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Diagnostic Characterization of Anopheles Freeborni and An. Hermsi by Hybrid Crosses, Frequencies of Polytene X Chromosomes and T

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Diagnostic Characterization of Anopheles Freeborni and An. Hermsi by Hybrid Crosses, Frequencies of Polytene X Chromosomes and T 198 JouRrveloF THE At'lnRrceN Moseurro Cotrnol Assocrerror'l Vol. 7, DIAGNOSTIC CHARACTERIZATION OF ANOPHELES FREEBORNI AND AN. HERMSI BY HYBRID CROSSES,FREQUENCIES OF POLYTENE X CHROMOSOMESAND TDNA RESTRICTION ENZYME FRAGMENTS G. N. FRITZ,I S. K. NARANG,' D. L. KLINE,' J. A. SEAWRIGHT,' R. K. WASHINO.I C. H. PORTER' lr.ro F. H. COLLINS3 U.S. Department of Agriculture, Agricultural ResearchSeruice, Medicat and Veterinary EntomologyResearch Laboratory, P. O. Box 14565,Gainesuille, FL 32604 ABSTRACT. A qolytene chromosomeanalysis was prepared ftom Anophelesfreeborni collectedfrom 25 Iocations in north and central California, and parts of Washington and Oregon.The X chromosome banding pattern, thought previously to be specifii to An. hcrmsi, *as co--otr-in mosquitoescollected from_foothill regions in California, and in all samplesfrom Washington and Oregon. Ai some of these lo-cation-s,many mosquitoeshad heterokaryotypesfbr the inversion thlt distinguishls the X chromosome of An. freeborni from that of An. hermsi. Use of rDNA restriction site analysis, and the results from crossingof different strains bearing either type of X chromosome,showed thai An. hermsi doesnot have a unique.or diagnosticX chromosome. Anopheleshermsi wascollected in San Mateo County, CA, which is now the northernmostknown limit of this species.Crossing studies, or the examinationof 1DNA restriction enzymeprofiles, are presently the only meansof identifying An. herrnsi. INTRODUCTION cording to the description of An. hermsi (Barr and Guptavanij 1988),the only characterthat A new speciesof anopheline mosquito,Anoph- distinguishesboth speciesis a simple inversion eleshermsi, was describedrecently by Barr and on the polytene X chromosome.The banding Guptavanij (1988). At present, An. hermsi is patterns on the autosomesof both speciesare thought to be limited to areasof southern Cali- either identical (Baker 1965,6Fritz 19897)or too fornia south of the Tehachapi Mountains (Cope similar (Menchaca19865) to be useful as diag- et al. 1988).This species has beenimplicated in nostic characters.Baker and Kitzmiller (1963) recentautochthonous cases of vivax malaria in first describedthe X chromosomeof An. hermsi southern California (Maldonado "southern et al. 1990). (then referredto as occidentalis")and Prior to 1988, mosquitoes now known as An. noted that it had unmistaken homology with hermsi were believed to be a coastal strain of that of An. Both types of X chromo- Anopheles Aitken (Aitken freeborni. freeborni 1939,1945), some appearedto differ only by a single inver- a geographic variant of Anopheles occidentalis sion (In(X)A) at subzones2B-5C (Baker 19656). (Lewallen Dyar and Knab 1957)or a new species As a part of a study on the population genetics (Baker and Kitzmiller 1963, Fujioka 1986,a of An. freeborni,we examinedthe polytenechro- Menchaca19865). mosomes of specimens collected throughout present, At there are no reliable anatomical north and central California, and parts of Ore- charactersthat distinguish eggs,Iarvae or adults gon, Washington and Utah. The purpose of this of.An. hermsi from those of An. Ac- freeborni. cytogenetic survey was to distinguish An. free- borni from An. hennsi, record chromosomalpol- ymorphisms and use these polymorphisms as 1 Department of Entomology, University of Califor- genetic markers to determine population struc- nia. Davis. CA 95616. ture and speciesdistributions. ' USDA, ARS, Medical and Veterinary Entomology Our initial survey showed that the X chro- Research Laboratory, P. O. Box 14565, Gainesville, mosome banding pattern described as specific FL 32604. present 3 to An. hermsi was in some samples Malaria Branch Division of Parasitic Diseases, collectedin northern and central California. and Centers for Disease Control. Atlanta. GA 30333. nFujioka, parts of Oregon and Washington. Furthermore, K. K. 1986. Hybridization and electro- phoretic studies of 3 members of the North American Anopheles maculipennis complex (Diptera: Culicidae). Ph.D. dissertation, University of California, Los An- o Baker, R. H. 1965. Cytogenetic evidence for the geles. evolutionary relationships among the Nearctic rnocu- u Menchaca, D. M. 1986. The cytogenetic study of lipennis speciesof anopheline mosquitoes.Ph.D. dis- an undescribed member of the North American sertation, University of lllinois, Urbana. A nophe Ies maculipennis (Diptera: Culicidae) complex. 7Fritz, G. N. 1989. Mass rearing and population Ph.D. dissertation, University of California, Los An- genetics of Anopheles freeborni. Ph.D. dissertation, geles. University of Florida, Gainesville. PoLYTENEX Csnonaosoun Table 1 Collection site number and location for samples of Anoph.elescollected in California, Oregon and Washington. Site no. Location California 1 NevadaCo., Wolf Creek& Highway 49 2 El DoradoCo.. Camino. Carson Rd. 3 El DoradoCo., Pleasant Valley Rd. 4 SacramentoCo., Sloughouse SacramentoCo., Folsom t) Sutter Co.,Highway 99 & HowsleyRd. 7 Yolo Co.,Capay Valley, Guinda 8 Yolo Co.,Knights Landing q ColusaCo., Millers Landing 10 Colusa Co., Highway 20 near Williams 11 Sutter Co.,west of Yuba City on Butte HouseRd. 12 Butte Co.,Chico IJ TehamaCo., Tehama, Gyle Rd. t4 Glenn Co.,east of Willows on Highway 162 15 Lake Co.,Clear Lake 16 SonomaCo.. Sonoma. Huichica Cr. 1a SacramentoCo., Highway 99 & Twin Cities Rd. 18 San Mateo Co.,Jasper Ridge Preserve on Sand Hill Rd. 19 Kern Co.,Onyx, CanebrakeCreek 20 Inyo Co.,Big Pine 2l Inyo Co.,Bishop Oregon 22 JeffersonCo., Madras oa Umatilla Co.,Hermiston Washington 24 Benton Co.,Richland 25 Yakima Co.,Yakima many mosquitoesat someof theselocations were MATERIALS AND METIIODS heterokaryotypes (one copy of the standard X Insect sarnplesand chrornosonxepreparation: chromosome specific to An. freeborni and one copy of In(X)A). These unexpected results Larvae and adults were collected during July- (Table raised some important questions: October 1988 from 25 different sites 1. 1) Was An. herrnsi more widely distributed Fig. 1). In California, many areas throughout than thought previously? the SacramentoValley were sampled as well as locationsin the foothills of the Sierra Nevada, 2) Were An. freeborni and An. hermsi really distinct species? the coastalrange,2locations in the OwensVal- ley and one location the coast at Palo Alto. 3) Were individuals of An. freeborni and An. near h,ermsi mating with one another in sympatric The techniquesdescribed by French et al. (1962) wereused to preparethe polytenechromosomes. populations? "C 4) Was one species,both, or perhaps an uni- Bloodfedfemales were kept at 28 for approx- dentified third speciespolymorphic for In(X)A? imately 28-29 h. Ovarieswere dissectedin dilute To answer these questions, we have crossed Carnoy's and then transferred to 75Vo acetic various geographic strains having one or the acid. Chromosomes,banding patterns and in- other type of X chromosome.According to Fu- versions were identified using a photographic jioka (19864),crossing An. freeborni to An. map made of the best polytene chromosome hermsi will produce sterile hybrid male progeny. preparations.This map was used in conjunction In addition. we examined rDNA restriction en- with the description and standard map of ovar- zyme pattern variations, which were shown re- ian nurse-cell polytenes (Faran 19818).The cently to distinguishboth species(Collins et al. 1990).We report here the frequencyand distri- bution of both types of X chromosomesfound 8Faran, T. C. 1981.The adult ovarian nurse cell in our cybogeneticstudy, the results obtained chromosomesof Anopheles(Arcpheles) freeborni Ait- from crossing different strains and the results ken 1939(Diptera: Culicidae). M.Sc. thesis, University of the rDNA analyses. of Maryland, CollegePark. 200 JounNlr,oF THE Ar',tenrclN Moseurro CoN.rRor,Assocrlrrox VoL.7,No.2 WASHINGTON OYAKIMA CALIFORNIA O MADRAS .,9TEHAMA @(t 50MILES OREGON O @(l A " 'i: ^"O' 'u1; (9. , ?"ril' LOSANGELES Fig. 1. Frequency of the standard (white) and inversion (black) karyotype for the X chromosome at collection sites in California, Oregon and Washington. Marysville strain (which Faran usedto prepare ner describedby Collins et al. (1990). These her map) was obtained from the Walter Reed sites included those in which mosquitoes were Army Institute of Researchand usedas a stand- polymorphic for the inversion on the X chro- ard for comparing banding patterns. mosome,as well as those fixed for either homo- rDNA probe: The rDNA of An. freeborni and karyotype.Prior to the rDNA analysis,the po- An. hermsi have a restriction enzymesite differ- lytene X chromosomekaryotype of each female ence in the external and internal spacerregions mosquito was determined. Samples of An. (Collins et al. 1990). These diagnostic differ- h.ermsifrom 3 sites in southern California were enceswere usedto identify speciesin this study. alsoprobed. In a blind test, the rDNA of at least 3 individuals Crosses.'Crosses were made among several from each of 12 collection sites in California, geographicstrains presumedto be An. freeborni. Washington and Oregonwas probed in the man- To examine widely separatedgeographic strains JUNE 1991 AN. HEBMSTPoLYTENE X CgRouosolrp that also showedsome habitat and chromosomal RESULTS differences,a strain from Richland, WA (WASH strain, site 24,Table 1), was crossedto a strain X chromosome:In California, all mosquitoes from Davis (DAVIS strain) in the Sacramento from the SacramentoValley (sites4-14 in Table Valley, CA. Mosquitoes in the former site were 2, Fig. 1) and the Owens Valley (sites 20, 2L) polymorphic
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