kitzmiller and Kreutzer (1967) Kitzmiller and Baker (1965) Kitzmiller (1973) Sharma et al. (1968a) Chowdaiah and Seetharam (1972) Kitzmiller and Baker (1965) Chowdaiah et al. (1967) Kanda and Oguma (1970b) Kanda (1969) Sharma e t al. (1968e) Kitzmiller and Baker (1965) Kitzmiller (1973) Hamed e t al. (1973b) Kitzmiller and Baker (1965) Kitzmiller and Baker (1965) Kitzmiller and Kreutzer (1967) Kitzmiller (1973) Kanda (1969) Kanda and Oguma (1970a, 1973) Sharma (1971) Kanda (1969) Kanda and Oguma (1972a,b) Sharma et 01. (1968d) Amirkhanian (1973a) Sharma et al. (1966) Siddiqi and Aslamkhan (1973) Chowdaiah and Seetharam (1973) Kitzmiller and Kreutzer (1967) Kitzmiller (1973) Chowdaiah et al. (1967) Chaudhry (1971) Pasahan (1973) Kitzmiller and Baker (1965) Kreutzer et al. (1972a) Kitzmiller and Kreutzer (1967)

Some preliminary descriptions of culicine polytene preparations are reported. The best of these are in Kanda (1971). Others reported are: Aedes aegypti Culex fatigans Culex fuscocephalus Culex vishnui Alalayia genurostris Mansonia uniformis W y o m y i a smithii

Sharma et al. (1967~) Sharma et al. (1967b) Kanda (1969) Pasahan (1971) Chaudhry (1973a) Kanda (1969) Chaudhry (1973b) Kanda (1969)

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Until a few years ago almost all polytene chromosome preparations had been made from the anophelines. And with good reason: in spite of many efforts by many investigators (this reviewer among them) the salivary gland chromosomes of Culex, Aedes, and other genera just seemed unamenable to spread and coloration. This impasse was broken by Dennhofer (1968, 1974b), who published the first map for Culex pipiens, with good photographs of the complement. The key to succcss was evidently making the preparations during a very narrow time range, between 14 and 18 hours a t 2Oo-24OC after molting from third to fourth instar. Rearing a t lower temperatures (15OC) for longer times gave thicker, better-staining chromosomes. At the lower temperature a t least 22-24 hours, but not later than 40 hours, after beginning of fourth instar was the optimum time. Sharma et nl. (1969a) published a map of Culex fatigans and compared their map with Dennhofer’s. No pictures are published. The maps differ in several banded areas and in the centromere regions. Another map of C. fatigans (Kanda, 1970) showed many similarities and some differences from Dennh6fer’s map. The photographs of actual chromosomes are not of the best quality, but they are good for Culez. A fourth study, with reasonably good photographs (Tewfik and Barr, 1974a,b), summarizes the differences among the 4 maps and attributes the differences to techniques of preparation. This reviewer is of the opinion that (1) the known differences in the members of the complex probably would be reflected in real banding pattern differences; (2) the age of the larva from which the preparations are made could easily account for intensity differences in bands; and (3) artifacts due to technique certainly exist in the anophelines. Culez chromosomes will be difficult to work with becauee of several factors (greater length, poor spread, telomere association, etc.) but should be worth detailed studies. Different techniques, such as the acid alcohol-hydrolysis-cresyl violet (Amirkhanian, 1968), deserve thorough and persistent studies. Chromosomes from other tissues (Cambefort and Larrouy, 1971) hold promise and should be investigated more assiduously. See also Dennhofer (1975b,c). 2. Polymorphism

Detailed studies on the anophcline polytene chromosomes revealed the existence of considerable amounts of inversion polymorphism in some species, moderate amounts in others, and an apparent lack in a few. Relatively few inversions were found in X chromosomes. Anopheles albima?iw appears to be lacking in inversion heterozygosity both in the laboratory and in the field (Keppler and Kitzmiller, 1969) although inversions may be easily produced in the laboratory by radiation (Rabbani and Kitzmiller, 1974). Moderate amounts of naturally occurring inversion poly-

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morphism have been shown in Anopheles aquasalis (Kitzmiller and Chow, 1971) ; atropos (Kreutzer et nl., 1969a) ; albitarsis (Kitzmiller et al., 1974b) ; messeae (Kabanova et al., 1972b, 1973) ; nuneztovari (Kitzmiller et al., 197313) ; franciscanus (Smithson and McClelland, 1972) ; and pulcherrimus (Baker et al., 1968). Elevated amounts of inversion polymorphism have been found in natural populations of Anopheles bradleyi and crucians (Kreutzer and Kitzmiller, 1971a; Kreutzer et al., 1970) ; darlingi (Kreutzer et al., 1972b), freeborni (Smithson, 1970), gambiae (Coluzzi and Sabatini, 1967, 1968a,b, 1969b), maculatus (Narang et al., 1973b), perplezens (Kreutzer and Kitzmiller, 1971b), punctimacula (Kreutzer et al., 1969b), stephensi (Coluzzi et al., 1970a, 1973a,b,c), subpictus (Narang et al., 1974), and vestitipennis (Chowdaiah et al., 1966). By far the greatest amount of polymorphism in any species has been reported in Anopheles messeae (Belcheva and Mihailova, 1972), in which 27 different autosomal inversions, one duplication, one deficiency, one “homozygous inversion,” and one “deletion” were described from three localities in Bulgaria. Some of the chromosomes pictured clearly have inversion loops, but several just cannot be identified from the photographs. Few bits of hard data exist on the adaptive significance of chromosomal polymorphisms in the anophelines. Vector capacity (Kitzmiller et al., 197313) appears to be correlated with a fixed inversion in the X chromosome. White (1974a,b) has summarized we11 the present and possible future work to be done in adaptive polymorphism of the anophelines. 3. Evolutionary Relationships

One of the uses of inversion polymorphism is to trace the evolutionary relationships among related species. This has been done, with varying degrees of completeness, in several groups of anophelines, but principally in the subgenus Cellia. Perhaps the clearest analysis has been made between Anopheles superpictus and stephensi (Coluzzi et al., 1973a). The standard map of A. superpictus (Coluzzi et al., 1970a) may be used to describe the A . stephensi arrangement, which differs in three fixed inversions from A . superpictus: zones 12-13 on 2R; zones 22, 23, and 24 on 2L; and zones 44 and 45 on 3L. Ten strains of A . stephensi from different geographical localities were examined. The Karachi strains showed the typical three inversions and were designated as the basic sequence for A . stephensi. Six intraspecific inversions were found in the 10 A . stephensi strains. None involved the X or 3R, which were also not involved in the A . superpictus-A. stephensi evolutionary changes. Three overlapping inversions were found on 2R, two overlapping inversions on 3L, and one inversion on 2L. All ten strains can be described on the basis of the inver-

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sions in which they differ from the A . superpictus standard. Two of these A . stephensi strains (Coluzzi, 1972, 1973) were found to have different

emcrgencc times perhaps owing to inversion coadaptation. Inversion polymorphism is also correlated with egg length in A . stephensi (Coluzzi et al., 1972). The evolutionary relationships are also clear in the A . gambiae complex. Species A and B differ by extensive rearrangements of the X chromosome (no heterozygotes) but share five inversions in 2R, one in 2L, and one in 3L. These are found in different proportions and combinations in different populations (Coluzzi and Sabatini, 1967). Species C also has a distinct X chromosome, which differs from that of species B by three inversions in the long arm, one of which is overlapping and one of which is included. Species C and species A differ by one long paracentric inversion in the long arm of the X. Only one new autosomal inversion, in 2R,was found in species C. All others were found in A and B, but the C arrangements were more frequent in A, arguing for closer relationships betwecn C and A than C and B (Coluzai and Sabatini, 1968a). The two salt water species of the complex, A . m e w s and A . melas (Coluzzi and Sabatini, 1968b,c, 1969a,b) have X chromosomes that are different, as expected, but A . merus is most like A and A . melas most like C. I n the autosomes A . nierus differs from A or C by two overlapping paracentric inversions in 2R, necensitating a hypothetical intermediate form. In 2R A . melas differs from species B by three paracentric inversions, two found in A and C, one unique to A . m?elas. A long inversion, evidently unique in 3R, is found in A . melas. A tentative phylogenetic scheme shows B as the basic species giving rise to C, which in turn is ancestral to A and to A . melas. A . m e w is derived from A. The newly discovered species D has an X chromosome similar to species c, but differs in important details. The autosomes are considerably different, but no details of inversion differences have as yet been worked out (Hunt, 1972a, 1973.) The species described above, Anopheles superpictus, stephensi, and gambiae, all belong to the subgenus Cellia. Maps of A . pulcherrimus (Baker et al., 1968), A . subpictus (Narang et al., 1973a; Seetharam and Chowdaiah, 1974) and A . tessellatus (Narang et al., 1974) all also in the subgenus Cellia, reveal remarkable affinities in some cases with A . superpictus, A. stephensi, and A . gambiae; A . tessellatus stands apart. It has little polymorphism and no readily ascertainable affinities with the other species. The detailed comparisons for the other species may be found in Narang et al. (1973a,b). All X chromosomes are distinctive. Within the autosomes there are clear banding pattern similarities in all arms. The two closest species appear to be A . maculatus and A . stephensi, which differ only in parts of 2R and 3R. Anopheles pulcherrimus could

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have been derived from A . maculatus by a series of two interarm exchanges and paracentric inversions in chromosome 3 and a t least 11 paracentric inversions in chromosome 2. Verification of these tentative relationships must await detailed study of naturally occurring polymorphism in these species, similar to that observed in A . stephensi. It seems clear, however, that A . subpictus and A . gambiae have much in common (Narang et al., 1973a). In the autosomes, eight paracentric inversions differentiate A . subpictus and A . gambiue, three in 2R, three in 2L, and two in 31,. 3R in A . subpictus is not all similar to 3R in A . gambiae. The cytological picture appears to bear out the taxonomic one in which A . subpictus and A . gambiae are both in the series Pyretophorus while A . maculatus, stephensi, pulcherrimus, and superpictus are in the series Naocellia. The apparently totally dissimilar A . tessel2atus is in another series, Neomyzomyia. It is probable that the chromosomal events postulated to differentiate the species might also have been involved in the evolution of the species groups within Cellia. The excellent studies by Kabanova et al. (1972a,b, 1973) have demonstrated inversions in A . messeae in the X, in 2R, in 3R, and 3L. These are very close if not identical to some of the inversions.found in North American species (Kitzmiller et al., 1967). The X chromosome of A . messeae is almost identical with that of A . quadrimaculatus. The chromosomal relationships between the Palearctic (Frizzi, 1967) and Nearctic members of the A . maculipennis complex are strikingly verified. Detailed analysis of the similarities and differences between A . atroparvus and three North American species has been initiated (Kreutzer, 1973, 1974a,b). The A . messeae inversions occur in all combinations in Siberian populations (Kabanova et al., 1972b) and evidently have high adaptive values as heterozygotes in seasonal fluctuations (Kabanova et al., 1973). I n the North American A . maculipennis species, several papers detail the chromosomal evolutionary patterns. One inversion in the X, one in 2R, two in 3R, and one in 3L differentiate the closely related species A . crucians and A . bradleyi. Relationship with A . jreeborni is more complicated, but the A . crucians and A . freeborni 2L arrangements differ by five paracentric inversions, and the 3R arrangements by four (Kreutzer and Kitzmiller, 1968, 1970, 1971a; Kreutzer et al., 1970). Similarly, the species A . punctipennis and A . perplexens are related by inversion patterns. Somewhat of a rarity in the anophelines, four inversions are found in the short X of A . perplexens. Two inversions in 2R, and two in 3R, one included, differentiate the autosomes (Kreutzer and Kitzmiller, 1969, 1971b, 1972). Anopheles atropos shows strong chromosomal affinities with A . quadrimaculatus, but with extensive rearrangements due to paracentric inversions, perhaps as many as twenty (Kreutzer et al., 1969a).

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Anopheles wallceri is closely related, chromosomally, to A. atropos and A. quudrimaculatus (Kitzmiller et al., 1974s). A thorough summation of the genetic, cytogenetic, and crossing studies among the North American A . maculipennis species (Kreutzer, 1974b) has been combined with data from crosses between the North American species and A . atroparvus. Cytogenetic relationships were strongest between A . atroparvus and A . punctipennis, weakest between A . atroparvus and A. quadrimaculatus. The strong cytogenetic relationship between A . messeae and A . quadrimaculatus is confirmed by X chromosome and autosomal inversions. Some fascinating data from mammal distribution have inspired a revival of the hypothesis of the North American origin of the A . maculipennis complex and its subsequent spread to the Palearctic region. Four autosomal and one X inversion differences are found between A . pseudopunctipennis and A. franciscanus (Smithson and McClelland, 1972; Smithson, 1972). The chromosomes of a neotropical species, A. vestitipennis show close similarities to those of A . freeborni and A. quadrimaculatus (Chowdaiah et al., 1966), differing by 9 inversions from A . freeborni. In this paper it is suggested for the first time that Anopheles banding pattern arrangements might be on a subgeneric basis. Another Neotropical species, A. punctimacula, again shows the subgeneric pattern and differs from A . vestitipennis by a t least 8 inversions and from A. neomaculipalpus by the same number (Kreutzer et al., 196913). The sibling species of A . farauti from Australia and New Guinea (Bryan and Coluzzi, 1971) differ in two paracentric inversions in chromosome 2. The Neotropical subgenus Nyssorhynchus contains about twenty closely related, morphologically similar species, including most of the important New World vectors of malaria. The cytogenetic approach has been most useful in this “difficult” group, and probably is the only certain method of distinguishing these species with certainty (Kitzmiller, 1973; Kreutzer and Kitzmiller, 1974). Twelve species have thus far been mapped. All show unique banding patterns in the X chromosomc and strong homologies in the autosomes, especially at the free and centromere ends. The internal portions of each arm have been extensively rearranged by paracentric inversions. There is extensive chromosomal polymorphism in some species (Anopheles darlingi, evamae, albitarsis) , very little in others ( A . albimanus). Detailed comparisons between A. aquasalis, A . albimanza, and A . oswaldoi (Kitzmiller and Chow, 1971) list some of the relationships. The greatest amount of chromosomal polymorphism has been found in A . darlingi, a widespread important vector. Thus far,

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9 inversions, one in the X, three in 2R, one in 2L, and two each in 3R and 3L have been recovered (Kreutzer et al., 1972b). The frequency of these inversions differs in the north (Manaus) and in the south (SSio Paulo) of Brazil. Fully 90% of all individuals collected in the state of Amazonas were heterozygous for one or more inversions. Comparisons between A . darlingi and A . argyn'tarsis (Kreutzer et al., 1975) show two inversion differences in 2L, two in 3R, and complex differences in 3L. Similar comparisons between A . darlingi and A . albitarsis indicate three inversion differences in 2L, two in 3R, and a t least four in 3L. An evidently fixed inversion in the long arm of the X chromosome may be used to distinguish vector from nonvector populations of A . nuneztovari (KitEmiller et al., 1973b). An interesting comparison of inversion polymorphism in anophelines with Drosophiln and the chironomids is given by Martin (1969).

STUDIES B. KARYOTYPE The last 7 years have seen investigation of karyotypes in many additional species, but, with one exception, no startling new information. The diploid number of 6 is apparently universal in mosquitoes, and only minor differences in arm-length ratios and the presence of secondary constrictions have been noted. The exception is the report for the first time of B chromosomes in mosquitoes (Belcheva and Mihailova, 1971). I n Anopheles maculipennis and A . messeae these authors report supernumerary chromosomes and aneuploidy in salivary glands, testes, and ovaries. The meiotic pictures unfortunately are not clear enough to show for sure whether the B element exists in addition to the smaller heteromorphic region of the X. The following list includes species in which karyotype descriptions or figures have been published since 1966. An asterisk (") indicates that there was some description or mention in the literature prior to 1966. 1 . Species Described Aedes aegypti*

Aedes a l b o p i c t i d Aedes campestris Aedes cataphylla Aedes cinereus

Baker and Aslamkhan (19691, Mescher and Rai (19661, Kanda (1968), Sharma e t al. (1967d), Rai (1967~1,Bhalla (1971b) Baker and Aslamkhan (19691, Kanda (19681, Sharma and Toor (19701, Jost (1971) Mukherjee et al. (1966) Mukherjee e t al. (1966) Mukherjee et al. (19701, Kabanova and Kartashova (1972)

JAMES B. KITZMILLER

Aedes dianteus Aedes dorsalis Aedes fitchii Aedes hatorii Aedes hexodontus Aedes impiger Aedes implicatus Aedes increpitus Aedes japonicus* Aedes leucomelas Aedes mascarensis Aedes nigromaculis Aedes niphadopsis Aedes polynesiensis* Aedes pullatus Aedes punctor Aedes river& Aedes sierrensis* Aedes simpsoni* Aedes spenceri Aedes thomsoni Aedes togoi* Ae des t rise ria tus* Aedes varipalpus Aedes vexans* Aedes vittatus Aedes w-albus Anopheles albimanus* Anopheles annularis Anopheles Anopheles Anopheles Anopheles Anopheles Anopheles Anopheles

balabacensis barbirostrk bengahnsis culicijacies earlei* jarauti fluviatilis

Anopheles franciscanus* Anopheles jreeborni* Anopheles gambiae A* Anopheles gambiae B* Anopheles gambiae C* Anopheles maculatus Anopheles maculatus willmori Anopheles messeae* Anopheles mela** Anopheles merus* Anopheles nigerrimus

Kabanova and Kartashova (1972) Mukherjee et al. (1966), Mukherjee and Rees (1970) Mukherjee et al. (1970) Kanda (1968) Mukherjee el al. (1966) Mukherjee et al. (1970) Mukherjee et al. (1970) Mukherjee el al. (1966) Kanda (1968,1971) Kabanova and Kartashova (1972) Rai (1966) Mukherjee et al. (1966) Mukherjee el al. (1966) Rai (1966) Mukherjee et al. (1970) Kabanova and Kartashova (1972) Kanda (1968) Rni (1966),Mikherjee et al. (1970) Rai (1966) Mukherjec el al. (1966) Baker and Aslamkhan (1969) Kanda (1968) Cocke et al. (1973) Mukherjee et nl. (1970) Mukherjee el al. (19661,Kanda (1968) Rai (1966) Singh and Bhat (1970) Keppler et al. (1973) Aslamkhan and Baker (1969b),Narang et al. (1972b) Aslamkhan and Baker (196913) Avirachan e t al. (1969) Kanda (1968) Aslamkhan and Baker (196913) Mukherjee et al. (1970) Bryan and Coluzzi (1971) G. P. Sharma et al. (1968b,1970), Arirachan el al. (1969) Mukherjee et nl. (1970) Mukherjee et al. (1966) Coluzzi and Sabatini (1967) Coluzzi and Sabatini (1967) Coluzzi and Sabatini, (1968a,b,c) Narang e t al. (1972b). G. P.Sharma et al. (1968f,1970) Kabanova et a / . (1972a) Coluzzi and Sabatini (1969b) Coluzzi and Sabatini (1969h) Aslamkhan and Baker (1969131,Avirachan et al. (1969)

GENETICS, CYTOGENETICS, AND EVOLUTION OF MOSQUITOES

Anopheles Anopheles Anopheles Anopheles Anopheles Anopheles

plumbeus piilcherrimus sinensis sineroides splendidits stephemi*

Anopheles subpictiis Anopheles Anopheles Anopheles Armigeres

superpictus lessellatus vagus

subalbatus*

Culex annulus Culux apicnlis Culex bitaeniorhynchus Culex erythrothorax

Culex fuscanus

Culex Culex Culex Culex Culex Culex Culex

fuscocephala gelidus haynshii* neolitoralis orientnlis pipiens* pipiens fatiguns*

Culex pipiens pallens* Culex pipiens molestiis* Culex pseudovishnui Culex raptor Culex salinnriiis* Cnlex sitiens Culex tarsalis* Culex teriitans* Culex theileTi* Culex th~iomhiiz Culex t i itneniorhynchus* Culex voraxf Culiseta impatiens Culise 1 a incide ns Culiseta inornnta* Eretmapodites chrysognster* Ficalbia chamberlaini clavipalpus Ficalbia hybridi Ficalbia minima Malaya genurostris

353

Coluzzi and Cancrini (1971) Baker e l al. (1968) Kanda (1968) Kanda (1968) G. P. Sharma e l al. (1970, 1968~) Avirachan et al. (1969), Aslamkhan (1973a), Hafeez el al. (19731, Coluzzi el al. (1970a) Aslamkhan and Baker (1969b1, Avirachan el al. (19691, Narang el al. (1972b, 1973a) Coluzzi et al. (1970a) Narang el al. (1972b) Avirachan el al. (1969) Baker and Aslamkhan (19691, Kanda (1968) Baker and Aslamkhan (1969) Mukherjee et al. (1970) Baker and Aslamkhan (1969), Kanda (1968, 1971) Mukherjee el al. (1966) Baker and Aslamkhan (1969) Baker and Aslamkhan (1969) Baker and Aslamkhan (1969) Kanda (1968) Baker and Aslamkhan (1969) Kanda (1968) Mukherjee et 01. (1966), Jost (1971) Sharma el al. (1967a1, Baker and Aslamkhan (19691, Kanda (1968) Kanda (1968) Kanda (1968) Baker and Aslamkhan (1969) Baker and Aslamkhan (1969) Mukherjee el al. (1966) Baker and Aslamkhan (1969) Mukherjee el al. (19661, Asman (1974) Mukherjee el al. (1966) Aslamkhan and Baker, (1969b) Mukherjee el al. (1966) Baker (1968), Baker and Aslamkhan (1969), Kanda (1968) Kanda (1968) Mukherjee et al. (1966) Mukherjee et al. (1966) Mukherjee el al. (1966) Rai (1966) Aslamkhan and Baker (1969b) Aslamkhan and Baker (196913) Aslamkhan and Baker (1969b) Kanda (1968, 1971)

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Aslamkhan and Hyder (1972) Randa (1968) Cocke et al. (1973) Muklierjee et nl. (1970) Kanda (1968) Aslnmkhan (1971) Kanda (1968, 1971) Knnda (1971)

Mansonia uniformis Orthopodomyia anopheloides Orthopodomyia signifera* Psorophora signipennis Toxorhynchites yamadi Tripteroides nrnnoides liranotaenia bimaculata Wyomyia smithii

2. Chromosome Morphology

The small size of the mitotic chromosomes and the relatively small proportional lengths of the arms makes it difficult to distinguish different kinds of karyotypes. Rai (1963) proposed that the ratio of the shortest chromosome to the two longer ones (I/II+III~ might give a better indication of the karyotype; this ratio has been generally used by authors describing karyotypes and is especially useful in distinguishing those species with rclatively long first chromosomes. One such species is Anopheles melas (Coluzzi and Sabatini, 1969b), in which the X chromosome is considerably longer than the X in other members of thc complex, the additional length presumably being heterochroniatic.

C. GAMETOGENESIS Detailed studies of spermatogenesis have been done in Aedes aegypti (Krafsur and Jones, 1967; Mescher and Rai, 1966; Bhalla, 1971b) ; Aedes dorsalis (Mukherjce and Rees, 1970) ; Culez tarsalis (Asman, 1974) ; Aedes albopictus (Jost, 1971; Smith and Hartberg, 1974) and Culez pipiens (Jost, 1971; Jost and Laven, 1971). All these studies agree in the apparent lack of prcpachytene stages, presumably due to the somatic pairing present. Three species of anophelines (maculatus willmori, fluviatilis, and splendidus) were observed by G. P. Sharma et al. (1970). Prepachytene stages are rare or absent. Anaphasc I bridges were seen in all species, more frequently in Anopheles splendidus, and may indicate the presence of heterozygous inversions. Somewhat more detailed observations (Narang et al., 1972b) of mitosis and spermatogenesis are available in Anopheles annularis, maculatus, subpictus, and tessellatus. Lack of prepachytene stages, positive heteropycnosis of the scx-bivalent during pachytene, 1 to 6 chiasmata per bivalent, and duplication by pachytene are common to all four species. A different approach to gametogenesis in mosquitoes (V. P. Sharma et al., 1970) used tritiated thymidine in Culen pipiens, followed by autoradiography to measure the total elapsed time from spermatogonia and spermatocyte to functional sperm. About 3-4 days are used to form the primary spermatocyte, about 1 day for the two meiotic divisions, and about 5 days for spermiogenesis.

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Abstracts of mitotic or meiotic studies have been published for several species: Anopheles fluviatilia Anopheles maculutus willmori Anopheles splendidus Anopheles stephensi Aedes aegypti Aedes albopictus Culex fatigans

Sharma et al. (1988b) Sharma et al. (1968f) Sharma et al. (1968~) Hafeez el al. (1973) Sharma et al. (1967d) ; Rai (1971b) Sharma and Toor (1970) Sharma et al. (1967a)

An excellent outline for studying the effects of radiation and related techniques on meiosis (Rai, 1968) also includes the genetic uses to be made of cytogenetic markers. Chiasmata frequencies vary in Aedes aegypti from 3.0 to 5.2 per cell (Ved Brat and Rai, 1973b), are random with respect t o arms, and tend to be distal. Duplication-deficiency heterozygotes in A . uegypti (Ved Brat and Rai, 1974b) usually form rings a t meiosis with a high (91%) proportion of the cells of one heteroeygote showing chiasmata in the interstitial regions. Chiasmata frequencies were higher in translocation heterozygotes (Ved Brat and Rai, 1974a) than in wild stock with standard chromosomes. A compound structural arrangement (McGivern and Rai, 1974) involves a translocation and an inversion. There is directed segregation a t meiosis I and a distorted sex ratio.

D. CYTOLOGY Although not specifically germane to this review, several investigations on what might be considered classical cytology are of interest. Modified synaptonemal complexes (Fiil and Moens, 1973) in Aedes aegypti and C . fatiguns have a relationship with the synaptic structures and the nuclear membrane. Details of spermiogenesis and fine structure of sperm (Breland et al., 1968) have been revealed by electron microscope studies. In Aedes dorsalis Mukherjee and Rees (1969) measured the duration of the mitotic cycle in brain cells, using high-resolution autoradiography. The GI period was about 75 minutes, DNA synthesis about 7 hours, Gz about 1 hour, and mitosis about 45 minutes. Cytochemical investigations apparently have been few (Kuenetsova, 1968; Tandon et al., 1971; Tandon and Bhattacharya, 1972). Tissue culture cell lines have been reported in Aedes, Anopheles, and various culicine species. These investigations obviously utilize systems that may be expected to clarify many phases of mosquito cytogenetics. (For an intro-

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duction to the recent literature, see Nichols et al., 1972; Weiss, 1971; Bianchi et al., 1972a,b.) IV. Evolution and Speciation

Relatively few papers have been published since 1967 that deal specifically with evolutionary syntheses or with specific evolutionary factors. Of course the chromosomal polymorphism in the anophelines and ethological and behavioral factors are clearly components of the evolutionary process. Ecological aspects of mosquito evolution are thoroughly treated by Mattingly (1971). The hybridization studies also shed light on speciation and isolating mechanisms. Barr (1974a) effectively relates the “new” concepts to classical taxonomy. A. SIBLING SPECIE^

AND

GENEFLOW

A review of the problem of sibling species (Coluzzi, 1970) emphasizes the lack of morphological variation in the anophelines and points out the obvious advantages of chromosomal identification, especially within the Anopheles gambiae complex. Lack of gene flow among three members of the subgenus Celliu (Narang et al., 1972a) was chiefly due to lack of insemination and fertilization. These hybridization data support the taxonomic and cytological data used to define the series groupings in the subgenus Cellia. The hypothesis of the Neotropical origin of the Anopheles maculipennis complex (Kitzmiller, 1966b) is probably invalid. The chromosomal similarities upon which this hypothesis was based are much wider than the A. maculipennis group, and are probably characteristic . the Aedes comof the entire subgenus Anopheles (Kitzmiller, 1 9 6 6 ~ )In munis group of species, three sibling species have been defined, communis, nevadensis, and churchillen& (Ellis and Brust, 1973). They differ in several minor morphological and physiological characters and are essentially reproductively isolated, although some small amount of successful hybridization is possible between A. communis and A . churchillensis. In the Palearctic Anopheles maculipennis complex, A . maculipennis (9.8.) and A. messeae may be distinguished by different puffing regions in chromosome 1, by a large rearrangement in 3R,and in the frequencies of individual heterozygous inversions in chromosomes 1 and 3 (Stegniy et al., 1973). The two species are reproductively isolated. A short but thoughtful paper (McClelland, 1967a) pursues some of the complex evolutionary problems that are involved in the Aedes aegypti eradication program. Immunodiffusion and immunoelectrophoresis studies (Cupp and

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Ibrahim, 1973) were able to differentiate all three forms of the Culez pipiens complex as well as their hybrids.

B. HYBRIDIZATION Hybridization studies are of primary importance in estimates of degrees of genetic proximity between species. I n mosquitoes this parameter is especially valuable because the technique of forced mating permits the observation of a wide spectrum of results in the laboratory, which may then be compared with observations from nature. 1. Anopheles

Hybridization within the A . gambiae complex (Davidson et al., 1967) has defined the sibling species of that complex. (For a comprehensive review, see Davidson et al., 1967.) Recently a sixth species within the complex, species D, has been isolated on hybridization and cytological evidence (Davidson and White, 1972 ; Davidson and Hunt, 1973). Before the advent of chromosomal identification, hybridization was the only way to identify the members of the complex (Odetoyinbo and Davidson, 1968). The hybridization of the various sibling species has also permitted precise definition of the extent of inversions and chromosomal identification (Coluzzi, 1966; Coluzzi and Sabatini 1967, 1968c, 1969b; Hunt, 1972a,b, 1973; White 1973b). The production of sterile males by hybridization within the A . gambiae complex has opened up a new potential control mechanism (Davidson, 1969a,b, 1970a, 1971a, 1972a, 1973b,c; Davidson et al., 1969, 1970; Davidson and Kitzmiller, 1970) (see also Section VIII and Davidson et al., 1969, 1970). Hybridization between species A and species B was used by Bryan (1968, 1972b) to study the results of consecutive matings. She found that whether a female lays fertile eggs or not depends on whether she has been first mated to a fertile or to a sterile male. Later experiments refined the information ; only males with well-developed and active accessory glands induce the monogamy. One of the practical uses of the chromosomal differentiation in A . gambiae is the identification of naturally occurring hybrids. White (1970b, 1971a,b) demonstrated asynapsis and two kinds of X chromosomes (A and B) in four females in Tanzania. Cox (1973c), using sterility and morphological criteria, inferred hybridization between A . melas and A and between A and B. I n introgressive hybridization situations, humid conditions appear to favor species A, drier conditions favor species B. The sex ratio is normal in crosses between A and B. Multiple hybridiza-

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tions (Davidson, 1970b) are possible. Assortative mating (Ramsdale and Leport, 1966, 1967) may be assessed by scoring of hybrids in nature, and hybridization (Ramsdale, 1966) may be used to identify species in mixed populations. Hybridization studies have also clarified the taxonomic status of a number of populations in the Anopheles punctulutus complex. These populations (Bryan, 1970a,b, 1972a, 1973a,b; Bryan and Davidson, 1967) fall into four groups, interfertile in each group, but each reproductively isolated from the others. Detailed studies of crosses (Bryan, 1973a,b,c) showed that of the 12 possible crosses among the four populations, three produced larvae that failed to survive. I n the other 9 crosses, adults were produced but both sexes were sterile, although a few exceptional individuals survived. Competition experiments with sterile hybrid males and normal males of one species (Bryan, 1973c) indicated that the addition of sterile males did not lower the insemination rate of females, but that the addition of sterile females did. I n Anopheles stephensi a number of crosses among geographic strains (Rutledge and Ward, 1970; Rutledge et al., 1970) were essentially interfertile, although reduced fertility was observed in some crosses. Similar results were obtained (Coluzzi et al., 1973a) in crosses among strains from Iran, Iraq, India, and Pakistan and by Zulueta et al. (1968) with strains from Iran, Iraq, and India. Interstrain fertility was also the case in A . albimanus (Keppler and Kitzmiller, 1969) in which 20 parental, 40 F,, and 40 F, crosses all yielded viable offspring. The same kinds of results were obtained with A . pseudopunctipennis (Martinez-Palacios and Davidson, 1967). Bianchi (1968~)combined hybridization studies and enzyme differences to reaffirm the validity of A . atroparvus and A . labranchiae as distinct species. The close chromosomal similarity between A . stephensi and A . superpictus (Coluzzi et al., 1970a) prompted crosses between these species (Coluzzi et al., 1971d). Anopheles superpictus females could be inseminated by A . stephensi males in laboratory cages but the reciprocal cross could be done only by forced copulation. In the former cross, hybrid larvae survived until fourth instar, but no viable pupae were obtained. I n the latter cross, embryonated eggs either failed to hatch or died as first-instar larvae. I n the North American species, hybridization experiments have been done in several groups. Crosses between A . crucians and A . bradleyi (Kreutzer and Kitzmiller, 1971a) produced adults of both sexes although in numbers lower than the controls. F, males were uniformly sterile with atrophied testes. F, females backcrossed to either parent showed reduced

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fitness, about 30% of the embryonated eggs failed to hatch, production of adults from eggs averaged about 376, and the number of adult males was reduced. Preliminary data (Kreutzer, 1971) indicate lack of gene flow between either Anopheles crucians or bradleyi and atropos, freeborni, and quadrimaculatus. Kreutzer and Kitzmiller (1969, 1972) also hybridized A. punctipennis and A. perplexens. When A . perplexens was used as the female parent no eggs hatched, although 61% were embryonated. In the reciprocal cross a few females were produced. Although the salivary gland chromosomes are quite similar in banding pattern, most regions were asynaptic in the hybrid larvae examined (Baker and Kitzmiller, 1965a). In crosses between the closely related A. pseudopunctipennis and A . franciscanus, (Smithson, 1972) no viable eggs were produced and only a few eggs contained early embryos. Presumably the isolating mechanism is sperm inactivation. Within the A. hyrcanus species group (Reid, 1965), Kanda and coworkers have attempted several crosses. Crosses of different geographical strains of A. sinensis (Kanda and Oguma, 1970a, 1972a,b) were fertile, but the hybrid chromosomes partially asynaptic. One strain of A. sinensis from North Hokkaido is reproductively isolated from all other strains (Oguma and Kanda, 1972). Similar intraspecific strain crosses with A. sinensis, A. sineroides, and A. lesteri (Kanda and Oguma, 1972a) were fertile, but species crosses showed hybrid inviability of the adults, although enough larvae were produced to demonstrate similar banding patterns (but with complete asynapsis) in salivary chromosomes. Hybrids in A. sinensis x A . lesteri crosses from Korea and Japan died as pupae. The larvae had asynaptic autosomes. Additional hybridizations have also been made between A. labranchhe and A. atroparvus (Coluszi and Coluzzi, 1969). Two related species, A . litoralis and A. subpictus have been crossed by the forced mating technique (Darsie and Cagampang-Ramos, 1973). Reciprocal crosses were successful (5246% hatch in the F,), but no F, males produced offspring. The only backcross tried, to A. litoralis males, gave about 35% hatch. Morphological patterns in the F, resembled A . subpictus. 8. Aedes

A series of crosses within the subgenus Stegomyh of the genus Aedes demonstrated the nature of the reproductive barriers that help isolate these species. A thorough study of the barriers to hybridization between A . aegypti and A . albopictus (Leahy and Craig, 1967) indicates that hybridization apparently does not occur between these species, in contrast to previous reports. At least five barriers are involved : mating behavior, structural

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incompatibility of male genitalia, sperm inactivation, reduced ovulation in A . albopictus females when crossed with A . aegypti males, and inviable embryos when produced. An excellent review of the earlier literature is given. The authors conclude that earlier reports of hybridization are probably due to contamination. Negative results were also obtained by Thomas and Leng (1973). In crosses between A . aegypti and A . mascarensis (Hartberg and Craig, 1968, 1970, 1973) viable, fertile offspring are produced, as known previously, but some indications of reproductive isolation are shown by preferential mating of A . aegypti X A . aegypti and by partial hybrid breakdown in backcrosses. Introgressive hybridization evidently does not occur (Hartberg and McClelland, 1973). The three color forms of A . aegypti (type, fomnosus and queenslandensis) in Tanzania (Hartberg, 1969) appear to be simple populational polymorphisms rather than taxonomic entities. In crosses between A . woodi and A . simpsoni (Hartberg, 1970, 1972) copulation by means of forced mating occurs readily, sperm are transferred with difficulty, some females will produce eggs and the adults produced are fertile in some backcrosses. The adults from these backcrosses suggested single-locus control of morphological differences that are used to separate the species. Attempted crossings of each species with A . aegypti were unsuccessful. Males of A . albopictus mate readily with females of A . polynesiensk (Gubler, 1970a) ; sperm are transferred, but no embryonated eggs are produced. The reciprocal cross does not result in sperm transfer, and the barrier is presumed to be ethological. The A . albopictus males are quite aggressive, therefore the successful copulations might have been a t least partially due to the small cage size. I n large-cage trials (Ali and Rozeboom, 1971a) the A . albopictus males were also successful in inseminating A . polynesiensis females. The same authors (Ali and Rozeboom, 1971b) extended these data, crossing two strains of A . albopictus, three of A . polynesiensis, one each of A . pseudoscutellaris and a member of the A . scutellaris group from Bangkok. The A . albopictus strains were interfertile, as were the A . polynesiensis, but subtle strain differences exist in A . albopictus as indicated by different insemination success with different strains of A . polynesiensis. The A . albopictus males did not inseminate either A . scutellaris or A . pseudoscutellaris. The A . polynesiemis females are receptive to their own males even when large numbers of A . a2bopictus males are present (Ali and Roaeboom, 1973), perhaps owing to earlier sexual receptivity. In Aedes polynesiensis, strains of different geographic origin are generally completely interfertile, but some reduced hatch takes place in certain ones. I n reciprocal crosses among strains from Tahiti, Taiaro, and Samoa (Tesfa-Yohannes, 1973) all gave fertile offspring, but the Tahiti

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female X Samoa male cross was less fertile than the others. When these three strains were crossed with Aedes scutellaris malayensis (TesfaYohannes and Rozeboom, 1974) five of the six possible crosses produced nonhatching eggs, but the A . scutellaris female x Samoa male cross gave a few hatching eggs. The F, males from this cross were sterile, but F, females could be backcrossed to A. scutellaris males and subsequent backcross generations were fertile. Evidently some introgressive hybridization is possible between some populations of these species. Hybridization was also possible between Aedes polynesiensis and an autogenous species of the A . scutellaris group from Tafahi Island, Tonga (Hitchcock and Rozeboom, 1973). The T o X P 6 was successful with 67% hatch, but the reciprocal was not. The offspring were fertile through the F,, but had reduced fertility in successive generations. The backcrosses were successful, but with reduced fertility. Extensive crosses among several Mediterranean populations of the Aedes mariae complex (Coluzzi and Sabatini, 1968e) demonstrated vigorous F, hybrids, but the F, males were sterile, largely owing to pronounced disturbances during spermatogenesis. Evidence for a precopulatory mechanism was later found (Coluzzi and Bullini, 1971). The tree-hole breeding species Aedes triseriatus and Aedes hendersoni (Truman and Craig, 1968; Craig, 1970b) were successfully colonized and hybridized using forced mating. The F, males of A . triseriatus females x A . hendersoni males were infertile, with abnormal genitalia. Backcrosses suggest monofactorial inheritance for several taxonomic characters. The closely related species Aedes sollicitans, mitchellae, nigromaculis, and taeniorhynchus have been successfully hybridized. Crosses between A . mitchellae and A . sollicitans (O’Meara et al., 1974) succeeded in both directions, both F, males and females were fertile when backcrossed (no F, x F, crosses), and examined males had motile sperm. Hybrid larvae resembled A . sollicitans, but hybrid adults were intermediate between the two species. Preliminary data also indicate successful hybridization between A . sollicitans and A. taeniorhynchus. Almost complete gene flow is indicated in crosses between A . sollicitans and A . nigromaculis (Fukuda and Woodward, 1974a). Crosses were successful in both directions, and both sexes were fertile in the F, and produced viable fertile F, and backcross progeny. Again the larvae resembled the female parent in the backcross generations, but adults were intermediate in taxonomic characters. Presygotic mechanisms must account for most of the isolation since hybrids are evidently rare in natural populations. Crosses are also possible between A . taeniorhynchus and A . nigromamlis (Fukuda and Woodward, 1974b). Hybridization experiments with 4 taxa within the Aedes atropalpus

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group, (Brust, 1974) supported by scanning electron micrographs indicated that Aedes atropalpus is a distinct species but that Aedes epactius, perichares and nielseni are closely related genetically and probably should be grouped as one species, epactius. 3. Culex In Culex fatigans, colonies from six localities in Kenya and Tanzania (Magayaka and White, 1971, 1972) were interfertile and showed no evidence of incompatibility. Other crosses involving African C . fatigans, however, did show incompatibility (Laven, 1969c,d; Eyraud and Mouchet, 1970). Crosses between C. molestus and C. futiganr (Dobrotworsky, 1967) in Australia are generally successful, as are crosses among different geographical strains of C. molestus. Both species appear to be reproductively isolated from C. australicus. In western Australia four members of the complex (Paterson, 1972) coexist in several places, but produce few natural hybrids. Probably the most extensive series of crosses involving Culex fatigans, pallens, and molestus has been done by Sasa et al. (1966). Four geographical colonies of C . fatigans, four of C. pallens and three of C. molestus were crossed in various combinations. Within each taxon, bidirectional fertility was the rule, and crosses involving C. fatigans and either of the other forms were compatible either in one or in both directions. However, males of C. molestus were highly incompatible with females of either C. fatigans or C. pallens, but the reciprocal crosses were partially fertile. The F, egg rafts contained mostly embryonated but unhatched eggs, with some “exceptional” individuals. Crosses from eight widely spaced localities in Brazil (Espinola and Consoli, 1972, 1973) were fertile in all combinations tested. The authors suggest that in these populations of C . fatigans the usual cytoplasmic incompatibility agents do not exist. [For a summary of the occurrence and distribution of the Culex pipiens complex, see Barr (1967), and for a n excellent description of morphological and biological characteristics in C. pipiens in the USSR,see Vinogradova (1965) .] An incisive review of the systematics of the complex (Mattingly, 1967a) stresses the dynamic aspects of speciation mechanisms. V. Cytoplasmic Incompatibility

The generally accepted hypothesis for the incompatibility of strains within the Culex pipiens complex is that of Laven, who proposed that the observed results are due to cytoplasmically inherited factors (Laven,

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1957a,b, 1967b,c). The mechanism is evidently one in which the entrance of the sperm stimulates ovogenesis and parthenogenetic haploid development of the egg to a certain stage. The sperm itself is blocked, karyogamy does not occur (Jost, 1970a,b, 1971, 1972). Polyspermy is rare (Jost, 1970b), and there is a suggestion that mitochondrial DNA might be involved (Jost, 1971). Additional surveys of African strains (C. fatigans) produced the classical kinds of incompatibility in crosses (Laven, 1966a,b,c, 1969c,d; Eyraud and Mouchet, 1970; Hamon et al., 1972; Subra et al., 1968; Subra, 1972). Tests with 19 strains of Asiatic C. fatigans (Thomas, 1971) were completely compatible, as were strains from Brazil (Espinola and Consoli, 1972, 1973). Strains from Bahia, Brazil, were compatible with Indian strains of C. fatigans (Smithson and Conceiciio, 1973). The data of Sasa et al. (1966) strongly suggest cytoplasmic factors in C . fatigans, C . pallens, and C. molestus crosses (see Section IV, B) . An alternative hypothesis (McClelland, 1967c) supports nuclear rather than cytoplasmic inheritance of incompatibility. This rather complicated hypothesis requires a t least three alleles t o condition the cytoplasm and invokes meiotic drive for two alleles. Recently, two other hypotheses have been advanced. French (1970, 1971, 1973a,b,c,d) believes that randomly segregating cytoplasmic genes determine the plasmatypes and that nuclear genes are not involved. French’s evidence strongly indicates that the molecular bases for cytoplasmic male sterility in Culex are polymorphic differences in mitochondria. The second new hypothesis (Barr and Yen, 1972; Yen and Barr, 1971, 1973, 1974) suggests that a Rickettsia-like symbiont, Wolbachiu pipientis, is the causative agent. This symbiont, which appears to be a universal symbiont of C . pipiens, can be killed with tetracycline. Males of incompatible strains which had been “cured” with the antibiotic no longer displayed incompatibility reactions. This hypothesis must still explain, however, the origin of the many different incompatible strains known. The Yen-Barr hypothesis has received striking support from studies with the Australian members of the complex (Irving-Bell, 1974). Electron micrographs showed Rickettsia-like symbionts in Culex pipiens, fatigans, molestus, and pallens, but not in the two native members of the complex, australicus and globocoxitus. Laven’s data are not incompatible (to coin a word) with those of Yen and Barr, and both Jost and French suggest a mitochondrial (DNA?) mechanism. Perhaps an episome or episomelike mechanism is possible in higher organisms. There are many enticing parallels between the situation in Culex and the F+ factor in the Hfr bacterial strains. Barr (1970) advanced a hypothesis of “partial compatibility” based upon lack of complete incompatibility in certain crosses in California.

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Data from Japan (Hubert et al., 1971), Brazil (Rai, 1969a), and Africa (Subra, 1972) appear to support Barr’s concept. The age of the males is a factor, with older males being “more compatible,” producing more partial compatible matings and more hybrid larvae (Singh et aZ., 1974b). VI. Genetic Manipulation

With the availability of chromosomal aberrations, it was natural that they could be used in genetic manipulations of various kinds (Rai, 1967a; Rao, 1972). In addition to genetic control possibilities (see below), they have been used in more classical manipulations, such as crossover suppression in Aedes aegypti (Bhalla, 1970a, 1972). The sex-linked loci m, bz, and w were affected on both M and m chromosomes with varying degrees of suppression of crossing-over according to the suppressor strain used. Although no direct cytogenetic evidence is yet possible (poor salivary chromosomes), the presence of anaphase bridges and fragments suggest heterozygous paracentric inversions. One of these crossover suppressors reduces recombination in females, but a t the same time enhances recombination in males. This system, COSES (Bhalla, 1971a), contains a small reciprocal translocation involving chromosomes 1 and 2 and a paracentric inversion on chromosome 1. Detailed studies showed that COSES suppresses crossing-over to the right of bronze in females, but enhances crossing-over to the left of that locus. In males i t enhances crossing-over to the right of bronze and sex, but the region to the left remains unaffected. COSES evidently also enhances recombination in linkage group 3. Sex-linked paracentric inversions (Bhalla, 1970b) were also used to formulate a system for the detection of sex-linked lethals. An inversion-dominant marker strain (Sakai and Baker, 1972) similar to the ClB system in DrosophiZa has been synthesized in Culex tritaeniorhynchus and used for the detection of sex-linked recessives. VII. Behavior Genetics

It is obvious that the behavior of mosquitoes had profound effects upon disease transmission. There is no lack of literature in this area, but few hard facts or permissible generalizations. Ethological considerations are also obviously important in the evolutionary process in mosquitoes, and ultimately must be responsible for most precopulatory isolating mechanisms. The genetic basis for specific behavioral characteristics or for

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mechanisms that affect evolutionary processes is poorly known. Since the able and comprehensive reviews by Mattingly (1967b, 1969) relatively few papers have appeared that deal specifically with the genetics of behavioral mechanisms. Perhaps best documented is the important area of behavioristic responses to contact with insecticides. Selection is possible in Anopheles atroparvus for escape from tubes impregnated with DDT in Risella oil (Gerold and Laarman, 1964, 1967, 1968). With strong selection pressure, “escape-prone” and “non-escape-prone” strains are differentiated after about 10 generations, and differ principally in two behavior patternsflight time and the ability to pass through an opening (Gerold, 1968a,b, 1969). The D D T irritates both strains to fly, but the irritability releases two different sets of behavior patterns in the two strains (Gerold, 1969). The details of these experiments coupled with an extensive review of the literature and a critical analysis is given by Gerold (1970). Tests with A . gambiae, species A and B, (Gerold, 1973) show clear genetic differences in takeoff time, flying time, and in the ability to pass through a n opening. “Bite-and-run” individuals (which might be selected for by D D T residual sprays) differ genetically from other females in the same population. Flight activity patterns in A . stephemi appear to have a genetic basis (Jones and Gubbins, 1973; Jones, 1974; see also Elliott, 1968, 1972.). Most of the rest of the recent literature deals with Aedes aegypti. Spontaneous movement, the tendency to oviposit on damp surfaces rather than open water, and speed of vertical movement of the larvae (Schoenig, 1967, 1968, 1969) are genetically controlled and differ in laboratoryselected strains. The small antenna mutant (Dunn and Craig, 1968) reduces mating competitiveness in males and host-finding ability in females. Evidence for a sexual pheromone and a tarsal receptor (Nijhout and Craig, 1971) appears plausible as a precopulatory mechanism between A . aegypti and A . albopictus. Interesting observations on mating behavior (Hartberg, 1972; Jones and Pilitt, 1973; Jones, 1973b) probably indicate an underlying genetic basis. Stenogamy in A . taeniorhynchus (O’Meara and Evans, 1974) is dependent upon female behavior. VIII. Genetic Control

A. GENERAL With the development of resistance to the commonly used insecticides, the search for alternative methods of insect control has become of great

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importance (Rabb and Guthrie, 1970, Pal and Wharton, 1974). Various kinds of biological control have been proposed for a number of vector and pest insects (Williams, 1967), and genetic control of one sort or another (Davidson, 1974) is an important component. The first attempts a t genetic control naturally followed the screwworm control model (Knipling, 1967) but had deep roots in radiation biology (Lachance, 1967; LaChance et al., 1968) and in classical population genetics (Boesiger, 1971a). In the last 7 years a voluminous literature has accumulated on subjects connected with the genetic control of mosquitoes. Much of this has been published in the proceedings of symposia, which contain resum& of other types of biological control as well. Much as been written about the theoretical possibilities of genetic control, and considerable background work has been done on radiation dosages, lethality, and sterility. Inversions and translocations, the production of sterility, and the associated chromosomal changes due to chemosterilants all have been popular subjects. Some of the theoretical data have been utilized in laboratory experiments designed to test the hypotheses of eradication or population reduction, some computer models have predicted laboratory or field results under varying conditions, and a few preliminary field experiments have been conducted. Perhaps here is the place to emphasize the role that the World Health Organization (WHO) has played in the support of genetic control, and indeed of vector genetics in general. The WHO has been instrumental in promoting the idea of genetic control through symposia, convocation of scientific groups, financial support, and individual encouragement (World Health Organization, 1966, 1967a,b, 1968). The publications “Vector Genetics Information Service” and “Information Circular on Insecticide Resistance, Insect Behavior and Vector Genetics” have been most valuable in coordinating different research programs and making the literature easily available. The division of Vector Biology and Control has been especially active in the encouragement of activities in this area (Curtis, 1973, Pal, 1971, 1972, 1973, 1974a,b,c; Pal and LaBrecque, 1972; Pal and Lachance, 1974; Pal and Wright, 1968; Smith, 1967b,c,d, 1972). B. INTERNATIONAL SYMPOSIA A number of international symposia have dealt with biological control of insects, especially mosquitoes. The first of these in Rome, in 1968, sponsored by the Accademia Nazionale dei Lincei, included genetic control, hormonal systems, and integrated control systems among other

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topics. Craig (1969) summarized the prospects for genetic control. A major international symposium at Walter Reed Army Institute of Research (Ward and Scanlon, 1970) covered vector control, malaria control, resistance, systematics, sibling species in Anopheles (Coluzzi, 1970), control methodology, repellents, genetic control (Davidson and Kitzmiller, 1970), hormonal control (Craig, 1970a), parasites of anophelines, problems in population measurement, epidemiology as applied to malaria, and computer simulation. This symposium, easily available in the literature, will be of interest to all who are interested in mosquito genetics. Another international symposium, held at the University of Montpellier, France, in 1969, covered many aspects of the “Lutte Biologique contre les arthropodes h6matophages” (Annales de Parasitologie Humaine et Compare’e 46, 3 ) . Of especial interest are discussions of population genetics in relation to vector control (Boesiger, 1971b), sterilization procedures (Mouchet, 1971), problems linked to mass rearings (Coluzzi, 1971), a general review of genetic control mechanisms (Laven, 1971a), a resumk of the principles of cytoplasmic incompatibility in Culex and a description of the Okpo experiment (Laven, 1971b), a description and critical analysis of the Anopheles gambiae hybrid male release in Upper Volta (Davidson, 1971b) and the role played by WHO (Pal, 1971). An American symposium sponsored by the Government of El Salvador, the Pan American Health Organization, and the Center for Disease Control, USPHS, took a hard look at malaria in the Americas and covered many aspects of this disease, all of which are of interest to mosquito geneticists (Scholtens and NBjera, 1972). Session 5 was entirely devoted to vector biology and biologic control and contained papers among others on vector behavior (Elliott, 1972), genetic control (Kitzmiller, 1972), and chemosterilants (Weidhaas, 1972). Session 6 on chemical control, included a study of resistance in Anopheles albimanus (Georghiou, 1972). The papers presented a t a symposium held a t the 14th International Congress of Entomology, Canberra, have been compiled and edited by Pal and Whitten (1974). Part 1 deals with genetic control methods for insects of agricultural importance, and part 2 with genetic control of insects of public health importance. Chapters that deal with mosquitoes are as follows: The WHO/ICMR programme of genetic control of mosquitoes in India (Pal, 1974) ; Incompatibility in Culex pipiens (Barr and Yen, 1974) ; The current status of genetic methods for controlling Aedes aegypti (Rai et al., 1974) ; and Genetic studies on Culex tritaeniorhynchus (Baker and Sakai, 1974). Another major symposium held a t Edmonton in 1972 developed the theme of biting fly control and environmental quality (Hudson, 1973). The entire symposium is of interest; especially applicable to genetics are

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articles on juvenile hormone analogs (Strong, 1973) and on genetic control (Rai, 1973).

SCHEMES C. CONTROL A number of papers review genetic control of mosquitoes in a general way, many of them for nontechnical consumption (Baker, 1971; Craig, 1967b, 1968, 1969; Davidson, 1972a, 1973c; Hamon, 1970; Kitemiller, 1972; Knipling et al., 1968; Laven, 1966a, 1968, 1969, 1970, 1972a; McClelland, 1974; de Meuron-Landolt, 1972; Patterson, 1972; Patterson and Lofgren, 1968; Rai, 1971a,b; 1972a,b; Smith, 1971; Smith and von Borstel, 1972; Rukavishnikov, 1968, Weidhaas, 1972; White, 1967a). The Journal of Communicable Diseases (New Delhi) devoted the June 1974 number (Vol. 6, No. 2) to genetic control of mosquitoes. Several other papers also review mosquito genetics and include details of specific procedures or experiments. Rai (1969b; Rai et ul., 1974) reviews previous field trials, assesses recent promising developments, and calls attention to alternatives to male sterility, particularly emphasizing translocations. Having securely established cytoplasmic incompatibility, Laven (1969b, 1970) turned to translocations as an alternative method of genetic control ; Laven outlines briefly the theoretical possibilities and further proposes integrated control methods (Laven and Aslamkhan, 1970) for Culex pipiens, advocating a combination of cytoplasmic incompatibility and translocations. A critical appraisal of the role of genetics in pest management (Whitten, 1970) tilts a few sacred cows and also outlines a sophisticated system for combining translocation sterility and insecticide resistance and susceptibility. Whitten also includes a thoughtful discussion of karyotype evolution, fitness, displacement, and genetic manipulation. The paper contains an extensive bibliography. The same concepts are presented with additional sophistication (Whitten, 1971a,b), including a mathematical treatment of fitness, the effects of multiple translocations, a discussion of frequency-dependent phenomena, and the possibilities of compound chromosomes. Details of theoretical possibilities with compound chromosomes (Foster et al., 1972) include schemes for recovering compound chromosomes. The authors express optimism that compound chromosome schemes may indeed be feasible in many vector species. An outline of theoretical possibilities in one such species, Aedes aegypti (Rai and McDonald, 1972), gives considerable data on production and transmission genetics of translocations as well as competitive mating tests and computer simulation runs.

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1. Sterilization

The production of sterile males (and/or females) has been the immediate goal of studies aimed at eventual genetic control. Most experiments designed to induce sterility have been done with radiation or with chemosterilants (Lachance, 1967; LaChance et aZ., 1967, 1968; Byrdy, 1972). A popular review of the subject (White, 1967a) and a brief but systematic account (Mouchet, 1971) are available. The latter article contains an extensive bibliography. A Russian review (Zakharova, 1969) covers sterilization by chemicals and by radiation. The most complete reviews of induced sterilization are those of LaBrecque and Smith (1968), Stiiben (1969), Proverbs (1969), and Campion ( 1972) ; see also Berryman (1967). a. Aedes aegypti. Chemosterilization of pupae (White, 1966) with an alkylating agent was possible a t dosages from 100 to 2000 ppm for 24 hours. Adults survive well for a t least 2 weeks, and young males were at least as competitive as untreated males and sometimes hyperactive (White, 1970a). Selection with a nonalkylating chemosterilant (Hempa, hexamethylphosphoric triamide) at 1280 ppm induced chromosomal changes in larval brain cells (George and Brown, 1967), but the percentage of sterility decreased for five generations, then increased in the F,. Inbreeding and selection for recessive sublethals might account for the change. Hempa-sterilized males (Grover and Pillai, 1970a) were fully competitive. Screening tests with 14 aziridinyl compounds (Madhukar et al., 1971a; Mohiuddin and Qureshi, 1973) and 9 phosphoramides (Madhukar et al., 1971b) indicated that the alkylamino compounds were more active larval and pupal sterilants than the alkoxy compounds and that the phosphoric acid amides were more effective than the amides of phosphorothiotic acid in larvae, but the reverse was true for pupae. Toxicity varied greatly, and both sexes were affected. Other compounds have also been screened for antifertility effects (Naqvi et al., 1973). A possible means of bypassing the noncompetitive nature of radiated males might be by radiating in nitrogen rather than air (Hallinan and Rai, 1973). Nitrogenirradiated males were at least as competitive in cage experiments as normal males, but air-irradiated males were not. Low dosages of beta radiation (Guthrie and Brust, 1971) sterilized both males and females, but ultraviolet rays are not effective (Riordan, 1969). In addition to the sterilizing effects, the chemosterilants and ionizing radiation affect other tissues as well (Sharma and Rai, 1969) and induce often drastic histopathological and developmental effects. Apparently minimal function is possible to permit survival. Larval treatment with

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sterilizing doses of Apholate and Hempa (Madhukar et al., 1970) inhibited DNA synthesis in testes of Aedes aegypti, and Apholate decreased ovarian alkaline phosphatase activity (Turner and Maheswary 1969). In transplants of normal ovaries into chemosteriliaed females, and vice versa (Rai and Sharma, 19711, the data indicate that the sterilization effect is in the oocyte itself rather than a general effect exerted by the humoral factors. Judson (1967) noted inhibition of ovarian development after treatment with Apholate. Ionizing radiation of A . aegypti a t dosages ranging from 500 to 32,000 R (Asman and Rai, 1972) had effects upon developmental processes and fitness components in proportion to dosage. Affected were egg hatch, growth, viability, longevity, and cell division, as well as fecundity and fertility in both sexes. Apholate resistance in A . aegypti (Seawright, 1972) is evidently a polygenic trait with a marked paternal influence. Sterilization is also possible (J. W. Patterson, 1971, 1972) with juvenile hormone mimics. b. Culez. Preliminary experiments with alkylating agents and nonalkylating chemosterilants (Hempa and Hemel) (Grover et al., 1967; Mulla, 1968; Pillai and Grover, 1968) confirmed the results of earlier investigations (Wattal et al., 1970) showing that aquatic stages are easily sterilized, with pupae the stage of choice (McCray and Schoof, 1970). Further screenings with 8 alkylating agents (Pillai and Grover, 1969a) showed different toxicities in larvae and pupae and induced morphological abnormalities. Nontoxic doses of Apholate produced total sterility in larval treatment, but Tepa was better as a pupal sterilant. Sterility was long lasting, but some recovery was noted in Apholate-treated males (Pillai and Grover, 1969b). Similar screenings with phosphoramides and s-triazines (Grover and Pillai, 1969) include the interesting result that some compounds produced more sterility in treated females than in treated males. These chemosterilants induced dominant lethal mutations in C . fatigans (Grover and Pillai, 1972) affecting different life stages as did Tepa and Hempa (Hafez et al., 1970a). Thiotepa (Sharma et al., 1972n, 1973) was an effective sterilant for C . fatigans. Mass sterilizing techniques (Boston et al., 1970; Patterson et al., 1971) were effective. Metepa and Hempa produced sterile adults (Saito and Hayashi, 1967, 1968) by the use of baits and also by contact with sprayed residues. Metepa gave high sterility rates at 0.1 and 0.01%, but Hempa only at 0.1%. Contact with Hempa also required higher dosage, but had a longerlasting effect. Tepa-sterilized males (Patterson et al., 1972b) were equally competitive with normal males. Chemosterilized males, irradiated males, and cytoplasmically incompatible males show equal dispersal rates in the field, but the cytoplasmic strain (D3) had a slightly lower daily survival rate (Rajagopalan et d.,1973).

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Chromosome damage in somatic cells and in gonadal and embryonic tissues (Grover et al., 1972, 1973) are cited as the causes of chemically induced sterility in Culez fatiguns. Various types of chromosomal changes evidently cause embryonic death. The chemosterilants also appear to inhibit DNA synthesis, especially in testes (Grover et al., 1971). Treatment of C. tarsalis with gamma rays (Darrow, 1968) produced sterility in both sexes at dosages from about 12.5 to 15 Kr. Using C. fatigans, treatment of adults with Apholate dust (Das, 1967; Smittle, 1968; Smittle et al., 1968) was a more effective sterilant than gamma irradiation applied to pupae, and treated males were equally competitive. X-rays and gamma rays were equally effective (Smittle et al., 1970). Eggs were most sensitive to gamma rays (Koshy and Singh, 1970; Smittle et al., 1971a; DeOliveira et al., 1969) larvae were less sensitive, and little mortality ensued after irradiation of pupae or adults. Dosages of 5000 R to 20,000 R produced high, but not complete, sterility in males. Comparisons with fast neutrons (Smittle et al., 1971b) indicate that fast neutrons are about 4-7 times more effective. Gamma radiation a t 5000 R (Sharma et al., 1972b, 1973) produced sterility, and loss of vigor in C . fatigans males and in C. molestus (Abdel-Malek and Ahmed, 1972a,b). Treatment of young males or pupae caused less inhibition of vigor (Sonoda, 1972). X-rays, thalidomide, and PMS (Jost and Amirkhanian, 1971; Amirkhanian, 1970, 1972, 1973b,c) all produced dominant lethals and semisterility in C. pipiens. Gamma rays up to 5000 R induced dominant lethals in C . pipiens (Abdel-Malek and Ahmed, 1972a,b,c, 1973) with highest effect in the egg stage. Autochemosterilization is possible. A modified CDC light trap, treated with Tepa was able to sterilize both males and females of C. fatigans (Grant et al., 1970). Best results were obtained when the trap was baited with a baby chick (see also Sacch et al., 1971 and Seawright et al., 1971). c. Anopheles. Early experiments with Anopheles labranchiae (D’Alessandro et al., 1966; Sandesco et al., 1968) suggested that the observed sterility was due to changes in ovaries rather than in testes. In A . pharoensis, radiation from 500 R to 7000 R (Abdel-Malek et al., 1966) mainly affected total egg production and hatchability, which fell below 1% a t 5000 R and above. Lethals induced by irradiation (Tantawy et al., 1966) were proportional to dosage, eggs were most sensitive to irradiation, and embryos from parents which had been treated in the pupal stage died mostly as embryos. A dosage of 12,000 R (Abdel-Malek et al., 1967a) completely sterilized both sexes, and with labeled 32Pit was determined (Tantawy et al., 1967a,b; Abdel-Malek et al., 1967b) that laboratory males were just as efficient, in cage experiments, in inseminating wild

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females as were wild males. Similar results were obtained in A. albimanus (Ali and Rozeboom, 1972) in which dosages from 2000 R to 8000 R caused marked reduction in egg production by treated females and increasing sterility, with dosage, in treated males. Anopheles pharoensis can be sterilized with Apholate, Thiotepa, Tepa, and Hempa (Hafez et al., 1969, 1970a,b, 1971, 1972) and A . stephensi with sulfaquinoxaline (Venters, 1973). In A. albimanus, P,P-bis ( l-aziridinyl) -N-methylphosphinothioic acid is an excellent sterilant (Lofgren et al., 1972), is efficient in low dosages, is stable (Seawright et al., 1974), and is highly biodegradable (Seawright et al., 1973). Hempa and Thiotepa were effective sterilants in A. labranchiae (Sandesco, 1967a,b). 2. Chromosomal Rearrangements Radiation-induced chromosomal aberrations with attendant reduced fertility have been produced in abundance in several species (Dennhofer, 1 9 7 4 ~ )Most . of these were produced in studies aimed a t genetic control, others as a by-product of other investigations. Many have been useful in classical genetic investigations, such as linkage group-chromosome correlations and linkage-crossover studies. As an indication of the popularity of fields associated with genetic control, when this subject was last reviewed (Kitzmiller, 1967) three investigators and six papers were cited; this review cites more than fifty papers that deal primarily with inversions and translocations, not even including those utilizing aberrations in control experiments. This has been a natural reaction to the increasing difficulty with insecticidal control, but undoubtedly reflects the tendencies of granting agencies to provide support for this work. Most of the aberrations have been produced by X-rays or with a cobalt source, with dosages ranging from 2000 to 8000 R. Translocations, pericentric and paracentric inversions have been reported, but only one deficiency and no duplications. An excellent review of chromosomal mechanisms of sterility (Dennhofer, 1974a) related these mechanisms to vector control and Dennhofer (1975a) suggests that a single locus with multiple alleles controls the preferential segregation responsible for translocation sterility. a. Translocations. The bulk of the research has been done in Aedes aegypti by the dynamic group of investigators at Notre Dame, headed by Professor K. S. Rai, the pioneer investigator in this area. Preliminary studies (Asman and Rai, 1966; Asman, 1967; Rai and Asman, 1968) showed that A . aegypti males with translocations were from 50 to 80% sterile. Crossing-over in a stock heterozygous for two translocations produced a double translocation, resulting in a “new” karyotype with two long chromosomes and one very short one (McDonald and Rai, 1970d).

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A similar “new” karyotype was obtained in Culex tritaeniorhynchus owing to a pericentric inversion (Sakai et al., 1971a). Enhanced crossingover appears to be another result of the system. Study of these two translocations, plus the double heterozygote (McDonald and Rai, 1970a,b) and the linkage group-chromosome correlations, resulted in the renumbering of the A . aegypti chromosomes. Detailed cytogenetic analysis of these translocations (Rai et al., 1970a,b) shows that one of the translocations has breakpoints very close to the sex locus on chromosome 1, and another close to the spot locus on chromosome 2. The other breakpoints are close to red on chromosome 1 and to black tarsi on chromosome 3. A computer simulation of release strategies (McDonald and Rai, 1970c, 1971) under varying assumptions indicated the promise of translocation releases. Reviews of the possibilities of genetic control by means of translocations (Rai and McDonald, 1971, 1972) expressed cautious optimism that these aberrations might be useful in field tests. Crosses with appropriate marker stocks (Lorimer et al., 1972a,b) produced translocation homozygotes for one sex-linked and one autosomal rearrangement. These homozygotes were fully viable although they produced fewer viable off spring than did the controls. The investigators attribute this reduced fertility to the low fertility and high variability of the mutant stocks in which the translocations were produced. Certainly translocations offer an intriguing new dimension for control work (see below Culex). Seven sex chromosome-autosome translocations (Bhalla, 1973b) show fertility ranging from 18 to 71%. A double translocation heterozygote has been reported (Uppal et al., 1974). I n Culez pipiens, most of the work has been done by Professor Hannes Laven, a recognized leader in the field of sterility and control mechanisms. First experiments with 4000 R (Laven, 1969a) produced many lines with reduced fertility; in 7 of these lines the reduced fertility was transmitted only by males, with sterility from 26 to 82%. Cage experiments with two lines showed virtual elimination of the populations after 5 and 8 generations, and, perhaps more significantly, indicated that translocation males were more competitive than normal males. Further data were added by Laven and Jost (1971) and by Laven et al. (1971c), showing the production of dominant lethals in relation to dose and the sterility produced in males. Semisterility can also be produced in C . pipiens with chemosterilants and mutagenic chemicals (Jost and Amirkhanian, 1971; Amirkhanian, 1972, 1 9 7 3 ~ )Translocations . have also been used (Bhalla et al., 1975; Dennhofer, 1972) in linkage groupchromosome correlations. In Culez tritaeniorhynchus, radiation of 3000 R from a s°Co source produced 46 translocations (Sakai et al., 1971b), most of which involved the marked sex chromosome. Detailed sterility figures and B puzzling in-

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crease in crossover values in certain regions of the sex chromosome (cf. also McDonald and Rai, 1970c) are reported. Reirradiation of translocation stocks (Sakai, 1972; Sakai et al., 1972b) produced double translocation heterozygotes, with sterilities as high as almost 90%. Translocation semisterility was also produced (Selinger, 1972) in this species with X-rays. The most detailed report of translocations in Anopheles (Rabbani and Kitzmiller, 1972b) lists and illustrates six translocations in A . albimanus; autosome-autosome, X-autosome, and Y-autosome with sterility values. Preliminary reports of translocations in A . gambiae (Krafsur, 1972; Hunt and Krafsur, 1972; Krafsur and Davidson, 1973; Akiyama, 1973) and in A . stephensi (Aslamkhan and Aaqil, 1970) stress dominant lethals and sterility values. Without question, considerable impetus was given to translocation research by the timely paper of Serebrovsky (1968). b. Inversions. Paracentric inversions have confirmed the suppression of crossing-over in mosquitoes. Using X-rays Bhalla (1970b) produced two sex-linked inversions in Aedes aegypti and measured the reduction of crossing-over and the increase in sterility. Combining a paracentric, sex-linked inversion with a translocation, Bhalla (1971a) was able to demonstrate both suppresssion and enhancement of crossing-over, depending on the region of the chromosome. Later experiments (Bhalla, 1973a,b) produced eleven sex-linked paracentric inversions, covering most of the known genetic length of chromosome 1. Using gamma radiation, McGivern and Rai (1972) produced an autosomal paracentric inversion involving linkage group 2 of A . aegypti, with associated reduction in fertility in males. Increased crossing-over was noted in group 1 and group 3 markers. In Culex pipiens (Tadano and Kitzmiller, 1969) chromosomal changes associated with breakages, dicentric bridges, and fragments followed treatment with Tepa. Similar evidence (Kuzoe et al., 1966) suggests inversions in A . aegypti and C . fatiguns. Dennhofer (1972) analyzed the eight translocations and one pericentric inversion produced by Laven (1969a,b), Laven et al. (1971d), Laven and Jost (1971), and Jost and Laven (1971). Both inversions and translocations have been used in C. tritaeniorhynchus to correlate linkage groups with chromosomes.

D. LABORATORY TESTS After the establishment of the theoretical bases of genetic control, the next step was laboratory tests. Only limited success was obtained in Culex fatiguns (Patterson et aZ., 1968a,b) with Apholate-sterilized males in a large outdoor cage although males may be competitive (Patterson et al., 1972a). The next test, with C. pipiens (Laven, 1969a) involved transloca-

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tions. Cage populations were dramatically reduced after five generations with the addition of adjusted numbers of translocation males. With Aedes aegypti, cage experiments with genetically marked translocation stocks (Rai and McDonald, 1971) generally followed theoretical expectations and also indicated (as in several other species) that the translocation males may actually compete better in mating than the normal males. With Anopheles gambiae, sterile hybrid males produced from known crosses within the complex (Davidson, 1967a, 1969a,b) were added to cages with normal males and females in varying proportions. There was a significant reduction in female fecundity, and again the sterile males appeared to have, a t least under cage conditions, superior mating ability (but see Section VIII, F). Competitive Displacement One of the suggested types of vector control is competitive displacement of a vector species with a nonvector one. This might be effected in several ways, many of them involving genetic or sterility mechanisms, and might be especially effective in Aedes mosquitoes, in which competitive displacement has evidently occurred (Gilotra et al., 1967). I n the South Pacific, A . polynesiensis is a vector of filariasis, A . albopictus evidently is not. In areas in which A. polynesiensis is abundant but A . albopictus rare or absent, a successful introduction of A . albopictus could possibly invade the A . polynesiensis niche and completely, or almost completely, replace it, with a concomitant effect upon disease transmission. Males of Aedes albopictus compete strongly with A . polynesiensis males for A. polynesiensis females (Gubler, 1970a; Ali and Roaeboom, 1971a) and readily inseminate them. When this cross-insemination occurs during the first 2 days of female life, it effectively blocks insemination of these same females with conspecific males. The opposite cross does not succeed. When A . albopictus males are added to a cage colony a t the rate of 10 A . albopictus males to 1 A . polynesiensis male, the cage colony is eliminated. Refined experiments (Gubler, 1970b) with different proportions of the two species also resulted in elimination of A. polynesiensis. The induced sterility of the A . polynesiensis females as well as the higher reproductive potential of A . albopictus (Gubler, 1970c) appear to be the principal factors involved. The two species apparently have few if any differences in oviposition behavior (Gubler, 1971). Competition experiments in a larger cage more approximating natural conditions (Roaeboom, 1971) also resulted in elimination of A . polynesiensis. Larvae of A . albopictus were superior in laboratory experiments (Lowrie, 1973a) and indeed, when all life stages of the two species were allowed to compete freely in a single cage, A . polynesiensis was drastically reduced or

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eliminated (Lowrie, 197313). It would seem that A . albopktus would be an ideal candidate for displacement of A . polynesiensis in an island situation. Possible natural displacements (Steff an, 1970; Rozeboom and Bridges, 1972) of one species by another are of interest. Competitive displacement (DeBach, 1966) using cytoplasmically incompatible strains in Culez was suggested by Laven (1967a) as a followup to control programs. Successful laboratory tests with bidirectionally incompatible strains (Curtis and Adala, 1974) indicate that this method has practical control value.

E. COMPUTER MODELS A series of computer models (Cuellar, 1968a,b, 1969a,b; Davidson and Cuellar, 1967) attempts to calculate the population dynamics of anopheline sterile male releases. Estimating the most important biodynamic factors, the number of F, (sterile hybrid) eggs which would be necessary to obtain a critical level of interference is calculated. The failure of the hybrid male experiments in the field (Davidson et al., 1970) is analyzed in terms of the feedback from the field work (Cuellar, 1970, 1973b). The reassessment of thc data is not as bleak as originally thought, and indeed eradication might have been achieved if releases had been continued for a longer period. A delightful discussion of the pros and cons of computers (Cuellar, 1973a) stresses thc necessity for feedback from the experimenter. Net reproduction rate and mean length of a generation as parameters in sterile male releases are discussed by Cuellar and Cooper (1973; see also Conway, 1970; Cuellar, 1973d; Berryman et al., 1973).

F. FIELD TRIALS After all the theoretical models have been constructed, the computers run, and the laboratory tests completed, the ultimate value of genetic control methods lies in the hard outside world, in field tests. Considering the newness of genetic control, field tests have been, on the whole, remarkably encouraging, although there have been some enlightening, instructive failures. Earlier field trials, unsuccessful, have been summarized in Rai (1969b) and in Davidson and Kitzmiller (1970). The eminently successful Okpo experiment with Culez fatigans is described in Section VIII, F, 6. A field release of Apholate-sterilized C. fatigans males in Tanzania (Bransby-Williams, 1971) was a failure. Release of large numbers of sterile males during a period in which the population was declining had

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no effect upon fertility. The author concludes that the laboratory-reared males were not competitive in the field. 1. Anopheles Hgbrid Males

Utilizing the sterility of male hybrids in certain crosses in the Anopheles gambiae complex, a field trial against A . gambiae species A was carried out by Davidson and co-workers in Pala, a small, isolated village near Bobo-Dioulasso, Upper Volta. Although laboratory experiments (Davidson, 1969a,b) had shown that sterile hybrid males were highly competitive, in the actual field trials these sterile males did not compete nearly as well for the native females as did the native males (Davidson, 1968, 1969a,b, 1970a, 1971, 1973a ; Davidson et al., 1969, 1970). Evidently the hybrid males are competitive in the small confines of a laboratory cage, but not in nature. One possible reason for this failure may lie in differences in ethological behavior, which certainly must have been enhanced by using a hybrid between two species to attempt to control a third (Davidson, 1973a). These trials are well summarized in Davidson (1970a, 1971). 2. Culex

a . Seahorse Key-Chemosterilized Males. A successful field trial was carried out with chemosterilized Culex fatigans on one of the islands off the coast of Florida (Patterson et al., 1970a,b,c; Lowe et al., 1974). The initial experiment on Seahorse Key in 1968 utilized Tepa as a sterilant; when the test was discontinued after 8 weeks, 85% of the rafts collected were sterile. The next year males sterilized with Thiotepa, were released a t a rate of from 8400 to 18,000 per day. After six generations 95% of the rafts collected were sterile. Furthermore, the total number of egg rafts laid decreased by 96%. A less successful field trial with gamma ray-sterilized C. fatiguns (Patterson and Sharma, 1972) introduced about 25% sterility into a small village. b. France-Translocation Males. Also highly successful was an experiment releasing translocation males of Culex pipiens in southern France (Laven et al., 1971a,b, 1972). After documenting a rapid increase in egg production in the summer of 1970, translocation males were released a t the beginning of August. By the end of September, the percentage of semisterile rafts had reached 95% and the total egg production was down to about 10% of normal. No further releases were carried out in 1971 but the same area was monitored. The translocations not only survived the winter but the population remained consistently low throughout the summer of 1971 (Cousserans and Guille, 1973), a result which must be due in large part to the continuing effect of the translocations.

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3. Aedes-Translocations

An important field trial was done with Aedes aegypti (Rai, 1972; Rai et al., 1973). Theoretical control of populations through genetic means, such as a translocation, depends on whether the genetic mechanism can be incorporated into, and survive in, a natural population. Using the dominant marker Silver mesonotum, heterozygous males were released into a tire dump area, with heavy A. aegypti breeding. The Silver marker survived a t least three generations. Similarly, translocation heterozygote males were released in the tire dump for 22 days. T/+ males mated with into a tire dump area, with heavy A . aegypti breeding. The Silver marker normal females, as evidenced by egg batches with less than 80% hatch, which were still being collected a month after the cessation of the initial release. This persistence argues for establishment of the translocation for at least two, and possibly three, generations.

4. Anopheles-El Salvador-Chemosterilized Males The largest-scale field trial to date has been done in El Salvador, sponsored jointly by the Insects Affecting Man Research Laboratory, USDA, Gainesville, Florida, and the Center for Disease Control, Atlanta, Georgia. This trial, against Anopheles albimanus and using ehemosterilized males, was a resounding success and definitely proved that control of albimanus by this method is feasible, at least in a limited area. A thorough ecological and biological survey of the test site, Lake Apastapeque (Breeland et al., 1974), measured larval and adult densities, principal food sources, dispersal, climatic factors, other anopheline species densities and especially the seasonal variation in densities. The anopheline density is lowest in March and April a t the end of the dry season, then rapidly builds up to a peak in September and October. Large laboratory colonies were established from A . albimanus strains native to the area, reared in mass culture, and sterilized as pupae in P,Pbis (l-aziridinyl)-N-methylphosphinothioic amide, which produced 99% male sterility (Dame et al., 1974). About 4.3 million chemosterilized males were released over a 5-month period beginning in April 1972 a t a rate that varied from about 13,000 to 40,000 per day. The larval and adult populations were immediately affected and were reduced more than 99% by September. No fertile females were found from September 1 through the middle of October (Lofgren et al., 1974). The release of sterile males reduced the population to a very low level and completely prevented the usual autumn peak of A . albimanus abundance. Valuable data on the population dynamics of the species (Weidhaas

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et al., 1974) include an estimate of the population levels before treatment, daily emergence rates, and rates of increase during the wet and dry seasons (see also Weidhaas et al., 1972a,b). Trials in India A large-scale experiment with Aedes aegypti and Culex fatigans is

6. WHO/ICMR-Field

being carried out by the World Health Organization and the Indian Council of Medical Research, New Delhi (Brooks, 1974). Comparative studies were under way with chemosterilized, radiosterilized, and cytoplasmically incompatible C . futigans, with translocation males of A. aegypti and with various other control schemes. Few data are complete a t this time. Volume 6, number 2, of the Journal of Communicable Diseases, 1974 (New Delhi) is devoted to short articles on various phases of the program. This experiment has been discontinued. See Anonymous (1975). 6. Cytoplasmic Incompatibility and Eradication

As long ago as 1964 cytoplasmic incompatibility (see Laven, 1957a,b) was suggested (World Health Organization, 1964) as a possible mechanism for the eradication of Culex pipiens. This had been questioned (Barr, 1966, 1970) because of a phenomenon which Barr called “partial compatibility,” in which some, but not all, of the females produce nonhatching egg rafts. Perhaps the dilemma involves the difference between inbred laboratory strains and heterogeneous field populations. At any rate, Laven tested his incompatibility hypothesis successfully in the laboratory, then set up an extensive field experiment in Burma, the nowfamous Okpo experiment (Laven, 1967a, 1971b). A strain of C . fatigans with Paris cytoplasm and Fresno genome was produced in large numbers. Okpo, north of Rangoon, is an “island” community, completely surrounded in the dry season by dry rice paddies. Prior tests had shown that (‘native” females were incompatible with the ‘‘imported” strain (0.14% hatchability). The local population was estimated, and sterile males were released starting in February, 1967. At first small numbers of males were released, then from March 16 until May 6, 5000 males were released per day. In spite of a 10-fold increase in the number of rafts by the end of 1 month the percentage of fertile rafts steadily declined, almost in a straight line, from the end of March until the second week in May. On May 9 and 10, 100% of the 70 rafts obtained were infertile. The monsoons started on May 11 and the experiment was discontinued. In this experiment it was clearly shown that the incompatibility principle can operate in nature, given ideal conditions. In Okpo, the local population was eradicated in about 3 months. Laven’s paper in

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Nature (1967a) gives the “bare bones” of the experiment; his 1971 paper in “Annales de Parasitologie” gives many details and interesting insights into the experiment (Laven, 1971b). A (to this reviewer) semantic objection was voiced (Laird, 1967) stating that eradication was “not proven,” but only suppression of the population to a low level. The fact remains that in the given circumstances the population was in fact eliminated, the method was proved, and vector control was achieved. Large-scale experiments in India are under way as this is written, but only fragmentary data are available a t this time (Rajagopalan et al., 1972, 1973; Krishnamurthy, 1972, 1974; Krishnamurthy and Laven, 1972). For brief accounts of experiments utilizing cytoplasmic incompatibility, see also Chan 1972, Laven 1972b,c, Thomas 1972. Laven (1975a,b) sharply disagrees with the design and execution of the Indian experiments (see also Subbarao et al., 1974).

IX. Biochemical Genetics

Many biochemical investigations have utilized mosquitoes as experimental animals. Those papers are not reviewed here. I have included only those papers that clearly deal with the genetic basis of biochemical pathways, with evolutionary patterns, or with molecular genetics.

A. PATHWAYS The first red-eyed mutant to be described in Anopheles (Laudani et al., 1969) was analyzed for the presence of ommochromes and their precursors. In the normal-eyed fly, both xanthommatin and an ommin could be identified, but in the red-eyed mutant only the xanthommatin was present. The mutant “or” (occhi rossi, Laudani et al., 1969) when purified (Laudani, 1970) showed nonhomogeneous chromatographic behavior. The or mutant also affects the pterine system (Laudani and Lecis, 1970) confirming the pleiotropic effect. In Aedes aegypti a white-eyed mutant (Bhalla, 1968b) blocks ommochrome synthesis, preventing the formation of 3-hydroxykynurenine, and the bronze mutant (Bhalla and Craig, 1967) may block one of the steps in tyrosine metabolism although artificial tanning only slightly improved hatching and embryonic development. Yurkiewicz and Bhalla (1969) suggest that alterations in fatty acid composition might be involved. Yellow larval color in Aedes aegypti has been considered t o be due to lowered amounts of uric acid in the fat body, but Inwang (1971) shows

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that yellow larvae in fact have increased amounts of uric acid, and heterozygotes are somewhat intermediate in level. Xanthine dehydrogenase levels in yellow larvae were approximately double those in wild-type larvae. It is suggested that the excess uric acid inhibits xanthine dehydrogenase activity, permitting accumulation. I n Culex pipiens three color mutants, white, red, and ruby were analyzed (Dennhofer, 1971a). An ommochrome and a sepiapterin occurred in wild type and also in red; neither was present in white. The ruby mutant showed properties similar to the brown mutant in Drosophila. B. MOLECULAR GENETICS A welcome start has been made in this important area. Mosquitoes are good material for such studies, and further work should be encouraged. Kreutzer (1970) demonstrated nucleolar organizer regions on the X chromosomes in the ovarian nurse cells of Anopheles atroparvus, further confirmed by Farci et al. (1973), who also demonstrated thymidine incorporation in the nucleolus and in certain puffed regions; I n Culez pipiens molestus and Aedes aegypti (Laurence and Simpson, 1972; Sharma, 1966) mitosis and DNA synthesis in the ovary are related t o the blood meal and in A . aegypti to an arrest in the Gz phase of the mitotic cycle. Also in A . aegypti 26 S RNA dissociates over a narrow temperature range into 18 S RNA (Shine and Dalgarno, 1972, 1973). Nine species of Nematocera (Jost and Mameli, 1972) fell into three categories with respect to DNA content: a Culex group with about 1.0 pg of DNA per haploid set, an Anopheles group with about 0.25 pg of DNA per haploid set, and a third group (Dixa) with 0.16 pg. No differences were found among sibling species of Anopheles or between reproductively isolated Culex populations. I n polytene chromosomes of A . atroparvus (Tiepolo and Laudani, 1972) DNA synthesis coincides with the beginning of the third and fourth molts; and in 48-hour fourth-instar larvae, 10% of the nuclei are in active synthesis. Curious differential DNA synthesis (Tiepolo et al., 1974) in A . atroparvus )( A . labranchk hybrids appears to be due to different rates of replication in the two species. DNA labels have been used to obtain estimates of the time required for spermatogenesis (V. P. Sharma et al., 1970) and ovarian ribosomal RNA is mostly conserved up to 48 hours but turns over between 48 and 120 hours (Frelinger and Roth, 1971; Roth et al., 1968). Quantitative changes in DNA (Blevins, 1972) were utilized to follow total DNA during embryogenesis, larval, pupal, and adult life of Aedes aegypti and, by dividing by the amount of DNA per cell, the maximal number of cells during those periods. DNA increased t o a maximum on day 5 of larval life. The

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newly emerged larvae had about lo6 cells. The same peak for DNA as well as RNA was found by Pillai and Agarwal (1969), who also followed DNA and RNA synthesis in Apholate- and Hempa-treated insects. Neither chemosterilant affected the amount of DNA or RNA. Preliminary data in Culez pipiens (French, 1970, 1971, 1973a,b,c,d) suggest alternative hereditary forms of mitochondria with important possibilities for organization and regulation. C. BIOCHEMICAL TAXONOMY Several biochemical assays may be used to separate closely related species of mosquitoes. Differences in chromatographic patterns of fluorescing compounds were used (Micks e t al., 1966a) to distinguish strains of Aedes aegypti and also of Anopheles gambiae (Davidson et al., 1967; Micks et al., 1966b, 1967; Micks, 1968). Five of these fluorescing compounds were identified as pteridines (Micks et al., 1968). Several mutants of Aedes aegypti, Aedes mascarensis, and Culex pipiens (Bhalla, 1968a) reflect qualitative and quantitative differences. I n Culez, immunodiffusion techniques (Cupp and Ibrahim, 1973; Cupp et al., 1970) may be used to separate C. pipiens, C . fatigans, and C . molestus; C . molestus has a unique precipitin band. Schumann (1973, 1974) analyzed three strains of C . pipiens: Paris, Hamburg, and Oggleshausen, which represent three different crossing types within the pipiens complex. By gel diffusion, 5 common and identical antigens can be demonstrated in males in all three crossing types; an additional common antigen is present in females. Using disc electrophoresis, 24 protein fractions common to both sexes and an additional female one could be demonstrated in all three types. When eggs of the same three strains were used the Paris and Hamburg strains contained four common antigens ; Oggleshausen, three, and certain crosses, five. Disc electrophoresis demonstrated 10 bands common to all strains, and nine additional, different bands in various crosses. Characterization of the enzymes showed that all types shared glucose-6-phosphate dehydrogenase, glutaminate dehydrogenase, lactate dehydrogenase, and catalase. Malate dehydrogenase and leucine aminopeptidase occurred as two isozymes in each raft. Alkaline phosphatase occurred only in Og eggs but these Og eggs had only two acid phosphatases while others had three. Eight esterases were demonstrated. The results indicate considerable biochemical uniformity among these three types. Details of how cytogenetics, biochemistry, and cytochemistry can be applied to the microtaxonomy of Anopheles are given in Frizzi et al. (1968).

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REFERENCES Abdel-Malek, A. A., and Ahmed, S. H. 1972a. Induced dominant lethals in the immature stages of Culex pipiens molestus Forsk. by gamma irradiation. Egypt. J . Genet. Cytol. 1, 300-302. Abdel-Malek, A. A., and Ahmed, S. H. 1972b. Biological effect of gamma-irradiation from CO* on the developmental stages of Culez pipiens mobstus Forsk. (Diptera). Acta Entomol. Bohemoslov. 69, 365-372. Abdel-Malek, A. A., and Ahmed, S. H. 1972c. Induced dominant lethals in the immature stages of Culex pipiens molestus Forsk. produced by gamma-irradiation of the parental pupae. Vestn. Cesk. Spobcnosti Zool.36, 233-236. Abdel-Malek, A. A., and Ahmed, S.-H. 1973. Effect of radiosterilization on mating frequency, mating competitiveness and sperm activity of Culex pipiens molestus. Forsk. (Diptera). Acta Entomol. Bohemoslov. 70,323-327. Abdel-Malek, A. A., Tantawy, A. O., and Wakid, A. M. 1966. Studies on the eradication of Anopheles pharoensis Theobald by the sterile-male technique using Cobalt-60. I. Biological effects of gamma radiation on the different developmental stages. J . Econ. Entomol. 59,672-678. Abdel-Malek, A. A., Tantawy, A. O., and Wakid, A. M. 1967a. Studies on the eradication of Anopheles pharoensis by the sterile-male technique using Cobalt-60. 111. Determination of the sterile dose and its biological effects on different characters related to "fitness" components. J . Econ. Entomol. 60, 20-26. Abdel-Malek, A. A., Tantawy, A. O., and Wakid, A. M. 1967b. Studies on the eradication of Anopheles pharoensis by the sterile male technique using Cobalt-60. VI. Sperm activity in males irradiated with the sterilizing dose. J . Econ. Entomol. 60, 1300-1302. Akey, D. H., and Jones, J. C. 1968. Sexual responses of adult male Aedes aegypti using the forced-copulation technique. Biol. Bull. 135, 445-453. Akiyama, J. 1973. Further isolations of translocations in Anopheles gambiae species Trop. Med. Hug. 67,440-441. A. Trans. Roy. SOC. Alexander, M. P. 1970. Use of a bench vise as a source of pressure for flattening chromosome preparations. Stain Technol. 45, 244-246. Ali, S. R., and Rozeboom, L. E. 1971a. Cross-mating between Aedes (S.) polynesiensis Marks and Aedes (S.) albopictus Skuse in IL large cage. Mosquito News 31, 80-84.

Ah, S. R., and Rozeboom, L. E. 1971b. Cross-insemination frequencies between strains of Aedes albopictus and members of the Aedes scutellaris group. J . Med. Entomol. 8, 263-265. Ali, S. R., and Rozeboom, L. E. 1972. Observations on sterilization of Anopheles ( N . ) albimanus Wiedemann by x-irradiation. Mosquito News 32, 574-579. Ali, S. R., and Rozeboom, L. E. 1973. Comparative laboratory observations on selective mating of Aedes (Stegomyia) albopictus Skuse and A . (S.)polynesknsk Marks. Mosquito News 33, 23-28. Amirkhanian, J. D. 1968. A combined HC1-acetic alcohol fixation and hydrolysis followed by cresyl violet staining for mosquito chromosome spreads. Stain Technol. 43, 167-170. Amirkhanian, J. D. 1970. Thalidomide as a mutagenic agent in the mosquito (Culex pipiens molestus) Experientia 26, 796799. Amirkhanian, J. D. 1972. Chemically induced inheritable semisterility in the mosquito (Cubx pipiens molestus). Experientia 28,7W709.

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Amirkhanian, J. D. 1973a. The salivary gland chromosomes of Anopheles stephensi mysorensis of Iran. Proc. Int. Congr. Trop. Med. Malariol, 9th, Athens 2, 186. Amirkhanian, J. D. 1973b. Culez pipiens molestus as a convenient test system for the evaluation of mutagenic agents: based on dominant lethal assays, inheritable semi-sterility and cytogenetical studies. Genetics 74,s6. Amirkhanian, J. D. 1973c. Production of semi-sterility in the mosquito by chemosterilants and chemical mutagens. Proc. Int. Congr. Trop. Med. Malariol, gth, Athens 2, 185. Anonymous. 1972. Monogamous mosquito. Nature (London) 235, 366. Anonymous. 1975. Oh, New Delhi; Oh Geneva. Nature (London) 256, 355-357. Ariaratnam, V., and Georghiou, G. P. 1971. Selection for resistance to carbamate and organophosphorous insecticides in Anopheles albimanus. Nature (London) 232, 642-644. Ariaratnam, V., and Georghiou, G. P. 1973. Carbamate-resistance in Anopheles albumanus: Cross-resistance characteristics inheritance pattern and carbaryl metabolism. Proc. Int. Congr. Trop. Med. Malariol., Oth, Athens 1, 262. Aslamkhan, M. 1970. A gynandromorph of Culex fuscocephalus Theobald from East Pakistan. Mosquito News 30,271-273. Aslamkhan, M. 1971. Karyotype of Tripteroides aranoides (Diptera, Culicidae) . Pak. J . 2001.3,237-239. Aslamkhan, M. 1973a. Sex-determination and linkage groups in Anopheles stephensi. Pak. Sci. Conf., Proc. 24, D46. Aslamkhan, M. 197313. Morphological mutant in Anopheles stephensi. Pak. Sci. Conf., Proc. 24, D45-D46. Aslamkhan, M. 1973c. Formal genetics of the malaria mosquito, Anopheles stephensi. Proc. Int. Congr. Trop. Med. Malariol., 9th, Athens 1, 246-247. Aslamkhan, M., and Aaqil, M. 1970. A preliminary report on the 7-induced translocations and semisterility in the malaria mosquito, Anopheles stephemi. Pak. J . Sn'. Res. 22, 183-190. Aslamkhan, M., and Baker, R. H. 1969a. Gynandromorphism in Culez tritaeniorhynchus. Mosquito News 29, 127-132. Aslamkhan, M., and Baker, R. H. 1969b. Karyotypes of some Anopheles, Ficalbia and Culex mosquitoes of Asia. Pak. J. 2001.1, 1-7. Aslamkhan, M., and Hyder, T. 1972. Karyotype of a Mansonia mosquito. Pak. J . Sci. Res. 24, 324-327. Aslamkhan, M., and Laven, H. 1970. Inheritance of autogeny in the Culez pipiens complex. Pak. J . 2001.2, 121-147. Aslamkhan, M., Aaqil, M., and Hafeez, M. 1972a. Genetical and morphological variations in a natural population of the malaria mosquito, Anopheles stephensi from Karachi, Pakistan WHOIVBC 72.337, W H O I M A L 72.762. Aslamkhan, M., Aaqil, M., and Hafeez, M. 1972b. Genetical and morphological variations in a natural population of the malaria mosquito, Anopheles stephensi from Karachi, Pakistan. Biologia (Lahore) 18,29-41. Asman, M. 1967. Inherited semi-sterility in Aedes aegypti. Proc. Calif. Mosquito Contr. Ass. 35, 103. Asman, M. 1974. Cytogenetic observations in Culex tarsalis: Mitosis and meiosis. J. Med. Entomol. 11,376-382. Asman, M.,and Rai, K. S. 1966. Low hatch in a new strain of Aedes aegypti. Bull. Entomol. Soc.,Amer. 12, 257. Asman, M., and Rai, K. S. 1972. Developmental effects of ionizing radiation in Aedes aegypti. J. M e d . Entomol. 9, 468-478.

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Avirachan, T. T., Seetharam, P. L., and Chowdaiah, B. N. 1969. Karyotype studies in oriental anophelines. I. Cytologia 34, 418-422. Baker, R. H. 1968. The genetics of “golden”, a new sex-linked colour mutant of the mosquito Culex tritaeniorhynchus Giles. Ann. Trop. Med. Parasitol. 62, 193-199. Baker, R. H. 1969. White eye, a female-sterile and sex-linked mutant of C u b z tritaeniorhynchus. Mosquito News 29, 571-573. Baker, R. H. 1971. Genetic control of disease carriers. Pak. Med. Forum 6, 67-71. Baker, R. H., and Aslamkhan, M. 1968. Mutants of Culex tritaeniorhynchus Giles. WHOIVBC 68.84. Baker, R. H., and Aslamkhan, M. 1969. Karyotypes of some Asian mosquitoes of the subfamily Culicinae (Diptera: Culicidae). J . Med. Entomol. 6, 44-52. Baker, R. H., and Kitzmiller, J. B. 1965a. Chromosomal asynapsis in hybrid anopheline mosquitoes. Amer. Zool. 5, 204. Baker, R. H., and Kitzmiller, J. B. 196513. The salivary chromosomes of Anopheles occidentalk. Bull. W.H.O. 32, 575-580. Baker, R. H., and Rabbani, M. G. 1970. Complete linkage in females of Culez t d a eniorhynchus mosquitoes. J . Hered. 61,59-61. Baker, R. H., and Sakai, R. K. 1972a. The genetics of Delta, a dominant sex-linked mutant of the mosquito, Culex tritaeniorhynchus. Can. J . Genet. Cytol. 14, 353-361. Baker, R. H., and Sakai, R. K. 1972b. Genetic studies on C u b z tritaeniorhynchus. Proc. Int. Congr. Entomol., 14th, Canberra Abstr., p. 122. Baker, R. H., and Sakai, R. K. 1973a. Genetic studies of two new mutants in linkage group I11 of the mosquito Culez tritaeniorhynchwr. Ann. Trop. Med. Parasitol. 67, 467-473. Baker, R. H., and Sakai, R. K. 1973b. Genetics of rose, an allele of the white locus in a mosquito. J . Hered. 64, 19-23. Baker, R. H., and Sakai, R. K. 1974. Genetic studies on Culez tritaeniorhynchus. In “The Use of Genetics in Insect Control” (R. Pal and M. Whitten, eds.), pp. 133-182. North-Holland Publ., Amsterdam. Baker, R. H., Kitzmiller, J. B., and Chowdaiah, B. N. 1965. The salivary gland chromosomes of Anopheles pseudopunctipennis pseudopunctipennis. Bull. W.H.O. 33, 837-841. Baker, R. H., Kitzmiller, J. B., and Chowdaiah, B. N. 1966. The salivary gland chromosomes of Anopheles hectoris. Chromosoma 19, 126-136. Baker, R. H., Nasir, A. S., and Aslamkhan, M. 1968. The salivary gland chromosomes of Anopheles pulcherrimus Theobald. Parassitologia 10, 167-177. Baker, R. H., Sakai, R. K., and Mian, A. 1970. Stock list of genetic strains of Culez tritaeniorhynchus. WHO/VBC 70.246. Baker, R. H., Sakai, R. K., and Mian, A. 1971a. Linkage group-chromosome correlation in Culez tritaeniorhynchw. Science 171,585-587. Baker, R. H., Sakai, R. K., and Mian, A. 1971b. Linkage group-chromosome correlation in a mosquito. Inversions in Culez tritaenwrhyzchus. J . Hered. 62,3136. Barr, A. R. 1966. Cytoplasmic incompatibility as a means of eradication of Culez pipiens L. Proc. Calif. Mosquito Contr. Ass. 33, 32-35. Barr, A. R. 1967. Occurrence and distribution of the Culez pipiens complex. Bull. W.H.O. 37, 293-296. Barr, A. R. 1969. Divided eye, a sex-linked mutation in Culez pipiens L. J . Med. Entomol. 6, 393-397.

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