630 US ISSN 0271-9916 August 1987 RESEARCH EXTENSION SERIES 078

HOST LIST OF PLANTS SUSCEPTIBLE TOTOMATO SPOTTED WILT VIRUS (TSWV)

J. J. Cha, R. F. L. Mau, w. C. Mitchell, D. Gonsalves, and L. S. Yudin

HITAHR COLLEGE OF TROPICAL AGRICULTURE AND HUMAN RESOURCES UNIVERSITY OF HAWAII

The Library of Congress has catalogued this serial publicationas follows:

Research extension series / Hawaii Institute of Tropical Agri­culture and Human Resources.-[Honolulu, Hawaii]:

The Institute, [1980-v. : ill. ; 22 cm.

Irregular.Title from cover.Separately catalogued and classified in LC before and

including no. 044.ISSN 0271-9916 = Research extension series - Hawaii

Institute of Tropical Agriculture and Human Resources.1. Agriculture-Hawaii-Collected works. 2. Agricul­

ture-Research-Hawaii-Collected works. I. HawaiiInstitute of Tropical Agriculture and Human Resources.II. Title: Research extension series - Hawaii Institute ofTropical Agriculture and Human ResourcesS52.5.R47 630'.5-dc19 85-645281

AACR 2 MARC-SLibrary of Congress [8506]

ACKNOWLEDGMENTSWe wish to thank M. Barut. H. Bridgman. J. Lee. S. Matsubara. and J. Palos for their able

assistance. Investigations were supported in part by the USDA/CSRS Special Grants Program inTropical and Subtropical Agriculture No. 58-9AHZ-O-546 and Grant 84-1 from the State ofHawaii Governor's Agricultural Coordinating Committee.

THE AUTHORSJ. J. Cho is an associate plant pathologist. University of Hawaii. Maui County Research.

R. F. L. Mau is an associate specialist in entomology. University of Hawaii at Manoa.

w. C. Mitchell is a retired entomologist, University of Hawaii at Manoa.

D. Gonsalves is a plant pathologist. New York Agricultural Experiment Station. CornellUniversity.

L. S. Yudin is a graduate research assistant in entomology. University of Hawaii at Manoa.

CONTENTSPage

Introduction..................................................................................................................... 3Table 1. Hosts of Tomato Spotted Wilt Virus..................................................................................... 4Literature Cited......................................................................................................................... 9

HOST LIST OF PLANTS SUSCEPTmLE TO TOMATO SPOTTED WILT VIRUS (TSWV)

J. J. Cha, R. F. L. Mau, W. C. Mitchell, D. Gonsalves, and L. S. Yudin

INTRODUCTIONSpotted wilt of tomato was first described by

Brittlebank (1919) in Australia. Samuel et aI.(1930) characterized the pathogenic agent as avirus and named it tomato spotted wilt virus(TSWV). Since then the disease has been reportedin several tropical and temperate regions and isconsidered worldwide in distribution.

TSWV is unique because it is one of only twoplant viruses known to be transmitted by thrips.Pittman (1927) was the first to establish thatTSWV could be transmitted by the onion thrips.Six thrips species are known TSWV vectors:

1) the western flower thrips. Frankliniellaoccidentalis (Pergande) (Gardner et aI., 1935);

2) the common blossom thrips.Frankliniella schultzei (Trybom) (Samuel et aI.,1930);

3) the onion thrips, Thrips tabaci Lindeman;4) the tobacco thrips, Frankliniella Jusca

(Hinds) (Sakimura. 1963);5) Scirtothrips dorsalis Hood (Amin et aI..

1981); and6) Thrips setosus Moulton (Kobatake. 1984).Only the first three species are known to

occur in Hawaii. Of these, the western flowerthrips is the most common vector (Yudin et aI..1986. 1987). TSWV can be acquired fromdiseased plants only by larval thrips; after alatent period of three to 12 days it can betransmitted by both larval and adult stages.

TSWV produces a variety of diseasesymptoms including a range of chlorotic.necrotic. stunting. and enation symptoms in allparts of the plant (Ie, 1970). TSWV symptoms onlettuce generally start as necrotic brown leafspots on one side of the plant. becomingsystemic and extending to the heart leaves. Thisresults in a cessation of development, causingthe plant to twist to one side. The outer leaveswilt and eventually the whole head collapses. Ontomatoes. symptoms are characterized by initialchlorosis of leaves and terminal shoots.developing into bronZing and necrosis.Symptoms on mature fruits are blotchiness ofcolor and rugosity of the fruit surface.

In Hawaii. TSWV has seriously affectedproduction of lettuce, tomato, and bell pepper inthe major vegetable-production areas of thestate. These areas include the Lalamilo and

Pukapu areas on the island of Hawaii, the Kulaarea on Maui. and the Waianae area on Oahu.This disease has been quite devastating in recentyears. when losses of 50 to 90 percent have beenobserved in lettuce and tomato production,particularly during the warm, dry summermonths.

Development of feasible control procedureshas been difficult because TSWV has anextensive plant host range. Best (1968) lists 157dicotyledonous plant species in 29 families, andsix monocotyledonous species in five families.as hosts. Francki and Hatta (1981) added 11additional host species. Several of thesesusceptible hosts are commonly found growingalong field borders and among crops of Hawaii'smajor vegetable-growing regions.

TSWV-infected plant hosts invegetable-growing regions play an importantrole in the epidemiology of the disease.Forty-four plant species representing 16 plantfamilies have been found to be naturallyinfected with TSWV. Twenty-six plant speciesare conSidered important reservoirs of TSWV(see Table 1). These plant sources perpetuateTSWV. are important habitats for thrips vectors.and serve as foci for subsequent movement andinfection of susceptible crop plants. Ipomoeacongesta (blue morning glory) and Tropaeolummajus (nasturtium) are present throughout theyear on the islands of Hawaii and Maui. OnOahu, Amaranthus spinosus (spiny amaranth)is commonly found. A high percentage of thoseplants found on farmland harbor the virus andwere associated with abundant vector thrips.Elimination of these and other important TSWVreservoirs should be helpful in reducing diseaselosses.

Several additional new plant hosts havebeen identified since 1981. and this paperpresents an up-to-date listing ofTSWV-susceptible plant species. Severalreferences listed in the table cite originalarticles demonstrating TSWV susceptibility ofplant species or host lists of TSWV-susceptibleplants.

3

TABLE 1. HOSTS OF TOMATO SPOTTED WILT vm.US

Scientific nameAcanthospermum hisipidium DC.Ageratum houstonianum Mill.Amaranthus caudatus L.Amaranthus graecizans L.Amaranthus hybridus L..Amaranthus retroflexus L.Amaranthus spinosus L..Amaranthus viridis L..Amaryllis sp.Ananas comosus (L.) Merr.Anemone sp.

. Antirrhinum sp.Apium graveolens L.·Aquilegia vulgaris L.Arachis hypogaea L.Arctium lappa L. •Arum palaestinum BoissAster sp.Atropa belladonna L.Begonia x hiemalis FotschBegonia semperflorens Link & OttoBegonia sp.Belamcanda chinensis (L.) DC.Bidens pilosa L.·Bidens pilosa

var. minor (B1.) Sherf·Brassica campestris ssp. chinensis L.Brassica oleracea var. botrytis L.Browallia americana L.Browallia speciosa Hook.Calceolaria sp.Calendula officinalis L.Callistephus chinensis (L.) NeesCampanula americana L.Campanula pyramidalis L.Capsella bursa-pastoris (L.) Medic.·Capsicum annuum L.Capsicum frutescens L.Capsicum sp.Carica papaya L.Cheiranthus sp.Chenopodium album L.Chenopodium ambrosioides L.Chenopodium murale L.Chenopodium quinoa Willd.Chondrilla sp.Chrysanthemum morifolium

(Ram.) Hems!.Chrysanthemum sp.Cichorium endivia L.

4

Common name

ageratum

green amaranthredroot pigweedspiny amaranthslender amaranthBarbados lilypineappleanemonesnapdragoncelery. celeriaccommon columbinepeanutburdockcallaasterdeadly nightshadebegoniabegoniabegOniablackberry lilySpanish needle

Spanish needlewhite stem cabbagecauliflowerbrowallia

slipperwortcalendulaChina asterbellflowerchimney bellflowershepherd's pursebell pepperred pepperpepperpapayawall flowerlambs quartersMexican teanettleleaf goosefoot

chrysanthemumchrysanthemumendive

ReferenceGreber and McCarthy, 1977Gumpf and Weathers, 1972Best. 1968Best, 1968Cho et al.. 1986Milbrath, 1939Cho et al.. 1986Cho et al., 1984. 1986Gardner et a1., 1935Linford, 1932Smith, 1957Smith, 1957Best. 1968Smith, 1957Costa. 1941Cho et al., 1986Tompkins and Severin, 1950Smith, 1957Smith. 1932Barnes and Halliwell, 1985Gardner et a1., 1935Best. 1968Yamamoto and Ohata. 1977Greber and McCarthy, 1977

Cho et al., 1984. 1986Cho et al., 1986Gardner et a1., 1935Gardner et a1., 1935Smith, 1957Smith, 1957Best. 1968Best, 1968Gardner et a1., 1935Smith, 1957Best. 1968Sakimura, 1940Smith, 1932Best. 1968Cook, 1972Smith, 1957Paliwal, 1974Cho et al.. 1986Cho et al., 1986Paliwal, 1974Best. 1968

Gardner and Whipple, 1934Smith. 1957Sakimura, 1940

Cichorium intybus L.Cineraria cruenta Mass.Cirsium lanceolatum HillConvolvulus arvensis L.Convolvulus sp.Cordyline terrninalis (L.) Kunth.Coreopsis drummondii Torr. & GrayCoriandrum sativum L.Coronopus didymus (L.) SmithCosmos bipinnatus Cay.Crepis capillaris Wallr. Beitr.Crepis divaricata F. SchultzCrepis pulchra L.Crepis pumilaCrepis rhoeadifolia Bieb.Crotalaria incana L.Crotalaria mucronata Desv.Cucumis sativus L.Cyanra scolymus L.Dahlia pinnata Cay.Dahlia variabilis (Willd.) Desf.Datura ferox L.Datura stramonium L. *Datura wrightii Hort. ex RegelDelphinium cultorum VossDelphinium sp.Desmodium uncinatum (Jacq.) DC.Dichondra carolinensis Michx.Duboisia leichhardtiiEmilia sagittata DC.Emilia sonchifolia (L.) DC.Emilia sp.Erigeron bonariensts L.Erigeron canadensis L.Gaillardia aristata Pursh.Gaillardia sp.Galinsoga parviflora Cay.*Galinsoga quadriradiata (Raf.) BlakeGerbera sp.Gladiolus sp.Gloxinia sp.Godetia grandiflora Lindl.Gomphrena globosa L.Gypsophila paniculata L.Helipterum manglesii (Lindl.)

F. J. Muell. ex Benth.Helipterum roseum Benth.Hippeastrum sp.Hydrangea macrophylla

'Imaculata' Ser.Hydrocotyle asiatica Bert. ex UrbanHyosoyamus niger L.Ipomoea congesta R Br.*

chicory

bullthistlebindweedmorning gloryti, kitick seedcorianderswinecresscosmoshawksbeard

fuzzy rattlepodsmooth rattlepodcucumberartichokedahliadahlia

jimson weed

larkspurSpanish clover

Flora's paintbrushred pualele

hairy horseweedsmall-leaf horseweed

fUji grassPeruvian daisyAfrican daisygladiolagloxinia

globe amaranthbaby's breath

Swan River everlasting

Barbados lily

hydrangeawater pennyworthenbaneblue morning-glory

Sakimura. 1940Jones. 1944Best, 1968Sherf. 1948Best, 1968Cho et al.. 1986Smith. 1957Best, 1968Cho et al.. 1986Smith, 1957Best. 1968Best, 1968Best, 1968Best, 1968Best, 1968Cho et al.• 1984. 1986Cho et al.• 1984, 1986Ie. 1970Garcia and Feldman. 1978Holmes. 1948Gardner and Whipple, 1934Greber and McCarthy, 1977Smith. 1931Smith, 1957Gardner et al.. 1935Best, 1968Cho et al.. 1986Sakimura, 1961McCarthy and Greher, 1978Linford, 1932Sakimura. 1940Best, 1968Helms et al.• 1961Cho et al.. 1986Gardner et al.. 1935Best, 1968Cho et al.. 1986Cho et al., 1986Best. 1968Smith. 1957Best, 1968Gardner et al., 1935Paliwal, 1974Best. 1968

Best, 1968Best, 1968Best. 1968

Allen et al.. 1983Paliwal. 1974Best. 1968Cho et al.. 1986

5

Ipomoea hederacea'Superba' (L.) Jacq.

Lactuca sativa L.*Lactuca sativa var. capitata L.*Lactuca sativa var. longifolia Lam.*Lactuca scariola L.Lathyrus odoratus L.Layia elegans Torr. & GrayLeonotis nepetaefolia R. Br. *Lilium tigrinum Ker-Gawl.Limonium latifolium (Sm.) Ktze.Lobelia sp.Lupinus aIbus L.Lupinus angustifolius L.Lupinus leucophyllus

Dougl. ex Lindl.Lychnis coronaria (L.) Desr.Lycium ferocissimum MiersLycopersicon esculentum Mill.*Lycopersicon hirsutum HBK.Lycopersicon peruvianum (L.) Mill.Lycopersicon pimpinellifolium

(Jusl.) Mill.Malcomia maritima R. Br.Malva nicaeensis All.Malva parviflora L.*Malva rotundifolia L.Marrubium vulgare L.Martynia annua L.Medicago polymorpha L.Melilotus officinalis (L.) Lam.*Mesembryanthemum sp.Montia sp.Myosotis alpestris SchmidtNepeta cataria L.Nerium oleander L.Nicandra physaIodes (L.) Gaertn.*Nicotiana acuminata

(R C. Grah.) Hook.Nicotiana alata Link & OttoNicotiana angustifolia Mill.Nicotiana atropurpureum Hort.Nicotiana benthamiana DominNicotiana bigelovii (Torr.) S. Wats.Nicotiana bonariensis Lehm.Nicotiana calycifloraNicotiana caudigera Phil.Nicotiana chinensis Fisch.Nicotiana debneyi DominNicotiana etiguaNicotiana glauca GrahamNicotiana glutinosa L.Nicotiana goodspeedii WheelerNicotiana langsdorfIi Weinm.

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lettuce

romaine

sweet pea

lion's eartiger lilystatice

lupine

dusty millermatrimony-vinetomato

Virginia stockmallowcheeseweedcommon mallowhoarhound

bur cloveryellow sweet clovertricolor ice plant

forget-me-notcatnipoleanderapple of Peru

jasmine tobacco

tree tobacco

Sakimura. 1961Gardner and Whipple. 1934Snyder and Thomas. 1936Tompkins and Gardner. 1934Milbrath. 1939Snyder and Thomas. 1936Gardner et al., 1935Cho et aI.. 1986Best. 1968Cho et aI., 1986Smith. 1957Gardner and Whipple. 1934Smith,1932

Smith. 1957Best. 1968Smith,1957Brittlebank, 1919Smith, 1944Hutton and Peak. 1949

Sakimura, 1940Best, 1968Sakimura. 1961Best. 1968Milbrath. 1939Best. 1968Gardner and Whipple. 1934Cho et aI., 1986Cho et aI.. 1986Best, 1968Best. 1968Best. 1968Milbrath, 1939Francki and Grivell, 1970Gardner and Whipple. 1934

Gardner and Whipple. 1934Gardner and Whipple, 1934Best, 1968Smith. 1957GonsaIves and Trujillo. 1986Smith,1957Best. 1968Smith,1957Best,1968Smith. 1957Best. 1968Best. 1968Smith, 1932Smith,1932Best. 1968Gardner and Whipple. 1934

Nicotiana longiflora Cav.Nicotiana macrophylla Spreng.Nicotiana maritima WheelerNicotiana nudicaulis S. Wats.Nicotiana paniculata L.Nicotiana plumbaginifolia VivoNicotiana rustica L.Nicotiana sanderae W. Wats.Nicotiana solanifolia Walp.Nicotiana suaveolens Lehm.Nicotiana sylvestris Spegaz.

& ComesNicotiana tabacum L.Nicotiana tomentosiformis

GoodspeedNicotiana undulata Ruiz & Pav.Nicotiana wigandioides Koch & FintOenothera sp.Papaver nudiaule L.Papaver orientale L.Papaver sp.Penstemon hirsutus (L.) Willd.Peonia sp.Petunia hybrida Vilm.Petunia sp.Phaseolus vulgaris L.Phlox drummondii Hook.Physalis peruviana L.Physalis pubescens L.Pisum sativum L.Plantago major L.Polygonum convolvulus L.Polygonum sp.Portulaca oleracea L. •Primula malacoides Franch.Primula obconica HancePrimula sinensis Lindl.Primula sp.Quamoclit coccinea 'Hederifolia' L.Ranunculus sp.Salpiglossis sinuata Ruiz & Pav.Salvia splendens Ker-Gawl.Saponaria officinalis L.Saxifraga sp.Scabiosa sp.Schizanthus pinnatus Ruiz & Pav.Senecio cruentus (Mass.) DC.Senecio jacobea L.Sinningia speciosa (Lodd.) Hiern.Solanum aculeatissimum Jacq.Solanum capsicastrum LinkSolanum dulcamara L.Solanum laciniatum Ait.Solanum marginatum L.

long-flowered tobacco

wild tobacco

Turkish tobacco

evening primroseIceland poppyoriental poppypoppybread-tonguepeonypetuniapetuniasnap beanphloxcape gooseberryhusk tomatogarden peabroad-leaved plantain

jointweedpurslanefairy primrose

Chinese primroseprimrosestar ipomoeabuttercuppainted tonguescarlet sagebouncing betrockfollmourning bridepincushion flower

tansy ragwortgloxiniacockroach berry

bittersweet

Smith. 1957Best. 1968Best. 1968Costa and Forster, 1942Smith. 1957Best, 1968Smith,1932Gardner and Whipple. 1934Best, 1968Smith. 1957

Smith,1957Smith, 1931

Best, 1968Best. 1968Best, 1968Best. 1968Best. 1968Gardner et al.. 1935Smith. 1957Gardner et al.. 1935Smith. 1957Gardner and Whipple, 1934Smith. 1932Smith. 1957Best. 1968Best. 1968Gardner and Whipple. 1934Snyder and Thomas, 1936Smith, 1932Smith. 1957Smith. 1957Allen et al.. 1983Smith. 1957Gardner et al.. 1935Smith. 1957Best. 1968Sakimura. 1961Smith. 1957Gardner and Whipple. 1934Gardner et al.. 1935Best, 1968Best, 1968Smith. 1957Gardner and Whipple, 1934Gardner and Whipple. 1934Allen et al.. 1983Gardner et al., 1935Smith. 1932Smith. 1931Smith, 1932Smith. 1931Smith. 1932

7

Solanum melongena L.Solanum miniatum Bernh. ex Willd.Solanum nigrum L.Solanum nodiflorum Jacq.Solanum sanitwongsei Craib.Solanum seaforthianum Andr.Solanum sodomeum Hort. ex Dun.Solanum triflorum Nutt.Solanum tuberosum L.Sonchus oleraceus L.*Spinacia oleracea L.Stachys arvensis L.Stellaria media (L.) Cyril!. *Streptosolen j amesonii MiersTagetes minuta L.Trachelium caerulea L.Trachelium sp.Trachymene caerulea R. C. Grah.Tribulus terrestris L.Trifolium repens L.Trifolium subterraneum L.Tropaeolum majus L.*Tropaeolum sp.Troximon spUrtica dioica L.Verbena hybrida VossVerbena litoralis HBK.Verbena sp.Verbesina encelioides (Cav.)

Benth. & Hook.* .Vicia faba L.Vigna sinensis End!.Vinca rosea L.Xanthium saccharatum WalIr.*Zantedeschia aethiopica Spreng.Zinnia elegans Jacq.

eggplant

black nightshade

apple of Sodom

potatosowthistlespinachstaggerweedchickweed

marigold

throatwort

white clover

nasturtiumnasturtium

nettleverbenaoi

golden crown-beardbroad beancowpeaMadagascar periwinklecockleburcommon calla lilyzinnia

Smith, 1932Smith. 1957Smith. 1931Smith, 1931Smith,1957Smith, 1957Smith. 1957Allen et al., 1983Smith, 1931Best, 1968Sakimura. 1940Greber and McCarthy. 1977Holmes, 1948Smith. 1957Best, 1968Best, 1968Best, 1968Best, 1968Allen et al.. 1983Paliwal. 1974Helms et aI.• 1961Best, 1968Best, 1968Best, 1968Gardner and Whipple, 1934Gardner et al.. 1935Cho et al., 1984. 1986Best, 1968

Cho et aI.• 1984, 1986Best. 1968Paliwal, 1974Paliwal, 1974Cho et aI.. 1986Best, 1968Best, 1968

8

LITERATURE CITEDAllen. T. C.. J. P. McMorran. and E. A. Locatelli

1983. Isolation of tomato spotted wilt virusfrom hydrangea and four weed species. PlantDis. 67:429-431.

Amin. P. W.. D. V. R Reddy. and A. M. Ghanekar.1981. Transmission of tomato spotted wiltvirus, the causal agent of budnecrosis of peanut, by Scirtothrips dorsalisand Frankliniella schultzei. Plant Dis.65:663-665.

Barnes, L. W.. and R. S. Halliwell. 1985. Tomatospotted wilt virus infecting begonia hybridsin Texas. Plant Dis. 69:613.

Best, R. J. 1968. Tomato spotted wilt virus.In K. M. Smith and M. A. Lauffer, eds.Advances in virus research. Vol. 13.Academic Press. New York. pp. 65-145.

Brittlebank. C. C. 1919. Tomato diseases. J. Agr.Victoria 17:213-235.

Cho, J. J .. W. C. Mitchell, L. Yudin, and L.Takayam? 1984. Ecology and epidemiologyof tomato spotted wilt virus (TSWV) and itsvector, Frankliniella occidentalis.Phytopathology 74:866. (Abstr.)

Cho, J. J .. R. F. L. Mau, D. Gonsalves, and W. C.Mitchell. 1986. Reservoir weed hosts oftomato spotted wilt virus. Plant Dis.70:1014-1017.

Cook, A. A. 1972. Virus diseases of papaya.Vniv. Florida Agr. Exp. Sta. Bull. 750. 19 pp.

Costa. A. S. 1941. Vma molestia de virus doamendoim (Arachis hypogaea L.). Biologico7:249-251.

Costa, A. S .• and R. Forster. 1942. Lista dehospedeiras do virus de vira-cabeca.Bragantia 2:83-91.

Francki. R. I. B.. and C. J. Grivell. 1970. Anelectron microscope study of thedistribution of tomato spotted wilt virus insystematically infected Datura stramoniumleaves. Virology 42:969-978.

Francki, R. I. B., and T. Hatta. 1981. Tomatospotted wilt virus. In E. Kurstak, ed.Handbook of plant virus infections andcomparative diagnosis. American ElsevierPubI. Co.. Inc.. New York. Pp. 491-512.

Garcia. 0 .. and J. M. Feldman. 1978. Naturalinfection of artichoke by tomato spottedwilt virus. Plant Dis. Rep. 62: 1076-1077.

Gardner. M. W., and O. C. Whipple. 1934. Spottedwilt of tomatoes and its transmission bythrips. Phytopathology 24: 1136.

Gardner. M. W.. C. M. Tompkins. and O. C.Whipple. 1935. Spotted wilt of truck cropsand ornamental plants. Phytopathology25:17.

Gonsalves. D.. and E. TrujillO. 1986. Tomatospotted wilt virus in papaya and detection ofthe virus by ELISA. Plant Dis. 70: 501-506.

Greber. R. S .. and G. J. P. McCarthy. 1977. Adisease of Duboisia caused by tomato spottedwilt virus (TSWV). Australian Plant Path.Soc. NewsI. 6:54-55.

Gumpf, D. J .. and L. G. Weathers. 1972.Identification and purification of tomatospotted wilt virus isolated from Ageratum.Plant Dis. Rep. 56:859-872.

Helms. K.. N. E. Grylls, and G. S. Purss. 1961.Peanut plants in Queensland infected withtomato spotted wilt virus. Australian J. Agr.Res. 12:239-246.

Holmes, F. O. 1948. Elimination of spotted wiltfrom a stock of dahlia. Phytopathology38:314.

Hutton. E. M.. and A. R Peak. 1949. Spotted wiltresistance in tomato. Australian Inst. Agr.Sci. J. 15:32-36.

Ie. T. S. 1970. Tomato spotted wilt virus.Commonwealth Mycol. Inst,fAssoc. AppI.BioI. Description of Plant Virus No. 39.

Jones. L. K. 1944. Streak and mosaic ofCineraria. Phytopathology 34:941-953.

Kobatake, H.. T. Osaki, and T. Inouye. 1984. Thevector and reservoirs of tomato spotted wiltvirus in Nara Prefecture. Ann. Phytopath.Soc. Japan 50:541-544.

Linford. M. B. 1932. Transmission of thepineapple yellow-spot virus by ThripstabacL Phytopathology 22:301-324.

McCarthy. G. J. P.. and R. S. Greber. 1978.Tomato spotted wilt virus in Duboisia.Queensland Agr. J. 104:336-338.

Milbrath, J. A. 1939. Tomato tip-blight virus.Phytopathology 29:156-168.

Paliwal. Y. C. 1974. Some properties and thriptransmission of tomato spotted wilt virus inCanada. CanadianJ. Bot. 52:1177-1182.

Pittman. H. A. 1927. Spotted wilt of tomatoes. J.Australian Counc. Sci. Indus. Res. 1:74-77.

Sakimura. K. 1940. Evidence for the identity ofthe yellow-spot virus with the spotted-wiltvirus: experiments with the vector, ThripstabacL Phytopathology 30:281-299.

9

Sakimura. K. 1961. Field observations on thethrips vector species of the tomato spottedwilt virus in the San Pablo area. California.Plant Dis. Rep, 45:772-776.

Sakimura. K. 1963. Frankliniella fusca. anadditional vector for the tomato spotted wiltvirus. with notes on Thrips tabacf, anothervector. Phytopathology 53:412-415.

Samuel. G.. J. G. Bald. and H. A. Pittman. 1930.Investigations on "spotted wilt" of tomatoes.Australian Counc. Sci. Indus. Res. Bull. No.44. 64pp.

Sherf, A. F. 1948. Occurrence of tomato spottedwilt in Nebraska in 1948. Plant Dis. Rep.32:509.

Smith. K. M. 1931. Studies on potato virusdiseases. VIII. On a ringspot virus affectingsolanaceous plants. Ann. AppI. BioI.18:1-15.

Smith. K. M. 1932. Studies on plant virusdiseases. XI. Further experiments with aringspot virus: its identification withspotted wilt of the tomato. Ann. AppI. BioI.19:305-330.

Smith. K. M. 1957. A text book of plant virusdiseases. 2nd ed. Churchill. London. P. 572.

Smith. P. G. 1944. Reaction of Lycopersicon spp.to spotted wilt. Phytopathology 34:504-505.

Snyder. W. C.. and H. R. Thomas. 1936.Spotted wilt of the sweet pea. Hilgardia10:257-262.

Tompkins. C. M.. and M. W. Gardner, 1934.Spotted wilt of head lettuce. Phytopathology24: 1135-1136.

Tompkins. C. M., and H. H. P. Severin. 1950.Spotted wilt of white. yellow. and pinkcallas. Hilgardia 20:207-232.

Yamamoto. T .. and K. Ohata. 1977. Someproperties and electron microscopy oftomato spotted wilt virus isolated fromblackbeny lily (Belamcanda chinensis DC.).Bull. Shikoku Agr. Exp. Sta. 30:39-47.

Yudin. L. S .. J. J. Cho. and W. C. Mitchell. 1986.Host range of western flower thrips.Frankliniella occidentalis (Thysanoptera:Thripidae). with special reference toLeucaena glauca. Environ. EntomoI.15: 1292-1295.

Yudin. L. S .. W. C. Mitchell. and J. J. Cho. 1987.Color preference of thrips (Thysanoptera:Thripidae) with reference to aphids(Homoptera: Aphididae) and leafminers inHawaiian lettuce farms. J. Econ. EntomoI.80:51-55.

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