Morphology and Taxonomy of Species of Phomopsis on Asparagus Author(S): F

Mycological Society of America

Morphology and Taxonomy of Species of Phomopsis on Asparagus Author(s): F. A. Uecker and Dennis A. Johnson Source: Mycologia, Vol. 83, No. 2 (Mar. - Apr., 1991), pp. 192-199 Published by: Mycological Society of America Stable URL: http://www.jstor.org/stable/3759934 . Accessed: 05/03/2014 11:45

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This content downloaded from 134.121.161.15 on Wed, 5 Mar 2014 11:45:57 AM All use subject to JSTOR Terms and Conditions Mycologia, 83(2), 1991, pp. 192-199. ? 1991, by The New York Botanical Garden, Bronx,NY 10458-5126

MORPHOLOGY AND TAXONOMY OF SPECIES OF PHOMOPSIS ON ASPARAGUS

F. A. Uecker

SystematicBotany and MycologyLaboratory, USDA, AgriculturalResearch Service, BeltsvilleAgricultural Research Center,Beltsville, Maryland 20705

AND

Dennis A. Johnson

WashingtonState University,Irrigated Agriculture Research and Extension Center, Box 30, Prosser, Washington99350

ABSTRACT

Phomopsisjavanica, a new species on Asparagus fromJava, is described and illustrated.It differs fromP. asparagi by producingparaphyses among the conidiophoresand conidiogenous cells, and is the firstPhomopsis described with these structures.Phomopsis javanica was more virulentthan P. asparagi when inoculated on asparagus foliage and stems. Phomopsis asparagicola is considered a synonymof P. asparagi. P. asparagi and P. javanica are compared in a series of photographs. Key Words: Phomopsisjavanica sp. nov.,Phomopsis asparagi, Phomopsis asparagicola, paraphyses

A serious disease of asparagus (Asparagus of- Thailand (Anon., 1961). Reifschneider and Lopes ficinalis L.) plants in Java, Indonesia was found (1982) reported it in Brazil. Few of these papers to be caused by a Phomopsis. Comparison of this contain much information on the morphology fungus with the two other Phomopsis species pre? and taxonomy of the fungus and it is possible viously reported on asparagus revealed it to be that two or more fungi are involved. an undescribed species. Saccardo (1878) de? Bausa Alcalde (1952) described a second spe? scribed Phoma asparagi Sacc. on putrescent stems cies, Phomopsis asparagicola Bausa Alcalde on of A. officinalisat Padua, Italy. Although Bubak branches of Asparagus plumosus Baker. It dif? (1906) transferredthis fungus to Phomopsis, it is fered from P. asparagi because the alpha conidia still being reported as Phoma asparagi in current of P. asparagi were oblong, 7-8 x 3 /umaccord? literature (Liu and Hwang, 1988; Tanaka et al, ing to Saccardo (1878) and oblong or spindle- 1987). shaped and 5.5-9 x 2-2.5 /nm according to Phomopsis asparagi (Sacc.) Bubak is well Bubak (1906) whereas they were ellipsoid or known in Europe (da Camara and da Luz, 1943; fusiform, 7-9.5 x 2-2.5 in P. asparagicola; co? Engler and Prantl, 1900; Hruby, 1928; Lind, 1913; nidiomata were subepidermal in P. asparagi ac? Roumeguere, 1889; Solla 1915), in the Middle cording to both Saccardo and Bubak but sub? East (Engler and Prantl, 1900), Puerto Rico (Sherf epidermal then erumpent in P. asparagicola; and and MacNab, 1986) and eastern U.S. (Farr et al, the beta conidia of P. asparagi were reported to 1989) but has not been considered an important be shorter and thinner (18-25 x 1 jtm) than those pathogen in any of these places. of Phlyctaena asparagi Fautrey & Roum. (30 x The disease caused by P. asparagi on aspara? 2 jum), which was considered by Bausa Alcalde gus was first described in India by Kheswalla to be the beta form of P. asparagi. (1936) and later by Galloway (1936), Sohi et al A Phomopsis was recently collected on stems (1975), and Tripathi (1985). It attacks asparagus of asparagus in central Java in Indonesia. It dif? in China (Teng, 1932; Yao et al, 1987), Japan fers from the two previously mentioned fungi in (Tanaka et al, 1987), Korea (Nakata and Taki- that the alpha conidia are more variable in length moto, 1928; Choi et al, 1981), Taiwan (Hsu and and are consistently broader than those of the Sun, 1969; Wu, 1970; Yang et al, 1970), and other two, it produces paraphyses among the

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This content downloaded from 134.121.161.15 on Wed, 5 Mar 2014 11:45:57 AM All use subject to JSTOR Terms and Conditions Uecker and Johnson: Phomopsis on Asparagus 193 conidiogenous cells, and it is much more virulent illuminator equipped with 4 x and 11 x objec- on inoculated asparagus than is an isolate of P. tives. asparagi. Study ofthe conidiogenous apparatus was ac- This paper describes and illustrates the new complished by using a modification ofthe phlox- fungus on asparagus from Java and compares it ine-KOH method introduced by Martin (1934). with isotypes of Phomopsis asparagi and other A single conidioma was allowed to expand in 3% herbarium specimens and with three ATCC cul? KOH. A small portion ofthe conidiogenous layer tures from Taiwan. A lectotype of P. asparagi is was stained with 1% aqueous phloxine, the designated and photographs illustrating this spe? phloxine was replaced with KOH, and the spec? cies for the firsttime are provided. Epidemiology imen was flattened. All photographs ofthe conid? of P. javanica will be discussed in a separate iogenous apparatus were made from specimens paper. stained with phloxine-KOH, which was then re? placed with 0.2% cotton blue in 85% lactic acid MATERIALSAND METHODS (LA-CB) to extend the useful life of the prepa? ration. The original collection of Phomopsis javanica Conidia were collected on cover slips by touch- was made by one of us (DAJ) in central Java in ing the cover slip to the mucilaginous drop con? January, 1989. A culture (FAU-477) from this taining conidia at the mouth of the ostiole. A collection was grown routinely on autoclaved one- small drop of water was added and the conidia inch long pieces of stem of Asparagus officinalis, were spread in an even layer across the cover alfalfa (Medicago sativa L.), soybean [Glycine max slip. In cases where conidia were not yet extruded (L.) Merr.], Stokes aster [Stokesia laevis (Hill) in a drop, conidiomata were dissected in a small Greene], kiwi (Actinidia chinensis Planchon), and drop of water on a slide and the conidia were cranberry (Vaccinium macrocarpon Ait), and on spread upon the slide. In either case, after air- grains of oat (Avena sativa L.) on water agar (WA) drying the conidia were mounted in a small drop plates. Specimens on each host material were of LA-CB. fixed in formalin-aceto-alcohol (FAA), embed? Phomopsis javanica was also grown on pieces ded in Paraplast Plus1 and sectioned at 7 jum. of stem of asparagus, alfalfa, and grape (Vitis Slides were run through a standard series of xy? labrusca L.) on WA plates and the conidiogenous lene and ethanol to water and were stained in apparatus was observed at intervals of one to methylene blue-azure II (Humphrey and Pitt- five days. Photographs were first taken on day man, 1974, omitting the basic fuchsin). Although five after inoculation, the last ones on day 40. this stain was originally applied to specimens The purpose of this series was to determine embedded in plastic, it is also useful for paraffin whether the conidiogenous apparatus in conid? sections. Used at full strength, it overstains iomata on asparagus varied morphologically at heavily. When diluted to 25% of the original any stage of development from those at other strengthand applied for 5-15 min at room tem? stages and from those on the other substrata. perature staining was more easily controlled. To compare the virulence of P. javanica with Slides were then dipped in water to remove ex- that of P. asparagi, potted asparagus plants of cess stain, blotted nearly dry on absorbent paper, cultivars Mary Washington and WSU-1 were in? and dehydrated and differentiated one to two h oculated with P. javanica and P. asparagi in three in tertiary butanol. After two 10-min baths ih separate tests. The isolate of P. asparagi (Fau- xylene, sections were mounted in Permount. 499) was derived from naturally infected aspar? Isotype specimens of Phomopsis asparagi were agus plants grown at Rutgers Research and rehydrated for several h on moist filter paper, Development Center, Bridgeton, New Jersey. In? fixed in FAA, and furthertreated as above. Co? oculum was increased on potato dextrose agar in nidiomata were measured on living or dried Petri dishes placed under continuous fluorescent specimens using a Leitz Ultropak incident light light at 20 to 23 C for 11-14 days. Conidiomata were scraped and washed from the agar, conidia 1 of a or Mention trademark,proprietary product, were filteredthrough cheesecloth and inoculated vendor does not constitutea guaranteeor warrantyof onto four to seven plants per test. One drop of theproduct by theU.S. Departmentof Agricultureand ml does not imply its approval to the exclusion of other Tween 20 was added to 500 distilled water productsthat may also be suitable. and one drop of the dilution was added to the

This content downloaded from 134.121.161.15 on Wed, 5 Mar 2014 11:45:57 AM All use subject to JSTOR Terms and Conditions 194 Mycologia inoculum. Concentration of inoculum was ap? aline, short or elongate, more or less tapered to? proximately 1.8 x 106, 2.8 x 106, and 1.5 x 106 ward apex, septate both at the base and above, alpha conidia, respectively, for the three tests. 10-40 x 2-3 Mm but up to 6 jum wide at the Inoculations were accomplished by wetting the base; except for the terminal cell each cell ofthe foliage of asparagus plants with inoculum. Shoots conidiophore produces a short or long lateral 60-80 cm in length were laid horizontally and branch just below the septum and becomes con? moved around on a 60 x 68 cm plastic sheet idiogenous; conidiogenous cells 10-20 x 2-4 fim, that contained 60-100 ml of conidial suspension. phialidic, integrated or discrete, hyaline, aper? Separate plastic sheets were used for each fungus. ture apical on the lateral branch, channel and After inoculation, plants were placed in a plastic collarette minute, periclinal thickenings of vari? mist chamber for 72, 48, and 64 h, respectively, able thickness; conidia acropleurogenous; alpha for the three tests. Plants were then placed in a conidia (Fig. 5) hyaline, aseptate, usually bigut- greenhouse with temperatures of 21-24 C during tulate but sometimes with one large guttule, the day and 17 to 19 C at night. broadly elliptic or elliptic or fusiform-elliptic,6- 12 x 3-4 Atm;beta conidia (Fig. 6) hyaline, asep? results tate, not guttulate, straight or curved or sigmoid, 12-22 x 0.9-1.3 ixm; sometimes a continuum of Phomopsis javanica Uecker et D. A. Johnson, sizes exists between alpha and beta conidia; pa? sp. nov. raphyses (Fig. 7) hyaline, septate or aseptate, Conidiomata brunnea,simpliciter eustromatica, im- arising from the terminal cell ofthe conidiophore dissita raro mersa,plerumque confluentia,ampullifor- or from the layer that gives rise to conidiophores, mia vel complanata, loculo solitari interdumconvo- up to 60 x 2-4 ^m, freeat tips, freeends rounded luto, 160-430 fimlongo x 85-315 alto; paries fuscus or in apicem versus, ad latera et infimumjuventute palli- expanded, some conidiomata abundant but dior, postea fuscansubique, 10-14 ixmcrassus vel tot in others sparse. quot 25 fimprope ostiolum, textura angularis; ostiolum circulare, plerumque solitarium, 15-20 fim; conidi? HOLOTYPE: BPI, USO 1102577, on sterilizedstems ophora hyalina,brevia vel elongata,plus minusve de- ofAsparagus officinalison wateragar. Isotypesin NY, crescentia,et basi et super septata,praeter terminalem DAOM, and IMI (abbreviationsfrom Holmgren et al., ramo lateralelongo vel breve infraseptum omnes cel? 1981). lulae conidiophoriconidiogenentes, 10-40 x 2-3 j^m Other specimens examined: BPI: USO356160, la- sed usque 6 /tmad basim; cellulae conidiogenae en- beled Phoma asparagi Sacc. on Asparagus officinalis teroblasticae,phialidicae, integrataevel discretae,hy? L., Taipeh, Taiwan, K. Sawada 15/XI/1922. alinae, aperturain ramo lateraliapicali, canale et col- Culture: ATCC 24624, on Asparagus, Taiwan, S. lulo minutis,spissitudine periclinali crassa vel non, 10- K. SunPA-12. 20 x 2-4 fim;conidia acropleurogena;conidia alpha hyalina,aseptata plerumque biguttulata,late elliptica Symptoms consisted of elliptical to round le? vel vel 6-13 x 3-4 elliptica fusiformia-elliptica, j^m; sions (less than 1 mm to over 45 mm in length) conidia beta hyalina,aseptata, non guttulata,recta vel with either tan to light brown or white centers curvatavel sigmoidea, 12-22 x 0.9-1.3 fim,interdum continuo interconidia alpha et beta; paraphyses hy? and brown to reddish brown margins. Coalescing alinae, septataevel aseptatae,e cellula terminaleconid? and large lesions girdled branches and stems iophori vel e cellulas iisdem atque conidiophoris,us? causing the fern and shoots to blight. Conidio? x que 60 2-4 fim,in conidiomatibusaliquot abundantes mata formed in lesions and in dead tissue. In sed in alterissparsae. January 1989, 20 to 80% ofthe shoot fern in the Conidiomata (Figs. 1, 2) brown or smoky or asparagus-growing area in central Java (23 ha in more heavily pigmented, simply eustromatic, five locations) had disease symptoms on 5 to immersed, usually separate but sometimes con? 100% ofthe surface area ofthe foliage. fluent,ampulliform or flattened,circular or elon? Lesions on inoculated plants became initially gate in outline, papillate or not, with a single evident five to six days after inoculation with P. locule that is often convoluted, (160-)250-350(- javanica and eight to ten days after inoculation 430) fim long x 85-315 fim high; wall (Fig. 2) with P. asparagi. Mean number of lesions per dark around the ostiole, lighterat sides and below inoculated plant were 23, 30 and 18 after inoc? when young but becoming darker with age, 10- ulation with P. javanica and 2, 1, and 0.5 after 14 fim thick, to 25 fim thick near the ostiole, inoculation with P. asparagi, respectively, forthe textura angularis; ostiole usually single, circular, three tests. Differences were statistically signifi? 15-20 fim diam; conidiophores (Figs. 3, 4) hy- cant (P = 0.01) using Student's Mest.

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Figs. 1-7. Phomopsisjavanica. 1. Habit on Asparagusstem, x 45. 2. Sectionthrough conidioma on Asparagus stem, x 325. 3. Portionof conidiogenouslayer showing conidiophores with septa and withconidiogenous branch emergingfrom below septa, x 1000. 4. Conidiophores withconidiogenous cells producingbeta conidia, x 1000. 5. Alpha conidia, x 1000. 6. Alpha, beta, and intermediateconidia, x 1000. 7. Conidiophores,conidiogenous cells, and paraphyses, x 1000.

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Phomopsis javanica produced conidiomata into a moist chamber and beta conidia were ob? abundantly on pieces of stem of asparagus, al? served after two more weeks. falfa, soybean, and stokes aster on WA. Produc? Phomopsis asparagi (Sacc.) Bubak, Bull. Herb. tion of conidiomata was sparse on kiwi and cran- Boissier Ser. 2, 6: 408. 1906. berry stems and on oat grains. Sizes of conidiomata were similar on all these substrates. = Phoma asparagi Sacc, Michelia 1: 257. 1878. = The conidiomata continued to enlarge for about ^Phomopsis asparagicola Bausa Alcalde, Anales Inst. Bot. Cavanilles 237. 1952. two wk after inception and continued conidium 10(2): production for about six wk. Conidiomata de? Saccardo (1878) described Phoma asparagi veloped in a subepidermal position but the os? with conidiomata gregarious, covered by the epi? tiole apex soon became erumpent through the dermis, globose-depressed, conidia oblong, 7-8 epidermis. Young conidiomata were yellowish x 3 fim, two-guttulate, hyaline, beta conidia not with a dark area surrounding the ostiole. Conid? seen. Based on a specimen from Yugoslavia rath? iomata became darker in color as they aged, the er than on type material, Bubak (1906) trans? color spreading downward from the area near ferred the name to Phomopsis as Phomopsis as? the ostiole. Conidiomata were sometimes nearly paragi (Sacc.) Bubak. He redescribed it as spherical but more commonly somewhat flat? extremely variable, with or without stromata, tened. They were ostiolate and more or less pa? stromata elongate or in a streak, subepidermal, pillate with great variation in the length of the 0.5-1 mm long and 300-400 ixmwide, rupturing papilla. The wall was thinner on the bottom and by a longitudinal split, black, dull, light brown sides, thickest near the ostiole (Fig. 2). within and black-brown above, pycnidia or sin? Sporulation was profuse on asparagus, alfalfa, gle complete or incomplete chambers mostly lens- and soybean stems. In the few conidiomata shaped and depressed and elongated, rarely formed on kiwi and cranberry stems and on oat hemispheric; conidiophores rod-shaped, 10-15 grains conidium production was much less pro? fim long, 1-1.5 fim wide, straight, hyaline; co? fuse. nidia elongate or spindle-shaped, 5.5-9 x 2-2.5 A more or less tightly packed, palisade-like fim or often with Septoria-shaped conidia, elon? layer of conidiophores and paraphyses lined the gate conidia rounded at the ends, spindle-shaped entire cavity of the conidiomata up to the point ones attenuated, appearing falsely two-celled be? where the ostiole begins. Each conidiophore was cause of two large oil drops; with a range of shapes attached at the base to a cell on the inside of the between the spindle-shaped and Septoria-shaped wall. No layer of morphologically specialized cells ones. was evident between the conidiophores and wall. Conidiomata of both the isotypes (Figs. 8, 9) As soon as conidiomata could be distinguished and of Bubak's specimens are black, stromatic, on stem pieces on WA, 4-5 days after inocula? ampulliform or somewhat flattened, elliptic or tion, conidiogenous apparatus and conidia were broadly elliptic to subcircular in outline, im- already present. mersed, separate or confluent, 190-390(-660) fim were firstobserved in conidiomata Paraphyses long x 115-240 fim wide, larger conidiomata on stems on WA about after inocu? eight days often found in one area on a stem and smaller lation. Paraphyses were longer and more nu? ones in an adjoining area on the same stem or merous in conidiomata 13 or more days after sometimes mixed; wall of the conidioma is tex? inoculation. After about six wk neither paraph? tura angularis (Fig. 9); conidiophores hyaline, yses nor conidiogenous apparatus were function- septate, tapering toward the apex, to 40 x 5 pm; al any longer. Paraphyses were often difficultto cells find in sections but were evident in nearly every conidiogenous (Fig. 10) phialidic, integrated or or squash mount. discrete, hyaline, cylindric tapered, to 20 x 4 Alpha conidia were variable in length from one pm, aperture apical on terminal cell or crop to the next on the same host (compare Figs. on lateral extensions of other conidiogenous cells, 5 and 6). Beta conidia (Figs. 4, 6) have so far periclinal thickenings usually visible; alpha co? been seen on only one occasion, afterinoculation nidia 6-8 fim in Saccardo's specimens (Fig. 11), ofa living asparagus plant. Branches excised from 6-9 in Bubak's; beta conidia were not seen in that plant after incubation for 15 days were put any of these specimens.

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Figs. 8-11. Phomopsisasparagi. 8. Habit on Asparagusstem, x 43.9. Section throughrehydrated conidioma of isolectotype,x295. 10. Short conidiophoreswith conidiogenouscells, x 1000. 11. Alpha conidia, x 1000.

The following specimen is here designated as 2940, III/1893, Newfield,New Jersey;labeled Phoma lectotype of Phoma asparagi Sacc.: media Ellis & Everhart:US0357121, on Asparagus sp., C. E. Fairman, no date, Lyndonville, New York; LECTOTYPE: Phoma (Diaphorte) [sic] asparagi US0357135 on Asparagus officinalisL., W. J. Young Sacc.?on Asparagus stem, X/1876 Padua, Italy. My- 14/IX/1923,Lancaster, Ohio; labeled Phoma micros? cotheca Veneta No. 932, Sbarbaro Collection (undis- pora Berk. & Curtis: US0357173, on Asparagus sp., tributed),in BPI. H. W. Ravenel FungiAmericani Exsiccati 538, no date, Otherspecimens examined:?BPI: labeled Phomopsis Aiken, South Carolina. asparagi (Sacc.) Bubak: US0358286 on Asparagus ver- IMI: 215726, on Asparagus,E. Niemann 5/IX/1975, ticillatus,F. Bubak 10/IV/1903,Fungi Montenegrini, Bako, Ethiopia; 190975, on Asparagus,H. S. Sohi 28/ Rijeka, Montenegro,Yugoslavia; US0358285 on A. 1/1975,Bangalore, India; 185760, on Asparagus sp., S. officinalis,EHettrich-Kalkoffl/1915, Arco, southTyrol, Ahmad 22/11/1965,Changa Manga, Pakistan; 22601, Italy; labeled Phoma asparagi Sacc: US0356156, on on Asparagus (inoculatedstems), Imperial mycologist, Asparagus, Saccardo Mycotheca Veneta No. 932 (Is- IMI, 31/X/1938, Pusa, India; 22600, on Asparagus olectotype);US0356161, on A. officinalis,G. W. Carv? (naturallyinfected), Imperial mycologist,IMI, 31/X/ er 378, 14/IV/1936,Tuskegee, Alabama; US0356159, 1935, Pusa, India; 292222, isol. ex Asparagus officina? on A. officinalis,K. Sawada 15/XI/1912,Taipei, Tai? lis, Chong P.P. 2841/60, 13/XII/1984,Tuaran, Malay- wan; US0356164, on A. officinalisJ. T. Rogers 27/11/ sia; 292893, isol. ex Asparagus officinalis,Bong. P.P. 1919, Washington,D.C; US0356158, on A. officinalis, 2866/60, 3/1/1985,Kundasang, Malaysia. Henry Banks 31/X/1927, Marion, Arkansas; MICH: Ellis & Everhart2940 (two specimens) and US0356157, S. C. Teng 770, 16/VIII/1931,Nanking, 2156, cited above. China; US0356163, on A. officinalis,Ellis & Everhart CUP: 35064, Phoma asparagi Sacc. on Asparagus NorthAmerican Fungi Second Series 2156, XII/1888, sp., L. Ogilvie 3/111/1926,Pomander Gate, Paget,Ber- Newfield,New Jersey;US0356155, on Asparagus sp., muda; CUP-A: Ellis & Everhart2940 and 2156, cited Ellis & EverhartNorth American Fungi Second Series above; CUP-F: Mycotheca Fairmani 3967, labeled

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Phoma media Ellis & Everharton Asparagus sp., R. and we believe that they should also be men- Latham New labeled Phoma 14/11/1915,Orient, York; tioned when they occur in Phomopsis. Such a Berk.& Curt.:H. W. Ravenel Amer- microspora Fungi distinctive character is welcome in the of icani Exsiccati 538, Aiken, South Carolina. study NY: labeled Phoma asparagi Berk. & Curt., on As? a group noted for a dearth of such characters. paragus sp., ex Herb. E. M., no date, ChesterCounty It was noted above that paraphyses in P. ja? (PA?); labeled Phoma asparagi Sacc: de Thuemen, vanica may arise either from the cells that pro? universalis 1585 on A. H. W. Mycotheca officinalis, duce conidiophores or from various cells of the Ravenel, 1877, Aiken,South Carolina; Ellis & Everhart the terminal cell. 2940 and 2156 (two specimens),cited above; ex Herb. conidiophore, especially Except A. Commons 1064, on decayingstems oiPhytolacca for one paraphysis shown developing from a low? decandra L., 5/XI/1889,Wilmington, Delaware; on A er cell of the conidiophore in Aschersonia aley- G. W. Carver officinalis, 418, 12/VIII/1897,Tuskegee, rodis, all those illustrated by Sutton (1980) de? Alabama; labeled Phoma asparagi f. tami: C. Rou- velop from the cell layers that give rise to the meguere,Fungi SelectiExsiccati 5764, on Tamus com? munis,Eug. Niel., no date,Ferrierres pres Broglie (Eure). conidiophores or conidiogenous cells. Especially MA: 12046, labeled Phoma asparagi Sacc. on As? in the case of those arising from the terminal cell paragus albus L., D. G. Sampaio 2/IV/1921, Faro, of a conidiophore, one might question why these 7080 labeled Phoma Sacc. on As? Portugal; asparagi structuresare not considered simply as elongated paragussp., P. D. Unamuno IX/1924,Santander, Spain; or cells that have 10779, labeled Phomopsis asparagi (Sacc.) Trav. & conidiophores conidiogenous Spessa on Asparagusplumosus,Silva TeixieraXII/1932, not yet become conidiogenous. It is possible and Horto Stellae Olissiponis, Portugal. even probable that some of them do become Cultures: ATCC S. K. Sun PA-7 on 24623, aspar? conidiogenous. They are sometimes septate, and agus, Taiwan; ATCC 24625, S. K. Sun PA-23 on As? have rounded ends in contrast to the acute apices paragus, Taiwan. of the conidiogenous cells. The paraphyses are We were unable to obtain the type of Phomop? present early in development, become more con? sis asparagicola. Types of four ofthe species which spicuous for a time, and some are still present Bausa Alcalde described in the same publication fortydays after inoculation. with P. asparagicola are available from MA but Three species of Phomopsis have been de? the types of Phomopsis asparagicola and P. ca- scribed on asparagus stems. We believe that P. talpicola were not found (F. Pando, Curator of asparagi and P. javanica are distinct and we con- Cryptogamic Herbaria, Jardin Botanico, Ma- sider P. asparagicola a synonym of P. asparagi. drid, personal communication). Comparison of Comparing the isotypes of P. asparagi and Bu- the type, if found, with that of P. asparagi will bak's specimens with the descriptions of P. as? probably show that both are the same species. paragicola indicates that there is no outstanding character to separate the two and that the conid? discussion ial shape and dimensions offeredfor P. asparagi? cola are not unlike those for isotypes of P. as? Sutton (1980) defined paraphyses as sterile hy? paragi. If the type of P. asparagicola or other phae which are free at the apex and are often authentic specimens are discovered, the question produced among fertile conidiophores or conid? can be reevaluated. iogenous cells. Among Phialostromatineae, the suborder which includes Phomopsis, Sutton de? ACKNOWLEDGMENTS scribed and illustrated paraphyses in Ascherso- nia, Plectophomella, Titaeospora, Amerospor- We thankDonald P. Rogers forcorrecting the Latin ium, Phaeocytostroma, and Massariothea. description.Stephen A. Johnsoncollected for us the fromwhich isolate FAU-499 was derived. S. C. Among Phialopycnidiineae, only Coleophoma plants Jongprovided the ATCC isolatesthrough the exchange and Pseudorobillarda exhibit paraphyses. The process. We thank the curatorsof the followingher? only previous indication that such structures baria forsending specimens: B, CUP, IMI, K, L, MA, might occur in Phomopsis is an illustration of a MICH, NY, UC, and WSP. single paraphysis in P. theae Petch (Punithalin- gam and Gibson, 1972). No mention is made of LITERATURECITED this structure either in the text or legend for the 1961. for to We consider to be Anonymous. QuarterlyReport January figure. paraphyses unreported March, 1961 ofthe Plant ProtectionCommittee previously in Phomopsis. Paraphyses have been for South East Asia and Pacific Region. F.A.O. considered worthy of description in other genera Publ., Bangkok,Thailand. 18 p.

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Bausa Alcalde, M. 1952(1951). Algunos micromi- Nakata, K., and S. Takimoto. 1928. Diseases of cul? cetos recolectados por el Prof. Caballero Segares tivated plants in Korea. /. Agric. Exp. Sta. Gov. en Valencia. Anales Inst. Bot. Cavanilles 10(2): Gen. ChosenNo. 15: 1-146. 229-255. Punithalingam,E., and I. A. S. Gibson. 1972. Pho? Bubak, F. 1906. Zweiter Beitrag zur Pilzflora von mopsis theae. Commow. Mycol. Inst. Descript. Montenegro.Bull. Herb. Boissier Ser. 2, 6: 393- Pathog. Fungi Bact. No. 330. 408. Reifschneider,F. J. B., and C. A. Lopes. 1982. Phoma Camara, E. da Sousa da, and C. G. da Luz. 1943. asparagi on Asparagus. FAO Pl. Protect.Bull 30: Mycetes Aliquot Lusitaniae. VI. Agron. Lusit. 5: 157. 119-146. Roumeguere,C. 1889. Fungi selecti exsiccati. Cen- Choi, J. K., Y. S. Kwon,and Y. H. Yu. 1981. Studies turieL. Rev. Mycol. (Toulouse) 11: 127-136. on the controlof stem blightof Asparagus caused Saccardo, P. A. 1878. Fungi Veneti novi vel critici by Phoma asparagi Sacc. Korean J. Pl. Protect.20: vel Mycologiae Venetae addendi. Series VIII. 83-86. Michelia 1:239-275. Engler, A., and K. Prantl. 1900. Die natiirlichen Sherf, A. F., and A. A. MacNab. 1986. Vegetable Pjlanzenfamiliennebst ihren Gattungen und wich- diseases and theircontrol. 2nd Ed. John Wiley & tigerenArten. Teil 1, Abt. 1, Leipzig, Verlag von Sons, New York. 728 p. Wilhelm Engelmann. 570 p. Sohi, H. S., B. A. Ullasa, and S. S. Sokhi. 1975. Stem Farr, D. F., G. F. Bills, G. P. Chamuris, and A. Y. blightof Asparagus officinaliscaused by Phomop? Rossman. 1989. Fungi on plants and plant prod? sis asparagi (Sacc.) Bubak. Curr.Sci. 44: 722-723. ucts in the United States. St. Paul, APS Press. Solla, R. 1915. In Italian in den Jahren1912-13 auf- 1252 p. getreteneschadlische Pilze. Z. Pflanzenkrankh.25: Galloway, L. D. 1936. Report ofthe Imperial my? 344-351. cologist. Sci. Rep. Agric. Res. Inst., New Delhi, Sutton,B. C. 1980. The Coelomycetes.Fungi imper- 1935-1936: 105-111. fecti withpycnidia, acervuli, and stromata. Com? Holmgren, P. K., W. Keuken, and E. K. Sehoneld. monw. Mycol. Inst.,Kew, Surrey,England. 696 p. 1981. Index herbariorum.I. The herbaria ofthe Tanaka, K., S. Tsuchida, and F. Nonaka. 1987. Stud? world.7th ed. Regnum Veg. 106: 1-452. ies on stem blightof Asparagus caused by Phoma Hruby,J. 1928. Die Pilze Mahrens und Schlesiens. asparagi Saccardo in Saga Prefecture.Proc. Assoc. Ein Versuch der Gliederungder Pilzdecke dieser Pl. Protect.Kyushu 33: 66-70. Lander. Hedwigia 68: 119-190. Teng, S. C. 1932. Fungi of Nanking II. Contr.Biol. Hsu, C-h., and S-k. Sun. 1969. Stem blightof As? Lab. Chin. Assoc. Advancem.Sci., Sect. Bot. 8: 5- paragus in Taiwan. I. Distributionof the disease 48. and culturalcharacters and spore germinationof Tripathi,K. C. 1985. Diseases of Asparagus in Gar- the causal organism, Phoma asparagi Sacc. Pl. hwal Hills. Indian J. Forestry8: 236. Protect.Bull. Taiwan 11: 47-60. Wu, F. S. 1970. Etiological surveyof Asparagus dis? Humphrey,C. D., and F. E. Pittman. 1974. A simple eases in Taiwan. /. Taiwan Agric.Res. 19(4): 71- methyleneblue-azure Il-basic fuschsinstain for 81. epoxy-embedded tissue sections. Stain Technol. Yang, I. L., S. M. Hour, and Y. L. Chen. 1970. Stud? 49: 9-14. ies on the stem blight of Asparagus caused by Kheswalla, K. F. 1936. A Phoma disease of Aspar? Phoma asparagi. II. Physiologicalcharacteristics agus. Indian J. Agric.Sci. 6: 800-802. and chemical control.J. Taiwan Agric.Res. 19(4): Lind, J. 1913. Danish fungi as representedin the 60-67. herbariumof E. Rostrup. Copenhagen, Gylden- Yao, W. Y., Y. Y. Zhu, S. Q. Zhang, D. W. Xu, and dalske Boghandel-NordiskForlag. 648 p. R.Y.Yang. 1987. Preliminarystudy on the stem Liu, T. M. E., and J. S. Hwang. 1988. Survival and blightin Asparagus. Shanghai Agric.Sci. & Tech- overwinteringof Phoma asparagi. Pl. Protect.Bull. nol No. 2: 37-38. Taiwan 30: 24-30. Martin, G. W. 1934. Three new heterobasidiomy- Accepted forpublication December 3, 1990 cetes. Mycologia 26: 261-265.

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