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    Diseases of Chickpea, Lentil, Pigeon Pea, and Tepary Bean in Continental United States andPuerto RicoAuthor(s): Walter J. KaiserSource: Economic Botany, Vol. 35, No. 3 (Jul. - Sep., 1981), pp. 300-320Published by: Springer on behalf of New York Botanical Garden PressStable URL: http://www.jstor.org/stable/4254300

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    Diseases of Chickpea, Lentil, Pigeon Pea,and Tepary Bean in Continental UnitedStates and Puerto Rico1WALTERJ. KAISER2

    The Leguminosae (pea or bean family)are composed of some 690 generaand18,000 species (Purseglove, 1968).It is the second largestfamily of seed plants(followingthe Gramineae) Aykroydand Doughty, 1964).Within the Legumino-sae, thereare 18-20 species thatare cultivatedwidely for their edible seeds whichare high in protein (17-25+%) (Aykroyd and Doughty, 1964).The seeds of le-gumesaresecondonly to cereals as the most important ource of food for humansand animals(NationalAcademy of Sciences, 1979).The term food legumegenerallyis given to species of Leguminosae,the seeds,pods, and/or leaves of which are eaten by humans. The word pulse is used insome countries colonizedby GreatBritain, ike IndiaandPakistan,to denote thedry, matureseeds which are consumedby humans.A chronic proteindeficiency exists in most developingcountries of the world(Mayer, 1976).In these countries food legumes usually providethe main, and attimes the only, source of proteinand essential aminoacids in the diets of poorerinhabitants or social, economic, or religiousreasons. Legumesare an importantcomplementto diets heavily dependenton high carbohydrate oods (cerealsandroot and tubercrops) (NationalAcademyof Sciences, 1979).

    In the United States and its territories,several food legumes are grown on acommercial scale. After soybean [Glycinemax (L.) Merr.]and peanut (Arachishypogaea L.) (which are also classified as oil crops), bean (Phaseolus vulgarisL.) is the most important ood legume cultivated in the United States (USDA,1979a). The seeds of some food legumes, like bean, pea (Pisum sativum L.),soybean, and lentil (Lens culinaris Medik.)are exported in large quantitiesandaid in correctingthe nation's sizeable balance of trade deficits (USDA, 1979a).In most developed countries of the world, with the exception of Japan, foodlegumes are consumedin smallamounts, and, therefore,contributeminimally osatisfying daily protein requirements.Protein from animalsources satisfies mostof that need, but this is a very inefficientmethod of producingprotein (Mayer,1976).With the recent dramaticrise in the cost of energy and nitrogenous ertil-izers, a searchwill be made to findcheaper,energy-savingmethods of increasingfood production.Food legumes will undoubtedly assume a more importantrolein providinga largershare of the proteinrequirementsof the inhabitantsof manydeveloped countries, includingthe United States, particularlyas the price of1 Received25 August 1980;accepted28 December 1980.Presentedat the Symposiumon Legumes

    at the Twenty-firstAnnualMeetingof the Society for Economic Botany,Bloomington, ndiana,June16-17, 1980;symposiumorganizedby Dr. A. DouglasKinghorn.Mention of a trade name or pro-prietaryproductdoes notconstitutea guaranteeorwarranty f the productby the U.S.D.A. and doesnot implyits approval o the exclusion of otherproducts hat may also be suitable.2 ResearchPlantPathologist,RegionalPlantIntroductionStation,U.S.D.A., SEA, AR, 59 JohnsonHall, WashingtonStateUniversity, Pullman,WA 99164.

    Economic Botany, 35(3), 1981,pp. 300-320(O 1981, by the New York Botanical Garden, Bronx. NY 10458

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    302 ECONOMICBOTANY [VOL.35TABLE 1. CONTINUED.

    Type of diseaseFoliageSeed

    or Mosaic Stem Podseed- Yel- and/or and/or spotling Root low defor- Spot- petiole orCrop Pathogen rota rot Wilt ing mation ting blight rotPhoma sp. + + +Phomopsis sp. + +Physalospora sp. +Phytophthora parasitica +Pleonectria megalospora ?Rhizoctonia ferrugena +Sclerotium rolfsii + + +Uredo cajani + ?Uromyces dolicholi + ?

    Virus or virus-like:Rhynchosia mosaic + +Witches'-broom + +Nematodes:Criconemoides sp. +Helicotylenchus dihystera +Hoplolaimus galeatus +Meloidogyne arenaria +M. javanica +Pratylenchus brachyurus +P. schribneri +Rotylenchulus reniformis +Trichodorus christiei +Tylenchorhynchus

    claytoni +Tepary bean Fungi:(Phaseolus Fusarium solani f. sp.acutifolius phaseoli +var. Sclerotinia sclerotiorum + + + +latifolius) Uromyces phaseolivar. typica + ?

    Bacteria:Pseudomonas phaseolicola + + ? +Xanthomonas phaseolicola + + ? +

    Virus or virus-like:Alfalfa mosaic ? +Bean common mosaic +Bean golden mosaic + +Curly top +Pod mottle +Whitefly-transmittedagents + +

    a+, pathogen capable of producing disease in host; +, pathogen may or may not produce disease in host; ?, pathogenicity ofmicroorganism in doubt.

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    19811 KAISER: LEGUME DISEASES 303

    animalprotein increases and people become more concerned about conservingtheir nation's scarce naturalresources and dwindlingenergy supplies.To anticipate some of the problemsthatmay be encountered n expandingthearea under cultivationof different ood legumes, it willbe necessaryto investigatethe different factors which may adverselyaffect production.Diseases frequentlylimit yields and reduce quality of edible legumes in the United States and else-where.The diseases of some of these crops, like bean(SilbernagelandZaumeyer,1973; Zaumeyerand Meiners, 1975), pea (Hagedorn, 1974, 1976)and soybean(Sinclairand Shurtleff, 1975)have been reviewed elsewhere. It is the purposeofthis paperto discuss the diseases of 4 less common food legumes that are culti-vated in the continentalUnited States and PuertoRico. These crops arechickpea(Cicer arietinum L.), lentil, pigeon pea [Cajanus cajan (L.) Huth], and teparybean(Phaseolus acutifoliusA. Gray).Foreasy reference,the pathogensaffectingthese 4 crops in the continental United States and PuertoRico have been listedaccordingto the type of disease(s) producedin each crop (Table 1).

    CHICKPEA (CICER ARIETINUM L.)Chickpea, also called garbanzo,gram, or Bengal gram,is an annual, self-pol-linated food legume that is cultivatedin manycountries of the world, frequentlyunder semiaridconditions (van der Maesen, 1972).In 1978, chickpeaswere cul-tivated worldwide on 10,481,000ha (26,202,500acres) with over 90% grown in

    India(FAO, 1979).Chickpeas are growncommercially n the United States, pri-marilyin the central coastal areas of California.In 1978, 5,200 ha (13,000acres)were harvested in California California, 1980). However, in the same year, im-ports of chickpea seeds into the United States exceeded $7,500,000 (USDA,1979b). Currently, here is an interest in testing chickpea as an alternativecropin the drylandareasof other states, e.g., northern daho andeasternWashington(Anonymous, 1980;W. J. Kaiser, unpublished).Diseases

    Differentdiseases affect chickpeawhen they are grownunder drylandas com-pared to irrigatedconditions. In some countries, like India or Iran, diseases ofthis cropare an important actorcontributingo low and erraticyields andquality(Kaiser and Danesh, 1971b;Nene et al., 1978;Sen Gupta, 1974).Much of ourknowledgeof chickpeadiseases in the United States is from researchcarriedoutin Californiaduring he last 25-30 yr.Cultivation of chickpeas in the southerncoastal counties of Californiadatesback to the foundingof the Spanishmissionsover 175yr ago (Smithet al., 1950).By the mid-1930s,chickpeas had become a commercialcrop in this regionwithan annualproductionof some 2,500,000lb (1,136,364 kg) (Smithet al., 1950).Ofthe several diseasesreportedto affect chickpeasin this region,Erwin andSnyder(1958)considered virus yellows and Fusarium wilt to be the most serious andwidespread. Symptomsof both diseases in chickpea consisted of yellowing andwiltingof the foliageand death of infectedplants. Symptomsof differentwilt-likedisorders of chickpea are illustrated and described in a recent publication byNene et al. (1978).At least 3 viruses were isolated fromyellows-affected plantsin California Snyder et al., 1956).These were identifiedas bean yellow mosaic,

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    304 ECONOMIC BOTANY [VOL. 35

    1S~~~~~~~~~~~~~~

    Fig. 1-4. Fig. 1. Fusarium root rot (right) s a serioussoilborne disease of chickpea worldwide.Roots of infectedplants(right)are dark n colorandrotted;healthy plants(left).Fig. 2. Lentilplants(lightcolored) n a fieldplantingaredyingdue to rootrot causedby a complexof soilbornepathogens,includingPythium ultimum andRhizoctonia solani. Fig. 3. Infectionof pigeon pea foliageby the rustpathogenmay resultin prematuredefoliationand reducedyields. Fig. 4. Stuntedteparybean plant(left) is infectedwiththe NY 15 strain of beancommon mosaic virus;healthy plant(right).

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    alfalfa mosaic, and pea enation mosaic viruses. Bean yellow mosaic virus wasisolated most frequentlyfrom plants exhibiting symptomsof virus yellows. In-cidence of virus yellows varied fromyear to year but in some plantingsas manyas 50%of the plantswere infected (Snyderet al., 1956).Yields of virus-infectedchickpeaare usuallyreducedby more than50%(KaiserandDanesh, 1971a)andmortality may be high, particularlywhen plants are infected early (Kaiser andDanesh, 1971a;Snyderet al., 1956).The pathogen causing Fusarium wilt of chickpea in the south coastal areas ofCaliforniawas identifiedas Fusarium lateritiumNees emend. Snyder& Hans. f.sp. ciceri (Padwick)Erwin (Erwin, 1958a). The fungus produced typical wiltsymptomsof the shoot and discolorationof the vascular elements. Erwin(1957,1958b) also identified another soil-borne pathogen, Verticillium albo-atrumReinke & Berth., as causing a chickpea wilt disease of minor importanceinCalifornia.The isolates of V. albo-atrum from chickpea were microsclerotialtypes.In recent years, the commercialchickpea-producing reas in Californiahaveshifted northward o the central coastal regionof the state (Phillips, 1979;Wes-terlundet al., 1974).In a survey of 7 chickpea plantingsin California'scentralcoastal area between 1971 and 1972, Westerlundet al. (1974)identified the fol-lowing fungi as causing root rot and wilt diseases of chickpea: Fusariumoxy-sporumSchlecht. f. sp. ciceri (Padwick)Matuo and Sato, F. solani (Mart.)Appel& Wr. f. sp. pisi (E. R. Jones) Snyder & Hans., Macrophominaphaseolina(Tassi) Goid., Pythium ultimum Trow., and Rhizoctonia solani Kuhn. Virus dis-eases are less a problemthan in the south coastal areas(Westerlund t al., 1974).Fusariumsolani f. sp. pisi (Fig. 1)is animportant oot rotpathogenof chickpeain California,as it was shown earlierto be in Washington Kraft, 1969). Wester-lund et al. (1974) solatedF. solani f. sp. pisi from47%of wilted chickpea plants,but F. oxysporumf. sp. ciceri from only 6%. Also, they failed to isolate F.lateritiumf. sp. ciceri from diseased chickpeas and they were unable to provethe pathogenicityof severalof Erwin'soriginal solates of thisfungusto chickpea.However, Erwin's originalisolates that were tested by Westerlundet al. (1974)may have lost theirpathogenicity o chickpeaafterbeing kept in culture for over15 yr. Both Fusariumwilt and root rot were severe in fields previously croppedto chickpea, but they were not found in fields planted to chickpea for the firsttime (Westerlundet al., 1974).Pythium ultimum and Rhizoctonia solani caused pre- and post-emergencedamping-offand necrosis and death of small chickpearoots in California Smithet al., 1950;Westerlund et al., 1974). Both pathogens were isolated most fre-quently from diseased plants on land with no previous history of chickpea pro-duction. Pythiumultimum s also the cause of a seriouspre-emergencedamping-off disease of white-seededchickpeasin the Palouse region of eastern Washington(Kaiser and Hannan, 1981).Emergenceof nontreatedchickpeaseeds was reducedby 60-80% or more by P. ultimumin Palouse soils with no previous history ofchickpea cultivation.Macrophominaphaseolina may induce a wilting disease of chickpea plantsgrownunderdrylandconditions,butonly where plants receiveless than adequatewater (Westerlundet al., 1974).This pathogen generally is not a problem in thefield when adequatesoil moisture is maintained.

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    306 ECONOMIC BOTANY [VOL. 35

    Erwin (1958a)found 4 chickpea lines that had high levels of resistance to F.lateritiumf. sp. ciceri. These lines need to be tested for their resistance to Fu-sariumwilt causedby F. oxysporum . sp. ciceri. In 1978,Phillips 1979)screened279 lines from the InternationalCropsResearchInstitute for the Semi-AridTrop-ics (ICRISAT), Hyderabad,India. Thirty-fouraccessions remained free of wiltsymptoms.Theprevalence of Fusariumdiseases only infieldspreviouslycroppedto chick-pea suggests the importanceof crop rotation(Westerlundet al., 1974).Both F.solani f. sp. pisi (Westerlund t al., 1974)andF. oxysporum . sp. ciceri (Hawareet al., 1978)are carriedon the seed which is one methodby which Fusariumwiltand root rot can be introduced nto new areas. Haware et al. (1978)showed thatthe wilt Fusarium was eradicated from seed by treatmentwith a mixture ofBenomyl (methyl-1-[butylcarbamoyl]-2-benzimidazolecarbamate)nd Thiram(bis-[dimethylthiocarbamoyl]disulfide).Ascochyta blight [Ascochyta rabiei (Pass.) Lab.] is a potentially devastatingseedborne disease of chickpea which until recently was not reportedfrom theWestern Hemisphere. In 1974, Ascochyta blight caused extensive damage toexperimentalchickpeaplantingsat Saskatoon, Canada(Morralland McKenzie,1974).Wet, cool weather favors disease development.Apparently,the pathogenwas introduced ntoCanadaon importedchickpeaseed. Since A. rabiei is readilyseedborne, care shouldbe exercised when chickpea seeds are imported nto theUnited States for research or commercialpurposes. Additionalresearch is re-quired to determinewhetherA. rabiei can be eradicatedfrom chickpea seed bythermotherapyand/or chemical means. Preliminaryresults indicated that treat-ment of Ascochyta-infected chickpea seed with certain chemical compounds,especially the benzimidazoles, greatlyincreasedemergenceand reduced diseaseincidence and severity in seedlings (Kaiser et al., 1973).

    LENTIL (LENS CULINARIS MEDIK.)The cultivated entil is a self-pollinated,cool-season annual hat is of OldWorld

    originand one of the first food crops to be cultivatedby man (Zohary, 1976).Inthe United States, lentilsaregrowncommercially n the Palouse region of easternWashingtonand northern Idaho in rotation with wheat (Triticumaestivum L.)anddry-ediblepeas. Lentils werefirstplanted n the Palouse in 1916(Youngman,1968), and have since become an importantcash crop. In 1978,lentils were har-vested from 52,400 ha (131,000acres) (Idaho, 1979;Washington, 1978).Much ofthe lentil crop producedin the Palouse is exported. The value of lentil exportsin 1978exceeded $15,000,000(USDA, 1979b).Diseases

    Consideringthe importance of lentil as an alternatecash crop in the PacificNorthwest, little informationhas been published on its diseases in the UnitedStates. The firstpublishedreportof fungusdiseases affecting entils in the PacificNorthwest was by Wilson and Brandsberg n 1965.They made numerousisola-tions from diseased, field-grown lentil seedlings over a 2-yr period and foundseveral soilbornefungito be incitantsof root (Fig. 2) and stem diseases of lentilin the Palouse region. Fungipathogenic in theirgreenhouse andfield inoculation

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    19811 KAISER: LEGUME DISEASES 307

    studies included Ascochyta pinodella L. K. Jones, Botrytis cinerea Pers. ex Fr.,Fusarium oxysporum (possibly F. oxysporum f. sp. lentis Vasudeva & Sriniva-san), Fusarium roseum (Lk.) Snyd. & Hans. 'Gibbosum,' Rhizoctonia sp., Scle-rotinia sclerotiorum (Lib.) d By., and Verticillium albo-atrum. In the Palouseregion, Rhizoctonia solani has been isolated from discolored roots of diseasedlentils and isolates of the funguswere pathogenicto germinating eeds androotsof inoculatedlentils (W. J. Kaiser, unpublisheddata). With high moisture con-ditions, S. sclerotiorumcan cause severe damageto the lentil crop in this region(Amer. Dry Pea & Lentil Assoc., 1980).Fusarium wilt caused by F. oxysporum . sp. lentis is one of the most importantdiseases of lentil in some lentil-producing ountries, such as India (Sen Gupta,1974). Wilson and Brandsberg(1965) isolated cultures of F. oxysporumfromdiseased Palouse lentils that were similarin cultural and morphologicalcharac-teristics and in pathogenicityto a known isolate of F. oxysporumf. sp. lentis.Their pathogenicisolates of F. oxysporumcaused a root rot andvascularwilt ofinoculated lentils.AlthoughBotrytiscinerea was isolated infrequently rom diseased lentil seed-lings, Wilson and Brandsberg 1965)consideredit to be a potentially importantpathogen of lentil in the Pacific Northwest. Recent studies have demonstratedthat B. cinerea is transmitted n a low percentageof seed of field-grownPalouselentils (W. J. Kaiser, unpublisheddata).Foliar diseases of lentil are usually of minorimportance n the Palouse. Fungiassociated with foliar diseases include Alternaria tenuis Nees. [=A. alternata(Fries) Keisslers], Cladosporium herbarum Pers. ex Fr., and Stemphylium bot-ryosum Wallr. (Wilson and Brandsberg, 1965). In Canada, Alternaria sp. andCladosporiumsp. also were associated with leaf blight diseases of lentil (Mc-Kenzie and Morrall, 1975).Lentils are a cool season crop with an indeterminategrowthhabit. High tem-peraturesadversely affect growth and floweringof lentils and may predisposeplantsto disease. For instance, in greenhousestudies, Lin andCook (1977)dem-onstratedthat F. solani f. sp. pisi, normallya root rot pathogenof pea (Pisumsativum), caused a black root rot of lentil at high (25-30 C), but not at low (

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    308 ECONOMIC BOTANY [VOL. 35

    tified diseases of a crop. An example of this phenomenonwith lentils was dem-onstratedrecently by Lin and Cook (1977).They showed how a previouslyun-diagnosed root rot disease of lentil caused by Fusarium roseum 'Avenaceum'increased in importancewhen nontilled lentils were direct-seeded into undis-turbed(zero tillage),dead bluegrasssod in easternWashington.The lentil crop in the Pacific Northwest is free from the ravages of a fewimportant and destructive diseases. Notable among these is Ascochyta blight(Ascochyta sp., possibly A. lentis Bond. & Vassil.), a seedborne disease thatattacksthe leaves, stems, pods, andseeds of lentil. Ascochyta blightwas reportedrecently in Canada(Slinkardand Drew, 1980), where it is considered to be apotentially importantdisease of lentil (R. A. A. Morrall, personal communica-tion). Every effort should be made to exclude this and other potentiallydevas-tating diseases fromlentil-growingareas of the United States.In Iran, lentils are naturally infected by alfalfa mosaic (AMV), bean yellowmosaic (BYMV),cucumbermosaic, andpea leaf roll viruses(Kaiser, 1973),whilein Italy, they are a naturalhost of pea enationmosaic virus (PEMV)(Vovlas andRana, 1972).Information s meageron the distribution, ncidence,andimportanceof virusdiseases of lentil in the United States. In the Palousearea,several viruseshave been isolated from diseased lentils, includingAMV, BYMV-typeisolates,PEMV, and pea streak virus (R. 0. Hampton,personal communication).Inci-dence of virus infection appearsto vary greatly from year to year and may berelatedto migrationsof aphidvectors andthe proximityof virus-infectedalternatehosts to lentil plantings.Virus diseases of lentil is an area that meritsadditionalresearch.Pea seedborne mosaic virus (PSbMV) may be a potential threat to the lentilcrop in the PacificNorthwest (Hamptonand Muehlbauer, 1977).However, pri-maryor secondaryinfection of lentilby PSbMV has not yet been observed in thefield. PSbMV is easily transmitted rompeas, a naturalhost of the virus, to lentilsby artificialmeans and is then seed-transmitted n lentil at frequencies of 32-44%(Hampton and Muehlbauer,1977). Four lentil lines in the USDA plant introduc-tion (PI) collection were found to be immuneto PSbMV(Haddadet al., 1978).

    PIGEON PEA [CAJANUS CAJAN (L.) HUTH.]Thepigeon pea is a woody, short-livedperennialshrub hat is cultivatedwidelyin the tropics and subtropics, often as an annual. It is a drought-tolerant ropfrequently grownin semiaridregions on poor soils (De, 1974; Purseglove, 1968;Royes, 1976). Pigeon peas, also called arhar, red gram, Congo pea, and gandul,are cultivatedon some 3,000,000ha (7,500,000 acres) worldwide, with over 90%of the production n India (Rachieand Roberts, 1974).In the Americantropics,the pigeon pea is an important ood cropwhere the seeds areconsumed as green(immature)or dry (mature)peas. In some countries, it is also used as forage foranimals or as a greenmanureor cover crop (Purseglove, 1968;Morton, 1976).The Commonwealthof Puerto Rico is the only location in the United Stateswhere pigeon peas are presentlygrownas a commercialcrop. Pigeon pea is themost important ood legume cultivatedin Puerto Rico, where an importantcan-ning industry has developed aroundthis crop. During 1975-1976, pigeon peaswere harvested from approximately6,000 ha (15,000 acres) (ConjuntoTechno-l6gico, 1977).However, due to anincreaseddemand or fresh, canned, andfrozen

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    pigeon peas, over 870,000 kg were imported nto PuertoRico andthe continentalUnited States during1978-1979(USDA, 1979c).Diseases

    A number of diseases of varying severity affect the roots, stems, petioles,leaves, pods, and seeds of the pigeon pea in tropicaland subtropicalareasof theUnited States, includingFlorida, Hawaii, PuertoRico, andTexas (Barnes, 1973;Nene, 1978; Spence, 1975;USDA, 1960).The diseases are caused primarilybyfungi, althougha few disordersof unknownetiology with virus- or mycoplasma-like symptomshave been observed in Puerto Rico (Maramorosch t al., 1974a,b;Birdet al., 1975;Vakiliand Maramorosch,1974).Much of the pioneeringresearch on pigeon pea diseases in the WesternHemi-spherewas done by researchers ocated at the MayagiiezandRlo Piedrasexper-iment stations in Puerto Rico, beginningin the early 1900s. A majorityof theearly studies dealt with diseases caused by fungi, particularly hose incitingdis-eases of the foliage, stems, and pods.Over 60 yr ago, Stevenson (1917) reportedon the etiology and occurrenceofseveral diseases of pigeon pea in PuertoRico. These were leaf spots caused byCercospora cajani P. Henn. [=Mycovellosiella cajani (P. Henn.) Rangel &Trotter], Uromyces dolicholi Arth. and an unknownstem canker disease, pos-sibly of fungaletiology.

    Two rust diseases have been reportedto infectpigeon pea in the United States(CommonwealthMycologicalInstitute, 1968;Nene, 1978;Spence, 1975;Tucker,1927a;USDA, 1960).These rusts were identifiedas Uredocajani Syd. and Uro-myces dolicholi. Some confusion exists concerningthe proper identificationofthese 2 fungion pigeon pea foliage. Differentiations possibleonly whenthe telialspore stage is foundandthis has yet to be observed in U. cajani (CommonwealthMycological Institute, 1968).Rust is a potentially mportantdisease of pigeonpeain the Caribbean Fig. 3). In Jamaica and Trinidad, Spence (1975) observed thatsevere rust infection resulted in the shedding of infected leaves and may beresponsible for significantyield reductions. Many of the dwarfpigeon pea vari-eties developed by the University of the West Indies are highly susceptible torust infection. Pigeon pea varieties recommendedfor cultivation in Puerto Ricoare tolerant to rust (ConjuntoTechnologico, 1977).Differentstrains of U. cajanimay exist in the Caribbeanarea (Spence, 1975).Two Cercospora species, C. cajani and C. instabilis Rangel, cause leaf spotdiseases of pigeon pea in Puerto Rico (Nene, 1978;Stevenson, 1917, 1918;Tuck-er, 1927a;USDA, 1960).Infection may result in prematuresheddingof leaves.The Cercospora lesions on pigeon pea leaves are similar to those caused byColletotrichum cajani. Cercospora leaf spots are common diseases of pigeon peain Puerto Rico and are generallyof minorimportance(Tucker, 1927a).Anthracnosecaused by Colletotrichum ajani is an important oliar disease ofpigeon pea that was first describedin Brazil in 1916 (Rangel, 1916). In 1925, C.cajani caused a destructive disease of the leaves, pods, and seeds of pigeonpeain Puerto Rico (Tucker, 1927a,b).The disease was most severe duringperiodsofheavy rainfall. Infection by C. cajani frequently resulted in spotting, necrosis,deformation, and/orsheddingof pigeon pea pods and leaves, and shrivelingandreductionin size of infectedseeds (Tucker, 1927a,b).Symptoms of C. cajani on

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    young pigeon pea leaves and pods were similar to those frequentlyobserved onbeans (Phaseolus vulgaris) infected with anthracnose caused by C. lindemuthian-um (Sacc. & Magn.) Briosi & Cav. However, the bean anthracnosepathogendoes not infect pigeon pea nor vice versa.Periodically, stem and crown canker diseases of pigeon peas have occurredinepidemic proportionsin Puerto Rico. A Phoma sp. was reportedby AlvarezGarcia (1960)and Lopez Rosa (1969)to cause a serious stem canker disease ofpigeon pea in commercialplantingson the island. The pathogen,which is trans-mittedin a low percentageof seeds (Lopez Rosa, 1969)affects both seedlingsandolder plants, sometimes resulting in death of infected plants (Aponte Aponte,1963). Pycnidia of the pathogen developed in lesions on stems, branches, andpods (Alvarez Garcia, 1960). Phoma isolates appearedto vary greatly in theirpathogenicity to pigeon pea. A few were highlyvirulentto Kakipigeon pea, butthe majoritywere avirulent(Lopez Rosa, 1969). Lopez Rosa (1969)observed theperfect stage of Phoma on agar media, and found it to be a heterothallicasco-mycete. Ascospore culturesformedpycnidiain 4-5 days. No mention was maderegarding he identityof the perfectstagenor whether it was pathogenic o pigeonpea. Earlier, Leach and Wright (1930) had reportedthat a collar rot and stemcanker disease of pigeon pea in Trinidadwas caused by Physalospora sp., anascomycete, which had2 pycnidialstages belonging o Phoma andMacrophoma.Phytophthoraparasitica Dastur is a potentially importantpathogen of pigeonpea in Puerto Rico (Kaiserand Melendez, 1978;Abramset al., 1978). Necrotic,depressed cankers developed on stems, branches, and petioles of naturally n-fected and artificially noculatedplants. Cankersup to 6 cm in lengthfrequentlygirdledstems, causing branches above infection sites to wilt and die. Several foodcrops grown in rotation with pigeon pea in Puerto Rico were also hosts of P.parasitica, notably eggplantand tomato(KaiserandMelendez, 1978).Pigeon peaisolates of the pathogenwere more virulent o pigeon pea thanthose from eggplantand tomato. Woundingof pigeon pea tissues favored infection. In India, otherPhytophthora spp. have been identified as the cause of a stem blight disease ofpigeon pea (Amin et al., 1978; Kannaiyanet al., 1980; Pal et al., 1970; Williamset al., 1975).Stevenson (1917, 1918) solated several fungi fromthe dead wood andbark ofpigeon pea plants in Puerto Rico. He believed that some of these fungi wereinvolved in the death and rotting of stems of mature plants, particularlyMega-lonectria pseudotrichia (Schw.) Speg., Creonectria grammicospora (Ferd. &Winge) Seaver (=Nectria grammicospora Ferd. & Winge), and Pleonectria meg-alospora Speg. [=Thyronectria megalospora (Speg.) Seaver & Chardon] (Ste-venson, 1917, 1918).However, pathogenicity of these fungi to pigeon pea wasnot tested. Tucker (1927a)reported that perithecia of Botryosphaeriaxantho-cephala (Syd. & Butl.)Th. & Syd. were found in the bark of diseased pigeon peaplants in Puerto Rico. AlthoughB. xanthocephala was associated with a seriouscanker disease, Tucker (1927a)did not demonstrate its pathogenicityto pigeonpea.Tucker (1927a)reportedRhizoctonia errugena Matz. to cause a pre- and post-emergence damping-offdisease of pigeon pea seedlings in PuertoRico and thatthe disease was most destructive during wet weather. He found the host rangeof R. ferrugena was extensive and includes several vegetable crops that were

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    grown in rotation with pigeon pea. Parmeterand Whitney(1970)observed thatan isolate of R. ferrugena obtained fromthe Centraalbureau oor Schimmelcul-tures, The Netherlands,had clamp connections and formed sclerotia similartothose of Sclerotiumrolfsii Sacc. It is possible thatTuckermayhave been dealingwith a fungusresemblingS. rolfsii. Sclerotiumrolfsii was isolated fromdead anddying pigeon pea seedlingsfrom Isabela, PuertoRico, and the fungusproved tobe pathogenic to seedlings of this crop in greenhouseinoculationstudies (W. J.Kaiser and P. L. Melendez,unpublisheddata).Recently in PuertoRico, the role of internallyseedbornefungion germinationand emergence of seeds in the fieldwas revealed (Ellis and Paschal, 1977, 1979;Ellis et al., 1977, 1978).During 1976,emergenceof seed in several commercialplantings was low (28-45%)and it appeared hat the high incidence of internallyseedbornefungiwas a primary actorcontributing o poor seed germination, huslimitingcommercialproductionof pigeonpea in Puerto Rico (Ellis and Paschal,1979).Fungi frequently solated frompoor qualitypigeonpea seed (as measuredby seed germinationand seedling emergence)were Alternariatenuissima (Fr.)Wilts., Fusarium semitectum Berk. & Rav., Lasiodiplodia theobromae Pat.,Phomopsis sp., Aspergillus sp., Cladosporium sp., and Penicillium sp. Therewas a negativecorrelationbetween the occurrence of A. tenuissima,F. semitec-tum, L. theobromaeand Phomopsis sp. and in vitro germinationandfieldemer-gence of pigeon pea seed (Ellis et al., 1977).Pigeon pea seed quality also wascorrelatedwiththe site of production nPuertoRico. Thequalityof seed producedat Isabela, PuertoRico, a high-rainfallarea, was poor in comparisonwith seedproduced at Fortuna,Puerto Rico, an arid, low-rainfallarea (Ellis et al., 1978).Delaying the harvest of seed past maturityfrequently resulted in a significantincrease in the incidence of Phomopsis sp., A. tenuissima, F. semitectum, andL. theobromae. Treatmentof pigeon pea seed from Isabela, Puerto Rico, withCaptan (N-trichloromethylthio-4-cyclohexene- ,2-dicarboximide)and Thiram(protectantfungicides) or Benomyl (systemic fungicide) significantly ncreasedgerminationand field emergence over nontreatedseeds. Likewise, foliar appli-cations of Benomyl several times during he growing season significantly educedsubsequentinfection of seed by several internallyseedbornefungi that played amajorrole in reducingseed quality.Recent investigationswith internallyseedbornefungihave emphasizedthe im-portantrole that these microorganismsplay in reducingqualityof seed of pigeonpea and other food legumes producedin Puerto Rico (Ellis et al., 1977).Addi-tionally, they point out the need of selecting suitable sites in Puerto Rico, likethe south coastal area, for the productionof high quality seed for use in estab-lishingnew plantings(Ellis et al., 1978;ConjuntoTechnologico, 1977).A few virus- andmycoplasma-likediseases of uncertainetiology affect pigeonpea in Puerto Rico and other islands in the Caribbean Bird and Sanchez, 1971;Birdet al., 1975;Maramorosch t al., 1974a,b).Since the late 1950s,Bird andhiscolleagues (1971, 1975) n PuertoRico have studiedvarious 'rugaceous'diseasesthat affect differentfood crops and weed species in the Caribbean.These ruga-ceous diseases aretransmittedby whiteflies(Bemisiatabaci Genn.). One of thesediseases, Rhynchosia mosaic, infects pigeon pea naturally n Puerto Rico, pro-ducinga yellow mosaic symptom(Bird et al., 1975).The primaryreservoirof theyellow mosaicpathogen s RhynchosiaminimaDC., a wildlegume. Several high-

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    yielding pigeon pea lines developed in Puerto Rico have resistance to yellowmosaic (Birdet al., 1975).A witches'-broomdisease of pigeonpea has been reportedfrom several coun-tries, includingPuertoRico, DominicanRepublic,and Haiti (Hirumiet al., 1973;Maramorosch t al., 1974a,b;Nene, 1978;VakiliandMaramorosch,1974).Symp-toms of witches'-broomfrequentlyconsisted of a dwarfingof leaves and a pro-liferationof shoots at the nodes. Electronmicrographsof pigeonpea tissues fromwitches'-broom-infectedplantsrevealed the presence of mycoplasma-likeorgan-isms andrhabdovirus-like articles(bullet-shaped)n phloemcells (Hirumiet al.,1973;Maramorosch t al., 1974a,b).The vector of the disease is unknown. Therewas a highincidence of witches'-broom n Haiti. This disease is a potentialthreatto pigeon pea cultivationin PuertoRico (Vakiliand Maramorosch,1974).In the Caribbean,surveys of agricultural oils have shown that various plantparasiticnematodes are associated with the roots of pigeon pea (Ayala, 1962a,b;Singh, 1975). High populationdensities of differentecto- andendoparasiticnem-atode species aroundpigeon pea roots result in damageto the roots which con-tributeto poor growthand reducedseed yields from diseasedplants. As earlyas1911, pigeon peas were reported to be a natural host of rootknot nematodes[Heterodera radicicola (Greef) Muller (=Meloidogyne spp.)] in the United States(Bessey, 1911). In Puerto Rico, pigeon pea roots are attackedby Meloidogynejavanica (Treub.) Chitwood, Rotylenchulus spp. (including R. reniformis Linford& Oliveira), Criconemoides sp., Helicotylenchus sp., and Hoplolaimus sp. (Ay-ala, 1962a,b). Rotylenchulusreniformis appearsto be the most importantnem-atode pathogenof the roots of pigeonpea and several other food crops in PuertoRico (Ayala, 1962a;Ayala and Ramirez, 1964;Steiner, 1960).In Hawaii duringthe 1930s,severallegumes, includingpigeon pea, were tested as potentialrotationand trap crops for pineapple on land that was heavily infested with rootknotnematodes. Pigeon peas provedto be one of the most rootknot-resistantegumestested (Godfrey, 1928).In greenhouse studies, pigeon peas were shown to be a good host for severalecto- and endoparasiticnematode species when planted in soils from cotton,peanut, and soybean fields in Alabama(Ingramand Rodriquez-Kabana,1977;Rodriquez-Kabana ndIngram, 1978).These includedHelicotylenchusdihystera(Cobb) Sher, Hoplolaimus galeatus (Cobb) Thorne, Meloidogyne arenaria Chit.,Pratylenchus brachyurus (Godfrey) Filipjev and SchuurmansSteckhoven, P.schribneri Steiner, Trichodorus christiei Allen, and Tylenchorhynchus claytoniSteiner.

    TEPARY BEAN (PHASEOLUS ACUTIFOLIUS A. GRAY)The teparybean is a drought-tolerant nnualthatoriginated n the New World,probably in the southwestern United States and northwesternMexico (Evans,1976;NabhanandFelger, 1978;National Academyof Sciences, 1979;Purseglove,1968).From archeologicalremains, it appearsthat the teparybean was a domes-ticated crop in Mexico over 5,000 yr ago. Tepariesare still found growing in thewild at elevations up to 1,650 m as far south as Guatemalaand as far northascentralArizona(Nabhanand Felger, 1978). Cultivated ines of P. acutifoliusareclassified as var. latifolius Freeman.Tepary beans are adapted to hot, ariden-vironments and will usually produce a crop in 60-90 days under these harsh

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    conditions where other Phaseolus species, like bean (P. vulgaris) would fail.They will not tolerate frost or waterloggedsoils. Hendry (1918) reported thattepary seeds germinatedrapidlyin soils with low moisturecontent, but rottedquickly in cold, moist soils. Teparieshad become an important ield crop in thearid Southwest by 1918 where in Californiaalone they were cultivated on some6,800 ha (17,000 acres) (Hendry, 1919), but their importancedeclined rapidlythereafter,due to a combinationof factors, includingsmall seed size, flavorandodor when cooked, changing and-usetrends, and culturalpreferences(Hendry,1918;Nabhan and Felger, 1978;National Academy of Sciences, 1979). Todayteparybeans are cultivated for their edible dry seeds on a limited scale by com-mercial and subsistencefarmers n southwesternNorth America(Nabhan, 1979;Nabhan and Felger, 1978).Teparies are a potentiallyimportanthigh-protein ood legume suited to culti-vation in the arid andsemiaridregionsof the world. It maybe possible to transfersome of the teparybean's desirablecharacteristics,such as tolerance to droughtand heat, to other Phaseolus species by interspecific hybridization(NationalAcademy of Sciences, 1979;Waines, 1978).Diseases

    Very little information s available on the diseases affectingwild or domesti-cated teparybeans undernaturalconditionsin the aridregionsof southwesternNorth America. In one of the earliest reportson cultivation of this crop in theUnited States, Freeman(1912, 1913) mentions that tepary beans differed frombeans in being more resistant to heat, drought,and insect pests. However, nomention was made of diseases. Subsequently, several reportshave appeared nthe literatureon the susceptibility or resistance of tepary bean accessions todifferent diseases. A majorityof these reportsare based on infectionof tepariesby variouspathogens n field orgreenhouse rials,butnot undernatural onditionsin farmers' fields. A recent report by the National Academy of Sciences (1979)on tropical legumes states that while some tepary bean accessions are highlyresistant to the bean common blight bacterium[Xanthomonasphaseoli (E. F.Sm.) Dows.], they areonly mildlyresistant to most otherdiseases andpests. Ourknowledge of the diseases affecting tepary beans in nature is fragmentaryandincomplete.Tepary beans are reportedto be infectedeitherexperimentallyor naturallybyseveral viruses. In 1922,Nelson (1932)observed natural spreadof a seedbornebean virus, presumablybean commonmosaic virus (BCMV)fromvirus-infectedbeans to differentPhaseolus species, including eparybean, in field trials at EastLansing, Michigan.Severaltepary bean accessions were shown to be susceptibleto natural and artificial nfection in PuertoRico and Iran, respectively, by 2 ormore strainsof BCMV(Kaiser and Mossahebi, 1974;W. J. Kaiser, unpublisheddata). The virus was also transmitted n 20-25% of the seed harvested fromthePuerto Rican BCMV-infectedtepary bean plants (W. J. Kaiser, unpublisheddata).ProvvidentiandCobb (1975)reportedseed transmissionof BCMVranging rom7-22% in 4 P. acutifoliusPI (USDA plantinventory)accesssions. All inoculatedplantsof 15 P. acutifoliusaccessions were susceptiblein greenhouseinoculationstudies to 3 isolates of BCMV from P. vulgaris (Provvidentiand Cobb, 1975).

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    Narrowing, mottling, deformation,and twistingof the leaflets and stuntingweresymptoms frequentlyobservedin teparybeanplantsinfectedwith BCMV(Kaiserand Mossahebi, 1974;Provvidentiand Cobb, 1975) (Fig. 4).In addition to BCMV, teparies were experimentally nfectedby other viruses,notably alfalfamosaic (ZaumeyerandThomas, 1957),bean yellow mosaic (Zau-meyer and Thomas, 1957), curly top (USDA, 1960), pod mottle virus (Zaumeyerand Thomas, 1957), and at least 4 whitefly-transmitted Bemisia tabaci) mosaicdiseases of Phaseolus lunatus L., Jacquemontia tamnifolia Griseb., Merremiaquinquefolia Hall., and Rhynchosia minima (Bird et al., 1975; G'amez, 1971). Themosaic disease of P. lunatus, also called bean golden mosaic or bean goldenyellow mosaic, is an importantdisease of bean andlima beanin severalcountriesof the Caribbeanand Centraland South America(Birdet al., 1975;Costa, 1975;

    Gamez, 1971). Granillo et al. (1975) reportedthe high incidence of a whitefly-borne disease of P. acutifolius in varietytrialsin the coastal areaof El Salvador.The disease was believed to be due to bean golden yellow mosaic.The roots of tepary bean are susceptible to Fusarium solani f. sp. phaseoli(Burkh.) Snyd. & Hans., a soilbornefungusthatcauses a seriousroot rot diseaseof bean worldwide(USDA, 1960;Zaumeyerand Thomas, 1957).Fusarium rootrot could become an importantdisease of P. acutifolius if they are plantedonland with a history of bean root rot. This Fusarium disease is frequently moreserious on beans that aregrowingunder water stress (SilbernagelandZaumeyer,1973)as may occur when beans and teparybeans are grownunderdrylandcon-ditions in arid and semiaridregions of the world. Teparybeans are susceptibleto white mold (Sclerotinia sclerotiorum)in California(J. G. Waines, personalcommunication).Teparieswill likely prove to be hosts to other soilbornefungiand nematodes that are pathogens of bean, notably Pythium spp., Rhizoctoniasolani, Thielaviopsis basicola (Berk. & Br.) Ferr., Sclerotium rolfsii, and Me-loidogyne spp. (USDA, 1960; Silbernageland Zaumeyer, 1973; Zaumeyer andThomas, 1957).Three of the diseases reportedto affect the foliage of teparies are bean rust[Uromycesphaseoli (Pers.)Wint. var. typica Arth.] (USDA, 1960;ZaumeyerandThomas, 1957),bean commonblight (X. phaseoli) (Zaumeyerand Thomas, 1957),and bean halo blight [Pseudomonasphaseolicola (Burkh.)Dows.] (M. L. Schus-ter, personal communication).The latter2 bacterial pathogens are seedborne inbean and probablywouldbe transmitted n seed of susceptible tepary bean lines.Spreadand disease developmentof both bacterialdiseases are favored by rainand humid weatherconditions.The leaves and pods of many P. acutifolius accessions are highly tolerant tomost isolates of X. phaseoli (Schuster, 1955;Schuster et al., 1973).Honma (1956)succeeded in makingan interspecificcross between P. vulgaris andP. acutifoliusby employingembryoculturetechniques.He obtained4 F1plants which appearedsimilar to the P. vulgaris parent. Progenyfrom these 4 F1plants varied greatlyin their reactionto X. phaseoli (Honma, 1956). Field and greenhouse screeningof the progenyfromthis interspecificcross eventually resulted in the release ofthe variety Great Northern(GN) Nebraska #1, Sel. 27, which exhibited a highlevel of tolerance to X. phaseoli isolates from the United States (Coyne andSchuster, 1974;M. L. Schuster, personal communication).Recently, GN Ne-

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    braska #1, Sel. 27 was shown to be slightly susceptibleto isolates of X. phaseolifrom Uganda and Colombia(Schusteret al., 1973).This variety is also tolerantto P. phaseolicola races 1 and2 (CoyneandSchuster, 1974).Since it was releasedinthe mid-1960s,GN Nebraska #1, Sel. 27 hasbeen used as a source of resistanceto common bacterial blight and halo blight by bean breedersin many countries(M. L. Schuster, personal communication).In Puerto Rico, Vakili (1976) screened 151 P. acutifolius lines (many of whichwere PI accessions) for resistance to several diseases, includingbacterialblight,bean commonmosaic virus, root rot, andrust. He selected several lines that hadmoderate o high resistanceto one or more of these diseases. Seed of the multiple-disease resistant selections were being increased for release to interested re-searchers.

    STRATEGIES FOR CONTROLLING DISEASES OF FOOD LEGUMESVarious methods are used to control diseases affectingedible legumes in theUnited States and its territories.These include the use of chemicals, resistantvarieties, and culturalpractices.Economic and environmentalconsiderationsprobablywill lead to a significantreductionin the applicationof pesticides to control foliarand soilborne diseasesof many crops like the food legumes. However, applicationof fungicidesto seedshould continueto be an importantmethod of controllingdamping-offand other

    seedlingdiseases.Culturalpracticescontinue to be used extensively to controlsoilbornediseasescaused by fungi and nematodes that attack the roots of edible legumes. Croprotationis one method commonly used to control soilborne diseases that have anarrowhost range, e.g., the Fusariumwilt or rootrot diseases of chickpea.Othercultural methods, like eradication of overwintering and/or alternativehosts ofcertain pathogens or removal of crop residues may be requiredto achieve asatisfactorylevel of controlof certain diseases.Host plant resistance offers the most promisingandpotentially mportantmeth-od of controllingdiseases of many food crops. Disease resistance has proveduseful in controllingmanydiseases of edible legumes (National Academy of Sci-ences, 1972, 1979),but much more remains o be done in developmentof multipledisease- and pest-resistantcultivarsof these food crops. It is possible that moreemphasiswill be given to host plantresistance as costs of othercontrol measuresthat depend heavily on the use of fossil fuel energy and other scarce resourcesrise. This method of control requires that researchershave access to a large,variable source of germ plasm of differentfood legumes to use in their breedingprograms.Plantgerm plasmcollections of numerous ood crops have been main-tainedat various locationsin the United Statesfor many years. These germ plasmcollections are an integralpartof the National PlantGermplasmSystem (NPGS).The NPGS is a coordinatedeffort among federal, state, and private organizationsin the United States to collect, introduce, maintain,evaluate, catalog, and dis-tribute all types of plant germ plasm (USDA, 1977).Most or all plant germ plasmavailableto researchers romgerm plasm centersin the United States have PI (USDA plant inventory)accession numbersassignedto them. Thegerm plasmcollections for the 4 legume cropsreviewed in this paper

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    are located at 2 of the 4 state-federalregional plant introductionstations in theUnited States:Pullman, Washington:chickpea, lentil, tepary beanExperiment, Georgia:pigeon peaSignificantstrides have been made in the last 30 yr in controllinga numberofimportantdiseases of food legumes andother crops. However, much remainstobe done in preventingcrop losses due to diseases and pests, particularly n thedeveloping countrieswherechronic food shortagesare a common occurrenceandcould worsen as the populationsof these countries increase. New approachesofdisease control must conserve resources and energy and promote a clean envi-ronment. The research challenges are great and the opportunities or success arepromising.

    LITERATURE CITEDAbrams,R., A. Morales,and F. J. Julia. 1978. Statusof research on pigeonpeas in Puerto Rico.Trop.GrainLegumeBull. 11& 12: 17-21.Alvarez Garcia,L. A. 1960. Phomacanker of pigeonpeasin Puerto Rico. J. Agric. Univ. PuertoRico 44: 28-30.Amer. Dry Pea & Lentil Assoc. 1980. Bull. 51 (July 18, 1980).Amin,K. S., B. Baldev,and F. J. Williams. 1978.Phytophthora ajani, a new speciescausingstemblighton Cajanuscajan. Mycologia70: 171-176.Anonymous. 1980. WashingtonFarmer-Stockman05(20):12A.AponteAponte, F. 1963. El cultivo de gandulesen PuertoRico. CaribbeanAgric. 1: 191-197.Ayala, A. 1962a. Pathogenicityof the reniformnematode on various hosts. J. Agric.Univ. PuertoRico 46: 73-82.. 1962b. Occurrenceof the nematodeMeloidogyneavanica on pigeonpea roots in PuertoRico. J. Agric. Univ. PuertoRico 46: 154-156., and C. T. Ramirez. 1964. Host-range,distribution,andbibliography f the reniformnema-tode, Rotylenchulus eniformis,with specialreference to PuertoRico. J. Agric. Univ. PuertoRico 48: 140-161.Aykroyd,W. R., and J. Doughty. 1964. Legumes n HumanNutrition.FAONutritionalStudiesNo.19. Food andAgricultureOrganization f the UnitedNations, Rome.Barnes,R. F. 1973. A preliminaryist of literatureon pigeonpea [Cajanus cajan (L.) Millsp.]plantpathology. Univ. West Indies,Dept. Biol. Sci. Bull. No. 1.Bessey, E. A. 1911. Root-knotand its control. U.S. Dept. Agric.BureauPlantIndustry,Bull. No.217.Bird,J., and J. Sinchez. 1971. Whitefly-transmittediruses in PuertoRico. J. Agric.Univ. PuertoRico 55:461-467.1 , R. L. Rodriquez,and F. J. Julia. 1975. Rugaceous(whitefly-transmitted)iruses inPuerto Rico. In J. Birdand K. Maramorosch, ds., TropicalDiseases of Legumes,pp. 3-25.AcademicPress, New York.CaliforniaCropandLivestockReportingService. 1980. Field cropreview. Vol. 1, Jan.28, 1980.CommonwealthMycological nstitute. 1968. Distributionmapsof plantdiseases. MapNo. 236, 3rdedit.ConjuntoTechnol6gicopara a Producci6nde Gandules.Su Situaci6nEcon6micay sus Perspectivas.1977. Estaci6nExp. Agric., Univ. PuertoRico. Publ. 116.Cook, R. J., J. W. Sitton,and J. T. Waldher. 1980. Evidence for Pythiumas a pathogenof direct-drilledwheatin the PacificNorthwest.P1.Dis. Reporter64: 102-103.Costa,A. S. 1975. Increase nthepopulationdensityof Bemisiatabaci, a threatof widespreadvirusinfectionof legumecrops in Brazil.In J. Birdand K. Maramorosch, ds., TropicalDiseasesof Legumes,pp. 27-49. AcademicPress, New York.Coyne, D. P., andM. L. Schuster. 1974. Breedingandgenetic studies of toleranceto severalbean(PhaseolusvulgarisL.) bacterialpathogens.Euphytica23: 651-656.

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    De, D. N. 1974. Pigeon pea. In J. Hutchinson,ed., EvolutionaryStudiesin WorldCrops, pp. 79-87. CambridgeUniv. Press.Ellis, M. A., S. R. Foor, and P. L. M6lendez. 1976. Effect of internallyseed-bornefungion ger-minationof pigeon pea in PuertoRico. (Abstr.)Mem. Soc. Puertorriquena ienciasAgric.2:8-9., and E. H. PaschalII. 1977. Methodsof controlling nternally eedborne ungiof pigeon pea(Cajanus cajan). Proc. Amer.Phytopathol.Soc. 4: 176., and . 1979. Effect of fungicideseed treatmenton internally eedbornefungi, germi-nationandfield emergenceof pigeon pea (Cajanuscajan). Seed Sci. & Technol. 7: 75-81.and E. Rosario. 1977. Similarities n the internally eedborne ungiof four eguminouscrops. Proc. Amer.Phytopathol.Soc. 4: 176.1 , E. J. Ravalo,and E. Rosario. 1978. Effect of growing ocationon internally-seed-borne fungi, seed germination,and field emergenceof pigeon pea in Puerto Rico. J. Agric.Univ. Puerto Rico 62: 355-362.Erwin,D. C. 1957.Fusariumand Verticilliumwilt diseases of Cicerarietinum. Abstr.) Phytopath-ology 47: 10.. 1958a. Fusarium lateritium f. ciceri, incitant of Fusarium wilt of Cicer arietinum. Phyto-pathology48:498-501.1958b. Verticilliumwilt of Cicerarietinumn southernCalifornia.P1.Dis. Reporter42: 1111.and W. C. Snyder. 1958. Yellowingof garbanzobeans. Calif.Agric. 12:6, 16.Evans, A. M. 1976. Beans. In N. W. Simmonds, ed., Evolutionof Crop Plants, pp. 168-172.Longman,London.Food and AgricultureOrganizationFAO)of the UnitedNations. 1979. ProductionYearbook.Vol.32. Rome.Freeman, G. F. 1912. Southwesternbeans and teparies.Univ. Arizona Agric. Exp. Sta. Bull. No.68: 573-619.1913. The tepary,a new cultivated egumefrom the Southwest. Bot. Gaz. 56: 395-417.Gamez,R. 1971. Los virus del frijol n Centroamerica.. Transmisi6npormoscas blancas(Bemisiatabaci Gen.) y plantashospedantesdel virus del mosaico dorado. Turrialba 1: 22-27.Godfrey,G. H. 1928. Legumesas rotationandtrap cropsfor pineapple ields.Hawaii Assoc. Pine-appleCannersExp. Sta. Bull. No. 10.Granillo,C., A. Diaz, M.Anaya,andL. A. Bermudezde Paz. 1975. Diseases transmitted y Bemisiatabaci in El Salvador.In J. Birdand K. Maramorosch, ds., TropicalDiseases of Legumes,pp. 51-53. AcademicPress, New York.Haddad,N. I., F. J. Muehlbauer, ndR. 0. Hampton. 1978. Inheritanceof resistance to pea seed-bornemosaic virus in lentils. CropSci. 18:613-615.

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    van der Maesen, L. J. G. 1972. Cicer L., a monograph f the genus, with special referenceto thechickpea (Cicer arietinum L.), its ecology and cultivation. Meded. Landbouwhogeschool,Wageningen72-10.Vovlas, C., and G. L. Rana. 1972. Le virosi delle plante ortensi in Puglia.VII. Lens esculentaMoench. ospite naturaledel virus del mosaico con enazioni de Pisello. Phytopathol.Medit.11: 97-102.Waines,J. G. 1978. Proteincontents, grainweights,andbreedingpotentialof wildanddomesticatedtepary beans. Crop Sci. 18: 587-589.WashingtonCropandLivestockReportingService. 1978. WashingtonAgricultural tatistics.Westerlund,F. V., Jr., R. N. Campbell,and K. A. Kimble. 1974. Fungal root rots and wilt ofchickpeain California.Phytopathology 4: 432-436.Williams,F. J., K. S. Amin, and B. Baldev. 1975. Phytophthorastem blightof Cajanus cajan.Phytopathology 5: 1029-1030.Wilson,V. E., andJ. Brandsberg.1965. Fungi solated from diseased lentil seedlings n 1963-64.P1.

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    Plant Classification.Lyman Benson. 2nd ed. 901 pp. illus. D. C. Heath and Company,Lexington,Massachusetts,1979.$22.95.The first edition of Plant Classification s outstanding.Benson has now come out withanupdatedandcompletelyrevised book thatbears littleresemblance o theoriginaleditionexcept for the splendidandprofuse black-and-whiteine drawingsmost of whicharenew.Dependingupon the previous training of the student, this volume could be used as acomprehensive text for a one-semester course or for a less concentratedtwo-semestercourse in planttaxonomyandclassification.As an authorityon Ranunlculus and Cactaceae in particular,Benson has numerousexamplesof the former n the section on keyingin which he uses examplesof keys from13 differentbooks coveringpractically all areas of the United States. He comparesthevarious ways by whichthe differentauthorskey a plant to a species of Ranunculus n allexcept one, Long and Lakela's Flora of Tropical Florida. Ranunculus does not occur inthe area, and a species of Citrus is the exampleused in this instance.I particularlyike Benson's discussionof the majorsystems of classification romTheo-phrastus o Cronquist.Coverageof families andorders of dicotyledonsand of morphologyandeconomic plantsof the groupsis especially good. Thereis a minimumof factualerrorin this magnificentvolume. My main regret is that the publisher'sproof readers let so

    manyprinter'serrorsthrough.This attractiveandsplendidlyboundandprintedbook shouldbe owned by allbotanistsregardlessof theirspecialization.EDWARD T. BROWNE, JR., MEMPHIS STATE UNIVERSITY, MEMPHIS, TN 38152