Enfermedades de Cereales y Pastos - Disease of Cereals and Grasses - Roderick Sprague PH.D 1950

DISEASES OF
CEREALS AND GRASSES

IN NORTH AMERICA
( F u n g i , E x c e p t S m u t s and R u s t s )
By
RODERICK SPRAGUE, Ph.D.
PATHOLOGIST, AGRICULTURAL E X P E R I M E N T STATION
T H E STATE 'COLLEGE OF •WASHINGTON
THE RONALD PRESS COMPANY ~9- NEW YORK

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T H E RONALD PRESS 'COMPANY
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The text of this publication or any part

thereof may not be reproduced in any
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P R I N T E D I N T H E U N I T E D STATES OF AMERICA
PREFACE
This book has been written to supply a source of centrally assem-

bled information concerning the fungi occurring on members of the
grass family and to serve as a contribution to a field in which there
is still a great need for fundamental study. It is an outgrowth of my
own special interest in the diseases of small grains and of the need
for an up-to-date and adequate source of information on the mor-
phology of those fungi causing leaf spot, blights, ergots, scalds, and
ro.ot rots in members of the grass family. In recent years, particularly
since the publication of Hitchcock's manual on the grasses (1935),
our knowledge of the diseases of range and pasture grasses has ma-
terially increased. The field, once covered to some extent by Stevens'
definitive text (1913), is now almost too large for any one person to
summarize. Descriptions of many fungi causing such diseases are, of
course, to be found scattered through the massive twenty-five vol-
umes of Saccardo's Sylloge Fungorum (1882-1931). While this work
is indispensable to the taxonomic mycologists, it is somewhat less
adapted to the needs of plant pathologists working in North America.
And so much has been written since the last volume of Sylloge
Fungorum was published that its value is steadily declining.
I am indebted to a number of my colleagues of earlier and current
years for aid in assembling material and literature related to this
book. John A. Stevenson, William Diehl, Edith Cash, C. L. Lefebvre,
A. G. Johnson, F. J). Bailey, William Bridge Cooke, George W.
Fischer, Jack P. Meiners, Charles Gardner Shaw, C. R. Orton, H. C.
Greene, John Hardison, George A. Rogler, and W. G. Solheim have
all given me necessary aid to permit completion of the work. My
family, especially Mary Willis Sprague, has aided me from the begin-
ning in the assembling of the manuscript. These pages are dedicated
to Aaron G. Johnson, my colleague for two decades..
The courtesy of Oregon State College through Editor D. M. Goode,
in permitting use of a number of figures from Oregon State College
Botanical Memoirs No. 6 is greatly appreciated, as are like courtesies
from the editors of Thytopathology, Mycologia, and the U.S.D.A.
Bureau of Plant Industry. RODERICK SPRAGUE
Wenatchee, Washington
June, 1950
CONTENTS
PAGE
INTRODUCTION 3
THE FUNGI
PHYCOMYCETES . . . .
Key to the Phycomycetes 5
Aphanomyces camptostylus Drechsl. 5
Cladochytrium graminis Buesgen 7
Hyphochytrium catenoides Karling 8
• Lagena radicioola Vanterpool and Ledingham 9
Olpidium brassicae (Wor.) Dang. . 10
Physoderma maydis Miyabe . 11
Phytophthora cactorum (Leb. and Cohn) Schroet. 13
Phytophthora parasitica Dastur 14
Polymyxa graminis Ledingham 15
Pythium root rot and seed rot diseases 16
Keys to species of Pythium isolated from Gramineae in North America 18
Pythium aphanidermatum (Edson) Fitzpatrick 20
Pythium artotrogus (Mont.) deBy. var. macracanthum Sideris 21
Pythium butleri Subrm. . 22
Pythium debaryanum Hesse 23
Pythium dissotocum Drechsl. 27
- Pythium graminicola Subrm. . 28
Pythium hypogynum Middleton 33
Pythium irregulars Buisman 34
Pythium rhonospermum Pringsh. 35
Pythium nagaii S. Ito and Tokunaga 36
Pythium oligandrum Drechsler 36
Pythium ostracodes Drechsl. . 37
Pythium periilum Drechsl. 39
Pythium rostratum Butl. 39
Pythium splendens Braun 40
Pythium tardierescens Vanterpool . 41
Pythium ultimum Trow 42
' Pythium volutum Vanterpool and Truscott 43
Pythium spp., mostly saprophytes . 44
Rhizophidium graminis Ledingham 46
47
Rhizopus spp. . . . . .
47
Sclerospora farlowii Griffiths , . 48
Sclerospora graminicola (Saco:) Schroet. 50
Sclerospora macrospora Sacc. . 51
Synchytrium graminicola sp. nov. .
ASCOMYCETES . . . . 53
Key to the Ascomycetes . 53
Balansia claviceps Speg. . 54
Balansia clavula (Berk, and Curt.) Lloyd 55
vi CONTENTS
ASCOMYCETES (Cont.) PAGE
Balansia hypoxylon (Pk.) Atk. 66
Balansia pilulaeformis (Berk, and Curt.) Diehl 58
Balansia trinitensis Cooke and Massee . 59
Ceratostomella paradoxa, see Endoconidiophora paradoxa 59
Claviceps cinerea D. Griff. . . ' 59
Claviceps paspali Stev. and Hall . . .i 60
Claviceps purpurea (Fr.) Tul. . . . . 61
Claviceps ranunculoides A. Moll 68
Claviceps rolfsii Stevens and Hall . . . . 66
Claviceps tripsaci Stevens and Hall 67
Claviceps yanagawaensis Togashi . . . . 67
Cochliobolus heterostrophus Drechsl. 6S
Cochliobolus miyabeanus, see Helminthosporium oryzae 69.
Cochliobolus sativus, see Helminthosporium sativum 69
Cochliobolus setariae, see Helminthosporium setariae 70
Cryptoascus sp. . . . . . 70
Dothichloe Atkinson . . . . 70
Dothichloe aristidae Atk. 71
Dothichloe limitata Diehl 71
Dothichloe nigricans (Speg.) Chardon 72
Dothidella aristidae (Schw.) Ell. and Ev., 73
Dothidella minima Sacc. and Sydow 73
Endoconidiophora adiposa (Sartoris) Davidson 74
Endoconidiophora paradoxa (De Seynes) Davidson 75
Endodothella traoyi (Ell. and Ev.) Theiss. and Syd, 76
Epichloe typhina (Fr.) Tul 77
Erysiphe graminis DC. . . . . . . 78
Gibberella fujikuroi and G. moniliformis, see Fusarium moniliforme 82
Gibberella zeae, G. pulicaris, and G. saubinetii, see Fusarium graminearum 82
Gnomonia iliau Lyon . . . . 82
Leptosphaeria avenaria, see Septoria avenae 83
L. avenaria f. sp'. triticea, see Septoria-avenae f. sp. triticea 83
83
Leptosphaeria herpotrichoides de Not.
Leptosphaeria sacchari Van Breda de Haan
Leptosphaeria salvinii Catt.
1 84
84
83
Lophodermium arundinaceum (Schrad. ex. Fr.) Chev.
87
Mycosphaerella calamagrostidis Greene .
8S
Mycosphaerella holoi Tehon 88
Mycosphaerella tulasnei (Jancz.) Rothers 93
Myriogenospora aciculisporae Vizioli 91
Myriogenospora paspali Atk. . 92
Ophiobolus graminis Sacc. 95
Ophiobolus oryzinus Sacc. 95
Ophiobolus sativus, see Helminthosporium sativum 93
Phialea temulenta Prill, find Del. . 100
Phyllachora spp. (key to species) . . . / 103
Phyllachora ammophilae Orton . . . ' 104
Phyllachora arundinariae Orton 104
Phyllachora boutelouae Rehm 104
Phyllachora chardonii Orton . . . . 105
Phyllachora coloradensis Orton . '. . - 105
Phyllachora congruens Rehm . . . -."'"! 105
Phyllachora cornispora Atk 106
Phyllachora diplocarpa Ell. and Ev. 106
Phyllachora epicampis Orton . . . . 107
Phyllachora eragrostidis Chardon •^ •
CONTENTS
PAGE
Phyllachora erianthii Orton 107
Phyllachora graminis (Pers.) Fckl, 107
Phyllachora guianensis Stev. . 108
Phyllachora heterospora P. Henn. 109
Phyllachora insularis Chardon 109
Phyllachora lasiacis Syd. 109
Phyllachora leptochloae Chardon 110
Phyllachora luteo-maculata (Schw.) Orton 110
Phyllachora maydis Maubl. 110
Phyllachora nervisequia (Schw.) Orton . 111
Phyllachora oryzopsidis Theiss. and Syd. 111
Phyllachora pammelii Orton . 111
Phyllachora parilis Syd. . . . . 112
Phyllachora paspalicola P. Henn. . 112
Phyllachora phalaridis Orton . 112
Phyllachora punctum (Schw.) Orton 113
Phyllachora quadraspora Tehon 113
Phyllachora serialis Ell. and Ev. 114
"Phyllachora silvatica Sacc. and Speg. 114
Phyllachora spartinae Orton 115
Phyllachora sphaerosperma Wint. . 115
Phyllachora texensis Orton 115
Phyllachora vulgata Theissen and Sydow 116
Phyllachora wilsonii Orton 116
Physalospora rhodina (Berk, and Curt.) Cooke 117
Physalospora tucumanensis Speg. 117
Physalospora zeae Stout . . . . 119
Physalospora zeicola Ell. and Ev. . 120
Placostroma bambusae (Turc.) comb, nov 121
Placostroma sporoboli (Atk.) comb. nov. 121
Pleospora sp 122
' Pyrenophora avenae, see Helminthosporium avenae 122
Pyrenophora bromi, see Helminthosporium bromi 122
Sclerotinia homoeocarpa F. F. Bennett . 123
BASIDIOMYCETES 125
Key to the Basidiomycetes . . . . 125
Corticium fuciforme (Berk.)' Wakef. 126
Discellomyces gloeosporus Olive 127
Entyloma spp 127
Marasmius sacchari Wakker . . . . 128
Pellicularia filamentosa (Pat.) D. P. Rogers . 130
, Pellicularia rolfsii (Sacc.) E. West . 136
Rhizoctonia oryzae Ryker . . . . 137
Rhizoctonia solani, see Pellicularia filamentosa 138
Rhizoctonia zeae Vorhees . . ., . 138
Sclerotium Tode 139
Sclerotium hydrophyllum Sacc. apud Rothert . 139
Sclerotium oryzae, see Leptosphaeria salvinii < 140
Sclerotium rhizodes Auersw. 140
Typhula spp. . . ' . 141
Typhula idahoensis Remsb. 142
Typhula itoana Imai 143
Nonsporulating Basidiomycetes 145
CONTENTS
FUNGI IMPERFECTI
PAGE
SPHAEROPSIDALES 149
Key to the genera of Sphaeropsidales 149
Apiocarpella agropyri 150
Apiocarpella minor 150
Ascoehyta spp. (key) 151
Asoochyta agropyrina (Fairman) Trotter 152
Ascoehyta avenae (Petrak) Sprague and A. G. Johnson 153
Ascoehyta brachypodii (Sydow) Sprague and A. G. Johnson 153
Ascoehyta desmazieri Cav. 155
Ascoehyta graminea (Sacc.) Sprague and A. G. Johnson 155
Ascoehyta hordei Hara . 156
Ascoehyta maydis Stout . 158
Ascoehyta missouriensis Sprague and A. G. Johnson 158
Ascoehyta oryzae Catt. . 159
Ascoehyta phleina Sprague 159
Ascoehyta sorghi Sacc. 159
Ascocliyta sorghina Sacc. 161
Ascoehyta subalpinus Sprague and A. G. Johnson 163
Ascoehyta utahensis Sprague . 163
Ascoehyta zeae Stout . . . . 164
Coniothyrium psammae Oud. . . X, 165
Coniothyrium zeae Stout 165
Cytospora saechari Butl 165
Darluca filum (Biv. ex Fr.) Cast. 168
167
Davisiella elymina (Davis) Petrak .
168
Dilophospora alopecuri (Fr.) Fr.
169
Diplodia bambusae Ell. and Langl. .
'169
Diplodia frumenti, see Physalospora zeicola
170
Diplodia macrospora Earle
171
Diplodia natalensis Pole-Evans
171
Diplodia zeae (Schw.) Lev. 172
Hendersonia spp. /
173
Hendersonia spp., keys 174
Hendersonia crastophila Sacc 176
Hendersonia culmicola Sacc. 177
Hyalothyridium calamagrostidis Greene 177
Leptostromella cynodontis Sacc. 177
Leptostromella panici' Dearness 178
Leptothyrium avenae Tehon . 178
Leptothyrium eylindricum Atk. 179
Leptothyrium zeae Stout 179
Macrophoma Sacc. (key) 180
Macrophoma oblongata Tehon 181
Macrophoma phlei Tehon and Stout / 182
Macrophoma secalina Tehon . 183
Macrophoma zeae, see Physalospora zeae 183
Macrophomina phaseolina (Tassi) G. Gold. 184
Melasmia setariae Atk. . . . • . / 185
Microdiplodia sp 185
Phaeoseptoria spp. (key) 187
Phleospora Wallr 187
Phleospora graminearum Sprague and Hardison 188
Phleospora idahoensis Sprague 188
Phleospora muhlenbergiae Sprague and S^lheim in Solheim
. CONTENTS IX
PAGE
Phoma sp 190
Phyllosticta Pers. ex Fr. (key) 191
Phyllosticta anthoxella Sprague 192
Phyllosticta avenophila Tehon and Daniels . 193
Phyllosticta glumarum (Ell. and Tr.) Miyake 193
Phyllosticta healdii Sprague 194
Phyllosticta helenae Sprague 194
Phyllosticta minutaspora Sprague 195
Phyllosticta miyakei Sydow 195
Phyllosticta owensii Sprague 195
Phyllosticta panici E. Young 196
Phyllosticta rogleri Sprague 198
Phyllosticta sorghina Sacc. 197
Phyllosticta spp. 198
Pyrenochaeta terrestris (Hansen) Gorenz, Walker, and Larson 198
Scaphidium boutelouae Clements 200
Selenophoma Maire (key) 201
Selenophoma bromigena (Sacc.) Sprague and A. G. Johnson 201
Selenophoma donacis (Pass.) Sprague and A. G. Johnson 203
Selenophoma donacis var. stomaticola (Baueml.) Sprague and .. G John-
son . . . 206
Selenophoma everhartii (Sacc. and Sydow) Sprague and A. G Johnson 209
Selenophoma obtusa Sprague and A. G. Johnson 211
Septori a Fr. em. Sacc 211
Septori a Fr. em. Sacc. (key) . . . . 212
Septor: a agropyrina Lobik . . . . 215
Septori a andropogonis J. J. Davis . 216
Septori a andropogonis var. sorghastri H. C. Greene and Spraigue 218
Septori a andropogonis forma sporobolicola Sprague 218
Septor a arctica Berk, and Curtis 219
Septor: a arechavaletae Wint. . . . . 220
Septori a avenae Frank 220
Septor: a avenae Frank f. sp. triticea T. Johnson 222
Septor a brevispora Ellis and J. J. Davis 224
Septor a bromi Sacc 225
Septor a bromi var. phalaricola Sprague 225
Septor a calamagrostidis (Lib.) Sacc. 227
Septor a calamagrostidis f. koeleriae (Cocc. and Mor.) Sprague 229
Septor a calamovilfae Petrak 230
Septor: a, carricerae Fairman . . . . 230
Septor a cenchrina J. J. Davis 231
Septor: a cjTiodontis Fuckel . . . . 231
Septori a digitarivbra Sprague . . . . 232
Septor a elymi Ell. and Ev 232
Septor: a on Agropyron and Elymus, key 233
Septor a glycericola Sprague . . . . 235
Septor a infuscans (Ell. and Ev.) Sprague 237
Septor: a jaculella Sprague . -. 238
Septori a loligena Sprague 239
Septor: a macropoda Pass! . . . . 241
Septori a macropoda var. grandis Sprague 242
Septori a macropoda var. septulata (Gonz. Frag.) Sprague 242
Septori a melicae Pass 243
Septori a mississippiensis Sprague 243
Septori a munroae Ell. and Earth. . 244
Septori nodorum (Berk.) Berk. . 244
X CONTENTS
SPHAEROPSIDALES' (Cont.) PAGE
Septoria oryzae Catt. 246
Septoria oudemansii Sacc. 246
Septoria pacifica Sprague 247
Septoria passerinii Sacc. 249
Septoria pertusa Heald and Wolf 251
Septoria poliomela Sydow 251
Septoria quinqueseptata Sprague 252
Septoria secalis Prill, and Del. 253
Septoria secalis var. stipae Sprague 254
Septoria spartinae (Trel.) Sprague 254
Septoria stipina Died. . . . . 255
Septoria tandilensis Speg. 257
Septoria tenella Cke. and Ell. 259
Septoria triseti Speg. em. Sprague . 261
Septoria tritici Rob. in Desm. 261
Septoria tritici f. avenae (Desm.) Sprague 265
Septoria tritici f. holci Sprague 267
Septoria tritici var. lolicola Sprague and A. G. Johnson 267
Stagonospora Sacc. (keys) 269
Stagonospora agrostidis f. angusta Sprague 270
Stagonospora arenaria Sacc. . . ./ 271
Stagonospora brachyelytri Greene . , 272
Stagonospora bromi A. L. Smith and Ramsb.'. 273
Stagonospora curvula Sacc, Bomm. and Rouss. 274
Stagonospora foliicola (Bres.) Bubak 274
Stagonospora glycericola Sprague 276
Stagonospora graminum Sacc. and Scalia. 276
Stagonospora intermixta (Cooke) Sacc. 276
Stagonospora ischaemi Sacc. ,277
278
Stagonospora maculata (Grove) Sprague
278
Stagonospora maritima Sydow
279
Stagonospora paspali Atk.
279
Stagonospora simplicior Sacc. and Berl.
280
Stagonospora spartinicola Sprague . / 281
Stagonospora subseriata (Desm.) Sacc. 282
Stagonospora vexatula Sacc. and other saprophytic species
MELANCpNIALES . .' . . ,. 285
Key to the Melanconiales . . . . . 285
Colletotrichum falcatum Went, see Physalospora tucumanensis- 285
Colletotrichum graminicola (Ces.) Wils. . 286
Cylindrosporium bambusae Miyake and Hara 288
Cylindrosporium calamagrostidis Ell. and Ev. 288
Cylindrosporium glyceriae Ell. and Ev. . 289
Gloeosporium bolleyi Sprague 289
Gloeosporium graminum Rostr. . [ . 292
Gloeosporium meinersii Sprague 292
Melanconium iliau Lyon, see Gnomonia iliau Lyon 293
Melanconium sacchari Massee apud Speg. 293
Septogloeum bartholomaei (Pk.) Wr. 294
Septogloeum oxysporum Sacc, Bomm. and Rouss. 294
Septogloeum spartinae (Ell. and Ev.) Wr. ' . 297
MONILIALES 299
Key to the Moniliales 299
Alternaria tenuis auct. sensu Wiltshira .'' 301
CONTENTS XI
PAGE
Botrytis cinerea Pers. ex Auct. 302
Centrospora bromi, see Ramulispora bromi 303
Cephalosporium acremomum Corda sensu Fresenius 303
Cercospora spp. . . . 305
Cercospora agrostidis Atk. .' . 308
Cercospora asprellae Ell. and Gall. nom. nud. 308
Cercospora boutelouae Chupp and Greene 306
Cercospora bromi Sprague, see Ramulispora bromi (Sprague) comb. , 307
Cercospora caespitosa Ell. and Ev. 307
Cercospora echinochloae J. J. Davis 307
Cercospora festucae Hardison 307
Cercospora fusimaculans Atk. 309
Cercospora longipes Butler 309
Cercospora muhlenbergiae Atk 310
Cercospora oryzae Miyake 310
Cercospora panici J. J. Davis 311
Cercospora paspali Ray . 311
Cercospora soolecotrichoides Atk. 312
, Cercospora seminalis Ell. and Ev. 312
Cercospora seriata Atk. . 313
Cercospora setariae Atk. 313
Cercospora setaricola Tehon and Daniels 313
Cercospora sorghi Ell. and Ev 314
Cercospora tessellata Atk. 315
Cercospora vaginae Krueger 316
Cercospora zeae-maydis Tehon and Daniels 316
Cercosporella spp. (key) . 317
Cercosporella herpotrichoides Fron 317
Cercosporella holci Sprague 321
Cercosporella poagena Sprague 322
Cerebella andropogonis Ces. . 323
Cladosporium herbarum Lk., see Mycosphaerella tulasnei 324
Coniosporium arundinis Sacc, see Papularia sphaerosperma 324
Coniosporium shiraianum, see Papularia sphaerosperma 324
Curvularia spp. (key) 324
Curvularia geniculata (Tracy and Earle) Boed 324
Curvularia inaequalis (Shear) Boed. 327
Curvularia lunata (Wakker) Boed. . 327
Curvularia trifolii (Kauf.) Boed. 328
330
Dactylaria graminum (Schw.) Sacc.
330
Ellisiella caudata (Pk.) Sacc. .
331
Epicoccum spp
331
Fusarium spp. - •: '
332
»Fusarium spp., key
333
Fusarium acuminatum Ell. and Ev. 336
Fusarium avenaceum (Fr.) Sacc. 337
Fusarium culmorum (W. G. Sm.) Sacc. 339
Fusarium equiseti (Corda) Sacc. 340
Fusarium graminearum Schw. . 341
Fusarium moniliforme Sheldon 344
Fusarium nivale (Fr.),Ces. 346
Fusarium oxysporum (Schl.) em. Snyder and Hansen 347
Fusarium poae (Pk.) Wr. 349
Fusicladium alopecuri Ell. and Ev. 349
Fusicladium destruens Peck 350
Gloeocercospora sorghi D. Bain and Edg.
xii CONTENTS
MONILIALES (Cont.) 'PAGE
Gliocladium spp 351
Hadrotriohum lineare Peck . . . . .| . . . , 351
Helicoceras oryzae Linder and TuUis . . . . . . . 352
Helminthosporium . . . . •, • • 352
Helminthosporium avenae Eidam . . .i . . 353
Helminthosporium bromi Dreohsler . . . . . . . 355
Helminthosporium buchloes (Ell. and Ev.) Lefebvr^ and A. G. Johnson 356
Helminthosporium californicum Maokie and Paxton •-->*. . 357
Helminthosporium carbonum Ullstrup . . . . . • . 357
Helminthosporium catenarium Drechsl 358
Helminthosporium cyclops Drechsl. . . . . . . . 359
Helminthosporium cynodontis Marig. . ' . . 360
Helminthosporium dematioideum Bubak and Wrob. . . . . 361
Helminthosporium dictyoides Drechsl. . . . . . . . 361
Helminthosporium erythrospilum Drechsler . . . . . . 362
Helminthosporium giganteum Heald and Wolf . . . . . 363
Helminthosporium gramineum Rab. . . . . . . . 363
Helminthosporium hadrotriohoides Ell. and Ev. . . . . . 365
Helminthosporium halodes Drechsl. ^ . . 366
Helminthosporium leersii Atk. . . . . . . . . 367
Helminthosporium maydis, see Cochliobolus heterostrophus . ' . . 367
Helminthosporium micropus Drechsl. .' . . . . 367
Helminthosporium monoceras Drechsl 369
Helminthosporium nodulosum Berk. E^nd Curtis . N . . . 369
Helminthosporium oryzae V. Breda de Haan . . . . . . 370
Helminthosporium poae Baudys . . . - . . . . 372
Helminthosporium ravenelii Curt, and Berk. . . . . . . 373
Helminthosporium rostratum Drechsl. . . . ~. . 373
Helminthosporium sacchari (Van Breda de Haan) Butler . . 375
Helminthosporium sativum P.K.B ' 376
Helminthosporium setariae Saw. . . . . . . . . 381
Helminthosporium siccans Drechsl. 383
Helminthosporium sigmoideum Cav., see Leptosphaeria salvinii . 384
Helminthosporium sigmoideum var. irregulare Craliey and Tullis . 384
Helminthosporium sorghicola Lefebvre and Sherwm . . . . 385
Helminthosporium stenacrum Drechsl. • , . . . . . 386
Helminthosporium stenospilum Drechsl. 387
Helminthosporium teres Sacc. . . . . • . . . . 387
Helminthosporium tetramera McKinney . . . . . . 389
Helminthosporium triseptatum Drechsl ^^~ .^^ . . 390
Helminthosporium tritici-repentis Died. . . . "» . . 391
Helminthosporium tritici-vulgaris Nisikado . . . . . . 392
Helminthosporium turcicum Pass 393
Helminthosporium vagans Drechsl. . -. . . . 394
Helminthosporium victoriae Meehan and Murphy . . . . 395
Helminthosporium zeicola Stout . . . ; 397
Helminthosporium spp • 397
Heterosporium avenae Oud. . . . . . . . . . 399
Heterosporium phlei Gregory 400
Mastigosporium Riess ap. Fres. . . . . . . . . 401
Mastigosporium, key . 402
Mastigosporium cylindricum Sprague • _^ - 402
Mastigosporium rubricosum (Dearn. and Barth.) Nannf. . . • . 402
Napicladium arundinaceum (Cda.) Sacc. . . . . . . 405
Nigrospora oryzae (Berk, and Br.) Petch 406
Nigrospora sphaeijica (Sacc.) Mason 407
CONTENTS xui
PAGE
Ovularia hordei (Cav.) Sprague 407
Ovularia lolii Volkart . . . . . 409
' Ovularia pusilla (Ung.) Sacc. and D. Saco. 409
Papularia sphaerosperma (Pers.) von Hoehnel 411
Penicillium expansum (Lk.) em. Thorn 411
Penioillium oxalicum Currie and Thorn 413
Periconia circinata (Mangin) Sacc. . 413
Piricularia grisea (Cke.) Sacc. 415
Piricularia oryzae Briosi and Cavara 416
Piricularia parasitica E. and E. 417
Ramularia graminicola Pk. 418
Ramulispora bromi (Sprague) comb. nov. 418
Ramulispora sorghi (Ell. and Ev.) Olive and Lefebvre 420
Rhynchosporium spp. (key) . " . 421
Rhynchosporium orthosporum Caldwell . 421
Rhynchosporium secalis (Oud.) J. J. Davis • 422
Soolecotrichum graminis Fckl. . . . . 424
Scolecotrichum maculicola Ell. and Kell. 429
•Sperfeaospora avenae (Sprague and A. G. Johnson) comb 430
Spermospora subulata (Sprague) Sprague 431
Sporotrichum columbiense Sprague 433
Sporotrichum peribebuyense Speg. 434
Stemphyllium botryosum Wallr. 435
Stejnphyliium consortiale (Thuem.) Groves and Skolko 435
Ustilaginoidea virens (Gke.) Tak. 436
GLOSSARY 437
LITERATURE CITED . 445
INDEX OF FUNGI . 525
INDEX OF CEREAL AND GRASS HOSTS 531
ILLUSTRATIONS
FIGURE PAGE
1. Aphanomyces camptostylus . . . . . 7
2. Lagena radicicola . . . . . . . 10
3. Olpidium brassicae . . . . . . . 11
4. Polymyxa grammis . . . . . . 16
5. Rhizophidium graminis > . . . . . '47
6. Balansia hypoxylon . . . . . . 57
7. Rhizoctonia solani . . . . . . . 133
8. Ascochyta agropyrina, A. avenae, A. sorghi, A. graniinea, A. braohypodii 154
9. Ascochyta utahensis, A. hordei . . . . 157
IQ. Ascochyta phleina, A. hordei, A. phase of Stagonospora simpUcior, A
missouriensis, A. subalpinus . . . . 157
11. Ascochyta sorghina . . . . . . 162
12. Cytospora sacchari . . . . . . 166
13. Diplodia macrospora, D . zeae, D . frumenti 170
14. Macrophoma phlei . . . . . . 182
15. Phaeoseptoria phalaridis, P. festucae, P . airae 183
16. Phleospora graminearum, P . muhlenbergiae . 187
17. Phleospora graminearum . . . . . 189
18. Phleospora idahoensis . . . 190
19. Phyllosticta anthoxella, P. sorghina, P . rogleri, P . healdii, P . minuta-
spora, P. helenae . . . . . . 192
20. Scaphidium boutelouae . . . . . . 200
21. Selenophoma donacis, S. bromigena, S. obtusa 202
22. Selenophoma donacis . . . . . . 204
23. Selenophoma donacis . . . . . . 205
24. Septoria agropyrina . . . . . . 216
25. Septoria andropogonis f. sporobolicola, S. mississippiensis 217
26. Septoria arctica . . . . . . . 220
27. Septoria avenae, S. melicae, Stagonospora arenaria 221
28. Septoria sp., S. avenae, S. avenae f. sp. triticea 224
29. Septoria bromi . . . . . . . ,226
30. Septoria calamagrostidis . . . . . 22S
31. Septoria calamovilfae . . . . . . 230
32. Septoria digitarivora ^ . . . . . . 232
33. Septoria elymi, S. agropyri, S. elymi f. elymina 234
34. Septoria glycericola, S. nodorum . . . . 235
35. Septoria infuscans . . . . . . . 236
36. Septoria infuscans . . . . ^. 238
37. Septoria jaculella, S. bromivora, S_. brevispora, S. brom 240
38. Septoria macropoda, S. annua . . . . 241
39. Septoria macropoda var. grandis . . . . 242
40. Septoria munroae . / . 244
41. Septoria inodorum . . . . . . . 245
42. Septoria oudemansii . . . . . . 247
248
43. Septoria elymi-europaei, S. pacifica, S. agropyrina .
250
44. Septoria passerinii, S. microspora . . . .
252
45. Septoria poliomela . . . . . .
253
46. Septoria quinqueseptata, S. calamagrostidis f. koeleriae
xvi ILLUSTRATIONS
FIGURE
\
PAGE
47. Septoria secalis . . . . . .
255
48. Septoria spp. . . . . . . .
255,
49. Septoria tenella, S. festucina ( = S. tenella), Phjllosticta spjp., S. festucae
( = S. tenella) '\ 258
50. Septoria triseti, S. secalis var. stipae, S. phleina, S. stipina 262
51. Septoria tritici, S. tritici f. avenae, S. negleota 264
52. Septoria tritici, S. tritici f. avenae, S. avenae . 266
53. S. tritici var. lolicola, S. tritici var. holoi 268
54. Stagonospora agrostidis f. angusta . 270
55. Stagonospora arenaria . . . . . 272
56. Stagonospora bromi . . . . . 273
57. Stagonospora foliioola, St. isohaemi 275
58. Stagonospora paspali, St. maculata, St. bromi, St. subseriata, Septoria
avenae or St. arenaria 279
59. Stagonospora simplicior var. andropogonis, St. subseriata, Stagonospora
maculata . . . . 281
60. Cylindrosporium calamagrostidis 289
61. Gloeosporium boUeyi 291
62. Melanconivim sacchari 293
63. Septogloeum o.xysporum 293
64. Septogloeum oxysporum
/ 297
65. Cercospora festucae (C. apii) 3^8
66. Cercospora sorghi . . . .- 315
67. Ceroosporella herpotrichoides 318
68. Cercosporella herpotrichoides 319
69. Cercosporella holci 322
70. Cercosporella poagena 323
71. Curvularia geniculata 325
72. Curvularia trifolii, Brachycladium spiciferum 329
73. Helminthosporium rostratum . 374
74. Mastigosporium rubricosum . 404
75. Ovularia hordei 408
76. Ramulispora bromi 419
77. Rhynchosporium orthosporum 421
78. Rhynchosporium secalis . 424
79. Scolecotrichum graminis 427
80. Spermospora subulata 433
81. Sporotrichum columbiense, S. peribebuyense 434
DISEASES OF CEREALS AND GRASSES
IN NORTH AMERICA
INTEODUCTION
The fungous diseases of the grass family are often better known
than the causes of these diseases. It is the primary aim of this manual
to aid the worker in the identification of these parasitic fungi and the ,
diseases caused by them. Therefore, the technical description, including
disease symptoms, host range by states or provinces, world distribution,
and citations to important literature'are given for each parasitic fungus.
Since the writer is a plant pathologist he has added, in each case, not
only data on the fungus but on the disease of which it may at least in
part be the cause. These data are strongly reflected in the literature
citations which refer not only to the literature of mycology but to
reports of plant pathology in even greater numbers.
While the manual deals with parasitic fungi it includes frequent
mention of saprophytic fungi sometimes associated with the parasitic
ones. It also includes borderline instances where the parasitic or sapro-
phytic nature of the organism has not been definitely established. The
problem in these instances is putlined. The area covered includes all the
mainland of North America, the Central American countries, and
Hawaii but not the islands of the West Indies.
The fungi in this manual are listed alphabetically under the four
classes of fungi: (1) Phycomycetes, (2) Ascomycetes, (3) Basidio-
mycetes, and (4) Fungi Imperfecta The manual contains discussions
of 384 species and subcategories of species of parasitic fungi, as well as
a number of associated saprophytes found growing in the region. The
number given, 384, is, of course, an approximation, as no two individ-
uals would segregate_^this bulk of fungi in exactly the same manner.
There are approximately 34 Phycomycetes, 88 Ascomycetes, 15 Basidio-
mycetes, and 247 species and subcategories of Fungi Imperfecti in-
cluded. In addition, there are discussed a large number of species inci-
dental to those which are known to occur in North America. It was
felt that mention should be made of foreign fungi in as many instances
as space permitted, to acquaint the routine worker with these forms on
the chance that these species might be encountered on' this' continent.
In most cases such fungi are mentioned only, or their morphology is
given in brief outline form. However, there are some instances where
the foreign species is discussed in more detail because of its similarity
to local forms or because it is likely to reach our shores within a life-
time.
In the descriptions of the species given herein the original, often

scanty, has been amplified where possible through the utilization of
3
4 DISEASES OF CEREALS AND GRASSES
additional reliable descriptive data. In, only a few cases the descrip-
tions seemed overly long and needlessly •w'ordy. In the earlier pub-
lished works the descriptions equal or approach the status of nomina
nuda in some cases. Within the bounds of rules laid down or sug-
gested by the several International Congresses on botanical nomew-'
clature (cf. Bisby, 1945), the endeavor has been to present the descrip-
tions in usable form. As mentioned above, the genera are listed alpha-
betically under the several classes. The species discussed are also
listed alphabetically under each genus.
Keys to the genera represented under the several classes are given
under each of these classes.
PHYCOMYCETES
A key to the genera of fungi discussed and represented is as follows:

A. Mycelium scanty or lacking, consisting of a few delicate hyphae; propa-
gated by amoeboid cells and spores or by sporangia and zoospores; sex--
cells rare
B. Mycelium lacking
C. Zoosporangia elongate, relatively large, with exit tubes; in root hairs
Polymyxa
CC. Zoosporangia sac-shaped, attached to cell wall by a collar; in root
hairs and roots Lagena
CCC. Zoosporangia or sori sphaeroidal on above-ground parts
Synchytrium
BB. Myceliuni scanty but usually evident in part of life cycle
C. Zoospores anteriorly uniflagellate, usually formed outside the sporan-
gium -. Hyphochytrium
CC. Zoospores posteriorly uniflagellate, formed inside sporangium
D. ThaUus holocarpic . . . / Olpidium
D D . ThaUus eucarpic
E. ThaUus monocentric Rhizophidium
EE. ThaUus polycentric, in part at least
F. ThaUus endobiotic, polycentric Cladochytrium
FF. Sporangia epibiotic, thallus monocentric; resting spores endo-
biotic, thaUus polycentric Physoderma
AA. Mycelium present, well developed, sex cells usually present
B. Conidia in globose stalked sporangia Rhizopus
BB. Conidia, if present, on naked conidiophores
C. Sporangia filamentous, producing by evacuation many zoospores
Aphanomyces
CC. Sporangia not filamentous
D. Causes a leaf shredding and mUdew of leaves Sderospora
D D . Causes a_ soft rot of leaves and roots Phytophthora
D D D . Conidia or gemmae not well differentiated from mycelium;
causes root decay Pythium
Aphanomyces camptostylus Drechsl.—Root Necrosis
Mycelium hyaline, 2.5-9.5 /A diam., sparingly or moderately

branched, with very scanty aerial development in pure culture.
Sporangia formed from vegetative hyphae, the delimiting septa occur-
ring as narrow cross walls or as curved or irregular partitions; sporangia
sometimes very extensive, including segments of the axial h y p h a e more
t h a n 3 m m . in length, together with numerous ramifying elements, t h e n
often provided with plural evacuation h y p h a e ; the latter 2 m m . or
5
more long, usually rather stout, often 6-7.5 /t diam., not markedly
narrowed toward the apex, bearing at times op the distal portion one
to four lateral protuberances or branches up to 30 /* long, which, like
the tip of the axial filament, may open to permit egress of zoospores.
Zoospores few to 300, diplanetic, on encystment usually .7-10 fi diam.,
developing a papilla approximately 2-2.5 /x diam., the cylindrical wall
of which persists on the empty cyst membrane. Oogonia terminal on.^
short branches or on longer hyphal elements, subspherical, smooth,
19-26 ij. (average 22.9 ju) diam., wall 0.6-1.2 /x (average 0.9 /J.) thick,"
with somewhat irregular inner contour; the oogonial stalk regularly
making contact with an antheridial branch, the stalk itself, and often
one or several branches arising from it or from the parent filament,
frequently coiling about the antheridial hypha. Antheridia as many as ,
four, composed of an inflated portion, 8-11 X 4.5-9 fi, frequently with
narrower, apical, proximal, or lateral extensions; regularly functional,,
communicating with the interior of the 'Oogonium by a very short fer-
tilization tube; borne terminally on supporting branches, 2-3 /.<. diam.
The latter branches usually diclinous in origin, the mycelial connec-
tion with the oogonia rarely demonstrable; encircling oogonium ex-
tensively and, when crowded, often coiled with the antheridia.
Oospores single, subspherical, colorless or chlorinous, mostly 16-21 /x
(average 18.8 jn) diam., surrounded by a wall 1.1-2 ju (average 1.5 /x)
thick, and containing at early maturity a slightly eccentric reserve
globule 9-13 ju, (average 11 /x) diam., germinating regularly either by the
direct production of mycelium or by formation of an unbranched germ
sporangium as long as 400 /x, discharging 10 to 17 zoospores, in most
instances. ,
On: Avena byzantina K. Koch, Oreg.—A.: sativa L., Minn., Mont., Oreg.,
Wise.—Festuca elatior L., N. Dak.—F. rubra L., N. Dak.—Setaria viridis
(L.) Beauv., Mont.—Stipa viridula Trin., N. Dak.
Very little study has been made of' this fungus during the years
since Drechsler presented his detailed taxonomic hiveatigations of this
and other species of Aphanomyces (1929,a). A. camptdstylus is not
readily isolated by the usual techniques. It sometimes develops on
water agar from water-washed pieces of .roots. Drechsler used distilled
water to facilitate isolation.
This fungus is probably of some economic importance in western
Oregon on red oats [A. byzantina) and common oats {A. sativa). It
occurs in the northern Great Plains and in Minnesota, where it is rela-
tively abundant in certain areas in the central and northern parts of
the state. It is particularly common in acid soils in northern Minnesota
(Sprague, isolation records). - "^ -
A. camptostylus has not been definitely associated with any specific
symptoms. In Oregon, where it is one of the components of root rot in
acid soils in the coastal region, oats aje stunted and possess thickened,
PHYCOMYCETES
yellowed roots. This condition has been called "brittle root" (Sprague,
1938,a).
" The genus Aphanomyces is characterized by the filamentous spo-
rangia which produce, under certain conditions, large numbers of
uniseriately arranged, diplanetic zoospores
(Fig. 1).
Range: United States, Japan.
References: Drechsler (1929,a, pp. 335-42,
Figs. 9-11); Ideta (1909); Sprague (1938,a).
Cladochytrium graminis Buesgen—"Damp-

ing-Off" (after Clinton)
Mycelium thread-like, with swellings.
Resting spores 30 X 40 /A; no further de-
scHption given (Fischer, A., 1892).
On: Agrostis alba L., Conn.
Krieger issued this species as Fungi
Saxonici No. 441 in 1888. The fungus was
vaguely described by Buesgen in 1887 while
Fischer listed it in 1892. Clinton (1934)
states that this organism was associated
with damping-off of redtop [Agrostis alba)
in Connecticut. It occurred in company with
Helminthosporium sp. and Pythium iii part
of his material. Clinton found it on the roots
and sheaths and all evidence seems to indi- FIG. 1.—Fragment of
cate a somewhat dubious parasitic nature. filamentous sporangium
of Aphanomyces campto-
Sparrow (1943) does not mention this spe- stylus extruding zoospores.
cies in his text on aquatic species of Phy-
comycetes, but he does give the following description of C. tenue
Nowakowski which has been reported on Glyceria spectabilis M. and
K. in Germany.
Sporangia intra-, or extramatrical, formed by a swelling of the
rhizoid or by enlargement of a segment of a septate turbinate cell,
when the sterile segment either remains at the base as an empty ap-
pendage or eventually becomes a sporangium, spherical or somewhat
pyriform, up to 66 /* diam. (average about 18 /*), with a niore or less
prolonged regular or irregular rather stout discharge tube (rarely sev-
eral) the tip of which is either extramatrical or penetrates a neighbor-
ing cell, wall smooth, colorless, proliferating, the secondary sporangia
smaller; rhizoidal system extensive, branching, the rhizoids 1-2 /j.
diam., with spindle-shaped or spherical swellings which are often trans-
versely septate and divided into two equal parts (occasionally three);
zoospores spherical, 5 /x, diam., with a colorless eccentric globule and a
posterior flagellum, emerging upon the deliquescence of the tip of the
discharge tube and forming a temporary motionless mass imbedded in
"slime," movement amoeboid or swimming; resting spores not observed.
D. J. C. Jones (1937) mentions damping-pff of young grass seedlings
by C. graminis (? C. caespitis, of. Cook, W.lR. I.,;R. A. M'. 13:520) in
England. Small yellow spots develop on the young lawns during wet
weather. Dry conditions check the disease. Cheshunt compound (1 ozT.
in 2 gals, of water) is sometimes used as a drench of infected turf.
The relation of these species to other members of the Chytridiales
is discussed by Sparrow but since this group is somewhat doubtfully
parasitic on Gramineae the above descriptions and discussions are ade-
quate for our purposes.
Range: United States, Germany.
References: Buesgen (1887); Clinton (1934, p. 260); Fischer, A. (1892);
Jones, D. J. C. (1937); Sparrow (1943).
Hyphochytrium catenoides^ Karling—On Corn Roots

Thallus predominantly polycenfric and intramatrical; usually hy-
pha-like, and consisting of a linear series, up to 500 [i in extent, of in-
tercalary and terminal swellings and sporangia, connected by tubular
hyphae or isthmuses 1-38 ju long' and 2.2-4 fi diam.; occasionally mono-
centric, oval, elongate, spherical, and Olpidium-hke. Zoosporangia ter-
minal or intercalary, delimited by cross septa, hyaline, smooth, spheri-
cal and 10-35 jn diam., or oval and 10 X 12 /x to 18 X 22 /x or broadly
spindle-shaped, 6 X 10 /x to 9 X 18 ft or elongate and sometimes slightly
irregular, with 1-4 single or branched, straight, curved, coiled, or irreg^-
ular exit tubes, 5-250 /x long, 3-6 ju. diam. Contents of sporangia usually
emerging to the outside of the host as nak6d masses and undergoing
cleavage into zoospores; occasionally;undergoing complete or partial
cleavage in the sporangium whereafter the segments glide out in suc-
cession. Zoospores anteriorly uniciliate, slightly flattened, oval, and
elongate, 3.5 X 1-5 /x to 3 X 2 /x, with .several small, slightly refractive
granules. Sexuality and resting spores unknown.-^
On: Zea mays L., N. Y.
This chytrid was described by Karling (1939) as a weak parasite'
on corn and cryptogamic plants. Because it grows very slowly on
nutrient agar he could not free the feebly growing fungus from bac-
teria. It is characterized by anteriorly uniciliate zoospores, a hypha-
like thallus, and elliptical to globose sporangia connected by hypha-
like isthmuses. The fungus, as illustrated, is somewhat Pythium-Vike.
Karling places the species in the Hyphochytriaceae. Our observations
indicate that fungi similar to this occurôn the roots of grasses in
North Dakota.
Range: United States.
References: Karling (1939); Sparrow (1943).
PHYCOMYCETES 9
Lagena radicicola Vanterpool and Ledingham—Rootlet Parasite

Mycelium lacking. Zoospores darkly granular, bean-shaped, with
two flagella attached in the lateral depression; about 7 X H /* when
motile and 6 n diam. when at rest; rounding up on surface of host and
penetrating it by germ tube which enlarges and develops into a coeno-
cytic, sac-shaped, sometimes curved, occasionally elongate-lobulatc,
multinucleate thallus. Thallus occupying the cortical and epidermal
cells, as well as the root hairs of the finer roots of the host. Mature
thalli occurring singly or aggregated in one cell; varying considerably ^
in size, average 14 X 35 /j., sac-shaped, attached to cell wall of host hy
a collar which in surface view appears as two concentric circles; func-
tioning either as zoosporangia or coenogametangia. Zoosporangium dis-
charging its contents extramatrically through a single emission tube
(10-15 X 4 ;u) into a vesicle where the zoospores, from a few to over
50, are formed and liberated. Contents of male thallus passing via a
conjugation tube, into a female thallus, where a smooth, double-walled,
subspherical resting spore (oospore) with central globule and refringent
spot, is formed; oospore varying from 10 to 25 p. diam. A single oospore
or several may be present in each host cell. External conditions largely
determine whether zoosporangia or oospores will be formed; Both
zoosporangia and oospores may be formed concomitantly in the life his-
tory of this organism. Oospore germination not noted.
On: Agropyron repens (L.) Beauv., Ont.—Hordeum vulgare L., Ont., Sask.
—Secale cereale L., Ont., Sask.—Triticum aestivum L., Ont., Sask.—T.
durum Desf., Sask.—Zea mays L., Ont., Sask.
Vanterpool and Ledingham (1930) erected a new genus to include
this fungus which belongs with the lower or primitive Phycomycetes.
The fungus was originally associated with a root browning (1930) but
now it is considered a minor parasite on rootlets of cereal and grass
plants. The infection of the rootlets occurs after zoospores germinate,
sending infection threads into the host. A thallus is formed which re-
mains attached to the host cell wall.- The thallus becomes the sporan-
gium which sends ite'contents through the collar (Fig. 2,^) or point of
attachment to the outside of the host cell. The vesicle outside expands
into a spherical body, forms zoospores, and finally breaks. The sexual
stage may occur when two thalli are found in the same host cell. A
fertilization tube or antheridium .grows from one thallus to the other.
A thick-walled oospore (Fig; 2,5) which resembles Pythium develops
following union of the two gametangia. This fungus is said to be closest
to Lagenidium of the Ancylistaceae (Lagenidiales cf. Sparrow, 1943).
Truscott (1933) noted some differences in the material he obtained in
Ontario from that in Saskatchewan. The organism is discussed by
Karling (1942,a).
Range: Canada.
References: Karling (1942,o); Truscott (1933); Vanterpool and Leding-
ham (1930). I
FIG. 2.—Lagena radicicola: A, developing sporangium showing collar attached

to host at narrowed exit end; B, resting spores (after Truscott).
Olpidium brassicae (Wor.) Dang.—Rootlet Necrosis

Olpidiuster radicis (de Wild.) Pascher
Asterocystis radicis de Wild.
Olpidium radicicolum de Wild.
Zoosporangia solitary or aggregated in the host cell, thin-walled,

varying from spherical, 12-20 /J. diam., to elongate, 25-220 X 20-45 ju,,
opening by 1-4 exit tubes which v a r y in length according t o the dis-
t a n c e from the host surface. T h e uniciliate zoospores are 3 [i diam.
Resting sporangia usually spherical, 8-25 /A diam., oval, u p to 30 /*
long. Exospore coarsely wrinkled with^ ridges up to 3.5 ju, high, show-
ing a stellate form with 6-9 points in optical section. Endospore is
t h i n and smooth, free from the exospore. I
On: Agropyron trachycaulum (Lk.) Malta, Sask.—Avena sativa L., Sask.—

Hordeum vulgare L., Sask.—SecaZe cereale L., Sask.—Triticum aestivum
L., Sask.—Zea mays L., Sask. *'
This is another obligate parasite" on- the roots and rootlets of
higher plants (Fig. 3 ) . I t has been noted by workers in western Europe
on various crops, especially Gramineae and Cruciferae. Sampson
(1939,b) has concluded t h a t t h e fuijgus on Gramineae is t h e same as
PHYCOMYCETES 11
the one that is well known on Brassica and all evidence indicates that
her diagnosis is correct. She also described (1932) 0. agrostidis.
Among those who have studied this fungus, Fron and Gaillat
(1925) reported it on a turf composed of bent and fescue grasses.
The disease occurred as more or less extensive brown patches of the
turf. Inoculation trials were positive on Lolium, Agrostis, and Festuca;
slight infection of oats was obtained but wheat and flax did not be-
FiG. 3.—Olpidium brassicae: A, sporangia; B, resting spore (semi-diagram-

matic).
come infected. Guyot (1927,a) could readily infect oats in France

but could infect wheat only when growing in very heavy soil. He
found 0. brassicae associated with chlorotic wheat and oats in low,
wet ground in early.spring.
Range: Belgium, Canada, France, Italy, Morocco, Wales.
References: Fitzpatrick (1930); Fron and Gaillat (1925); Guyot (1927,o);
Karling (1937,a,&); Peyronel (1922; 1923;- 1924; 1926,6); Sampson
(1939,6); Rives (1925); Vanterpool (1930); de Wildeman (1893); Woro-
nin (1878).
Physoderma inaydis Miyabe—^Brown Bundle Disease

Cladochytrium maydis Miyabe nom. nud.
Physoderma zeae-maydis Shaw
Occurs within the parenchymatous cells of the culms, the outer
sheaths, and the leaves, particularly in the region of the midribs, pro-
12 DISEASES OP CEREALS AND GRASSES
I
ducing numerous, orbicular, elliptical or linear spots; spots mostly
small, but numerous and confluent, brown pv fuliginous, light in color
near the margin, much deeper colored at.the center; resting spores
ellipsoid-ovate or globose, deep brown, with a thick, smooth wall,
slightly flattened on one side, which is provided with a circular cap
or lid, 18-24 X 20-30 /x; lid of the resting spote opening after winter
incubation, liberating swarm spores 3-4 X 5-7 n, with one long polar
cilium; swarm spores producing fine fibrous hyphae on germination. .
Sporangia epiphyllous, irregular in outline, mostly slipper-shaped,
variable in size, apically papillate, proliferating, anchored to the host
cell by a coarse, branched rhizoidal system arising from a small basal
apophysis; sporangia liberating innumerable (more than 300) zoospores
(gametes) through a broad pore resulting from the deliquescence of
the papilla; zoospores (gametes?) smaller than those escaping from the
resting spores.
On: Zea mays L., Del. and N. J., southward in great abundance to the Gulf
of Mexico; Ark., 111., Ind., Iowa, Kans., Ky., Md., Minn., Nebr., Ohio,
Okla., S. Dak., Tenn., Tex., W. Va. Rare in the far west but reported
from California, once from Philomath, Oreg., from Puyallup, Wash., in
1947, and from Pullman, Wash^,- in 1948, Costa Rica, Guatemala, El Sal-
vador. Distribution map shown by Tisdale (1919, Fig. 1, p. 138).
Miyabe first described P. maydis in 1909 (Ideta, 1909, ed. 4) from
material collected in Japan by K. Sengoku (1901). The fungus was
called Ckidochytrium sp. by Sengoku (1901) and then C. piaydis
Miyabe (nom. nud.) in 1903 and 1904 (Ideta, 1903, ed. 3). I t was called
P. zeae-maydis Shaw in 1912 (Sydow, Sydow, and Butler, 1912), the
• name by which it is generally called today. However,^ there appears to
be no fundamental difference between the/Japanese material of Sen-
goku, Miyabe, and others and the Indian material of Shaw which W. H.
Tisdale (1919) considered the same| as the specimens Tisdale had
studied in the United States. It appears reasonable to accept the
earlier, adequately described binomial, P. maydis, which, incidentally,
is shorter than the one proposed by Shaw. " --,,
AV. H. Tisdale gave a detailed account of the life Mstory of Physo-
dernia maydis (1919). Sparrow (1934) supplemented this study with
the notation that true sporangia occur-in the species. He (1943, 1947)
determined that the bodies which developed such neat trap-door oper-
cula or lids were resting spores rather thap actual sporangia.. Tanaka's
English-language review of earlier Japanese articles is helpful (1922),*'
while Heald's Manual gives a good student's summary of the disease
and its causal organism (Heald, 1933, pp. 475-479).
W. B. Tisdale (1931) reported that thfii.most effective way to pro-
duce infection in the field was to pour a 'water suspension of resting
spores into the top of young corn plants when they were 3-4 feet tall.
He failed, however, to infect sorghums, millet, and certain other warm-
temperature grasses. Eddins (1933)'found that 38 southern varieties
PHYCOMYCETES 13
of dent and flint corn were susceptible. He found that resting spores
survived for 2-3 years under a variety of conditions. These results indi-
cate that control of this organism is difficult. Fortunately it appears to
be somewhat spasmodic in its outbreaks.
J. A. Stevenson (1946) described Physoderma paspali on Paspalum
flicatulum from Puerto Rico. I t has golden brown, smooth, resting
spores, 18-33 X 15-24 /x, which are larger than those of P. maydis or
p. gerhardti.
What appears to be a species of Physoderma was obtained on ne-
crotic leaves of Beckmannia syzigachne (Steud.) Fern, at West Fargo,
North Dakota, and near Moorcroft and Lonetree, Wyoming (Sprague
and Fischer, 1948). Thirumalachar and Dickson (1947) reported a
Physoderma disease of quack grass. The fungus had nonerumpent sori
and sporangia, 20-40 X 20-34 fi. Those on Beckmannia were perhaps
snialler. Childers (1948) reported Physoderma graminis on Agropyron
repens from Ottawa. The diseased plants were dwarfed and bore yel-
lowish to brown stripes. They were conspicuous in close-mown fields
by reason of their stiil, erect leaves and general cblorotic appearance.
Range: United States, China, Costa Rica, El Salvador, Gold Coast, Guate-
mala, India, Japan, probably widely distributed.
References: Heald (1933, pp. 475-479); Miyabe in A. Ideta (1909, ed. 4);
Shaw in Sydow, Sydow, and Butler (1912); Sparrow (1934, 1943, 1947);
Sengoku (1901); Tanaka (1^22); Tisdale, W. H. (1919, 1920); Tisdale,
W. B. (1931); Vorhees (1933,6), Wellman (1949).
Phytophthora cactorum (Leb. and Cohn) Schroet.—Rot

Oogonia pyriform or subspherical, usually with a broad, funnel-
shaped base; oospores straw-color to paler, smooth and thick-walled,
filling most of the interior of the oogonia; antheridia usually paragy-
nous, attached near base of the oogonium; oogonia and their oospores
produced profusely on various media, averaging 26-30 /i (oogonia)
and 24-27 /x (oospores). Typified by profuse production of oogonia with
paragynous antheridia, the absence of typical chlamydospores, the
small and often inconspicuously papillate sporangia, and by its growth
on cornmeal agar at 27.5° C. (Tucker, 1931).
On: Avena fatua L., Calif., Oreg.—Echinochloa ctusgalli (L.)-Beauv., S. Dak.
—Panicum capillare L., Mont.
P. cactorum and certain undetermined collections of Phytophthora are
sometimes found associated with soft rots of the leaves and roots of grains
and grasses. In addition'to the host and geographical range given above for
P. cactorum undetermined species of Phytophthora have been reported on
the following hosts from the states indicated: Agropyron repens (L.) Beauv.,
Minn.—A. sibiricum (Willd.) Beauv., N. Dak.—Avena sativa L., Calif.—
Hordeum jubatum L., N. Dak.—Lolium multiflorum Lam., Oreg.—Poa vasey-
ochloa Scribn., Oreg.—Stipa viridula Trin., N. Dak.—Triticum durum Desf.,
' I
N. Dak.—Zea mays L., Ariz., Hawaii, S. Dak. P. 'parasitica Dastur was
recently found on greenhouse corn in Indiana Ijy Ullstrup and Tucker (1948).
McMurphy (1916) noted a species similar to P. calocasiae Rac. on
oats in California (see also Tucker,,1931); The infected areas were yel-
lowish when young, whitish later, and finally brown or reddish brown.
We have seen scanty material of P. cactorum on Avena fatua from near
The Dalles, Oregon,'and on the same host from the coastal region of
northern California. No symptoms were evident other than a water-,
soaked appearance of the leaves which, incidentally, were in contact
with the soil. The most abundant material that the writer has obtained
was in stunted plants of barnyard grass (Echinochloa) growing near
Belle Fourche, South Dakota. This material was identified as P. cat-
torum. The North Dakota isolations of Phytophthora spp. were made
from the roots of seedlings. Usually other fungi were also present and
frequently the Phytophthora represented only a minority of the isola-"
tions made. P. cactorum was isolated in moderate quantities from'roots
of corn growing at Newell, South Dakota, in June. In the greenhouse
at Mandan, North Dakota, at the/Northern Great Plains Field Station
artificial inoculations with this iungus, added to the soil at seeding
time, caused 88 per cent loss injTurghai proso {Panicum miliaceum L.),
only 6 per cent loss in crested wheat grass {Agropyron cristatum L.)
Gaertn., scarcely a trace in wheat, 70 per cent in Ladak alfalfa, 71 per
cent in blue grama [Boutelova gracilis (H. B. K.) Lag.], but only a
trace in corn. The type of injury was mainly seed rot at the tempera-
ture range used (48° to 70° C ) . In general, Phytophthora is'not im-
portant on Gramineae. Helena de Bruyn (1922) has shown that species
of Phytophthora can live for some time as soil saprophytes.
Range: United States, Hawaii, probably widely, but perhaps irregularly,

distributed on Gramineae. |
References: Blackwell (1943); de Bruyn (1922); McMurphy (1916);
Sideris (1932); Sprngue (1946,e); Tucker (1931, 1933).
I
Phy'tophthora parasitica Dastur—CTrmvn Jlot

Causes a wilting, with subsoil-line, slowly spreading crown rot,
decay firm, deep brown, slightly flepressed; fungus produces an
abundance of sporangia and zoospores in sterile distilled water after"
previous nourishment on sterile pea broth, zoospores biflagellate, kid-
ney-shaped, sporangia 47-48 X 21-46 fi, 'chlamydospores 18-40 /n.
On:- Zea mays L., Ind.
Ullstrup and Tucker (1948) identified, this sp/cies from corn plants
growing in the greenhouse. . "
Reference: Ullstrup and Tucker (1948).
PHYCOMYCETES 15
Polymyxa graminis Ledingham—Root Hair Necrosis

Resting spores spherical or polyhedral, 5-7 /A diam., smooth, with
yellow-brown outer walls. Zoosporangia large with persistent smooth
thin walls; zoosporangia formed by septation of the mature thallus.
Discharge tube segmented. Zoospores numerous, 4-5 n diam., difiagel-
late and heterokont, discharged without vesicle formation. Nonhyper-
trophying, obligate parasite.
On: Agropyron repens (L.) Beauv., Ont.—Hordeum vulgare L., Ont.—Secale
cereale L., Ont.—Triticum aestivum L., Ont.
Ledingham created the monotypic genus Polymyxa and placed it in
the Plasmodiophorales. It has cystosori similar to those of Lignierq but
the large elongate zoosporangia with conspicuous exit tubes distinguish
it from the latter genus. The mature thalli are developed by progressive
lobular outgrowths from uninucleate amoebae and always have a thin
surrounding wall. The mature thallus is transformed by septation into
zoosporangia. In the formation of resting spores naked, multinucleate
myxamoebae develop first, these segment to form spore clusters or
cystosori, which lack a soral membrane. Both zoosporangia and resting
spores produce identicalzoospores with two flagella of unequal length.
Ledingham erected the new genus because the zoosporangial character-
istics differed from those of any described genus in the Plasmodiopho-
rales (Fig. 4).
This species invades the root hairs (Fig. 4,A) and smaller roots of
Gramineae but apparently does slight damage. Ledingham first noted
it in the fall of the year in Ontario. I t was found in various soils and
appears capable of withstanding long periods of drought or cold.
Schwartz (1911) described a similar fungus, Sorosphaera graminis
on Poa anniiu L. from England. Maire and Tison (1911) proposed
Ligniera n. gen. as a member of the Plasmodiophorales. The genus was
characterized by its failure to cause host hypertrophy. They trans-
ferred (1911) Sorosphaera junci Schwartz (1910) to Ligniera and later
W. R. I. Cook (1926) placed S. graminis [L. graminis Schwartz, 1914)
under L. junci on the basis of cross-inoculation studies. L. pilorum Fron
and Gaillat (1925) is another synonym according to Cook. Palm and
Burk (1933) go still further arid place Sorosphaera (sensu Schroeter,
non Fitzpatrick), Spongospora Brunchorst, Ligniera, Sorodis(yus
Lagerh. and Winge, Ostenfeldiella Ferd. and AVinge, Clathrosorus Ferd.
and Winge and Membranosorus Ostenfeld and Petersen under one
genus which would presumably' be Plasmodiophora Wor. Guyot
(1927,6) reported a fungus on Bromus and Festuca referable to L.
verrucosa.
Karling (1942,6) reported that J. T. Barrett found material on
roots of Poa annua at the University of California which resembled
Sorosphaera radicalis Cook. In addition to cystosori and resting spores
he observed thin-walled sporangia which produce biflagellate, hetero-
kont zoospores. Cook and Schwartz (1929) 'contended that S. radicalis
is distinct from L. junci.
Matz described a dry top disease of sugar cane as due to Plasmo-
diophora vascularum in 1920 (J. Dept. Agn Puerto Rico 4:41). M. T.
Cook (1929) transferred the causal organism to. Ligniera, calling it
L. vascularum (Matz) M. T. Cook. W. R. I. Cook studied its life
history (1932). On the basis of his observations, he decided that dry
EiG. 4.—Polymyxa graminia: A, zoospores on root hairs; B, young sporangium

in root hair; C, maturing, mature and evacuated sporangia in root tissues show-
ing zoospore formation and the evacuation! tubes (after Ledingham).
top of sugar cane was caused by two distinct organisms which should
be considered amoeboid protozoans. He thereforfr-etected a new genus,
Amoebosponis, and designated the two species as A, ifascularum and
A. saccharinum. M. T. Cook (1937) later rejected'the findings of
W. R. I. Cook and agreed with Palm and Burk that the fungus prob-
ably belonged in Sorosphaera.
Karling (1942,6) excluded Plasmodiophora (Ligniera) vascularum
from the Plasmodiophorales. '
Range: Canada.
Reference: Karling (1942,&); Ledingham (1933, 1939); Sparrow (1943).
Pythium Root Rot and Seed Rot Diseases

The soil-borne species of Pythium cause severe injury to cereals
and grasses throughout the world. Injury is of five types:
f /'
PHYCOMYCETES 17
1. Seed decay, before emergence. Such injury is caused by Pythium
debaryanum Hesse, P. ultimum Trow and P. irregulare Buisman. Seeds
decayed through the action of Pythium spp. are injured, in most in-
stances, just as they start to develop roots, that is, as they start to
sprout.
2. Root necrosis. Decay of roots after emergence is common and is
caused by a number of species, including those causing seed rot.
3. Daiivping-off. This is a condition wherein the seedlings collapse
from a soft stem rot and root rot soon after emergence. P. debaryanum
and P. ultimum are two common causes of damping-off, especially in
nongrass hosts. Damping-off in the Gramineae is not nearly as common
as the slower manifestation, referred to above as root necrosis.
4. Root browning, or browning root rot, in final stage called seedling
blight. P. graminicola Subrm. and related species cause this condition
which differs somewhat from root necrosis, caused by P. debaryanum,
and other organisms. Root browning is characterized by a relatively
firm, brown rot of the roots. Sometimes only short segments of roots
are browned; more often the entire system is so affected that death by
blighting occurs about six weeks after seeding. The plants then shrivel
and are blown away by winds and storms, leaving thin stands which
are partially replaced by weeds. Root browning also develops on older
plants of perennial grasses, resulting in premature death or "going-out"
of grasses in the plains country of the United States and Canada. Root
browning causes stunting of sorghum, wheat, oats, barley, and some-
times corn.
5. Stalk rot of corn and cane crops. This is due to such warm-
temperature fungi as P. aphanidermatum (Edson) Fitzpatrick and P.
butleri Subrm. However, a stalk rot of corn due in part to P. grami-
nicola and associated fungi has also been noted at Mandan, North
Dakota.
Isolation of Pythium spp.—In isolating species of Pythium from
roots of Gramineae, surface sterilizing with chemicals is undesirable
because the delicate mycelium of Pythium is readily killed by such
materials. One of the standard methods by which Pythium spp. are
readily isolated employs running water as a washing medium, followed,
after five or six hours,'by plating roots and crown parts on nonnutrient
water agar. After two or three days of incubation at cool room tem-
peratures or in the refrigerator at 40° F., the developing mycelia are
transferred to nutrient agar. Other practices include variations of this
technique and modern improvements in direct isolations from adjacent
soil. Propylene oxide as a sterilizing agent of organic media has proved
to have some merit but has not been used widely. The most generally
employed medium for'routine growth of Pythium is cornmeal agar.
Some isolates which fail to sporulate on this medium may be induced
to do so on grated carrot agar (Johann, 1928,6). Zoospore formation is
favored by placing the mycelium in distilled water with sterilized, ex-
cised wheat root tips (Vanterpool, 1938), hemp seeds (Mathews, 1931),
or carrot gratings. The addition of sterilized, excised roots of wheat to
the surface of water agar or nutrient agar often results in the produc-
tion of oogonia adjacent to or on the excised roots after the mycelium
has spread over them. Many isolates, however, never produce oogonia
while some produce scarcely any sporangia or even gemmae.
Many ways have been suggested for clearing cultures of Pythium
from the persistent presence of associated bacteria (Machacek, 1934;
Meredith, 1940). The method suggested by Sleeth (1945) has been
satisfactory and in teaching isolation techniques in plant pathology
laboratories we have used it with good results.
In the northern United States, particularly in the areas with com-
paratively late springs, some cultures may be obtained in late February
in open weather, but usually root necrosis and seed decay fungi do not
appear in isolates until April, May, and early June. Root browning
rot, seedling rot', and blight fungi become dominant in June but some-
times are prevalent until July. In the Pacific Northwest areas with
early spring weather, root browning appears at times in February.
However, in the adjacent Palouse,region these fungi appear later.
Root browning disappears in iummer, except in warm-temperature
crops such as proso millet [Panicum miHaceum L.) and various grass
weeds. It also develops on fall^seeded grasses and, less often, on cereals
during warm September weather. In many areas of the Pacific North-
west, after early spring has passed, the relatively dry summers prevent
root browning from being serious.
There have been some papers (Peyronel, 1922) on the association of
pythiaceous fungi with mycorrhizal condition in Gramineae but the
evidence is inconclusive.
A key has. been prepared to aid in determining isolates of Pythium
spp. from cereals and grasses. The write^ has modified his key some-
what from that of Mathews (1931)' in that he has not attempted to
separate the lobulate sporangial group (e.g., P. graminicola) and the
sphaerosporangial group (e.g., P. debaryanum) (cf. Sparrow, 1931).
Rather, he has employed, wherever possible, means of identification
which are mordreadily observable to the rouJihe-wQrker. He depends,
to some extent, on the appearance of the cultures on cornmeal agar
and cautions the reader to use Mathews' key, and any other available
as well. Usually P. graminicola produces a flat, wet, "pebbly" growth
(due to the numerous oogonia) as contrasted with the cottony aerial .
growth of the sphaerosporangial forms. Final determination of the
species must often rely on tedious studies using excised roots in dis-
tilled water and other techniques needed to induce fruiting and zoospore
discharge.
KEYS TO SPECIES OP PYTHIUM ISOLATED EROia GRAMINEAE I N NORTH AMERICA
A. Mycelial growth on cornmeal agar typically cottony, although sometimes

scanty, sporangia, if present not lobulate (except P. monospermum, P. but-
leri and P. wphanidermatum)
PHYCOMYCETES 19
B. Sporangia filamentous or lobulate, oospores filling oogonium
C. Antheridia not swollen nor broadly appressed
P. monospermwm Prings. (rare on Gramineae)
CC. Antheridia swollen, broadly appressed
D. Oogonia averaging 22 M diam.
P. aphanidermatum (Edson) Fitzpatrick
DD. Oogonia averaging 27 ," diam.
P. hutleri Subrm.
BB. Sporangia," if present, not filamentous but spherical (sphaerosporangial
forms)
. C. Oogonia seldom or never present
D. Oogonia' never present; mycelial growth moderately profuse
, P. intermedium deBy. (saprophytic) ''
DD. Oogonia rarely present; mycelial growth profuse, "conidia" (gem-
mae) abundant, large, average 36 /x
P. splendens Braun (rare on Gramineae)
. CC. Oogonia commonly present, at least on grated carrot agar or excised
wheat roots
D. Outer oogonial wall irregular, at least in part
E. Oogonial wall briefly rostrate, oogonia often in chains
P. rostratum Butler (rare on Gramineae, probably sap-
rophytic)
EE. Oogonial wall with several denticulate extensions but not' typi-
cally spiny
P. irregnlare Buisman (common on necrotic roots)
EEE. Oogonial wall spiny
F. Spines narrow, quill-like
P. megalacanthum deBy. (parasite on Pythium spp.)
FF. Spines, stout, few
P. artotrogus (Mont.) deBy. var. macracanthum Si-
deris (rare on Gramineae)
FFF. Oogonia stoutly warty-spiny, 17-35 M
P. oligandrum Drechsl.
DD. Outer oogonial wall not irregular or scarcely so
E. Antheridiurh always originating from the same hypha as the
oogonium
F. Antheridium entering oogonium through oogonial stalk
- - P. hypogynum Middleton
FF. Antheridium usually not as above
P. vltimum Trow (parasite on many hosts, causes seed
rot, root necrosis, damping-off)
EE. Antheridia frequently originating' at points distaifT from the
oogonia
F. Oogonia with a thick shell-like exterior
P. ostracodes Drechsl. (reported from Texas)
FF. Oogonia not as above
G. Antheridia widely appressed to the oogonia, widely flattened
P. vexans deBy. (saprophytic on Gramineae)
GG. Antheridia not as above, oogonia commoner, mycehal
growth strong
P,. deharyanum Hesse (parasitic on many hosts, caus-
ing seed decay, root necrosis) (P. nagaii has only one
antheridium, P. deharyanum has several)
A.A. Mycelial growth on cornmeal agar is fiat, "pebbly," aerial mycelium
scanty, lobulate sporangia formed [
B. dogonia averaging about 28 M diam., mycehal growth not particularly
scanty '
P. graminicola Subrm. (causes root browning) (P.
volutum Vant. tends to have antheridia coiled about
the oogonial stalk)
BB. Oogonia averaging about 24 /* diam., growth, scanty
P. tardicrescens Vant.
BBB. Oogonia averaging 17-20 M diam., often wrapped by hyphae, growth
scanty
P. periilum Drechsl. (weak parasite, rare in northern
U. S.)
AAA. MyceUal growth scanty, filamentous sporangia formed
P. dissotocum Drechsl.
/
Pythium aphanidermatum (Edson) Fitzpatrick—Sugar Cane Rot
Mycelium well developed on cornmeal agar and boiled Swiss chard
or similar material. Aerial mycelium moderately profuse on cornmeal
agar; hyphae branched, 2-8 /u, diam.; sporangia filamentous, composed
of a moderately lobulated, inflated mass of branches cut off by cross
walls from the remainder of the mycelium, with long or short discharge
tubes. Zoospores averaging 9 /J,, reniform, difficult to induce on arti-
ficial substrate. Oogonia smooth, spherical, terminal, averaging 22 /x
diam.; oospores spherical, smooth, not filling/the oogonium, 12-25 ju,
diam., averaging about 17.5 /i. Broadly appraised, moderately swollen
antheridia frequently produced immediately adjacent to oogonia but
often an adjacent antheridium fertilizes some more distant oogonium.
On: Saccharum officinarum L. (and other species of the sugar cane complex),
La. There is-also mention made of this fungus on sugar cane, in Hawaii but
in this case the fungus was probably P. graminicola (see Mathews, 1931,
p. 58). Middleton (1943) listed Agrostis stolonifera L. as a host, citing
Monteith (1933).
Drechsler (1934,a) treated P. aphanidermatum and P. butleri
Subrm. as distinct species although Mathews (1931) had earlier placed
P. butleri in synonymy with the former. According to Drechsler, of one
hundred isolates with lobulate sporangia with broadly appressed an-
theridia five were P. aphanidermatum. Three of these were obtained
from sugar cane in Louisiana. He states, "the generally smaller zoospo-
rangia here (of P. aphanidermMtum) are derived from elements much
less extensive (than are those of P. butleri), less swollen and less elabo-
rately branched. The zoospores are smaller, their diameter, after en-
cystment, averaging about 9 /* instead of 11 /x; and their production is
PHYCOMYCETES • 21
difficult to induce on artificial substrata. The sexual structures are
smaller, the oogonial diameter averaging about 22 /* instead of 27 /i,
the oospore diameter about 17.5 /i instead of about 22.5 /A. The smaller
antheridium is much more frequently produced immediately adjacent
to an oogonium, though here also an adjacent antheridium generally
fertilizes some legs directly connected oogonium." He concluded that
the smaller organism was P. aphanidermatum, the larger, P. butleri.
On this basis it is necessary to include both P. aphanidermatum and
P. butleri in this text. Both species are apparently favored by high
temperatures and are essentially habitants of warm temperature and
subtropic zones. As yet, P. aphanidermatum does not appear to be of
very great importance on Gramineae.
Range: Widely distributed on nongrass hosts in the tropics, scattered on
sugar cane in the same general areas. On Festuca in Cyprus, Secale in
China, corn in the Philippines, sorghum in Hawaii.
Re-ferences: Drechsler (1934,a); Mathews (1931).'
Pythium artotrogus (Mont.) deBy. var. macracanthum Sideris—Rot

Mycelium intra- and extracellular, in culture media (such as
papaya) exhibiting a strong aerial development; hyphae branching,
3-6 [x diam., laterals occasionally-with allantoid bodies; asexual repro-
duction mostly metasporic; ."conidia" (gemmae) spherical to sub-
spherical, 20-30 IX diam., mostly intercalary; oogonia spiny with large,
somewhat bluntly tapered spines, terminal or very often intercalary,
20-30 fi diam., produced intra- and extracellularly; antheridia unicel-
lular, cylindrical to club-shaped, tapering gradually toward the de-
limiting septum, 10-14 /x long, 5-7 /x wide, hypogynous or epigynous,
mostly one in, relation to a single oogonium; oospores echinulate, 15-25
fx diam., less spines, or 30-45 /A diam. including spines, cell wall 1.5-
2.0 IX thick, spines 7.5-10 /* long, 1.0-1.5 /* wide. Said to differ from the
species proper in having some epigynous antheridia, in the greater
length of the spines of its oospores, and in the greater amount of aerial
mycelium produced on culture media.
On: Panicum purpurascens Raddi, Hawaii—Saccharum officinarum L., Ha-
waii. Apparently neither host is known to be attacked in nature; they
were infected artificially.
Sideris (1932) isolated the fungus originally from pineapple roots
in Hawaii but obtained infection of the roots of sugar cane and grasses.
This fungus is characterized by coarsely spiny oogonia which remind
one somewhat of a "cedar apple" in full fruit. Since this fungus is
capable of attacking Gramineae it is included here.
Range: Hawaii.
References: Parris (1940); Sideris (1932).
Pythium butleri Subrm.—Stalk Rot of jCorn, Turf Spot

Mycelium well developed on cornmeal agar and on boiled Swiss
' chard in distilled water. Aerial mycelium moderately abundant on corn-
meal agar. Hyphae branched, 2-8 /* diam.; sporangia filamentous, com-
posed of a mass of lobulate,-inflated branches, cut off by cross walls
from the remainder of the mycelium, with a long or short discharge
tube. Zoospores 15-40 or more to a vesicle, reniform, bicSliate, 9 X 12-
14 iji. Oogonia terminal, smooth, spherical, 20-34 pi. (average 27 fi.) diam.
Oospores spherical, smooth, not filling the oogonium, 12-28 fx (average
22.5 /x) diam.; in the mature stage the typical structure consists of a
heavy wall and a central reserve globule surrounded by a granular layer
of protoplasm in which a small refractive body is embedded. Antheridia
stalked or intercalary, hypogynal or diclinous, usually one, rarely two
to an oogonium, characteristically broadly appressed and swollen.
On: Agrostis palustris Huds., D. C, 111., Mo., Penn,, Va.—Zea mays L., Tex,,
Va. Listed on sugar cane from Hawaii./
Elliott (1943) reported a stalk rot'of certain inbred lines of corn at
Arlington (Farm), Virginia. The stalks in late July and August broke
over with a rot just above the ground line. The roots and tops remained
healthy, often recovering after hot weather passed. Elliott stated that
"the outer rind of the corn stalk, the epidermis, the lignified cells of the
hypodermis, and inner parenchyma cells are softened, disorganized and
destroyed, leaving only the separate browned strands of vascular
bundles, which cannot support the plant."
P. biitleri, according to Drechsler (1934,a—see P. aphanidermatum,
above), is a distinct species. Elliott determine^ that the material she
had was P. butleri. Isolates from the corn stalks caused typical soft
rots of cucumber and summer squash. Roots of corn were not injured,
the injury being confined to the stalk base. To be actively parasitic
on corn, the fungus requires high summer temperatures accompanied
by high humidity. It quickly subsides after hot wea^Br-has passed.
Monteith (1933) described a disease of creeping bent {A. palustris)
caused by P. butleri. Affected patches up to 4 inches in diameter first
appeared blackened and water-soaked, later becoming shriveled and
brown. They often coalesced and destroyed large areas of tuff. The
fungus can attack plants over a wide temperature range, the optimum
being 35° C. Partial control was obtained by'spraying the grass with
mercuric chloride solution.
At times a stalk rot of field corn is common at Mandan, North
Dakota, especially in some hybrid dent varieties. It is' also present in
selections of Northwestern Dent. The plants-dfe prematurely, the ears
become flaccid, and the kernels are undersized and pale in color. The
bases of the culms are somewhat discolored and sometimes partially
collapsed. Many fungi have been isolated from this material, including
P. graminicola Subrm.
PHYCOMYCETES 23
Drechsler (1947) obtained germination of the oospores of P. butleri
by transferring 60-day-old oospores to a shallow layer of distilled
water. The thick oospore wall is reduced to a thin shell by reabsorption
of the thicker, inner layer. The granular contents, on being discharged
into a vesicle at the tip of a simple, or sometimes branched, evacuation
tube, 40-225 X 2.5-5.0 /A diam., break into approximately 10 motile
zoospores.
Range: Southern United States and northward at times; India; probably
widely distributed on various hosts in the tropics and subtropics.
References: Drechsler (1934,a; 1947); Elliott (1943); Monteith (1933). '
Pythium debaryanum Hesse—Seed Rot, Damping-Off and

Root Necrosis
• Mycelium well developed in host tissues and on nutrient agar.
Hyphae branched, usually about 5 ^ diam., often septate in old cul-
tures. Sporangia few, spherical to oval, terminal or intercalary. Conidia
(gemmae) numerous, same size as the sporangia, germinating by a
germ tube soon after being formed while still attached to the hyphae
or after falling off, and after a long or short rest period. Oogonia smooth,
terminal or intercalary, uusally spherical, 15-25 /x diam. Oospores
smooth, not filling the oogonium, 10-18 ii diam., germinating directly
after a rest of several months. Antheridia 1-6 to an oogonium, diclinous,
androgynous and hj^pogynal, when androgynous often arising at some
distance from the oogonium.
On: Aegilops cylindrica Host., Wash.—A. triuncialis L., N. Dak.—Agwpyron
caninum ,(L.) Beauv., N. Dak.—A. ciliare (Trin.) Franch, N. Dak.—A.
, cristatum.' (L.) Gaertn., Ida., Minn., Mont., Nebr., N. Dak., Oreg., S. Dak.,
Wash., Wyo.—A. desertorum (Fisch.) Schult., N. Dak., S. Dak.—A. inerme
(Scribn. and Sm.) Rydb., Oreg., Wash.—A. intermedium (Host.) Beauv.,
N. Dak., S. Dak.j-^Vash.—A. michnoi Roshev, N. Dak., S. Dak.—A. mon-
golicum Keng, N. Dak.—A. repens (L.) Beauv., Ida., Minn., Nebr., N.
Dak., Oreg., S. Dak., Wash., Wyo.—A. riparium Scribn. and Sm., N. Dak.,
Wash.—A. semicostatum (Steud.) Nees, Wash.-Â. sibiricum (Willd.)
Beauv., Minn^ Nebr., N. Dak., Oreg., S. Dak., Wash., Wyo.—A. smithii
Rydb., Mont., Nebr., N. Dak.—A. subsecundum (Lk.) Hitchc, Wyo.— -
A. trachycaulum (Lk.) Malte, Mont., Nebr., N. Dak., Wash.—A. tri-
chophorum (Lk.) Richt., N. Dak.—Agrostis alba L., .Nebr., N. Dak.—
A. scabra Willd., N. Dak.—Andropogon furcatus Muhl., Nebr., N. Dak.,
S. Dak., Wyo.—A. hallii Hack;, N. Dak., S. Dak.—A. scoparius Michx.,
N. Dak., S. Dak.—Anhenatherum elatius (L.) Beauv., N. Dak., Wash.—
Avena byzantina K. Koch, Iowa, Minn., Nebr., N. Dak., Oreg.—A. fatua
L., N. Dak., Oreg., Wash., Wyo.—A. nuda L., N. Dak.—A. sativa L., Colo.,'
Iowa, Minn., Mont., Nebr., N. Dak., Oreg., S. Dak., Wash.—Beckmannia
syzigachne (Steud.) Fern., N. Dak.—Boutelona curtipendula (Michx.)
Torr., N. Dak.—B. gracilis (H. B. K.) Lag., Mont., N. Dak., S. Dak.—
Bromus arvensis L., N. Dak.—B. carinatus Hook, and Arn., N. Dak.,
Wash., Wyo.—B. erectm Huds., Wash.—B. inermis Leyss., Minn., Nebr.,
N. Dak., Oreg., S. Dak., Wash., Wyo,—B. japonicus Thunb., Wash.—B.
madritensis L., Wash.—B. tectorum L., Ida., ISfebr., Wyo.—Calamovilfa
longifolia (Hook.) Scribn., Nebr., S. Dak.—Cynddon dactylon (L.) Pers.,
N. Dak.—Dactylis glomerata L., S. Dak., "Wa.&h.-r:Distichlis stricta (Torr.)
Rydb., S. Dak.—Echinochloa crusgalli (L.) Beauy., Minn:, N. Dak., Wyo.
—Elymus canadensis L., N. Dak., S. Dak., Waih., Wyo.—E. dahuricvs
Turcz., N. Dak., Wash.—E. ex,celsus Turcz., N. Dak.—E. glaixus BuckL,
Wash.—E. interruptus BuckL, N. Dak.—E. ju7iceus îsch., N. Dak.—
E. macounii Vasey (hybrid), N. Dak.—'E. sibiricus L., N. Dak., Wash.—
E. virginicus L., N. Dak.—Festuca elatior L., N. Dak., Wash., Wyo.—
F. elatior var. arundinacea (Schreb.) Wimm., Mont., Nebr., N. Dak.—F.
megalura Nutt., Oreg.—F. octoflora Walt., Nebr., N. Dak,, Wash.—F.
ovina L., Wash.—F. pacifica Piper, Wash.—F. rubra L., N. Dak.—F. rubra
var. commutata Gaud., .N. Dak.—Hordeum brevisubidatum (Trin.) Lk.,
N. Dak.—H. bulbosum L., N. Dak., Wash.—H. distichon L., Minn., N.
Dak., Oreg., S. Dak., Wyo.—H. jubatum L., Minn., N. Dak., S. Dak.—
H. vulgare L., Colo., Iowa, Manit., Minn., Nebr.,"N. Dak., Oreg., Sask.,
S. Dak., Wash.—Koeleria cristata (L.) 'Pers., Wyo.—Lolium remotum
Schrank, Wash.—Oryzopsis hymenoides/ (Roem. and Schult.) Rick., N.
Dak., S. Dak., Wash.—0. mtKacea (L.) Benth. and Hook., N. Dak.—Pani-
cum capillare L., Minn., N. Dak.—F. miliaceum L., Minn., N. Dak., Greg.,
Sask., S. Dak., Wash.—P. virgatum L., Nebr., N. Dak.—Phalaris arundi-
nacea L., N. Dak.—Phleum pratense L., Minn., Wash.—Phragmites com-
munis Trin., N. Dak.—Poa ampla Merr., N. Dak., Wash., Wyo.—P. bidb-
osa L., N. Dak.—P. nevadensis Vasey, Wash.—P. palustiis L., Minn., N.
Dak., S. Dak.—P. pratensis L., Minn., N. Dak., S. Dak., Wash.—P.
secunda Presl, N. Dak., Wash.—Saccharum officinarum L., Hawaii—
Schedonnardvs panicidatus (Nutt.) Trel., Mont., N. Dak., Wyo.—Secale
cereale L., Mont., Nebr., N. Dak., S. Dak., Wash.—S. cereale X S. mon-
tanum Guss., Wash.—S. montanum Guss., Wash'.—Setaria. italica (L.)
Beauv., N. Dak., Sask.—S. lutescens (Weige;!) FJT. Hubb, N. Dak., S.
Dak.—S. viridis (L.) Beauv., Minn., N. Dak., Sask., S. Dak.—Sitanion
hystrix (Nutt.) J. G. Smitli, Wyo.—Sorghum vulgare Pers,, Minn., Nebr.,
N. Dak., S. Dak.—S. vulgare var. sudanense (Piper) Hitchc, Nebr., N.
Dak., S. Dak.—Sporoboltts asper (Michx.) Kunth., Mogt.—Stipa comata
Trin. and Rupr., Sask.i, Wyo.—S. sibirica (L.) Lam., N. Dak:—^. spartea
Trin., S, Dak,—S, viridula Trin., Minn., Mont., N. Dak., S. TD&k.—Triti-
cum aestivum L., Ariz., Cahf., Colo., Iowa, Kans., Manit., Minn., Mo.,
Mont., Nebr., N. Dak., N. Mex,, Oreg., Sask.,..S. Dak., Utah, Wash., Wyo.
—T. dicoccum Schrank, N. Dak.—T. durum Desf., Minn., N. Dak,, S.
Dak., Wash.—T. timopheevii Zhuk., N. Dak.—Zed mays L., Hawaii, Iowa,
Minn., Mont., N. Dak., Okla., Ont., S. Dak., Wyo.
The host range of this common cause of root necrosis and damping-
off is obviously very extensive. Most of the above citations arc based
on isolations made by the writer while at Mandan, North Dakota.
Many of the citations for North Dakota, South Dakota, Minnesota,
Nebraska, Wyoming, and Montana have not been previously reported
although a number of them -were included in data given to Middleton
for inclusion in his t^xt (1943). Current învestigations at Pullman,
PHYCOMYCETES 25
Washington, indicate that P. debaryanum is parasitic on the roots of
many grass seedlings in the Palouse prairie region.
P. debaryanum has been known as a parasite of cereals and grasses
for many years but most workers have forgotten that fact. In the
original discussion of this species Hesse (1874) listed Panicum mili-
aceum L. (proso millet) and corn (Zea mays L.) as original hosts.
Eriksson (1912) and also Gram and Rostrup (1922) reported it on
barley, and there are a number of citations on cane crops, including
corn (Ho, 1944) and oats (Welch, 1945). In the northern Great Plains,
in adjacent areas, and in the Pacific Northwest, as mentioned, P. de-
baryanum and related species (i.e., P. ultimum Trow, P. irregulare
Buis.) are associated with root necrosis of many grasses and cereals.
Seed rot and sometimes damping-off occur as a consequence of the
parasitic activity of these fungi, notably in wet soil in the spring
(Sprague, 1944,6). Sometimes cereals as well as grasses are severely
injured when growing in low wet ground or when moldy seed is planted
in heavy soil. The injury in most cases to both grasses and cereals
occurs just as the seed sprouts. The rootlets are unable to emerge from
the seed. In the greenhouse, under artificial conditions, some grasses
have the roots reduced to yellow or brown stubs even after emergence.
There are various strains of P. debaryanum, some of which will
attack certain hosts more readily than others. Some will attack certain
cereals, such as wheat, barley, and oats more severely than other
cereals; some strains from grasses will attack many species of grasses
but are only mildly parasitic on nongrass crops; in the case of other
strains the reverse is true. In general, however, the host range is so
great that crop rotation offering control of this disease complex would
be very difficult to plan. Since legumes, notably alfalfa, are very sus-
ceptible to P. debaryanum (Buchholtz, 1942,6) and the use of summer
fallow seems to favor its increase, the control of P. debaryanum be-
comes more difficult than that of P. graminicola, which is largely con-
fined to the grass family. However, P. debaryanum is seldom an im-
portant parasite on wheat and usually does not cause the complete
destruction of grasses such as may result from attacks by P. gramini-
cola manifested as seedling blight. Buchholtz (1942,a) has shown that
seedling blight' symptoms are produced by P. graminicola. We have
found that recovery from infection by P. debaryanum is very rapid
while recovery from infection by P. graminicola is very slow. This
difference is particularly noticeable when seedling blight, which nor-
mally occurs in June, is severe. P. debaryanum is, therefore, mainly
important as a seed rot but can cause some root necrosis in Gramineae.
Oats are somewhat more susceptible to P. debaryanum, especially dur-
ing cool, wet weather (Welch, 1940; Gram and Ilostrup, 1922; Beau-
mont, 1927; Buchholtz, Melhus, Welch, and Murphy, 1947). Welch
(1945) found that P. debaryanum was the principal cause of root ne-
crosis of oats in Iowa. The fungus was favored by a temperature below
25° C., the optimum being between 8° and 15° C. The fungus was iso-
lated during May and early June. Of 232 varieties which Welch tested,
Coast Black, Black Algerian, Early Red Rustproof, Red Algerian,
Ruakura, and Flughafer were the most resistant, although in all these
the tolerance was only moderate. From observations made by the writer
some years ago, the indications are that/ some' of these varieties or
others very similar to them were certainly not resistant in the acid soil
of coastal Oregon.
P. debaryanum, P. irregulare, and P. ultimum belong to the sphaero-
sporangial group of Pythium spp,, which includes the species with
spherical or lemon-shaped sporangia. P. graminicola, on the other hand,
produces irregular, lobed or lobulate swollen bodies of mycelia, from
which stalked, spherical sporangia later develop. P. debaryanum is by
far the most common of the sphaerosporangial species of Gramineae in
North Dakota and surrounding regions.
P. debaryanum is distinguished from its occasional associates, P.
irregulare and P. ultimum, by several characteristics. P. irregulare has
irregular oogonial walls with irregular projections. P. debaryanum pro-
duces zoospores from the sporangia while P. ultimum generally germi-
nates directly by sending out germ tubes from the sporangia. In addi-
tion P. -ultimum has antheridia which arise from the oogonial stalk,
while those of P. debaryanum, which are frequently several in number,
are likely to be produced from distinct, distant hyphae. Van Luijk
(1934) questions if the recognized differences are of critical value in
distinguishing these fungi from one another.
P. vexans de Bary is another sphaerosporangial species which 'is
sometimes encountered on grasses and cereals. It is more or less readily
distinguished from the others by its broadly appressed antheridial at-
tachment. It has a tendency to remain sterile in/culture. The cultures
are generally less cottony than in P. debaryanum and are nonparasitic
on Gramineae in all trials which we have conducted to date. P. vexans
is more likely to be isolated very early in the season from the rhizo-
sphere of plants growing in cold, wet soil. I
P. hypogynum Middleton (1941) is possibly a variant.oTP. ultimum
but will be discussed separately here. This fungus is characterized by
an antheridium which grows through the oogonial stalk into the
oogonium. The writer isolated this fungus from wheat growing in low,
wet soil at Carrington, North Dakota, and from Avena sativa at Minne-
chaduza Creek, Nebraska. / /
P. iwayamai Ito (Ito and Tokunaga, 1935) was said to cause a snow
rot disease of cereals in Japan (Iwayama, 1933). This fungus has not
been seen in the United States, or at least it has not yet been recog-
nized. Iwayama (1933) reported that the disease he noted was particu-
larly injurious on early-sown, heavily manured^crops on poorly drained
soils under a thick snow panoply. The fungus grew well on potato and
barley decoction agars, on which it was characterized by pale yellow,
granular hyphae, 2.9-8.9 ju. diam. (average 6.6 /x), becoming septate
when mature; spherical, ellipsoid, oval, lemon-shaped or irregular, thin-
PHYCOMYCETES 27
walled, smooth, pale olive-yellow, terminal sporangia, 28-48 X 26-44 /*
on the host, 33-39 /A diam. in culture; flask-shaped intercalary oogonia,
with a single spherical, pale olive-yellow oospore, 19-24 /* (average
21 /x) diam.; and clavate antheridia arising from the same hypha as
the oogonium or from another one. In culture the fungus grew best at
a temperature between 15° C. and 18° C. Inoculation experiments con-
sistently gave positive results on cereal plants under snow.
Biraghi (1936) reported P. polymorphon Sideris (1932) from Italy
on cereals. He may have been dealing with P. irregulare, but the iden-
tity of his fungus is not certain. In Italy, de Paolis (1931) caused stunt-
ing of wheat seedlings by watering planted seed with a water extract
of a culture of Pythium isolated from wheat near Padua.
Range: General on Gramineae (see Middleton, 1943).
References: Beaumont (1927); Biraghi (1936); Buchholtz (1942,a,6);
- Buchholtz, Melhus, Welch, and Murphy (1947); Butler (1913,a); Car-
penter (1919); Dreohsler (1927,a); Eddins (1930,a); Eriksson (1912);
Gram and Rostrup (1922); Granfield, Lefebvre, and Metzger (1935);
Hesse (1874); Ho (1944); Ho and Melhus (1940); Hoenk (1932); Ito
and Tokunaga (1935); Iwayama (1933); Mason and Powell (1947);
Mathews (1931, PI. 17, Fig. 17-22,24,25); McLaughlin et al. (1943);
Melhus et al. (1940); Meredith (1938, 1940); Middleton (1941; 1943);
Miura (1920); de Paolis (1931); Petri (1930); Ramakrishnan (1941);
Richardson (1942); Sideris (1931, 1932); Sprague (1943,a; 1944,6,c);
Spyshneva (1895, 1897); Summers (1948); Van Luijk (1934, 1938);
Vanterpool (1942); Welch (1940,-1942, 1945).
Pythium dissotocum Drechsl.—Weak Root Rot of Sugar Cane

P. oryzae Ito and Tokunaga
Aerial mycelium scanty on all but the richer media and then not
plentiful, submerged mycelium somewhat lustrous with a tendency for
the mycelium to radiate from the center of the culture in nearly parallel
strands. Spoi'angia filamentous, resembling those of P. monospermum
Prings., sometimes entire mycelial structure will produce zoospores on
irrigation with sterile water. The spherical, sporangial vesicles or other-
wise differentiated sporangia may contain as many as 125 zoospores.
Zoospores given to itinerant swimming, then encysting and germinating,
sometimes by a germ tube. Oogonia terminal or subterminal, but often
intercalary, 12-32 /i, commonly 17-24 /*; oospores ll-27>" diam., com-
monly 16-20 fx,; anthridia 1-3, inflated, crook-neck in type, borne more
or less indiscriminatelyôn the same hyphae producing oogonia, or on
adjacent or moderately distant hyphae.
On: Saccharum officinarum L., La.
Under average field conditions this species seems to be considerably
less important and less prevalent than P. graminicola in Louisiana. The
fungus has also been isolated from roots of peas, beets, spinach, and
Pilea. I t resembles P. monospermum in some ways, but Drechsler illus-
trates the oogonia and associated structures as being definitely more
rugged. The sexual spores also are described as being somewhat larger
than those of P. monospermum.
Petri (1930) discussed a root rot and foot rot (basal culm rot) of
wheat occurring in northern Italy. Among other fungi, a species of
Pythium was present in the lesions. On carrot agar (Petri, 1930) the
mycelium formed a snow-white, superficial, cottony layer of bundles of
regularly branched, aerial hyphae showing false dichotomy, often un-
dulating at the extremities, occasionally thickened at the point of
branching, and measuring 2-5.5 ji diam. The very numerous, apical or
intercalary oogonia measured 21-26 /x, each being fertilized by a single
antheridium which arose from the same oogonial hypha. Sometimes the
antheridia were septate at their base. The smooth oospores filled the
oogonia (monosporic) and were 16-24 /i diam. Lobulate "presporangia"
were present. While Petri placed this fungus in the P. gracile group,
the description indicates that it might be close to P. dissotocum, since
the oogonia and oospores are somewhat large for P. monospermum.
The description of the culture, however,'is not the same in all respects
as Drechsler gives for P. dissotocum. Petri's fungus appeared to have
somewhat whiter and less submerged mycelium in pure culture than
that of Drechsler. However anyone who has worked with the group
recognizes that there is considerable variance within species. Curzi
(1929) described the symptoms and the fungus morphology of a simi-
lar or identical form on corn at Padua the year previous to the work
by Petri.
Ito and Tokunaga (1935) describe P. diclinum [P. gracile Schenk
var. b (eta) Butler] on rice from Japan. This species may also be very
similar to P. dissotocum. It has filiform sporangia;; sphaeroidal, smooth
oogonia, sometimes with a sharp point near the apex, 18-25.2 /j, diam.,
each with 1-3, mostly 2, diclinous, ovoid or clavate, often curved an-
theridia and solitary, spherical or broadly ellipsoid oospores, 14.4-
22.8 /x diam.
Range: United States, Japan. ,
References: Drechsler (1930, 1940); Ito and Tokunaga (1933); Middle-
ton (1943); Rands and Depp (1938).
Pythium graminiccla Subrm.—Root Browning of Cereals,

Seedling Blight of Grasses
P. arrhenomanes Drechsl.
P. aristosporum Vanterpool
Mycelium well developed on cornmeal agar. Hyphae irregularly
branched, 2.5-7.2 jn diam. Sporangia consisting^ ol irregular inflated ele-
ments, cut off from the remainder of mycelium by cross walls. Dis-
charge tube slender, often long. Zoospores 15 to 40 or more to a vesicle,
reniform, biciUate, encysting as spherical or ellipsoid bodies, 9.6-12 /*
PHYCOMYCETES 29
diam. Conidia (gemmae) spherical, 24-36 /x (usually about 30 fi diam.)
germinating by a germ tube in old cultures. Oospores usually filling
oogonia, spherical, rarely elliptic, 21.6-32 ju. diam. (usually about 28 /x),
each a t m a t u r i t y with a heavy wall and a central reserve globule sur-
rounded b y a granular layer of protoplasm, in which a small, refrac-
tive b o d y is embedded. Antheridia often curved, 1-6, or more, t o a n
oogonium. Antheridial filament often long and irregular, as m a n y as
five antheridia arising from one filament, which in t u r n m a y arise from
the h y p h a bearing the oogonium or from a more distant one (from
M a t h e w s , 1931).
O n : Aegilops triuncialis L., N. Dak.—Agropyron angustiglume Nevski, N .
Dak.—A. ciliare (Trin.) Franch, N. Dak.—A. cristatum (L.) Gaertn.,
Manit., Mont., Nebr., N. Dak., Sask., S. Dak., Wash.—A. dasystachyum
• (Hook.) Scribn., N. Dak., Wash.—A. desertorum (Fisch.) Scliult., Manit.,
Mont., N. Dak., S. Dak.—A. elongatum (Host.) Beauv., Nebr., N. Dak.—
, A. inerme (Scribn. and Sm.) Rydb., N. Dak., W a s h . ^ 4 . intermedium
(Host.) Beauv., N. Dak., S. Dak., Wash.—A. michnoi Roshev, N. Dak.,
S. Dak., Wash.—A. mongolicum Keng, N. Dak.—A. orientale var. lasi-
anthum (Boiss) Boiss, Wash.—A. pendulinum (Nevski) ined., fide Swallen,
{ttoegneria pendidinum Nevski), N. Dak.—A. pubicalum var. pilipes
(Keng) ined., fide Swallen, N. Dak.—A. repens (L.) Beauv., Minn., Nebr.,
N. Dak., Sask., S. Dak., Wash.—A. riparium. Scribn. and Sm., N. Dak.—
A. semicostatum (Steud.) Nees, N. Dak.—A. dbiricum (Willd.) Beauv.,
Minn., Nebr., N. Dak., S. Dak.—A. smithii Rydb., Mont., N. Dak., Sask.,
Wash.—A. spicatum (Pursh) Scribn. and Sm., N. Dak., Wash.—A. trachy-
caulum (Lk.) Malte, N. Dak., Sask., Wash.—A. trichophorum (Ik.)
Richt., N. Dak., Wash.—Agrostis alba L., Sask., Wyo.—Ammophila are-
naria-{L.) Lk., N. Dak.y-Andropogon furcatus Muhl., N. Dak.—A. hallii
Hack., N . Dak.—Aristida purpurea Nutt., Minn.—Arrhenatherum elatius
(L.) Beauv., N. Dak., Wash.—Avena byzantina K. Koch, Iowa, N. Dak.
— A . fatuaL., Alta., N. Dak., Sask., Wyo.—A. nuda L., N. Dak.—A, sativa
L., Alta., Iowa, Minn., Mo., N. Dak., Oreg., Sask., S. Dak., Wyo.—Boute-
loua curtipendida (Michx.) Torr., N . Dak.—B. gracilis (H. B. K.) Lag.,
N. Dak., Wash.—Bromus arvensis L., N. Dak.—B. brizaeformis Fisch. and
Mey., Wash.—B. carinatus Hook, and Arn., N. Dak., Wash., Wyo.—B.
erectus Huds., N. Dak., Wash.—B. inermis Leyss, Iowa, Minn., Nebr.,
N. Dak., Sask,, S. Dak., Wash.—B. japoniciis Thunb., S. Dak.—B. madri-
tensis L., Wash.—B. pumpelliamis Scribn., Wyo.—B. tectorum L., Nebr.,
S. Dak., Wash.—Calamovilfa longifolia (Hook.) Scribn., N. Dak., Wash.
—Cenchrus pauciflorus Benth., Minn., Nebr., N. Dak.—Dactylis glome-
rata L., N. Dak., Wash.—Echinochloa crusgalli (L.) Beauv'., Minn., N .
Dak., S. Dak., Wyo.—Elymus^ canadensis L., N. Dak., Sask., Wash.,
Wyo.—E. condensatus Presl, N. Dak.—E. dahuricus Turcz., N. Dak.—
E. excelsus Turcz., N.'Dak.—E. gigantevs Vahl, N. Dak.—E. glaucus
Buckl., N. Dak.—E. interruptus Buckl., N. Dak.—E. junceus Fisch.,
Nebr., N. Dak., S. Dak.—E. macounii Vasey (hybrid), N. Dak.—E. sibi-
ricus L., N. Dak.—E. virginicus L., N. Dak.—Eragrostis cilianensis (All.)
Lk., N . Dak.—E. curvida (Schrad.) Nees, N . Dak.—Festuca elatior L.,
N. Dak., Sask., Wyo.-^2^. elatior var. arundinacea (Schreb.) Wimm.,
Mont., N. Dak.—F. idahoensis Elmer, Wash.-^F. kingii (S. Wats.) Cas-
sidy, Wyo.—F. octoflora Walt., N, Dak.—Hordeum brevisubulatum
(Trin.) Lk., N. Dak., Wash.—H. distichori t . , Iowa, Minn., Nebr., N.
Dak., Sask., S. Dak., Wash., Wye—.ff. jubatum L., Minn., N. Dak., Sask.,
S. Dak.—H. murinum L., Wash.—H. vulgare ,L., Alta., Calif., Colo., Iowa,
Manit., Minn., Nebr., N . Dak., Sask., S. Dak.,' Wash.; Wyo.—Melica sca-
brosa Trin., N. Dak.—Muhlenbergia racemosa] (Michx.) B. S. P., Minn.,
N. Dak., S. Dak.—Oryza sativa L., Hawaii—Oryzopsis hymenoides (Roem.
and Schult.) Rick., Mont., N. Dak., S. Dak., Wash.—Panicum capillare
L., Minn., N. Dak.—P. miliaceum L., Manit., Minn., N. Dak., Sask., S.
Dak.—P. purpurascens Raddi, Hawaii—P. subvillosum Ashe, Minn.—P.
virgatum L., N. Dak.—Phalaris arundinacea L., Sask.—Phleum. pratense
L., Minn., N. Dak., Sask., Wash.—Phragmites communis Trin., N. Dak.—
Poa compressa L., Sask.—P. juncifolia Scribn., N. Dak.—P. nevadensis
Vasey, N. Dak.—P. palustris L., Mont.—P. pratensis L., N. Dak., Sask.—
P. secunda Presl, Wyo.—Redfieldia flexuosa (Thurb.) Vasey, Nebr., S,
Dak.—Saccharum barberi Jeswiet, La.—S. officinarum L., Hawaii, La.—
S. spontaneum L., La.—Schedonnardus paniculatus (Nutt.) TreL, Mont.,
N. Dak., S. Dak.—Scolochloa festucacea Willd., N. Dak.—Secale cereale
L., Minn., N. Dak., Sask., S. 'Da.k.-rSetaria italica (L.) Beauv., Minn.,
N. Dak., S. D a k . ^ S . lutescens (Weigel) F . -T. Hubb, N. Dak., Sask., S.
Dak.—S. viridis (L.) Beauv., Minn., Mont., Nebr., N. Dak., Sask., S. Dak.
—Sorghastrum nutans (L.) Nash; N.'Dak.—Sorghum vulgare Pers., Calif.,
Colo., Iowa, Kans., Minn., Nebr., N. Mex., N. Dak., Okla., Tex.—S. vul-
gare var. sudanense (Piper) Hitchc, Calif., Iowa, Nebr., N . Dak., S. Dak.
—Sphenopholis obtusata (Michx.) Scribn., N. Dak.—Sporobolus cryptan-
drus (Torr.) A. Gray, S. Dak., Wyo.—Stipa baicdensis Roshev, N. Dak.
—S. comata Trin. and Rupr., Mont., N. Dak.—S. sibirica (L.)' Lam.,
N. Dak.—S. sparfea Trin., N. Dak.—S. viridula Trin., Mont., N. Dak.—
S. vnlliamsii Scribn., N . Dak.—Triticum aestivum L., Alta., Cahf., Colo,,
Hawaii, Iowa, Manit., Minn., Mont., Nebr., N. Dak., Okla., Ont.,' Sask.,
S. Dak., Wash.—T. dicoccum Schrank, jN. Dak.—7". durum Desf., Minn.,
N. Dak.—T. timopheevii Zhuk., N. Dak.—Uniola latifolia Michx., Mich.
—Zea mays L., Colo., D. C , Hawaii, 111., Iowa, Ky., Manit., Minn., Mo.,
N. Dak., Ont., Sask., S. Dak., Wise.
Root browning is characterized by relatively, firm, dark, brown

lesions on the small roots of grains, especially wheat anci'barley, in late
M a y and J u n e in the northern G r e a t Plains and adjacent regions and,
a t comparable times, depending on the latitude, in areas to the north
a n d south of the plains. W h e n severely infected, the plants are pale
green in contrast with healthy ones. T h e fungus is very -yvidespread in
all of the northern plains states, the C a n a d i a n prairie provinces, and
occurs south to Louisiana. I t is also present in the Sacramento Delta
region of California, in H a w a i i , and was recently found in t h e Pacific
Northwest. I t is well known as the cause of a root rot of sorghum,
especially Milo varieties in the central, s t ^ e s . Recent studies (Leukel
a n d Pollack, 1947) indicate t h a t it is not the sole cause, or possibly
n o t the main cause, of the "milo disease," a stunting disease of sorghum
in the latter region (cf. Periconia circinata).
PHYCOMYCETES 31
In parts of the grassland areas of the Dakotas and adjacent states
P. graminicola causes a blight of grass seedlings. During some years
P. graminicola causes almost complete destruction in seedings of crested
wheatgrass, slender wheatgrass {Agropyron trachycaulum (Lk.) Malte)
and smooth brome at Brookings, South Dakota, and at numerous other
places within the area. This seedling blight consists of a general root
browning of the April-seeded grasses which die, shrivel up, and blow
away in late May or June.
Subramaniam (1928) described P. graminicola from India. Drechsler
contends (1936) that his Pythium arrhenomanes (1928,b) is distinct
from P. graminicola because "the close mycelial connection between
oogonium and antheridium, very frequent in P. graminicolum is rare in
P. arrhenomanes; and in parallel cultures the sturdy, more substantial
membranous parts of the sexual apparatus of the former species remain
discernible long after the evanescent antheridial envelopes and support-
ing branches of the latter have become nearly or wholly invisible." On
the basis of this distinction the material in the northern Great Plains
would be P. arrhenomanes [except for that referable to P. aristosporum
Vanterpool, 1938, Fig. 2, PI. X X I I ; 1942; which has the long-lived
traits of P. graminicola in pure culture]. Mathews (1931) and Middle-
ton (1943) both recognize P. graminicola and P. arrhenomanes as dis-
tinct. In recent literature, however, workers are more or less divided
on which fungus they are working with. After endeavoring to keep
these two species and also P.' aristosporum separated, the writer has
finally concluded that they cannot be readily distinguished and, for all
practical purposes, should be considered as one species. While retaining
an open viewpoint on the matter, in this text all are referred to the
earlier name, P. graminicola. Subramaniam (1928) and others used the
ending icolum but, as my colleague Charles Gardner Shaw has pointed
out, the Iowa' State College workers are correct in using the ending
icola. Ainsworth and Bisby (1945, p. 70) state that "the suffix -icola is
indeclinable."
Root browning in Saskatchewan (Vanterpool, 1939; 1940,a,6) is
partially checked by judicious use of fertilizers containing a proper
balance between nitrogen and phosphorus. Hanson (1944,a) has ob-
tained striking results"in the control of root rot at Lamberton, Minne-
sota, by the use of phosphate or phosphate and nitrogen fertilizers on
wheat. The writer has obtained similar control of the root-rot complex
in oats in acid soils in coastal Oregon. While at the Northern Great
Plains Field Station, Mandan, North Dakota, he conducted a number
of detailed trials to determine-the' effect of fertiUzers on the root-rot
•tomplex, which included root browning. The results in this somewhat
arid region were usually conflicting, but ammonium phosphate (16-
20-0) in pellet form showed definite promise when 100 lbs. per acre
were applied. It was especially promising in lighter soils at McCanna,
North Dakota, and in some of the sandy loams at Mandan. Ammonium
nitrate showed residual effect in the year following its application (100
lbs. per acre) b y reducing the incidence of root browning in year-old
stands of crested wheatgrass, smooth brome, (and Intermediate w h e a t -
grass. T h e "going-out" disease of m a t u r e grass stands appears to be
checked by balanced fertilizers. T h e going-out disease, which is widely
known to the rancher, if not to the pathologist, is due to various fungi,
particularly P. graminicola, a t t a c k i n g the sbd-bound roots of older
plants. T h e plants fail to set seeds, the crowns and roots become brown
and rotten, and the plants die prematurely. Such plants can be readily
kicked out of the ground because their root systems are usua41y com-
pletely decayed. B y adding nitrogen and phosphorus fertilizers in mod-
erate quantities, starved plants often are given a new.lease on life and
t h e competing fungi no longer prey heavily on the root systems. While
from a commercial viewpoint fertilizers are not a paying proposition
on dry land ranges, they are practical in wheat lands and in irrigated
pastures, as well as in some fields where grass is grown in rows for seed.
Therefore, the fertilizer studies conducted a t M a n d a n during t h e years
1940-46 should be continued, or duplicated in a similar area, and are
mentioned here for t h a t reason. /
Since P. graminicola, in general, is restricted to Gramineae, r o t a -
tions including nongrass plants ^reduces the injury. Among cereals,
oats are most tolerant. I t is less desirable to include proso millets
(Panicum miliaceum L.) or sorghums in t h e rotations. While fallow
does not appear to increase the a m o u n t of Pythium, it creates the same
effect by starving out other organisms, permitting the Pythium forms
t e m p o r a r y ascendancy. Small-seeded grasses, such as blue grama,
should not be started on fallow if it can be avoided, a t least n o t in the
spring of the year. Crested wheatgrass, however, can be sown on
fallow under most conditions. F a l l seeding is/preferred to spring seed-
ing where conditions permit. E v e n winter seeding of crested wheatgrass
is preferred, if periods of open weather occur, as is sometimes the case
even as far north as N o r t h D a k o t a .
Little resistance or tolerance has been found in susceptible species
of cultivated cereal, crops. F o r instance, Semeniuk^Wallin, and Melhus
(1947) found t h a t in 988 collections of corn only 6 w * e highly re-
sistant to the fungus. Of course, if these maintain their resistance to
widely collected strains of t h e fungus t h a t is a definite step forward.
I n other hosts still less promising results have been noted.
Leukel (1942) controlled root rot in Sorghum by using^chloropicrin
b u t perhaps as indicated b y recent work (Leukel and Pollack, 1947;
Leukel and Johnson, 1948) the fungus inhibited was Perihonia circinata
and not Pythium graminicola. D . A. Cooke (1933), however, has con-
trolled P. "aphanidermatum" in sugar cane with cbloropicrin used a t
t h e r a t e of 200 lbs. per acre. _ _, - '
R a n g e : United States, Brazil, England, Hawaii, India, Italy, Malaya, Mau-
ritius, Philippines, Puerto Rico; probably widely scattered.
R e f e r e n c e s : Andrews (1943); Bourne (1922); Bowman,'et al. (1937);
Branstetter (1927); Buchholtz (1942,tt); Buchholtz, Melhus, )Velch, and
PHYCOMYCETES 33
Murphy (1947); Carpenter (1919, 1921, 1934); Conners (By Yanter-
pool, 21st Ann. Rept. Canad. PL Dis. S. pp. 6-8, 1941); Conners and
SavUe (1942); Cooper and Chilton (1947); Drechsler (1928,6; 1936);
Edgerton, Tims, and Mills (1929); Elliott et al. (1937, Figs. 1-8, 17, 21);
Flor (1930); Hanson (1944,a); Hawk and Welch (1948); Ho and Melhus
(1940); Ho, Meredith, and Melhus (1941); Ho and Koepper (1942);
Hoffmaster (1942); Hoppe (1949); Johann (1928,6); Johann, Holbert,
and Dickson (1928); Kendrick and Briggs (1939); Leukel (1942, Fig. 1);
Mathews (1931); Melchers (1942); Melchers and Lowe (1940, 1943);
Melhus, Martin, and Murphy (1943); Middleton (1943); Rands and
Dopp (1933, 1934 1938,a,6); Richardson (1942); Robertson (1931,6);
Roldan (1930, 1^32); Sallans (1942); Semeniuk, Wallin, and Melhus
(1947); Simmonds, Russell, and Sallans (1935); Sprague (1943,a,rf;
1944,6; 1946,c,e)j Sprague and Atkinson (1942); Subramaniam (1928);
Summers (1948); Valleau, Karraker, and Johnson (1926); Vanterpool
(1933,-1935, 1938, 1939, 1940,a,6; 1942); Vanterpool and Sprague (1942);
-Vanterpool. and Truscott (1932); Wagner (1936).
Pythium hypogynum Middleton—Root Necrosis

Mycelium nonseptate when young, irregularly septate when old,
measuring 1.5 to 8.3 /x,, averaging 5.1 jii diam. Sporangia terminal, rarely
intercalary, spherical to subspherical, smooth- and thin-walled, 6.5-
34.6 IX,, average 22.1 /x diam., sp'orangium germinating by means of germ
tube or zoospores. Oogonia terminal, spherical to subspherical, smooth-
and thin-walled, 10.2-35.1 /x, average 22.0 /x diam. Antheridium hypogy-
nous, single, delimited within the oogonial stalk at a distance varying
from 5-30 /t below the oogonium; antheridial cell small, 2.8-8.3 /x,
average 5.4 /x wide, 3.0-11.1 fx, average 6.6 /x long, supplied with a
tenuous fertilization tube that just penetrates the oogonial wall.
Oospore plerotic, smooth-walled, containing a single reserve globule
and a prolate ellipsoidal or flattened refringent body. Oospore germina-
tion not observed.
On: Avena sativa L., Nebr.—Hordeum vidgare L., Mo.—Triticum aestivum
L., N. Dak.
Middleton (1941) reported that this species caused a root rot of
barley in poorly drained soils in Missouri. The infected roots were dis-
colored with considerable cortical decay accompanying-the discolora-
tion. Diseased plants are usually stunted and chlorotic. As mentioned
previously (page 26), we have isolated this fungus a few times from
oats and wheat growing in wet soil in the plains region. This species
needs additional study'to determine whether the hypogynal nature of
the antheridia is a regular and specific trait of this fungus or whether
it represents only a variation of the condition found in P. ultimum.
Range: Known only from the central United States.

References: Middleton (1941, 1943); Sprague (1946,e).
34 DISEASES OF CEREALS AND, GRASSES
P y t h i u m i r r e g u l a r e Buisman:—Root Necrosis
Mycelium-well developed on n u t r i e n t agar. H y p h a e 2-6 /A diam., side

branches numerous. Sporangia spherical to pjlriform, terminal or inter- -
calary, 10-30 ju, diam., not a b u n d a n t in most cultures, forming spores in
a vesicle or germinating directly as conidia. I Discharge t u b e short or
equal to the diameter of the sporangium; zoospores as m a n y as 15 or
more, 4-6 X 10-12 jn. Oogonia terminal or intercalary, sessile or stalked,
spherical to cylindric, v a r y i n g greatly in shape; the wall smooth or
with several projections varying in length, which^may or m a y not be
cut off b y a cross wall, diameter of oogonium 14-26 (i without t h e p r o -
jections. Oospores smooth, not filling the oogonium, 10-20 /* diam.,
each of typical structure with a h e a v y wall at m a t u r i t y and a central
reserve globule, surrounded b y a granular layer in which a refractive
b o d y is embedded. Antheridia stalked, 1-3 to an oogonium (usually
only one), androgynous or diclinous, very rarely hypogynous, usually .
arising from the oogonial stalk (Mathews, 1931, pp. 96-97).
On: Agropyron cristatum (L.) Gaertn., N. Dak.—A. intermedium (Host.)

Beauv., N. Dak., S. Dak., Wash.^-k. trachycaulum (Lk.) Make, N. Dak.
—Andropogon hallii Hack., Nebr.—Arrhenathenim elatius (L.) Beauv.,
Wash.—Avena byzantina K. Koch, Iowa, Minn., N. Dak.—Avena nuda
L., N. Dak.—Avena sativa L., Iowa, Minn., N. Dak., Oreg.—Boutelouw
gracilis (H. B. K.) Lag., S. Dak.—Bromus carinatus Hook, and Arn., •
Mont., Wash.—B. inermis Leyss, Minn., Nebr., N. Dak.—Bromus tecto-
rum L., Nebr.—Elymus junceus Fisch., N. Dak.—Festuca elatiorlh., N : '
Dak.—F. elatior var. arundinacea (Schreb.) Wimm., Mont.—Hordeum
bidbosum L., Wash.—H. vulgare L., N. Dak.—Saccharum officinarum L.,
La.—Sporobolus cryptandrus (Torr.) A. Gray^ Nebr.—Stipa comata Trin.
and Rupr., Wyo.—Triticum aestivum L., Minn., Nebr., N. Dak., S. Dak.
P. irregulare causes a root necrosis which is virtually identical with
that caused by P. debaryanum. While it is much less prevalent than the
latter,.P. irregulare is a frequent isolate from the roots of Gramineae
in the plains region of the northern United States^-particularly on oats
in June. It is readily distinguished by its small oogohia with their
irregular, pseudo-spiny walls. Even under the lower magnifications on t
a compound microscope the crisp, somewhat glacial appearance of the
smaller oospore is readily distinguishable. Macroscopically, cultures of
P. irregulare resemble those of P. debaryanum. They are ,cottony when
free from bacteria.
All isolates of P. irregulare tested for pathogenicity were parasitic
on at least some species of grasses. Certain isolates were strongly para-
sitic, and a few races or isolates caused se^vere root necrosis on wheat
and oats under artificial moculation- tJonditioiis (Sprague, 1944,6;
1946,e; and unpublished notes). Green vegetables, such as beets, cab-
bages, beans, and peas, were also injured by isolates 'Of this fungus
originally obtained from Gramineae.
PHYCOMYCETES 35
R a n g e : Northern United States, Europe, probably widely distributed on

Gramineae, as well as being prevalent on many nongrass hosts.
R e f e r e n c e s : Buisman (1927); Mathews (1931); Middleton (1943); Sprague
(1944,6; 1946,e).
P y t h i u m m o n o s p e r m u m Pringsh.—Root Necrosis
P. complens A. Fischer
Mycelium well developed on ordinary media. Hyphae branched,
1.6-7.3 ju diam., often with many bud-like outgrowths. Sporangia un-
branched and arising from ordinary filaments, or composed of highly
branched elements cut off from the remainder of the mycelium by cross
walls. Zoospores few to 50 or more to a vesicle, 7.9-9.6 jx long in swim-
ming stage. Oogonia abundant on agar cultures, smooth or with a
papilla, terminal or intercalary, 15.6-21.6 /A diam. Oospores smooth,
usually filling the oogonium, each possessing at maturity a very thick
wall and a central reserve globule surrounded by granular protoplasm
in which a refractive body is embedded. Antheridia one or two to an
oogonium, club-shaped, androgynous or diclinous.
O n : Hordeum vulgare L., Mo.—Panicum miliaceum L., N. Dak.—Saccharum
officinarum L., La.—Sporobolus_ cryptandrus (Torr.) A. Gray, S. Dak.
This .fungus has filamentous or semilobulate, branched sporangia.
The paucity of antheridia distinguishes it from P. graminicola.
In general, P. monospermum is considered saprophytic (Mathews,
1931). However, the isolations which we assigned to this species were
parasitic under artificial inoculation trials started at seeding time
(Sprague, 1946,e). We obtained this fungus in pure culture only twice
during the course of somewhat extensive studies in North Dakota and
vicinity, once from mildly necrotic roots of Turghai proso millet at
Mandan, North Dakota, and once from Sporobolus cryptandrus on the
range near Buffalo, South Dakota. Pathogenicity trials averaged 62 per
cent loss in the generally root rot-resistant crested wheatgrass, 100 per
cent kill in alfalfa and 28 per cent in oats. Blue grama, which is usually
very susceptible to Pythium spp. in the seed-sprouting stage, suffered
only 31 per cent loss, and corn was resistant with 4 per cent injury.
On nongrass hosts Chinese cabbage was almost completely destroyed
while tomatoes were wiped out. Drechsler, on the other hand-(1925,a),
could not infect cabbage heads with an isolate of this species. There is
need for further study of this organism.
Range: As a grass and cereal parasite it has apparently been reported only
from the northern Great Plains, La., Mo., U. S., and on rice from Japan
(Ito and Tokunaga, 1933). Apparently not uncommon in soil over wide
areas, however.
References: Drechsler (1925,a); S. Ito and Tokunaga (1933); Mathews
(1931); Middleton (1943); Rands and Dopp (1938,o).
^ - -1
P y t h i u m nagaii S. I t o and Tokunaga—:Leaf and Sheath Spot
H y p h a e 1,5-4 /A (average 2.5 JA) in breadth, at the tip of which are

produced ovoid, pyriform, or rarely spherical, proliferous sporangia,
24-36 X 20-26 /x. Zoospores 12 X 7.2 /x, (8.2-9.6 /* when encysted).
Sporangia also germinating directly b y means of a germ-tube from the
apex. Oogonia sphaeroidal t o irregular, smooth, 14-22 /i diam., each
having a single ovoid, globoid, or clavate antheridium. T h e spherical,
smooth, guttulate oospores measure 12-19 ft, diam., their pale t o dark
yellow wall being less t h a n 0.8 /i wide.
O n : Oryza sativa L., Ark.

This species has been reported from Arkansas (Edson and Wood,
1937) as a leaf and sheath spot (cf. also Middleton, 1943, pp. 82-83).
Investigations have also been in progress in Louisiana on a root disease
of rice with an associated undetermined species of Pythium present.
Ryker and Douglas noted (1941) that when root maggots {Lissorhop-
trus simplex) were also present injury was appreciably increased.
The status of P. nagaii as a^distinct species "is not clear. Ito and
Tokunaga have described a number of "new" species on Gramineae. In
the same article in which P. nagaii was described (1933) they also de-
scribed P. oryzae and P. echinocarpum on rice as well as noting P.
monospermum and an undetermined nonsporulating species oh this ,
host.
Range: The United States and Japan. .
References: S. Ito and Tokunaga (1933); Middleton (1943).
i
Pythium oligandrum Drechsler—On Wheat
Growth on corn meal agar somewhat arachnoid, finally pebbly or
crustose due to accumulation of oospores. MycMiumJias many fine-
side branches which give it a delicate appearance. Subspherical sporan-
gial enlargements develop in sterilized water; these consist of single '
subspherical bodies or of two or more globose enlargements together
with connecting and adjoining hyphal parts. Evacuation tube 15 p. to
225 /x long, terminating in a vesicle whifch produces a' number of
zoospores. Oogonia formed early, each with one or two antheridia, or
often formed parthogenetically, stoutly warty-spiny, when mature with
lumpy protoplasm, 17-35 /* diam., commonly 24-30 a; oospores 15-30 i^
germinating after some months, when bathed-in sterile water, producing
a globular, stalked vesicle with zoosporan'îa or sometimes developing
vegetatively.
On: Stiya sp.; no state citation (Middleton, 1%^?,)—Triticum aestivum L.,
Tex.
PHYCOMYCETES 37
Drechsler received cultures of this species from A. A. Dunlap who
isolated it from wheat roots growing in the Panhandle region of Texas.
Middleton (1943) listed it on Stipa sp. Drechsler reported it from a
number of hosts including beets, cucumbers, sweet peas, garden peas,
and watermelons (1946,a). He believed it to be a secondary invader in
most instances, although sometimes it was a parasite on other Pythium
spp. He found that it could cause a soft rot of watermelon. Drechsler
worked with several strains of the fungus.
Drechsler discussed in detail (1946,a) the taxonomical and nomen-
clatural complexities of the fungicolous species of Pythium with spiny
oogonia. On the basis of present information it would appear that this
species of Pythium with spiny oogonia is of slight consequence as a
parasite of Gramineae.
We isolated a species of Pythium with spiny oogonia from Distichlis
stricta growing in blue mud near Sulfur, South Dakota, and this fungus
may have been P. oligandrum, although this South Dakota isolation
may possibly have been closer morphologically to P. acanthicum
Drechsl. Drechsler isolated the latter species from watermelons (1930;
1939, Figs. 2-4, pp. 398-400).
Reference: Drechsler (1946,a).
Pythium ostracodes Drechsl.—On Wheat Roots

Intramatrical mycelium in transparent agar media frequently of
somewhat lustrous, radiate appearance, capable of approximately 15
mm. radial extension ("/*" in Drechsler, 1943, typographical error) in
24 hours at 24° C.; hyphae mostly 1.8-6.5 /^ diam. Aerial mycelium
sometimes absent, but at other times moderately or even rather
abundantly developed, especially under not too humid conditions,
persisting long withoiit collapse. Sporangia under aquatic conditions
formed terminally on long, simple, or sparingly branched extramatrical
hyphae, mostly 2.5-4.5 /x wide; sporangia subspherical, prolate-ellip-
soidal, or lemon-shaped, 25-55 X 16-38 fx.; discharging contents into a
vesicle through an evacuation tube which is often reflexed, 2-45 X 4-9
/x; discharge tube usually formed as a prolongation of an apical papilla,
but sometimes arising from a lateral position, especially after frus-
tration of an apical tube; sporangia proliferous in moderate measure,
either through subsporangial 'branching, or through uniaxial elonga-
tion of the supporting filament. Zoospores 5-35 /x, kidney-shaped, lat-
erally biciliate, rounding to form cysts mostly 10-12.5 n diam. which
germinate usually by putting forth a germ tube about 2 /* long.
Oogonia sometimes formed terminally on lateral branches less than
25 IX long, but more often borne directly on long filaments in terminal,
lateral, laterally intercalary, and mesially intercalary positions; in
some instances protruding noticeably toward the antheridium, yet gen-
erally subspherical, 14-43 (mostly 30-41, average 35.4) ;«, diam.; wall
0.5-1.0 /x thick. Antheridia one or two per oogonium, usually formed
terminally or somewhat laterally on a branch 2 to 100 jx from the
oogonium, but occasionally of diclinous origin and then usually borne
terminally on a branch 50-250 n long arising ;from a distant filament;
antheridia elongate-cylindrical, clavate, or saccate-shaped, often with
somewhat wavy contours, 10-20 X 4-7 /*, sometimes attached to the
oogonium apically, but at other times attached lengthwise for some
distance below the apex, if not for the entire distance from apex to base,
producing a fertilization tube usually 1-3 X 1-1-5 /*. Oospore usually
distinctly yellowish, subspherical, 13-41 (mostly 28-39, average 33.5)
ji. diam.; provided with a wall 1.5-4.3 (mostly 3.2-4, average 3.6) ju, in
thickness; containing 10 to 75 reserve globules mostly 3-4 /A diam. and
5-20 refringent bodies 2-3 fx diam.; germinating readily by producing
a mycelium, or under aquatic conditions, by extending a simple or
somewhat branched sporangiophore, 75 to 800 jx long, whereon 1 to 3
sporangia are borne terminally, often proliferating.
/
On: Triticum aestivum L., Tex.
This species was described from a culture sent by A. A. Dunlap to
Dr. Drechsler for identification. It was isolated from wheat roots ob-
tained from the Panhandle region of Texas. The features of this species
are included in portions of the summary given by Drechsler which is
perhaps simpler than his detailed description noted above: "Its, pro-
duction of subspherical zoosporangia at the tips of hyphae, together
with its development of oospores having multiplicate internal organi-
zation, characterizes it as a member of the helicoides series. In this
series it is distinguished more especially by the very simple make-up
of its sexual apparatus; . . . When bathed in fresh water its oospores
germinate very readily by producing, often in part through proliferous
development, one to three terminal, subspherical, zoosporangia. During
germination the very thick, shell-like^ colored, outer layer of the
oospore wall becomes conspicuously honeycombed'with^radial pockets
and canals as its substance undergoes progressive resorption."
Except for the fact that this species was isolated from wheat root-
lets there is no evidence available regarding its pathogenicity. If it is
at all parasitic it must be largely confined to a more southernly area
than in the regions worked by Buchholtz, Melhus, Sprague, and others
as no fungus resembling this has been recognized in the northern plains
area and prairie country. Unusual forms, however, appear not infre-
quently in all of this region and perhaps it may have been overlooked
or does not lend itself to ready isolation.
Range: Known only from the Texas Panhandle.
Reference: Drechsler (1943).
PHYCOMYCETES 39
Pythium periilum Drechsl.—Root ]S[ecrosis

Mycelium on corn meal agar weak, flat, with scattered tufts or
rosettes of growth; swollen digitate, lobulate sporangia formed from
which elongate evacuation tubes arise. Spherical sporangial vesicles
formed at apex, later discharging biciliate zoospores by disintegration
of the vesicular membrane. Oogonia long-lived, developing readily on
corn meal agar, frequently intercalary, less often subterminal or ter-
minal. Oogonia usually enwrapped with supporting hyphae on which
are borne 2 to 5 antheridia which become delimited by basal septa;
oogonium with a central reserve globule surrounded by a dense gran-
ular parietal layer in which is embedded a single refringent body, sub-
spherical or oblate-ellipsoidal in shape, 2 oospores sometimes occur in
one oogonium; size variable, 16-22 [i, more often 17-20 ju.'
On: Agropyron repens (L.) Beauv., Minn.—Saccharum officinarum L., La.
Stevenson and Rands (1938) reported that this species was a feeble
parasite on sugar cane. The isolations made by the writer from roots
of Agrapyrart were obtsined from a gray ghcinl soil near Bemidji,
Minn. The fungus grew very feebly on potato-dextrose agar and on
cornmeal agar. It showed the characteristic enwrapments about the
oogonium which Drechsler illustrated (1940, Fig. 5). The isolates from
Bemidji died before their pathogenicity could be tested; but it is
doubted, from the relatively vigorous condition of the host, that the
fungus was very parasitic.
Range: Known only from the United States.
Reierences: Drechsler (1930, 1940); Rands and Popp (1938,a); Sprague
1946,e); Stevenson and Rands (1938).
Pythium rostratum Butl.—Root Necrosis

Mycelium well developed, hyphae irregularly branched, up to 6 or
8 fji diam., with a few cross walls when old. Sporangia terminal or
intercalary, spherical to oval, 23-34 jx diam. (average about 28 ju.),
appearing before'oogonia. Discharge tube lateral, large, usually thick-
ened in the middle, length about equal to diameter of sporangium.
Conidia (gemmae) similar to sporangia except in germination. Oogonia
usually about 21 fx. in transverse diam., larger longitudinally. Oospore
smooth, usually completely filling the oogonium, 12-26 /x diam. (aver-
age 21 fx). Sometimes 2 or. 3 oogonia formed in a short series. Antheridia
usually single, arising from the oogonial hyphae; in some cases hy-
pogynal or a short lateral process formed immediately below the
oogonium (Mathews, p. 73).
On: Avena sativa L., Wash. Middleton, without specific location, reports the
species on oats, barley, rice, and wheat in the United States (1943).
40 DISEASES OF CEREALS AND] GRASSES
T h i s species is generally considered to be a soil saprophyte. I t is
doubted if the fungus is parasitic, a t least to any extent. I t was asso-
ciated with a mild root necrosis during c o o l w e t weather a t Pullmanj
Washington, in M a y , 1942. I t has been reported from Iowa soils as well
as from F r a n c e (Butler, 1907), H a w a i i , and|from Georgetown, South
Carolina (Mathews, 1931, p. 73, pi. 17). T h e fungus is characterized
by the somewhat rostrate oogonia.
R e f e r e n c e s : Butler (1907); Mathews (1931, p. 73, pi. 17); Middleton
(1943).
P y t h i u m s p l e n d e n s B r a u n — R o o t Necrosis
Mycelium profuse on media, branched, 3.5-9.2 ix (average 6.4 ju)

diam. Conidia always terminal, smooth, spherical, 21.7-48.9 /* diam.
(average 36.2 /x), germinating by t h e formation of 1 to 6 germ tubes.
N o sporangia. Sickle-shaped, clavate bodies formed in agar cultures.
Oogonia rarely formed, smooth, spherical, 25.5-34.7 /i diam. (average
31.7 ji). Oospores not filling the oogonium, smooth, 21.3-29.8 fx diam.
(average 26.6 jx) in m a t u r e stage, with a heavy wall and a central
reserve globule surrounded b y a layer of granular protoplasm in which
a small refractive body is embedded. Germination direct after a resting
period. Antheridia 3 to 8 to an oogonium, arising from the oogonial
stalk or from distant h y p h a e . Antheridial cell large, clavate, fused at
t h e t i p with the oogonial wall.
On: Saccharum officinarum L., Hawaii—Triticum aestivum L., Hawaii—T.

durum Desf., N. Dak. .
This species, as recognized, is rare on Gramineae and possibly more
or less accidental. We isolated it once froin mildly necrotic roots of
durum wheat at Mandan, North Dakota. Another isolate of P. splendens
from garden peas {Pisum sativum L.), at Mandan caused seed decay
and root necrosis in wheat, oats, crestedjWheatgrass, blue grama, alfalfa,
proso,' and sorghum under artificial inoculatie.n conditions. Braun
(1925)', who isolated P. splendens from Pelargonimn^QXiitmgs, proved
that the species is pathogenic to coleus and begonia cuttings and also
to cucumber and radish seedlings. Braun distinguished P. splendens
from P. debaryanum and allied species because of the larger size of
the fruiting bodies of the former, the distinctive growth in pure cul-
ture, preponderance of conidial production, preference for higher tem-
peratures, and by its tendency not to produce oogonia. in pure culture.
I t also fails to produce zoospores, the conidia always germinating
directly. ' /
Range: United States, Hawaii, Japan, Netherlands,
References: Braun (1925); Mathews (1931); Middleton (1943).
PHYCOMYCETES 4]
Pythium tardicrescens Vanterpool—Root Rot, Root Browning

Mycelium with a flat, radiate, nonlustrous growth on agar; hyphae
mostly 2.5-5.0 jx diam., irregularly branched with fine laterals; knob-
like appressoria often present; radial growth on carrot-maize-meal
agar about 11 mm. in 24 hrs. at 22° C. Toruloid buds or a moderate
development of lobulations infrequently produced in old cultures on
the surface of plain agar containing wheat root tips, or in sterile water
root-tip cultures, occurring naturally intracellularly in-living tissues;
buds never complex, rarely exceeding 12 /i diam.; zoospore discharge
not observed, instead germ tubes terminating in a dark conidium are
produced; spherical walls of germinated conidia frequently conspicuous
in culture. Oogonia smooth, terminal on short branches or rarely inter-
calary, 17-30 fj. (average 24.1 n), commonly with large, lustrous oil
globules; oogonia forming readily in plain agar containing wheat root
tips or in water root-tip cultures, but more sparsely on agar medium
alone; empty oogonial "shells" common following failure to mature,
though occasionally oogonia become filled with tangled hyphae; de-
limiting septum frequently visible in the supporting hypha below the
base of the oogonium. Antheridia as many as 6 but more^ often 2-3,
club-shaped or crook-necked, averaging 6-§.5 /* wide, 10.5 /JL frorh apex
to curve, and 5.5 /x from curve to basal septum, fertilization tube 2 /x
and making fairly narrow to medium contact with the oogonium;
antheridia commonly arise from the oogonial stalk or a branch but all
may come from neighboring branches. Oospores smooth, spherical or
subspherical, usually free within the oogonium, 16-26 /x (average 20.3
/JL) diam., with a central globule averaging 11.1 /x which is usually clear
and embedded in a finely granular matrix; oospore wall 1.25-2 /*;
oospores forni best in solid or liquid media containing wheat root-tips
(Vanterpool, 1938).
On: Avena fatua L., Wash.—Echinochloa crusgalli (L.) Beauv., Minn.—

Triticum aestivum C, Sask.
Vanterpool found this species relatively common on wheat in
Canada and in England (1938) but it has be^fjn reported from the
United States only three times, once on wild oats at Pullman, Wash-
ington, during cool wet weather in May, once on barnyard grass grow-
ing in poor soil at Graceton, Minnesota, and once by.I)rechsler (1947)
from Virginia on the nongrass host Bidens. It produces a firm, dark
brown rot or root browning, and in our trials had approximately the
same host range and relative pathogenicity as P. graminicola. The
oogonia generally average smaller than those of the latter species and
have a characteristic subglobulate content. The fungus produces weak,
scanty growth in culture on cornmeal agar, and in our isolates, usually
only a few oogonia matured.
This species needs further investigation in the United States and
should be critically compared with some of the Japanese species de-
42 DISEASES OF CEREALS A^ND, GRASSES
scribed by Ito and Tokunaga (1933, 1935). Because of its relativelj^
feeble growth it is difficult to isolate, a^ fact that may account for its
not having been observed to any extent except by Vanterpool whose
experience with the group permitted him to obtain a higher percentage
of survival among his isolates. |j
Drechsler (1947) obtained zoospore formation from oospores that
were taken from cultures 112 days old. The cultures had been-jsolated
from a blackened root of Bidens sp. obtained at Arlington (Farm),
Virginia. He stated that oospores germinated by producing about 10
zoospores in a vesicle at the slightly reflexed orifice of a simple or
occasionally somewhat branched evacuation tube, which, commonly
measured 60-350 X 2-3 /i. After discharge the remnants of the oospore
wall enclosed a separate sporangial envelope.
Range: United States, Canada, England.
References: Drechsler (1947); Sprague ,(1943,a; 1944,&; 1946,e); Vanter-
pool (1938, Fig. 1, PI. XXII; 1939; 1942).
Pythium ultimum Trow—Root Necrosis

Mycelium well developed on^media, branched, septate in old cul-
tures, 1.7-6.5 IX (average 3.8 /x). Conidia usually terminal and spherical,
12-28 [J. diam. (average 20 ju,), intercalary, barrel-shaped forms, 14-
22.9 X 17-27.8 fi, also formed in cultures; conidial germination direct,
zoospores not formed. Oogonia smooth, usually terminal, rarely inter-
calary, spherical, 19.6-22.9 /x diam. (average 20.6 /x). Oospores not fill-
ing the oogonium, spherical, 14.7-18.3 /t diam. (average 16.3 /x), in
mature stage with a heavy wall and a central reserve globule sur-
rounded by a granular layer of protoplasm iji which a small refractive
body is embedded; germination is direct by one or more germ tubes.
Antheridia usually one to an oogonium 'arising from the oogonial stalk
immediately below the oogonium (Mathews, 1931).
On: Agropyron caninum (L.) Beauv., N. Dak.—A.-cjistatum (L.) Gaertn.,
N. Dak., Wash.—A. desertorum (Fisch.) Schult., NT^Qak.—A. inter-
medium (Host.) Beauv., N. Dak., S. Dak.—A. repens ( t j Beauv., Minn.,
Wash.—A. semicostatum (Steud.) Nees, Wash.—A. smithii Rydh., N. Dak.
—Agrostis alba L., N. Dak.—Andropogon furcatus Muhl., Nebr.—Aveha
sativa L., Oreg., Wash.—Bromus arvensis L., N. Dak.—B. carinatus Hook,
and Am., Wash.—B. erectiis Huds., Wash.-f-5. inermis Leyss, Nebr., N.
Dak., Wash., Wyo.—Dactylis glomerata L., Wash.—Echinochloa crus-
galli (L.) Beauv., N. Dak.—Elymus canadensis L., Wash.—E. dahuricus
Tufcz., Wash.—E. virginicus L., Wash.—Hordeurn vulgare L., N. Dak.—
Panicum miliaceum L., N. Dak.—Phragmites commy/nis Trin., N. Dak.—
Poa ampla Merr., N. Dak., Wash., Wyo.;Ê'. 'canbyi (Scribn.) Piper, N.
Dak., Wyo.—P. secunda Presl, N. Dak.—Secale cereale L., N. Dak.—
Sorghum vulgare L., CaUf.—Triticum aestivum L., Minn., Mont., Nebr.,
N. Dak., S. Dak., Wash.—T. durum Desf., N. Dak.—Zea mays L., N.
Dak.. Sask., S. Dak.
PHYCOMYCETES 43
Middleton (1943) mentions that this species is the most prevalent
one of the genus on plants that suffer from damping-ofi. On Gramineae,
on the other hand, it is apparently less common, in most regions, than
P. debaryanum. Usually it seems to be somewhat less parasitic on
cereals than the latter. This is, however, only a generalization, for we
have worked with isolates of P. ultimum which caused severe root
necrosis and seed decay in wheat and oats under artificial inoculation
conditions (1944,6 and unpublished notes).
The main distinction between P. ultimum and P. debaryanum is
based on the concept that in P. ultimum the antheridium originates on
the oogonial stalk usually just beneath the oogonium while in the other
the antheridium, or, more often the antheridia, may originate from
distant hyphae. This point is often difficult to determine in those cul-
tures which produce few oogonia.
In addition, it has frequently been said that the oospores of P.
ultimum do not produce zoospores, but germinate directly, as do the
conidia. Drechsler (1946,5), however, vfas able to produce zoospores
from three-month-old oospores from cornmeal cultures within 18 hours
after transfer to a shallow layer of water in Petri dishes maintained
at 10° C. An evacuation tube, usually 3-6 ix wide beyond the frequently
broadened base, was pushed through the oogonial wall and grew to
a total length of 10-40 /*, when it formed a papilla through which the
protoplasmic contents soon migrated into a vesicle where, usually,
eight to ten zoospores wel-e formed. When encysted they measured,
mostly 8-10 (rarely up to 13 fj.) ju diam. and germinated by the emission
of one or two hyphae, usually 2-3 /^ wide, but they also often showed
repetitional emergence. At 25° C. the zoospores germinated by hyphae
even without the addition of fresh water, after removal to a glass slide.
This is said to be the first report of zoospore formation in the species.
Drechsler does not believe that zoospores are important in the spread
of the disease-causing fungus in the soil. For all practical phytopatho-
logical purposes, P. debaryanum and P. ultimum could be considered
as a unit, although morphologically, perhaps, they are distinct.
' In the northern Great Plains, as indicated, P. aebaryanum is much
more important on grasses and cereals than P. ultimum, but it should
be noted in passing that P. ultimum seems to be sometimes more preva-
lent during early spring in cool, wet weather in the Palouse region of
Washington.
Range: Widespread in the temperate zones throughout the world.
References: Drechsler (1925,a;'1927,a; 1946,5); Mathews (1931); Middle-
ton (1943); Sprague (1944,&); Trow (1901).
Pythium volutum Vanterpool and Truscott—^Root Browning
Mycelium nonseptate, lustrous, with an aerial tendency in culture,

radial growth on agar of approximately 16 mm. in 24 hrs. at 22° C ;
44 DISEASES OF CEREALS AND, GRASSES
/ I
appressoria consisting of lateral, falcate structures with rounded ends;
mostly intracellular in host tissue.--Sporangia consisting of small lobu-
lations or toruloid buds formed only rarely'in aqueous culture; dis-
charge tube usually about 50 /x long and 3-4 /* wide; zoospores bi-
flagellate, bean-shaped, about 10-14 /x. Oogonia smooth, subspherical,
dark brown, terminal on short side stalks orlrarely intercalary, formed
copiously in culture but a large percentage remain sterile, average diam.
30 /J,; antheridia 3-10 to each oogonium, crook-necked, sometimes
curved or straight, with narrow apical contact, usually arising from
adjacent hyphae, each of which supplies one to four antheridia, or
more rarely arising from oogonial stalk; antheridial branches, com-
monly entwining about the oogonial stalk in liquid media, but less fre-
quently on solid media. Oospores smooth; spherical to oblong, usually
free within the oogonium, average diameter 27.7 jn, central globule
14.2 fi, refringent spot 8.5 X 2.2 /x, oospore wall 2.0 jn. Oblong oospores
(average 36.9 X 19.2 ix) with two reserve globules are sometimes pres-
ent in the host cells.
On: Avena sativa L., Sask.—Triticum jxestivum L., Sask.
This appears to be allied to P. drrhenomanes, or P. graminicola but
Vanterpool and Truscott pointedôut some differences which they con-
sidered diagnostic. The antheridia tend to coil about the oogonial stalk
which is not typical of P. graminicola. The fungus grows faster on
carrot-corn meal agar; the formation of lobulate sporangia is feeble
or absent in P. volutum; and, as reported, P. volutum is more aggres-
sively parasitic on oats than either P. arrhenomanes or P. graminicola.
All of these traits, except possibly the tendency to coil about the
oogonial stalk, are relative, and future studies may indicate that P.
volutum should be considered a form or phaê of P. graminicola.
Range: Canada in areas in northern aiid in southern Saskatchewan, Nether-
lands, England. ,
Reference: Van Luijk (1938); Vanterpool and Truscott (1932).
Pythium spp.—Mostly Saprophytes '

In the course of investigations on root rots of many grasses and
cereals (Sprague, 1944,b, and unpublished notes) a large number of
isolates of Pythium species were not determined, or, if determined,
proved to be saprophytic forms which were of little or ho further in-
terest from a pathological standpoint. How.ever, several of these identi-
fied forms are included in the key to the species (pp-. 19-20), because
they can be confused with the parasitic ones. One ,6i these, P. vexans
de Bary, is especially common. It produces'an off-white, somewhat flat
or scant growth on nutrient agars usually'without the formation of
oogonia, although chains of conidia are often formed. When oogonia
are formed, on carrot agar, or on wheat tips in agar or water, they are
PHYCOMYCETES 45
distinguished from those of P. debaryanum by the broadly appressed
antheridia (Drechsler, 1946,a, b). The antheridia often encircle nearly
Yg of the circumference of the oogonium. We have never found satis-
factory evidence that this species is parasitic on Gramineae, even
though sometimes it is common on necrotic roots of cereals and grasses.
Another species, P. intermedium de Bary, produces a relatively
scanty growth on cornmeal agar. Conidia are produced in chains but
no other fruiting bodies occur. It is entirely saprophytic on grasses.
It is sometimes isolated in early spring, before P. ultimum or P. de-
baryanum appear in isolations, from roots of Gramineae in the north-
ern Great Plains.
Many cultures of Pythium fail to produce sexual bodies on either
potato-dextrose agar or cornmeal agar. Their somewhat luxuriant
growth and their ability to cause at least moderate root necrosis sug-
gests that they may be either P. ultimum or P. debaryanum. When
grown with excised wheat roots or in shredded carrot agar, however, a
number of these do form oogonia and are identifiable as parasitic
species of the sphaerosporangial group (e.g., P. debaryanum, P. ulti-
mum, etc.).
There have been a scattering of isolations of Pythium which have
not been determined to species because they appeared to be different
from those known on Gramineae. One of these was isolated from roots
of Puccinellia nuttalliana (Schult.) Hitchc. found growing in alkaline,
blue mud (odor that of hydrogen sulphide) on the shore of an alkaline
lake near Crystal Springs, North Dakota. A similar fungus was iso-
lated from Salicornia rubra Nelson growing in the same type of odorous
alkaline mud near Sulfur, South Dakota. This fungus produced a some-
what lustrous, flattened, "pebbly" growth on cornmeal agar. Oogonia
were large, each with several attached antheridia. The fungus was
typified by many vacuolated bodies on short twisted branches, or in
chains. These bodies may have been aborted oogonia. The weird, dis-
torted hyphae and its general sterility may have been due to the ex-
treme difference in environment encountered by the fungus on being
transferred from an alkaline mud "flat" to nutrient agar. The species'
did not resembleâny known to the writer and it is probably unde-
scribed. Its parasitic proclivities were not determined.
P. megalacanthum de Bary has spiny oogonia. It was sometimes
found as a parasite on other species of Pythium, usually P. vexans or
P. ultimum. It was nonparasitic on grasses and cereals in artificial
inoculation trials (Sprague, unpublished reports).
A species of Pythium belonging to the sphaerosporangial group was
isolated from the roots of Poa compressa L. at Powder River Pass, in
the Big Horn Mountains, Wyoming. I t was not assignable to any of
the common species on the basis of brief study. A species resembling
P. pemiciosum Serbinow was isolated from Bromus tectorum near
Lilargent, Washington, in February, 1947.
40 DISEASES OF'CEREALS AND GRASSES
/I
An isolate from Beckmannia syzigachhe (Steud.) Fern., obtained
from a roadside ditch at Alzada, Montana, appeared to be similar to
P. gracile Schenk which is a parasite of green algae growing in a simi-
lar habitat. Middleton (1943) isolated P. gracile from wheat -and
cites other reports on Gramineae. Another isolate, from the roots of
Scolochloa festucacea (Willd.) Lk. also growing in a swampy habitat
near Medina, North Dakota, resembled P. splendens. None of these
fungi appeared to be at all parasitic.
Ling (1947) reported Pythium sp. on barley from Chengtu, Szech-
wan Province, China. Altson (1926) reported Pythium sp. from
graminicolous hosts in British Guiana. Fenne (1941) reported that
Pythium sp. was isolated from wheat, oats, barley, and rye seedlings
showing root rot in Virginia. From 15 to 20 per cent of the stand was
dead from disease in fields of barley and wheat. W. A. Jenkins (1948)
has recently contended that root rot fungi in Virginia on small grains
are secondary to nemas.
References: Drechsler (1930; 1940; 1943; 1946,a); Ling (1947); Mathews
(1931); Middleton (1943); Sprague (1944, also unpublished data includ-
ing notes on pure culture isolations, isolation numbers 1-4329 representing
over 40,000 pure cultures). /
Rhizophidium graminis Ledingham—Rootlet Necrosis

Zoosporangia on roots, 10-75-100 p. diam., globose to ellipsoid, with
smooth walls, solitary or gregarious, giving rise at the base to fine
rhizoids which ramify within the host cells. Zoospores 3 X 1-8 /*, uni-
guttulate and uniciliate, the cilium 10-12i fx. long, liberated in large
numbers by the rupturing of the apex pf the sporangium. Resting
spores 6-12 jn diam., globose, with smooth endospore a n d ' rough,
scabrous exospore wall, and oily protoplasmic contents, sexual fusions
occurring between rhizoids of two thalli. Germination of the resting
spore takes place by the extrusion of the cell Qontents to form, a thin-
walled sporangium. ^"^ -f
On: Panicum sp., Mass.—Triticum aestivum L., Mass., Oht.
Unlike Polymyxa, which is placed in the Plasmodiophorales
(Karling, 1942), this is classed as a member of the Chytridiales. The
amoeboid zoospore attaches itself to the rootlet or root hair and de-
velops a rhizoidal system into adjacent host tissue (Fig. 5,B). This is
accompanied by development of the sporangia on the surface of the
root. These vary in size from tiny bodies about 10 [i diam. to ones ten
times as broad and contain from 8 to/100 zoospores. The sporangia
release the spores by suddenly burstingT The zoospores swim away,
leaving the collapsed sporangium (Fig. 5,^) which persists for several
days. The abrupt and violent ejection of the zoospores is said to dif-
PHYCOMYCETES 47
ferentiate this species from others in Rhizophidium which discharge the
spores through pores.
The sexual process was believed to have been initiated by. the
fusion of rhizoids from which the globose resting spores (Fig. 5,C)
occurring on the surface of the rootlet subsequently develop.
Sparrow (1943) gives a detailed discussion of aquatic species of
Bhizophidium, merely mentions B.
graminis since its habitat on grass '\^—->-N /C~~\ R
roots is not aquatic. ' V^ /
Range; United States, Canada.
Reference: Ledingham (1936).
Rhizopus spp.—Scutellum Rot

of Corn
Koehler (1927) reported that Bhi-
zopus tritici Saito and B. nodosus ^
Namysl., but not B. nigricans Ehrenb., „ J-- j^^^^^Zl^
caused scutellum rot in the corn ger- sporangium; B, developing spo-
minators in the Middle West. He rangium showing rhizoidal devel-
, ,~- , . . /. TIT . opment in root hairs; C, resting
tested a number oi species oi Khizopus spores (after Ledingham). '
and found several of them capable of
causing a scutellum rot disease, but apparently the common species ob-
tained in his isolations were the first two mentioned. Adams and Rus-
sell (1920) did pioneer work with the scutellum rot disease while Hol-
bert, et al. (1924) published data on the field performance of plants
grown from seed susceptible to the fungi and named the symptoms
"scutellum rot." Species of Rhizopus, Mucor, Penicillium, Aspergilliis,
and Fusariwn, among others, are instrumental in causing scutellum
rot in the soil. Since Mucor and Bhizopus include species of fungi
which are considered to be rather weakly parasitic, itîs not deemed
practical to discuss them individually. Taxonomically these two genera
are considered difficult. The work by Lendener (1908) is helpful in de-
termining Mucor while Oilman (1945) is also valuable for the group.
Range: General, at-least in all but the driest regions.
References: Adams and Russell (1920); Hurd (1921); Koehler (1927);
Lendener (1908); Manns and Adams (1923); Taubenhaus (1920); Trost
(1922).
Sclerospora farlowii Griffiths—^Downy Mildew

Infected areas on the leaf sheaths, rarely on the leaf blades, irregu-
lar in outline, usually elongate, brownish with a darker border, up to
1 cm. or more in length; conidiophores not reported; oospores globose,
28-45 ix] epispore slightly wrinkled, very opaque, reddish brown, often
appearing black.
On: Chloris virgata Swartz, Ariz.—Cynodon flqctylon (L.) Pers., Okla.
Griffiths collected this at Cochise,, Arizona, and this, plus a collec-
tion reported on Bermuda grass from Oklahoma, appears to be the only
reported specimens of this species. Wilson (J1907) lists the distribution
as Arizona and Sonora (Mexico). He exaniined material sent by Grif-
fiths fro.ni the type locality. Wilson states | "This species is very dis-
tinct from S. graminicola, from which it differs in the slightly larger
and mpre opaque oospore with its lighter and smoother epispore. The
disintegrating influence of the fungus upon the host is also absent in
the present species, while in S. graminicola this is very pronounced."
Range: United States and probably adjacent Mexico, known range con-
fined to Arizona and Oklahoma.
References: Griffiths (1907); Saccardo (Sylloge, 1912, v. 21, p. 862); Wilson
(1907).
Sclerospora graminicola (Sacc.) Schroet.—Downy Mildew

Protomyces graminicola Sacc. (1876, p. 172)
Peronospora graminicola Schroet. in Zopf. and Sydow,
Myc. March. 9. 1881
Peronospora graminicola Setariae-italicae Traverso
(1902)
Causing (on corn) discolored areas on the leaves, or (on Setaria)
discolored spots or streaks with the white downy mildew covering areas
involved; conidiophores fugaceous, nonseptate, stalk-like with a'slightly
bulbous foot with short apical branches, conidia on short sterigmata,
2-6 on a branch, conidiophores 100 X 10-12 fi but those developed later
in the season are larger (Tasugi, 1933), c6nidia 16-22 X 12-;16 /M on
Setaria viridis; 19-31 X 12-21 /A on Pennisetum typhoideum (India);
13-36.9 X 11-24.9 /* on Setaria magna; mature conidia having a papilla
or dehiscence prominent at the apex^ germinating by the production
and release of 3-12, or more, zoospores which are irregularly kidney-
shaped with 2 cilia on the concave side, 9-12 /x^diain. Oogonia 30-60 /i
diam. with heavy uneven walls which are 4-12 [i thicl^,' reddish brown
or amber brown; oospores pale brown, 22.5-35 /x diarii., averaging 32 fi
bu,t sometimes as much as 45 /J. diam., .germinating by a germ tube.
On: Panicum miliaceum L., Iowa, Minn., N. Dak., S. Dak.—Setaria italica
(L.) Beauv., Iowa, Mich., Minn., N. Y., pask., Wise.—S.'lutescens (Wei-
gel) F. T. Hubb, Iowa, Nebr., N. Y., Wise—5. magna Griseb., Fla.—S.
viridk (L.) Beauv., B. C., Iowa, Minn., Mont., Nebr., N. Mex., N. Y.,
N. Dak., Sask., S. Dak., Wash., Wise.—Zea mays L.,. Iowa, Okla., Wise.
This fungus is favored by high hunaidity and moderate night tem-
peratures which permit the production of conidia, and hence rapid local
spread. Tasugi (1933) found that conidia were produced in daylight
hours if the humidity was high. Corn is not particularly susceptible
PHYCOMYCETES 49
(Melhus and Van Haltern, 1925), or at least is seldom seriously injured
in the better corn-growing areas of the country (Weston and Weber,
1928). In some regions Setaria viridis is injured so severely that the
fungus must be a factor in reducing the spread of this grass weed. This
condition prevails at times in North Dakota and in granitic soil in the
Okanogan area of Washington and British Columbia. The same con-
dition prevails in South Russia (Borghardt, 1932). The fungus is also
common in Iowa and Minnesota during most years. Wilson (1907)
gives the range in this country as Vermont to South Dakota and
Kansas.
This fungus is said to be systemic in the perennial plants of Setaria
magna in Florida (Weston and Weber, 1928). Although this plant was
widely infected in the state Weston and Weber did not find corn in-
fected even when the Setaria weed was closely associated with corn
fields (1928).
The disease on pigeon grass (Setaria viridis) is recognizable by the
yellowish, sickly appearance of the plants and the white, somewhat
downy growth in elongate lesions on the sharply pointed, somewhat
inrolled leaves. Diseased areas eventually turn brown and the infected
plants frequently die or fail to head. Symptoms are similar on culti-
vated Setaria millets. On Pennisetum typhoideum L., in India, the
fungus causes a "green ear" disease, so named because the usually
solid spikelet becomes at least partly transformed into a loose green
head composed of a mass of twisted leaves. The material on sorghum
from India is placed in S. sorghi (Kulk.) Weston and Uppal (1932).
Uppal and Desai (1932) reported the same fungus on maize in India.
Chaudhuri (1932) placed material on Pennisetum in *S. graminicola.
In older material the leaves and inflorescences of the hosts shred
and in these lesions the brownish, resinous-appearing oogonia are pro-
duced in great numbers. Oogonia survived for as long as 600 days
(Tasugi, 1933).
Another downy mildew, Bremia graminicola Naoumoff (Bull. Myc.
Fr. 13:274, 1913) occurs on Arthraxon ciliaris in the Primorye region
of the Ussuri River in Far Eastern Siberia, the type area, and in Japan
and China (Miyake in Bot. Mag. Tokyo 1914). Togashi (1926) noted
the fungus at Kyoto, Japan. The conidiophores were hypophyllous,
rarely epiphyllous, caespitose, flocculose, at first whitish, later grayish,
generally 460-760 /x long (often up to 825 /x) 7.5-10 n broa,d with some-
what swollen bases, and five to six times dichotomously branched, the
branches being more or less recurved and with swollen ends bearing
four to five sterigmata. The, conidia are papillate and measure 11-15 /x
diam. The fungus produces large, yellowish, later brownish, spots at
first delimited by the veins but finally occupying the whole surface of
the leaf and causing desiccation.
Range: Common in the United States, general in Africa, Asia, Europe, and
the Western Hemisphere, reported from Canada, China, Czechoslovakia,
/j
Germany, Hawaii, Hungary, India, Italy, |Japan, Korea, S. Africa, Tan-
ganyika, U.S.S.R.
References: Butler (1918); Evans and Hiirrar (1930); Halsted (1889);
Hirata and Takenouti (1932); Hiura (1^30,a,6; 1935); Hunt .(1940);
McDonough (1937, 1938); Melhus and,Van Haltern (1925); Melhus,
Van Haltern, and Bliss (1928); Pammel, HVeems, and Scribner (1901);
Patel (1949); Prillieux (1884); Sohroeter (1886 in Cohn; 1879); Steven-
son and Rands (1938); Tasugi (1933, 1934); Takasugi and Akaishi
(1933); Traverse (1902, a,b); Uppal and Desai (1931, 1932,a,b); Wang
(1936); Weston (1924, 1929,a); Weston and Weber (1928); Wilson
(1907); Yu (1944).
Sclerospora macrospora Sacc—Wheat and Oats Downy Mildew

*S. kriegeriana Magnus
S. oryzae Brizi
Mycelium intercellular, aggregating near vascular bundles; conidio-
phores seldom formed and abortive; conidia rare but reported from at
least Italy and Japan, papillate, 60-70 X 38-52 ju, borne on short
peduncles, germination by means' of zoospores; oogonia and antheridia
developing abundantly on hyphae parallel to the vascular bundles;
antheridia formed on side branches in the vicinity of the oogonia;
oospores invested by the thick, persistent, oogonial wall with narrow
lumen between; oogonial wall generally smooth, sometimes with knobs
or projections, pale yellow, wall not folded over as in S. graminicola,
wall averaging 4.2 fx thick; oospores spherical to oblong, walls of the
oospore always smooth, lacking the color of the oogonial wall, 35-80 n
diam., average 55-65 ix. Oospore germinating by means of a short
peduncle on the apex of which a single lemon-shaped macroconidium
75-80 X 55-60/A is formed; macroconidiun/producing zoospores in turn.
i '
On: Avena byzantina K. Koch, Miss.-rA. sativa L., Ida., Ind., La., Mich.,
Miss.—Bromics commutatus Schardj, Ky., Tenn.—Hordeum vulgare L.,
Calif., Okla.—Triticum aestivum L.,iDel., Ky., Miss., N. Y., Tenn.
This species has larger oogonia than S. graMinioola. I t occurred in
low moist flooded ground on oats in Mississippi (Miles and Epps, 1942),
although sometimes it was found on upland-grown oats as well. In
general, however, the fungus seems to persist in land which is subject
to flooding. Weston (1921,a) reported infected wheat in Kentucky and
Tennessee displayed excessive tillering, stccompanied by the death and
browning of many shoots. There was distinct dwarfing of shoots, yel-
lowing, thickening, stiffening, twisting, and curling of the leaves.
Peglion (1931) found that some of the oospores germinated after
15-20 days in moist chambers at temperatures not exceeding 18° C ,
each forming a large macroconidium wbicli at maturity was 76-80 X 55-
60 fi. This germinated by production of zoospores.
Peyronel (1929) was able to induce conidia formation on bits of
infected wheat culms and leaves in water. Conidia have also been
PHYCOMYCETES 51
found on plants in Japan. The conidiophores emerged through stomata
in groups. The conidia produce zoospores.
The parasite is seldom important in this country and is feared more
than is usually warranted. Part of this fear stems from the serious
losses encountered in tropical countries on sugar cane and maize which
are caused by congeneric species. Some ten or more additional species
have been reported on graminicolous hosts, mostly on these tropical
species. For a discussion of these species the reader is referred to the
original papers by Weston (1920; 1921,b,c; 1923; 1929,6,c; 1942).
Range: United States, Abyssinia, AustraUa, Bulgaria, France, Germany,
Italy, Japan, Spain.
References: Arnaud (1915); Brizi (1912); Caldwell, Stanton, and Mulvey
. (1949); Kent, Tyler, and Jensen (1947); Mackie (1930); Miles and
Epps (1942); Noble (1934); Peglion (1931); Peyronel (1929); Stanton
and Down (1943); Stanton, Rodenhiser, and Stevens (1949); Weston
(1921,a); Yamada (1911).
Synchytrium graminicola sp. nov.—Purple Leaf Speckle

Maculis minutis, ambitus, diffusis, purpurascens v. amethysteus,
centro'stramineis v. dilute brunneis v. alutaceis; amyceliis, arrhizus;
sorus subepidermibus, globosis v. sphaeroideis, buUatis, operculatis v.
ostiolatis, hyalinis, solitaris v. subgregariis v. cohaerens; sporangia et
sporis (?) destitutis, sorus evac'uatis, peridium firmibus, hyalinis.
Hab. in foliis vivis Gramineae in humidulus pasci, prati et silva in
hiemem.
Spots small, edge diffuse but becoming purple to amethyst colored,
center straw-colored to light brown or yellow gray; without mycelia or
rhizoids; sori subepidermal, erumpent, globose or sphaeroidal, some-
what swollen or bubble-like, with very slight gall formation, operculate
to definitely ostiolate, hyaline, solitary but sometimes gregarious or a
few closely packed; sori^ empty; wall of sorus firm, evident, hyaline.
In living leaves of many grasses in wet pastures, meadows, open
woods, and lawns in winter in western Oregon and Washington.
On: Agrostis aZ6ai., (Dreg.—A. canina L., Oreg.—A. exarata Trin., Oreg.—
A. hallii Vasey, Oreg.—A. palwtris Huds., Oreg. (type)—A. tenuis Sibth.,
Oreg., Wash.—Anthoxanthum odoratum L., Oreg.—Bromus carinatus Hook,
and Arn., Oreg.—Dactylis glomerata L., Oreg.—Elymus glaucvs Buckl.,
Oreg.—Festuca rubra var. commutata Gaud., Oreg.—Poa annua L., Oreg.
—P. jundfolia Scribn., Oreg.—P. prdtensis L., Oreg., Wash.—P. secunda
Presl, Wash.
The unilocular bodies appear to be vacuolated sori of a species of
Synchytrium. I t also has some of the appearance of Eurychasmidium
Karling (1942,a). The writer is puzzled as to why these bodies are
always empty when observed. He has hesitated for a considerable
number of years before describing so unsatisfactory material as this
/
/ i
and perhaps his final decision to name it/ may be unnecessary. How-
ever, the genus Synchytrium does not appear to be mentioned on this
family and the possible relation of this species'to any described form
is vague. The sori resemble some illustratecl by Cook (1945) and the
lack of definite gall formation does not appear to be sufficient to ex-
clude S. graminicola from the genus. There is localized cell hypertrophy
sufficient to cause some slight swelling. '
The fungus is relatively common but very obscure during wet, open
winter weather in the Willamette Valley, Oregon. It has been collected
a few times in Washington, once on the east slopes of the Cascade
Mountains on Poa secunda growing near Easton. Its host range is rela-
tively extensive although the most abundant material occurs on bent
grasses. It sometimes attacks bent grass lawns at Corvallis, Oregon.
Most of our material was sent to Harvard University, Cambridge,
Massachusetts, for study a number of years ago. The remainder is
filed at Oregon State College.
Range: United States in western Oregon and Washington.
References: Cook (1945); Karhng (i942,a); Sprague (1942,a,6; 1946,e).'
ASCOMYCETES
There are a number of important diseases of Gramineae caused by
members of the Ascomycetes. The Ascomycetes are a somewhat difficult
group to present adequately since they include so many species, espe-
cially in the order Sphaeriales, which are genetically connected with
better known conidial' stages. From the pathologist's standpoint one
finds it difficult to decide whether to discuss such species within the
Ascomycetes or the Fungi Imperfecta Generally speaking, the imper-
fect stages are better known to the plant pathologist; indeed, in many
cases the ascomycetous stage has been found by only one or two in-
vestigators. In some such cases the supposed genetical connection be-
tween the imperfect stage and the perfect stage must be considered as
still subject to question. The writer's policy has been to cite the fungus
both in the Ascomycetes and in the Fungi Imperfecti, if both stages
occur, but to include the bulk of the information in the group which
in his opinion is the best known to the general reader.
A key to the Ascomycetes to be discussed is as follows:

A. Fruiting body an apothecium
B. Apothecium arising from a sclerotium Sclerotinia
BB. Apothecium not arising from a sclerotium Phialea
AA. Fruiting body a perithecium
B. Perithecia' fleshy or sunken in a .black stroma
C. Perithecia fleshy (Hypocreales)
D. Stromata encircling culms or sheaths Epichloe
DD. Stromata not as above, spores several-septate, broadened, usually
perithecial stage of Fusarium Gibberella
CC. Perithecia in black stroma (mostly Dothidiales)
D. Stromata stalked or somewhat capitate
E. Stromata somewhat capitate and stipitate, on culms and other
plant parts Balamia
EE. Stromata stalked, capitate, arising from black horny sclerotia
which replace ovary of the flowering host-parts .-.^,,, .-^^Claviceps
DD. Stromata flattened, diffuse, not as above
E. Ascospores 1-celled, hyaline Phyllachora
EE. Ascospores 2-celled
F. Spores hyaline
G. Paraphyses present
H. Spores medianly septate Endodothella
HH. Spores septate near base Placostroma
GG. Paraphyses absent . . .- Dothidella
EEE. Ascospores fUiform, hyaline
53
/i
F. Stroma innate, forming a clyj)eus Myriogenospora
FF. Stroma diffuse (Sphaeriales) ,i , ^Dothichloe
BB. Perithecia carbonaceous or parenchymatous, separate
C. Perithecium without a definite ostiole Erysiphe
CC. Perithecium with a definite ostiole ' j
D. Ostiole an elongated cleft ,1 Lophodermium
DD. Ostiole more or less circular '
E. Perithecia not strongly beaked [short beaks in Ophiobolus (in
part), Cochliobolus, Pleospora, and Pyrenophora]
F. Ascospores filiform, hyaline
G. Spores strongly coiled in the ascus Cochliobolm
GG. Spores not strongly coiled in ascus ,. Ophiobolus
FF. Ascospores 2-celled, hyaline, paraphyses absent
Mycosphaerella
FFF. Spores 1-celled, hyaline, ovoid, paraphyses present
Physalospora
FFFF. Spores several-celled, broadened, yellow or pale brown
Leptosphaeria
FFFFF. Spores muriform, brown
G. Perithecia hairy . . ./ Pyrenophora
GG. Perithecia smooth/ Pleospora
EE. Perithecia beaked
F. Spores 1-celled
G. Spores hyaline Endoconidiophora
(Ceratostomella)
GG. Spores brown, few, lying free in the primitive perithecia
Cry'ptoascus
FF. Spores 2-celled ^ Gnomonia
Balansia claviceps Speg.—Inflorescence Blight

I '
Spikes and rachis somewhat stunted, rachis thickened; stromata
white inside, dark to black on outside, Ihemispherical or globose, arising
from the spikelets, 1.5-3 mm. diam.,'stipe is 1.5-3 mm. long X 1-1-5
mm. thick; perithecia subcortical in cap of j-he stromata, densely-
grouped, distinctly obovate, 200-220 X 120-140 /i,~êFforate ostiole;
asci cylindrical, apices rounded, truncated, bases moderately attenu-
ated, briefly rounded, 150-160 X 5-6 /x, without paraphyses, 8-spored;
ascospores filiform, fascicled, 0.6-0.8 > across, transversely septate,
hyaline. Conidial stage not know^n but a similar species of Balansia
from Cuba bore spores of Ephelis mexicana Fr. (Diehl, 1930,a).
On: Cenchrus echinatus L., Fla., Va.
This fungus causes a blight of the inflorescence of Setaria and
Pennisetum in South America (Spegazzini, 1885, Fungi Guar. Pugill.
I. n. 253). Numerous, small, black, stalked, stromata arise on spikelets
of the inflorescence. Gaumann and Dodge (1928) consider B. claviceps
as the most advanced species of the genus and place it next to Claviceps
in complexity. Claviceps differs in tjiat true sclerotia with thick rinds
ASCOMYCETES 55
are not formed. They say (p. 246), "In B. claviceps, widely dispersed
in the tropics on Setaria and Pennisetum, the spikelets are surrounded
as in Balansia diaderma but, in contrast to that species, the stromata
are sclerotic as in those of B. ambiens."
B. claviceps is reported by Weiss (1945) from this country and is,
therefore, included in this book.
It is inferred from Diehl's article that Ephelis mexicana is probably
the conidial stage of a species of Balansia (1930,a, p. 763). However,
Diehl says that no conidial stage of B. claviceps is known. E. mexicana,
it might be mentioned here, grows on various grasses in Mexico, Florida,
and the West Indies. Diehl had material on Cenchrus echinatus from
Lake City, Florida. He says, "The Ephelis fructification develops from
certain points beneath the black, laccate rind of the sclerotium as a
globular, hairy protrusion which opens, becoming discoid. The rim and
lower disk are an exciple covered with long hairs 3-4 ^ in diam. The
upper surface of the disk is a grayish layer of needle-shaped conidia
borne as a palisade upon conidiophores at first simple, then verticil-
liately branched. . . . When mature the fructifications are usually but
not always discoid." This pycnidium-like structure belongs perhaps in
the Excipulaceae although, as Diehl points out, except for the exciple,
the body resembles a sporodochiuna. '
Weber reported E. mexicana and a Balansia similar to B. hypoxylon
(Pk.) Atk. on Cenchrus echinatus near Gainesville, Florida (1924). He
briefly discussed the development of the perithecial stage.
Range: Tropical S. America and northward in the United States to Vir-

ginia; widely scattered in the tropics, Senegal.
References: Diehl (1930,a); Gaumann and Dodge (1928); Marchionatto
(1947,a); Saccardo (Sylloge, 1891, v. 9, p. 997); Spegazzini (1885); Weber
(1924); Weiss (1945).
Balansia clavula (Berk, and Curt.) Lloyd—Black Crust

Xylaria clavulus B. and C. N. Amer. F. No. 825
Gregarious, seriate, short-stalked, convex, stromata, 2-3 mm. diam.,
on culms, not varnished, rather firm.
On: Paspalum ciliatifolium. Michx., Fla., Va. Type was on putrescent culms
of a dead grass in Texas.
This inadequately described species is listed by Weiss as present on
Paspalum, but nothing is.known about its parasitic nature. Possibly it
does not warrant inclusion here. Lloyd (1924) examined the fragmen-
tary material of the type and stated that it appeared to be the only
true specimen of Balansia which he had seen from this country.
Another species evidently not a Balansia, described as Xylaria
graminicola Gerard (26th Kept. N. Y. State Museum, p. 85), is listed
by Ellis and Everhart (1887,6, p. 101; see also, 1892,6, p. 669) with the
following description: "Club slender, cylindrical, simple, 3-5 cm. tall,
at first greenish pulverulent, then blackish brown, roughened by the
prbminent, globose perithecia, tips sterile, acuminate, stems smooth,
straight or flexuous, brown, 1.5 cm. long, arising from a brown, felt-like
subiculum; perithecia up to 300 /x diarneter, afeci cylindrical, stipitate,
55-60 X 5 /*, with paraphyses; spores uniseriate, unequally elliptical,
5-10x3.5-4.0 ^ (.0004 X .0002 inches). Stalks about 2 inches-tall,
said to be parasitic on the roots of languishing grasses. Poughkeepsie,
New York." This species is not known to the writer but is included here
for reference. Along the same line, Xylaria rhizophila Cooke and Massee
was described on the roots of grasses (Grev. 22, p. 37) from Australia.
The stromata were clavate, 2-6 divided, 3-5 cm. tall, 5-7 mm. wide,
the asci were cylindrical, and the spores brown, 8-10 X 2-3 fj..
Range: Southern United States.
References: Ellis and Everhart (1892,6, p. 667); Lloyd (1924); Saccardo
(Sylloge, 1882, v. 1, p. 323); Weiss (1945).
Balansia hypoxylon (Pk.) Atk.—Black Choke

Epichloe hypoxylon Pk'.
Sclerotia formed in the fruiting axis of the ho.st, curved and irregu-
lar, 5-20 mm. or more in length, grayish or blackish; stromata black,
prominent, pulvinate or subhemispheric, 1-5 mm. diam., several spring-
ing from the same sclerotium, minutely roughened by the slightly pro--
trading perithecia; perithecia immersed, 100-150 X 175-250 jx, elongate;
asci cylindric, 150-200 X 4-5 fi, with a pedicel at the base as much as
20 ju long; spores filiform, 1 fi thick, at maturity breaking into segments
3-4 ju long. J
Conidial [Ephelis borealis E. and E.). Young sclerotium small,
whitish, grayish buff, to grayish green, surrounded by immature in-
florescence which it mummifies. Surface of young sclerotium composed
of palisade-like, swollen conidiophores with sterigmata-like apical pro-
jection bearing an ovate conidium..(amerospore). Young^sclerotiuni at
first gelatinous, later cartilaginous, but never horny (as *m E. mexi-
cana). Spores acicular, nearly straight, often bent in the~middle, hya-
line or yellowish, 15-25 X 0.75 /x (Ellis and Everhart, 1885, p. 86).
On: Danthonia compressa Austin, Me. to Tenn., Ohio—D. sericea Nutt.,
Ga.—D. spicata (L.) Beauv., Me. to Va., Jowa, Mo., N. Car., Ohio,
Tenn.—Stipa leucotricha Trin. rfnd Rupr., Tex.—S. pidchra Hitchc,
Calif.—S. viridula Trin. Tex. The range for the fungus on D. compressa
and D. spicata is taken from Weiss (1945) who has lumped the range of
the fungus on these two hosts together. Sinpe.the known geographical
ranges of the hosts do not coincide in all cage'sT'some interpreting of their
respective ranges was necessary.
Very little information is available on the prevalence (jr importance
of B. hypoxylon. It appears to be a sputhern form which sometimes
ASCOMYCETES 57
ranges north to Maine, Iowa, Ohio, and west to California. The type,
however, is from Sandlake, New York. Seaver (1911) gives the range
as Maine to South Carolina, Texas and Iowa. Gaumann and Dodge
(1928) consider this species more primitive than B. claviceps since it
does not produce stromata approaching the sclerotial condition of
Claviceps. Heald and Wolf (1912) report this blight as "Quite common
during the last part of April and the first part of May" on Stipa leuco-
6.—Balansia hypoxylon on Stipa leucotricha from Austin, Tex. (Photo

by F. D. Heald, State College of Washington.)
tricha (Fig. 6). "It destroys the entire spike, forming around the vas-
cular tissue of the spikelets a pseudosclerotium which is grayish or
bluish black on the outside, whitish within. The black pulvinate
stromata which contain the flask-shaped perithecia project promi-
nently from the sclcrotia."
- Diehl (1930,6) reported Ephelis-Yike conidia on Aristida glauca
(Nees) Walp. from Texas. The dwarfing of infected plants was ex-
treme and included the entire inflorescence. The individual spikelets
were uniformly dwarfed and the glumes were only % as long as normal
ones. The resultant inflorescence was scarcely recognizable as belonging
to the same species as healthy ones. The fungus mass in sterile florets
was sclerotioid and similar to Ephelis borealis E. and E. (1885, p. 86),
the conidial stage of B. hypoxylon as above noted. Mature sclerotia
were rarely over 2 mm. in length and varied in color from gray through
58 DISEASES OF CEREALS AND. GRASSES
brown t o black. Diehl mentions t h a t these' m a t u r e bodies differ from
sclerotia hitherto recognized in Ephelis in t h a t t h e entire central por-
tion of t h e Texas material is a cavity filledl with a mass of acicular
conidia of t h e t y p e found in a normal Ephelis fructification. These
conidia are of a dark greenish cast when in mass, b u t hyaline when
single, 20-23 X 1-2 ii., rarely u p t o 29 /x, long; borne on short conidio-
phores n o t exceeding 3 x 1 / ^ - Although not clearly demonstrated, Djehl
felt t h a t t h e spores would become 3-segmented.
Also we m a y note in passing t h a t Ephelis japonica P . Henn., t h e
conidial stage of Balansia sp. on Eriochloa polystachya H.B.K., is
known from P u e r t o Rico and m a y perhaps be linvolved in t h e complex
of this k i n d found in t h e southern United States.
Ephelis oryzae Sydow t h e conidial stage of Balansia oryzae occurs
in I n d i a and China, causing a disease called I-chu-hsiang or'incense
rod ( T a i and Siang, 1948).
Ellis and E v e r h a r t (1892,6) mention Hypocrea subviridis B . a n d C.
on dead grass leaves from South Carolina. T h e y quote from Têrkeley
and Curtis (Grevillea 4 : p . 1 5 ) : "Effused. Perithecia pale dull green,
tomentose, crowded, seated on a white mycelium. A curious species."
R a n g e : United States.
R e f e r e n c e s : Atkinson (1905); Diehl (1930,a,6); Ellis and Everhart (1885;
1892,6); Giiumann and Dodge (1928); Heald and Wolf (1912); Peck
(1875); Schwarze (1917); Seaver (1911).
B a l a n s i a pilulaeformis (Berk, and Curt.) Diehl '

Hypocrella hypoxylon v a r . pilulaeformis B. and C.
Dothidea pilulaeformis B . and (y. (Grev. 4 : p . 105)
Similar t o B. hypoxylon (Ellis a n d E y e r h a r t , 1892,b) b u t t h e

s t r o m a t a are tubercular or otherwise irregular.
On: Paspalum pubescens Muhl., Ga. !
Diehl (1921) hsted Hypoxylon pilaeforme Berk, and Curt, from the
Sandwich (Hawaiian) Islands as No. 31 of the fungi^coljected on the
Wilkes Expedition. He mentioned that in the 1874 edition by Berke-
ley and Curtis No. 31 of the Wilkes fungi is illustrated. However,
Ellis and Everhart (1892,&, p. 91) under a discussion of Hypocrella
hypoxylon (Pk.) list Dothidea pilulaeformis B. and C. and also
refer to "Var. pilulaeformis B. and C." in t^he discussion. Apparently,
therefore, B. pilulaeformis is derived from Dothidea pilulaeformis
and not H. pilaeforme.
Range: Georgia. /
References (including the above citations to^ypoxylon): Berkeley and
Curtis (1851, 1862, 1874); Diehl (1921) ,Êllis and Everhart (1892,6).
ASCOMYCETES 59
Balansia trinitensis Cooke and Massee—Char Fungus

Stroma globose, firmly, attached, 1.0-1.5 mm. diam., hard, black,
papillate, stipe erect, scaly, 2-4 mm. long, 5 mm. wide, black; peri-
thecia embedded in the periphery at the head of the stroma, crowded,
obovate, ostiole scarcely perforated; asci cylindrical, base attenuate;
120-130 X 9-10 iJL, 8-spored, spores filiform, continuous, flexuose, hya-
line, 90-100 X 1-5 fj, [Ephelis trinitensis Cooke and Massee).
On: Panicum sp., Guatemala.
This was collected as W. A. Kellerman 6079 on Sierra de Mico be-
tween Los Amates and Izabal at an altitude of 1200 feet in February,
1907. It was found in a moist ravine on a large grass which was
severely injured by the fungus.
Range: Guatemala, Trinidad.
References: Cooke and Massee (1889); Kellerman (1907).
Ceratostomella paradoxa (De Seynes) Dade—See Endoconidiophora

paradoxa (De Seynes) Davidson
Claviceps cinerea D. Griff.—Ergot on Hilaria

Sclerotia clavate, gradually tapering upward, straight, curved,
twisted, or contorted, 1.5-3 cm. in length by 1.75-2.5 mm. diam. at the
base, very viscid while developing, the base permanently invested by
the flowering glumes of the host, which are smooth, shining black, and
closely adherent; sclerotia somewhat reticulate above, dark gray to
lighter at apex. Stroma erect, erumpent with a cylindrical or usually
slightly fusiform, short, stout, almost white stalk and a subglobose
head usually slightly flattened below and overlapping the upper end of
the stalk, 1.75-2.75 mm. diam.; head light gray, almost smooth, viscid,
punctiform with small darker points indicating positions of perithecia.
Perithecia sunken, not projecting above stromatic mass, ovate to very
slightly pyriform, 190-225 X 60-90 ju. Asci 8-spofed, fasciculate, nar-
rowly cylindrical, slightly narrowed below into rather long, stout
pedicels, and slightly enlarged at attachment, rounded above, 135-
150 X 4-5 fi. No paraphyses. Spores nearly parallel, filiform, coarsely
but rather indistinctly guttulate, 100-120 X 1-1-5 M-
On: Hilaria cenchroides (Steud.) Nash, Ariz.—H. mutica (Buckl.) Benth.,
Ariz., N. Mex., Okla. -
Griffiths (1901) found this ergot in, great abundance in the vicinity
of Cochise, Arizona, in 1900. Some heads had as many as 20 sclerotia
in them. He planted sclerotia in building sand on October 26 and one
month later mature ascospores were formed, some having developed in
20 days. The long horn-like sclerotia are distinctive.
It is expected that this fungus is variable in its prevalence because
of the erratic rainfall of southern Arizona. The author traveled over
the same area in June, 1947, during an exceedingly dry period, and
found no evidence of ergot in Hilaria. Griffiths indicates in his discus-
sion that the occurrence of the fungus is dependent upon local rainfall.
j
Range: United States in southern Arizona, adjacent New Mexico, and prob-
ably Sonora, Mexico. Reported from Oklahoma.
Reference: Griffiths (1901).
Claviceps paspali Stev. and Hall—Ergot

Sclerotium paspali Seavers p.p.
Sclerotia yellowish to gray, globose, roughened when mature; about
3 mm. diam., one to several stromata per sclerotium; head of stroma
dull yellow; stipe short to medium, usually not more than 1 cm. long
(3-15 mm.); perithecia completely covering the head, numerous, ovoid,
340 X 119 /*; asci cylindric, 150-175 ju long; spores filiform, 101 X 0.5-
1^. /
On: Paspalum caespitosum Fliigge, Fla.—P. ciliatifoUnm^Michx., Fla., Ga.,
Va.—P. dilatatum PoiR, Alabama, Arlf., Fla., Ga., Hawaii, La., Md., Miss.,
N. Car., Okla., S. Car., Tex., Va.—P. distichum L., Miss., Okla., S. Car.,
Va.—P. floridanum Michx., Fla., Ga., N. Car., Va.—P. giganteum Baldw.,
Fla.—P. intermedium Munro, Va.—P. laeve Michx., Ark., Md., N. Car.,
Okla.—P. langei (Foum.) Nash, Va.—P. longipilum Nash, Va.—P. mono-
stachyum Vasey, Tex.—P. notatum Fliigge, Ariz., Fla., Ga., La.—P. oV-
biculare, Hawaii—P. •plicatulum Michx., Fla., La., Tex.—P. pubescem
Muhl., Va.—P. pubiflorum Rupr., Ga., Tex., Va.—P. quadrifarium Lam.,
Va.—P. supinuM Bosc, Va.—P. urvillei Steud., ¥\k., Md., N. Car., Okla.,
Tex. j '
This species has been reported as causing considerable damage in
the southern United States (Brown and Raiick, 1915). It has also been
reported as sometimes bothersome in South Africa. It occurs in New,
Zealand, where it is at' times very abundant. ^"^ -.^'
Lefebvre (1939) conducted artificial inoculation trials in the field
and greenhouse at Arlington Farm, Virginia, using 23 species and
strains of Paspalum. Although most of these were infected readily
enough, some were resistant. The following could not be infected! P.
notatum Flugge, P. lividum Trin., P. malaqophyllum Trin. and P.
supinum Bosc. Ergot however was seen in small quantities, on P. nota-
tum at Trfton, Georgia.
Rosen (1920) reported Paspalum floridanum Michx. and P. laeve
Michx. infected with ergot at Fayetteville, Arlânsas./He reported cor-
respondence with Dr. Stevens in which Stevens 'pointed out that it is
necessary to germinate sclerotia of C. paspali and C. roljsii in order to
distinguish the two species.
Hauman (1922)' described C. deliquescens on Paspalum, dilatatum
ASCOMYCETES g^
from Argentina. Soriano (1928) considered this species identical with
C. paspali. Another species, C. liitea A. Moller (Saccardo, Sylloge, v.
16, p. 609), on Paspalum sp. from Brazil was said by Barger (1931) to
be a distinct species. The pale yellow sclerotia of C. lutea have stipes
up to 4 cm. long. Shepherd reported the conidial stage of an ergot on
P. dilatatum and on P. maximum in Mauritius (1928). This was prob-
ably C. paspali.
Range: Ijnited States, Argentina, Australia, Brazil, China, Costa Rica, El
Salvador, Hawaii, Mauritius (?), Mexico, New Zealand, Panama Canal
Zone, Puerto Rico, South America.
References: Brown (1916); Brown and Ranck (1915); Burton (1945);
Burton and Lefebvre (1948); Hauman (1922); Lefebvre (1939); Rosen
(1920); Seaver (19U); Soriano (1928); Stevens and Hall (1910).
Claviceps purpurea (Fr.) Tul.—Common Ergot

Sclerotia 2-25 mm. long,_ horn-shaped^ pseudoparenchymatous,
purple black on surface, white in the interior, replacing ovaries; in the
spring of the year from 1 to 60 flesh-colored, stalked stromata develop
from each sclerotium, stromata 5-25 mm. long; flask-shaped perithecia
imbedded in the knob-like apex of the stromata, perithecia slightly pro-
truding at ostiolar end, 150-175X200-250 fj.; asci hyaline, slightly
curved, club-shaped with somewhat tapering ends, 100-125 X 4 /*,
paraphyses present; ascospores'8 per ascus, filiform, finally septate,
50-76 X 0.6-0.7 fi. Spores sometimes forcibly discharged, sometimes
exuded in a mucilaginous material. After infection through the open
flowering glume of the host, a whitish slimy mycelial mass replaces
ovary; sphacelial or conidial stage produces "honeydew" masses of
elliptical, aseptate spores 4-6 X 2-3 ju. Secondary infections develop
from conidia and secondary spores.
On: Agropyron albicans Scribn. and Sm., N. Dak.—A. caninum (L.) Beauv.,
Mich., N. Dak., Wash.—A. cristatmn (L.) Gaertn., Alta., B. C, Ind.,
Manit., Minn., Mont., Mich,, N. Dak., Ont., Oreg., Sask., S. Dak., Wash.,
Wyo.—A. dasystachyum (Hook.) Scribn., Alta., Ida., Manit., Mont.,
N. Dak., Wash.,_WyD.—A. desertorum (Fisch.) Schult., Manit., Mich.,
N. Dak., Wash.—A. gamelinii, N. Dak.—A. griffithsii Scribn. and Sm.,
Alta.-;-4. inerme (Scribn. and Sm.) Rydb., Mont., N. Dak., Oreg., Utah,
Wash.—A. intermedium (Host.) Beauv., N. Dak., Sask., S. Dak.—A.
junceum (L.) Beauv., Manit.—A. michnoilloshey, N. Dak., Sask.—
A. repem (L.) Beauv., Alta., Iowa, Kans., Manit., Mass., Micli., Minn.,
Mo., Mont., Nebr., N. B., N, Y., N. Dak., N. S., Ohio, Okla., Ont., Oreg.,
P. E. Is., Que., Sask., S. Dak., Wash., W. Va., Wise.—A. riparium Scribn.
and Sm., 'Ida., N. Dak., Wash.—A. semicostatum (Steud.) Nees, Mich.,
N. Dak., Wash.—A. sibiricum (Willd.) Beauv., Mich., Minn., Mont.,
N. Dak.—A. smithii Rydb., Alta., Ariz., Colo., Iowa, Manit., Mich., Minn.,
Mo., Montr, Nebr,, N. Mex., N. Dak., Okla., Sask., S. Dak., Tex., Wash.,
Wise, Wyo.—A. spicatum (Pursh) Scribn. and Sm., Ida., Iowa, Mont.,
N. Dak., Oreg., Wash., Wyo.—A. suhsecundum (Lk.) Hitchc, Ida., Iowa,
Manit., Mich., Minn., Mont., S. Dak., Uitah, Wash.—A. trachycaidum
(Lk.) Malte, Alta., Ariz., Ida., Iowa, Manitj, Mont., N. Mex., N. Dak.,
Oreg., Sask., S. Dak., Utah, Wise, Wyo.—A., trichophorum (Lk.) Richt.,
N. Dak., Wash.—Agrostis alba L., IlL, Iowa, Minn., N . Dak., Oreg.—
A. canina L., Que.—A. exarata Trin., Alaska^ Oreg.;—A. palustris Huds.,
Oreg.—A. scabra Willd., N. Dak., Okla., Wise,.—A. stolonifera L., N. S.—
A. tenuis Sibth., Oreg., Wash.—Alopecuris aequalis SoboL, N. S.—A. geni-
culatus L., N. Dak.—A. pratensis L., Mont., N. Dak., Oreg.—Ammo-
phila arenaria (L.) Lk., Wise.—Andropogon furcatus Muhl., Okla., Tex.,
Wise.—A. hallii Hack., Okla., Tex.—A. scoparius Michx., Okla.—Arc-
tophila fulva (Trin.) Ander., Alaska—Arrhenatherum elatius (L.) Beauv.,
Mo., Mont., Ohio, Oreg., Wise.—Averia hookeri Scribn., Sask.—A. sativa
L., Alta., B. C , Iowa, N. Dak., Wash., scattered in the East—Bouteloua
curtipendula (Michx.) Torr., Nebr., Okla.—B. gracilis (H. B. K.) Lag.,
Okla.—Brachiaria externa Chase, Fla.—Bromus carinatus'S.odk. and Arn.,
Ida., Ind., Mont., Oreg., Utah, Wash., Wyo.—B. ciliatm L., Mont., Wise.—
B. erectus Huds., Manit., Minn., Mont.—B. inermis Leyss, Alta., Ariz.,
B. C , Colo., D. C , Ida., III., Iowa, Manit., Md., Mich., Minn., Mo.,
Mont., Nebr., N. B., N. Dak., P. E. Is., S. Dak., Wash., Wise., Wyo., gen-
eral eastward—B. pumpellianus Scribn., Alta., Minn., Mont.—B. secalinus
L., Ind., Wash.—Calamagrostis canadensis (Michx.) Beauv., Alta., B. C ,
Iowa, Me., Manit., Mich., Minn., N. S., Oreg., -Wash., Wise—C. cana-
densis var. scabra (Presl) Hitchc, Alaska—C. inexpansa A. Gray> Manit.,
Mont., N. Dak., Sask.—C. neglecta (Ehrb.) Gaertn., N. Dak., Wise.—
C. nutkaensis (Presl) Steud., Alaska, Oreg.—Cinna arundinacea L., Wise.
—Dactylis glomerata L., Alaska, Ariz., Conn., Ky., Me., Manit.,. Mich.,
Minn., Mont., Okla., Oreg., Tenn., Wash., Wise, general in the lEast^—
Danthonia parryii Scribn., Colo.—D. spicata (L.) Beauv., Mich.—Des-
champsia caespitosa (L.) Beauv., Oreg.—Distichlis spicata (L.) Greene,
Md., Wash.—Elymus ambiguus Vasey and Scribn., Mont.—E. arenicold
Scribn. and Sm., Oreg.—E. canadensis L..' Alta., Ida., Iowa, Manit.,
Minn., M c , Nebr., N. Mex., N. Dak.,,'Okla., Oreg., S. Dak., Tex., Utah,
Wash., Wise, Wyo., general in the East—E. canadensis var. robustus
(Scribn. and Sm.) Mackenz. and Bush, Iowa—E. condensatus Presl,
Alta., B. C , Colo., Ida., Mont., N . Dak., Oreg., Utah, Wash., Wyo.—
E. curvatus Piper, Manit.—E. dahuricus Turcz.,>ti. Dak., Wash.—E. gi-
ganteus Va_hl, N. Dak., Wash.—E. glaucus Buckl., ~B~ (2., Calif., Ida.,
Oreg., Wash., Wyo.—E. innovatus Beal, Sask.—E. junceus Fisch., Mont.,
N. Dak., Wash.—E. macounii Vasey (hybrid), Manit., Mont., N. Dak.,
Wash.—E. mollis Trin., Alaska, Oreg.—E. pseudoagropyroft (Griseb.)
Trin., Mont.—E. salsuginosus Turcz., N. Dak.—E. triticoid'es Buckl.,
Cahf., Nev., Oreg., Wash.—E. virginicus /L., Iowa, Kans., Mich., Mo.,
Nebr., N. Dak., Okla., Tex., general eastivard—Erianthus alopecuroides
(L.) Ell., Alabama, Okla.—Festuca elatiorL., Me. to Va'., N. Dak., Okla.,
Oreg.,-Wash.—F. elatior var. arundinacea (Schreb.) Wimm., Ky., N. Dak.,
Oreg., Wash.—F. idahoensis Elmer, Ida., Mont., N. Dak., Oreg.—F. kingii
(^. Wats.) Cassidy, Colo., Wyo.—F. octcyflom Walt., Mich.—F. ovina L.,
Mo., Wise—F. rubra L., Alta., Ky., Minn.; N . Y., N. Dak., Ont., Oreg.,
Wash.—F. rubra var. commutata Gaud., Ky., N. Dak., Wash.—Glyceria
ASCOMYCETES 63
borecdis (Nash) Batchelder, Ida., Me., Mich., Que., Wise.—G. canadensis
(Michx.) Trin., Me.—G. grandis S. Wats., Iowa, Manit., N. Dak.—
G. pauciflora Presl, Alaska—G. septentrionalis Hitchc, Ind., Wise.—
G. striata (Lam.) Hitchc, Okla., Wash., Wise.—Hierochloa odorata (L.)
Beauv., N. Dak.—Holcus lanatus L., Oreg.—Hordeum jubatum L., Iowa,
Manit., Mont., N. Dak., Okla., Oreg., Wise.—H. jubatum var. caespito-
sum (Scribn.) Hitchc., Mont., Wash.—H. nodosum L., Alaska, Minn.,
Wash.—H. nodosum var. boreale (Scribn. and Sm.) Hitchc, Alaska, Colo.
—H. vulgare L., Alta., B. C, Colo., Ida., Manit., Minn., Mo., Mont.,
N. B., N. Dak., N. S., Okla., Ont., Oreg., P. E. Is., Que., Sask., Wash.,
Wyo., Yukon, general in the. East—Hystrix patula Moench, Ind., Iowa,
N. Dak.—Koeleria cristata (L.) Pars., Ariz., Ida., Iowa, Kans., Mo.,
Mont., N. Dak., Okla., Wise.—Lolium multiflorum Lam., Calif., Ky.,'
Okla., Oreg., Wash.—L. perenne L., B. C, Calif., Ida., Kans., Md., Mich.,
Minn., Nebr., N. Dak., Okla., Oreg., S. Dak., W. Va., Wise, Va.—L.
rigidum Gaud., Manit.—Phalaris arundinacea L., Alta., 111., Manit., Minn.,
Mont., N. Y., N. Dak., Okla., Que., Me. to Va., Wise—P. canariensis L.,
Alta., Mich., Minn.—Phleum pratense L., Alaska, Alta., B. C, Conn., and
scattered in the East, Iowa, Manit., Minn., N. Dak., Oreg., P. E. Is.,
Wise.—Phragmites communis Trin., Iowa, N. J., N. Dak., Okla.—Poa
ampla Merr., Ida., Mont., N. Dak., Wash.—P. annua L., Mo., Okla.—
P. arida Vasey, N. Dak.—P. canbyi (Scribn.) Piper., Ida., Manit., Mont.,
N. Dak., Wash., Wyo.—P. compressa L., Manit., Mo., Minn., Mont.,
N. Dak., Okla., Wash., Wisc^P.. fendlenana (Steud.) Vasey, Utah—
P. glaudfolia Scribn. and Williams, Manit.—P. juncifoUa Scribn., Oreg.—
P. longifolia, Trin., Utah—P. 'nemoralis L., Wash.—P. nervosa (Hook.)
Vasey, Wyo.—P. nevadensis Vasey, Mont., N. Dak., Oreg., Wash.—P.
palustris L., Minn., N. Dak., S. Dak.—P. pratensis L., Alta., Iowa, Ky.,
Manit., Minn., Mont., N. Y., N. Dak., Ohio, Okla., Penn., Utah, Wash.,
Wise, Wyo.—P. pratensis XP- arachnifera (hybrid). Mo.—P. scabrella
(Thurb.) Benth., Utah—P. secunda Presl, Ida., Wash.—P. stenantha
Trin., Wash.—Puccinellia distans (L.) Pari, Wash.—Secale cereale L.,
general in the U. S. and Canada, less prevalent in the drier parts of the
Far West—S. cereale y.S. montanum Guss. (hybrid), B. C, Mont., N.
Dak., S. Dak., Wash.—Setaria macrostachya H. B. K., N. Mex.—S. viri-
' dis (L.) Beauv., N. Mex.—Sitanion hystrix (Nutt.) J. G. Sm., Oreg.—
Sorghastrum nutans (L.) Nash, Kans., Okla., Tex.—Spartina alternijolia
Lois, Del., La., Me., Mass., Miss., N. J., N. Y.—S. cynosuroides (L.)
Roth., N. Y.—S.^ gracilis Trin., Sask.—S. patens (Ait.) Muhl., Md.—
S. pectinata Lk., Iowa, Mont., Nebr., Okla., S. Dak.—Stipa Columbiana
Macoun, Ida., Utah—S. columbiana var. nelsonii (Scribn.) Hitchc, Wash.
—S. comata Trin. and Rupr., N. Dak.—S. lettermanii Vasey, Wash.—S.
robusta Scribn., Mont,—S. spartea Trin., N^ Dak.—S. viridula Trin.,
Manit., Minn., N. Dak., Sask.—Tripsacum dactyloides L., Okla. (C trip-
sacil)—Trisetum canescens Buckl., Ida.—Triticum aestivum L., Alta.,
Ariz., B. C, Iowa, Kaiis.,'Manit., Minn., Mo., N. B., N. Y., N. Dak.,
N. S., Ohio, Okla., Oreg., Penn., P. E. Is., Que., Sask., Wise—Triticum
dicoecum Schrank, N. Dak.—T. durum Desf., Manit., N. Dak., Sask.—
Zizania aquatica L., Iowa, Me., Manit., Md., Minn., Nebr., N. B., N.
Dak., N. S., Ont., Wise—Z. aquatica var. angustijolia Hitchc, N. B.
Ergot is extremely widespread on many species of grasses. Our list
above includes 146 host names for C. purpurea. Barger (1931) lists
about 200 for all species for the world. The literature relative to ergot,
and the serious disease ergotism, is extfensive,, some of it dating back
long before plant pathology was a recognized, Science. While the cause
and prevention of ergotism in humans is now generally known, ergotism
in livestock is still a serious factor. It causes heavy losses in range
cattle and other livestock in the southern plains region of the "United
States (Chester and Lefebvre, 1942).
Fetch (1937) has determined that there is no, genetic difference be-
tween C. purpurea and C. microcepiiala (Wallr.) Tul. and that the
difference in sclerotial size reported for the two species is directly de-
pendent upon the size of the flowering parts attacked. Sclerotia of C.
purpurea range in size from the tiny ones produced on Agrostis tenuis
to the giant horns found on rye and on such grasses as Elymu^ con-
densatus Presl.
Stager (1903-23) distinguished several races of this fungus. This is
now recognized as one of the earlier classics on physiological specializa-
tion in fungi. Barger (1931) gave numbers to these races:
Race Pi—attacks rye {Secale), wheat, barley, Festuca elatior, Bromus
sterilis L., and four species of Poa. On Anthoxanthum, the conidial
stage only occurs.
P2—attacks Brachypodium sylvaticum only but the conidial stage oc-
curs on some others.
P3—attacks Lolium spp. and Bromus erectus. '
Mastenbroek and Oort (1941) added to this information when they
isolated race P4. This race was obtained from Festuca arundinacea,
Lolium perenne and Bromus erectus. In inoculation tests it was able to
attack the hosts attacked by both race 1 Pi and P3. Mastenbroek and
Oort also worked with inoculum from rye sent from Spain, the Nether-
lands, Poland, and Canada. These all Iwere readily identified as Pi
except the Canadian isolate. This was capable, of mildly attacking
Lolium.'They were'not certain as to whether thisBelonged in a new
race ("P5"), or a subrace of Pi. Mastenbroek and 0"ort, as well as
Stager, found Lolium and rye heavily infected in nature in close prox-
imity to one another. Possibly this race should be known as Pi-a.
Baldacci and Porlani (1948) report "P5" as a new strain attacking
wheat, durum wheat, T. turgidum, rye, barley, ryegrasses, and Agro-
pyron sp.
Stager also inoculated 17 species of grasses with C. wilsonii Cooke
and found that it could only attack Glyceria fluitari^ (1903). He also
found that C. sesleriae Stager could infect only îesleria coerulea Ard.,
another species of the same genus, and could- also produce the conidial
stage on Melica spp. 1
Stager also recognized two races of C. microcephala which modern
opinion would add to C. purpurea. One, which could be called Pe,
ASCOMYCETES 65
occurs on Phragmites, Aira, Molinia and Nardus, in Switzerland. The
other race which we call P7 appeared to be confined to Poa annua.
Along this same line McFarland (1921) cross inoculated rye and wheat
with conidia'from Bromus inermis, Agropyron repens, Poa pratensis,
and Arrhenatherum elatius. Wheat was occasionally infected by inocu-
lum from rye, Arrhenatherum, Poa pratensis, and A. repens. Rye was
more readily infected. An exhaustive study of the races involved on
all known hosts of C. purpurea would no doubt produce a maze of races
and subraces which would be far more complicated than that disclosed
by our present restricted knowledge of this species. Brentzel (1947)
recently outlined such a study. His article also contains practical, sum-
marized information on ergot and its control. The most useful readily
available summaries of this fungus are those by Atanasoff (1920) and
Barger (1931), and such texts as Heaid (1933), Wolf and Wolf (1947),
and Dickson (1947).
It Jias 'been the observation of the writer that in some cases involv-
ing severe losses the ergot never develops beyond the conidial stage
because of the fact that the particular hosts normally mature before
the ergot sclerotia do. Poa ampla, P. secunda, and Festuca rubra at
Mandan, North Dakota, were often almost devoid of germinable seed
because of ergot, and yet no ergot sclerotia were present in early July,
when the seeds of healthy grass were-.ripe. There were very few symp-
toms other than sterile heads and a general dusty appearance of the
heads, due to accumulation of pollen and dust in the honeydew. How-
ever a ready test for ergot is to taste the heads for the tell-tale, sweet,
honeydew flavor.
It was noted that after dusting Poa canbyi with gray "gas house"
sulfur for control of powdery mildew, a considerable increase in yield
was obtained in ihe treated plot while but little mildew developed in
the check. The honeydew stage of C. purpurea was prevalent in the
check plot but was greatly reduced in the treated plot. The difference
in yield was, therefore, attributed to this treatment in controlling ergot.
This work, done in the Nursery Division plots, Soil Conservation
Service, Mandan, North Dakota, was never repeated but the indicated
control may have some-^protective value in growing grass seed stock
where the conidial stage of ergot seriously reduces yield.
There are many papers dealing with the commercial culture of ergot
for medical use. Bekesy (1944) has develope,d^ a .small, inoculating
machine. He found that in Hungary there were two races of ergot, one
rich in alkaloids, one poor.
Range: United States, Algeria, Argentina, Australia, Austria, Belgium, Bul-
garia, Canada, China, 'Czechoslovakia, Denmark, England, Finland,
France, Germany, Greece, Hungary, Ireland, Italy, Japan, Mexico,
Netherlands, Norway, Palestine, Peru, Poland, Portugal, Romania, Soot-
land, South Africa, Spain, Sweden, Switzerland, Turkey, U.S S.R., Yugo-
slavia. General in temperate areas.
References: Anderson (1889,a); Atanasoff '(1920); Avizohar-Hershenson
and Plaut (1947); Baldacci and Forlani (1948); Barger (1931); Bekesy
(1938, 1944); Bondartzeff (1929); Bonns ( i t e ) ; Brpntzel (1947); Brown
(1948); Chester and Lefebvre (1942); Ducellier (1922); Forlani (1947);
Fyles (1915); Heald (1933); Hecke (1922, ll923); KiUian (1919); Kirch-
hoff (1929); Krebs (1936); Lepik (1935);' Lewis' (1945); Markhasseva
(1936); Mastenbroek and Oort (1941); McCrea, A. (1931); McFarland
(1921, 1922); Pammel, Weems, and Scribner (1901); Fetch (1937);
Sampson and Western (1941); Schweizer (1941); Seymour and McFar-
land (1921); Stager (1900, 1903, 1905, 1907, 1908, 1910, 1912, 1922,
1923); Steinmetz and Wright (1943); StoII and Brack (1944); Tanret
(1922); Vincens (1917); Weniger'(1924); Whetzel and Reddick (1911);
Wolf and Wolf (1947).
Claviceps ranunculoides A. Moll.—Ergot

Sclerotia horny, oblong horn-shaped, curved, blue black, conidia
7-8 X 3-4 jx, borne on conidiophores,' 12 X 2 /A; stromata stipitate, yel-
lowish; perithecia almost free in the stromata, 400-500 jw long; asci
300 X 4 /x; ascospores 160 /* long. '
On: Setaria macrostachya H. B. K., Ariz., N. Max.,
This species was originally described on Setaria sp. from southern
Brazil. It is listed as distinct by Barger (1931). He says that the
perithecia are almost free, so that the capitulum recalls the syncarp
of Ranunculus. Both Setaria macrostachya and S. viridis are listed as
hosts for C. purpurea from New Mexico and no doubt these collections
should be checked again.
Range: United States, Brazil. /
References: Barger (1931, p. 109); MoUer (1901).
I
Claviceps rolfsii Stevens and Hall—Ergot
Sclerotium rolfsii Seayer p.p.
Sclerotia yellowish to graTy, globose, ro'ughened whên mature, about
3 mm. diam., head dull yellow; stipe fihform, 1-1.5 cm: long; perithecia
few in head and mostly upon the extreme distal portion, cylindric-
ovate, 8 1 6 X 2 2 5 m; asci cylindric, 3 7 5 X 3 /x; spores filiform, 260-
275 X 0.5-1.0 /x. / /
On: Paspalnm laeve Michx., N. Car. ,
This species is said to have much longer spores than C. paspali. It
has not received very much attention because ihost of the material
which has been studied is readily assignable to C. paspali. While Wolf
and Wolf (1947) found considerable variation in the size of different
collections of sclerotia on Paspalum laeve, P. dilatatum, and P. florida-
num in North Carolina, they assigned all of their • collections to C.
ASCOMYCETES 67
paspali on the basis of perithecia and contents. It would appear there-
fore that the status of C. rolfsii is in some doubt.
Range: United States, reported from North Carolina only.
References: Seaver (1911); Stevens and Hall (1910); Wolf and Wolf
(1947).
Claviceps tripsaci Stevens and Hall—Ergot

Sclerotia smooth, white to dark brown or black, nearly conical,
4-5 mm. in diam. at the base; heads gray to grayish white; stipe thick,
white to purplish white, 1-1.5 cm. long; perithecia numerous, ellipsoid
in longitudinal section, with a short beak toward the surface of the
head, 390 X 153-187 yn; asci cylindrical, 145-175 X 2-3 [i.; ascospores
filiform, 130 /* long.
On5 Tripsacum dactyloides L., Ga., Md., Miss., N. Car., Va.
Information on this species is scanty. It differs from C. purpurea in
that it appears to have longer perithecia with proportionately longer
ascospores. A conidial stage once recognized is now considei'ed an asso-
ciated species of Fusarium.
Range: United States in the southern coastal region.
References: Seaver (1911); Stevens and Hall (1910).
Claviceps yanagawaensis Togashi—Ergot

Sclerotia 2.5-15 mm. X 0.5-1 mm. wide, flattened, usually slightly
curved, and for the most part grayish-violet, host glumes remain at-
tached to sclerotia; stroma one, or rarely two stromata, stipes 4-16
mm. long and 0.3-0.5 mm. wide, heads 0.3-1.0x0.4-1.5 mm.; peri-
thecia 180-320 X 70-190 fi.; asci 85-165 X 4.0-8.0 fi. and ascospores
75-1.35 X 1.0-2.25 p.. ,^
On: Zoysia japonica Steud., on imported seed in U.S.D.A. laboratory, Wash-
ington, D. C.
This species probably does not belong in a discussion of the fungi
on Gramineae in the United States because it has not been reported
as growing naturally in North America. However, Lefebvre reported it
on ergot-infested seed intercepted in a shipment from Japan (1942).
For that reason it is included as it is likely to occur in the United
States.
Lefebvre (1942) was unable to infect rye hybrid with inoculum
of C. yanagawaensis. He killed the sclerotia by treating the seed in
75 per cent sulfuric acid for 20 to 30 minutes.
Range: Japan, intercepted in the United States on seed imported from
Japan.
References: Lefebvre (1942); Togashi (1936).
Cochliobolus heterostrophus Drechsl.—Leaf Spot

Ophiobolus heterostrophus Drechsl.
Helminthosporium niaydis Nisik. and Miyake
Causing, on leaves, numerous dead cinnamon-buff or purplish areas
surrounded by a darker reddish brown riiargin, and often'delicately
variegated with brownish zonate bands; the lesions longitudinally
elongated, first elliptical, later long-rectangular, typically limited to a
single intervascular region, usually 1-3 X 5-15 mm., often coalescing
to form more extensive dead portions. Perithecia developing on dis-
integrating host tissues, usually early erumpent, black, often bearing
a variable number of conidiophores, but no differentiated sterile setae;
ascigerous portion subglobose or more frequently somewhat ellipsoidal,
0.4-0.6 mm. transverse diam., approximately 0.4 mm. tall; ostiolate
beak well defined, subconical or paraboloid, approximately 0.15 mm.
diam. at base and 0.15 mm. tall; interior composed of colorless pseudo-
parenchymatous tissues consisting of vertically oriented appressed fila-
ments, diminishing usually with ythe development of the asci. Asci
numerous, short-stipitate, with rounded apex, subcylindrical but some-
times becoming more inflated before discharge,; 160-180 X 24-28 /x,
containing 1-4, rarely 8 spores,'^but typically 4 spores. Ascospores fila-
mentous, fuliginous, thin-walled; in somewhat immature condition, of
uniform diameter of 6-7 /<., except at extremities which are somewhat
attenuated; later becoming 5-9 septate, the septa usually associated
with perceptible constrictions, the delimited segments becomin'g some-
what swollen so as to attain in places a diameter up to 9 /x; thrusting
firmly against apex and into base of stipe in multiple, heterostrophic,
helicoid arrangement with approximately/ four turns to each spore
(corkscrew arrangement); 130-340 /j. long( discharged simultaneously,
often with mucous envelope; germinating promptly by producing in-
discriminately from any or all segments, either laterally or terminally,
germ tubes up to 8 in number, from 3.5-5 fi. diam.
Conidiophores arising singly or in groups ol.2_or 3'from stomata in
center of killed ifoliar parts; olivaceous, septate atTiniervals of 15 to
60 /x; bearing the first conidium after attaining a length of 50 ;u. or
more; points of attachment of successive spores marked by scars oc-
curring at intervals from 10 to 40 /x ^t geniculatibns not always pro-
nounced; in nature measuring usually 120-170 ju. in length, but under
moist conditions occurring as irregularly branched sporophoric fila-
ments, often exceeding 1 mm. in length. Conidia developed at 25° C.
on diseased maize leaves in damp chambers or -in pure culture on arti-
ficial media, fuliginous to light olivaceous, 30-J.15 X 10-17 ju.; often
strongly curved, usually widest near thejniddle and tapering percep-
tibly toward the rounded ends; periphâl wall thin, especially in the
apical and basal regions; basal scar broad, not conspicuous, contained
within rounded contour; germinating promptly by the production of
two polar gerni tubes.
ASCOMYCETES 69
On: Zea mays L., Fla., Ga., Md., Miss., N. Car., Penn., S. Car., Tenn., Tex.,
Va.
Drechsler (1934,6) placed the Ofhioholus-Vik.e forms with corkscrew
ascospore arrangement and bipolar-germinating Helininthosporium
spores (conidial) in his new genus Cochliobolus. This is described in
detail and well illustrated in his earlier article (1925,6). H. maydis, the
conidial stage, is an important fungus in warm regions such as the
southern United States, Japan, and the Philippines. Ullstrup once as-
signed a fungus from the Middle West to H. -maydis (1941,a) but later
made a new species of this fungus which he called H. carbonum (1944).
H. maydis has regularly curved, almost lunate spores. Ullstrup found
that both his fungus and H. maydis produced light, straw-colored spots
on leaves with a tendency towards zonation but the lesions caused by
H. maydis had a purplish brown margin which was absent in H.
carhonum.
H. maydis shows some resemblance to species.of Helminthosporium
on rice and sugar cane but Drechsler found that these hosts were
scarcely or not at all attacked by the fungus from corn (1925,5).
Drechsler also illustrated H. cwvulum as possibly related to the corn
disease but the material on tassels from Malay belongs to Curvidaria
(Groves and Skolko, 1945, pp. 99-100).
Range: United States, China, Gold Coast, Japan, N. Caucasus, Philippines.
References: Boning (1938); Drechsler (1925,f); 1927,b; 1934,6) Nisikado
(1927,a); Nisikado and Miyake (1926,a); Young, George Y. (1943).
Cochliobolus miyabeanus (Ito and Kuribayashi) fide Dickson (1947)

Ophiobolus miyabeanus Ito and Kuribayashi
See Helminthosporium oryzae V. Breda de Haan for a discussion of
this fungus. For a summary of the literature on it see Motte (1947).
Cochliobolus sativus (Ito and Kuribayashi) Drechsl. fide Dickson

(1947)
Ophiobolus sativus Ito and Kuribayashi
C. tritici Dastur
See Helminthosporium sativum P.K.B. for a discussion- of this
fungus. Drechsler (1934,6) inferred that the species of Ophiobolus with
spores spirally coiled in the ascus and with a species of bipolar-germi-
nating Helminthosporiy,m as"the conidial stage, should all be placed in
his newly creaited genus Cochliobolus. Except for C. heterostrophus he
did not actually make the changes. Dickson (1947) credited some of
these to Drechsler. One, C. miyabeanus, Dickson has credited to Ito
and Kuribayashi although the latest pubUshed paper (1931) which
Dickson cited for these authors was published 3 years before the genus
I
Cochliobolus was created. This combination should, perhaps, be cred-

ited to Dickson, the others to Drechsler fide Dickson.
Cochliobolus setariae (Ito and KuribayasM) Drechsl. fide

Dickson (1947) I
Ophiobolus setariae Ito and Kuribayashi
See Helminthosporium setariae Sawada for a discussion of this
fungus.
Cryptoascus sp.—On Roots
Perithecia develop in clusters on a limited stroma with the spores
lying free in the cavity of the perithecium. In pure culture the few
perithecia which were formed produced small evanescent asci usually
containing two spores, sometimes one or three.
On: Avena sativa L., P. E. Is.—Hordeum vvlgare L., P. E. Is.—Triticum
aestivum L., P. E. Is. /
Whether this fungus is parasitic has not been proved. The material,
first found at Malpeque, Prince Edward Island, by R.'R. Hurst on July
29, 1936, was so heavily rusted that the stunting which prevailed may
have been due to the rust. The fungus was associated with a root
necrosis, usually of wheat and has been found a number of times in
the same general region of Prince Edward Island.
CryptoUscus Petri is a primitive genus of the Sphaeriales with col-
ored,-rionseptate spores. I have never encountered any fungus similar
to %hi^/'bne. It is apparently undescribed. ,
Microasciis trigonosporus Emmons andj Docjge occurs in the seed
coat of various plants including cereals in the United States. It causes
little or no injury (Whitehead, Thirumalachar, and Dickson in Phyto-
pathology 48:968-973. 1948).
Range: Known only frqm Prince Edward Is. ^..^^^^
References: Conners (in 17th Ann. Kept. Canad. PL I)is7~Snnvey, 1937,
pp. 5-6 and subsequent reports).
Dothichloe Atkinson
This genus is said to differ from Balansia infnot possessing'-a pseudo-
sclerotium composed of host and fungus tissue (Chardon, 1921). This
means, in part at least, that the stroma is distinct and more or less
superficial. The fungus has been placed in both the Hypocreales
(Stevens, 1913; Theissen and Sydow, 1915) and in, the Dothideales
(Chardon, 1921). •^ '
Diehl (1930,a) distinguished the two genera Balansia and Dothi-
chloe on the basis of the form and position of the ascogenous fructi-
fication. The stroma of Balansia is capitate and stipitate' and that of
Dothichloe is effuse. ,
ASCOMYCETES 71
Dothichloe aristidae Atk.—Black Choke

Stroma dimorphic, sterile portion confluent, forming a thin black
layer, in the specimens seen entirely surrounding the culm. Fertile por-
tion much thicker, confluent or interrupted, forming small perpendicular
elevations on the sterile portion, projecting apices of the crowded peri-
thecia more or less confluent in an irregular manner giving a rugulose
or convolute appearance to the stroma.
On: Aristida purpurascens Poir, Alabama—A. stricta Michx., Ga.
This appears to be another of those vague fungi which may be a
legitimate species, if we can judge by the literature, but which do not
seem to even have a relatively usable technical description available.
Earle (1900) placed this fungus in Ophiodothis. The stroma completely
surrounds the culm, hence the common name of black choke.
Dothichloe hemicrypta Diehl is listed on Aristida from Texas
(Weiss, 1943).
Range: Southern United States, Puerto Rico.
References: Atkinson (1894); Chardon (1921); Diehl (1930,a); Earle
(1900).
Dothichloe limitata Diehl—Leaf Disease
Hypostroma effuse, as a flat, thin, white, prosenchymatous layer or
film becoming black with age, on adaxial surfaces of unrolling leaves,
rarely exceeding 15 /A in thickness (vertical diam.) and 200 mm. hori-
zontal diam., rarely separated into smaller units. Scolecosporous coni-
dial fructification ephemeral, white to gray, covering the surface of the
hypothallus wjith a palisade of conidia upon short, simple conidiophores,
rarely swollen and branched; conidia 18-30 X 1-2 /x. Amerosporous
conidial fructifications developing under conditions of extreme mois-
ture upon the immature ascostroma as a layer of simple to spatulate
conidiophores, each with an apical sterigmata bearing a single conidium;
conidia obovate, up to 4 X 3 /*. Ascostroma uniformly effuse, maculi-
form, usually 10-15 mm', in horizontal diameter, rarely exceeding 200 /x
in thickness, sometimes separated into pulvinuli by sterile areas; sur-
face uniform to rugulose, black, punctate from the slightly emergent
ostioles; context white; perithecia not crowded, ovate_to lageniform,
180-315 X 80-130 /x, ostioles short; perithecial walls 6-12 /i thick; asci
105-160 X 4-6 iJ.; ascospores 90-120 X 1-1-5 fi.
On: Andropogon scoparius Michx., Va.—A. virginicus L., Ga.—A. sp., Mo.—
Calamagrostis canadensis (Michx.) Beauv., Minn., Wise.—C. inexpansa
A. Gray, N. Dak.—Chloris petrae Swartz, F!a.—C. virgata Swartz, Fla.—
C. sp., Mex.—Ctenium aromaticum (Walt.) Wood, N. Car.—Eragrostis
capillaris (L.) Nees, S. Car.—E. Ursula (Michx.) Nees, Ga.—E. refracta
(Muhl.) Scribn., Fla., Ga., Va.—E. sp., Fla., S. Car., Gymnopogon am-
biguus (Michx.) B. S. P., Ga., N. Car., Va.—Oryzopsis hymenoides (Roem.
and Schult.) Rick,, Penn.—Panicum agrostoides Spreng., Miss., Mo,—
P. meridionale Ashe, N. Y.—P. virgatum L., E^.—P.. sp.. Miss.—Pas-
palum conjugatum Bergius, Va,—Sporobolus poireiii (Roem. and Schult.)
Hitohc, Alabama, Ark., Fla., Ga., La!, Miss., N'., Car., Va.
Diehl (1939) reports that this fungus is systemic. I t does not cause
leaf discoloration and would not be considered to be parasitic were it
not that infected plants often failed to produce flowers.
The symptoms of this disease are somewhat obscure. The fructifi-
cations are first white. The conidial stage is prpminent only during
periods of moist weather but the mature black ascostromata are more
conspicuous, especially after the leaf has inrolled, leaving the area
adjacent to the stroma still flattened. The black stromata later slough
off with little trace left of them on the leaf.
Diehl (1939) suggested that low temperature inhibited the fructi-
fications of the parasite and thus limited its northward spread.
Range: Southeastern United States, Cuba,'Pvierto Rico, the West Indies,
and south to Brazil, north in the United States sometimes to North
Dakota. '
References: Chardon (1921, PL 14, Fig. 12); Diehl (1930, a. PL 2, D, E,
and Fig. 3, B; 1939, PL 1); Earle (1900); Weese (1919).
Dothichloe nigricans (Speg.) Chardon—Black Ring

Epichloe nigricans Speg.
Black stroma completely surrounds internode of culm, short, 5-10
mm. long; perithecia immersed or locule-like, elliptical ovate, base
somewhat truncate, 140-150 X 50-80 /x, ostiolat'e; asci cylindrical to
subfusoid-cylindrical, 9 0 - 1 0 0 x 4 - 5 /x; without'paraphyses, 8-spored,
sporidia fascicled, filiform, 70-80 X 0.6-0.8j/t, septate, straight, hyaline.
On: Axonopiis affinis Chase, Fla.—Bovteloua curtipendula (Michx.) Torr.,
Kans.—Panicum aciculare Desv,, Ga.—P. clandestinum L., N. Y., Penn.—
P. commutatum Schult., Fla.—P. hemitomym Schuit.7'Fla.::^P. scoparium
Lam.,'Ga.—P. scribnerianum Nash, Nebr.—P. sp., Va. , ''
This tropical species has been reported from as far west and north
as Nebraska (Weiss, 1945, p. 331). The salient points of Spegazzini's
lengthy description are given above. Chardon transferred this to
Dothichloe, rejecting -D. nigricans (Speg.) Seaver as being'a distinct
species which he called D. subnodosa Chardon <(Chardon,.I921).
Range: In the United States as far north as New York and Nebraska, south
to Brazil, Puerto Rico, and the West Indies in generalf^
References: Chardon (1921, 1927); Saccardo (Sylloge, 1891, v. 9, p. 1001);
Spegazzini (1885); Theissen and Sydow (1915T; Weiss (1945, p. 331).
ASCOMYCETES 73
Dothidella aristidae (Schw.) Ell. and Ev.—Char Spot
Sphaeria aristidae Schw.
Phyllachora aristidae (Schw.) Sacc.
Euryachora{l) aristidae (Schw.) Theiss. and Sydow
Stroma effused, thin, black, interruptedly continuous over both sides
of the leaves and enveloping the culms for several cm. in extent, causing
them to appear charred. Ascigerous cavities small, mostly less than
75 /It diam., subglobose and seriate between the parallel nerves of the
leaves. Asci subglob.ose, 15-25 /x diam., sessile, 8-spored. Spores in-
ordinate, subelliptical, or ovate-oblong, obtuse, hyaline, uniseptate,
8-12 X 5-6 ;a.
On: Aristida dichotoma Michx., N. Car.—A. purpurascens Poir, Miss.—A.
stricta Michx. (listed in Seymour, 1929)—Distichlis spicata (L.) Greene,
Gahf., Oreg.—D. 'stricta (Torr.) A. Gray, N. Dak., Greg., S. Dak., Tex.,
" Utah, Wash.
The prominent "black char spot" is very common on salt grass in
eastern Oregon and \¥ashington. Ellis and Everhart (1892,b) mention
{hat the Oregon material is mature. Much of that collected in recent
years is not mature. As pointed out by the writer (1941,a), the issue
is considerably confused by similarity in symptoms to the advanced
lesions caused by Septogloeum oxysporum Sacc, Bomm., and Rouss.
At this time the char spot oh Distichlis and Aristida is not recognized
as the same species which occurs on Agrostis, Calamagrostis, and
Elymus spp. This situation will be discussed in more detail under
Septogloeum oxysporum of the Fungi Imperfecta
D. aristidae appears to be virtually saprophytic while Septogloeum
oxysporum is actively parasitic.
References: Ellis and Everhart (1892,&); Sprague (1941,a); Theissen and
Sydow (1915).
Dothidella minima Sacc. and Sydow—Leaf Spot

Stroma sparse to more or less gregarious, mostly hypophyllous,
small, elliptical, up to 500 X 250 /x, innately erumpent, applanate,
irregularly oblong, black, locules numerous, globose, 45-60 ju, diam.;
asci terete-clavate, apex rounded, briefly nodulbse-stipitate, 45-
50 X 8.5-9 /x, •8-spored, without paraphyses, spores one- or two-seried,
fusoid, oblong, 11-12 X 2.5-3.0 jx, 1-septate in center, nonconstricted,
hyaline.
On: Arundinaria gigantea (Walt.) Chapm., La.
Range: La.
References: Saccardo (Sylloge, 1905, v. 17, p. 851); Saccardo and Sydow
(1904); Theissen and Sydow (1915, p. 316).
• I
Endoconidiophora adiposa (Sartoris) Davidson—Black Rot of

Sugar Cane "^ ~ I
Sphaeronema adiposum Butl.
Ceratostomella adiposa (Butl.) Sartoris
Dematium scahridum Oilman and Abbott (R.A.M..9:612)
Causes a black rot or mold of the seed pieces of cane stalks and of
old canes, discoloring interior pith in reddish streaks, the pith later
becoming muddy brown color and drying. Discolored pith filled with
grayish brown mycelium; conidia formed on exposed surface or in
cavities in the pith, formed on the end of short, special, lateral branches
(conidiophores), of which two or three may be united at the base, more
frequently single. Conidiophores septate, usually composed of a row of
2 or 3 cells, the apical being the longest and bearing the spores; endo-
spores emerge from the apical cell, being cut off internally; endoconidia
unicellular, extremely variable in size and shape, ranging from elon-
gated, smooth, hyaline spores, to large, round, deep brown, spiny ones.
The hyaline type are usually formed first and/the brown, larger, spiny
ones later. The spiny spores emerge with a thick, jelly-like wall which
darkens and hardens rapidly into radial bands of spines; endoconidia
9-25 X 4.5-18 fi, the smaller being hyaline. Perithecia initiated by a
coil of mycelium, at first spherical, later with a very long, hollow,
bristle-like neck up to 50 /J. wide and as much as 600 /* long, with a
short fringe of hairs or appendages around the neck, basal spherical
portion of the perithecium sometimes 200-270 /A; asci cylindrical to
pyriform, 10-20 X 12-25 fi., somewhat evanescent, ascospores--thickly
falcate or angled, 3.5-6.5 X 4-8 ju..
On: Saccharum ojficinarum L., La.
Sartoris (1927) reported that the Sphaeronema stage discussed by
Butler (1918) was in actuality a member of the Sphaeriales, the long-
necked fruiting bodies being perithecia, not pycnidia. Davidson (1935)
placed all Ceratostomella with endospores in Endoconidiophora. Weiss
accepts the transfer, citing the fungus as E. adiposum (Sartoris)-D.^vid-
son (1945). Wiltshire (1947) however prefers Ceratostomella. ' _
The causal organism is a weak parasite or mold which does exten-
sive damage to seed pieces under certain conditions. Ideal conditions
for the fungus, according to Sartoris (1927) are frequent rains, with
intermittent periods of warm weather, and loosely p'acked, lumpy soil,
which allows air pockets to exist around the seed pieces. He cited one
case where the stand was reduced to 20 per cent of normal under such
conditions. The pieces that escaped the attack of the fungus.produced
normal appearing stools. Butler (1918) reported that-the fungus was
a wound parasite of a low order, active when -tfte^stalks were near
maturity and, as in Louisiana, was of greatest importance as a mold
of seed pieces. The heaviest damage was done to setts (seed pieces) left
in earth pits for about a iprtnight before planting. '
ASCOMYCETES 75
Ascospores are protruded in a foamy substance of a fatty nature
which is expelled at the ostiole by hydrostatic action. Sartoris (1927)
gives details of the cytology of the fungus.
Apparently E. adiposa is less widely recognized than is the pine-
apple rot fungus, E. paradoxa. In recent reviews (R.A.M.) the latter is
frequently mentioned in connection with cane sett treatment while E.
adiposa is not reported.
Range: United States, India, widely scattered in the tropics. •
References: Butler (1906, 1918); Davidson (1935); Sartoris (1927).
Endoconidiophora paradoxa (De Seynes) Davidson—Pineapple Rot

Sporoschisma paradoxum De Seynes
Ceratostomella paradoxa (De Seynes) Dade
Thielaviopsis paradoxa (De Seynes) v. Hoehnel
T. ethaceticus Went
Mycelium in the interior of culms at first hyaline, later becoming
brown or nearly black, coarse, intracellular, penetrating the cell walls
by narrow pegs, sometimes forming haustoria-like swellings. Conidia of
two types :^Macroconidia black, thick-walled, oval-elliptical, produced
in a chain by septation at the tip of hyphae, microconidia endogenous,
short-cylindrical, or oval, in chains. Perithecia flask-shaped, light col-
ored at .base, 200-350 ju,, immersed or nearly so, with a black shining
beak, 800-1200 X 30-40 fx. with a fimbriate apex; asci clavate and stipi-
tate, spores irregularly biseriate, or later indefinitely arranged, ellipti-
cal, frequently more convex on one side, 7-10 X 2.5-4 /A.
On: Saccharum officinarum L., Hawaii, La.
The name pineapple rot is derived from the fruity odor of freshly
split, diseased canes. Later, after this name had been given to the
fungus, it was found that it actually attacked pineapple. The name,
therefore, has double significance.
The fungus causes a blackening of the core of the culm. The black
area may be surrounded by reddened cells. The fungus is primarily
serious as a decay of ripe cane and of seedling cuttings (setts). Nowell
(1923) discusses practical control which includes use of healthy stock
and dipping in well-agitated Bordeaux mixture. Wiehe (1947) used
aretan with good results. Edwards, W. H. (1927) reported from Mauri-
tius that treating the setts with Burgundy mixture reduced the inci-
dence of pre- and post-germinative mortality by 46 and 58.3 per cent
respectively. Bordeaux mixture was given as an alternate remedy.
McMartin (1944) in South Africa obtained good control by dipping ends
of cuttings in either Agrosan, Ceresan, or Thiosan. Next year (1945,b)
he obtained good control with Ceresan and Agrosan and excellent con-
trol with 1 per cent Abavit. He also (1945,a) found that application of
fertilizers to the soil increased the yield only 3 per cent over that of
the check. That treated with Ceresan showed 124 per cent increased
yield, while Ceresan plus fertilizer yielded 31 |per cent higher than the
checks. Agrosan alone yielded an 18 per cent increase, but Agrosan plus
fertilizer gave a 43 per cent increase in yield. •
This fungus enters through wounds, leaf scars, and insect borings.
Edgerton and Moreland (1921) reported this species as one of a number
causing rot of cane seed stock in Louisiana where, at the northern limit
of sugar cane growing, the host suffers appreciably from rots, especially
in cool periods. M. T. Cook (1932,a) reported this fungus as causing
an important rot of seed cane in cold, wet, clay soil.
Range:_ United States, Brazil, British Guiana, Colombia, Egypt, Gold Coast,
Java, Mauritius, Natal, Philippines, Puerto Rico, St. Vincent Is., S. Africa,
Sumatra.
References: Averna-Sacca (1933); Bovell, J. R. (1921); Cook, M. T.
(1932,a,d); Dade (1928); Davidson (1935); Edgerton and Moreland
(1921); Edwards, W. H. (1927); Evans and Wiehe (1947); McMartin
(1944; 1945,a,6); Wiehe (1947).
Endodothella tracyi (Ell. and Ev.) Theiss. and Syd.—Tar Spot

Phyllachora tracyi Ell. and Ev..
Ascigerous cavities small, 35 n, in flat, black, char-spot, subasto-
mous, subglobose, of fine cellular texture, fused together in a thin
stroma either continuous in patches extending nearly or quite across
the leaf and reaching longitudinally for 0.25-0.5 cm. or forming longi-
tudinal series interrupted at frequent intervals and extending along the
entire length of the leaf; asci subglobose, 15-20 /x diam.; spores crowded,
8 in an ascus, subelliptical or ovate-oblong, obtuse,
tus hyaline, 1-septate,
8-12 X 5-6 IX. J
On: Distichlis spicata _(L.) Greene, Greg.—;D. stricta (Torr.) Rydb., N.
Dak., Okla.
Ellis and Everhart (1888,6, p. 63) state' that this has much the same
appearances as what'we list under Dothidella aristidae^W. and Ev.
The fungus differs from Phyllachora in the 1-septate spores. Theissen
and Sydow place it in the genus Endodothella (1915, p. 583) but
Clements and Shear (1931) place this genus under Placostroma Theis-
sen and Sydow (1915, p. 407). Whether the combination Placostroma
tracyi (Ell. and Ev.) has been made and whether it is justified must
await work by qualified students of this group.
Theissen and Sydow (1915, p. 583) also list Phyllachora diplocarpa
Ell. and Ev., Homostegia diplocarpa Ell. and Ev. and pothidella tracyi
Sacc. as synonyms for their E. tracyi. HoweverJDrton gives Phyllachora
diplocarpa as a good species of Phyllachora (1944). One cannot but
wonder about the authenticity of some of the various char-spots which
were studied by Theissen and Sydow. ,
ASCOMYCETES 77
Range: Western United States.
References: Ellis and Everhart (1888,6, p. 63); Theissen and Sydow (1915,
p. 583).
Epichloe typhina (Fr.) Tul—Choke (Cat-Tail Disease)

{Typhodium typhinum (Fries) Seaver)
Stromata cream-colored, later dark orange, surrounding sheaths or
culms of living grasses, 2-5 cm. Jong, conidia 3-9 X 1-3 /t, ovoid, hya-
line; followed by perithecia, soft, carnose-membranaceous, yellow, 300-
600 II high, 250 ju, diam., ostiole prominent, immersed, somewhat elon-
gate; asci cylindrical, 150-230 X 6-9 /x; spores 8 in an ascus, filiform,
yellowish, nearly as long as the asci, 1.5-2 /^ diam., septate at intervals
of 8-12 /*.
On: Agropyron inerme (Scribn. and Sm.) Rydb., Ida.—A. smithii Rydb.,
Ariz., Mont., N. Dak.—A. spicatum (Pursh) Scribn. and Sm., Mont.,
Wash.—A. trachycavlum (Lk.) Malte, Mont., N. Dak., Wash.—Agrostis
alba L., Que.—A. scabra Willd., Wise.—Andropogon furcatus Muhl., Wise.
—A. scoparius Michx., Wise.—Bouteloua gracilis (H. B. K.) Lag., Iowa,
Kans., Nebr.—Calamagrostis canadensis (Michx.) Beauv., Minn., Wise.—
Cinna arundinacea L., Ark., Md., Va.—Dactylis glomerata L., N. J.—
Danthonia spicata (L.) Beauv., Md., Va.—Elymus canadensis L., Wash.—
E. dahuricus Turcz., Wash.—E. virginicus L., Iowa, Minn., N. Dak.,
Okla., S. Dak., Wash.—Festuca rubra L., Penn. (imported)—Glyceria cana-
densis (Michx.) Trin., Mich.—G. septentrionalis Hitchc, Mich., N. Y.
—G. striata (Lam.) Hitchc, Ga., 111., Iowa, Md., Mich., Miss., Wise.—
Hystrix patvla Moench, Mich., Minn., Mo., Wise.—Koeleria cristata (L.)
Pers., N. Dak., Penn., S. Dak., Wise.—Leersia oryzoides (L.) Swartz,
N. Y.—Melica bulbosa Geyer, Wash.—Phleum pratense L., Me., Nebr.,
Penn.—Poa compressa L., N. Dak.—P. palustris L., Del., N. Y.—P. pra-
tensis L., Del., Iowa, Nebr.—P. secunda Presl, N. Dak.—P. stenantha
Trin., Wash.—Sphenopholis nitida (Spreng.) Scribn., Md.—S. obtusata
(Michx.) Scribn., Ga., Miss., N. Gar.—S. pollens (Spreng.) Scribn., Ohio.
This fungus is systemic, living in the crown and rhizomatous parts

of the plant and developing on the surface of the host during the sum-
mer months. Longitudinal sections of culms stained with cotton blue
disclose the sparsely septate intercellular strands of mycelium. In early
spring the mycelium forms a white weft on young tillers. Sometimes in
late grasses, such as timothy, tillers may be trapped and delayed or
destroyed. The fungus increases in thickness, becoming first yellow,
then orange, forming the characteristic spongy collar around the sheath
or culm. Conidia are first developed and later numerous immersed
perithecia with papillate ostioles form.
Epichoe has been grown in pure culture but only conidia form under
such conditions. Benedict (1929) found that conidia germinated in 15
to 18 hours at 18° to 21° C. It has not been proved that the conidia can
cause the disease. Sampson and Western (1941) state that Vladimir-
skaja has claimed to have obtained infection with them b u t Sampson
and Western question the work. T h e y do report t h a t spread of t h e
disease a n d the fungus is often due to clonal division of infected peren-
nial grasses and b y t r a n s m i t t a l on seeds of perennial grasses such as
red fescue. T h e bulbils of Poa bulbosa also cariry the fungus, according
to these authors. There seems to be no record of: the fungus on this host
in N o r t h America. E. typhina is much more important in E u r o p e t h a n
in N o r t h America. I t is relatively common on Koeleria cristata in t h e
prairie regions of the United States and in isolated instances has been
locally a b u n d a n t on Poa compressa, P. secunda, Agropyron smithii, a n d
A. trachycaulum. ,
Infection from seed-transmitted material is sometimes high, as
much as 99 per cent has been reported in red fescue at Aberystwyth,
Wales. T h e fungus occurs on all p a r t s of the flower, the palea, lemmas,
stamens, and young ovaries. I n this case it is similar to t h e systemic
fungus, Apiosporina collinsii (Schw.) v.^ Hoh., a perennial parasite of
Amelanchier spp. '/
T h e economic importance of E. typhina lies in its detrimental effect
on seed-producing stands. W e r n h a m ^ ( 1 9 4 2 ) , incidentally, reported its
introduction on varieties of red fescue from H u n g a r y .
N o adequate control has been'developed although roguing is p a r -
tially successful.
I n a Fitzgerald gives synonymy of this fungus in her recent paper
(1949). T h e preferred name is Typhodium typhinum (Fries) Seaver
(Seaver, 1910, Fries, 1823) according to one school of thought. If a c -
cepted b y the next Botanical Congress this name will replace the well-
known Epichloe typhina (Ann. Sci. N a t . IV. 1860).
R a n g e : United States, Canada, China, Denmark/England, Germany, Hun-
gary, Netherlands, Poland, Sweden, Switzerland, U.S.S.R., Wales.
R e f e r e n c e s : Benedict (1929); Fitzgerald! (1949); Fries (1823); Ingold
(1948); Fetch (1938); Sampson (1929,i 1933); Sampson and Western
(1941); Seaver (1910); Stevens (1913)1; Vladimirskaja (1928); Wern-
ham (1942). , —-^
E r y s i p h e g r a m i n i s D C . — P o w d e r y Mildew
Usually epiphyllous, b u t sometimes amphigenous; mycelia more or
less persistent, effused or forming scattered patches, at first^ white, fre-
quently becoming buff, pale brown or weathering to g r a y ; perithecia
(cleistothecia) large, 135-280 n, mostly about 200 /i diam., scattered or
gregarious, globose-depressed, becoming concave, usually more or less
immersed in the lanuginose persistent mycelium, which is .formed of
sparingly branched, curved, rather rigid, tMck-walled, or solid, shining,
interlaced h y p h a e , 4-5 jj. wide, cells of perithecium obscure; appen-
dages rudimentary, few or numerous, very short, simple or sparingly
branched, pale brown; asci numerous, 9-30, usually from 15-20, v a r y -
ASCOMYCETES 79
ing from cylindrical to ovate-oblong, more or less longly pedicellate,
70-108 X 25-40 z^; spores 8 (or rarely 4), 20-23 X 10-13 /*, seldom pro-
duced on living host plants. Conidial stage (Oidium monilioides Desm.)
light gray, ovoid, in chains arising from simple conidophores, 25-30 X 8-
10 jn, from superficial myce!ia. Haustoria directly penetrate the hosts
cuticle forming a few branches in the epidermal cells, conidial forma-
tion chiefly diurnal.
On: Agropyron cristatum (L.) Gaertn., Wash.—A. dasystachyum (Hook.)
Scribn., Ida., N. W. Terr., Oreg.—A. inerme (Scribn. and Sm.) Rydb.,
Mont., Oreg., Wash.—A. repens (L.) Beauv., Alta., Calif., Ida., Iowa,
Manit., Mich., Minn., Mont., Nebr., N. B., N. S., N. Dak., Oreg., P. E.
Is., Que., S. Dak., Utah, Wash., Wise, general in the eastern United
States—A. riparium Scribn. and Sm., Ida.—A. sibiricum (Willd.) Beauv.,
Wash.—4. smithii Rydb., Colo., Mont., Nebr., N. Dak., S. Dak., Wye—
A. spicatum (Pursh) Scribn. and Sm., Colo., Nev., Oreg., Wash.—A. stria-
tum (Steud.) Nees ex Hook., Wash.—A. subsecundum (Lk.) Hitchc,
Colo., Minn., Wise, Wyo.—A. trachycaulum (Lk.) Malte, Alta., Calif.,
Colo., Manit., Minn., Mont., N. Dak., Oreg., S. Dak., Utah, Wash., Wise,
Wyo.—A. trichophorum (Lk.) Richt., N. Dak., Utah—Agrostis alba L.,
Ida., Iowa, Minn.—A. diegoensis Vasey, Cahf.—A. exarata Trin., Alaska,
Calif., Idaho, Mont., Oreg., Wash., Wyo.—A. verticillata VilL, Oreg.—
Avena byzantina K. Koch, Cahf., Ga., Oreg., Wash.—A. fatua L., Calif.,
Oreg.—A. sativa L., Alta., B. C, Cahf., Ga., Mo., Oreg., Que., Tex., Utah,
Wash., "northern states" (Wejss, 1943)—Beckmannia syzigachne (Steud.)
Fern., Ida., Minn., Mo., Mont., S. Dak.—Bromus breviaristatus BuckL,
Mont., Wash., Wyo.—B. carinatus Hook, and Arn., Colo., Ida., Mont.,
Oreg., Wash., Wyo.—B. catharticus Vahl, Ga., Okla., S. Dak., Tex.—B.
commutatus Schrad., Wash.—B. erectus Huds., Wash.—B. inermis Leyss,
Minn., Wash.—B. mollis L., Wash.—B. racemosus L., Oreg.—B. rigidus
Roth, Wash.-^5. sterilis L., Oreg.—B. suksdorfii Vasey, Wash.—B. tectorum
L., Nebr., Wash.—B. vulgaris (Hook.) Shear, Ida.—Buchloe dactyloides
(Nutt.) Englm., Okla.—Calamagrostis canadensis (Michx.) Beauv., Ohio
-;-C. rubescens Buck!., Mont.—Catabrosa aquatica (L.) Beauv., Mont.,
S. Dak.—Cinna ariindinacea L., Iowa—Cynodon dacttjlon L., Calif.—
Dactylis glomerata L., B. C, Ida., Iowa, Mich., Minn., Mo., Mont., N. Y.,
Ohio, Ont., Oreg., Penn., Utah, Wash., Wise.—Digitaria sanguinalis (L.)
Scop., Tex.—ElymuS'Canadensis L., Iowa, Mich., Minn., Mo., Mont., Nebr.,
Penn., Wash., Wise.—E. condensatus Presl, Alta., Calif., Ida., Nev., Oreg.,
Wash., Wyo.—E. dahuricus Turcz., Mich.—E. glaucus Buck!., Mont.,
Utah, Wash.—E. junceus Fisch., Alta., Mont., N. Dak., Wash.—E. sibiricus
L., N. Dak.—E. triticoides BuckL, Mich.—E. villosus Muhl, Wise.—E.
virginicus L., Minn., Mo.—Festuca idahoensis Elmer, Wash.—F. ovina
L., Wash.—F. rubra L., Mich.—i?". rubra var. commutata Gaud., Oreg.—
Glyceria striata (Lam.) Hitch., Mont.—Hordeum brevisubulatum (Trin.)
Lk., N. Dak.—H. disiichon L., Calif., Ida., 111., Iowa, Ky,, Minn., Mo.,
Nebr., N. Y., Ohio, Oreg., Penn., Utah, Va., Wash., Wise—F. jubatum
L., Alta., Colo., Iowa, Manit., Minn., Miss., Mo., Mont., N. Dak., N. W.
Terr., Wise.—H. nodosum L., Colo., Oreg.—H. pusillum Nutt., Ga., Kans..
Okla.—H. vulgare L., Alta., Ariz., B. C, Calif., Colo., Conn., D. C, Ga-
80 DISEASES OF CEREALS AND I GRASSES
/ 1
Ida., Iowa, 111., Ky., Manit., Mo., Nebr., N] B., N. S., Ohio, Okla., Ont.,
Oreg., Penn., P. E. Is., Que., Sask., Utah, Va., Wash., Wise.—Hystrix
patula Moench, Wise.—Koeleria cristata (L.) Pers., Wash., Wise.—Melica
calijornica Scribn., Calif.—Milium cfjusum lli., Mich.—Phalaris arundi-
nacea L., Wash., Wise—Phleum pratense L.,'Alta., Manit., Minn., Penn.,
Wise.—Poa al-pina L., Wyo.—P. ampla Men., Mont., Oreg., Wash., Wyo.
—P. arachnifera Terr., Iowa—P. arida Vasey, Mont., N. Dak.—P. canbyi
(Seribn.) Piper, Colo., Mont., N. Dak., Oreg., S. Dak., Utah, Wash.,
Wyo.—P. compressa L., Manit., Minn., Mont., Okla., W. Va., Wise.—
P. cusickii Vasey, Mont., Wyo.—P. epilis Seribn., Wash.—P. glaucifolia
Seribn. and Wils., N. Dak.—P. gracillima Vasey, Oreg., Wash., Wyo.—
P. interior Rydb., Me., Minn., Mont., S. Dak., Wash., Wyo.—P. junci-
folia Seribn., Oreg.—P. leptocoma Trin., Oreg.—P. longifolia Trin., Wash.
—P. nemoralis L., Mont., S. Dak., Wash., Wyo.—P. nervosa (Hook.)
Vasey, Utah, Wyo.—P. nevadensis Vasey, N. Dak., Wash.—P. palustris
L., Colo., Iowa, Manit., Mich., Mont., N. Dak., Oreg., S. Dak., Utah,
Wash., Wise., Wyo.—P. pratensis L., Alta., Alaska, B. C, Cahf., Colo.,
Del., Ida., Iowa, Manit., Mich., Minn., Mo., Mont., Nebr., N. Mex.,
N. Dak., Ont., Oreg., S. Dak., Utah,'Wash., Wise, Wyo., general in the
East (Weiss)—P. scabrella (Thurb.) Benth., Mont., Oreg., Wash.—P.
secunda Presl, Calif., Colo., Mont., Oreg., Wash.—P. silvestris A. Gray,
Wash.—P. vaseyochloa Seribn., Oreg., Wash.—Polypogon monspeliensis
(L.) Desf., Calif., N. Mex., Okla., Oreg., Wash.—Puccinellia distans (L.)
Pari, Wash.—Secale cereale L., Alta., Ariz., B. C, Conn., Iowa, Manit.,
Mich., Minn., Okla., Ont., Oreg., Que., Sask., Wash., Wise.—Sitanion
hansenii (Seribn.) J. G. Sm., Calif.—S. hystrix (Nutt.) J. G. Sm., Calif., •
Nev.—S. jubatum J. G. Sm., Utah—Sphenopholis obtusata (Michx.)
Seribn., Iowa—Sporobolus giganteus Nash, Ariz.—Stipa calijornica Merr.
and Davy, Calif.—Triticum aestivum L., Alta., B. C, Calif., Colo., Del.,
Ga., Ida., Ill, Iowa, Manit., Md., Minn., Mo., Mont., Nebr., N. B., N. J.,
N. S., Ohio, Okla., Ont., Oreg., P. E. Is., Que/ Utah, Wash., Wyo.
!
The host range of this species is large, as indicated by the above list.
A number of workers have determined (hat E. graminis is composed of
a number of distinct physiological varieties. Marchal (1902, 1903) 'dis-
tinguished seven of these, some of which were sharply restricted in their
host range to certain genera of grasses. These included varieties on
Triticum, Hordeum, Secale, Avena (and Arrhenatherwn), Poa, Agro-
pyron, and Bromus. Salmon (1904) found that the varieties of Marchal
could, as in "bromi," be broken down further into physiological races.
This was further elaborated on by Mains and Dietz (1930) working
with .E. graminis hordei and Mains with.E. graminis tritici (1933).
Hardison (1944,1945,a) found increasing evidence that/numerous races
were present which did not follow the clearly defined varietal lines
given by Marchal. The conclusion reached after reading Hardison's
papers is that Marchal's varieties are now at least/partially obsolete.
Considerable study is now being conducted on varietal resistance to
powdery mildew in cereals. Sampson and Western (1941) among others
have noted varietal resistance in pasture grasses. Honecker (1931)
found that Pflug's Intensiv barley wa^ resistant because of its low albu-
ASCOMYCETES 81
men content and high' reserve of starch. There seems to be consid-
erable difference in native rangeland Poa spp. in their susceptibility
to mildew and there are also clonal differences in the same species in
some cases.
Eraser (1947) reported that Cascade, a new Canadian spring wheat,
was highly resistant to powdery mildew.
Mehta (1930) found that powdery mildew survived on plants grow-
ing in the hills of India in the conidial stage and the spores spread to
the foothills and to some extent onto the hotter plains by wind dispersal
in favorable seasons. Montemartini (1930) suggests that in Mediter-
ranean countries the disease may have become able to overwinter
solely in the conidial stage. Gorlenko (1942) found that wheat mildew
overwintered in the Voronezh district of Russia as brown mycelial pads
on winter wheat varieties, as did barley mildew. The fungus over-
wintered on Bromus arvensis by means of numerous surviving peri-
thecia. On Agropyron repens the fungus overwintered as mycelium. On
perennial Poa it survived as conidia in the winter rosettes while on the
dead plants of an annual species of Poa it survived as perithecia. Gor-
lenko concluded that on annual plants powdery mildew tended to
produce perithecia while on perennial grasses perithecia were not neces-
sary and hence were seldom formed.
Meyer-Hermann (1935) reported that unoiled calcium cyanamide
used at the rate of 1 doppelzentner per hectare apparently controlled
Erysiphe graminis. "Pearl" (pellet) calcium cyanamide used at the rate
of 2 doppelzentner per hectare also proved beneficial. Generally, an
abundance or excess of nitrogen is considered as tending to increase
infection by Erysiphe graminis.
Homma (1937) found that infection tubes from germinating spores
of E. graminis penetrated into the cell wall of both immune and sus-
ceptible varieties of wheat but formed haustoria only in the susceptible
varieties. Homma also described Uncinula bifurcata on oats.
Yakoubtziner (1934) reported that Triticum timopheevi Zhuk. was
resistant to various fungi including Erysiphe graminis. This emmer-like
host is widely used in experimental breeding work. Along the same line
Nover (1941) found a few plants of winter wheat grown from seed
obtained in Hiffdu Kush that were resistant to two races of mildew.
Freisleben and Lein (1942) found 19 mildew-resistant plants of the
spring barley Haisa among thousands of seedlings of the progeny of
seed which had been exposed to Rontgen rays at dosages of 4,000 to
14,000 r. The X2 progeny produced one vigorous seedling which was
mildew-free in the field. What became of this one survivor is not known.
Newton and Cherewick (1947) reported that nine physiologic races
of E. g. hordei and E. g. tritici have been isolated in Canada. Discussion
of these races was detailed.
The physiology of E. graminis has been the subject of many papers.
One of the more available of these is the study by Graf-Marin (1934).
/j
See Yarwood and Cohen (1949) for recent paper 'on diurnal response
to nutrients.
Range: General, except in some tropical areas;
References: Allen and Goddard (1938,a,6); Anderson, W. F. (1889,6)
Bjorling (1946); Briggs (1935, 1938); Briggs and Stanford (1938, 1943)
Brodie (1942, 1945); Clierewick (1944); Comer (1935); Dietz (1930)
Favret (1947); Foex (1924); Fraser (1947); Freisleben and Lein (1942)
Garofalo (1947); Onrlenko (1942); Graf-Marin (1934); Grainger
(1947); Hardison (lb'44; 1945,a); Homma (1929, 1937); Honecker
(1931, 1934, 1938); Johnston, Fellows, and Melchers (1937); Kostoff
(1938); Lowig (1933); Mackie, Jane R. (1928); Mains (1925, 1933,
1934); Mains and Dietz (1930); Marchal (1902, 1903); Mehta (1930);
Montemartini (1930); Newton and Cherewick (1947); Never (1940);
Pape and Rodemacher (1934); Pratt (1938); Reed (1905, 1909, 1912,
1916, 1920); Roemer, et al. (1938); Rosenstiel (1938); Salmon (1900,
1904); Sarasola, Favret, and Vallega (1946); Schhcthng (1939); Schulz
(1930); Shands, R. G. (1939); Tapke ,(1948); Tidd (1937); Tracy and
Galloway (1888, p. 35); Trelease and Trelease (1928, 1929); Vallega and
Cenoz (1941); Vallega and Favret (1947); Vik (1937); Wolff (1875);
Yakoubtziner (1934); Yarwood (1945); Yarwood and Cohen (1949).
Gibberella fujikuroi (Saw.) Wollenw. (and var. subglutinans

Edwards)
G. moniliformis (Sh.) Wineland

See Fusarium moniliforme Sheldon for a discussion of the peri-
thecial stage known by both of the above names. It seems more satis-
factory to the writer to discuss this group from the standpoint of the
well-known conidial stage. ; /
Gibberella zeae (Schw.) Fetch.

G. pulicaris (Fries) Sacc.
I G. saubinetii (Mont.) Sacc. ^--~. ..^^
• •
See Fusarium graminearum Schw, for a discussion Df this 'fungus.

The exact name for the perithecial stage is not settled but see Fetch
(1936) and Fitzgerald (1949). For the same reasons mentioned under
G. moniliformis, above, it also seems simpler to discuss this fungus
under the genus Fusarium. , '
/
Gnomonia iliau Lyon—Iliau Disease of Sugar Cane
Melanconium iliau Lyon ^ /
Produces a cementing mat of white "mycelium at base of sheaths,
binding culms at times. Ferithecia globose, flattened but with greatly
elongated necks which protrude from the culm or leaf for some dis-
tance; horny, 325-480 X 240-340 /i, bgak 350-550 /j.; asci clavate, thin-
ASCOMYCETES 83
walled, 60-80 X 8-14 ft with well-developed pore at apex; spores hya-
line, 1-septate, constricted, slightly curved, spindleform, 22-30 X 5-7 /x.;
acervuli or pycnidia, 500-700 fx., somewhat thin-walled, globose, semi-
chambered, broadly ostiolate, conidia ellipsoid-ovoid, obscurely brown,
15-28 X 7-10 ju, borne singly on conidiophores, spores filled with char-
acteristic large globules.
On: Saccharum officinarum L., Hawaii, La.
Faris and Allison (1927) reported that this fungus was nonpara-
sitic. It may, perhaps, be a wound parasite. As indicated by Edgerton
(1913) the imperfect stage is not a good Melanconium as the fruiting
bodies are thin-walled or loosely formed pycnidia with an ostiole
usually sessile with the leaf surface.
The disease, or the condition, is favored by cold, backward weather
and poor growing stock. The Louisiana disease appears to be caused
by-a strain of fungus different from that found on sugar cane in Hawaii.
Freise (1930) reports that this disease is called "colla-folha" or
leaf gluing in Brazil from the fact that the leaf sheaths surround and
adhere firmly to the stalk which turns reddish-blue and very flaccid.
Where not severe, Freise (1930) controlled the disease by applying
stagnant black water from virgin forests. Apparently the colloidally
suspended humic acids destroyed the fungus. The canes recovered
within a few days after application of the black water.
Range: United Sjtates, Australia, Barbados, Brazil, Cuba, Hawaii, West
Indies.
References: Edgerton (1913); Faris and Allison (1927); Freise (1930);
Nowell (1923).
See Melanconium sacchari Mass. apud Speg. in the Fungi Imperfecta
Leptosphaeria avenaria Weber, see Septoria avenae Frank
L. avenaria f. sp. triticea T. Johnson, see Septoria avenae f. sp.

_ triticea T. Johnson
Leptosphaeria herpotrichoides de Not.—Secondary Root Rot

Perithecia sphaeroid-depressed with a short neck- which extends
through the leaf sheath, black, up^ to 200-250 /x diam.; asci subcylin-
drical, 80-100 X 10 /*; spores-fusoid-elongate, somewhat curved, 25-
40 X 4-6 [I, yellowish, 6-9' septate, constricted at most of the septa,
paraphyses present. '
On: Secale cereale L., N. Dak.—Triticum aestivum L., Alta., Wash.
Leptosphaeria herpotrichoides was, for many years, believed to be
the main cause of straw-breaker foot rot of cereals, especially wheat,
in Europe. Research has shown that it is actually only a saprophyte
which may have some weak ability to invade maturing plants (Sprague,
1931,a; 1934,6; Sprague and Fellows, 1934). Iti is seldom mentioned in
recent literature although Dickson (1947, p. 215) still lists this fungus
as apparently comparable in im^-rtance to '.Cercosporella herpotri-
choides, relying on early records which were liot based on any critical
pathogenicity studies. i
In North America McKinney (1925) reported this fungus on wheat
from Spokane County, Washington, in material now known to have
been parasitized by Cercosporella herpotrichoides (Sprague and
Fellows, 1934). Henry and Foster found the fungus on wheat in
Alberta but did not follow this further, evidently concluding that the
fungus was of slight importance. The writer has seen this species on
dead rye plants in North Dakota and has observed similar, if not
identical, saprophytic material on alpine grasses in Wyoming and
Washington. In all instances which the writer has observed this species
it has no particular association with any other root-rot fungus. Inocu-
lation trials with a culture of this fungus sent by the late Dr. Foex
indicated that L. herpotrichoides was scarcely parasitic under the cli-
matic conditions prevailing during the season of the year when straw-
breaker foot rot is developing in Northern Europe. ^
Range: United States, scattered in the cooler parts of the Northern Hemi-
sphere including Belgium, Canada, Denmark, Finland, France, Germany,
Italy, Poland.
References: Delacroix (1901); Dickson (1947); Henry and Foster (1929);
Mangin (1899); McKinney (1925,a); deNotaris (1865); Sacc'ardo
(Fungi Italici, 1879, Fig. 503); Sprague (1931,a; 1934,6); Sprague and
Fellows (1934).
Leptosphaeria sacchari Van Breda de Haan—Secondary Leaf Spot

Phyllosticta saccharicola Hennings '
See discussion incidental to Helminthosporium sacchari (Van Breda
deHaan) Butler.
Leptosphaeria salvinii Catt.—Stem Rot of Rice -

Sclerotium oryzae Catt.
Helminthosporium sigmoideum Cav.
H. irregulare var. microsphaeroides Nakata ^
See also H. sigmoideum var. irregulare Cralley and Tullis
Mycelia slender, white, 2-6 /j. diam., sclerotia black, smooth, shiny,
circular to somewhat irregular in outline (but see H. sigmoideum var.
irregulare) more or less spherical, 150-500 /* diam.,/usually small and
approximately 200-275 /A on culms; conidi'a- (-Helminthosporium, sig-
moideum Cav.) borne on dark-colored conidiophores which are, septate,
erect, simple or sparsely branched, 100-175 X 4-5 /*; conidia borne
singly on sharp pointed sterigmata, fusiform, typically 3-septate,
ASCOMYCETES 85
simply curved or slightly sinuous; intercalary cells Prout's brown,
densely granular; terminal cells lime green, less granular than inter-
calary cells; apical cell frequently longer and less acutely pointed than
basal cell; spores occasionally constricted at middle septum, 29-49 X 9.9-
14.2 fi, mostly 34-40 X 11-12.5 y.. Perithecia dark, globose, embedded
in outer tissues of sheath, 202-481 n, average diam. about 381 [n, beak
rather short (30-70 /x), frequently half the diameter of the perithecium
in width. Beak nonprotruding, tip flush with surface of outer epidermis
of sheath, visible to naked eye; asci narrowly clavate, walls almost
invisible and deliquescing by the time spores mature, short-stalked,
90-128 X 12-14 n, mostly 103-125 X 13-5 jx] ascospores biseriate, nor-
mally eight in ascus (rarely four), 3-septate, when mature, usually
somewhat constricted at septa, particularly at middle septum, brown,
two end cells usually lighter in color and contents less granular than
middle cells, fusiform, somewhat curved, 38-53 X 7-8 n, mostly 44-
48 X 8 /t.
On: Echinochloa colonum (L.) Lk., Aik.—Oryza saliva L., Ark., Calif., La.,
N. Car., Tex.—Zizaniopsis miliacea (Michx.) Doell., and Asch., Ark., La.,
Tex.
The disease first appears in late July or early August as small, dark,
discolored areas on the leaf sheaths. Invasion of the stem follows'. Ac-
cording to Tullis and Cralley after the fungus reaches the culm (1933)
it produces abundant appressoria over the area covered by the my-
celium. These are visible as minute black dust. Small irregular sclero-
tial masses are formed on the' culm. Invasion and decomposition con-
tinue until the culm collapses. The sclerotia are by then very numerous
in the diseased tissue.
The optimum growing temperature for the stem-rot fungus in pure
culture lies between 25° C. and 30° C , the minimum growing tempera-
ture is 13° C , and the maximum temperatures at which growth will
occur is 32° C. to 35° C.
The conidial stage- develops on the host in early August. Large
numbers of conidia are produced from then until the time that the
water is draiiied from the fields in late fall, shortly before the rice
becomes mature.. The -ascigerous stage develops in later August and
perithecia apparently survive until the following year. These may be
a source of infection but the abundant long-lived sclerotia are prob-
ably more important. The conidia are important sources of secondary
infection, being readily spread by the water in the fields. Red rice,
because it matures early, is an, additional source of conidial inoculum
(Tullis and Cralley, 1933),
The control of stem'rot is difficult but the most promising practise
has been to withdraw the standing water from the fields for a period
prior to maturity. Some Japanese varieties have shown some resistance
to stem rot (Cralley, 1936). No control by use of fertilizers was ob-
tained by Cralley (1939). Tullis and Cralley (1941) found that
sclerotia of L. salvinii could remain viable in uncultivated rice^soil for
at least six years. Rotation periods of 4 to 6, years apparently will not
eliminate the disease. Park and Bertus arrive^ at somewhat the same
conclusions (1932,b) as did Nisikado and Hirata (1937).
Range: United States, Bulgaria, Burma, Ceylon,'China, .Dutch East Indies,
India, Indochina, Italy, Japan, Madagascar, Phijippines, Portugal, Turkey.
References: Baldacci (1947-48,a); Baldacci and Ciferri (1936); Bouriquet
(1948); Butler (1913,6); Cattaneo (1876, 1879); Cavara (1889); Cralley
(1936, 1939); Cralley and TuUis (1935, 1937); Metcalf (1907); Miyake
(1910); Mundkur (1935); Nisikado and Hirata (1937); Nisikado and
• Miyake (1922); Park and Bertus (1932,6; 1934,a);' Shaw (1913); Tis-
dale (1921, 1922); TuUis (1932; 1933,a; 1940); TuUis and Cralley (1933,
1941); TuUis, Jones, and Davis (1934); Wei (1934,6).
Lophodermium arundinaceum (Schrad. ex. Fr.) Chev.—On Dead '

Culms and Leaves
Perithecia scattered or gregarious, on pale spots, innate erumpent
and prominent, lying in the direction of the axis of the stem, elongated
or elliptical, straight, simple, obtuse or acute, brown or black, with
sharp, sometimes white-margined lips, finally opening lightly to leave
a narrow crevice through which the pale brown disk is partly visible,
Yz-l mm. long X / i - % mm. wide. Asci clavate, subsessile, 70-80 X 8-10
/t (75-100 X 9-12 fi, after Rehm), with filiform paraphyses longer than
the asci and mostly undulate above. Ascospores 8 in an ascus, fili-
form, straight or slightly bent, continuous, multinucleate, hyaline| 40-
70 X 1-5-2 /i. The pycnidial stage (spermogonial as given by Ellis and
Everhart), that is, Leptostromella hysterioides Sacc, have curved,
subulate, continuous spores, 16-18 fj. long. Tw() varieties are given by
Ellis and Everhart (1892,6): var. vulgare Ecki. has scarcely prominent
perithecia, which are brownish elliptical with a very narrow cleft,
l%-2 X %-% mm- and ascospores 70-75 jx long. This occurs on
Phragmites communis in Europe. The other variety is var. culmigenum
(Fr.) Fckl. based onHysterium culmigenum Fr.'It'has strongly promi-
nent perithecia which are mostly elliptical, obtuse, black, Opening with
a decided cleft, % - l % X "14-/4 mm. and ascospores 45-75 X 1.5 fi,
subacute. This is also listed sometimes as L. arundinaceum (Fr.)
Karst.
On: In Weiss's check list the hosts are Arundinaria gigantea (Walt.) Chapm.,
Alabama, Ga., S. Car.—Bouteloua curtipenduia (Michx.) Torr., Kans.
Ellis and Everhart (1892,6) report var. culmigenum on the dead culms
and sheaths of Poa compressa L., Phleum pratense L., and Andropogon
virginicus L. as common near Newfield, N. J., .and on wheat straw in Ohio.
In the alpine regions of the West this" apparent saprophyte is com-
mon on many species of grasses. The scattered large elliptical black
perithecia or hysterothecia (Gaumann and Dodge, 1928,, p. 296) occur
ASCOMYCETES 87
as relatively prominent, elongate black bodies on the white, bleached,
overwintered leaves and stems of many alpine and subalpine species of
grasses in Utah, Arizona, Washington, and California. These perithecia
are as large as 700 X 300 ^ and have the characteristic slit-like ostiole.
The following host range was noted by the writer (1948,c; 1949,a):
Agropyron inerme (Scribn. and Sm.) Rydb., Utah—Agrostis oregonensis
Vasey, Wash.—Calamagrostis breweri Thurb., Calif.—C. canadensis var.
scabra (Presl) Hitchc, Wash.—Danthonia intermedia Vasey, Calif., Colo.—
Festuca arizonica Vasey, Ariz.—F. ovina var. brachyphylla (Schult.) Piper,
Calif., Colo., Mont., Wyo., Wash.—Muhlenbergia sp., Cahf.—Oryzopsis
exigua Thurb., Wash.—Poa alpina L., Colo., Wyo.—P. arctica R. Br., Wash.
—P. compressa, L. Calif.—P. epilis Scribn., Calif.—P. fendleriana (Steud.)
Vasey, Ariz., Utah, Wyo.—P. juncifolia Scribn., Ariz.—P. leibergii Scribn.,
Calif.—P. longUigula Scribn. and Wils., Ariz.—P. pratensis L., Ida.—P.
rupicola Nash, Calif.—P. secunda Presl, Ariz.—Sitamon hystrix (Nutt.) J.G.
Sm., Ariz., Calif., Wash.—Trisetum spicatum (L.) Richt., Calif., Colo., Mont.,
Wyo.
An examination of herbaria would add a large number of additional
hosts but the above citations will be sufficient to show that the fungus,
where it occurs, is more or less indiscriminate in its host range. Since it
seems to be strictly saprophytic it probably scarcely warrants discus-
sion in this text. It does occur with snow mold in the high country in
the West and since it appears in phytopathological literature to some
extent it is, with considerable hesitation, included as an associated
saprophyte in this work.
Range: General, in the mountains.
References: Ellis and Everhart (1892,6); Fuckel (1869, Symb. Myc. pp.
256-257).; Sprague (1948,c; 1949,a).
Mycosphaerella calamagrostidis Greene—Leaf Spot

On growing leaf-tips and leaf sheaths, perithecia subcuticular,
seriate, subglobose, ostiolate, black, component cells pseudoparen-
chymatous, 125-160 ;* diam., asci numerous, 50-60 X 12-15 y.; width
nearly equal throughout, ascospores hyaline, subfusiform, flattened on
one side, septum median, 15-17 X 4-5 /*.
On: Calamagrostis canadensis L., Wise.
Greene (1945) reported that this fungus occurred on living plants at
the height of the growing season and he believed that it was parasitic.
Mycosphaerella calamagrostidis Volk was issued, without diagnosis,
in Rehm's Ascomycete Exsicc. on C. varia from Switzerland. I t dif-
fered, it was said, from Sphaerella tassiana de Not. mainly in the shape
of the perithecia. Greene's material, collected in August 1943 at Madi-
son, Wisconsin, has asci and spores much smaller than M. tassiana.
The perithecia of M. calamagrostidis are vaguely subcuticular, rather
than subepidermal. This species should be compared with several de-
scribed on Gramineae from Japan.
Range: Known from Wisconsin but possibly com'parable material occurs
in Japan.
Reference: Greene (1945).
Mycosphaerella holci Tehon—Leaf Spot

On leaves, inhabiting very extensive tan to straw colored red-
bordered areas; perithecia innate, developed in and occupying the
mesophyll, spherical, membranaceous, 60-100 /i, (iiami., opening hypo-
phyllously; ostiole papillate, usually about 15 /<, high, the pore 7-10 /*
wide. Asci cylindrical to clavate, 30-45 X 8-12 ^, aparaphysate, 8-
spored; ascospores 12-15 X 5-6 /*, biseriate, 1-S6ptate, constricted at
the septum, hyaline, upper cell somewhat the larger.
On: Sorghum vulgare L. (broomcorn), 111.
Tehon (1937) also noted an associated Phyllosticta stage with
spares 6-7 X 3-3.5 /* which he considered near P. sarghina Sacc. He
suggests that his fungus be compared .'with SphM^rella ceres Sacc. We
have not seen Tehon's material and jthe name is included here because
he states that the fungus, or fungi, occur on living leaves of broomcorn.
Range: United States (possibly also Italy, if a synonym of >S. ceres).
Reference: Tehon (1937).
Mycosphaerella tulasnei (Jancz.) Rothers—Leaf Mold

Cladosporium herbarum Lk.
Perithecia thick-walled, black, typically bro^clly flask-shaped with
a short neck, erumpent, 150-250 X 100-15p fx,; asci cylindrical, fusoid
or slightly tapering at the ends, 80-120 Xi 15-20 n; ascospores oblong,
bluntly pointed, 1-septate, upper cell somewhat larger than the lower
one, 18-28. X 6-8.5 i^. '
Conidial stage [Cladosporium herbarum) forming dense tufts of
velvety olive-black color. The conidiophores are; more or^Tess erect,
septate, sparsely branched; spores often in chains of 2 or~3, subcylin-
dric, pale olive, l-(2-3) septate, 10-15 X 4-7 /^.
On: Many species of Gramineae but chiefly saprophytic. There are some
indications that in early spring it may be parasitic on the leaves of cer-
tain grasses, notably western wheat grass {Agropyron smithii Rydb.) at
Mandan, North Dakota, and on Idaho fescue {Festuca idahoensis Elmer)
in the Far West. It produces brown flecks on the young leaves and forms
a brown to olive-black mold on any of the abovegrounfi parts after the
plant dies. There is some doubt about whether it is parasitic on even
the young leaves of western wheat grass at Mandan- because it usually
follows low-temperature conditions in early spring, especially after a
sharp frost or late snow flurry. In addition to its presence on grass in the
ASCOMYCETES 89
spring it is also frequently associated with leaf rusts and with such leaf
parasites as Septoria nodorum Berk. It is seldom isolated from the roots
of cereals and grasses (Sprague, isolation records). However, it is
almost omnipresent on leaves of winter wheat injured by Typhula ida-
hoensis Remsb. in Douglas County, Wash.
Bennett (1928,a), in England, has presented a critical piece of work,
with C. herbarum, which is of use to the plant pathologist. He con-
cluded that the fungus was of little importance in causing thinning out
and "deaf ears" in wheat. Grintescu in Romania (1923) decided that
Cladosporium blackening of wheat was saprophytic.
C. herbarum is possibly one of several species of this genus on
Gramineae, but without sufficient critical modern study, the writer
follows precedent in assigning them all to C. herbarum. C. graminum
Cda. is frequently used also. Its distinction from C. herbarum is not
clear. Schnellhardt and Heald (1936) report C. malorum Ruehle as
common on wheat seeds in Washington. Heald and Ruehle (1931) also
presented evidence that the microsporous stage, Hormodendrum clado-
s-porioides Sacc, is not conclusively part of the life cycle of M. tulasnei.
They (Ruehle, 1931; Heald and Ruehle, 1931) verified the work of
Janczewski (1894) who determined that M. tulasnei was the peri-
thecial stage of C. herbarum. The writer obtained M. tulasnei in ai pure
culture of C. herbarum in 1943, isolated from Stipa viridula Trin. in
western North Dakota. Fosterjs (1944) described C. moldavicum, said
to be parasitic, on all the aerial organs of Festuca ovina and Avenas-
trum planiculme in Romania. It has straight, simple, continuous or
uniseptate, rarely biseptate, conidiophores, dark brown at the base, be-
coming paler towards the apex, 75-200 X 3-5 /* with acrogenous, oval
or elongated, finally punctate, yellow-brown conidia, continuous spores
4-14 X 4-5 /x, uniseptate spores 6-14 X 4-6 /j., biseptate 4-18 X 5-7 fi.
Cladosporium herbarum is probably not important in its role as a
mold of shocked or standing grain or cut hay. When seed grain is moldy,
however, it is usually jlesirable to treat the seed with New Improved
Ceresan because the presence of C. herbarum so often indicates the
occurrence of more virulent species.
Pammel, Weems, and Scribner (1901) discussed the early history of
C. herbarum in this country. Fusicladium destruens Pk. (Rpt. N. Y.
State Museum Nat. Hist. 43. 1890) described on oats is probably simi-
lar. Peck also described C. zeae Pk. C. velutinum Ell. and Tracy (1890)
was described on Phalaris canariensis L. from Mississippi as forming
velvety patches on the leaves. The conidiophores were 50-75 X 4-5 n
and the 1- to 3-septate coiiidia'were 8-20 X 4-5 p., subhyaline, varying
from elliptical to oblong or cylindrical. C. piricularioides Dearness and
House was said to be parasitic on Panicum boreale Nash (N. Y. Mus.
Circ. 24. 1940).
In the Canadian Plant Disease Survey reports, the name C. grami-
num is used. For instance Cormack (Conners, 19th Ann. Rpt. Canad.
PI. Dis. Survey, p. 14, 1939) states that C. graminum is moderately
90 DISEASES OF CEREALS AND .GRASSES
severe on several varieties and strains of winter wheat in the plots at
Edmonton. This fungus, and possibly the somewhat more spiny Hetero-
sporium avenae Oud., becomes commoner toward the north where
spring frosts or periods of low temperature prevail. C. herbarum is also
reported as the cause of a floral sterility of PHleum pratense in Alberta.
In addition to M. tulasnei, M. tassiana (de Not.) Johans. has been
found on grasses in the Far West but is evidently not related to the
C. herbarum complex. Cash (1934) listed M. longissima (Fckl.) v. Arx
(1949) on Bromus secalinus L. from Sitka, Alaska, but this also ap-
peared to be saprophytic. Ellis and Everhart (1892,6, p. 297)' list a
number of additional species on grasses, all given under Sphaerella.
These include S. pilochorta Cooke, spores 13-17 X 3 /x on "some grass"
from Maine; S. muhlenbergiae Ell., spores 11-15 X 3-3.5 ix on Muhlen-
bergia from New Jersey; S. californica Cke. and Hark., spores 8 X 2 / / , ,
apparently associated with a Septoria on native grass; S. spartinae E.
and E., ascospores 12-16 X 4.5-5.5 ft on dead leaves of Spartina cyno-
suroides (L.) Roth, Nebraska; S. crus-galli E. and K., spores 14-16 X
4 /x on Echinochloa, Kansas; S. zizaniae (Schw.) Ell., spores 15-20 X
5-6 n on Zizania aquatica L., Delaware, Pennsylvania. If these prove
to be autonomous species most of them will be transferred to Myco-
sphaerella. The last mentioned was described as Sphaeria zizaniae
Schw. (Syn. N. Amer. 1750). Ellis and Everhart (1892,b) Hst Sphae-
rella zizaniae Cke. as a synonym under their S. zizaniae (Schw.). This
species seems to have smaller ascospores than M. tulasnei and M. tas-
siana and for the present can be considered distinct. I cannot find' any
reference to its transfer to Mycosphaerella although it is surprising if
this has not been done. To clarify the record we propose Mycosphae-
rella zizaniae (Schw. em. Ell. and Ev.) comb' nov. We are assuming
that Ellis and Everhart were correct in assigning their material to the
species represented by the sterile ones of Schweinitz' type. The latter
is essentially a nomen nudum. i
In routine examination of decadent material of grasses, especially
those from alpine habitats, the writer has noted fragmentary material
of Mycosphaerella many times but the asci are associated with so
many fungi that their relationship to any one asexual stage is not read-
ily determinable. A great deal of study is-needed with this group.
Range: General, especially in the cool temperate zone and northward.
References: v. Arx (1949); Bennett (1928,a); Bockmann (1933); Cash
(1934); Grintescu (1923); Heald and Ruehle (1931, Figs./9; 13,0,P,Q);
Janczewski (1894); Pammel, Weems, and Scribner (1901); Ruehle (1930,
1931); Schnellhardt and Heald (1936). .
^ Myriogenospora aciculisporae Vizioli—Tangle Top

Forms black scabs on the leaves varying from minute dots on longi-
tudinal stripes to irregular lesions affecting more than a' square centi-
ASCOMYCETES 91
meter of the lamina. White layers of closely interwoven compact
hyphae cover the spots, extending over the surface of the leaf and
resembling pseudoparenchymatous stromata, especially when situated
between two new leaves or within furled leaves which are then ad-
hered. Hyphae hyaline at first or tinged with yellow, later pale gray
and somewhat fleshy, septate, branched, containing numerous oil drops,
3-6 /t wide. Loculate, uniseriate, flask-shaped perithecia 250-380 X
200-300 //., completely immersed in the stroma and delimited by a zone
of nearly black tissue, ostiolar; asci fusiform, 210-250 X 10-16 [i, asco-
spores in bundles, numerous acicular, hyaline, nonseptate, straight or
slightly curved, 18-25 X 1-5 ;«..
On: Saccharum ofjicinarum L., La.
Abbott and Tippett (1941) reported this species on one variety of
sugar cane in Louisiana. It appears doubtful whether this species is
distinct from M. paspali (Diehl, 1934). The type area is Sao Paulo,
Brazil (Vizioli, 1926).
Range: United States, Argentina, Brazil.
References: Abbott and Tippett (1941); Freise (1930); Vizioli (1926).
Myriogenospora paspali Atk.—^Dwarfing and Fasciation

Stroma one to two centimeters long, seated upon one side of the
young culms in the cases observed, and partly enclosed on the sides by
the equitant leaf blades. Perithecia large, 200-300 jx diam., oval or
obovate, projecting above the stroma but little by a small conical apex
producing a slight unevenness of the surface. Asci large, 200-250 X
15-20 ju,, tapering each way to slender ends, more strongly so at the
base, so that they appear oblanceolate in side view. Spores very numer-
ous, several hundred, narrowly apiculate, 15-25 X 0.5-0.8 /* fusoid,
straight or slightly curved.
On: Andropogon scoparius Michx., La.—Cymbopogon citratus (DC.) Stapf,
Fla.—Paspalum laeve Michx., Alabama.
Diehl (1934) reported somewhat incomplete dwarfing and fascia-
tion of Andropogon as a result of a partial systemic association of the
fungus with otherwise healthy plants. In a high-temperature green-
house the plants eventually outgrew the fungus^ Most of the injury is
due to retardation of normal leaf unfolding because of the binding
action of the fungus stroma. The effect is similar to that recorded for
the "pokkah-boeng" disease.of sugar cane. Inoculation trials were un-
successful. M. paspali is said to be similar to M. bresadoleana P. Henn.
and M. aciculisporae Vizioli but the final taxonomic assignment of the
species was not made (1934).
Range: United States, Brazil.
References: Atkinson (1894); Diehl (1934); Freise (1930); Vizioli (1926).
, I
Ophiobolus graminis Sacc.—^Take-All
0. cariceti (Berk, and Br.) S^cc.
Ophiochaeta graminis (Sacc.) K. Hara
Mycelium permeating the roots of the hd^t, causing them to be-
come very brittle and easily broken away, developed profusely above
the crown of the plant in and about the leaf sheaths and forming a
definite thick plate between the inner leaf sheath and the culm; the
mycelial plate (usually adhering to the culm when the leaf sheaths are
stripped away) composed of coarse, dark brown hyphae, three to six
microns in diameter, which frequently run rather definitely parallel
to one another forming broad, flat, ribbon-like strands resembling
somewhat compressed rhizomorphs; perithecia membranaceo-carbona-
ceous, dark brown to black, smooth, rostrate, ostiolate, occurring on
the roots of the host- especially within thick wefts of fine rootlets de-
veloped abnormally about the (frown of the plant, but more frequently
observed arising from the mycelium êneath the outer leaf sheath,
standing singly or in groups, the individuals in a group occasionally
fused laterally but no true stromatic' tissue developed, firmly bound
to the leaf sheaths by numerous strands of mycelium attached both
to the neck and ascigerous portion^ often developed in great numbers,
more than one hundred individuals having been counted on a single
culm; when young hidden from view but at maturity the beaks pro-
truding and prominent, and by the shredding of the leaf sheath the
upper hemisphere of the perithecium often exposed to view; perithecial
beak developed obliquely and in the beginning lying more or less
parallel to the surface of the culm, later, curving sharply outward,
penetrating the leaf sheath and protruding, the obliquely attached
curved beak so characteristic of the species as/to be almost diagnostic;
ascigerous portion of the perithecium globose or subglobose, though
sometimes compressed between the leaf' sheaths, 330-500 /x (usually
about 425 /x) in diameter, narrowing gradiially into the truncate-conoid
to cylindrical beak which frequently atiJains a length as great as the
diameter of the ascigerous cavity; asci numerous,^seicled, elongate-
clavate, straight or curved, short stipitate to subsessile'j_90-115-X 10-
13 /J., rounded at the apex, 8-spored, thin-walled; paraphyses abundant,
thread-like, ilexuous, unbranched, hyaline; ascospores fascicled to sub-
biseriate, hyaline, as viewed together in the ascus faintly yellowish,
linear, curved, broader at the middle and tapering gradually towards
the ends, the upper end rounded, the basal end more acute and some-
times more sharply curved, 60-90 (chiefly 70-80) X 3 /*, when young
continuous and multiguttulate, at maturity 5- to 7-septate, not reach-
ing morphological maturity until late autuma or winter.
On: Agropyron repens (L.) Beauv. Alta., N. Y.—A. trachycaulum (Lk.)

Malta, Alta.—Agrostis alba L,, Ida.—A. exarata Trin., Ida.—Avena by-
zantina K. Koch, Calif., Oreg.—A. sativa L., Oreg.—Bromus carinatus
ASCOMYCETES 93
Hook, and Am., Ida.—B. japonicus Thunb., Ida.—B. orcuttianus Vasey,
Oreg.—B. secalinus L., Ark., N. Y.—B. tectorum L., Ida.—B. vulgaris
(Hook.) Shear, Wash.—Deschampsia danthonioides (Trin.) Munro, Ida.
—Elymus glaucus Buckl., Ida.—E. villosus Muhl., Okla.—Festuca der-
tonensis (All.) Aesohers. and Graebn., Oreg.—F. elatior L., Ark.—F.
megalura Nutt., Oreg., Wash.—F. myuros L., Oreg.—F. octoflom Walt.,
Ark.—Holcus lanatus L., Oreg.—Hordeum distichon L., Calif., Ida., Oreg.,
Wash.—H. pusillum Nutt., Ark.—H. vulgare L., Alta., Calif., Ida., Kans.,
Mo., Okla., Oreg., Sask., Wash.—Phleum pratense L., Ida., N. Y.—
Poa canbyi (Scribn.) Piper, Wash.—P. compressa L., N. Y.—P. pratensis
L., Ida.—Secale cereale L., N. Y., Okla.—Setaria geniculata (Lam.)
Beauv., Ark.—Triticum aestivum L., Alta., Ark., B. C, Calif., Ida., 111.,
Ind., Kans., Manit., Md., Mo., Nebr., N. Y., N. Car., Okla,, Ont., Oreg.,
Penn., Que., Sask., Tenn., Tex., Va., Wash., Wise.
Take-all apparently has been present in wheat fields in the eastern

United States since Colonial days. It was rediscovered in Missouri
during World War I and has since been found from Maryland and
Virginia to New York and westward in the winter wheat area.
In the Pacific Northwest take-all occurs in Oregon, Washington,
Idaho, and northern California. During the historically open winter
of 1933-34 it was found on the Idaho prairies near Nezperce, where
it occurs but is so inhibited during most years as to be unrecognized.
It was widespread there and in eastern Washington in 1948, a year of
excessive rainfall (Sprague 1948,d; 1949,6). I t is abundant in western
Oregon and it also occurs in trace amounts in eastern Oregon in the
Grande Ronde valley. In 1930 it was found on irrigated spring wheat
at Jerome, Idaho (Sprague, 1931,6).
Take-all appears to be native in the Pacific Northwest, although it
was not observed by experiment station workers until 1901, at Albany,
Oregon (Cordley, 1902). This material, still unlabeled, was seen by
the writer in 1933. It was definitely 0. graminis. The writer has seen
specimens of Holcus lanatus infected with take-all taken from virgin
soil from high in the hills of the Coast Range in Oregon and similar
material of Bromus orcuttianus Vasey has been found in the Siskiyou
Mountains, Ovegou.-Ophiobolus graminis is particularly common on
annual fescues [Festuca dertonensis (All.) Aschers. and Graebn., F.
megalura Nutt., F. myuros L.) in the Willamette Valley, in Newberg
sandy loam, and in the greenhouses at Corvallis, Oregon, it is' a nui-
sance in unsterilized soil. While 0., graminis is important mainly on
winter wheat and winter barley, it has been collected also on winter
oats in acid soil in coastal'Oregon and California. It was relatively
common on these hosts during the year 1933-34 and was found in the
Willamette Valley as well as along the coast. A trace of take-all was
noted on Poa canbyi (Scribn.) Piper at Pullman, Washington, and
small quantities were noted on wheat in the Palouse region in 1947
but, as mentioned, was common in late season there in 1948 during a
!\
year of heavy rainfall. Fraser, Simmonds, and Russell (1926) believed
that 0. graminis was indigenous to north-central Alberta.
The work of Fellows, Garrett, and the Canadian and Australian
workers has made 0. graminis one of the most jihoroughly studied fungi
in the cereal root-rot complex. Garrett's text (l944,a) deals thoroughly
with this fungus and the disease it causes. |
White (1947) found that take-all is favored.by dry top soil because
the diseased plants suffer more from drought than healthy ones which
have more extensive root systems. He found that two years of bare
fallow reduced the loss while cropping four years to wheat increased it.
One year's fallow, or oats in rotation with wheat, or oats-fallow-wheat
rotation decreased the severity and increased the yield. Increasing the
carbon content of the soil to activate biological antagonism was detri-
mental to Ophioboliis. Recently Garrett and Buddin (1947) reported
that take-all in continuously cropped spring-sown barley in England
could be controlled by the F. P. Chamberlain method: adding 10 lbs.
of trefoil and 10 lbs. Italian ryegrass/to the barley at seeding time,
plowing under in the fall. This results in low nitrogen between crops,
high during crops.
Until the modern work on take-all cleared up the confusion of
earlier observations even the cauSal organism was improperly said to
be the saprophyte 0. herpotrichus (Fr.) Sacc. While such popularizers
as Frank (1895, 1900) recognized the importance of take-all they in-
sisted on using the name 0. herpotrichus which simply compounded
some mistakes by Cugini (1890), Morini (1886), and others. Even as
late as 1908 Krueger used the name, Criiger employed it much later
(1929) and it occurs today in current literature from central Europe.
Foex and Rosella (1930,1931,a), van de Laar ('1931), Schaffnit (1930),
Fellows (1928,a,6; 1941), McKinney (.1925,4), Russell (1930), and
Garrett (1944,a) have all very conclusively laid the saprophyte 0.
herpotrichics to rest. I
There are various species of Ophioh'olus described on leaf sheaths
and dead culms of various grass hosts in additionJ:o those mentioned.
None of them seemfe to be of any importance and sbiiie-ipay prove to
be synonyms of 0. graminis. Ellis and Everhart (1892,'^ lists 0. me-
dusae Ell. and Ev. on Andropogon from Louisiana and several others
which are apparently saprophytes. 0. jestucae Tracy and Earle occurs
on dead leaves of fescue grass in Colorado. Drechsler (1934,6) reports
0. herpotrichus as being abundant on deadfquack grass iii Maryland,
the District of Columbia, Virginia, Wisconsin, and New York.
Holm (1948) believes that 0. graminis is a member of the Gnomo-
niaceae and probably is closely allied to Plagiosphaera.
Range: United States, Argentina, Australia',—Austria, Belgium, Canada,
Chile, Czechoslovakia, Denmark, E. Africa, "England, France, Germany,
India, Italy, Japan, Kenya, Morocco, Netherlands, New Zealand, Nor-
way, Poland, Scotland, Spain, Sweden, Switzerland, Uruguay, Wales.
References: Adam and Colquhoun (193^); Akerman, et al. (1934); Angell
ASCOMYCETES 95
(1947); Blanchard and Carrera (1933); Broadfoot (1933,6,c; 1934,a);
Brommelhues (1935); Buddin and Garrett (1941); Bussman (1936);
Carne and Campbell (1924); Clark (1942); Cordley (1902); Criiger
(1929); Dana (1919); Davis, R. J. (1925); Dennis (1944); Fellows
ll928,a,b; 1937,a,b; 1938); Fellows and Ficke (1934; 1939); Fitz-
patrick, Thomas, and Kirby (1922); Foster and Henry (1937); Fraser
(1924); Fraser, Simmonds, and Russell (1926); Garrett (1934,a,6,c;
1936; 1937,0,6; 1938; 1939; lMO,a,b,c; 1941,a,&; 1942; 1943; 1944,a,b;
1946); Garrett and Dennis (1943); Garrett and Mann (1948); Glynne
(1935); Griffiths (1933); Hara, K. (1916); Henry (1932, 1935); Henry
and Gilpatrick (1947); Holm (1948); Hynes (1937,o); Kirby (1922,
1925); Krebs (1933); Laar, van de (1931); Lai (1939); Ludbrook
(1942); Ludwig and Henry (1943); Mackie (1923); Mangin (1899)
McAlpine (1904); McKinney (1925); MoKinney and Davis (1925)
Melchers and Sewell (1926); Moore (1948); Moritz (1931, 1932)
MuUer-Kogler (1934, 1938); Nattrass (1943); Noll (1938); Ogilvie and
CoUett (1948); Osborn (1919); Padwick (1935, 1936,a,&); Padwick and
Henry (1933); Putterill (1924); Robertson (1932); Rosen and Elliott
(1923); Russell (1928, 1929, 1930, 1934, 1939); Samuel (1924, 1937);
Samuel and Garrett (1933); Sanford (1926, 1939, 1946); Sanford and
Broadfoot (1931); Sewell and Melchers (1924); Simmonds (1939,a,6);
Simmonds, Russell, and Sallans (1935); Sprague (1931,6; 1948,d;
1949,rf); Stevens (1919); Stumbo, et al. (1942); Suneson and Oswald
(1948); Turner (1940,a,6); Walker (1945); Weigert and Weizel (1935);
White (1941, 1942, 1945, 1947),; Winter (1939; 1940,a,6,c; 1942; 1945).
Ophiobolus oryzinus Sacc.—Black Sheath R o t of Rice

Perithecia gregarious, globose to subglobose, embedded, grayish at
first, becoming black to blackish brown a t m a t u r i t y , 187 to 375 /x diam.,
beak erumpent, hairy in culture only, 125 to 375 /* long; asci colorless,
clayate or subcylindric, tapering a t t h e ends with t h e apex slightly
larger t h a n the base, elongate, 8-spored, 96-115 X 12.5-14.5 /*; p a r a -
physes present, hyaline; spores fasciculate, long and slender, 3 - to
5-septate, often multiguttulate, usually crescent-shaped and often
slightly twisted spirally, slightly constricted a t the middle septum,
apex rounded, base-tapering, 79-112 (mostly 86-111) X 2-3 /x, buff to
white in mass to the naked eye but colorless when observed micro-
scopically.
O n : Oryza sativa L., Ark., La., Tex.
Lesions first appear on the culms in late J u l y or in August. Affected
plants show a brown discoloration of the sheaths from the crown of
the p l a n t to considerably above t h e water line. Finally the sheaths
but not the culms are rotted away. T h e perithecia numbering from a
few t o several hundred per plant lie with their beaks protruding
through the sheaths. Invasion of the host occurs by direct mycelial
penetration of the epidermis of the basal leaves. Appressoria are
. /I .
formed which aid in the invasion of the tissue, lying under the basal
leaves. Seedling injury also occurs.
Tullis (1933,6) also reported 0 . oryzde Miyake (1910) on rice in
the vicinity of Crowley, Louisiana. Drechsler (11934,6) indicated that
this species is close to 0. graminis. Hara reported 0. graminis on rice
from Japan (1916). Wei (1934) and Ling (1948) listed 0. oryzae from
China. Hara placed 0. graminis in Ophiochaeta graminis (Sacc.) Hara
(1916) because of the presence of bristly hairs on the beak of the peri-
thecium. He reported perithecia, 240-480 /x diam. with asci 80-120 X
8.5-16 /x and ascospores 72.1-100 X 3.2-4 /*.
Tullis and Adair (1947) reported that black sheath rot was present
and caused breaking over at the nodes, not at the internodes as in
Sclerotium rot. The following varieties were to some extent resistant:
Nira, Bluebonnet, Cody, Asohi, Fortuna, Kamrose, Texas Patna and
Hill Patna. While this disease has been present in Arkansas since at
least 1923 it had not caused appreciable loss until 1946.
Ito arid Kuribayashi (1927) described 0. miyabeanus on rice from
Japan. This is said to be the perithecia,l stage of Helminthosporium
oryzae and will be mentioned under the conidial stage. It has large
perithecia 560-950 X 368-777 /x and ascospores 250-468 X 6-9 /x. The
tremendous length of the ascospores is due in part to their corkscrew
placement in the ascus, [Cochliobolus miyabeanus (Ito and Kuribaya-
shi) Drechsler] the spores being appreciably longer than the ascus.
Range: United States, Italy, Malaya, Philippines. ^
References: Baker (1916); Hara (1916); Ito and Kuribayashi (1927);
Miyake (1910); Roemer, et al. (1938); Tullis (1933,&); Tullis and Adair
(1947).
/
Ophiobolus sativus Ito and Kuribayashi. See Helminthosporium
sativum P.K.B;.
Phialea temulenta Prill, and Del.—Blind Sefid^ Disease

P. mucosa Gray ~~]" ,»•
Apothecia small, fleshy, arising singly or in small numbers, 1-7^
usually 1 or 2, from colorless, septate, intertwining hyphae, 3-4 /A wide,
ramifying throughout the pericarp, testa, and endosperm; discs pale
pinkish cinnamon, darkening to cinnamon when old, 1-3.5 mm. in
diam., most frequently 2.5 mm., at first almost closed, opening, into a
cup shape and finally becoming flat or slightly recurved, with a srhooth
margin; stalks cylindrical, 1-8 mm. long, 0.4 mm. diam...smooth; struc-
ture fairly uniform throughout, consisting of hyaline,/parallel hyphae,
occasionally intertwining and seldom branched, forming a "textura
porrecta" with cells in the stalk 20-30 X 3-4 ju, grading into the more
interlaced hyphae of the excipulum, with cells 18-24 X 4-6 /j., passing
gradually into the hypothecium, which is 22-27 ^ deep, composed of
ASCOMYCETES 97
fine, interlacing hyphae, 2 /x broad; asci cylindrical-clavate, very little
thickened at the apex, the pore not staining blue with iodine, 66-116 X
3.3-7 [X, most frequently 73 X 6 /*, 8-spored; ascospores smooth, uni-
cellular, ellipsoidal, with pointed ends, biguttulate, hyaline, 7.6-12 X
3-6 /i, most frequently 9.5 X 4.5 fi, obliquely uniseriate in the upper
half to three quarters of the ascus; on germination each ascospore pro-
duces first a terminal germ tube and then a second, frequently lateral
in position and usually constricted at the point of origin; paraphyses
simple, filiform, hyaline, not swollen at the apex, 2-4 /x broad.
Macroconidia unicellular, uninucleate, cylindrical to slightly cres-
centic, with rounded ends, usually biguttulate, hyaline, 11-21 X 3.3-
6.0 fi, most frequently 16 X 4 /x, developed in summer in very large
numbers in succession from the apices of short outgrowths from the
hyphae on the pericarp, forming a pink slime on the surface of the
ovary; in germination a transverse septum forms, two germ tubes are
developed, one from each cell, the first terminal, the other terminal
or lateral, or the conidium may remain unicellular and produce only
one terminal germ tube.
Microconidia in pink pulvinate sporodochia, 1-1.5 X 0.5 mm. on
the surface of caryopses; conidiophores septate, guttulate, hyaline, two
or three times branched; microconidia unicellular, uninucleate, ovoid,
guttulate, hyaline, 3.4-4.8 X 2.7-3.2 n, most frequently 4 X 3 /x,' the
first formed by a constriction below the apex of the conidiophore, the
rest developed in succession inside a tube, 5 X 3 /x, formed by the
terminal portion of the conidiophore; germination not observed;
apothecia on dead or sometimes germinated seeds, in June; macro-
conidia on the ovary through the summer; microconidia on seed in the
soil, usually in February and March, sometimes later. (Description
from that of P. mucosa Gray, 1942.)
On: Aim caryophyllea L., Oreg.—Alopecurus geniculatus L., Oreg.—Bromus
racemosus L., Oreg.-—Cynosurus echinatiis L., Oreg.—-Danthonia cadi-
fornica Boland, Oreg.—Deschampsia caespitosa (L.) Beauv., Oreg.—
Festuca elatior L., Oreg.—F. myuros L., Oreg.—Glyceria borealis (Nash)
Batohelder, Oreg.—Holcus lanatus L., Oreg., Wash.—Hordeum gits-
soneanum Pari., Oreg.—Lolium perenne L., Oreg.—L. temidentum L.,
Oreg.—Phleum'prate'nse L., Oreg.
Fischer (1944), by using the New Zealand centrifuge method, found
that the conidial stage of this fungus occurred on many grass seeds
in western Oregon. He also reported apothecia on Lolium perenne
and L. temulentum, and a few on'Hordeum gussoneanum. The writer
found apothecia on L. perenne at Brookings, Oregon, in 1934 but
at that time the organism was not identified. It probably has been
present for a long period of time in western Oregon and probably in
western Washington. Hardison (1948), following suggestions by
Fischer, used a haemacytometer in counting conidia obtained by soak-
ing 30 ml. of seed in 30 ml. of water for 30 minutes. He segregated the
samples into 9 classes depending on the nuniber of conidia found in
a sample measuring ^ g o cu. mm. Thus class 1 had no conidia and the
seed averaged 93 per cent germination during two years; class 3> "very
light," had only 4-6 spores and the seed germinated 91.5 per cent;
while class 5, "moderate," had 16-30 spores i but germinated only 87
per cent. Class 7 was "heavy" with 50-100 spores and 76.5 per cent
germination, and class 9, "extremely heavy,"• had 200-600 spores or
more and germinated 57 per cent. Only classes 1 and 2 (trace) were
safe to leave for another crop. The others were generally plowed under
and another crop was not risked.
Hardison (1948) concluded that seed treatrnent was impractical in
Oregon although vapor heat treatment did control the fungus. There
is sufficient disease-free seed and two-year-old seed to meet the needs
of growers for planting fields for seed production in Oregon. Seed 24
months old in the fall or 20-22 months old in the spring does not pro-
duce apothecia and is safe for seeding (Hardison, 1946, 1947). Hardi-
son also recommended seeding ryegrass with a drill to a depth of at
least one-half inch to eliminate apothecia (1946, 1947, 1948). Burning
fields was recommended in some cases but generally old heavily in-
fested stands should be plowed under. Along this line, elimination of
potentially serious though only mpderately infected fields by plowing
is important because a moderate infection one year may result in heavy
infection the next. Cooperation by the grower in plowing up heavily
diseased fields is important. ''
The identity of the causal organism and the etiology of the disease
itself have been the subject of considerable study within the past dec-
ade. As a result of this work a great deal more is known about the
fungus and the. disease it causes than was known before these several
workers investigated the trouble. /
. The endoconidial stage of this fungus was described as Endoco-
nidium temulentum by Prillieux and Delacroix (1891) and a year
later they described the apothecial stage i as Phialea temulenta (1892).
Some workers confused the conidial stage with the yeast-like fungus
Pullularia pullulans ,but the latter has been found""to beân associated
but very distinct organism which appears to be saprophytic (Wilson,
Noble, and Gray, 1945). Phialea temulenta produces a~ white, fluffy
mycelium, followed by a light to dark chocolate-brown conidial slime,
containing an oily substance which gives it an iridescent appearance,
the oily consistency of the slime distinguishing it from Pullularia pul-
lulans. The same workers (Wilson, Noble, arid Gray, 1945) found that
ascospores and macroconidia gave rise to a branched, septate mycelium
with numerous intertwining coils. After two or three days macroconidia
produced hyphal outgrowths of varying length. They^reraain clustered
around the apex, or become detached aadUre against the hyphae, so
that they may seem to be budded off directly from many' points on
the hyphal wall. In culture the conidia are cylindrical, hyaline, with
rounded ends, 16-29 X 3-5.3 ju (average 21 X 4-3 /j.), are straight or
ASCOMYCETES 99
slightly allantoid; and contain several often polar oil globules. In cul-
ture and on the host, the endogenous conidia occur in pale pink sporo-
dochia. They are unicellular, 3.4-4.8 X 2.7-3.2 /x. Germination of the
microconidia was not noted." These writers found that cold wet con-
ditions favored the disease by prolonging apothecial formation and
the pollination and ripening of ryegrass and increasing secondary
conidial infection. Also they found that the more often the glumes
open before pollination, the more often the ovary is exposed to direct
infection by wind-borne ascospores. They found that if the scutellum
becomes infected, the seed usually fails to germinate. An endophyte
was repeatedly isolated by the authors (see also Peyronel, 1930, among
others) from the pith of flowering shoots of L. perenne and could be
distinguished from Phialea temulenta within the grain by its finer
hyphae and its restriction to the layer immediately outside the aleu-
rone layer, whereas P. temulenta has coarser hyphae and penetrates
to any part of the seed. The authors point out that six fungi showing
afhnities have been recorded on Lolium species. These endophytic
fungi (cf. McLennan, 1926) have been discussed in literature for some
time and have been confused with the blind seed disease itself. They
have been credited with the ability to cause fluorescence under screened
ultraviolet light in L. multifioru7n but not in L. perenne. Wilson, Noble,
and Gray (1945) found that the presence of the blind seed fungus
does not alter this fluorescent activity which they state is caused by
-the radicles of L. multiflorum. They found that heavily infected seed
is generally opaque on the diaphanoscope while slightly infected seed
is generally translucent.
Giinnewig (1933) isolated another seed-borne fungus from three
species of Lolium and this fungus was identified as Chaetomium kun-
zeanum Zopf. The perithecia were 140-200 X 90 X 135 ju,, asci 30-40 X
14-18 fjL, and olive-brown ascospores 9.5-10.5 X 6-8 /t. Inoculation trials
were inconclusive.
Muskett and Calvert (1940) found, in Ireland, that the blind seed
disease and Pullularia were different entities. Calvert and Muskett
(1944) found that Pullularia was nonparasitic on ryegrass. With
Phialea however they obtained heavy infection at flowering time and
comparatively little after seed had set. They isolated the fungus from
sheep's fescue, ryegrasses, velvet bent, Agrostis palustris, Kentucky
bluegrass, velvet grass, and crested dogtail. They obtained infection
on Festuca elatior, Poa trivialis, and ryegrass'fforn inoculura'obtained
from sheep's fescue and the two bent grasses. Seed treatment trials
employed a four-hour tepid steep, which was followed by a hotter steep
at 50° C. for 15 minutes.. Complete control of the disease was obtained.
Apparently no injury followed the treatment if the seed had been
properly and promptly dried. However in field trials reinfection seemed
to occur in many cases from nonseed sources. They concluded that
the organism was not important except as it reduced seed germination.
In a more recent summary of this work (1945) they indicate that seed
' ! • •
kept over a year will be largely free from l)lind seed inoculum. They
found the fields and hedgerows were infested with inoculum which
made seed treatment unreliable.
Some observations have been made on varietal resistance to this
fungus. The workers in England and Irelandj and especially in New
Zealand have reported some indications that there is sofne difference
in susceptibility to this parasite among lines of ryegrass. Corkill and
Rose (1945) report that four lines of ryegrass from Southland (New
Zealand) were appreciably more resistant than plants of certified
origin. Breeding for resistance was suggested.
Gorman (1940) found that the best control of blind seed disease
in New Zealand employing crop management practices stressed early
crops and those accompanied by a dense growth of a pedigree strain
of white clover.
Range: United States, England, France, Ireland, New Zealand, Scotland,
Sweden, Tasmania, Victoria, Wales.
References: Calvert and Muskett (1944,'1946); Corkill and Rose (1945);
Fischer (1944); Freeman (1906); (Glasscock (1940); Gorman (1940,
1945); Gray (1942); Hardison (1945, 1946, 1947, 1948); Hyde (1938,
1945); Muskett and Calvert (1940)'; Neill (1940, 1941, 1942); Neill and
Hyde (1939, 1942); Prillieux -and Delacroix "(1891, 1892); Rose
(1945,a,b); Sampson (1935, 1937, 1939,a); Sampson and Western (1941);
Wilson, Noble, and Gray (1940, 1945).
Phyllachora spp.—^Tar Spots
The graminicolous Phyllachora spp. have been recently mono-
graphed by Orton. Most of the available information is obtained from
his paper.
OpA SPP.
KET TO SPECIES OP PHYLLACHORA
I
(After Orton, 1944)
1. Ascospores arranged uniseriately in the apcus
2. Ascospores spherical or subspherical ,
3. Ascospores averaging 7.5 X 10 /i ^..^^
' P. sphaerospSrmfi-
3. Ascospores averaging 8 X 12 M _
P. ammophilae
3. Ascospores averaging 10.5 X 16 M
P. spartinae
2. Ascospores ovoid / /
3. Ascospores broadly ovoid, average ratio 6f width to length less than
one to two
P. spartinae-
3. Ascospores narrowly ovoid, average ratio of width' to length one to
two or greater ' - /
4. Asci eUiptical, 12-20 /^ wide " - ' '
P. quadraspora
4. Asci cylindrical
P. eragrostidis
ASCOMYCETES 101
2. Ascospores ellipsoid
3. Ascospores broadly ellipsoid, ratio of width to length less than one
to two
4. Ascospores small, 7.5-11 t^ long
5. Clypei oval to elliptical in outline
P. boutelonae
5. Clypei elliptical to fusiform in outline
P. phalaridis
5. Clypei irregular in outline
P. insvlaris
4. Ascospores of medium size, 9-13 M wide
5. Asci narrowly cyhndrical, 8-12 AI wide
6. Asci 60-80 M long
7. Clypei circular to oval in outhne, 0.5-1X0.5-1.5 mm.
P. parilis
7. Clypei smaller, 0.1-0.3 X 0.2-0.8 mm.
* P. paspalicola
7. Clypei circular in outline, large
P. maydis
7. Clypei elliptical to linear
P. graminis
6. Asci 80-115 j« long, clypei elliptical to linear, large
P. luteo-maculata
5. Asci broader, 10-15 M wide
P. serialis
4. Ascospores larger, mostly 12-17 /a long
5. Asci 10-15 M wide
P. ammophilae
5. Asci 15-20 fi wide
P. nervisequia
3. Ascospores narrowly ellipsoid, average ratio width to length 1-2 or
greater
4. Ascospores small, 7.5-10 M long
P. guianensis
4. Ascospores medium-sized, 9-13 j" long
5. Clypei circular to broadly oval in outline
P. paspalicola
6. Clypei not more than 0.5 mm. wide
7. On Paniceae
P. puncturri
7. On Festuceae
P. eragrostidis
6. Clypei up to 1.0 mm. wide
P. vulgata
5. Clypei oval to fusiform in outline, up to 2,0 mm. long
P. wilsonii
5. Clypei elliptical to linear in outline
6. Clypei large, up to 5 mm. long / ,
P. I graminis
6. Clypei small, not more than l^mm. long
P. 'serialii
4. Ascospores large, mostly 12-20 IJ. long
5. Clypei circular to oval in outline
P. \silvtttica
5. Clypei oval to elliptical in outline '
P. erianthii
6, Clypei long-elliptical to linear in outline
6. Clypei amphigenous
7. Ascospores 13-16 /J. long
P. texensis
7. Ascospores 11-14 /^ long
P. coloradensis
6. Clypei chiefly epipliyllous, ascospores 11-14 /i long
P. oryzopsidis
2. Ascospores ovate-acuminate
3. Ascospores narrow, 5-6 /t wide • /
P. leptochloae
3. Ascospores broader, 6-8 /* wide /
4. Ascospores 11-16 ," wide (8-sp'ored form)
y P. quadraspora
4. Ascospores 15-23 M long
5. On Paniceae
P. cornispora
5. On Agrostidae
P. epicampis
3. Ascospores larger, 20-26 X 7.5-9,5 M (4-spored form)
P. quadraspora
2. Ascospores fusiform
3. Ascospores narrow, 4.5-6 <« wide /
4, Ascospores 10-14 t^ long j
P. pammelii
4. Ascospores 14-19 A* long
P. leptochloae
3. Ascospores broader, 5.5-8.5 /t wide
4. Ascospores 15-22 M long
P. cornispora _
4. Ascospores 12-16 f>- long
X P. silvatica
1. Ascospores arranged biseriately or inordinately
2. Ascospores ovoid /
3. Ascospores narrow 4-5 M wide
P. lasiacis
3. Ascospores broader, 6-9.5 M wide
P. quadraspora
4. Ascospores 11-16 /* long (8-spored form^ ^ _ '
P." quadraspora
4. Ascospores 17-26 /* long (4-spored form)
P. quadraspora
ASCOMYCETBS 103
2. Ascospores ellipsoid
3. Ascospores broadly ellipsoid, ratio width to length less than 1 to 2
P. chardonii
3. Ascospores narrowly ellipsoid, ratio width to length 1-2 or greater
4. Clypei mostly hypophyllous
5. Ascospores 4.5-5.5 /i wide
P. diplocarpa
6. Ascospores 6.0-7.5 /J. wide
P. silvatica
4. Clypei amphigenous
5. Clypei not more than 0.5 X 1.0 mm.
P. arundinariae
5. Clypei as much as 1.0 X 2.0 mm.
P. heterospora
2. Ascospores ovate-acuminate
4. Clypei small, up to 0.5 X 1.5 mm.
P. cornispora
4. Clypei larger, up to 1.0X3.0 mm.
P. leptochloqfi
3. Clypei elliptical to Unear in outline
P. epicampis
2. Ascospores fusiform
3. Ascospores narrow, 4.5-6 M long
4. Asci short, 40-70 /x long
P. congruens
4. Asci of medium length, 70-95 M
5. Ascospores 10-14 f. long
P. pammelii
5. Ascospores 15-19 /J. long
P. leptochloae
3. Ascospores of medium width 6.0-8.5 /i
4. Clypei' 0.1-0.4 X 0.1-0.8 mm.
P. arundinariae
4. Clypei 0.4-0.8X0.5-1.0 mm.
P. silvatica
Phyllachora ammophilae O r t o n — T a r Spot

Clypei amphigenous but more conspicuous on upper leaf surface,
scattered, linear, 0.2-1.0 X 1.0-5.0 mm. long, brownish black; fructifi-
cation compound with numerous ascocarps; asci cylindrical, 90-115 X
10-15 11] 8 ascbspores, broadly ellipsoid, 10-15 X 7-9 /n, often appearing
subspherical, uniseriate.
O n : Ammophila arenaria (L.) Link, Mass., N. J., N. Y.
T h i s species was described by Orton from t y p e material collected
at Southampton, N . Y.
Reference: Orton (1944).
/'
/ I
Phyllachora arundinariae Orton^-Tar Spot
Clypei small, amphigenous, scattered, or sometimes gregarious, oval
to elliptical in outline, 0.1-0.4 X 0.1-0.8 mm., black, fructification sim-
ple or compound with few ascocarps; aspi narrowly ellipsoid, 75-100 X
12-18 z^; 8 ascospores, fusiform to narrowly ellipsoid, 15-20 X 6-8.5 n;
biseriate or more rarely uniseriate. '
On: Arundinaria tecta (Walt.) Muhl., Alabama, Ga., S. Car.
This was described from Ravenel's Fungi Amer. 389 from Darien,
Ga. See also P. arundinellae Doidge (Bothalia 4:427. 1942).
Range: United States, Alabama to Georgia; and the West Indies.
Phyllachora boutelouae Rehm—Tar Spot

P. boutelouicola Speg;,
P. chloridicola Speg.
P. minima Chardon
/
Clypei amphigenous but more ^conspicuous onûpper leaf surface,
oval to elliptical in outline, scattered, sometimes confluent, 0.3-1.0 X
0.5-2.0 mm., brownish black; fructification generally compound with
few to numerous ascocarps; asci cylindrical, 75-95 X 9-12 fj.; opercu-
lum prominent; 8 ascospores, broadly ellipsoid, 8.5-11 X 5-6 fi.
On: Bouteloua curtipendula (Michx.) Torr., 111., Iowa, Okla., Tex., Wise.—•
B. gracilis (H. B. K.) Lag., Nebr., N. Mex., N. Dak., S. Dak.—5. hirsuta
Lag., Wise.—Buchloe dactyloides (Nutt.) Englm., Ark., Kans., Okla.,
Tex.—Chloris chloridea (Presl) Hitche., Tex.—C. orthonoton DoelL,
Guatemala—C. submutica H. B. K., Coahuila—C. virgata Swartz, Lower
Calif.
I
Range: United States, Wisconsin west to North Dakota, south to Mexico,
Central America, and in South America. " ^--^__^_^
References: Chardon (1912); Orton (1944); Rehm (ISg?); Spegazzini
(1909, pp. 415, 416). -
Phyllachora chardonii Orton—Tar Spot

Clypei chiefly epiphyllous, scattered, oval to circular' in outline,
0.2-0.3 X 0.3-0.5 mm., black; fructification simple or compound, peri-
thecia flattened, globose; asci ellipsoid, 65-110 X 12-22 ju,; ascospores
broadly ellipsoid, 12-15 X 6.5-8 /x, usually biseriate or inordinate.
- • - /
On: Panicum geminatum Forsk., Fla. - ~^_ -

Range: Southern U. S., West Indies and Philippine Is.
References: Orton (1944); Seaver and Chardon (1926). ,
ASCOMYCETES 105
Phyllachora coloradensis Orton—Tar Spot

Clypei amphigenous, scattered or gregarious, ellipsoid to linear in
outline, 0.2-0.5 X 0.5-1.5 mm., sometimes forming larger groups up to
5 mm. long, dull black; fructification usually compound; asci cylindri-
cal, 70-85 X 9-12 /x, 8 ascospores, narrowly ellipsoid, 11-14 X 5.5-6 /t,
uniseriate, conidia present in similar fructifications, fusiform, 15-20 X
5.5-7 fi, four-celled, hyaline.
On: Muhlenbergia montana (Nutt.) Hitchc, Colo.
Known only from the type locality. Pikes Peak, Colorado, and
-Buckhorn Creek, Colorado (A.S. 17,077). Orton recognized four species
on Muhlenbergia of which all but the common P. vulgata are restricted
in host and geographical range. The material we found along Buck-
horn Creek was growing on a dry bushy slope.
Range: Reported only from Colorado.
Phyllachora congruens Rehm—Tar Spot

P. microstroma Chardon ,
Clypei amphigenous or sometimes epiphyllous, circular or oval in
ôutline, scattered, 0.1-0.3 X 0.15-0.5 mm., sometimes confluent to form
linear series up to 1 mm., black; fructifications simple or compound
with few ascocarps, asci ellipsoid, 40-70 X 10-15 /x, 8 ascospores, fusi-
form or sometimes rounded at one end, 11-15 X 4.5-6 fx, biseriate.
On: Panicum Mans Ell., Alabama—P. longijoUum Torr., N. J.—Paspalum
saccharoides lêes, Panama.
Range: Eastern United States, West Indies, South America, S. Africa and
Philippine Is.
References: Chardon (1928); Orton (1944); Rehm (1914).
Phyllachora cornispora Atk.—Tar Spot

P. acuminata Starb.
P. chaetochloae F. L. Stevens
PJcornispora-necrotica Chardon
P. ortonii Chardon
P. murilloi Garces
Clypei amphigenous but_ often more prominent on upper leaf sur-
face, circular .or oval to ellipsoid in outline, 0.15-0.5 X 0.5-1.5 mm.,
sometimes confluent to form lines up to 7 mm. long, occasionally
forming oval necrotic spots several millimeters long, black; fructifi-
cation usually compound with numerous ascocarps, asci cylindrical,
rarely ellipsoid or saccate, 65-125 X 10-18 /x; 8 ascospores, ovate-
acuminate, more rarely fusiform, 15-22 X 5.5-8.5 /A, biseriate or uni-
/\
seriate, conidia fusiform, often curved, 25-35 X 3-4 jn, 2-celled; sper-
matia (?) cylindrical, filiform, 17-25 X 0.5 [i, continuous.
On: Panicum agrostoides Sprang., Alabama, 111.—P. anceps Michx., Ga.—
P. glutinosum Sw., Canal Zone—P. longifolium Torr., Alabama—Paspalum
distichum L., Fla.—P. jasciculatum 'Willd.,. Costa Rica'—Paspalum laeve
Michx., Fla.—P. virgatum L., Guatemala, Panama.
Range: Southern U. S., Central Americas, S. America, and West Indies.
References: Atkinson (1897); Chardon (1928, p. 116; 1929); Orton (1944);
Stevens (1923).
Phyllachora diplocarpa Ell. and Ev.—Tar Spot

P. nutalliana Fairm.
Clypei hypophyllous, chiefly oval to elliptical in outline, 0.2-0.5 X
0.5-1.0 mm., rarely up to 2.0 mm. long, scattered or sometimes conflu-
ent, forming larger groups, dull black; fructification compound with
few ascocarps, flattened in the scanty mesophyll; asci clavate or some-
what saccate, less often cylindrical,/50-60 X 11-14 jn, often with promi-
nent pedicel; 8 ascospores, narrowly ellipsoid, 13-17 X 4'.5-5.5 /*, uni-
seriate or partially biseriate, or more generally inordinate.
On: Distichlis spicata (L.) Greene, Calif.—D. stricta (Torr.) Rydb., Calif.,
Colo., Kans., N. Mex., N. Dak., Okla., S. Dak., Tex.—Spartina patens
(Ait.) Muhl., S. Car.
Range: On the plains in the United States and along the coastal area.
References: Ellis and Everhart (1897); Orton (1944)._
Phyllachora epicampis Ortoii—Tar Spot

Clypei amphigenous but more corispicuous on upper leaf surface,
scattered or gregarious, elliptical to linear in outline, 0.2-1.0 X 1.0-3.0
mm., dull black or grayish, often confluent to form patches up to 2 cm.
long;'fructifications compound with numerous ascocarps; asci cylin-
drical, to clavate,' 95-150 X 10-14 /A, with long pedicel^, .8 ascospores,
ovate acuminate, 19-23 X 6-8 y., uniseriate or biseriate.
On: Muhlenbergia emersleyi Vasey, Ariz.—M. longiligula Hitchc, Ariz.—
M. rigens (Benth.) Hitchc, Ariz., Calif.
The host sometimes known as Epicamhes rigens Benth. is the basis
for the specific name of this fungus which Orton spells' as epicmnpedis
in his key but epicampis in the technical description (1944). Solheim
collected the fungus on two new hosts in southern Arizona.
Range: Southwestern United States and ôseibly adjacent Mexico.
ASCOMYCETES 107
Phyllachora eragrostidis Chardon—^Tar Spot

Clypei amphigenous, scattered, oval to elliptical in outline, 0.1-0.4
X 0.5-1.5 mm., sometimes confluent to form wider and longer patches,
black; fructification compound with numerous ascocarps; asci cylin-
drical, 80-100 X 8-10 {x.; ascospores narrowly ovoid to ellipsoid, 10-13
X 4.5-6 /x, uniseriate; form on Triodia flava with ascospores 6.0-8.0 X
4.5-5.0 IX.
On: Ei'agrostis capillaris (L.) Nees, Alabama, Nebr., Tex.—E. hirsuta
(Michx.) Nees, Ga.—E. lugens Nees, Okla.—E. -palmeri S. Wats, Tex.—
E. sp., N. Mex.—E. variabilis, Hawaii—Triodia albescens Vasey, Tex.—
T. flava (L.) Smyth, Ga., Tex.
Phyllachora eragrostidis Petrak (1947) was described from Kwan-
tung Province, China, on E. jerruginea. It caused no spots, had asci
50-90 X 8-12 jx; the spores were ellipsoid to ovate, 10-15 X 6-8 {x.. Cer-
tainly the descriptions of these two forms sho^y• similarity. We wrote
to Dr. Petrak on receipt of his paper pointing oiit that his species was
a homonyii.
Range": United States, Georgia to Texas, and South America.
References: Chardon (1939); Orton (19.44).
Phyllachora erianthii Orton—Tar Spot

Clypei amphigenous, scattered or sometimes gregarious in large
patches, chiefly oval in outline, rarely confluent, 0.1-0.6 X 0.2-1.0 mm.,
black; fructification simple or compound with few ascocarps; asci
cylindrical, 75-115 X 9-15 fi; 8 ascospores, narrowly ellipsoid, 13-18 X
6-8 /x; uniseriate; conidia borne in similar fructifications, cylindrical,
3-septate, 23 X 4 û. Spermatia (?) thread-like, 15-20 X 0.5 /x, often
curved.
On: Erianihiis alopecuroides (L.) Ell., S. Car.—E. brevibarbis Michx., Fla.,
Ga.—E. contortus Ell., Ga.—E. giganteus (Walt.) Muhl., Alabama, Fla.
The type is based on Ravenal, Fungi Am. 388.
Reference: Orton (1944). ^
Phyllachora graminis (PersT) Fckl.—Common Tar Spot

(For numerous- synonyms see Orton, 1944, pp. 46-47)
Clypei aihphigenous, scattered, long elliptical to linear, often fusi-
form in outline, 0.1-1.0x0.2-5.0 mm., frequently confluent, black;
fructifications compound with many ascocarps; asci cylindrical, 70-100
X 8-10 /J.; 8 ascospores, ellipsoid, 9-12 X 4.5-6 /x, uniseriate.
O n : Agropyron cristatum (L.) Gaertn., N. S.—A. repens (L.) Beauv., Alta.,
Iowa, Me., Manit., Mass., Minn., N.. H., N. Y.,' Ont., Penn., Que., R. I.,
Vt., W. Va., Wise.—A. smithii Rydb., -Nebr.f-A. subsecundum (Lk.)
Hitchc., Wise.—A. trachycaulum (Lk.) Malte, Manit., N. Y., Wise.—•
Agrostis alba L., Calif., N. Y., Va.—Brachyelytrum erectum (Schreb.)
Beauv., Nebr., Vt.—Bromns ciliatus L., Ind., Wise.—B: purgans L., Penn.
•—B. trinii Desv., Calif.—Calamagrostis canadensis (Miohx.) Beauv.,
Mich., Nebr., N. Y., Que., Wise.—Cinna arundinacea L., 111., Kans., Md.,
Va.—Ehjmus canadensis L., D. C , 111., Ind., Iowa, Kans., Manit., Md.,
Mich., Minn., Miss., Nebr., N. Y., N. Dak., Ohio, Ont., Penn., S. Dak.,
Vt., Va., W. Va., Wise.—E. condensatus Presl, Calif.—E. glaucus Buckl.,
Calif., Mont., N. Y.—E. riparius Wieg., N. Y., Penn., W. Va., Wise—
E. triticoides Buckl., Calif.—E. virginicus L., Ark., D. C , Ga., 111., Ind.,
Iowa, Kans., Ky., Md., Mass., Mich., Minn., Mo., Nebr., N. Y., N. Car.,
N. Dak., Ohio, Ont., Penn., S. Dak., Va., W. Va., ^Yisc.—Festuca elatior
. L., Me.—Hordeum jubatum L., Wise.—Hordeum sp., Calif.—Hystrix
patula Moench, III, Ind., Iowa, Ky., Mich., N. J,, N. Y., Ohio, Ont.,
Penn., Vt., Va., W. Va., Wise.—Panicum virgatum L., Iowa, Kans., Ky.,
Md., Nebr., N. Y., N. C , Okla., R. I.—Pappopharum mucronvlatum Nees,
Tex.—Phleum pratense L., Me., P. E. Is.—Schizachne purpurascens (Torr.)
Swallen, Wise.—Triticum aestivum L., Iowa—Uniola laxa (L.) • B.S.P.,
Ga.
At one time most of the collections of t a r spot on grasses were in-

discriminately assigned to this species b y routine workers. While Orton
has removed m a n y of the collections of t a r spots from this species, it is
still the most common species of the genus on grasses in the temperate
zone. I t is especially a b u n d a n t on grasses in ungrazed thickets,or at t h e
edge of woods in partially shaded areas of relatively" high humidity.
I t is much less prevalent in pastures and in plots in the open while
it is r a r e in grain fields. Orton records only one collection on wheat.
Range: United States, Austria, Belgium, China, England, France, Germany,
Japan, Netherlands, Sweden, widely distributed in the north temperate
zone.
References: Fuckel (1863); Orton (1944) ; ' P e t r a k (1947,a, p. 143); Samp-
son and Western (1941). ^"^—--^
Phyllachora guianensis Stev.—^Tar Spot

(See Orton, 1944, p. 34 for list of synonyms)
Clypei amphigenous, but more conspicudus on upper leaf surface,
scattered, oval, rarely confluent, 0.15-0.3 X 0.2-1.0 mm.,^Klack; fructi-
fication generally simple, occasionally compound; asci cylindrical, 55-
70 X 8-10 /J.; 8 ascospores, narrowly ellipsoid, 7.5-11 X 3.5-5.0 /i, uni-
seriate; conidia sometimes present in similar fructifications, narrowly
ellipsoid, 2-celled, 9.5-12 X 2.0-2.5 /i hyaiineT • '
O n : Paspalum ciliatifolium Miehx., Fla.—P. laeve Michx., La.
ASCOMYCETES 109
Range: Gulf States, West Indies, Central Americas, South America.
References: Chardon (1928); Orton (1944); Theissen and Sydow (1915).
Phyllachora heterospora P. Henn.—Tar Spot

P. raciborskii Theiss. and Syd.
P. seriata Theiss. and Syd.
P. vanderystii Theiss. and Syd.
Clypei amphigenous, gregarious, circular to oval, rarely elliptical in
outline, 0.2-1 mm. wide by 0.2-2 mm. long, often arranged in linear
series up to 5 mm. long, black; fructification simple or compound with
numerous ascocarps; asci ellipsoid to saccate, 12-22 X 75-125 /*; 8 as-
cospores, narrowly ellipsoid, 6.5-7.5 X 13-19 fi, uniseriate or biseriate
and sometimes inordinate.
On: P_ennisetum_ distachyum (Four.) Rupr. Costa Rica.
Range: Congo, Costa Rica, Java, Philippines, South America.
References: Orton (1944); Theissen and Sydow (1915).
Phyllachora insularis Chardon—^Tar Spot

Clypei amphigenous, but more conspicuous on upper leaf surface,
scattered, margin indefinite, irregular in outline, 0.1-0.5 X 0.2-1.6 mm.,
black; fructification compound "with numerous ascocarps; asci cylin-
drical, 60-85 X 8-12 (U,; 8 ascospores, broadly ellipsoid, 8-11 X 4.5-6 ju,
uniseriate, conidia in similar fructifications, lunate, 26-29 X 3-3.6 jn,
one celled, hyaline.
On: Trichachne patens Swallen, Tex.
Range: Southern United States, West Indies, So\ith America.
Phyllachora lasiacis Syd.—Tar Spot

Clypei amphigenous, gregarious, interveined, elliptical to linear in
outline, 0.1-0.2 mm. wide by 0.1-1.2 mm. long, sometimes laterally con-
fluent; fructification simple or compound with numerous ascocarps;
asci long, ellipsoid, 9-12 X 55-75 /i; 8 ascospores, narrowly ovoid, or
ovate-acuminate, 4-5 X 9.5-13 /*, biseriate; conidia (?) present-in simi-
lar fructifications, ellipsoid, 1-1.5 X 3.5 /*, hyaline, continuous.
On: Lasiacis divaricata (L.) Hitchc., Costa Rica.
The type locality is San Jose, Costa Rica. The fungus was also col-
lected on L. swartziana Hitchc. in Puerto Rico.
Range: Costa Rica, Puerto Rico.
References: Orton (1944); Sydow (1925, Ann. Myc. 23:374).
/l
Phyllachora leptochloae. Chardon—Tar Spot
Clypei amphigenous, oval to elliptical, sometimes linearly extended
to produce an irregular outline, scattered, Irarely confluent, 0.3-1 X
0.7-3 mm., black, somewhat shiny; fructifications compound with few
to numerous ascocarps; asci cylindrical, 75-95 X 9.5-13 /t; 8 ascospores,
ovate-acuminate or fusiform, 14-19 X 5-6 /.i; chiefly uniseriate; sper-
matia sinous, 15-20 X 1-0 /*, continuous.
On: Leptochloa virgata (L.) Beauv., Canal Zone, Honduras.
Range: Central Americas to Venezuela.
Reference: Orton (1944). See also Chardon, J. Dept. Agr. Puerto Rico
16:176. 1932.
Phyllachora luteo-maculata (Schw.) Orton—Tar Spot

Sphaeria androipogonicola'^chw.
S. luteo-maculata Schw.
Clypei amphigenous but more'conspicuous on upper leaf surface,
elliptical to linear, scattered, but sometimes confluent, 0.2-1.2 X 0.5-
6.0 mm., black; fructifications compound; asci cylindrical, 75-115 X
9-12 /x; 8 ascospores, broadly ellipsoid, 9-12 X 5-7.5 /i, uniseriate; co-
nidia in similar fructifications, 3-septate, cylindrical, 18-20 X 3-3.5 fi.
On: Andropogon furcatm Muhl., Ida., 111., Mass., Mich., Nebr., N. Y., Okla.,
S. Dak., Wise.—A. hallii Hack., N. jMex.—A. perforatus Trin., Tex.—
A. saccharoides Swartz, Okla., Tex.—A. scoparius Miohx., Ga., Mass.,
Minn., Okla., Penn., Va—A. virginicus L., Alabama, La., N. J., N. Car.,
Va.—A. sp. S. Car.—Sorghastrum nutdns (IJ.) Nash, Iowa.
Range: Eastern half of the United States from the Great Plains to the
Atlantic. I
References: Orton (1944); Schweinitz (1832).
Phyllachora maydis Maubl.—Tar Spat

Clypei amphigenous, generally circirlar in outline, 0.5-2.0 mm. wide
and sometimes confluent to form stripes up to 10 mm. long, black;
fructification compound, ascocarps nearly spherical, immersed in the
mesophyll; asci cylindrical, 80-100 X 8-10 /x., with a short pedicel;
8 ascospores broadly ellipsoid, 9-12 X 5.5-7 /* uniseriaite.
On: Zea mays L., Canal Zone, Guatemala, Mexico. /
Range: Mexico, West Indies, Central Americas, Colombia, and adjacent
South America, Peru.
References: Maublanc (1904); Orton (1944); Weiss (1945, p. 663).
ASCOMYCETES 111 "
Phyllachora nervisequia (Schw.) Orton *—Tar Spot

Sphaeria nervisequia Schw.
S. andropogoni Schw.
Phyllachora andropogoni Ell. and Ev.
Clypei chiefly epiphyllous, scattered, oval to elliptical, black, 0.3-
0.5 X 0.5-1.3 mm., rarely 1 mm. wide and confluent to form stripes up
to 4.5 mm. long; fructification compound; asci cylindrical to narrowly
ellipsoid, 95-125 X 15-19 /x; 8 ascospores, broadly ellipsoid, 12-17 X
7-8.5 ij., usually uniseriate.
On: Andropogon elliotii Chapm., S. Car.—A. glomeratus (Walt.) B. S. P.,
Fla., Ga., Tex.—A. longiberbis Hack., Alabama, Fla.—A. ternarivs Michx.,
Alabama, Ga.—A. sp., La., Miss., Penn., Va.
Range: Atlantic Coast of the United States and in the West Indies.
References: Ellis and Everhart (N.A.F. 2828); Orton (1944); Schweinitz
"(1832).
Phyllachora oryzopsidis Theiss. and Syd.—Tar Spot

P. graminis f. oryzopsidis Rehm
Clypei chiefly epiphyllous, slightly evident on under surface of
leaf, scattered, elliptical to short linear in outline, 0.2-0.4 X 0.3-1.0
mm., black, shiny; fructification compound with numerous ascocarps;
asci cylindrical, 70-100 X 9-10 /j,; 8 ascospores, narrowly ellipsoid, 11-
14 X 5-6.5 /i, uniseriate.
On: Oryzopsis asperijolia Michx., Mass., Mich., Minn., N. Y., Ont., Que.,
S. Dak., Vt.
Orton (1944) lists Rehm's name as a hyponym and a synonym of
P. oryzopsidis (Rehm) Theissen and Sydow. Theissen and Sydow
(1915) describe this fungus as new and make only vague reference to
Rehm's collection. It-is indicated therefore that Rehm's name should
be omitted from the authorities. Bartholomew (F. Col. 3536) credits
P. graminis var. oryzopsidis to Rehm, which is apparently another
hyponym as Rehm apparently listed the fungus as a forma.
Range: United States, Canada.
References: Orton (1944); Theissen and Sydow (1915).
Phyllachora pammelii Orton—Tar Spot

Clypei amphigenous, scattered, elliptical to linear in outline, 0.2-
0.6X0.5-2 mm., black, not shining; fructification compound with
* In recent correspondence Orton points out that this name is a homonym of
P. nervisequia Winter (Hed. 24:29. 1885. Also see Cooke Grev. 13:70. 1885). With-
out critical study it would appear that Orton's fungus could be referred to P.
andropogoni Ell. and Ev. Orton is working on the group.
numerous ascocarps; asci cylindrical, 70-80 X 8-12 p., fusiform, 10-
14 X 4.5-5 11, uniguttulate, uniseriate or occasionally biseriate.
On: Disthichlis stricta (Torr.) Rydb., Colo. Known only from the type local-
ity at Ft. Collins, Colo. • |
Range: United States, as above noted. ''
Reference: Orton (1944). !
Phyllachora parilis Syd.—Tar Spot

Clypei amphigenous but more conspicuous on lower leaf surface,
circular to oval in outline, scattered, 0.5-1.0 mm. wide X 0.5-1.5 mm.
long, brownish black; fructification compound; asci cylindrical, 9-
12 X 60-80 p.; 8 ascospores, broadly ellipsoid, 6.5-7.5 X 10-12 /j. uni-
seriate; spermatia (?) curved or cylindrical, 0.5-1.0 X 15-20 ^.
On: Paspalum candidum (H. & B.) Kunth., Costa Rica.
Type locality is Asseri, Costa Rica. '
Range: Costa Rica to South America. '
References: Orton (1944); Sydow (1927).'
y
Phyllachora paspalicola P. Henn.—Tar Spot

P. vaginata Chardon
Clypei amphigenous, circular or oval in outline, scattered, 0.1-0.3
mm. wide X 0.2-0.8 mm. long, black, shining; fructification simple or
compound with few ascocarps; asci cylindrical, 8-12 X 65-80 ju; 8 asco-
spores, ellipsoid, 4.5-7 X 9-14 /t, uniseriate; conidia in similar fructi-
fications, fusoid, sometimes curved, 2-3 X 20-30 /i, mostly two-celled,
hyaline. ;
On: Paspalum conjugatum Berg., Costa Rica—P. plicatulum Michx., Canal
Zone, Panama—P. tenellum Willd., Panama—-P. sp., Fla.
Originally described ;from Para, Brazil. ^-,^
Range: Central and S. America, West Indies.
References: Chardon (Jour. Dept. Agr. Porto Rico 1932); P7 Hennings
(Hedwigia 48:106. 1908); Orton.
Phyllachora phalaridis Orton—Îj'ar Spot

Clypei amphigenous but more conspicuous on upper leaf surface,
scattered, ellipsoid to fusiform in outline, 0.2-0.5 X 0.3-1.0 mm. long,
shiny black; fructification simple or compound with fdw ascocarps;
asci cylindrical, 70-80 X 7.5-9 ;«,; 8 ascosporea,^broadly ellipsoid, 7.5-
10 X 4-5 [I, uniseriate.
On: Phalaris arundinacea L., Mass. Known only from Southwiek (Seymour
250).
ASCOMYCETES 113
Reference: Orton (1944, p. 39).
Phyllachora punctum (Schw.) Orton—Tar Spot

Sphaeria punctum Schw.
S. panici Schw.
P. panici Sacc.
(See Orton, 1944 for list of synonyms)
Clypei amphigenous, scattered, oval to elliptical, often arranged in
linear series, sometimes coalescing to form lines several millimeters
long, 0.1-0.4 mm. wide by 0.15-1.5 mm. long, less often gregarious,
black, shining; fructification simple or more generally compound, con-
taining several ascocarps; asci cylindrical, 75-90 X 8-10 JA", 8 asco-
spores, ellipsoid, 4-5.5 X 9-12 fx, generally with a prominent guttule,
uniseriate; conidia in similar fructification, fusiform, often curved, 1-3
septate, 12-20 X 2-4 fi; spermatia (?) small, 7.5-9.5 X 0.5 jx, often
curved.
On: Leptoloma cognatum (Schult.) Chase, Tex., Wise.—Oplismenus bur-
manni (Retz.) Beauv., Costa Rica—0. hirtellus (L.) Beauv., S. Car.—
0. humboldtianus, Costa Rica-—0. setarius (Lam.) R. and S., Fla., La.—
Panicum boreale Nash, N. H., Wise.—P. boscii Poir, Fla., Ga., lU., Ky.,
Mich., Mo., N. Y., Wise—P. clandestinum L., Del., D. C, Ga., Ind.,
Towa, Md., N. J., N. Y., Ohio, Penn., R. I., Va., W. Va.—P. commutatum
Schulter, Ga., Miss., Okla.—P. consanguineum Kunth, Ga.—P. depau-
peratum Muhl., Va.—P. dichotomum L., 111., Mich., N. Y., Tex.—P.
euchlamydeum, Wise.—P. flavovirens Nash, Fla.—P. helleri Nash, Tex.—
. P. huachucae Ashe, Ark., Del., Okla., Va., Wise.—P. implicatum Scribn.,
Ont., Wise.—P. joori Vasey, Fla., La.^—P. lancearium Trin., Fla.—P.
lanuginosum EH.', Ark., Del., La., Va.—P. latijolium L., Del., 111., Ind.,
Iowa, Ky., Mich., Mo., N. Y., Ohio, W. Va., Wise.—P. lindheimeri Nash,
Ind., La., Wise.—P. linearifolium Scribn., Ont., Tex.—P. microcarpon
Muhl., Alabama, Ga., N^ Car., Va.—P. pacificum Hitche. and Chase,
Calif.—P. pedicellatum Vasey, Tex.—P. scabriusculum Ell., Alabama, Ga,
—P. scoparium Lam., Alabama—P. scribnerianum Nash, Alabama, Nebr.,
Okla., Wise.—P. sphaerqcarpon Ell, Penn., Va., W. Va., Honduras—
P. tennesseense Ashe, Iowa, Okla., Ont., Wise.—P. wrightianum Scribn.,
N. Y.—P. xalapense H.B.K., Ga., Guatemala, Miss.
Range: United States, Argentina, Canada, Costa Rica, Cuba, Dominican
Republic, Guatemala, Haiti, Honduras, Jamaica, Puerto Rico, St.
Croix Is.
References: Orton (1944); Schweinitz (1832); Stevenson (1918).
Phyllachora quadraspora Tehon—Tar Spot

Clypei amphigenous but more conspicuous on lower surface, scat-
tered or gregarious in areas up to 2 cm. in length, elliptical in outline,
0.1-0.3 X 0.3-1.0 mm. long, fructification simple or compound; asci
114 DISEASES OF CEREALS AND (GRASSES
narrowly ellipsoid to cylindrical, 75-110 X ;15-20 /x; 8-ascospore form
with uniseriate, biseriate, or inordinate arrangement, ascospores nar-
rowly ovoid, 11.5-16 X 6-8 /*; 4-ascospore form with uniseriate or bi-
seriate arrangement, ascospores narrowly oypid to ovate acuminate,
20-26 X 7.5-9.5 /n; conidia lunate, 1-septate, '23-40 X 2:5-3.0 /x.
On: Andropogon fastigiatus Sw., Mexico—A. sp., Fla.—Rotboellia rugosa
Nutt., Miss.
Range: United States, Dominican Republic, Mexico, Panama, Puerto Rico.
Orton includes South America in the range.
References: Orton (1944); Tehon (1919).
Phyllachora serialis Ell. and Ev.—Tar Spot

Clypei poorly developed, amphigenous but more conspicuous on
under surface of the leaf, elliptical to linear in outline, 0.1-0.2 X 0.5-1.0
mm., gregarious, often confluent to form lines up to 5 mm. long; fruc-
tification simple or rarely compound; asci cylindrical, 70-80 ><; 10-15 \>.;
8 ascospores, ellipsoid, 10-13 X 5-6 ^x!, uniseriate.
On: Elymics triticoides Buckl., Calif.—Spartina leiantha Benth., Calif.
W. B. Cooke collected this fungus on E. triticoides a t Point Reyes,
California, within recent years, which appears to be the first time it
has been found since it was originally collected many years before.
Orton (in a personal letter) indicates that he believes that the type
host is misnamed and is probably the same host as Cooke found.
Range: Known only from the coastal region of California in the vicinity of
San Francisco. /
References: Ellis and Everhart (1902); Orton/(1944).
Phyllachora silvatica Sacc. ajnd Speg.—Tar Spot

Clypei mostly hypophyllous, scattered or often^gregarious, roundish
or oval in outline,'0.4-0.8 X 0.5-1.0 mm., black; 7fucti^cation com-
pound with few ascocarps; asci cylindrical or narrowly_elliptical, 75-
100 X 10-15 [J.; 8 ascospores, ellipsoid to fusiform or ovoid, 12-16 X 6.0-
7.5 fi, biseriate or uniseriate.
On: Festuca dertonensis (All.) Aschers. and Qraebn., Oreg.—i^. idahoensis
Elmer, Calif.—F. megalura Nutt., Oreg.-—F. occidentalis Hook., Oreg.—
F. rubra L., B. C, Calif., Oreg.
This species is relatively common in western'Oregon on fescues in
woodlands, waste areas, and sometimes on creeping red fescues in uncut
lawns. • -'T -
The classical spelling of the species name is properly silvatica, as
given by Orton, although Saccardo and Spegazzini (1878) used the
medieval spelling^ sylvatica. The recommendations as listed by Bisby
ASCOMYCETES 115
(1945) are somewhat ambiguous but taken literally it appears per-
missible to use the spelling silvatica in this ease, if preferred.
Range: United States, Canada, England, France, Italy, Wales.
References: Guyot (1932); Orton (1944); Saccardo and Spegazzini (Sac-
cardo, 1878, p. 410); Sampson and Western (1941).
Phyllachora spartinae Orton—Tar Spot

Clypei chiefly epiphyllous, slightly noticeable on under leaf surface,
elliptical to linear 0.5-1.0 X 2.0-3.0 mm., scattered, grayish to shining
black; fructification compound with numerous ascocarps; asci cylindri-
cal to narrowly ellipsoid, 90-110 X 15-20 /t; 8 ascospores, broadly ovoid
to broadly ellipsoid, often appearing nearly spherical, 15-19 X 9-12 ja,
uniseriate.
On: S-partina alternifolia Lois, Fla., Ga., Md.
Range: East coast of the United States.
Phyllachora sphaerosperma Wint.—Tar Spot

P. cenchricola Speg.
Clypei amphigenous, broadly oval, rarely elliptical in outline, often
thickly scattered over considerable areas of the leaf, 0.15-0.4 X 0.2-0.8
mm. long, black; fructification generally compound; asci cylindrical,
65-100 X 10-13 ix; 8 ascospores, nearly spherical, 8-11 X 7-8.5 y., uni-
seriate; conidia associated in similar fructifications, fusoid to lunate,
1- to 3-septate, 24-34 X 3-4 /*.
On: Cenchrm echinatus L,, Bahama Is., Calif.—C. incertus M. A. Curtis,
Miss.—C. viridis Spreng., Canal Zone;
Range: United States, Bahama Is. (Berry Is.), Brazil, Central Americas,
Cuba, Jamaica, Mona Is., Puerto Rico.
References: Orton (1944); Spegazzini (1909, p. 245); Stevens (1927);
Winter (1884, p. 170).
Phyllachora texensis Orton—^Tar Spot

Clypei amphigenous, scattered or sometimes gregarious,- elliptical
to fusiform in outline, 0.2-0.4 X 1.0^1.5 mm., dull black; fructification
compound with numerous ascocarps; asci cylindrical, 100-120 X 7-11
[I, pedicels long; 8 ascosp'ores, narrowly ellipsoid, 13-16 X 6.5-7.5 /*,
uniseriate. ,
On: Muhlenbergia reverchonii Vasey and Scribn., Tex.
Range: Known only from Ft. Worth, Tex.
/
/ i
Phyllachora vulgata Theissen and Sydow—Tar Spot
Sphaeria agrostidis Schw. .~ j
Clypei amphigenous, scattered, oval to ellipsoid in outline, 0.1-
1.0 X 0.2-2.0 mm., dull black, fructification simple or compound, gen-
erally with few ascocarps; asci cylindrical, 65-HOl X 7-11 fi; ascospores
ellipsoid, 9-12 X 4-5.5 ja, uniseriate. '
On: Blepharoneuron tricholepsis (Torr.) Nash, N. Max.—Muhlenbergia
asperifolia (Nees and Mey.) Parodi, B. C, Colo., Mont., Nebr,, N. Max.,
N. Dak., Oreg., Wash.—M. californica Vasey, Calif.—M. capillaris (Lam.)
Trin., Okla.—M. cuspidata (Torr.) Rydb., Ind., Iowa, Minn., Nebr.,
N. Dak., S. Dak., Wyo.—M. foliosa (Roem. and Schult.) Trin.,'Penn.,
Wise.—M. glauca (Nees) Mez., Ariz.—M. mexicana (L.) Trin., 111., Ind.,
Iowa, Md., Mich., Nebr., N. Y., Ohio, Ont., Penn., S. Dak., W. Va.—
M. montana (Nutt.) Hitchc, Ariz., Colo.—Af. pauciflora Buckl., N. Max.
—M. porteri Scribn., Ariz.—M. racemosa (Michx.) B. S. P., Colo., Ga.,
111., Kans., Mich., Nebr., N. Y., N. Dak., Ont., S. Dak.^ Wise'.-M!
reverchonii Vasey and Scribn., Md.—M. schreberi Gmel., Ga., 111., Iowa,
La., Md., Miss., N. Y., N. Dak., Penn., Tehn., W. Ya-.—M. silvatica Torr.,
Kans., Ma., Mo., Wise—M. sobolifera' (Muhl.) Trin., Iowa, Penn.-
M. spiciformis Trin., Mexico—M. squarrosa (Trin.) Rydb., Colo., Mont.,
N. Mex., N. Dak., Wye—M. tenuifolia (Willd.) B. S. P., Wise—M.
uniflora (Muhl.) Fern., Yt.—M. sp., Alabama, R. l.—Sporobolus airoides
(Torr.) Torr., Colo.—S. argutus (Nees) Kunth, Tax.—S. asper (Michx.)
Kunth, 111.—S. cryptandrus (Torr.) A. Gray, Kans., Okla., Tex.—S. sp.,
S. Dak.
This species is very abundant on the plains in species of Muhlen-
bergia. It is moderately important on some grasses such as M.
cuspidata. /
Range: United States, Canada, Mexico. Type locality is Saline River, Kans.
References: Orton (1944); Schweinitz (1832); Theissen and Sydow (1915).
Phyllachora wilsonii Orton—Tar Spot"--- ,_^^

Clypei amphigenous, scattered, oval to fusiform in outline, 0.2-
1.0 X 0.3-2.0 mm., cinerous to black; fructification compound with
numerous ascocarps; asci cylindrical, 60-85'X 9-12 /*; 8 ascospores,
ellipsoid, 9.5-13 X 4.5-7 [n.; uniseriate; spermatia (?) in similar fructi-
fications, filiform, 12-18 X 0.5-1.0 ju; continuous; conidia in' similar
fructifications, 23-30 X 2 /i, 3-septate, slightly curved.
On: Panicum obtusum H. B. K., N. Mex., Okla.—Paspalum ciliatifoUum
Michx., Alabama, Fla., Ga.—P. muhlenbergii l«Jadi, Aliiiama, Va.—P.
pubescens Muhl., 111., Mo., N. Car., Va.—R.-pUbiflorum var. glabrum
Vasey, Tax.—P. setaceum Michx., Alabama, D. C., N. J., S. Car.—
P. stramineum Nash, Nebr., Okla., Tax., Wise—P. supinum Bosc, Mo.—
P. sp.. La., Miss.
ASCOMYCETES 117
Physalospora rhodina (Berk, and Curt.) Cooke—Stalk and Ear Rot

D. tubericola Ell. and Ev. (but see Eddins and Vorhees, 1933).
D. gossypina Cooke
Pycnidia simple or compound, varying in size and complexity with
the host, from 1 to 6 or 7 mm. diam., pycnospores at first one-celled and
hyaline, becoming dark brown and septate usually with irregular, longi-
tudinal striations, 20-33 X 10-18 /*, mostly 24-28 X 12-15 fi. Perithecia
gregarious, black; 250-300 /i diam.; asci typically 8-spored, 90-120 /^
long; ascospores hyaline, 24-42 X 7-17 fi, mostly 30-35 X 11-14 (i.
On: Sorghum vulgare (L.) Pars., Tex.—Zea mays L., Tex.
This species which has a number of synonyms, sometimes occurs
on the grass family, although it appears to be better known on various
tropical and subtropical nongrass hosts. I t was originally collected on
Rosa rubiginosa in North Carolina. Eddins and Vorhees (1933) found
that P. rhodina, P. seicola, and D. tubericola grew and formed pycnidia
on 31 plants when these plants were inoculated in the branches, stems,
or other plant parts with mycelium from pycnospore cultures. These
hosts included peach, fig, sweet potato, watermelon, castor bean, avo-
ca(3o, cotton, papaya, orange, grapefruit, and mango. They suggest
reclassifying the Diplodia spp. Later Vorhees (1942) studied a large
number of isolates of P. rhodina from a considerable range of hosts.
An indefinite number of races were involved. He found that identical-
appearing imperfect fungi had different Physalospora or Physalospora-
like stages.
Range: Widely distributed in the tropics on many hosts.
References: Eddins and Vorhees (1933); Stevens, N. (1926); Vorhees
(1942).
Physalospora tucumanensis Speg.—Red Rot, Leaf Spot, Anthracnose

Conidial stage is Colletotrichum jalcatum Went. (C. lineola Corda
var. halepense Heald and Wolf). Spots circular to slightly elongate, 2
to 5 mm. or more long, margin bright red with gray or dirty-yellow
center bearing a central cluster of black acervuli, red rotting of in-
ternodal stem tissue, acervuli on both surfaces, not abundant, conidio-
phores are short, 15 /^ long, nonseptate, setae pointed with slightly bulb-
ous base, 1- to 2-septate and up to 75 /i long; spores falcate, acute-
pointed, hyaline, 20-27 X 4-5 fi with 1 to 3 granules.
Perithecia mostly scattered, sometimes 2 or 3 together, on various
parts of host plants but abundant on leaf sheaths and leaf blades,
nearly submerged with only a small portion of ostiole protruding,
'!
locally confined, in the main, to the areas between the fibrovascular
bundles and often entirely filling the ap_ace between the bundles, black,
100-260 /J. (wide) X 65-250 fx (tall), the width in longitudinal section of
leaf greater than in cross section. Asci numerous, clayate, thickened at
the apex, 50-118 X 7.4-19.2 /*,, usually ranging from 70-90 X 13-18 ix.
Ascospores irregularly biseriate," single-celled, hyaline, straight or
somewhat fusoid, elliptical to ovate at maturity and, while fresh, con-
taining a hyaline area near center, 12.5-30 X.5-0-11.1 /x, usually rang-
ing from 18-22 X 7-8 /*. Paraphyses very abundant, delicate walls,
septate, usually unbranched, filled with conspicuous granules or oil
droplets, extending to ostiole.
On: Erianthus giganteus (Walt.) Muhl. La., Miss.—Leptochloa dubia (H.
B. K.) Nees, La.—L. filiformis (Lam.) Beauv., La.—Saccharum barberi
Jeswiet, La.—S. ojficinarum L., Fla., Hawaii, La., Miss.—S. ro6iistum, La..—
S. sinense Roxb., La.—S. spontaneum L., La.—Sorghum halepense (L.)
Pars., Alabama, Fla., Ga., La., Miss., N. Car., S. Car., Tex.—S. vvlgare
Pers., Alabama, Fla., Ga., La., Minn., Miss., S. Car., Tex., W. Va.—iS. vvl-
gare var. sudanense (Piper) Hitchc, Miss., Tex.
Went described the conidial stage/of this fungus in 1893 but it was
not until 50 years later that its perithecial stage was reported on
(Carvajal and Edgerton, 1943, 1944). This fungus fruits on all parts
of the plants but the conidial fruiting bodies are particularly common
on the midribs of the leaves. Perithecia form in abundance on crowded
plants, usually some time later than the conidial stage. The perithecia
are submerged in the leaf tissue with only the ostiole protruding. Qross
inoculation trials proved the pathogenicity of the perithecial and conid-
ial strains.
Edgerton and Carvajal (1944) reported on the histology and cytol-
ogy of host-parasite relations in some detail. Abbott (1933) found three
races of the fungus on sugar cane in Louisiana.
Steib (1948) found that apparently h|ealthy sugar cane seed stalks
harbored the fungus and attempts to sterilize such seed stalks were not
entirely successful. ^.^^
Zeman (1921) reported P. phalaridis on Phalarisûlbosa from
Argentina, describing it as new. _
Kiryu (1940) classified the resistance of sugar cane varieties on the
basis of the size of lesions and on the numl3er of nodes of cane involved
in each one. He used a simphfied scale modified from the McKinney
method. >
Forbes, Mills, and Dunckelman (1944) found that seed cane left in
windrows until spring varied according to varieties in their suscepti-
bility to red rot. C.P. (Canal Point) varieties 29/120, 33/310, and
29/103 were said to be nearly ideal for tbe-purppse but 33/243 and.
34/20 were susceptible.
Abbott (1946) reported that red rot is considered the most serious
cane disease in Louisiana. The commercially desirable strains are not
ASCOMYCETES 119
sufficiently resistant. C.P. 28/11 is very resistant but not popular
otherwise. Other resistant recent releases include C.P. 36/105 and
C.P. 36/13.
Khanna and Chacravarti (1949) found that red rot reduced the
sucrose content of cane in Bihar.
Range: United States, Antigua, Argentina, AustraUa, Barbados, Belgian
Congo, Bihar, Brazil, British Guiana, Burma, Central America, China,
Colombia, Cuba, Dominican Republic, Egypt, Fiji, Formosa, Gold Coast,
Grenada, Guadeloupe, Haiti, Hawaii, India, Indochina, Jamaica, Japan,
Java, Madagascar, Madeira, Mauritius, Mexico, Montserrat, Natal, Peru,
Philippines, Puerto Rico, Reunion Is., St. Lucia, St. Vincent, S. Africa,
Sumatra, Surinam, Tanganyika, Trinidad, Uganda, West Indies, general
in the tropics.
References: Abbott (1933, 1935, 1936, 1938, 1946); Atkinson (1939);
Bijl (1921); Brandes and Sartoris (1936); Caravajal and Edgerton
(1943, 1944); Chona and Padwiok (1942); Edgerton (1910,a,6; 1911);
Edgerton and Caravajal (1944); Edgerton, Forbes, Mills, Dufrenoy, and
Luke (1942); Edgerton, Tims, and Mills (1934); Evans (1941); Evans
and Wiehe (1947); Forbes, Mills, and Dunokelman (1944); Khanna and
Chacravarti (1949); Kiryu (1940); Lohman and Stokes (1944); Lucas
(1942); North (1923); Ramakrishnan (1941,6); Rands (1926); Rands,
Abbott, and Summers (1936); Roldan and Tecson (1935); Spegazzini
(1896); Steib (1948); Tims, Mills,..and-Edgerton (1935); Weiss (1945);
Went (1893). (See also Le Beau in Phytopath. May, 1950.)
Physalospora zeae Stout—Gray Ear Rot

Conidial {Macro-phoma zeae Tehon and Daniels)—spots diaphyl-
lous, very extensive, 5-10 X 3-4 cm., irregular in shape, tan with a
slightly darker margin, unlimited. Pycnidia abundant, occurring
throughout the spot, round to oval or even lenticular in cross section,
spherical to applanate in long section, developing in and occupying the
mesophyll, membranaceous, becoming carbonaceous with age, 65-120 /x
diam.; ostiole protruding hypophyllously, usually through a stoma,
nonrostrate, oval, 15-17 X 28-35 /x. Spores continuous, hyaline to green-
ish, fusiform, 17-31 X 6.5-8.5 /i.
Perithecial—^Foliicolous in very large spots, perithecia located in
the mesophyll, opening by a minutely papillate ostiole, externally carbo-
naceous, but microscopically a dark reddish brown, with a pseudo-
parenchymatous wall which is continuous with an- inner • structure of
hyaline pycnosclerotial pseudoparenchyma enclosing the hymenium
and from which the latter appears to arise, globose, 75-235 ju, diam.;
ostiole rounded, 12-30 jx across. Asci cylindrical, straight to curved,
stalked, double-walled, a tiny pore sometimes apparent at the apex,
the inner wall fitting closely to the spore column, the outer wall thick-
ened, especially at the apex, and colorless so as to be seen with difficulty
except for its obscure outer boundary, 85-150 X 13-22 jx. Paraphyses
obscure, hyaline, filamentous. Spores eight per ascus, arranged sub-
/ i
seriately, hyaline t o very dilute olivaceous, one-celled, narrow-ellipsoid,
often tapering to narrow, rounded ends, sometimes presenting one flat-
tened and one curved side, 19-25 X 6.5-8 ft. ' i
O n : Zeo mays L., III.,, Ind. I
Stout (1930) distinguished P . zeae from P. zeicola in t h a t the former
had much larger asci with longer narrower spores. Stout found P . zeae,
Macrophoma zeae a n d a pycnosclerotial form associated. Recently
Ullstrup (1946) determined t h a t a gray rot of corn was the same as
P . zeae and t h a t M. zeae was the pycnidial stage of P . zeae. H e also
found a microspore stage. The ear rot injured as much as 10 per cent
of the crop in some areas in I n d i a n a in 1944. I t began as a premature
bleaching of t h e husks, followed by cementing of these husks. A white
felty mycelium develops on and between the kernels and, when the ear
is broken, the pith has a characteristic slate gray discoloration dotted
with black sclerotia. T h e mycelium on the kernels becomes speckled
with black sclerotia a n d the kernels often s h o w black streaks starting
from the base. B e n e a t h the seed coats of such kernels black stroniatic
layers of mycelium often form. Completely rotted ears have dark gray
husks, and cobs and kernels are reduced t o ' a dry, gray-black, pithy
mass. Ullstrup (1946) mentions t h a t the^ symptoms caused b y - t h i s
fungus are easily confusable with Diplodia ear rot b u t it m a y be dis-
tinguished as follows:
1. Both the presence of sclerotia and the slate gray color in the cob
in early stages are distinctive in Physalospora.
2. Gray mycelium between the husks and the black sclerotia be-
tween the pericarps of the kernels are further distinguishing features.
3. In advanced stages, the gray-black color and dry pithiness of
completely rotted ears are distinct from Diplodia symptoms.
Ullstrup based the name "gray ear rot" on j the distinctive gray
color of the husks and the appearance of the interior of the cob.
TJllstrup iound identical gross morphology of cultures isolated from
M. zeae, P. zeae, and the gray ear-rot fungus. All three grew at the
. same rate and all produced niicrospores, 2-4.8 X 1-2 /* and pycnospores
of M. zeae on sterilized corn stalks. Artificial inoculation of ears frofn
the various isolates all produced gray ear rot. —
References: Stout (1930); Tehon and Daniels (1927); Ullstrup (1946).
/
I
Physalospora zeicola Ell. and Ev.—Stalk and Ear Rot /
Diplodia frumenti Ell. and Ev. (Fig. 13,C)
Causes an ear and stalk rot, the invaded kernels__become black,
mycelium brown, pycnidia are embedded in stroma.tfc'^masses of my-
celium on the ears and husks; pycnospores short cylindric to subellip-
tical, 1-septate, brown, 21-29 X 10-14 [JL, mostly 25-27 X 12-13 /*.
Perithecia gregarious, 250-330 m, covered by the cuticle of the 'stalk
ASCOMYCETES 121
which is shghtly raised and pierced by the obtusely conical, black and
shining ostiole. Asci clavate-cylindrical, 75-80 X 12-15 /* (95-140 X 10-
13 IX in Florida material) nearly sessile, with abundant paraphyses,
ascospores crowded-biseriate, elliptical or almond-shaped, hyaline,
granular, 1^-20 X 8-10 /i.
On: Zea mays L., Fla., La., Okla.
Eddins reported D. jrumenti on corn from Florida (1930,a). He
stated that it was morphologically very close to D. natalensis (see P.
rhodina, above), D. tubericola, and D. gossypina. Eddins and Vorhees
(1933) later showed that D. -jrumenti was the conidial stage of Physalo-
spora zeicola. P. zeicola differs from P. rhodina in having shorter and
narrower ascospores.
Range: United States, Tanganyika, probably more widely distributed than
records show.
References: Eddins (1930,a,6,d); Eddins and Vorhees (1933); Ellis and
Everhart (1886, p. 103; 1890).
Placostroma bambusae (Turc.) comb. nov.—On Culms

Scirrhodothis bambusae (Turc.) Trotter
Melanconium bambusae Turc.
Scirrhia bambusae Turc.
Stromata ellipsoid elongate, sublinear, 1-2 X 0.5-1 mm. diam., gre-
garious, somewhat seriately arranged, locules in the stromata numerous,
mostly 2-3 seriate, subglobose to ovate, 180-200 ft. diam., ostiole obscure
to briefly prominently papillate; asci cylindrical-clavate, obtusely
rounded at the upper part, lower part subattenuate or bluntly stipitate,
paraphyses filiform, guttulate, 8-spored, spores in 2 rows to obliquely
one-rowed, oblong-clavate, downward attenuate to more or less curved,
distinctly 1-septate near base, granulose, guttulate, hyaline 20-24 X 8-
12 /t. (From Sylloge v. 24.)
On: Bamboo, Fla., Ga.
The conidial stage {Melanconium) is said to be associated with a
blight of bamboo shoots.-Little information appears to be available on
this fungus. Clements and Shear (1931) placed Scirrhodothis under
Placostroma, hence the above combination.
Range: United States, Italy.
References: See Saccardo (Sylloge Pungorum v. 24, p. 611); Clements and
Shear (1931).
Placostroma sporoboli (Atk.) comb. nov.—On Culms

Scirrhia sporoboli Atk.
Stromata 1-3 X 0.2-0.5 mm. diam., narrow, black, white inside,
erumpent, locules small, 120-150 jn, very briefly rostrate, asci clavate.
122 DISEASES OF CEREALS AND GRASSES .
somewhat a t t e n u a t e at base, 50-70 X 8-9 /x, paraphyses filiform; spores
in 2 series, fusoid, inequilaterally 1-septate, hyaline, 12-16 X 3-4 fj..
O n : Muhlenbergia asperifolia (Nees and Mey.) Parodi, Ok\a,.—Sporobolus
asper (Miohx.) Kunth, Alabama.
Preston (1947) reported this fungus from Oklahoma. It is doubted
if the species is parasitic. Atkinson (1897) said' that the fungus dif-
fered from S. rimosa in the prominent pBrithecia and in the size and
shape of the spores and asci. This fungus apparently belongs in Placo-
stroma.
References: Atkinson (1897); Preston (1947); Theissen and Sydow (1915,
p. 418).
Pleospora sp.—Brown Turf Spot
Dunlap (1944) reported an undetermined species of Pleospora on
Poa pratensis L. in Texas. Brown spots'4-5 inches in diameter were
first noted in August in new lawns at .^marillo and small dollar-size
spots at Seguin, Texas, in November, 1943. Perithecia comparable to
Pleospora were found in both collections. The material from Seguin
showed perithecia on dying leaves indicating a possible parasitic na-
ture, while those in the specimens from Amarillo were on dead leaves.
The spores from this material in February were larger than those from
Seguin. The cause of this condition remains uncertain.
References: Dunlap (1944). See also Clinton (1934, p. 215).
Pyrenophora avenae Ito and Kuribayashi—Oat Leaf Blotch

This fungus will be discussed under \ Helminthosporium avenae
Eidam. Pleospora avenae Schaffnit and Rathschlag is apparently the
same fungus although this point needs further stu4y^ Most workers
appear to favor use of'the Japanese name for the perfectTBtage of this
species. Wehmeyer (1949) gives Pleospora avenae Died, as-a synonym
of Pleospora trichostoma (Fr.) Ces. and de Not. Pyrenophora tritici-
repentis (Died.) DrechsL, Py. bromi (Died.) Drechsl., and Py. teres,
(Died.) Drechsl. get the same treatment. .
Pyrenophora bromi (Died.) Drechsler—Leaf Spot of Brome

Recent work by Chamberlain and Allison (1944, 1945) indicates
that the perithecial stage of this species is ve£y~important in its life
history. Because it is so well known under its'corlidial stage, Helmintho-
sporium bromi Died., it too will be discussed under Helminthosporium.
See Wehmeyer's recent paper (1949).
ASCOMYCETES 123
Sclerotinia homoeocarpa F. T. Bennett—Sclerotinia Turf Rot,

Dollar Spot
Rhizoctonia monteithianum F. T. Bennett
Forms small spots on turf about 2 inches in diameter, producing on
nutrient agar reddish brown, abundant aerial mycelia;; sclerotia of
parchment-like thickness, variable in size; apothecia illy formed, pro-
ducing asci; ascospores 16-17 X "5-2-6.5 /i. Sometimes cupulate bodies
produce hyaline, oblong, uniseptate conidia averaging 19.5 X 4.7 /* with
slight median constriction and tapering ends.
On: Agrostis alba L., Mass., Penn., R. I., Va.—A. canina L., Mass., R. I., Va.—
A. palmtris Huds., D. C, Mass., Penn., R. I., Va.—A. tenuis Sibth., D. C,
Mass., Ont., Penn., R. I., Va.—Festuca ovina L., Mass.—F. rubra L., Mass.
—Poa annua L., Mass.—P. pratensis L., Mass., Penn., R. I. Other reports
not specific, widely distributed in northeastern United States. While it is
alleged to occur in the Pacific Northwest we have never had any material
available for isolation.
The small brown, later bleached turf spots are sometimes known as
dollar spot (Monteith and Dahl, 1932). The disease is said to be com-
mon in the eastern United States and occurs westward at least to
Minnesota (Tervet, 1942). It is probably less important in this country
than snow mold [Fusarium nivale {FT.) Ces.] or brown patch (Rhizoc-
tonia solani Kuehn). Sampson And Western (1941) are of somewhat
the same opinion regarding the fungus in Great Britain. It appears to
have been unduly publicized although it is disfiguring to well-kept,
closely cropped lawns and greens.
Tervet (1942) was able to prevent dollar spot from spreading on
unspecified turf of golf courses in Minnesota in 1941 and 1942 by using
Thiosan (50 per'cent tetra-methyl-thiuram-disulfide) at the rate of 1
lb. per 6,000 sq. ft. of turf at 2-week intervals. He was considerably
less successful in eliminating well-established stands of dollar spot and
brown patch. He was able to control the diseases by using the standard
mercuric chloride and mercurous chloride mixture (1942).
There appear to be several races of this species in Great Britain.
Further study is needed in this country.
Brien (1935) reported Sclerotinia trijoliorum on turf in New
Zealand. I t developed in small circular patches 6 to 8 inches in di-
•ameter, at first yellow, later b^rown, and finally dying -out. -It was
sometimes associated with brown patch and pink patch. I t was con-
trolled by using 3 oz. of mercuric chloride in 50 gal. of water applied
to 2,000 sq. ft. of lawn.
Sclerotinia gramineariim Elen. is widely known in U.S.S.R. as a
snow mold of winter grain and grasses. Tupenevitch and Shirko dis-
cussed control measures (1939). Solkina reported on life cycle studies
(1939), Yakovleff on its biology (1939), and Fokin on meteorology
(1939). This organism has not been reported in North America but is
124 DISEASES OF CEREALS AND GRASSES'
/I
likely to be found. Ekstrand (1946) report^ that Sclerotinia borealis
was common in Sweden, Finland, and. U.S.S.R. in the northern parts of
these countries on winter cereals. Timothy, rye, and some grasses were
injured, sometimes severely.
Range: United States, Australia, England, Scotland and Wales. Probably
widely distributed in continental Europe in edoler regions and in Canada
in localized areas.
References: Anderson, Carl (1944); Bennett (1935,a; 1937); Howard and
Keil (1947); Keil (3944); Monteith and Dahl (1932); Sampson and West-
ern (1941); Sprague (1946,e); Struble (1943); TTervet (1942);'Tyson
(1936).
BASIDIOMYCETES
This group includes a number of well-known parasitic fungi,
especially some of the sclerotium-forming species which cause root
rots and culm rots. They tend to be less restricted in their host speciali-
zation than some other groups and they are often, because of their
sclerotia, difficult to control. While the Basidiomycetes are common in
the tropics and in the warm temperate zones some forms prefer very
low temperatures, scarcely above freezing. Considerable taxonomic
work has been done with this group during recent years. However,
some of the sterile forms are still unclassified.
The following key to the Basidiomycetes includes such a motley
array of genera that it is by necessity somewhat involved:
A. Fruiting body a mushroom with gills (Agaricales)
B. Spores hyaline Marasmius
BB. Spores ochraceous to dark ferruginous Naucoria
BBB. Spores dark or black Coprinus
AA. Fruiting body a delicate stalked obclavate structure without gills, aris-
ing from a sclerotium, basidia 'external, sclerotia formed in late winter,
the basidiophore the next late fall Typhida
AAA. Fruiting body, if present, not as above
B. Forming a subepidermal black leaf or blister smut, vaguely resembling
a tar spot, hence included in this text Entyloma
BB. Not as above
C. Forming a yellow, firmly gelatinous body on leaves; basidia with
2 sterigmata ; Discellomyces
CC. Otherwise
D. Forming coral red gelatinous growths on prostrate grasses in wet
and cool weather Corticium (fuciforme)
DD. Otherwise
E. Basidiospores produced on delicate thin-hypochnoid growth,
hyphae branching at right angles, basidia with 4 sterigmata (See
Rhizoctonia, its well-known myceUal stage) Pellicidaria
EE. Fruiting body not present
F. Hyphae relatively coarse, branching at right angles and often
pinched at branches, stromata formed are brown, pink or darker
Rhizoctonia (see Pellicularia)
FF. Sclerotia firm, black rind, interior of sclerotia paler
Sclerotium *
FFF. Hyphae delicate or variable
Undetermined sterile Basidiomycetes
* See also Typhula and such Asoomycetes as Leptosphaeria salvinii and Sclero-
tinia. Sclerotium as a form-genus belongs in the Fungi Imperfect! but is treated
here because most of the remaining unassigned forms appear to be basidial.
125
l\
Corticium fuciforme (Berk.) Wakef.—Pink Patch
Tufts slender, filiform, bright coral pink, somewhat fleshy algal-like
growth with a delicate, effused, somewhat gelatinous, coral pink basal
incrustation on leaves and culms, forming more or less circular patches.
Basidia slightly tinged pink, clavate, with 2-4; stout, curved sterigmata,
5.5-7 jx diam. Spores pip-shaped, one-sided, depressed, hyaline, apicu-
late, 11-12.5 X 5-6 jn. Hyphae tinged pink, with clamp connections,
thin-walled and rather vacuolate, 2-4' /x diam. Walls of hyphae slightly
gelatinous.
On: Agrostis alba L., R. I.—A. canina L., R. I.—A. palustris' Huds., R. I.,
Wash.—A. tenuis Sibth., R. I.—Briza media L., R. I.—Festuca myuros
L., Oreg.—F. ovina var. duriuscula (L.) Koch, R. I.—F. rubra L., R. I.—
F. rubra var. commutata Gaud., Oreg.—Poa annua L., R. I., Wash.—
P. pratensis L., R. I.
In England pink patch is common on red fescue and it also occurs
on Agrostis tenuis, Poa annua, Holcu's lanatus, Lolium perenne, and
others. It forms a bright coral pink, thinly gelatinous growth in small
areas on fescue lawns in western Oregon during spring and fall wet
weather. We have seen fragmentary material from Bremerton, Wash-
ington, in late June on thin sod, composed mostly of Poa-annua. It de-
veloped in early spring at Pullman in 1949, associated with Fvârium
nivale. The gelatinous growth in better material covers leaves and
culms and is very conspicuous when the lawns are wet. The growth con-
sists of parallel branching hyphae bound together by anastorrioses and
gelatinous walls. The strands of bound hyphae are readity recognized
even in fragmentary material of the fungus. The thin gelatinous layer
may bear outgrowths nearly half an inch lon^/resembling spear points
but often with short fucus-like branches. Fine hyphae emerge from the
edge of the layer, spreading concentrically, increasing in thickness, and
finally bind the grass in a wet, pink mat! The basidia are borne on the
surface of the mass, forming a nearly complete hymenium. The fungus
is spread by the multitudinous spores and by bits oTmycelium, either
wet or dry. McAlpine (1906) discussed the fungus in some detail.
Various control measures have been proposed for the control of the
disease. Bennett (1935,c) proposed promoting vigorous growth and
using a suitable turf drench or dust applied at the correct time. He
suggested use of mercurial treatments or a 'combination of malachite
green and Bordeaux mixture. More recent investigations with asso-
ciated fungi, such as work conducted at Rhode Island Agricultural Ex-
periment Station, no doubt also held this fungus in check, although
there is very little specific information on tliifparticular fungus.
Erwin (1937) reported that the optimufn temperature ior growth of
this species was between 18° and 20° C , the temperature range for
growth being 1° C. to 30° C.
BASIDIOMYCETES 127
Range: United States in New England and the Pacific Northwest, Austraha,
England, New Zealand, Scotland, Wales.
References: Bennett (1935,c); Berkeley (1873,a,6); Erwin (1934, 1937,
1941); Howard and Keil (1947); McAlpine (1906); Sampson and West-
ern (1941); Sprague (1940,a); Sydow and Sydow (1906); Wakefield
(1916).
Discellomyces gloeosporus Olive—Leaf Spot

Fructification hypophyllous, on discolored spots, small, yellow,
firmly gelatinous, pulvinate to nearly discoid, with blunt base inserted
in the leaf, hyphae penetrating into the leaf tissue, measuring 440-
700 X 485-955 /x, and 194-353 /x tall, becoming dark brown and shrunken
with age, hymenium covering entire upper surface of the fructification,
composed of closely packed, long-stalked, clavate to nearly pyriform
probasidia, 4.1-6.3 /* diam., the latter persistent and giving rise to
narrow basidia of varying lengths; basidia 2.4-4.1 X 9-25.2 /x, produced
externally to the gelatinous surface; scattered among basidia there are
numerous, very slender unbranched paraphyses with knob-like swell-
ings at the tips; basidia give rise to 2 relatively short sterigmata,
1.8-4.0 /ji. in length; basidiospores allantoid, 2.7-4.5 X 8.6-11.9 /*, pro-
duced in adherent pairs, several pairs often clinging together in a 'ball;
apiculus not apparent, basidiospores frequently becoming 1-4 septate,
often budding out small, more, or less globose conidia.
On: Arundinaria tecta (Walt.) Muhl., Ga., N. Car.
The first symptom is the appearance of small yellow spots on the
leaves. They enlarge, becoming dark brown with a yellow halo. The
small yellow fructifications develop in these spots. In severe infesta-
tions badly diseased leaves may become brown and dead. The fruits
may occur singly or in groups. They arise from within the leaf, rup-
turing the overlying leaf epidermis as the fungus fruit grows. The
fruiting bodies are composed of an inner sterile area and an outer
hymerial layer which covers most of the exposed surface.
Olive (1945) placed this fungus in the Dacryomycetales creating
the new genus Discellomyces to accommodate it.
Range: S. E. United States.
References: Olive (1945, 1947).
Entyloma spp.—Blister Smut on Leaves

The blister smut of grasses, especially Poa spp., should be men-
tioned here because it' can be and has. been confused with tar spot
fungi such as Phyllachora spp. The stroma-like subepidermal chlamydo-
spores form gray-black, blister-like areas on the leaves of Poa pratensis,
P. compressa, P. juncifolia, and P. annua in Oregon and adjacent Wash-
ington. The conidial phase of the fungus forms a fine, dried, amorphous
i\
series of dots scattered seriately along the surface of the bUsters. The
Uamulana-Yike. sporidia of Entyloma irregulare Johans. from Poa
annua germinated readily on potato-dextrose agar and produced flesh-
colored mucose masses of sporidia (Sprague, 1946,e).
Entyloma sp. {E. bingenends Zundel, in herb.) occurs abundantly
on Poa vaseyochloa Scribn. along the rocky, warm slopes near Bingen,
Washington, toward Lyle, Washington. The fungus is found on new
growth in March and was found once in February (Sprague, field
notes).
Two possibly identical species occur widely in the United States on
Poa grasses, E. crastophilum Sacc. and E. irregulare Jdhan. (?) E.
spragueanum is a fragment of doubtful specificity (Zundel, 1942).
There are other species of Entyloma on other species of grasses
(Fischer, 1950) but for the purpose of this text the group on Poa is
sufficiently illustrative.
Reference: Fischer (1950).
Marasmius sacchari Wakker—Associated with Root Necrosis

Bases of the fragile white pilei gregarious or fascicled, pilei variable,
carnose-membranaceous, persistent,'^tardily campanulate, then becom-
ing dirty white, flattened to cup-shaped, 15 mm. diam., lamellae white,
simple to bifurcate; stipe central, white, 15 mm. long, apex tubiform,
base villose; mycelium white; spores hyaline, continuous, irregularly
oblong, somewhat tapering toward the rounded end, 16-20 X 4-5 jn. 1
The var. hawaiiensis Cobb has spores 12-16.4 X 4.2-4.9 /A (Sac-
cardo, Sylloge v. 23, p. 161).
On: Saccharum officinarum L., Hawaii, La. /
Injured plants make poor growth; the lower leaves die and are often
bound together and bound to the base of the plant by a white mycelial
weft extending both above and below the soil level. The young canes
are sometiimes killed and the roots onînjured plantgjaecome necrotic.
There appears to be no completely convincing infornration on the
role of this and various other associated fungi as the cause of decay
in growing sugar cane. Cook, M. T. (1932,c), however, found-that the
fungus could attack and often destroy the leaves, stems, and roots of
young plants. Nowell (1923) discusses the diseases in some detail in
his text which deals with the fungi on crop plants in the Lesser Antilles.
Wilbrink (1923) found some evidence that hot water steep tended to
reduce injury in setts. ,
A number of other toadstools and related forms onyGramineae have
been described which will be briefly mentioiied"here. Tehon (1924) re-
ported Marasmius sp. (near M. scorodonius Fr.) growing directly out
of the basal node of dead wheat culms in Illinois. He believed that the
fungus was parasitic. The next year Young, P. A. (1925) described
BASIDIOMYCETES 129
M. tritici on wheat, rye, barley, quack grass, and an undetermined
grass in central Illinois. He found that there was no injury to the crop
but he believed that the fungus developed while the host was still
green. Boewe (1938) described Naucoria cerealis as a weak parasite
of wheat, barley, and rye in Illinois. He later (1939) issued a key to all
species of Naucoria known or associated with living plants. We have
collected Naucoria spp. and other mushrooms, growing on grass in
widely scattered parts of the United States. These forms spring up
after rains and seem to be saprophytic, utilizing decayed plant parts
for growth. Similarly we encountered Pholiota dura (Bolt.) Fries and
P. praecox (Pers.) Fries attached to cereals in Oregon (Sprague,
1938,e). Inoculation trials with these fungi showed that they decom-
posed cellulose and increased the yield of inoculated plants over that
obtained from checks. Of course when some of these forms produce
fairy rings they interfere with growth of grass, causing disfiguring
circular bands in lawns or pastures seeded to perennial grasses. Bayliss
(1926) disagrees with Shanz and Piemeisel (1917), who were of the
opinion that injury was occasioned by the drought produced by the
presence of excess mycelium in the soil in the fairy ring areas. Bayliss
considered that the death of the grass in the presence of Marasmius
oreades was due to parasitic activity. She also said that both Clitocybe
gigantea and Tncholoma gambosuni affected grass in England similarly.
Mains (1925,6) found clumps of Festuca capillata Lam. dying
'during the summer in Indiana. He reported a mushroom on these
clumps which resembled M. interstitians.
Vincens (1921,a) claimed that Schizophyllum commune Fries was
an actual parasite on sugar cane in Indochina.
Faris (1927) described a zonate basal culm rot of sugar cane in
Cuba with Fomes sp. associated.
Hanna (1939) reported Coprinus lirticaecola on stems of Marquis
wheat. We have noted a minute delicate mushroom with a campanu-
late pileus growing on dead sheaths of Bromus mollis which were grow-
ing in dense grass stands in exceedingly wet weather at Pullman, Wash-
ington, in mid-June, 1948. The fruiting bodies were approximately 6
mm. tall and possibly 4 mm. wide, pale fawn-colored.
Thompson (1940) reported Marasmius semiustus (M. stenophyllu^)
as the cause of death of dry-land rice in Malaya. Poor soil conditions
favored infection. Petch (1947) reported M. paspdli n. sp. asprobably
parasitic in Ceylon.
Range (of M. sacchari): United States, Tropics and subtropics, Argentina,
Australia, Barbados, Brazil, Burma, Egypt, Hawaii, Indochina, Java,
St. Lucia Is., St. Vincent Is., Trinidad, West Indies in general.
References: Cook, M. T. (1932,c,d); Fulton (1908,a); Howard (1903);
Johnston and Stevenson (1917); Nowell (1923); Saccardo (Sylloge,
V. 14, p. 115; V. 23, p. 161); Vincens (1921,b); Wakker and Went (1898);
Wilbrink (1923); Williams (1921).
i\
Pellicularia filamentosa (Pat.) D. P. Rogers
Brown Patch, Root Rot, Sharp Eyespot
Rhizoctonia solani Kuehn j
Fructification forming a delicate separably] flaky: or thin-hypoch-
noid pellicle; when dry it is white to buff-colored or semi.translucent;
hyphae branching at right angles and with some cruciform cells, with-
out clamps, 4.5-14(17) /A diam., the basal ones long-celled and fre-
quently tinted, the subbasidial ones hyaline, short-celled; frequently
with doleiform segments, branching abundantly and bearing basidia
in small imperfectly symmetrical cymes; basidia subcylindric and
doleiform, obpyriform or clavate, 12-18 X 8-11 /x, bearing 4 sterigmata
which arise as blunt knobs, later horn-shaped, 3-20 X 1.5-4.5 j«,, mostly
5.5-12 X 1.5-3.5 jLi; spores ellipsoid or oblong-ellipsoid, thin-walled,
flattened on the inside, a little broader below the middle, truncate-
apiculate, 7-12.5 X 4-7 fi, occasionally .germinating by a stout pro-
mycelium on which is born a similar secondary spore.
The vegetative stage (R. solani) fprms tan- to brown mycelium in
pure culture, 4-15 jx diam., with rightrangled branching and character-
istic constriction at the septa; stroma somewhat .cheesy, later firm,
flattened to subglobose, pale buff, brown to dark chestnut brown, my-
celium variable in diameter but usually 2-5 /t.
On: Agropyron angustiglume Nevski, N. Dak.—4. ciliare (Trin.) Franch,
N. Dak.—A. cristatum (L.) Gaertn., Mont., Nebr., N. Dak., Oreg,, S.
Dak., Wash., Wyo.—A. dasystachyum (Hook.) Scribn., Oreg.—A. deser-
torum (Fisch.) Schult., N. Dak., Oreg.—A. elongatum''{Host.) Beauv.,
Nebr., N. Dak.—A. intermedium (Host.) Beauv., N. Dak., S. Dak.—
A. michnoi Roshev., N. Dak., S. Dak.-—A. monjolicum Keng, N. Dak.—
A. repens (L.) Beauv., Ida., Minn., Nebr., N. Dak., Oreg., S. Dak.,
Wash.—A. riparium Scribn. and Sm., N. Dak.—A. sibiricum (Willd.)
Beauv., Mont., N. I)ak., Oreg., S. Dak., Wash.—A. smithii Rydb., Mont.,
Nebr., N. Dak., Oreg., S. Dak., W:'o.—A. subsecundum (Lk.) Hitchc,
Ida.—A. trachycaulum (Lk.) Malte, Mont., Nebr^^N. Dak., S. Dak.,
Wyo.—A. trichophorum (Lk.) Richt., N. Dak.—Agrostis-alba L., Conn.,
D. C, Md., Minn., Nebr., N. Dak., Ohio, Oreg., Penn.;_Va., Wash.—
A. canina L., D. C , Oreg., Penn., R. I., Va.—A. exarata Trin., Oreg.,
Wash.—A. hallii Vasey, Oreg.—A. palmtris Huds., Calif., Conn., Del:,
D. C , 111., Md., Mass., Mo., N. J., N . Y., Ohio, Ont., Oreg., Penn.' R. I.,
Sask., Va., Wash., W. Va., Wise.—A. stolonifera L., Oreg., R. I.—A. tenuis
Sibth., Conn., D. C , Ont., Oreg., Penn., Que., R. I., Va., Wash.—
Arrhenatherum elatius (L.) Beauv., Calif., Oreg., Wash.—Avena byzan-
tina K. Koch, Oreg., Wash.—A. jatua L., N. Dak., Oreg., Wash.—A. sativa
L., Iowa, Minn., N. Dak., Oreg., S. Dak., Wash., Wyo^—Axonopus affinis
Chase, Hawaii, Southern United States (Weijg,-1943)—BoMteZowa curti-
pendula (Michx.) Torr., N. Dak., S. Dak:—B-. gracilis H. B. K., Mont.,
N. Dak., S. Dak.—Bromus arvensis L., N. Dak.—5. carinaius Hook, and
Arn., Mont., N . Dak., Oreg., Wash., Wyo.—5. catharticus Vahl, Oreg.—
B. erectus Huds., N. Dak., Oreg.—B. inermis Leyss, Ida.,'Minn., Nebr.,
BASIDIOMYCETES 131
Oreg., S. Dak., Wash.—B. japonicm Thunb., Minn., Mont., N. Dak., S.
Dak., Wyo.—B. madritensis L., Oreg.—B. mollis L., Oreg., Wash.—B.
pumpellianus Scribn., Wyo.—B. racemosus L., Oreg.—B. rigidus Roth,
Oreg., Wash.—B. secalinus L., Ida., Oreg., Wash.—B. tectorum L., Mont.,
Ida., Nebr., N. Dak., Oreg., S. Dak., Wash., Wyo.—Buchloe dactyloides
(Nutt.) Englm., Nebr., N. Dak.—Calamagrostis montanensis Scribn.,
S. Dak.—Cenchrus pauciflortis Benth., N. Dak.—Cynodon dactylon (L.)
Pers., Ariz., Hawaii, La., Tex.—Dactylis glomerata L., Mont., Oreg.—
Danthonia californica Boland, Oreg.—Deschampsia atropurpurea (Wahl)
Scheele, Wash.—Digitaria sanguinalis (L.) Scop., Ohio—Echinochloa col-
onum (L.) Lk., Fla.—E. crusgalli (L.) Beauv., Minn., N. Dak., Oreg.-—
Eleusine indica (L.) Gaertn., Fla.—Elymus antarcticus Hook, f., N. Dak.
—E. canadensis L., Ida., Minn., Nebr., N. Dak., S. Dak., Wyo.—E. dahu-
ricus Turcz., N. Dak.—E. giganteus Vahl, N. Dak.—E. glaucus Buckl.,
Oreg., Wash.—E. interruptus Buckl., N. Dak.—E. junceus Fisch., N. Dak.,
S. Dak.—E. macounii Vasey, N. Dak.—E. sibiricus L., N. Dak.—Era-
grostis curvula (Schrad.) Nees, N. Dak.—E. tricodes (Nutt.) Nash, N.
Dak.—Festuca dertonensis (All.) Aschers and Graebn., Oreg.—F. elatior
Ir., Mont., Oreg.—F. elatior var. arundinacea (Schreb.) Wimm., Md.—
F. idahoensis Elmer, Oreg., Wash.—F. megalura Nutt., Oreg., Wash.—
F. myuros L., Oreg., Wash.—F. octoflora Walt., N. Dak., Oreg., S. Dak.,
Wash.—F. ovina L., Ida., Md., Mass., N. Dak., Ohio, R. I., Wash.—
F. rubra L., Alta., Md., N. Dak., Ohio, Oreg., R. I., Va., Wash.—i^. rubra
var. commutata Gaud., Conn., D. C., Ida., Ind., Ky., Md., Mass., Mich.,
Mo., N. Y., N. Dak., Ohio, Oreg., Penn., R. I., Va., Wash., Wise—
Hierochloa odorata (L.) Beauv., Wyo.—Holcus lanatus L., Oreg., Wash.—
Hordeum brevisubulatum (Trin.) Lk., N. Dak.—H. distichon. L., Minn.,
Nebr., N. Dak., Oreg., S. Dak., Wyo.—H. gussoneanum Pari., Oreg.—
H. jubatum L., Minn., N. Dak., Oreg., S. Dak.—H. murinum L., Oreg.—
H. nodosum L., Oreg.—H. pusillum Nutt., Mont., Nebr., S. Dak.—H. vul-
gare L., Minn., Mont., Nebr., N. Dak., Oreg., S. Dak., Wash., Wyo.—
Koeleria cristata (L.) Pers., Minn., N. Dak., Oreg.—Lolium multiflorum
Lam., N. Dak., Oreg., Wash.—L. perenne L., Oreg., Wash.—Muhlenbergia
racemosa (Michx.) B. S. P., Minn., S. Dak.—M. squarrosa (Trin.) Rydb.,
N. Dak.—Oryza sativa L., Ark., Calif., La., Tex.—Oryzopsis hymenoides
(Roem. and Schult.) Ricker, N. Dak., S. Dak., Wyo.—Panicum capillare
L., N. Dak.—P. miliaceum L., N. Dak., Oreg.—P. tennesseense Ashe,
Nebr.—P. virgatum L., S. Dak.—Phalaris arundinacea L., Minn., Oreg.,
S. Dak.—P. tuberosa: var., Oreg.—Phleum pratense L., Minn., Mont.,
N. Dak., Oreg., S. Dak., Wyo.—Poa ampla Merr., Mont., N. Dak., Wyo.—
P. annua L., Ida., Ont., Oreg., Wash.—P. bulbosa L., N. Dak., Oreg.,
Wash.—P. compressa L., Minn., N. Dak., Oreg., Wash.—P. palustris L.,
Minn., N. Dak.—P. pratensis L., Calif., Conn., D. C, Ida., 111., Iowa,
Ky., Md., Minn., Mo., Mont, Nebr., N. Dak., Ohio, Oreg., Penn., S. Dak.,
Va., Wash., Wyo.—P. secunda Presl, Wash.—P. trivialis L., D. C, Oreg.,
Va.—Redfieldia flexuosa (Thurb.) Vasey, Nebr., S. Dak.—Saccharum
officinarum L., La.—Schedonnardus paniculatus (Nutt.) Trel., S. Dak.—
Secale cereale L., Minn., Miss., N. Dak., S. Dak.—Setaria lutescens (Wei-
gel) F. T. Hubb., Minn., N. Dak.—;S. viridis (L.) Beauv., Minn., Nebr.,
N. Dak., S. Dak.—Sitanion hystrix (Nutt.) J. G. Sm., Wyo.—Sorghum
vidgare Pers., N. Dak., Okla., Tex.—S. vulgare var. sudanense (Piper)
/I
Hitchc, N. Dak.—Sporobolus cryptandrus (Tdrr.) A. Gray, Nebr., S.
Dak., Wyo.-—S. neglectus Nash, N. .Dak.—Stenotaphrum secundatum,
(Walt.) Kuntze, Tex.—Stipa comata Trin. andjllupr., Mont., N. Dak.,
S. Dak., Wyo.—S. sibirica (L.) Lam., N. Dak.-j-S. spdrtea Trin., Nebr.,
S. Dak.—S. viridula Trin., Mont., Nebr., N . qdk., S. Dak., Wyo.—Tri-
setum canescens BuckL, Wash.—Triticum aestivum ' IJ.,' Colo., Ida.,
Mont., Nebr., N. Dak., Oreg., S. Dak., Utah, W^sh., Wyo.—ST. dicoccum
Schrank, N. Dak., Oreg., Wash.—T. durum Desf., Minn., Neb~r., N. Dak.,
S. Dak., Wash.—T. monococcum L., Wash.—T. spelta L., Oreg.—T. timo-
pheevii Zhuk., N. Dak.—Zea mays L., Ariz., IlL, Iowa, Mass., Minn.,
Mont., N. Dak., Ohio, Okla., Oreg., S. Dak., Wash. .
D . P . Rogers (1943) placed the basidial stage of JR. solani in Pellicu-

laria filamentosa (Pat.) Rogers based on Hypochnus filamentosus Pat.
He gave Corticium vagum var. solani Burt ex Rolfs as a synonym, but
C. vagum Berk, and Curt, was recognized as a common mold on wood-
land debris and was assigned to Pellicul'aria vaga (Berk, and Curt.)
Rogers ex Linder. Because most of our materia] has not been connected
with P. filamentosa, it is easier to employ the well-known R. solani for
the purpose of this report. /
R. solani is prevalent in most soils,'particularly acid ones. It attacks
a large number of grass hosts during wet weather in the spring. The
damage usually is slight, mostly mild root necrosis or slight stunting ;-
or sometimes elliptical light-colored eye spots with brown borders occur
on the sheaths or culm bases (Fig. 7). Under favorable conditions, the
fungus may be rather destructive on cereals, causing deep-seated lesions
at the ground line, which under stress of wind cause straw-breaking at
this point. This condition was noted in one variety of wheat at Mandan,
North Dakota, in 1943 (Sprague, 1944,5). An .exceptionally virulent
race of R. solani causes considerable damage i^ acid soils in the coast
region in Oregon (Sprague, 1934,d; 1937,a; 1944,b). The fungus from
the coast was erroneously assigned to a different species by the writer
(1934,d). The Oregon material from the coast is definitely a distinct
race, both in its very strong pathological traits and the light, somewhat
fluffy type of growth' it makes in culture. The Oregon-strain or race
may be similar to Italian material; at least the Oregon material which
the writer sent to Italy is identical with the Italian material which
was later sent to him. Glynne and Ritchie ,(1943) report similar symp-
toms on wheat from England and Wales. They believe that Oort's
"sharp eye spot" (1936) is the same fungus. They mention that Blair,
working with some Canadian strains of R. solani, found some that
cause stem girdling. They state that sharp eye spot does not increase
with the frequency of wheat or barley in rotations.Tt is as common in
the first wheat crop as it is on old, arable l^nd- The writer has long
recognized that this fungus is difficult to coritrbl by crop rotation be-
cause of its ability to develop on so many hosts. In some places in
Oregon it is a common practice to plant cereals to clean up soil in
preparation for seeding iJ/iiôcionia-susceptible crops such as potatoes
BASIDIOMYCETES 133
in following years. This practice is liable to be unsuccessful. It is

interesting to note that much of the detailed study in recent years on
racial differences in R. solani (Houston, 1945) has not considered the
forms on Gramineae, the most widespread host family.
FIG. 7.—Rhizoctonia solani lesions on barley: A, showing culm with sheath

stripped away disclosing stromatic tissue on surface, X 3 ; B, eyespot lesions on
sheaths, X l - " "
Hynes described "purple patch" disease of cereals in Australia due

to Rhizoctonia solani (1933; 1937,fe) where the plants in small to ex-
tensive areas in fields are stunted, stiffened, and slightly purplish in
color. The roots are rotted. Dillon Weston and Garrett (1943) discuss
a similar condition in England and state that the culm is not attacked
as had been discussed by Glynne, Sprague, and also Oort as noted pre-
viously. Samuel and Garrett (1932) earlier reported Rhizoctonia on
cereals in Australia, and Dana (1919) mentions it from western Wash-
ington. Photographs of these isolations at Washington State College in-
/I
dicate that Dana may have been dealing with the same race as occurs
on cereals in western coastal Oregon.
Foex and Rosella (1937) described Sclerotiutn constantini Foex and
Rosella on wheat, which is the same as the stem-canker-causing strain
of R. solani (Sprague, 1944,6). Foex (1941) reported further on S. con-
stantini which Whetzel believed close to R. solani and also to Sclero-
tium rhizodes (see R.A.M. 25:336). ' " •
The writer has made a study of the races of B. solani on certain
field crops, mainly grasses (1947,c). He determined that there were at
least four races of R. solani on Gramineae. Race 1 was nonparasitic
on this group; race 2 was weakly parasitic on most of them. Race 3
was strongly parasitic on most of the grasses but was very weakly so
on alfalfa. This is the race from the coast region of Oregon which
caused cankering-off of cereals. Race 4 was relatively parasitic on
grasses, mainly as a seed and seedling rotter but it also attacked alfalfa
strongly. Race 4 was essentially a complex, some strains under cool
conditions segregating distinctly, depending on ability to attack proso,
sorghum, oats, and wheat.
Allison (1949) reported a serious lâf rot'of Alta fescue {Festuca
elatior var. arundinacea) on lawns at Plant Industry Station, Belts-
ville, Maryland. While the fungus {M. solani) resembled the race from
Oregon (race 3) it was able to attack legumes more readily than race 3
and is apparently distinct from it. This can be known as race 5.
Brown patch, the well-known disease of turf, is simply a case where
R. solani attacks a crowded host colony, the fungus spreading radially
from a central point, leaving dead brown areas of grass. Brown patch
is important in lawns in Minnesota, the coast regions of the Atlantic
and Pacific areas, and is important at times throughout much of the
United States and Canada. In Oregon, R.,solani is a common leaf rot
of a large number of species of grasses during the humid winter months.
It was a serious pest in spray-chamber inoculations at Corvallis during
the years the writer was studying Septoria^ leaf spots of grasses. It was
particularly destructive to leaves of Holcus lanatusÂgrostis alba, A.
tenuis, A. palustris, and barley. Control of brown patclTby—fungicides
is not always easy (cf. Rowell, 1948). —
In addition to the diseases of the cereals and grasses growing in
temperate climates there are various fungi mentioned in literature
which cause important diseases of subtropical, and tropical Gramineae
and which are similar if not identical to R: solani in part at least.
Hypochnus (Corticium) sasakii Shirai, for instance, is a common foot
rot parasite of rice in the Orient. It belongs with what we now call
R. filamentosa as based on observations made by Bhdo some years
ago (1930,i>). Palo, working with Philippinfi/material (1926) came to
the same .conclusions. Endo (1930,a) found that the Japanese and
Philippine strains of this fungus were most parasitic on the rice plant
at a temperature of 32° C. Ikeno (1933) found that H. sasakii caused
BASIDIOMYCETES 135
more .damage on rice subject to discontinuous wetting than those ex-
posed to continuous wetting.
Matsumoto, Yamamoto, and Hirane (1933) studied temperature
and humidity relations of 17 Formosan strains of H. sasakii together
with an Indian strain obtained from the banded sclerotial disease of
sugar cane. The optimum temperature for growth of all of the Formosan
strains lay between 28° C. and 31° C. The Indian strain reacted simi-
larly but the temperature requirement for a German isolate of R. solani
was different. High humidity favored the fungus. Matsumoto (1934)
retained Corticium sasakii and C. solani as separate species, not en-
tirely because of small morphological differences but because, in addi-
tion, the two "species" showed vegetative differences.
Padmanabhan (1946) reported that a leaf spot of sugar cane in
Bengal was caused by R. solani.
Rhizoctonia occurs on Bermuda grass, sugar cane and rice, among
otherg, in Louisiana (Ryker, 1939). On Bermuda grass infection de-
velops as rather definite, gradually spreading \ r e a s in the field, in
which every plant shows severe blight of the leaves and leaf sheaths,
which bear large, irregular, bleached spots with redoish-brown margins.
The roots are not injured and the stems are only sHghtly attacked.
Other grass hosts are parasitized. Infection spreads to sugar cane
plants. The injury is confined to the lower leaves and leaf sheaths of
older plants, but on young suckers which emerge from the base of the
stool all the aerial parts are affected. The diseased leaves bore large,
irregular, bleached areas with reddish-brown margins, which, where
whole leaves were affected, formed bands across the leaf. These bands
resemble symptoms reported by Bell from Queensland, Australia, for
"banded" sclerotial disease (R.A.M. 11:203) or banded sheath spot
which F. S. Earle stated was caused by R. grisea (R.A.M. 2:261). A
strain of i?., solani was noted on Bermuda grass growing along the edge
of a rice field. The fungus produced large, distinct, irregular, grayish-
white (later gray-black) sclerotia which were flattened on the bottom
and rounded on the top. This, according to Ryker (1939) is the same as
R. grisea Matz and probably H. sasakii. The large sclerotia distinguish
it from the common small-sclerotia strains of R. solani.
Baudys (1929) reported R. violacea on Agropyron repens but it was
not parasitic on corn, Poa annua, nor Festuca pratensis.
Whetzel (1922) reported a destructive shoot rot of bamboo in
Bermuda which he attributed to aspecies of Rhizoctonia. Viegas (1939)
thought that this might be the same as Tomentella bambusina Yiegas,
a fungus which occurs in Brazil. Tonientella is a member of the Thello-
phoraceae (R.A.M. 19:155). •
Range: United States, Australia, Barbados, Brazil, British Guiana, Canada,
Ceylon, China, France, Great Britain, India, Italy, Japan, Morocco,
Netherlands, Philippines, Sierra Leone, S. Africa.
References: Baldacci and Ciferri (1936); Bertus (1927); Blair (1942,
1943); Dahl (1933); Dana (1919); Dickinson (1930); Dillon-Weston
/
and Garrett (1943); Edwards and Newton (1937); Endo (1930,
1932,a,6); Exner and Chilton (1943); Foex :(1941); Foex and Rosella
(1937); Garrett (1944,a); Glynne and Ritchie (1943);; Hearn (1942);
Hemmi and Endo fl931, 1933); Hemmi and Yakogi (1928); Howard
. and Keil (1947); Hynes (1933; 1937,6); Ikata and Hitomi (1930);
Jaarsveld (1942); Matz (1921); Monteith (1926); Monteith and Dahl
(1928); Gort (1936); Palo (1926); Park and Bertus (1932,a; 1934,6,c);
Peltier (1916); Peyronel (1926,c); Piper and Coe (1919); Rogers (1943);
Rowell (1948); Ryker (1939); Samuel and Garrett (1932); Sanford
(1938); Shaw and Ajrekar (1915); Sprague (1934,d; 1937,a; 1939,c;
1944,6; 1946,e; 1947,6,c); Tilford (1925); Tyson (1936,a); Vorhees
(1934,6); Wei (1934,o,6); Weindling (1934,a,6).
Pellicularia rolfsii (Sacc.) E. West—Seedling Blight, BasaLSheath and

Stalk Rot
Corticium roljsii (Sacc.) Curzi
Sclerotium rolfsii (Sacc.) Curzi
Mycelium white, coarse, cells 150-25^0 X 2-9 /x, side branches smaller
(2 /i), producing small, nearly spherical sclerotia about 1 mm. diam.,
(0.5-1.5) tan colored on potato-dextrose agar; basidial hymenium very
coarsely areolate at first, consisting of clusters of^basidia arranged in
lines on a very tenuous subiculum; as basidia increase in number
hymenium becomes denser but never forms a continuous or fleshy layer;
when fully developed usually 30-40 /* thick, putty colored, becoming
gray; basidia obovoid, 7-9 X 4-5 /* wide, bearing 2 or 4 parallel or
divergent sterigmata, 2.5-4 /x or less often up to 6 /x long. Spores ellipti-
cal to obovate, rounded above, rounded or pointed a t t h e base, apicu-
late, 3.5-5 X 6-7 /A, hyaline and smooth. Cystidia or incrusted hyphae
not seen. /
I /
On: Arrhenatherum elatius (L.) Beauv., La.;—Avena sativa L., Ark.,,Miss.—•
Cynodon dactylon L., Fla.—Hordeum vulgare L,, Calif.—Oryza sativa L.,
La.—Saccharum officinarum L., Fla., Hawaii, La.—Sorghum vulgare Pars.,
Ga., La.—Triticum aestivum L., La.—Zea mays L., Alabama, Fla., Tex.,
Va. ' , '"-----^^
Tisdale reported a severe pre-emergence seed rot and seedling blight
of rice and wheat (1922). He also noted that a strain of the fungus from
soybean and tall oatgrass seemed to b6 less parasitic on rice. The
injury in rice tends to follow drill rows and the diseased seedlings have
a dark moldy appearance. The base of the stem and the roots of dis-
eased plants are dark colored and often appear frosty, due to the pres-
ence of wefts of mycelium of the causal organism. Brown sclerotia are
attached to the roots or the base of the culm of diseaised plants. After
irrigation water is let into the rice fields these plants which are still
alive usually recover.
Barrett (1934) reported finding the basidial stage of this fungus
in the United States. He tentatively referred the United States and
BASIDIOMYCETES 137
New Zealand isolates to Corticium rolfsii (Sacc.) Curzi. However West
assigned the fungus t o Pellicularia (1947). T h e description given for
this species is t a k e n from West's article. West mentioned t h a t both
Corticium centrifugum (Lev.) Bres. (Goto, 1930; Curzi, 1931) and C.
rolfsii (Sacc.) Curzi (Curzi, 1931; B a r r e t t , 1934; M u n d k u r , 1934;
Miltthorpe, 1941) h a v e been given as t h e basidial stage of Sclerotium
rolfsii Sacc. W e s t reported t h a t the spore measurements of the material
he studied, which w a s oil Ficus, were within t h e range of C. centrifugum
but such characteristics as color and thickness of t h e hymenium were
very different. Fifteen single-spore isolates included forms which were
typical of S. rolfsii while others were like JS. delphinii D . S. Welch
(West, 1947). All of these isolates were parasitic on Lupinus angusti-
folius L. While P. rolfsii has a wide host range it is n o t omnipresent on
the grass family in the South. I n 1931 Weber gave t h e known host range
as 189 species, 12 of these belonging t o the grass family.
Goidanich (1939) reported Sclerotium delphinii on sorghum from
I t a l y . S. delphinii, as indicated above, is sometimes considered to be
the same as S. rolfsii.
Range: United States, Argentina, Australia, Brazil, British Guihna, Ceylon,
China, Colombia, Dominican Republic, Formosa, India, Indocmna, Italy,
Madagascar, Malaya, New Zealand, Puerto Rico, Sierra Leone, TKîdad,
Uganda.
References: Baldacci and Ciferri (1936); Barrett (1934); Curzi (1931,
^ 1932); Endo (1932,a,b); Fischer'and Noll (1942); Godfrey (1918); Goto
(1930, 1933); Hemmi and Endo (1928, 1931); Higgins (1922, 1927);
Miltthorpe (1941); Mundkur (1934); Nakata (1926); Stakman, L.
(1923, R.A.M., 3:83); Tisdale (1922); TuUis (1940); Weber (1931);
West (1947).
Rhizoctonia oryzae Ryker—Root Rot, Sheath Spot

Mycelium superficial and submerged when grown in pure culture—
hyaline, white, granulate-^when young, gradually becoming clear when
old. The main mycelial strands are 6 to 10 /* wide, branched at an acute
angle with a slight constriction at the point of branching, and with a
septation a short distance from the point of constriction. The main
hyphae is also septate a short distance above the branch. Later short-
celled, much branching hyphae emerge at right angles from the main
branches and ramify through the substrate, sometimes forrniiig barrel-
shaped cells. Forms salmon-color sclerotial masses. Produces a sheath
spot.
On: Avena sativa L., N. Dak.—Bromus tectorum L., Ida.—Hordeum vul-
gare L., N. DRk.—Oryza'Sativa L., Ark., Calif., La., Tex.—Stenotaphrum
secundatum (Walt.) Kuntze, Fla.—Triticum aestivum L., N. Dak.
This species and B. zeae Vorhees may be pinkish variants of R.
solani derived from basidial segregates. The writer sometimes isolated
this pink form from roots of cereals and grasses during early and mid-
/I
summer in N o r t h D a k o t a while he once obtained it on roots of over-
wintered living cheat grass growing in sandy soil' near the foot of the
Lewiston grade in Idaho. T h e pinkish, later"~ orange-pink t o orange-
buff colonies have stroma and mycelium very similar, except for t h e
unusual color, to R. solani. T w o races were noted in N o r t h D a k o t a .
One isolated from w h e a t was associated with R. solani and Helmintho-
sporium sativum in a straw-breaker condition in w h e a t (1944,b) while
another with more active parasitic tendencies was found on common
oats.
References: Ryker (1938); Ryker and Gooch (1938); Sprague (1944,&;
1947,6).
Rhizoctonia solani Kuehn—See Pellicularia filamentosa (Pat.)

D. P. Rogers
Rhizoctonia zeae Yorhees—Root Rot

Sclerotia submerged and superficial when grown in pure culture,
0.5-1 mm. diam., mostly superficial on host, 0.1-0.5 mm. diam., white
when young, brown to dark brown when mature; homogeneous in
structure and color, hard at maturity, semigregarious or scattered,
often abundant, hyphae typically constricted at right-angle branches,
4-10 jj. wide; at first hyaline and granular, reddish brown in old cul-
tures, salmon pink on host, later dull gray, multiseptate, constricted |
at the septa.
On: Zea mays L., Fla.
It is likely that this species and R. oryzae will hp proved to be only
wild variants of R. solani arising from strains of Pellicularia filamen-
tosa. However, until this has been shown, it appears to be more con-
servative to recognize Vorhees' strain which! is distinguishable from'
R. solani, at least by the salmon pink color of the fungus in some
stages (Vorhees, 1934,6).. """""^-^-.^.^
R. zeae causes a sclerotial dry rot of corn ears. The ears become
shrunken and covered with pink, later gray, compacted rnycelium.
The general effect is like that caused by Diplodia zeae (Schw.) Lev.
The husks adhere tightly to the kernels. Sclerotia form on the outer
husks in some instances. / /
Seedling blight is readily induced by artificial inoculation with
R. zeae. The fungus is favored by a temperature of about 33° C. which
is the equivalent of a hot summer day (91° F.). This, according to
Vorhees, accounts for its ability to attack the earsjn direct sunlight.
The pink Rhizoctonia which the writer isolated,in -North Dakota
and Idaho may be the same as R. zeae to which he was inclined to
assign it (Sprague, 1944,6) but recently he placed these in R. oryzae
Ryker. '
BASIDIOMYCETES 139
A saprophytic fungus which produces small, hard, cinnamon or red-
dish stromata and cinnamon red thinly webby mycelium is distinct
from the forms just mentioned. Whetzel, in correspondence, indicated
that this was the same species as isolates obtained from forest soils
in the Appalachian Mountains and probably was not RMzoctonia at
all. This fungus is associated with roots of grasses in sandy soil at
Mandan and in other localities in North Dakota and adjacent states.
Pathogenicity tests with various isolates of this fungus on small grains
and some range grasses proved negative in the greenhouse at Mandan,
North Dakota. Bourne (1922), however, does mention a somewhat
similar fungus on sweet sorghum from Barbados. Pure cultures devel-
oped numerous reddish-brown sclerotia about 4 mm. in diam. He '
assigned this to RMzoctonia ferruginea Matz which had been de-
scribed from roots of sugar cane in Puerto Rico the preceding year
(Matz, 1921). Tucker (1927) found that isolates from sugar cane could
attack cabbage, carrots, beets, Swiss chard, cotton, sweet peas, and
other vegetables but strangely corn, sugar cane from cuttings, sweet
potatoes, and Cynodon were immune.
Range: S. E. United States.
Reference: Vorhees (1934,6).
Sclerotium Tode
" The genus Sclerotium belongs to the Mycelia Sterilia, consisting,
usually, of basidiomycetous white or sometimes tinted mycelia which
aggregate to form hard tubercles or balls of tissue called sclerotia.
These sclerotia are usually light colored on the inside, oxidized to dark
brown or black on the exterior. Some of these species have been shown
to have sexual stages such as Typhula, Claviceps, Sclerotinia, Lepto-
sphaeria, and others. Some species are' still listed in the form-genus
Sclerotium. Most of these species cause culm rots or leaf rots, fre-
quently at low temperatures. The genus however is no longer impor-
tant to the plant pathologist interested in diseases of Gramineae be-
cause most of the species are no longer referable to Mycelia Sterilia.
There are now only three species to be discussed in this text and only
S. rhizodes is at all well known. Even its importance is to some extent
based on confusion, in earlier years, with species we now know as
Typhula spp. The genus Sclerotium is discussed in the Basidiomycetes
because the remaining species appear to be logically referable to that
group.
Sclerotium hydrophyllum Sacc. apud Rothert—^Leaf Sheath Rot

Sclerotia (tubercles) globose, 0.3-0.7 mm. diam., numerous, dark,
with white interior, loose hyphae, generally floating or submerged, on
aquatic plants.
On: Echinochloa colonum (L.) Lk., Ark.—E. crusgalW (L.) Beauv., Ark.
This species is reported on Echindchloa spp.i from Arkansas in
Weiss's lists (1945) but the original source is not known to us. Rothert
reported this species, described by Saccardo (Rothjert, 1892) on water
plants from horticultural gardens in Gerniany and Kazan. Whether
barnyard grasses are "normal" hosts for this aquatic species is not
certain, and, in fact, the name is listed here with considerable hesita-
tion. The description is actually too brief to permit any critical com-
parison with other members of the genus. '
Weiss (1943) also lists Sclerotium sacidioides Speg. on Arundinaria
gigantea (Walt.) Chapm. from Virginia. No information is available
on the role of this species in the United States other than this report.
References: Rothert (1892); Weiss (1945).
Sclerotium oryzae Catt.-—See Leptosphaeria salvinii Catt.

/
Sclerotium rhizodes Auersw.—Leaf Blight or Rot
Infected leaves as they unfold remain more or less convolutely
rolled as in the bud, becoming white, dead, dry, and rigid; when nu-
merous, appearing as though frosted. Tissue covered with a thin, ob-
scure but dense felt of mycelium which is most marked on the inner
surface of the leaf, especially when only a part of the leaf is rolled^
laterally. Infected leaves are often caught and held by their tips in
the sheath, and sometimes the culms are stunted. Black sclerotia form
on the leaves, 1-5 mm. diam.
On:-Agropyron subsecundum (Lk.) Hitchc, Wise.—Agrostis alba L., Conn.,
Penn., Utah, Wye.—A. scabra Willd., Wise—5romMS ciliatus L., Wise.—
Calamagrostis can'adensis (Michx.) Beauv., Ida., Mass., N. H., N. J.,
Wash., Wise.—C. neglecta (Ehrh.) Gaertn., !Mey. and Schreb., Wise.—
Glyceria grandis S. Wats., Ida.—G. striata (Lam.) Hitehe., Wise.—
Hordeum jubatum L.,,Wise.—Phalaris arundinacea L^Wisc^—Phleum
pratense L., Wise.—Poa pratensis L., Penn., Wise.—Sphenopkolis pallens
(Spreng.) Seribn., Wise. ~
Whetzel indicated that this fungus was probably a species of Rhi-
zoctonia (cf. Remsberg, 1940,a, footnote, p. 178, and Foex, 1941).
Stout (1911) found that the fungus entered th4 roots from the soil
and could not infect the leaves of growing plants directly. He appar-
ently believed that the fungus was related to one of the mycorrhizal
forms such as discussed by Schlicht (1889). Sclerotium rhizodes was
confused with Typhula spp. (Hungerford, 1923 ;^Heald,/1924) and it
was not until the work of Remsberg had beeiCpublished (1940,o,b)
that the two groups were segregated. Therefore some of the early refer-
ences to S. rhizodes are not clear. However, Stout clearly described the
symptoms of S. rhizodes which causes a leaf tip or blade killing and
BASIDIOMYCETES 141
not a snow mold. He found the fungus in Wisconsin forming on the
plant in April, May, or, to some extent, later in the season. It prefers
cool, moist growing conditions but is by no means a snow mold that
requires near freezing conditions. In the Far West we have seen S.
rhizodes on Agrostis and Calamagrostis along mountain streams and
at the edge of thickets, sometimes in abundance as at Priest Lake,
Idaho, where it occurred in large areas in thick stands of Calamagrostis
canadensis. In Utah, Fischer, Meiners, and the writer found it in mid-
June at an elevation of about 5,000 feet along an irrigation ditch.
Kreitlow (1942) reported observations in Pennsylvania. Pape (1926)
suggested early cutting to reduce the amount of inoculum.
Baudys (1930) found that S. rhizodes was distinct from Typhula.
He found that the sclerotia of the former were globose, scarcely gla-
brous, rough and blackish. He was inclined to consider S. rhizodes as
belonging to Sclerotinia. He found that it showed affinity to S. menieri
Boud. and Sclerotium irregulars on rice but was distinct from S. oryzae-
sativaeîci. R.A.M. 10:191-192).
Goidanich (1939) reported a collar rot on sugar sorghum from Italy
caused by Sclerotium delphinii (but see Pellicularia roljsii).
Range:-United States, Austria, Belgium, Czechoslovakia, Denmark, Eng-
land, Germany, Norway, Poland, Sweden and probably widely distributed
in northern Europe and Asia.
References: Baudys (1930); Davis, W. H. (1933); Ekstrand (1938,6);
\ Flachs (1935); Foex (1941); Kreitlow (1942); Oettingen (1934); Pape
(1926); Sampson and Western (1941); Stirrup (1931); Stout (1911).
Typhula spp.
As mentioned above under Sclerotium the Typhula group has not
been well understood until recently. Most of the earlier references to
fungi which we now know as Typhula were once called Sclerotium. The
genus Typhula is characterized by small, hard, or firm sclerotia which
germinate later to produce a more or less club-shaped small sporophore
which may be rose colored, white, or darker, depending on the species.
Over thirty years ago Osmun and Krout (1918), to cite one in-
stance, described a sclerotium disease of lawn grasses, which from the
description could have been either Rhizoctonia or, perhaps, Typhula.
Much of this earlier literature is not helpful because of the confusion
necessarily prevailing in this complex and, at that time, poorly studied
group of turf diseases.
While Remsberg's studies (19^0,a,b) in the United States have
clarified the group considerably, they are still difficult to study be-
cause of their exacting requirements of temperature and light. Investi-
gation on their control has been under way for many years in Idaho
and for a number of years in the state of Washington, in cooperation
with the United States Department of Agriculture, in the latter in-
stance.
l\
Typhula idahoensis Remsb.—Speckled Snow Mold or Scald
Sclerotia light amber when young, chestnut brown when mature but
appearing nearly black when dry, commonly ontthe leaves and parts
above ground in the winter rosette stage of the host, sclerotia erumpent
to superficial, not coalesced, globose to somewhat flattened^ 0.5-0.9 X
1-2 diam., cortex reddish brown, 5-20 /x thick; sporophores clavate,
erect, straight to somewhat curved, simple to rarely ramose, glabrous
with thick apex, rounded to acute, fawn color to wood brown, clavula
elongate fusiform cylindrical, 5-10 mm. long, 0.5-1.5 mm. broad, vina-
ceous brown to leathery brown; basidia elongate, thicker at apex, 4-6-8
spored, 27-31.5 X 5.8-7.8 /x,; basidiospore ovate to ellipsoid, 8-14 X 3.8-
8 (h, average 10.5 X 4.5 /i. \
On: Agropyron cristatum (L.) Gaertn., Ida., Wash.—A. inerme (Soribn. and
Sm.) Rydb., Wash.—A. intermedium (Host.) Beauv., Wash.—A. smithii
Rydb., Wash.—Bromiis carinatus Hook, and Arn., Ida., Wash.—B. inermis
Leyss, Wash.-—B. tectorum L., Wash.—'Deschmnpsia elongata (Hook.)
Munro, Ida.—Hordeum nodosum L., Wâsh.—Stipa columbiana var. nel-
sonii (Scribn.) Hitchc, Wash.—S. comata Trin. and Rupr., Wash.^
Triticum aestivum L., Ida., Mont., Utah, Wash.
This funguS' is very common on cereals and some grasses' in the
Pacific Northwest. It grows best in culture at temperatures between
9-12° C. but the temperature range is 0° C. to 18° C. In the field it
does the most damage at a temperature scarcely above freezing during
the time when the snow is melting or before. Mycelial growth in cul-
ture is abundant, fluffy, and concentrically zoned according to Rems-
berg but, compared with other fungi which we have worked with, its
growth seems slow and its culture tedious.i Sclerotia, which Remsberg
states (1940,a) appear in 5-10 days, are! clustered or in concentric
rings, always single, never coalesced, and are chestnut brown when
mature. The mycelium under the snow is ofl-white or light gray. Rems-
berg (1940,a) determined that the surfaces of the sclerotia were com-
posed of irregular thiii walls while those of T. itoana wefe-composed
of irregular thick walls. —
According to Remsberg (1940,&) T. borealis Ekstrand is the same
as T. idahoensis. Ekstrand (1937) failed to'describe the sporophores of
T. borealis and Remsberg's name is therefore valid.
T. graminum Karst., according to Remsberg (1940,a) has not been
demonstrated as causing snow mold. It has white sporophores instead
of the rose-colored ones of T. itoana. ,
T. idahoensis is the most common Typhula on wintfer wheat in the
Dyer Hill area, Douglas County, Washingtenj in other parts of this
general region T. itoana is common. Characteristically all but the
innermost leaves of winter rosettes are diseased in heavy infestations.
The fields as viewed irom a short distance have a dead, straw-colored
BASIDIOMYCETES 143
appearance after the snow has left. The leaves become dry, bleached,
and peppered with the nearly black, numerous, flattened sclerotia.
During the moist year 1948 fields which were killed 100 per cent to
the ground recovered and yielded a good crop. There were spots in
the field which were thin. During drier years recovery is less certain.
Many growers become panicky and reseed to spring grain. This often
results in a mixture of spring and fall wheat with the spring wheat
suffering from competition with the fall grain. In general it appears
to be wiser to leave the fields to recover, especially since modern
weedicides permit destruction of weeds which tend to smother thin
stands of snow mold-injured fall-seeded wheat.
In Douglas County, Washington, observations by G. S. Holton in-
dicate that in most years, at least, early seeding favors recovery of
the injured grain. It develops a better root system when seeded in
August. This practice, of course, favors common root rot but in Douglas
County this is often the lesser of the two evils. In the Dyer Hill area
of Douglas County, most of the growers seed early with excellent
recovery, while in the LaMoine area in the same county they tend to
seed later and recovery has been less reliable according to C. S. Hel-
ton's observations (1948).
This fungus attacks range grasses in the vicinity of infested wheat
fields and occurs on miscellaneous grass hosts in the hills of Idaho.
The writer found the fawn-colored sporophores developed in vast
quantities on November 14, 1948 in the vicinity of Mansfield, Wash-
ington, following cold rains and light snows (Sprague, 1949,e) and on
the same date in 1949. In 1949 sporophores were found from Novem-
ber 14 to the time of the last visit on December 9.
Range: United States, Finland, Sweden, U.S.S.R.
References: Ekstrand (1937; 1938,a; 1946); Heald (1924); Holton and
Sprague (1949); Hungerford (1923); Miiller (1930); Remsberg
(1940,a,6); Remsberg and Hungerford (1933); Sprague (1942,6; 1949e).
Typhula itoana Imai—Speckled Snow Mold

Sclerotia tawny to hazel brown, in general reddish (or the color of
radish seeds when growing), often in the sheaths, crown, and near root
parts and less often on the leaves as in T. idahoensis, frequently falling
away from the substrate, single or coalesced, smooth at first, rough
when dry or old, sometimes spherical to subglobose but- on the host
often irregular or flattened or concave below and convex above, 0.5-2 X
1.5-4 mm. diam., rind golden to reddish brown, about 7.8 fj. thick, tough,
composed of a smooth gelatinous layer on the outer walls of enlarged
peripheral cells; medulla -paraplectenchymatous with hyphae loosely
interwoven, small sclerotia solid, large ones coalesced masses, often
hollow; sporophores clavate, erect, straight, or slightly curved, apex
broad, round, or slightly pointed, tapering into stipe, simple or
branched, usually one arising from each sclerotium, often slightly
I
pubescent at the base, 8-25 mm. tall, clavula rose color, opaque, stipe
white, translucent, becoming flesh color to bittersweet pink when dry,
clavula straight, erect, smooth, cylindrical, or j somewhat broadened
and flattened, tapering toward a round or slightly pointed apex, 10-15
mm. tall, 1-2 mm. broad, hydrangea' pink to palejvinaceous pink, com-
posed of a hollow cylinder of agglutinated hyphae bearing the hy-
menium which is continuous over the apex, crystals present in the
hymenium and subhymenium; stipe distinct in size and color from the
clavula, erect, straight, pubescent at the base with hairs clasping the
sclerotium, 5-10 mm. long, 0.5-1.0 mm. in diam., white, solid, hyphae
often slightly incrusted with flne crystals; basidia broadly clavate,
27.2-34.6 X 5.1-7.8 /A wide; basidiospores ovoid, flattened on one side
above a pointed apiculus, 4.3-7.8 X 11.3-14.8 fi, average 11.7 X 6-05 /x.
On: Agropyron spicatwn (Pursh) Scribn. and Sm., Wash.—Agrostis palustris
Huds., Ida., Mass., N. Y.—A. tenuis Sibth., Penn.—Bromus tectorum L,,
Wash.—Dactylis glomerata L., Conn., Ida.—Elymus glaucus Buckl., Ida.
—Hordeum vulgare L., Ida.—Poa pratehsis L., Ida., Minn., Wash.—
Secale cereale L., Wash.—Stipa columbian'a var. nelsonii (Scribn.) Hitchc,
Wash.—Triticum aestivum L., Ida., Wash.
According to Remsberg (1940,a) this species has the same optimum
temperature of growth as T. idahoensis, 9° C.-12° C, but Ekstrand
says that the fungus flourishes at a temperature slightly above the
range 0° C.-5° C , under or at the edge of melting snow (1938,a).
Remsberg's report is based on pure culture studies. The field observa-
tions of Holton and the writer have shown that T. itoana appears ,to
develop somewhat later in the season than T. idahoensis^. In" 1948 the
former seemed to be somewhat active in May, and even in late June
the sclerotia could be found in the crown of winter wheat. In the Dyer
Hill area the black sclerotia of T. idahoenns were almost omnipresent
in March and early April but later, after this material dried down, the
red sclerotia of T. itoana moved into the Icrowns. In addition it was
noted in Lincoln County that the April reports of snow mold were all
due to T. itoana, while those reported eaHier from^Douglas County
were due to T. idahoensis. Possibly some of this differBnceJs due to
the fact that the red sclerotial fungus develops mainly in'_the moister
habitat of the crown, while the black sclerotial fungus [T. idahoensis)
is soon inhibited by drying conditions on -the dead exposed rosettes.
While the season of 1948 was unusually wet and cool observations at
least indicate that the etiology of the two forn{s is somewhat-'different.
It was also noted that T. itoana could be isolated with less difficulty
than T. idahoensis and appeared to develop at slightly higher tempera-
tures. Study on the life history of these forms is nee'ded'in the state of
Washington with emphasis on the developmendrnder/field conditions.
Holton (1948) has made extensive field observations on the group with
emphasis towards practical field control. The writer found mature
sporophores of this fungus in great numbers on November 14, 1948.
BASIDIOMYCETES 145
Remsberg (1940,a) reported that the mycelial growth was abun-
dant, white, webby radiating, concentrically banded, and fan-shaped.
The sclerotia which appeared in 5-10 days were pinkish orange when
young, tawny to hazel-brown when mature, single or coalesced, and
tended to develop in concentric rings. Sterile white sporophores fre-
quently developed in culture from sclerotia. Kozhevnikova (1946) also
discussed symptoms (cf. R.A.M. 27:226-227).
Remsberg (1940,a) found that T. itoana was more abundant in
Idaho than T. idahoensis. In 1946 and 1947 T. idahoensis was much
more prevalent in Douglas County, Washington, than T. itoana. In the
wet year 1948, T. idahoensis was even more prevalent in the Dyer HilL
area of Douglas County, but T. itoana was very common in the
LaMoine-Waterville area of this county and near Deep Creek and
Davenport east of these areas. In 1949 sporophores grading from one
species to the other were noted.
Remsberg (1940,a,6) found that Sclerotium fulvum Fries was ap-
parently the same as T. itoana but T. graminum Karst. was, as men-
tioned, probably distinct.
Wernham and Chilton (1943) studied T. itoana and an undeter-
mined T.yphula on orchard grass and some other turf grasses in Con-
necticut. Search for resistant grasses did not disclose any promising
leads.
Zimmerman (1922) reported that top dressings with nitrate of soda
and sulphate of ammonia, together with thorough hoeing and harrow-
ing, greatly improved the condition of the crop of winter barley at-
tacked by Typhula in Germany. Wiirtenberger (1927), in discussing
Typhula graminum, which produced small (0.5-2 mm. diam.) brown
sclerotia, reported that top dressings of sodium nitrate, leuna saltpeter,
and ammonium sulphate, accompanied by hoeing and harrowing, were
officially recommended for the control of the disease.
Range: United States, Belgium, Denmark, England, Finland, Germany,
Japan, Norway, Poland, Sweden, U.S.S.R.
References: Ekstrand (1937, 1938,a); Helton and Sprague (1949); Hun-
gerford (1923); Imai (1929, 1936); Kozhevnikova (1946); Miiller (1930);
Remsberg (1940,0)6); Sprague (1949,e); Tasugi (1929, 1930, 1935);
Tervet (1941); Vang (1945); Volk (1937); Wernham (1941); Wern-
ham and Chilton (1943); Young (1937); Young and Morris (1929).
Nonsporulating Basidiomycetes—Snow Molds and Root

Necrosis Diseases
Broadfoot (1936) and later Broadfoot and Cormack (1941) re-
ported a snow mold-like gray fungus on 22 species of overwintered
grasses including crested wheatgrass, red fescue, winter wheat, and
a nongrass host, alfalfa. Smooth brome proved highly resistant in field
tests at Edmonton, Alberta. This fungus is very parasitic at very low
temperatures. They state, "This pathogen grows readily on most nu-
trient media at temperatures from 0° to 18° G., producing a fluffy,
white, sterile mycelium. Cardinal temperaturesi for growth are —4° C ,
15° C , and 26° C. Sclerotia, fruiting bodies, and spores have not been
found on the diseased host plants, and many attempts to produce them
in culture under varying conditions of temperature, light, moisture, and
nutrient media failed. The characteristic clarhp connections of the
mycelium and the occasional production of small rudimentary sporo-
phores on culture media indicate that the fungus is a basidiomycete."
Vanterpool (in Canad. 26th Ann. Rpt. PL Dis. Survey, p. 31) 'noted
that lawns treated with mercurials suffered little or no damage from
the low-temperature basidiomycetes. Cormack '(1947) reported the
fungus was widespread and severe on timothy, redtop and creeping
red fescue, less severe on Kentucky bluegrass and meadow fescue.
Another nonfruiting basidiomycete was isolated from crested
wheatgrass in Minnesota by Andrews (1944), from Agropyronsibiri-
cum (Willd.) Beauv., A. elongatum (Host.) Beauv., Elymus canadensis
L., and crested wheatgrass at Mandan, North Dakota (Sprague,
1944,5), and on slender wheatgrass [A., trachycaulum) near Medora,
North Dakota. It has also been isolated from crested wheatgrass col-
lected near AVibaux, Montana, and south of there in the Custer National
Forest, Montana. This fungus is a weak parasite on grassesj causing a
slow root necrosis and yellowing of the seedlings. It develops during
the summer and fall and is therefore distinct from the low-temperature
fungus of Broadfoot and Cormack. The summer species produces a
white cottony growth on potato-dextrose agar. It may, perhaps, be
related to mycorrhizal forms.
There is a very common white mold on the base of culms of Cala-
niagrostis rubescens Buckl. in the fir-spruce timber of Chelan National
Forest north of Winthrop, Washington. The white basidiomycetous
mold is prevalent on the sheaths, culms, ahd adjacent forest duff. The
fungus probably develops as a snow mold but appears to be somewhat
parasitic on pine grass (C. rubescens). It also occurs on other grasses
growing in association with colonies of pine grass.^Hher hosts seen
included Agrostis alba L., A. exarata Trin., Calamagrostis- canadensis
(Michx.) Beauv., and Poa pratensis L. These observations are based
entirely on extra-official notes and virtually nothing is known about
the etiology or taxonomy of this mold condition. A comparable fungus
was noted in 1948 on Dactylis glomerata L. in Logan Canyon, Utah
(Sprague and Meiners, 1949). | '
References: Andrews (1944); Broadfoot (1936); Broadfoot and Cormack
(1941); Cormack (1947); Sprague (1944,6).
FUNGI IMPERFECTI
FUNGI I M P E R F E C T I |
Because there are so many species of Fungi Imperfecti on Gra-
mineae and because some of them have such extensive host ranges, the
entire group represents a very complex problem,; considered from the
viewpoint of the plant pathologist as well as the mycologist. It will be
obvious to the reader that considerable fundamental work needs to be
done with this heterogeneous group. However, in the lifetime of the
writer, he has noted that the attitude toward this group has greatly
changed. The plant pathologist has needed information on^Fungi Im-
perfecti because so many plant disease problems involve the often
abundant conidial phases, " ,'
In order to bring related genera together the following standard
grouping is followed: '
A. Spores borne in pycnidia ,. . . ^ . Sphaeropsidales
AA. Spores borne in acervuli Melanconiales
AAA. Spores borne otherwise Moniliales
SPHAEROPSIDALES
Most of the pycnidial forms t h a t occur on all b u t the coarser grasses
and canes are simple, unilocular, ostiolate forms which generally have
globose pycnidia. W e do encounter more complex forms on the sub-
tropical and tropical bamboos and canes where nutrients and growing
conditions favor extensive fungus growth.
A key t o the Sphaeropsidales involved in this m a n u a l is as follows:
A. Parasitic on or associated intimately with fruiting bodies of other fungi
B., Parasitic on rust pustules Darluca
BB. In fruiting bodies of Phyllachora Davisiella
AA. On leaves or other plant parts, sometimes but only incidentally asso-
ciated with other fungi
B. Pycnidia multilocular or in a stroma
C. Pycnidia multilocular Cyt'os'pora
CC. Pycnidium in a stroma Melasmia
BB. Pycnidia separate
^ C. Pycnidium typically elongate with elongate ostiole
D. Pycnospores 1-celled, globose to oblong, hyaline . . . . Leptothyrium
DD. Pycnospores 2-celled, hyaline Scaphidium
DDD. Pycnospores filiform Leptostromella
CC. Pycnidia typically globose to ellipsoidal
D. Pycnidial wall somewhat dothidiaceous Macrophomina
DD. Pycnidial wall parenchymatous
E. Spore's hyaline or light yellow
F. Spores nonseptate
G. Pycnidia with stiff hairs in ostiolar region . . . Pyrenochaeta
GG. Pycnidia smooth
H. Spores strongly curved, falfcate to bluntly so
Selenophoma
HH.. Spores more or less straight
I. Spores filiform Septoria *
II. Spores broadened
J. Spores typically less than 15 M long
K. Pycnidia somewhat carbonaceous, often tardily os-
tiolate (probably not distinguishable from Phyllos-
ticta) ,-. Phoma
KK. Pycnidia parenchymatous, brown, relatively thin-
walled, ostiole formed early Phyllosticta
J J . Spores coarser, 20 /i or more in diameter
Macrophotna
FF. Spores septate
* In part (also Phleospora but with spores coarsely obclavate).
149
150 DISEASES OF CEREALS AND GEASSE^ |
G. Spores filiform •.I Septoria *
GG. Spores broadened ~ ^ '~ ' \
H. Spores appendaged . |. . .Dilophbspora
HH. Spores not appendaged
,1. Spores 1-septate
J. Cross wall medianly located ^ .1 Ascochyta
J J . Gross wall not medianly located . . . . ' . . . Apiocarpella
II. Spores 2- to multiseptate
J. Pycnidia well formed about ostiole . . . . .Stagonospora
J J. Pycnidia breaking open at apex, leaving ragged ostio-
lar area, spores typically obclavate ....... Phleospora
III. Spores muriform Hyalothyridium
EE. Spores colored
F. Spores nonseptate, globose Coniothyrium
FF. Spores septate
G. Spores 1-septate Diplodia
(See also Microdiplodia)
GG. Spores multiseptate "" /
H. Spores cylindrical, broadened . , Hendersonia
HH. Spores elongated, filiform, or obclavate . Phaeoseptoria
Apiocarpella agropyri Sprague in Greene—Leaf Spot

Spots round, elliptical, or elongate, coalescing into irregular lesions,
4-20 X 3-5 IX, d r a b brown, center buff brown or t a w n y , narrow margin
of darker brown, sometimes surrounded by wider yellow areas, pyc-
nidia globose, erumpent, ostiolate, golden brown, 70-130 /x, not promi-
nent, spores exuding in very short tendrils, narrowly boat-shaped or
fusiform, bases blunt, tapering to a pointed apex, septate about % from
blunted base, slightly constricted a t septum, 17-22.5 X ^ - 8 - 4 . 2 /*, sub-
hyaline. I
O n : Agropyron repens (L.) Beauv., Wise. '
H . C. Greene sent this material to t h e writer for description
(Greene, 1950). T h e spores are narrower t h a n those of A: rnacros'pora
(Speg.) Sydow and larger t h a n those of A. minor. T h e fungus isaTvery
active parasite. T h e lesions resemble those of Phleospora graminearum.
I t is surprising t h a t a species as evidently parasitic as this one should
escape notice in Wisconsin until this date. T h e explanation lies in the
obscurity of the grass hosts which have been ignored until recent years.
Range: Wisconsin. '
Reference: Greene (1950). ''
Apiocarpella minor Sprague in Greene—Leaf-Spot /

Spots elliptical or linear, gray or pale yellowish, ^dinetimes with
centers sordid whitish, margins purplish; pycnidia elliptical or subglo-
bose, erumpent, brown, parenchyma-like, ostiolate, 90-165 X 80-125 /x;
*In part (also' Phleospora but with spores coarsely-'obolavate).
FUNGI IMPERFECTI—SPHAEROPSIDALES 151
pycnospores ellipsoid-ovate with acute base, eccentrically septate (the
upper locule larger, the lower smaller, appendiculiform), hyaline, 13-
17 X 3.7-5 jot.
On: Calamagrostis canadensis (Michx.) Beauv., Wise.
H. C. Greene collected this fungus on living leaves of the host at
Madison, Wisconsin, July 22, 1947. A. minor has much smaller spores
than those of A. macrospora (Speg.) Sydow which has been found
growing saprophytically on Stiporysopsis bloomeri (Boland) Johnson
in California (Sprague, 1948,c). Recently A. macrospora was found in
fawn colored spots on Poa ampla in Colorado (Sprague, 1950,a).
Range: Wisconsin.
References: Greene (1948); Sprague (1948,c).
Ascochyta spp.
There has been comparatively little published on the Ascochyta
spp. on Gramineae with the exception of Davis' earlier report (1919,a)
and some preliminary notes by the writer during recent years (Sprague,
1946,e; 1948,c). A lengthy article on the group by A. G. Johnson and
the writer is now in press (1950) and summarized information is taken
from that article for this text. Ascochyta, as indicated earlier (1946,e),
includes hyaline and yellow spores {Ascochytula and Ascochyiella),
and^those assigned to Diplodina (see also Diedicke, 1912,6) which is
not distinct from Ascochyta.
The characters principally used in segregating the fungi studied in
this report are included here.
KEY TO ASCOCHYTA
Section I. Eu-Ascochyta: Spores hyaline

A. Spores chiefly less than 4.0 M in diameter
B. Spores rather narrowly fusiform-ellipsoid
C. Spores chiefly 2.5 IJ. or more in diameter
A. sorghi
CC. Spores chiefly less than 2,5 M in diameter
A. phleina
BB. Spores narrowly cylindrical with blunt ends
A. desmazieri _ _
AA. Spores chiefly more than 4.0 M in diameter
B. Spores chiefly less than 5.0 /t in diameter, rather broadly cylindrical
with tapering ends
A. brachypodii
BB. Spores oblong ellipsoid, up to 5.0 M or more in diameter with blunt ends
C. Spores usually up to 6 /^ in diameter
A. graminea
CC. Spores up to about 8 M in diameter
A. sorghina
Section II. Ascochytella: Spores slightly colored or tinted by
colored inclusions ^ j
A. Spores narrowly cylindrical, with blunt ends, chiefly 3 M or less in diameter
A. subalpinus
AA. Spores more than 4 /u. in diameter 'i
B. Spores broadly cylindrical with more or less blunt ends scarcely at all
constricted at the septum
C. Spores tapering rather abruptly to blunt ends '
D. Spores chiefly less than 6 /* in diameter
A. agropyrina
DD. Spores chiefly more than 6 M in diameter
A. utahensis
CC. Spores tapering graduaUy to rather blunt ends
D. Spores broadly fusiform to ellipsoid, seldom 2-septate
A. hordei
DD. Spores irregularly cylindrical to ellipsoid, some 2-septate
" A-. avenae
BB. Spores broadly eUipsoid with rather sharply pointed ends, shghtly
constricted at septum /
' A. missouriensis
Preston (1945) listed Ascochyta sppT (undetermined) on the follow-
ing hosts in Oklahoma: Aegilops cylindrica, Bouteloua curtipendiila,
B. gracilis, B. rigidiseta (Ste'ud.) Hitchc, Bromus commutatus, B. iner-
mis, B. purgans, and Calamovilja gigantea.
Ascochyta agropyrina (Fairman) Trotter—Leaf Rot

Ascochytula agropyrina Fairman
Pycnidia sparse in tawny, later brown, ^spotôr in stramineous
areas, globose, dark brown to black, 130-320 /x diam.; spores oblong,
cylindrical, straight, ends rounded, uniseptate, constricted slightly, at
first hyaline, then pale yellow, 17-23 X 4.5-6.2 /* but as wide as 7 ju in
winter material at Pullman, Washington. '
On: Agropyron bakeri E. Nels., N. Mex.—A. caninum (L.) Beauy.,^S[.'Dak.
—A. cristatum (L.) Gaertn., Mont., N. Dak., Wash.—A. michnoi Roshev,
N. Dak.—A. semicostatum (Steud.) Nees, N. Dak., Wash.—A. sibiricum
(Willd.) Beauv., N. Dak.—A. smithii Rydb., Mx)nt.—A. spicatum (Pursh)
Scribn. and Sm., Mont., Wash.—A. subsecundum (Lk.) Hitchc, N. Dak.,
Wash.—A. trachycaulum (Lk.) Malte, Colo., N. Dak., Okla., Sask., -Utah—
A. trichophorum (Lk.) Richt., N. Dak.—Elymus canadensis L., N. Dak.—
E. virginicus L., N . Dak.
This is a yellow-spored species which is a common hut relatively
unimportant component of the leaf spot complfix~on grasses in the
plains country. The spores and pycnidia of the material at Mandan,
North Dakota, and westward (Sprague and Johnson, 1950) are smaller
than the type but otherwise are similar to Fairman's collection which
was made on Agropyron bakeri. The spores>re typically stoutly cylin-
drical (Fig. S,A,B) and not related to the A. sorghi complex. Scanty
winter material on A. spicatuni collected by Jack Meiners in January,
1948, had lightly tinted spores, 17-23 X 4.8-7.0 fi..
The symptoms expressed by this species are variable. Often the pyc-
nidia occur in straw-colored, bleached areas. Developing spots, or ones
not weathered, are small, brown, later tawny colored. One collection
on A. subsecundum from Pullman, Washington, had fuscous spots with
red to vinaceous borders. The spores from this collection were particu-
larly pale yellow but had the same shape as those of A. agropyrina.
Somewhat similar material on other hosts from the Far West was
assigned to A. hordei Hara (Sprague, 1948,c).
References: Fairman (1918); Sprague (1946,e); Sprague and A. G. John-
son (1950); Trotter (1931).
Ascochyta avenae (Petrak) Sprague and A. G. Johnson-^Leaf Spot

Ascochytella avenae Petrak
Lesions somewhat elongated, gray brown, later straw colored or
vague^^pycnidia prominent, dark brown, somewhat flattened, ostiolate,
100-200 ju, diam., wall 5-10 /x thick; pycnospores irregularly cylindrical
to fusoid with rounded ends and coarse granular contents, 1- (some-
times 2-) septate, finally yellow, 17-26 X 6.2-7.0 ja.
On: Avena byzantina K. Koch, Oreg.—A. jatua L., Wash.
We collected this fungus on red oats in the early spring from over-
wintered plants near Troutdale, Oregon. The spores (Fig. 8,C) turned
yellow after a year in the herbarium. A leaf spot on A. jatua from
Pullman, Washington, appears to be summer material of the same
species. The latter material has an associated Mycosphaerella-like
fungus present, which is too undeveloped to determine.
The writer examined the type of this species on A. sativa, but in
the portion available there was nothing present except Puccinia coro-
nata Corda. The diagnosis is based therefore on the technical descrip-
tion by Petrak (1925).
Range: United States, Czechoslovakia.
References: Petrak (1925, pp. 107-108); Sprague (1946,e; 1948,c); Sprague
and A. G.Johnson (1950).
Ascochyta brachypodii (Sydow) Sprague and A. G. Johnson

comb. nov.—Leaf Spot
A. graminicola var. brachypodii Trail
Diplodina brachypodii Sydow
In vague or white spots, with or without margins; pycnidia erum-
pent, ostiolate, brown, 90-200 /*; spores hyaline, 1-septate, typically
fusoid or bent, seldom straight-cylindrical, 15-20 X 4-5.5(6.0) fj..
O n : Andropogon furcatus Muhl., N. Dak.—A. scoparius Michx., Ariz.—Avena
sativa L., N. Dak.—Chloris virgata Swartz, N. Dak.—Elymus macounii
Vasey (hybrid), N. Dak.—Festuca rubra !>., N. |Dak., Oreg.—Lolium
perenne L., Oreg.—Melica virgata, N. Dak.—Phdlards arundinacea L., N.
Dak.—Pleuropogon refractus (A. Gray) Benth., Oreg.—Poa cusickii Vasey,
Oreg.—Sorghastrum nutans (L.) Nash, N. Dak.—Siipa comata Trin. and
Rupr., Mont.—S. viridula Trin., Minn. j
The coarse fusoid spores (Fig. 8,I,J,K,L) of this species should be
recognized as distinct from A. sorghi. W e have noted spindle-shaped
spores on Andropogon and Sorghastrum (Fig. 8,J,K) which develop
in pure culture more like A. brachypodii.
FIG. 8.—Pycnospores of: A, Ascochyta agropyrina on Elymus virginicus,

Mandan, N. Dak.; B, A. agropyrina on Agropyron smithii, Forsythe, Mont.;
C, A. avenae on Aveiia byzantina, Troutdale, Oreg.; D, A. sorghi on Elymus
canadensis, near Laurel, Mont.; E, A. graminea orf Bouteloua gracilis, Pingree,
N. Dak.; F, A. graminea on Cynodon dactylon, Davis J c , Calif; G, A. sorghi
on Elymus condensatus, Pullman, Wash.; H, A. sorghji on Elymus canadensis,
Kingdom City, Mo.; I, A. brachypodii on Beckmannia syzigachne, Lisbon,
N. Dak.; J, A. brachypodii on Andropogon jurcatus, Mandan, N. Dak.; K,
A. brachypodii on Sorghastrum nutans, Mandan, N. Dak., in pure culture; L,
A. brachypodii on Stipa comata, Hebgen Lake, Mont.; M, A. sorghi on Bromus
inermis, Lexington, Ky.; N, A. sorghi on Hierochloa odorata, Mandan, N. Dak.;
O, A. sorghi on Lolium perenne, Troutdale, Oreg. JJ^ /
Range: United States, Germany, Scotland.

References:'Sprague (1946,e; 1950,a); Sprague and A. G. Johnson (1950);
Sydow (1916); Trail .(1887,a, p. 88).
Ascochyta desmazieri Cav.—Leaf Blotch

Septoria graminum var. lolii Mont.
S. lolii West.
Diplodina lolii Zimm.
Spots chocolate brown to purplish brown, later fading to straw color
in the center, pycnidia more or less grouped, compressed globose, im-
mersed, brown, 130-200 /x diam., ostiolate, spores narrowly cylindrical
(bacillar), 1-septate, 13-20 X 2-3.5 fi.
On: Lolium multiflorum Lam., Calif., Oreg., Wash.—L. perenne L., Calif.,
Oreg., Wash. -^
This common leaf and glume spot on ryegrasses in western Oregon
and Washington was the subject of a detailed taxonomic study of its
involved history. It was pointed out that it might be an Ascochyta
stage of Septoria loligena Sprague but no additional information has
been obtained on this point. It is distinguished from A. sorghi by its
truly cylindrical spores. If the type of Phoma lolii Pass, is in condition
to study it should be compared with A. desmazieri also.
A. desmazieri is prevalent enough in western Oregon to be of some
economic importance, and in the Puyallup Valley in AVashington is one
of the leaf spots which is being avoided in selection of ryegrass strains
for production.
Range: United States, Belgium, Denmark, France, Moravia.
References: Cavara (1893); Frandsen (1943); Sprague (1944,d); Sprague
and A. G. Johnson (1950); Westendorp (1857); Zimmermann (1914).
Ascochyta graminea (Sacc.) Sprague and A. G. Johnson

comb. nov.—Leaf Mold
Diplodina graminea Sacc.
Ascochyta cynodontis Unam. ,
A. boutelouae Fairman
Pycnidia on dead leaves in vague spots, minute or larger, often
elongate, black, composed of large, polygonal cells, 55-90 X 90-160 /x;
spores capsular or short cylindric, blocky, 1-septate, clear hyaline but
with a hint of smokiness in mass, 11-20 X 3.9-6.0 fi, sometimes as much
as 7 ju wide.
On: Bouteloua gracilis H. B. K., N. Max., N. Dak.—Cynodon dactylon (L.)
Pers., Calif., S. Car.
In spite of the unfortunately similar name this species is not to be
confused with A. graminicola which we now place under A. sorghi.
Probably A. graminea is a saprophyte, or at best, only a very weak
parasite. We can see no very good reason for retaining the species,
A. boutelouae. The spores in the type are slightly broader than most
of the material on Bouteloua (Fig. 8,S) in North Dakota and on the
collection of Cynodon (Fig. 8,F) from California, but otherwise the
group represents a morphological unit. The species is characterized by
its short, blocky, capsular spores.
Range: United States, Italy, Spain.
References: Diehl (1937); Fairman (1918); Sprague (1946,e); Sprague
and A. G. Johnson (1950); Unamuno (1929). I
Ascochyta hordei Hara—Leaf Spot

Spots few to several per plant, 3-11 mm. diam., round, oval, or
irregular, brown at first, sometimes with several rings, fading to whit-
ish yellow, clay colored, or vaguely pale, nearly white, later with
brown margins or surrounded by yellow outer areas, leaves finally
withering and dying; pycnidia spherical or subspherical, thin-swalled,
ostiolate, epiphyllous, erumpent, 100-172 fj.; pycnospores chlorinous or
very pale yellow to virtually hyaline, cylindrical with rounded ends or
less often slightly twisted or semifusoid but scarcely fusoid and not
tapering strongly toward either end (except on Holcus and cultivated
barley in Washington), 1- (seldom 2-) septate, 16-22 X 4.8-6.2 jx on
Festuca and Hordeum murinum, 17.8-21 X 5.3-7.1 m j on Bromus, 18-27
X 5.5-6.8 /x on Holcus and cultivated barley, all in the western United
States and 17-28 X 4-6 ii on cultivated barley in Japan (description
modified from Hara).
On: Bromus carinatus Hook, and Arn., Utah, Wash.—Festuca elatior L.,;
Utah—Holms lanatus L., Wash.—Hordeum murinum L., Ida.—H. vvl-
gare L., Wash.—Phleum phleoides (L.) Karst., Wash.—Siipa rohusta
Soribn., Colo. ,
Tsuruda (1915) mentioned a ring-spot disease of wheat and barley
in Japan. Hara described the fungus in 1916, apparently, and our de-
scription is taken, in part, from the text by Ideta (1926) and by Hara
(1930) and translated at Beltsville, Maryland, by Thomas K. Take-
shita and at Pullman, Washington, by J. Y. Park. The' Japanese fungus
which Ideta called "round spot" is said to have spores which are oval,
egg- or cocoon-shaped, colorless or yellowish. The description"which we
gave our western material is close to that of the Japanese material
except that the ring-spotting is lacking or very obscure. Another
fungus, A. tritici Hori and Enjoji (Journ. Plant Path, and Ent.,Japan,
August, 1930), has narrower spores, 14-21 X 3-6.5 fx. borne in spots free
of rings. Perhaps our material (Fig. 9,B,C,D; Fig.* 10,B,C) differs from
A. hordei, in which event it would either be assigned to A. avenue, A.
tritici, or described as new. A. avenae (Fig. 8,C) has somewhat coarser
spores than A. hordei. - -^Z - •
Before chancing on the Japanese discussion of A. hordei (Ideta,
1926) we. had described with A. G. Johnson in the herbarium, another
A. hordei which wfe have now placed under A. hordei Hara. The spores
FIG. 9.—Pycnospores of: A, Ascochyta utahensis, type on Agropyron inerme,

Logan Canyon, Utah (C.S. 3610); B, A. hordei on Festuca elatior, Logan Canyon,
Utah (C.S. 3656); C, A. hordei on Hordeum murinum, Riggins, Idaho (C.S.
3677); J), A. hordei on Bromus carinatus, Logan, Utah (C.S. 3852). (From
Mycologia.) (
in this Washington material on Holcus and barley are- fusiform-cylin-

^.drical (Fig. 10,JB,C) .
One western collection obtained on dead overwintered material of
Phleum phleoides at Pullman, Washington, had brown spores, 16-21 X
5-7 jj.. In order to keep this group from getting further complicated we
Af f\
^
]\]
-• 4 =s
\J
H • -J
FIG. 10.—Pycnospores of: A, Ascochyta phleina on Phleum pratense, type,

Niawa, Minn.; B, A. hordei on Hordeum vulgare, High Prairie, Wash.; C,
A. hordei on Holcus lanatus, Bingen, Wash.; D, Ascochyta phase of Stagono-
spora simplicior on Stipa columbiana var. nelsonii, Mt. Shasta, Calif.; E, A.
missouriensis on Panicum virgatum, Wilton, N. Dak.; F, A. subalpinus on Ory-
zopsis asperijolius, (Upper) Spearfish Creek, S. Dak.; G, A. subalpinus on
Deschampsia elongata, Mt. Rainier, Wash.; H, A. subalpinus on Distichlis stricta,
Marion, Mont.
have assigned this to A. hordei, considering it to'be winter spores which
approach Diplodia in morphology. Material on Stipa robusta Scribn.
from Cedaredge, Colorado, is similar. ~^ I
Range: United States, Japan. 1
References: Ideta (1926, see pp. 683-84 and' translation of same by Take-
shita which is filed at Dept. of Plant Pathology, Wjash. State College and
in Div. of Cereal Crops and Diseases, U. S. D. A., Beltsville, Md.); Hara,
S. (1916); Hara (1930); Sprague (1948,c); Sprague and A..G. Johnson
(1950); Tsuruda (1915).
Ascochyta maydis Stout—Leaf Spot

Pycnidia localized in a small patch on a large, effuse, translucent,
dead area of the leaf, developed subepidermally, opening £ither epi-
phyllously or hypophyllously by a minutely papillate ostiole, dark
brown, membranaceous, their walls composed of a pseudoparenchyma,
lenticular, 75-150 yu, diam.; ostiole rounde^d, 7.5-15 p. diam., the ostiolar
papillum composed of smaller cells and appearing darker than the wall
of the pycnidial body. Spores 2-celled, hyaline, long-ellipsoid to fusoid,
rarely lightly constricted at the septuni, 11-18 X 3-4.5 /* (also material
10-16 X 5 ;u).
On: Zea mays L\, 111.
This species has some of the appearance of A. brachypodii. Except
for the original study very little information is available on A. maydis.
A key to the Ascochyta species on corn is given under A. zeae.
Range: United States. /
Reference: Stout (1930). /
Ascochyta missouriensis Sprague and A'. G. Johnson—Leaf Rot

Spots none to gray, emarginate; pycnidia^ few, black, erumpe'nt, sub-
papillate, 80-120 fx; pycnospores lemon-shaped, apices^cute^ 1-septate,
hyaline (with yellow inclusions), 15-19 X 6-8 jn. - ^
On: Panicum dichotomiflorum Michx., N. Dak.—P. virgatum L., N. Dak.
This fungus is probably saprophytic. It has been collected in small
quantities several times in west-central North (Dakota in late season.
It is typified by the distinct lemon-shaped spores (Fig. 1Q,E) with
hyaline inclusions. I t may be the developing phase of a more complex
fungus. '
/
Range: United States. ^./-'~ '
References: Sprague (1946,e); Sprague and A. ,G-. Johhsoh (1950).
Ascochyta oryzae Catt.—On Discolored Culms and Leaves

Pycnidia on leaves and glumes, erumpent, parenchymatous, black,
ostiolate, spores linear, oblong, ends rounded, 1-septate, pale yellow,
biguttulate, 15 X 4 ft.
On: Oryza sativa L., Va,
This has been reported on discolored glumes of rice from Virginia.
Available information on this fungus is scarce.
Ling (1947) listed A. leptosj)ora (Trail) Hara on rice from Chengtu,
Szechwan Province, China. This species must have been created from
A. graminicola var. leptospora Trail (1887,a). This form has spores
12-15 X 2.5-3 fj. on Agrapyron repens in Scotland. We have never seen
material of var. leptospora but the measurements are about the same
as for A. elymi, now considered to be a phase of A. sorghi (Sprague and
A. G. Johnson, 1950).
Range: United States, Italy, Japan.
References: Cattaneo (1877); Weiss (1945, v. 29, p. 328).
Ascochyta phleina Sprague—Leaf Spot

Spots becoming tawny, pycnidia golden brown to brown, globose,
strongly erumpent, more or less grouped,' ostiolate, 100-135 /*, pycno-
spores hyaline, cylindrical, both ends rounded with a tendency to
narrow towards the apices from the center of the spore, contains few
small oil drops, nonconstricted at the one septum, 11-16 X 1.5-2.2 /x,
mean size 13 X 1.75 ix, ratio of length to width 7.4-1.
On: Phleum pratense L., Minn.
This fungus is probably moderately parasitic. The relatively narrow
spores (Fig. 10,A) distinguish this species from the A. sorghi group.
Reference: Sprague (1948,a).
Ascochyta sorghi Sacc.—Leaf Spot

• A. graminicola Sacc.
A. graminicola Sacc. sensu Davis
A. graminicola var. leptospora Trail
A. graminicola var. hold Sacc.
A. graminicola var. diedickeana Baudys and Picb.
A. elymi Teh on and Daniels
Lesions circular to elliptical, brown or fawn colored, usually with
red borders or thin brown discolored ones, mainly on the lower, dew-
covered leaves or on the tips of the foliage, often scanty; pycnidia sub-
globose, erumpent, ostiolate, 90-140 ^ diam. for the most part, golden
/ i
brown with d a r k e r oxidized cells adjacent to theiostiole; walls varying
from fuliginous t o paler; pycnospores variable in size, subcylindric to
somewhat ovate, fusoid, 11-21 X 1.6-4.0 /*. '
O n : Agropyron desertorum (Fisch.) Schult., N. Dak.-pA. repens (L.) Beauv.,
Mass.—A. smithii Rydb., Colo.—A. spicatum (Pprsh) Scribn. and Sm.,
Mont.—A. trachycaulum (Lk.) Malte, Ida.—Agrostis alba L., Ida., N. Y.
—A. canina L., Mass.—Arrhenatherum elatius (L.) Beauy., Oreg.—Avena
sativa L., Ga.—Beckmannia syzigachne (Steud.) Fern., N. Dak.—Bromm
carinatus Hook, and Arn., Oreg., Utah, Wash.—B. commutatus Schrad.,
Wise.—B. inermis Leyss, Ky., Minn., N. Dak.—B. japonicus Thunb., Wise.
—B. tectorum L., Minn.—Calamagrostis canadensis (Miehx.) Beauv.,
Wise.—Dactylis glomerata L., Del.—Deschampsia elongata (Hook.)
Munro, Oreg.—Elymus canadensis L., Mo., N. Dak., Okla., Wise.—E.
condensatus Presl, Ida., Mont., Wash.—E. glaucus Buckl.,'Ida.—E. ma-
counii Vasey (hybrid), Mont., N. Dak.—E. -virginicus .L., 111., Tex.—
Festuca ovina L., Wise.—F. rubra L_., N. Dak., Oreg.—Hierochlpa odorata
(L.) Beauv., N. Dak., Sask., S. Dak., Wisc-r—Hordeum jubatum L., Iowa—
H. murinum L., Ida.—H. nodosum t.,, Alaska—H. vulgare L., 111.—
Hystrix patula Moeneh, Wise.—Melica bulbosa Geyer, Wyo.—Muhlen-
bergia joliosa (Roem. and Sehult.) Trin., Wise.—Panicum capillare L.,
Wise.—Poa bulbosa L., Wash.—P. grdcillima Vasey, Oreg.—P. pratejisis
L., Colo., Iowa, N. Car., N. Dak., Ohio, Tex., Wash.; Wyo.—P. secunda
Presl, N. Dak., Wash.—Schizachne purpurascens (Torr.) Swallen, Wise.—
Sclerochloa dura (L.) Beauv., Ida.—Secale cereale L., III.—Setaria lute-
scens (Weigel) F . T. Hubb, Mo.—Sitanion hystrix (Nutt.) J. G. Sm.,
Oreg.—S. jubatum J. G. Sm., Wash.—Sorghum vulgare var. sudanense
(Piper) Hitchc., S. Dak.—Sphenopholis intermedia Rydb., Okla.—Stipa
occidentalis Thurb., Oreg.—S. viridula Trin., S. Dak.—Trisetum canescens
Buckl., Ida.—Triticum aestivum L., Okla. (fide Preston).
A. graminicola is the species t h a t most Ascochyta-like fungi on
Gramineae have been compared with and often indiscriminately a s -
signed to. Actually a majority of such material proved to be Darluca
fiXum (Biv.) Cast. T h e writer and A. G. Johnson (1950) assigned
the A. graminicola material (Fig. 8,D,G,H,M,N,0) to A. sorghi, an
earlier name. T h e modern concept of A. graminicola -was^based on
D a v i s ' description (1919,a) which amounts t o a n emendation t o in-
clude var. hold in t h e description of the species. This variety has some-
what larger spores t h a n the vague t y p e of A. graminicola. T h e emended
description, however, seems logical and is the basis of the writer's con-
cept of the group assigned to A. sorghi. I
While there are some 14 varieties of A. graminicola known, some of
these are. obvious duplications, several are Darluca filum', but several
could possibly be retained because of recognizable differences from t h e
species proper. While A. sorghi m a y some day_prove to be only an
artificial species created out of miscellaneou^'^Qrm-s'pecies it has not
yet been so proved. I n fact, the material included in D a v i s ' concept of
the species appears to represent a definite fungus with distinct ability
to a t t a c k certain grasses. '
The Canadian Plant Disease Survey contains reference to several
collections of A. graminicqla. Some collected by Gordon are appended
by spore measurements. A collection on Bromiis inermis Leyss had
pycnidia about 100 n diam., containing spores which were somewhat
irregular, 8.7-15 X 3-4 /x. Another collection on Agropyron trachy-
caulum from Ft. Garry, Manitoba, had much larger spores, 17.5-20 X 5
/x, and is probably distinct. Another Manitoba collection on red fescue
had smaller, narrower spores, 12.5-14 X 2.5-3.0 /i. The 1943 Canadian
report (Johnston as collaborator, cf. Conners) gives A. graminicola on
wheat from Alberta with spores 18.1 X 5.8 /j..
Hara, S. (1916) reported A. graminicola from Japan. Pycnidia were
given as 100-150 /j., dark brown, spores 1-septate, oval or spindle-shaped
with tapered ends, 10-14 X 3-4 ju.
A. elymi (Fig. 8,D) is assigned to A. sorghi because it is apparently
immature material of A. sorghi. Tehon and Daniels state (1927), "The
distinction between this species and A. graminicola Sacc. lies in its
pycnidia, the walls of which are very thin and scarcely distinguishable,
in contrast with the distinctly parenchymatic, fuliginous ones of Sac-
cardo's species." The writer has seen comparable material on seedlings
of Elypius canadensis along the Cannonball near Solen, North Dakota
(1946,e), and possibly the same on Agropyron repens near Eugene,
Oregon (1944,d). We have also seen fragmentary material from time
to time on various grasses, especially old leaves of Elymus condensatus •
'in the West. Material on A. smithii from Buckhorn Creek, Colorado,
had spores 12-18 X 2.0-3.5 /*. Most of these collections were repre-
sented by scattered pycnidia associated with other fungi.
A. graminicola var. diedickeana produces a dark brown or nearly
black general scalding of affected green parts in some cases. The
pycnidia are thin walled, 120-150 /*, and the spores subcylindrical to
slightly fusoid, 15-20 X 2.5-4.0 /x. It is not recognized because morpho-
logically it is within the A. sorghi group.
Range: General in North America, Europe, and Asia, at least in the cool
temperature parts; Czechoslovakia, Germany, Greenland, India, Japan,
Poland, Scotland, Spitzenberg, Turkey, U.S.S.R., Uruguay.
References: Davis (1919,a); Frandsen (1943); Greene (1942, pp. 92-93);
Saceardo (1875); Sprague (1946,e); Sprague and A. G. Johnson (1950);
Tehon and Daniels (1927).
Ascochyta sorghina Sacc.—Rough Spot

Spots elongate, fuscous with red borders, pycnidia densely gre-
garious, globose depressed, papillate; spores oblong-ellipsoid, 1-septate,
minutely pluriguttulate, hyaline, 20 X 8 j«-
On: Sorghum halepense (L.) Pers., Ga., La., N. Car.—S. vulgare Pers., Ala-
bama, Fla., Ga., Miss., N. Car., Okla., S. Car.^S. vulgare var. sudanense
(Piper) Hitchc, Ga.
The pycnidia are so large that they give the surface of the leaf a
1
rough feel to the touch, hence the name "rough spot" (Leukel, et al.,
1944) (Fig. 11).
A. sorghina has much coarser spores than A. sorghi. The fungus was
originally described from Italy. Diehl (1937) found sorghum infected
Fia. 11.—Enlarged portion of sorghum leaf showing Ascochyta sorghina, Ex-

periment, Ga. (Photo furnished by C. L. Lefebvre, U. S. Dept. of Agr.)
in Alabama. The leaves bore large necrotic areas up to 12 X 2 cm. con-

taining numerous pycnidia. The report by Diehl was the first citation
from North America. He noted that the fungus was apparently more
parasitic in this country than reports from Europe would indicate might
be expected.
Wallace (1934) reported that A. sorghina was a common parasite
of young sorghum in Tanganyika. It produced long, gray, dead areas
on the leaves. Older plants were resistant.
Chaves Batista (1946) recently described A. sacchari on Brazilian
sugar cane. The leaf spots were elliptical, chlorotic, purple bordered,
12-15 X 3-5 mm. diam. It produced amphigenous, glabrous, circular,
dark chestnut colored, immersed, later erumpent, ostiolate, pycnidia,
96 /t diam. with 1-septate, elliptical or cylindrical, hyaline pycnospores,
8-10 X 2-3 II.
It should be noted that A. sorghina and A. zeina Sacc. have virtually
the same descriptions (1878). A. zeina is listed as having lenticular
pycnidia and A. sorghina globose-depressed pycnidia. If these differ-
ences are not of specific rank, the fact that A. zeina appears two pages
earlier in the text would reduce A. sorghina to synonymy. This point
needs critical study.
Range: United States, Italy, Tanganyika.
References: Diehl (1937); Leukel, et al. (1944); Saccardo (1878); Wal-
lace (1934); Weimer, Burton, and Higgins (1937).
Ascochyta phase of Stagonospora simplicior

See St. simplicior (Fig. 10,D)
Ascochyta subalpinus Sprague and A. G. Johnson—Leaf Rot

This is a saprophytic, yellow-spore species having very narrow
spores (11-19 X 2-2.4 ^) (Fig, 10,F,G,H) which, in the extreme, could
• be placed in Septoria. I t has been collected on Distichlis stricta on Lost
Prairie, Marion, Montana, on Deschampsia elongata (Hook.) Munro,
Mt. Rainier, Washington, and on Oryzopsis asperifolia Michx. on
(upper) Spearfish Creek, South Dakota. The collection on Oryzopsis
represents a somewhat distinct form of the species, as the black pyc-
nidia are subsuperficial and the spores have characteristic guttulae at
either end (Fig. 10,F).
Reference: Sprague and A. G. Johnson (1950).
Ascochyta utahensis Sprague—^Brown Spot

Spots chocolate brown, 1-6 X 1-3 mm., mostly elongate or forming
irregular areas by fusion of several spots, sometimes fading to straw
color in centers; pycnidia few, scattered, obscure, pseudoparenchyma-
tous, brown, globose, ostiolate 110-148 /J.; pycnospores elongate-ellipti-
cal or cylindrical but tapering slightly to blunt ends, straight or more
or less bent or slightly lumpy in outline, contents hyaline to slightly
yellow, guttulate to granulose, 1-septate with a median septation, not
or slightly constricted at the septum, large, 22-29.2 X 6.6-10 fi.
On: Agropy'ron inerme Rydb. and Sm., Utah—A. trachycaidum (Lk.) Malta,
Colo., Wyo.
This species is definitely parasitic. Both the type material from
Logan Canyon, Utah, and material from Togwatee Pass, Wyoming,
/ !
show distinct lesions on living culms. Morph'ologically the fungus is
distinguished by its large spores which exceed in isize those of any other
oh Gramineae (Fig. 9,^1). The fungus is distinct from A. agropyrina
not only in morphology but in pathogenicity. i
References: Sprague (1948,c; 1949,a). .I
I
Ascochyta zeae Stout—Leaf Spot
Spots at first ellipsoid, becoming elongated and somewhat irregular,
sometimes confluent, the leaf veins tending to bound the lesions later-
ally, their margins brown, and well defined or faded, their interiors
becoming tan-cinereous. Pycnidia moderately abundant, developed sub-
epidermally, sometimes in rows between the microscopic leaf veins,
opening either epiphyllously or hypophyllously, often through the
stomata, by a minutely papillate ostiole, dark brown, membranaceous,
their walls composed of an indistinct pseudoparenchyma, lenticular,
55-160 /J. diam.; ostiole rounded, 6.5-18'/i across, the ostiolar papillum
appearing darker than the wall of the pycnidial body. Spores obscurely
uniseptate, the septum often apparently lacking, hyaline, oblong-
ellipsoid to somewhat irregular, rarely constricted at the septum,
8.5-13.5 X 3-4.5 fx.
On: Zea mays L., 111.
Stout (1930) proposed a key to the species of Ascochyta on Zea,
presented here. ;
KEY TO SPECIES OF ASCOCHYTA ON ZEA
On stalks: spores 6-8 X 1.5-2.0 ii

A. zeicola Ell. and Ev.
On leaves
Spores 8.5-13.5 X 3-4.5 /i
A. zeae Stout
Spores 11-18 X 3-4.5 fi
A. maydis Stout
Spores 18 X 7.5 fi
A. zeina Sacc. _
A. zeae is similar to A. sorghi but the writer has not seen the former
and cannot be certain that they are identical. A. zeicola is apparently
a saprophyte while it is not clear that A. zeina is known,to North
America. Lobik (1933) reported it from the North Caucasus. As men-
tioned above under a discussion of A. sorghina, A. zeina'may be the
same as A. sorghina. A. zeina apparently is distinct from A. maydis
because the largest spores listed for A. maydis just readh the minimum
length for A. zeina, while the width of the. spores of the latter is vir-
tually twice that of A. maydis.
Reference: Stout (1930).
Coniothyrium psammae Oud.-—Leaf Spot

Spots large, elliptical, light buff with prominent vinaceous borders;
pycnidia few, relatively prominent, spores brown, elliptical, 10-13 X 3-4
/*, borne by budding from the walls of the unilocular pycnidia on pycno-
phores varying from mere papillae 6 /* long to ones 10-12 ja long.
On: Calamagrostis nutkaensis (Presl) Steud., Oreg.
This unimportant spot occurs sparingly along the Pacific sea coast.
It is weakly parasitic and is favored by the high humidity, saline air,
and rigorous growing conditions endured by the host along the seashore.
Range: United States, Netherlands.
References: Sprague (1941,6; 1946,e).
Coniothyrium zeae Stout—On Corn

Spots at first ellipsoid, becoming elongated and irregular, some-
times confluent, leaf veins tending to bound the spots laterally, at first
tan colored, their margins brown and well marked to somewhat faded,
their interiors becoming lighter.
Pycnidia located in the mesophyll, opening by a minutely papillate
ostiole, brown, membranaceous, composed of a pseudoparenchyma,
globose, 130-150(230) jn diam., ostiole rounded, 12-30 ju, across, the
ostiolar papillum appearing darker than the pycnidial wall. Spores
1-celled, brown-olivaceous, their walls dark and well marked, long-
ellipsoid, often presenting one nearly flattened and one curved side,
8.5-13.5 X 4.4-6.6 /J^.
Stout (1930) placed this species in Coniothyrium instead of
Sphaeropsis because the spores were small and the pycnidia lacked
conspicuous conidiophores. He said that it differed from Sphaeropsis
ambigua Mont., also on corn, by its smaller spores, those of S. ambigua
measuring 15 X 5 j«. Sphaeropsis sp. has been reported as a rot of corn
in California. The only Sphaeropsis collection which we have made was
fragmentary leaf material on Poa pratensis L. from the South Roose-
velt Park, North Dakota.
The species of Coniothyrium on corn and the one on Calamagrostis
may be closely related although the spores of the species on corn are
broader.
Cytospora sacchari Butl.-^Stem Canker

Stromata wart-shaped, usually in lines, subcutaneous, erumpent,
pluriloculate, black, opening is elongate, spores small, cylindrical.
/I
curved, ends obtuse, hyaline, 3.5 X 1.0-1.5 /*, sporophores ramose, sep-
tate, 12-18 /x long. (Fig. 12.)
O n : Saccharum officinarum L., La.
Abbott (1931) reported that this species was fairly abundant in
Louisiana but it was said to be of little or no economic importance.
It was collected in the United States first in the autumn of 1930 on
Co. 281. He reported that certain interspecific hybrids were susceptible
(1939).
Nowell (1923) reported that the leaf sheaths of young plants af-
fected with this fungus were firmly bound together by
/^^ the mycelium. The exposed parts take on a dull, very
0 ^^§ dark red color, and these parts are profusely covered by
0 § GB the pycnidia. Their comparatively long necks project be-
^ i? 0 yond the surface sufficiently to be readily seen and to
FIG. 12.— give a rough sensation when the finger is passed ,over
Pyenospoi-es them. During wet weather a minute yellowish globule of
lacchari conidia exudes from the tip of the beak. The disease
ceased to be of any importance in Puerto Rico after
susceptible strains were eliminated.
Luthra (1936) found the fungus on the Coimbatore varieties 223,
312, 313, 385, 392 and 394 in the Punjab. The symptoms were con-
spicuous on the rind when the canes were dried or had been buried in
the soil. Luthra, Sattar, and Singh (1938) proved that the fungus was
parasitic. '
Kiryu (1937) reported that the temperature range for this fungus
on nutrient media was 7° C. to 37° C. with the optimum at 31° C.
Range: United States, Formosa, India, Puerto Ricy^ S. Africa.
References: Abbott (1931, 1939, see also R. lA. M. 9:808); Butler (1906,
Mem. Bot. Ser. 1, p. 31, T. VIII); Kiryu (1937); Luthra (1936); Luthra,
Sattar, and Singh (1938); Nowell (1923); Saccardo (Sylloge, 1913, v.
22,962). I
Darluca filum (Biv. ex Fr.) Cast.—-Parasitic on Rust

Pycnidia in rust pustules, often giving the pustule a black appear-
ance, gregarious, globose to mammiform, strongly erumpent, golden
brown, finally black, darker about the nipple-like, ostiolar region, tex-
ture somewhat thin but not fragile although i somewhat brittle, par-
enchymatous; spores suboblong-fusoid, straight, finally 1-septate,
broadest in the middle and tapering strongly toward each- end; on each
end are short mucous appendages which are readily dislpdged and diffi-
cult to see, often appearing as hyaline, fragmented ends as though the
spore wall at the tips of the spore was missing, 14-18 X 3-4 /*.
On: Many species of rust, especially on Puccinia rubigo-vera (DC.) Wint.,
P. coronata Corda. and P. poae-sudeticae (West.) Jorstd. '
This fungus is very common in open winter or late fall in western
Oregon and Washington as well as being common in widely distributed
areas in the Middle West and eastern United States. It often checks
rust attacks on lawn grasses in the Willamette Valley, Oregon.
This species germinated with great speed in water. Since its career
is fleeting, at best, this ability to germinate quickly is no doubt im-
portant in obtaining a foothold during moist weather. Fedorinchik
(1939) found that this fungus germinated in 3 hours at 15° C.-25° C.
under high humidity with the optimum temperature at 20° C.-21° C.
The incubation period varied from 4 to 7 days within the temperature
limits of 9° C.-30° C. He mentioned that the fungus had been grown
in pure culture up to the pycnidial stage. Chester (1946) reported that
the fungus was an obligate parasite. Keener (1934) found some indi-
cation that physiologic races of Darluca exist but wide cross inocula-
tions have been made (Chester, 1946; Hardison, 1942; Traylor, 1940).
- Ascochyta graminicola var. y Sacc. and var. coerulea Br. and H.,
A. contubernalis Oud., A. -pucciniophila Starb. and {!)Rendersonia
uredinicola Desm. are all considered synonyms of D. filum (Grove,
1935). Bivona-Bernardi (1813) described the fungus many years ago.
Fries^ (1823) called it Phoma filum and Castagne (1851) proposed
Darluca.
Range: United States, Austria, Belgium, China, Czechoslovakia, Denmark,
Finland, France, Germany,. Great Britain, Italy, Japan, Netherlands,
V Norway, Poland, Sweden, U.S.S.R., widely distributed.
References: Bivona-Bernardi (1813); Castagne (1851); Chester (1946);
Fedorinchik (1939); Fries (1823); Grove (1935); Hardison (1942);
Keener (1933, 1934); Traylor (1940).
Davisiella elymina (Davis) Petrak—On Tar Spots

Cytodiplospora elymina Davis
Pycnidia in loculi of Phyllachora spp., spherical to elliptical, 100-
135 z^; spores oblong, hyaline, often 4-guttulate, becoming uniseptate,
7-10 X 2.5-3 iJb, larger in some collections.
On: Andropogori- jurcatus Muhl., Wise.—Bouteloua curtipendula (Miohx.)
Torr., Wise.—Elymus canadensis L., Wise.—E. villosus Muhl., Wise.—
E. virginicus L., Wise.—Hystrix patula Moench, Wise.—Muhlenbergia
racemosa (Michx.) B. S. P., Wise.—Oryzopsis asperijolia Miehx., Wise.
This is found growing with Phyllachora spp. but there is little data
on whether it is a parasite ot Phyllachora, an associate or a conidial
stage. In describing Cytodiplospora elymina in 1919 Davis gave the
spores as 7-10 X 2.5-3> (Davis, 1919,c). He considered the fungus to
be a spermbgonial stage of the Phyllachora on Elymus virginicus L.
Petrak, after examining the Wisconsin material and creating the new
genus Davisiella, gave the spore measurements as 8-13 X 3-4.5 [i
(1924, p. 134). Davis (1926) reported a form on Calamagrostis with
spores 10-20 /x in length, which is similar, perhaps, to our western
material of Septogloeum oxysporum Sacc., Bomm. £ind Rouss.. (Sprague,
1941,a). Davis (1937) listed a specimen on Muhlenbergia racemosa
with spores 10-16 X 2-3 /u, and one on Oryzopsislmth spores mostly
17 y. long but not yet septate. A collection on Andropogon Jurcatus had
3-septate spores. Greene (1942) relisted this group of collections and
mentioned that 3-septate material on Hystrix had spores 15-17 X 4 /n.
It would appear that the size of the spores had increased with the
additional hosts reported or else more than one species is iriVolved.
Greene (1944,a) found Davisiella in Phyllacho'â boutelouae Rehm on
overwintered leaves of Bouteloua curtipendula. The conidia are ob-
scurely. 3-septate, almost cylindrical, tapering slightly at each end,
mostly about 17-22 X 3 /i.
References: Davis (1919,c; 1926; 1937); Greene (1942; lQ4i,a,d; 1949,c);
Petrak (1924). ,
I
Dilophospora alopecuri (Fr.) Fr.—Twist Disease

D. graminis Desm.
Causing twisted and malformed flowering shoots, or the leaves are
partially enveloped in a black, ill-formed stroma bearing black, globose
pycnidia, 120-300 p., spores 1- 3-septate, hyaline, narrowly fusiform to
cylindrical with branched or bristle-like appendages at each end,
spores 8-15 X 1.6-2.5 ii, appendages 5-7 p., less often 10 /t long and
0.5 p thick at the base adjacent to the body of the spore, 3-4 branched
with some of the branches trifurcate.
On: Agropyron inerme (Scribn. and Sm.) Rydb., Wash.—A. smithii Rydb.,
Ida.—A. spicatum (Pursh) Scribn. and Sm., I'Mont., Utah—A. trachycau-
lum (Lk.) Malta, N. Dak., Wyo.—Calamagrostis canadensis (Michx.)
Beauv., Wise.—Deschampsia caespitosa (L.) Beauv., Oreg.—Elymus
glaucvs Buckl., Wyo.—E. triticoides Buck!., ISTev.—Festuca ovina L., Wise.
—Holcus 'lanatus L., B. C, Calif., Oreg., Wash.—Hordswn nodosum L.,
Oreg.—H. vulgare L.,' Sask.—Leersia oryzoides (L.) Swartz,. Wise—
Melica bulbosa Geyer, Wyo.^—Phalaris arundinacea L., Wisc..r—Poa se-
cunda Presl, Utah—Secale cereale L., Oreg.—Sitanion hystrix (Nutt.)
J. G. Sm., Nev.—S. jubatum J. G. Sm.,«Wash.—Triticum aestivum
L., Va.
This disease somewhat resembles leaf smut because of the black
ill-formed stroma which develop in the twisted, malformed shoots and
flowering parts of the grass. Under conditions favorable to the fungus
the black stroma takes on a charred appearance. In Qther instances
there is not a great deal of stroma formed but'ihe white heads form,
or the shoots become arrested in the boot and variously deformed and
twisted. The remarkable spores are not confusable with any other
species, their claw-like appendages cannot be mistaken for any other
fungus. They are, however, small.
Twist is not of very great economic importance in this country as
it is often confined to low-value range grasses. It has been seen in
abundance a few times, such as on Sitanion jubatum in Klickitat
County, Washington. I t is also relatively common on meadow grasses
at times in eastern Oregon and in parts of Nevada and Wyoming.
Atanasoff (1925) presented evidence that the bristles of the spores
adhered to the bodies of wheat nematodes and were carried by these
organisms to the growing points of the hosts. In the regions where the
writer has observed D. alopecuri eelworms are invariably absent.
Schaflnit and Wieben (1928) found that eelworms were not necessary
to transmit the disease. Stieltjes (1933) also found that eelworms were
not necessary to cause infection in oats.
Sampson and Western (1938), Schaffnit and Wieben (1928) and
Sprague (1938,6) all found that there is no genetic connection between
D. alopecuri and Mastigosporium album Riess as had been claimed by
Rainio (1936). There also appears to be no connection with Dilophia
graminis Sacc. (Schaffnit and Wieben, 1928).
Twist disease is not sufficiently important in the United States to
have induced any work toward its practical control. In Europe the
disease is sometimes of moderate importance. Bevilacqua (1935) recom-
mended that the wheat be cut high, the stubble burned, and the 'seed
treated.
"Range: United States, Canada, England, Finland, France, Germany, Italy,
Netherlands, Wales, widely distributed.
References; Atanasoff (1925); Bessey (1906); Bevilacqua (1935); Grove
(1935); Kessler (1921); Kotte (1934); Rainio (1936); Sampson and
Western (1938); Schaffnit and Wieben (1928); Sprague (1938,6;
1946,e); Stieltjes (1933).
Diplodia bambusae Ell. and Langl.—On Shoots

Pycnidia hemispherical, 300-500 /x, papillate; pycnospores ellipsoid,
brown, 1-septate, 15-20 X 8-10 /*. Pycnidia mostly on the nodes.
On: Bambiisa sp.. La.
This species is reported on dead culms, and may not be parasitic.
The genus Diplodia is omnipresent in warm humid regions and this
poorly known species certainly should be critically compared- with con-
generic forms.
Range: United States, probably elsewhere but information not available.
Reference: Ellis and Langlois (1890).
Diplodia frumenti Ell. and Ev.—Dry Rot of Ears and Stalks

See Physalospora zeicola Ell. and Ev. (Fig. 13,C)
A m::
FIG. 13.—Pycnospores of: A, Diplodia macrospora; B, D. zeae; C, D. jrumenti.
Diplodia macrospora Earle—Dry Rot of Ears and Stalks

Pycnidia scattered, large, carbonaceous,, flask-shaped, buried but
with the ostiole erumpent, elevating and rupturing the epidermis, 150-
450 [I, pycnospores short, slender, hyaline; spores very long, dark
fuliginous, irregularly clavate or cylindrical-clavate, unequally uni-
septate but 2- or 3-septate forms do occur, scarcely constricted, each
cell often biguttulate, oozing out and blackening the epidermis (of old
stalks), 43-95 X 6-13 [i, mostly 60-75 X 8-10 /x, averaging 67.8 X 9.0 fi.
On: Zea mays L., Conn., Del., Fla., Ga., La., N. Car., Okla., S. Car., Tenn.
on the ears and stalks—as a leaf spot reported from Alabama, Md.
(rare), Miss. , ^---^^^^^ .
This species is seldom prevalent and even in the southeastern
United States D. zeae is much the commoner of the two. The report
from Maryland represents one isolate out of. many thousands of other
fungi from corn. Eddins examined 618 stalks of corn affected with
Diplodia in Florida (1930,c) and 496 of these /were infected with D.
zeae, 102 with D. macrospora, and 20 with D. jrumenti {Physalospora
zeicola).
Badly diseased ears affected with D. macrospora are shrunken and
shriveled and the husks are tightly glued to th§ ^kernels by the white
mold which spreads through the kernels. Eyemdia are present. The
fungus also attacks the leaf sheaths and sometimes forms yellowish,
later brpv,"n, spots on the leaves. Some pycnidia occur in the spots but
they are much more abundant on the stalks at the nodes. '
The three species of Diplodia, which are the commonest ones on
corn, have radically different spores (Fig. 13), one from another. D.
macrospora produces a light colored dry rot, similar to that caused by
D. zeae but very different from the black ear mold caused by JD. fru-
menti. D. macrospora grows much more slowly in pure culture than
does D. zeae.
Unamuno (1931) reported D. guineae on Lolium perenhe from
Spain. We have no information on this fungus.
Range: United States, Argentina, Brazil, Gold Coast, Philippines, Sierra
Leone, Tanganyika.
References: Earle (1897); Eddins (1930,c); Hoppe (1943); Johann
(1935); Kinsel (1937); Larsh (1938); Stevens, N. E. (1936, 1945).

D. tubericola Ell. and Ev.
See its perithecial stage Physalospora rhodina (Berk, and Curt.)
Cke. Weiss (1945) listed D. cacaoicola P. Henn. and D. theobromae
(Pat.) Nowell as conidial stages of P. rhodina on sugar cane in the
"Gulf states and tropical America." It was listed as the cause of a dry
rot of the canes and a seed cane rot. This fungus is apparently common
in the tropics and subtropics on many hosts but is not important on
Gramineae in the United States.
Diplodia zeae (Schw.) Lev.—Stalk Rot, White Ear Rot

D. maydis (Berk.) Sacc.
Producing a white mold on the ears, pycnidia globose or flask-
shaped, often multichambered and irregular, immersed but with a
definite beak protruding through the epidermis; pycnophores simple,
short and pointed, pycnospores ovate, 2-celled, straight or slightly
curved with rounded to bluntly tapered ends, 25-30 X 4.5-6.5 ju,, oliva-
ceous, slender, filiform scolecospores occur.
On': Zea mays L., Ark., Calif., Conn., Del., D. C, Ga., 111., Ind., Iowa, Kans.,
Ky., La., Md., Mass., Mich., Minn., Miss., Mo., Nebr., N. J., N. Y.,
N. Car., N. Dak., Ohio, Okla., Ont., Penn., R. I., Sask., S. Car., S. Dak.,
Tenn., Tex., Va., W. Va., Wise.
Dickson (1947) gives a lengthy list of synonyms and associated
names for Z>. zeae. Shear and Stevens (1935) present data indicating
that T). maydis is the correct name for this fungus but they prefer to
conserve D. zeae as being more practical, everything considered. Dick-
son has followed this suggestion, as does the writer.
Woronichin (1925) proposed Phaeostagonosporopsis zeae (Schw.)
Wor., erecting a new genus which is fundamentally a brown spored
Stagonosporopsis, or a Hendersonia with loosely constructed pycnidia.
Diplodia as a genus probably contains some forms better assigned to
Hendersonia. However D. zeae (Fig. 13,B) which even forms scoleco-
spores can just as well be left in Diplodia as being jplaced in t h e "mega-
nomial" Phaeostagonosporopsis.
Dry rot of the ears and stalk rot caused by this species does seri-
ous damage at times in the Middle Western corn belt. There has been
a great deal written about the fungus and the disease it causes. I t is
rare or unknown in the Far West, probably because of the relatively
dry summers and autumns. ' . •
Range: United States, Argentina, Australia, Belgian Congo, Brazil, Canada,
Mexico, S. Africa, Tanganyika.
References: Bijl (1916); Branstetter (1922); BurrUl and Barrett (1909);
Butler (1947); Clayton (1927); Dickson (1947, pp. 80-84); Durrell
(1923, 1925,a); Eddins and Vorhees (1933); Heald (1906); Heald, Wil-
cox, and Pool (1909); Hoadley (1942); Holbert, et al. (1924); Holbert,
Hoppe, and Smith (1935); Hoppe (1936); Hoppe and Holbert (1936);
Ireland (1928); Kent (1940, 1942); Kinsel (1937); Koehler and Holbert
(1938); Larsh (1938); Manns and Adams (1921,6); MeNew (1937);
Melhus and Durrell (1922); Raleigh (1930); Semeniuk (1940,a,6; 1941;
1942,a); Semeniuk, et aA (1940-41, 1944)'; Shear and Stevens (1935);
Smith and Trost (1934); Stevens (1945); UUstrup (1943, 1948); Val-
leau, et al. (1926); Wilson, W. E. (1942)/; Young, H. C, Jr. (1943).
Hendersonia spp.—Molds
Some of the pioneer invaders of dead or necrotic plant tissue are
species of Hendersonia Berk. em. Sacc. While these fungi are sapro-
phytes or very weak parasites, the scattered pycnidia will be encouuj-
tered by routine workers. Therefore they are mentioned in this text, at
least to the extent of including a key to the forms found in the United
States, especially those west of the Mississippi where we have con-
ducted spasmodic mycological studies with the genus during the past
twenty years. {
The multiseptate, brown spores of Hendersonia are broadened, not
filiform, nor elongate-vermiform as in some species of Phaeoseptoria
(Sprague, 1943,6). However we have h a d ' a difficult time with this
group because many collections have narrow spores which-,dp not fit
the description of Phaeoseptoria but which would perhaps outrage other
workers' ideas of Hendersonia. Most of these forms we place in Hender-
sonia but without any dogmatic statement that this is necessarily the
proper place for them. Not only have we found the group difficult to
segregate along genera and species lines but we have encountered more
than usual difficulty in obtaining type material for comparison. Fortu-
nately from the standpoint of the plant pathologist few, if &ny, of the
forms which we list in the key should be considered ^further. Two
species may be weakly parasitic, H. culmicola Sacc. and/H. crastophila
Sacc. (H. graminis, Wojnowicia graminis) and'ilherefore descriptions
of these are included.
Hendersonia molds are especially common on overwintering grasses
in the Pacific Northwest and Rocky Mountain areas and on grass in
FUNGI IMPERFECTI-^PHAEROPSIDALES 173
the late fall after killing frosts have occurred in the Northern Great
Plains. Among the common species are H. culmicola Sacc. in various
sizes, H. crastophila Sacc, H. calamovilfae Petr., H. stipae-pennatae
Fautr., and H. calospora Fautr. Some forms are apparently undescribed.
SIMPLIFIED KEY TO THE HENDERSONIA SPECIES ON GRAMINEAE
A. Spores when mature typically 7-septate

B. Pycnidia very large, 180-300 /* in diameter
C. Spores falcate or falcate-fusiform, 28-40 X 3.8-4.4 A
Hendersonia crastophila Sacc.
CC. Spores fusiform, vermiform, coarse, somewhat constricted at the
septa, 35-53 X 3.5-5 /*
H. calospora Fautr.
BB. Pycnidia smaller, 80-220 n in diameter
C. Spores regularly cylindrical, ends blunt, cross walls closely adjacent,
20-29 X 4-6 M
H. stipae-pennatae Fautr.
CC. Spores more or less fusiform, subcylindrical; ends or at least one
end somewhat to definitely tapered; septa not particularly closely
adjacent
D. Spores as much as 7 /» wide
H. calamovilfae Petr.
DD. Spores mostly 3-5 f* wide '
E. Spores brown, fusiform or falcate, flattened on one side, con-
stricted at the septa. ^
H. crastophila Sacc.
EE. Spores lighter colored; small, paired, regularly placed, promi-
nent guttulae; often constricted at septa, subcylindrical, ends coni-
cal, not stiffly flattened at one end
Hendersonia sp. (W. Oregon)
EEE. Spores shorter than in EE, rarely constricted at septa, more
often 5- than 7-septate
H. culmicola Sacc.
AA. Spores when mature typicafly 5-septate or less (see also H. cidmicola
above)
B. Spores sometimes more than 3-septate, cylindrical
C. Spores 3-, 5-, 7-septate, 30-38 X 3.1-4.2 /*
H. culmicola Sacc.
CC. Spores 3- to 4-septate, 20-27 X 3.2-3.4 n
H. culmicola (var. intermedia Sacc.)
BB. Spores not more than 3-septate
C. Spores with stout epispores, 6.2-7.8 j" wide
Hendersonia sp. (alpine)
CC. Spores with thin walls, less than 5 /* wide
D. Spores 28-48 )<; 3.8-4.3 /«
Hendersonia sp. (Coastal Oregon)
DD. Spores less than 30 M long or approximately that
E. Spores short-cylindrical, some at least 4 n wide, spores 22-29 X
2.8-4.6 !>•
H. culmicola (on Hordeae)
/I
EE. Spores subcylindric to fusiform, not over 3-5 M wide
F. Spores often 1-septate, finally 3-septate, sublanceolate at first,
17-22 X 2-2.6 M . •- ' i
H. culmicola (var. minor Sacc.)
FF. Spores mostly 3-septate ,j
G. Spores bacillar or sub-bacillar, 24-29 X 2.6-3.4 /*>
H. sp. on Calamagwstis inexpansa (alpine)
GG. Spores subfusiform or less uniformly cylindrical
H. Spores 20-25 X 2.3-3.8 /i.
H. sp. on Erharta undtdata
HH. Spores 23-28 X 2.1-2.4 u.
H. sp. on Melica sp. .
HHH. Spores 14-26 X 2.6-3.1 ii
H. culmicola (var. minor)
EEE. Spores navicular or broadened
F. Spores 11-16 X 3.9-4.4 n ^
H. culmiseda Sacc.
FF. Spores 12-15 X 6-8.5 /^ , ,
H. grantii Dearn.
/
Hendersonia crastophila Sacc.—Secondary Foot Rot, Rot of
Mature Straw
H. graminis McAlp.
Wojnowicia graminis (McAlp.) Sacc. and D. Sacc.
Saprophyte and weak parasite on the crown and culm lumen and as
a leaf rot without definite spotting; pyenidia sunken, basal portion
more or less globose but ostiolar region with a short neck which has
a few, scattered, stiff, spiny hairs about the ostiole, brown, firm-walled,
sometimes with considerable brown mycelium/ ramifying from the
pycnidium, 90-360 ft, sometimes large; pycnospores yellow-brown with
small inclusions often adjacent to the cross walls, slightly flattened on
one side, curved spindleform to nearly straight, usually 7-septate,
24-45 X 3-5 n, rarely as large as 43-51 X 4.5-6 fx (on Agropyron repens
in this latter case). ^ -^^
On: Agropyron cristatum (L.) Gaertn., N. Dak., Oreg., Wash.—A. inerme
(Scribn. and Sm.) Rydb., Wash.—A. intermedium (Host.J" Beauv., N.
Dak.—A. repens (L.) Beauv., N. Dak.—A, riparium Scribn. and Sm.,
Wash.—A. smithii Rydb., Mont., N. Dak., Wyo.—A. subsecundum (Lk.)
Hitchc, N. Dak.—A. trachycaulum (Lk.) Malê, N. Dak.—Avena sativa
L., Oreg., Wash.—Bouteloua gracilis H. B. K.,'N. Dak.—Bromus inermis
Leyss, N. Dak.—B. tectorum L., Ida., Oreg., Wash.—Calamagrostis
canadensis var. scabra (Michx.) Beauv., Alaska—C. rubescens, Buckl.,
Mont.—Calamovilfa longifolia (Hook.) Scribn., N. 'Dak.—Dactylis glo-
merata L., N. Dak., Wash., Wyo.—Distichlisp-stricta (Torr.) Rydb.,
N. Dak.—Elymus ambiguus Vasey and Scribn., 'N, 'Dak.—Festuca ida-
hoensis Elmer, Oreg.—F. pacifica Piper, Wash.—F. rubra L., Oreg.—
Hordeum jubatum L., N. Dak.—H. vvlgare L., Calif., Colo., Ida., N. Dak.,
Oreg., Wash.—Koeleria cristata (L.) Pers., Ida., Wash.—Phleum pratense
L., Minn., N. Dak., Wyo.—Poa ampla Merr., Wash.—P. bulbosa L., Wash.
—P. canbyi (Scribn.) Piper, N. Dak., S. Dak.—P. interior Rydb., Wyo.—
P. pratensis L., Wyo.—P. secunda Presl, Mont., N. Dak., Wash.—Se-
cale cereale L., Ida., Mont., Oreg., Wash.—Sorghastrum nutans (L.)
Nash, N. Dak.—Spartina pectinata Lk., N. Dak.—Stipa lemmonii
(Vasey) Scribn., Oreg.—S. viridula Trin., Mont., N. Dak.—Triticum
aestivum L., Alberta, Cahf., Colo., Ida., Kans., Mont., Oreg., Sask.,
Wash., Wyo.—T. dicoccum Schrank, Oreg., Wash.—T. durum Desf.,
Oreg., Wash.—T. monococcum L., Oreg., Wash.—T. spelta L., N. Dak.,
Oreg., Wash.
The fungus H. crastophila and Wojnowicia graminis appear to
be one species. We have placed this fungus in Hendersonia where it was
originally placed before the Saccardos transferred it to Wojnowicia.
The latter genus is said to differ from Hendersonia in the presence of
hairs on the neck of the pycnidium. A better differential character,
perhaps, would have been to emphasize the presence of a definite ostio-
lar "neck. However, this character does not seem to be too well fixed,
even in the species graminis. After long delay we have somewhat reluc-
tantly given up the now better-known Wojnowicia graminis and taken
up the old name Hendersonia because we do not believe that the genus
Wojnowicia is necessary.
H. crastophila is a weak parasite, noted largely as an associate of
Cercosporella herpotrichoides Fron in the prairies of the Columbia
Basin but it occurs far beyond this region into the dry wheat lands of
the western United States, Canada, and probably Mexico. It is common
in Kansas, Colorado, and westward. It is not abundant in the hard
red spring wheat area but occurs on perennial grasses in the Dakotas,
often on dead leaves and on the crowns. It is an associate of Ophiobolus
graminis in Australia and with various root-rot and straw-breaker
fungi in Europe.
Most of the pure culture isolates of H. crastophila are olive gray
and cottony with dark or sometimes brightly colored pigments in the
depths of the medium.- Pycnidia are produced infrequently on potato
dextrose agar incubated in darkness in electric refrigerators but many
cultures sporulate readily under north light at common storage tem-
peratures on various organic materials such as sterilized wheat heads
(Sprague, 1935,d).
Fully mature spores of H. crastophila are easily recognized by their
bright brown color, strongly marked septations, and the fiattened-on-
one-side appearance. In developing material, however, the spores are
paler, narrower, less strongly flattened.
While H. crastophila is feebly parasitic it is of definite economic
importance on account of its ability to invade the culms of plarits with
ripening grain and thus predisposing them to premature lodging. This
condition is common in the Palouse region of Washington and Idaho
where it increases the cost of harvesting and lowers the quality of the
grain. The fungus invades the lumen of the culm through the crown,
, I
spreading upwards through the pith cells for as great a distance as 6
inches above the soil line. The culms have a discolored, leaden color
from the outside. This saprophytic develdpmerit weakens the ripened
culm with its heavy seed load, causing lodging in the event heavy rains
and winds occur before harvest. This condition was formerly more
important than now because of the rich, then, still uneroded soil of
swales and areas where lush grain grew. I t was widespread during 1948,
however, a year of exceptionally heavy precipitation. This coftdition
has been mentioned earlier (Sprague, 1946,e) but there is little infor-
mation on the subject. Carroll (1943) discussed a comparable situation
in Eire.
Range: United States, Australia, Canada, France, Germany, Sweden, widely
distributed in Europe.
References: Broadfoot (1933,a); Ludbrook (1936); McKinney and John-
son (1921); Sprague (1935,d; 1946,c,e); Sprague and Fellows (1934).
Hendersonia culmicola Sacc.—Leaf Rot

Hendersonia simplex Schrtr. ^
Septoria simplex (Schrtr.) Sacc. (Sylloge Fungorum v. 3, p. 560)
Pycnidia in vague, brown, later straw-colored spots, sometimes with
brown margins, pycnidia gregarious but not abundant, globose but flat-
tened, nearly black, obscure in faded lesions, 100-185 [i, ostiolate;
pycnospores bacillar, sometimes curved at central septum,, slightly
acute at apex, more often rounded, yellow-hyaline, 0-, 3-, 5-, 7-septate,
finally light brown, 17-35 X 1.9-4.6 //,.
On: Agropyron repens (L.) Beauv., N. Dak.—A semicostatum, (Steud.)
Nees, N. Dak.—A. spicatum (Pursh) Scribn. a^d Sm., Oreg.—A. trachy-
cavlum (Lk.) Malte, Wash.—Bromus vulgaris L., Oreg.—Deschampsia
caespitosa (L.) Beauv., Oreg.—Elymus glaucus Buckl., Oreg., Wash.—
Festuca idahoensis Elmer, Calif., Oreg.^—F. ovina var-. brachyphylla
(Schult.) Piper, Mont.—F. pacifica Piper, Oreg.-—F. rubra L., Oreg.—
Koeleria cristata (}J.) Pers., Oreg.—Melica subulataîGriseh.) Scribn.,
Ida.—Oryzopsis asperijolia Michx., N. Dak.—Poa canbyiXÔTihn.) Piper,
N. Dak., S. Dak.—P. com,pressa L., Calif.—P. gracillimaâsey, Calif.,
Oreg., Utah—P. palustris L., S. Dak.—P. pratensis L., Minn., N. Dak.—
P. secunda Presl, Wash.—Schizachne purpurascem (Torr.) Swallen,
Minn.—Sporobolus airoides (Torr.) Torr., Wash.—S. asper (Michx.)
Kunth, Wash.—Trisetum spicatum (L.) Ricllt., Mont. '
This fungus may be weakly parasitic but is more probably only a
primary invader of necrotic or virtually dead tissue in late season. The
small-spored var. minor was discussed recently (Spra'gue, 1949,a) and
in these observations its pathogenicity was. ooRSidered of a low order.
All collections show hyaline juvenile spores but as septations in-
crease their color darkens and the spores also become larger. Aberrant
collections with tapered apices occur. >
FUNGI IMPERFBCTI—SPHAEROPSIDALES 177
Range: United States, England, France, Germany, Italy, Scotland, Sweden,
Wales.
References: Grove (1937); Saccardo (1884, v. 3); Sprague (1946,6,e;
1948,c; 1949,a).
Hyalothyridium calamagrostidis Greene—Leaf Spot

Spots amphigenous, ellipsoid or fusiform, sometimes confluent, paler
above with a narrow, dark purple border; 0.25-1.5 cm. long; pycnidia
rudimentary, subepidermal, scattered, deeply sunken in the host tissues,
amphigenous, globose, erumpent, ostiolate, 110-135 /* diam., wall com-
posed of pale brown, flattened, compressed cells, almost concolorous
with surrounding host cells; conidiophores not differentiated, but
sporiferous tissue occupying the lower half of the inner surface of the
pycnidium; conidia hyaline, muriform, many-celled, variable, narrowly
to broadly ellipsoid, ovate or obovate, 30-45 X 16-23 fi, usually with
a p'ointed, hyaline apiculum, 3-13 (mostly 4-6) X 1-5-2.0 /*.
On: Calamagrostis canadensis (Michx.) Beauv., Wise.
Greene has recently described this interesting fungus, a pycnidial
form with hyaline, muriform spores. It is the hyaline counterpart of
Camarosponum. ,
Reference: Greene (1946,a).
Leptostromella cynodontis Sacc.—Decline

Stromata linear, in parallel series, dark, erumpent, 0.5-0.8 mm. long,
spores bacillar, 70-75 X 2.5-3.0 /t, curved, hyaline.
On: Cynodon dactylon L., Alabama.
Said to be associated with a turf decay in the South. The black,
linearly arranged stromata are relatively prominent on the necrotic
leaves of the host. C. L. Lefebvre showed material of this fungus to the
writer.
Reference: Saccardo (1903).
Leptostromella panici Dearness—On Leaves

Pyncidia minute, 50-250 X 50. /x, innate, amphigenous but more
numerous on the upper side,,close together in lines between the veins;
pycnospores hyahne, linear oblong, 18-22 X 2.75-3.0 ix., 3-septate on
short pycnophores.
On: Panicum Mans Elliott, S. Car.
This species, which is not known to the writer, is included because
Dearness (1924) reported it on living and languishing leaves, indicating
/I
that it could be somewhat parasitic. It was collected at Orangeburg,
South Carolina, on August 15,1905, by R. Latham (No. 667).. Dearness
noted that it was externally similar to L. septorioides S. and R.^
Leptostromella Sacc. is a genus with about 20 species beloi^ging to
the hyaline scolecosporous section of Leptostroinataceae. The elongate
pycnidia are typical. i
Range: United States. ,
Reference: Dearness (1924).
Leptothyrium avenae Tehon—On Glumes and Pedicels

Pycnidia substomatal, in rows on the outer faces of glumes and on
pedicels, subcuticular, black, remaining immersed, with cells of the
outer wall aliform and radiate, for the most part broadly elliptical,
sometimes nearly round, 100-200 X 65-140 /x, pore minute, substomatal;
spores hyaline, nonseptate, oval, oblong or pediform, 16-23 X 4-8.5 /x,;
sporophores simple, short, acute. '
On: Avena sativa L., 111. /
The spikelets when attacked on the glumes and pedicels by this
fungus are said to develop poorly (Tehon, 1948) and bear small grains.
In addition the pedicels of infected plants are short and they spread
less widely from the axis of the panicle when compared with healthy
ones.
This species differs from Leptostroma avenae Brun. in the lack of
a longitudinal slit type of ostiolum and by its longer, wider spores.
Reference: Tehon (1948). /
iI •
Leptothyrium cylindricum Atk.—Leaf Spot
I
Spots gray white, finally elliptical \\îth a dilute brown margin;
pycnidia amphigenous, dimidiate, black, astombus^pore variable in
dehiscence, pycnidia 300-400 /* long, depressed; spores cylindrical, both
ends abruptly attenuate, straight or somewhat curved, 18-^5 X 3.5-4 /A.
On: Arundinaria gigantea (Walt.) Chapman,-Alabama.
There is no information available on this fungus in its role as a
parasite of Southern cane. The genus Leptothyrium belongs in the
Leptostromataceae in Section Hyalosporae while Leptostromella be-
longs in Section Scolecosporae of the same family. This family has the
pycnidia membranaceous, or carbonaceous, black, more or less dis-
tinctly dimidiate, scutiform, astomous, osti^ffte or cleft, eruhapent or
superficial.
Reference: Atkinson (1897).
FUNGI IMPERFEGTI—SPHAEROPSIDALES 179
Leptothyrium zeae Stout—Leaf Spot

Foliicolous; pycnidia hypophyllous, not gregarious but occurring in
small patches, sometimes on elongate-oblong, tan-colored spots which
are bounded laterally by the leaf veins, subcuticular and possibly sub-
epidermal, circular dimidiate, dark-brown, membrano-subcarbonaceous,
parenchymatically-reticulate, 55-225 /i diam., without ostiole; spores,
nonseptate, hyaline, irregularly globose, sometimes with one or more
flattened sides, 8,5-13.5 /A, their walls somewhat thickened.
This species was collected three times in November 1926 and 1927
in Illinois (Stout, 1930). Proof of its pathogenicity is lacking. One col-
lection was associated with Mycosphaerella zeicola Stout. However in
other material Phyllosticta zeae was associated with this ascomycete
and in still other material it occurred with Septoria zeae. Therefore the
evidence is against assigning any of these imperfects to this Myco-
sphaerella. Stout has described a large number of species on corn, some
of which it has been necessary to consider in this text. The fact that
much of this material was collected late in the fall leads the writer to
suspect that most of the described species are saprophytes, or' ones
capable of invading only necrotic, frost-injured leaves. This would be
at least partially verified by the fact that few of Stout's species have
been noticed by pathologists in states other than Illinois where his
mycological studies were made. Until the parasitic status of these
forms is clarified it will be necessary to include some of them in this
text, depending in our selection on evidence obtained in congeneric
forms or from the kinds of lesions produced on the hosts.
Leptothyrium spp. or similar forms are not uncommon on dead
grasses, especially from alpine areas. Cursory observations have always
indicated, however, that these forms are saprophytes.
Macrophoma Sacc.
This genus has elongate to elongate-ovate, hyaline spores usually
with pure white, somewhat opaque contents borne in relatively robust
pycnidia, often-with small ostioles. The spores are nonseptate, larger
than Phoma, usually 20 [i or more long and 4-10 fi wide. None of the
described species on Gramineae appear to be actively parasitic. One
name, M. fiennebergii' {Kuehn) Berl. and Vogl. is Septoria nodorum
Berk.
Tehon (1937) has prepared a key to the species of Macrophoma on
Gramineae. We are including this key, with additions. Tehon points
out that M. suspecta Pk. is Ascochyta graminicola Sacc.
/'
' I
KEY TO SPECIES OP MACROPHOMA ON GRAMINEAE
A. Pycnidia aggregated in crusty or stromatic, masses
B. Crusty masses extensive, pycnidia numerous, on wheat
M. crustosa Sace. and Berl.
BB. Stromata small and round, pycnidia few \
M. erumpens (B. and C.) Berl. and Vogl.
AA. Pycnidia separate and essentially discrete
B. Pycnidia oblong in outline, on Poa
M. oblongata Tehon
BB. Pycnidia round to oval in outline
C. Spores mostly more than 25 fi long
D. Spores 15 /» or more wide, on Zea
M. zeae Tehon and Daniels
DD. Spores 10-12 i^ wide
E. Spores mostly 30-40 /* long, on Saccharum
M. sacchari (Cooke) Berl.
EE. Spores mostly 25-30 M long,-on Koeleria
M. arens Davis
DDD. Spores 5-6 jf wide, on dead grass
M. melanostigma (Lev.) Sacc.
DDDD. Spores 8.5-11 M wide, on Elymus and Agrostis
M. phlei Tehon and Stout (but see below)
CC. Spores mostly less than 25 ii long
D. Spores mostly more than 20 /«long
E. Spore width constant, 6.5-7.5 /t, on Phleum
M. phlei Tehon and Stout
EE. Spore width variable, 6.5-10 /i, on Secale
M. secalina Tehon
DD. Spores less than 20 fi long
E. Spores 18-20 X 6-7 M on Sorghum, pycnidia gregarious
M. sorghicola Speg.
EE. Spores 7.5-19 X 5.6-8.5 f., variable. On Sporobolus
M. sporoboli Sprague
EEE. Spores about 4 /* wide, on Calamagrostis
M. graminella (Sacc.) Jierl. and Vogl.
As will be pointed out in the discussion of M. phlei thejdentification

of species of Macrophoma with this key will prove difficult unless ad-
ditional species are recognized or, better, -the dimensions of the spores
of a fewer number of species be increased in range to permit greater
variability than this key indicates. j
Macrophoma oblongata Tehon—^Leaf Mold

Pycnidia caulicolous but not in spots, oblong with bluntly rounded
ends, oriented parallel with the culm and' set in rows between the
sclerenchyma strands, dark brown, membranaceous, and translucent,
becoming carbonaceous and opaque, 95-135 ;u, wide, 200-500 /* long,
usually discrete but less often joined terminally. Ostiole central, round,
not papillate, the opening 20-25 ji diam.; spores hyaline, continuous,
oblong-ovate, with rounded ends, 19-26 X 6.5-8.5 /i, borne on tapering
conidiophores up to 10 [>, long and 2.5-3 jx wide.
On: Poa pratensis L., 111.
This may be a saprophyte. I t was collected in August, 1935, and the
type is 111. Ace. No. 25,381.
Macrophoma phlei Tehon and Stout—Leaf Mold

Foliicolous, no evident spots; pycnidia appearing abundantly on
dry, dead leaves from tip to sheath, subsequent to fruiting of host, not
gregarious but disposed in rows closely adjacent to the leaf veins,
membranaceous, subcarbonaceous or completely carbonaceous, devel-
oped in and occupying the mesophyll, rotund or applanate in long
section, often oval in outline, the long axis parallel with the vein when
viewed from above, 105-225 /t diam., erumpent hypophyllously only;
ostiole papillate, at first closed by a thin, cellular membrane which dis-
solves when the spores mature, leaving a very distinct, irregularly, cir-
cular stoma, 17-28 /j. in diam. Spores hyaline, continuous, oval, 18-
26 X 6.4-7.7 IX (on Phleum). ,
On: Agrostis -palustris Huds., Oreg.—Blymus triticoides Buckl., N. Dak.—
Phleum pratense L., 111.
This dubious parasite is characterized by the large, elongate, ovate,
nonseptate spores with pure white, opaque contents (Fig. 14). The
material on Elymus triticoides from Mandan, North Dakota, showed
diffuse straw-colored spots, or in some cases the pycnidia were formed
in undefined areas. The pycnidia were black, depressed, globose, 120-
135 X 120-204 [x, bearing elliptical to elongate-ovate spores, which
were slightly larger at one end, with pure white opaque contents, 28-
39 X 8.5-11.1 IX (Fig. 14:,A), mean size 32 X 9.4' [x. M. -phlei was also
found on Agrostis palustris at Waldport, Oregon, with spores 24-
3 1 x 9 . 5 - 1 1 / . (Fig. U,B).
Tehon and Stout (1929) state that this species resembles Sphaeropsis
phlei Ell. and Ev. except that the spores are colored in the latter.
We are convinced that the Macrophoma species are distinct from
Sphaeropsis.
It was necessary to modify Tehon's key to the graminicolous
Macrophoma spp. to permit including the collections of M. phlei on
Elymus and Agrostis. We question if it would be practical to add
further species having a spore range averaging about 30 /x length but
varying a few microns in width, one from another. I t would be better
to place them in the species to which they show the most resemblance.
While we may be somewhat arbitrary in selecting M. phlei it seemed
to us the most logical assignment for the- few collections which we
chanced on in routine examinations of grass foUage.
Davis (1924) described M. arens on Koeleria cristata (L.) Pers.
from Boscobel, Wisconsin, as follows: "Pycnidia'Ijlack, scattered, sub-
epidermal, globose, 130-150 ju, diam., sporules narrow, ovoid, becoming
subfuligenous, 27 X 10 /* at maturity. On more or less distal parts of
the leaves which become sere and involute." This fragment was later
FIG. 14.—Pycnospores of: A, Macrophoma phlhi on 'Elymus triticoides, Man-

dan, N. Dak.; B, M. phlei on Agrostis palusiris, jNewport, Oreg.
rejected by Davis (1942) who stated that,this species was "Stricken

out, the specimen being poorly developed and its character doubtful."
Based on Davis' description M. arens is probably the same~as-M. phlei
which it antedates by five years. However since Davis doubted his
diagnosis it is probably necessary to ignore M. arens on the basis of
available information. Our collections of Mabrophoma have been scanty
also.
/
References: Davis (1924, 1942); Sprague (1946,o,e); Tehon and Stout
(1929, p. 188).
Macrophoma secalina Tehon^-4jeaf Mold

Not in spots, pycnidia scattered and separate, amphigenous, on
leaves', spherical to. decidedly applanate or almost dimidiate, dark
brown to black, mem'branaceous and translucent to heavily carbonized
and opaque, 145-260 /x diam., ostiolate, apparently subcuticular and
developed from a subcuticular plate of aliform mycelium; ostiole circu-
lar, carbonized, somewhat papillate, 20-25 ju diam. Spores hyaline, ob-
long to somewhat pyriform, rounded above, with a short or well-
defined basal stalk, 19-30 X 6.5-10 /x; pycnophores simple, hyaline,
nonseptate, often hourglass shape, 8-14 X 2.5-3 ft wide, spores acrog-
enous, not catenu'iate.
O n : Secale cereale T-i., 111.
This fungus was found on the lower dead leaves and sheaths of rye
in two localities in Illinois. Type is 111. Ace. No. 23,913.
Macrophoma zeae Tehon and Daniels—See Physalospora zeae Stout
Macrophomina phaseolina (Tassi) G. Gold.—Charcoal Rot

Dothiorella phaseoli (Maubl.) Petrak and Sydow
M. phaseoli (Maubl.) Ashby
- Sclerotium bataticola Taub.
Macrophoma phaseolina Tassi
Produces a spongy rot and eventual shredding of the base of the
• culms and adjacent tissue, the softer cortical and pith cells decay,
leaving the vascular tissue and sclerenchyma cells; sclerotia variable,
small in general, black, numerous, scattered through the interior of the
diseased tissue, 50-150 /j., but on woody plants up to 1 mm. diam.,
Bome'ii.Tnes HTmieTOMS ^-tityagVi \,o grvs a cWxTed appBaTKtite", pjicnidia
with small ostioles, subcarbonaceous, mostly 100-200 /x. diam.; conidio-
phores arising from hyaline inner layers, 10-15 /x long, rod-shaped;
spores hyaline, cylindrical or ovate-elongate, nonseptate, 14-31 X 6-
11.2 II.
' On: Bouteloua curtipendula (Michx.) Torr., Okla.—Muhlenbergia mexicana
(L.) Trin., Ind.^M. schreberi Gmel., Va.—Sorghum vulgare Pars., Ark.,
Calif., Kans., Nebr., Okla., Tex.—S. vulgare var. sudanense (Piper)
Hitchc, Okla.—Zea mays L., Ark., Calif., Colo., Ga., 111., Iowa, Kans.,
Mo., Nebr., Okla., S. Car., Tex.
Charcoal rot occurs on a large number of cultivated plants in the
southern United States and north to Nebraska, Iowa, and northern
California. It attacks mature plants or ones well past the seedling
stage. It can cause seedling injury in the greenhouse, however
(Semeniuk, 1944). Dunlap (1938) reported symptoms of the disease
as follows: affected spots in the field are circular with most of the
culms broken over a few inches above soil level. Diseased plants are
dried, show premature ripening, under-developed kernels, disintegra-
tion of the pith in the lower stalk and collapse of the stem. The small
black sclerotia were prominent and numerous in the pith.
/I
Hoffmaster, McLaughlin, Ray, and Chester (l'943) propose control
measures as follows: increase crop vigor, lime, soil for sorghum, in-
crease organic matter, use resistant varieties which ..include sweet
sorghums, some of the kaffir varieties, and certain other sorghums.
Resistance in some corn hybrids is also known. ' | '
Ashby (1927) lists a large number of hosts, several synonyms, and
gives the geographical range of the fungus. He lists Macrophoma
phaseoli Maubl. (1905), Sclerotium bataticola Taub. (1918), Ma-
crophoma corchori Saw., M. cajani Sydow and Butler, Macrophomina
philippinensis Petr. (1923), Rhizoctonia lamellifera Small (1924), B.
bataticola (Taub.) Butl. (Briton-Jones, 1925), Dothiorella cajani Syd.
and Butler (Petrak and Sydow, 1925), and Macrophoma sesami Saw.
Ashby states that the pycnidia are membranaceous or subcarbonaceous,
at first immersed but later becoming more or less erumpent, globose,
or depressed-globose with an inconspicuous truncate ostiolu'm. No
stroma occurs. The walls of the pycnidium consist of 3-4 layers of
blackish brown, thin-walled, angular cells- about 9 /i thick and inside
these layers are about three layers of hjjjaline cells which give rise to
the pycnophores. The thin-walled pycnospores have granular contents,
are variable in shape, elliptical or oval, at times with irregular contour
and more or less bent. Along this Ime Petrak (1923) described the
genus Macrophomina as having heavy walls with a somewhat dothidi-
aceous aspect. Petrak is probably correct in segregating this unusual
fungus from Macrophoma which has truly parenchymatous pycnidia.
Recently Goidanich (1947) proposed the combination given herein
for this fungus. The genus is similar to Dothiorella Berl. and Vogl.
sensu Petrak and Sydow, differing in the absence in the pycnidium of
a typical extramatrical stroma. /
Range: United States, Ceylon, East Africa, Egypt, Formosa, India, Italy,
Palestine, Philippines, Romania, West Indies.
References: Ashby (1927); Briton-Jones (1925); Dunlap (1938); Goidanich
(1947); Hoffmaster, McLaughlin, Ray, and Chester (1943); Livingston
(1942,a,6; 1945); Maublanc (1905); Miller and-Wooli (1447, Fig. 2);
Petrak (1923); Semeniuk (1942,6, 1944); Shaw and Ajrekar (19T5); Small
(1924, 1926); Taubenhaus (1913); Uppal, Kohlatkar, and Patel (1936).
Melasmia setariae Atk.—Leaf Spot

Spots epiphyllous, elliptical-oblong, small, confluent or thickly scat-
tered over a large part of the leaf. Continuous membrane covering sev-
eral pycnidia or sporogenous groups composed of layers of' dark brown,
angular cells, 3-4 j«, diam. Sporogenous stroma occupying and com-
pletely filling the cavity of the epidermal cells^^ôres oblong-elliptical,
with rounded ends, hyaline, 12-14 X 2.5-3.0 /t7 Conidiophores short,
colored. ^
On: Setaria viridis (L.) Beauv., Alabama.
FUNGI IMPERFECTI-SPHAEKOPSIDALES 185
This is listed'as occurring on living leaves (Atkinson, 1897). I have
not seen the specimen. The genus is sometimes the conidial stage of
Rhytisma.
Reference: Atkinson (1897).
Microdiplodia sp.—Leaf Spot

Apparently, at this writing, a species of Microdiplodia on sorghum
from Louisiana is undescribed (Miller and Wood, 1947). It was found
in an experimental planting. I am not familiar with the genus. Clements
and Shear place it under Diplodia (1931).
On: Sorghum vulgare Pars., La.
Reference: Miller and Wood (1947, p. 9).
Phaeoseptoria spp.—Leaf Molds and Seconaary Invaders

This genus which belongs in the Scolecosporae of the Sphaeropsi-
dales is composed of a number of species which are saprophytes on
Gramineae. Because they will be encountered at times they will be
mentioned here. Phaeoseptoria species have coarse, black pycnidia and
light brown, multiseptate, elonga'te spores (Fig. 15). A key to this
group follows.
KEY TO SPECIES OP PHAEOSEPTORIA ON GRAMINEAE
A. Spores filiform-bacillar
P. elymi Sprague
AA. Spores clavulate-filiform
B. Spores yellow to light brown
C. Spores 60-75 ^ 2.5-3.0 i^
P. airae (Grove) Sprague (Fig. 15,G)
CC. Spores narrow, 50-77 X 2.1-2.4 M
P. poae Sprague
CCC. Spores wider, 55-64 X 3.7-4.5 /*
P. festucae var. muhlenbergiae Sprague (Fig.
15,-D)
CCCC. Spores 75-115 X 2.4-3.4 ^
P. festucae Sprague on Festuca -ovina var.
brachyphylla (Schult.) Piper
BB. Spores light brown, coarser '
C. Spores 5- to 7-septate, 20-90 X 4.5-6 M
P. urvilleana (Speg.) Sprague
CC. Spores 8- to 13-septate, narrower
D. Spores vermiform, obtuse, 55-71 X 4.3-5 M
P. calamagrostidis Sprague
DD. Spores obclavate-filiform, scarcely vermiform, acute at one end
or at least acuminate
FIG. 15.—A, pycnospores of Phaeoseptoria •phalaridis on Phalaris arundinacea,

Corvallis, Oreg. (O.S.C. 17); B, pycnospores of P. festucae on Festuca rubra,
Garibaldi, Oreg. (O.S.C. 8376, type); C, pycnospores of P. festucae on,F. rubra,
King's Valley, Oreg. (O.S.C. 202); D, pycnospores of, P. festucae var. muhlen-
bergiae, Lansing, Mich. (H. H. Hicks, 1211, type); E, segment of a cross section
of the pycnidial wall of P. festucae on F. rubra, Garibaldi, Oreg. (O.S.C. 8376,
type); F, segment of a cross section of the pycnidial wall of P: phalaridis on
Phalaris arundinacea, Corvallis, Oreg. (O.S.C. 17); G, pycnospores of P. airae on
Deschampsia caespitosa, Sneyd's Coppice, Worcestershire,_England, W. B. Grove.
(From Myeologia.) _ ^•''
E. Spores stiffly obclavate-filiform, apices acute, bases truncated,
scarcely constricted at the septa, pycnophores elongate
P. festucae Sprague (Fig. 15,B)
EE. Spores similar to above but somewhat constricted at septa,
pycnophores broadened or obscure
P. phalaridis (Trail) Sprague (Fig. 15,A)
It should be added that the recently described Phaeoseptoria multi-

septata Frandsen (1943) is virtually identical with Hendersonia cras-
tophila Sacc. as we understand the species (Sprague, 1943,6; 1946,6).
References: Spegazzini (1908); Sprague (1943,&; 1948,a,c).
Phleospora Wallr.
Some consider Phleospora to be an unnecessary genus, more or less
intermediate between Cylindrosporium and Septoria. We follow Grove
(1935) in recognizing it as a sometimes useful genus characterized by
having pycnidia with relatively thin apices which soon burst or eva-
nesce to release a mass of comparatively coarse spores. There are three
known species of fungi on grasses in the United States which should be
placed in this genus.
Phleospora graminearum Sprague and Hardison—Leaf Spot

Spots variable, elongate, brown, paler in center, margins sometimes
yellow or buff-colored; pycnidia subgregarious, at first immersed, non-
ostiolate, brown, later erumpent, prominent with small ostiole, sub-
globose, 90-160 jx, pale brown, upper layer finally breaking away", con-
text transluceiit and golden-colored; pycnophores blunt, short, 3-5 /*
wide, hyaline, pycnospores yellow-hyaline, obclavate, tapering to a
blunt point, base rotmded or blunt, variable, 30-55 X 3.3-5.6 jm, 1- to
6-septate (Fig. 16,4)...
FIG. 16.—Pycnospores of: A, Phleospora graminearum on Elymus canadensis

(type); B, P. muhlenbergiae (type).
O n : Agropyron repens (L.) Beauv., Mich, (typa host)—Elymus canadewis
L., Mich. - I
This species is actively parasitic (Fig, 17) and yf some local im-
portance. I t is surprising t h a t it has not been foundl elsewhere. .
I N
R a n g e : United States. '
R e f e r e n c e : Hardison and Sprague (1943).
P h l e o s p o r a i d a h o e n s i s Sprague—Leaf and Pedicel Blight

Spots black t o d a r k brown, diffuse, pycnidia immersed, subglobose
to cup-shaped, brown-black, gregarious, wall thin, somewhat variable
(12 fi or less), p a r e n c h y m a t o u s , finally ostiolate or opening a t the top
through stomata, 90-150 /J.; spore horns short b u t numerous, hyaline
or faintly t i n t e d ; pycnospores stoutly club-shaped, t o somewhat cylin-
drical, somewhat to distinctly curved, apices tapering, sometimes
pointed, bases blunt but the spores narrowed/toward t h e ultimate base,
rarely 1-septate, usually aseptate, contents'hyaline with coarse inclu-
sions, 44-53 X 4.6-5.6 ;u.
O n : Festuca idahoensis Elmer, Ida.
Paraffin-embedded material of the fruiting bodies of this fungus •
show that it is not in Septogloeum but in Phleospora. I t shows some
relationship to Septoria infuscans but the Phleospora type of pycnidia
with their coarse spores (Fig. 18) seems to militate against this spgcies.
P. idahoensis seems to be localized, as is the present known range
of the previously discussed species of Phleospora. However, there are
many species of fungi on grasses which are, relatîvely common and
some which are abundant but which have not been collected by more
than two or three persons. It should be adddd that a search for this
fungus in 1948 in the vicinity of Trude, Idaho, was unsuccessful al-
though Fischer, who collected it originally, helped in the search. It
therefore remains somewliat of a mystery, ^-
Range: United States. • •
Reference: Sprague (1946,e; 1948,a).
Phleospora muhlenbergiae Sprague and SolKeim in Solheim—

Leaf Mottle
Spots straw-colored without borders, pycnidia scattered to subgre-
garious and numerous enough to give a mottled appearance to the pale
host tissue, pycnidia black, subcarbonaceous, atjifest" immersed, without
ostiole, later subglobose or bluntly lenticular, 95-245 X 65-121 /*, prom-
inent, ostiole irregular or the apex of the pycnidia eroding away to
disclose the hyaline interior, 45-170 X 38-104 /*; pycnospores pbclavate,
curved, apex tapering to a softly blunted pointy base rounded, tapering.
FIG. 17.—Leafspots caused by Phleospora graminearum on Elymui canadensis

(type).
finally blunted, chlorine-yellow, context homogeneous, 1- to 2-septate,
21-45 X 2.6-4 /x, mostly 28-34 X 3.0-3.2 ^,
On: Muhlenbergia arizonica Scribn., Ariz,
The spores of this species were defi-
nitely smaller (Fig. 16,B,) thaiiN*^^hose of
P. graminearum (Fig. 16,A} and P. ida-
hoensis (Fig. 18) and are mostly 1-sep-
tate. The species did not for*n definite
spots as did the other two species of
Phleospora. The leaves which were
straw-colored formed a contrasting
background for the black pycnidia. The
ostioles consist of fractures in the apex
of the pycnidia, probably caused by
internal pressure or enzymatic action
of the spores. The scarcely ostiolate
FIG. 18.—Pyenospores of
Phleospora idahoensis from pycnidia with the obclavate yellowish
Festuca idahoensis (type). spores are typical for the genus as we
(From Mycologia.) interpret it.
Range: United States and possibly Mexico.
References: Solheim (1949); Sprague (1950,a).
Phoma sp.—^Black Stem j

Culms black as though burned, areas involved are sometimes sev-
eral inches long, covered with superficial thin layers of firm black
stromatic mycelia in which small, subsuperficial/pycnidia may occur,
sometimes absent, pycnidia carbonaceous, averaging about 50-60 (-90)
;«. diam., but variable, containing hyaline rnicrospores, 4-6 X 0.5-0.8 fi
diam. These microspores are spermatia-like. Some of the pycnidia ap-
pear to contain ascogenous initials. i
On: Festuca elatior L., Alaska, Ida.—F. idahoemis Elmer, Ida^-Mont.—F. oc~
ddentalis Hook., Mont.—F. rubra L., Alaska.
This fungus appeared in several places in northern Idaho, adjacent
Montana, and in southern Alaska in 1948. Several specimens were re-
tained but considerable quantities of it were thrown out earlier in the
year before the pycnidial material was seen. Specimens collected by
Lefebvre in Alaska in 1948 were seen and briefly examined. Study of
this Alaskan material might disclose perithecial material. Because most
of the specimens which have been studied have not beeri mature (or at
least they so appear), it is considered better, to-call this Phoma, sp. and
await opportunity to observe better specimens.
Other-species of Phoma are sometimes reported on grasses and
cereals. Most of the reports refer to P. herbarum or comparable sapro-
phytes. O'Gara (1915) reported an undetermined species of Phoma on
Agropyron smithii Rydb. from Utah. On the basis of the vague dis-
cussion it is impossible to identify this fungus satisfactorily. The
writer suspects that it may be Seleno-phoma obtusa Sprague and A. G.
Johnson.
Phoma gyneriicola Cke. and Massee is listed on Cortaderia selloana
(Schult.) Aschers and Graebn. from California (Weiss, 1945). I t is
probably saprophytic. P. lineata Sacc. {Coniothyrium lineare v.
Thuem.) is also reported on this host from the same state (Weiss,
1945). Pampas grass, from our observations, is highly resistant to leaf-
invading fungi in this country.
Range: United States and Alaska.
Reference: Sprague (1949,a; 1950,6).
Phyllosticta Pers. ex Fr.

A great deal of the fascimile material of Phyllosticta found on
grasses is developing spores of Ascochyta, Stagonospora, Septoria,
Hendersonia and other genera. However, some forms appear to be
fixed in the nonseptate, hyaline phase, at least until just before ger-
mination. There are eleven such species which are .recognized in' this
text. One of these, P. sorghina Sacc, has a growing list of synonyms.
,^ The species may be distinguished by the key presented here.
KEY TO SPECIES Oy PHYLLOSTICTA
A. Pycnidia in spots with pale centers, margins of spots usually darker

B. Spots small, white, spores 11-15 X 2.5-4.5 i^
P. rogleri Sprague
BB. Spots irregular, pale to discolored, with red to purple borders, spores
4-7 X 2-3 /*
P. sorghina Sacc.
BBB. Spots with brown borders, spores 3-4 X 2-2.5 IJ,
P. glumarum (Ell. and Tr.) Miy.
AA. Pycnidia in gray or vague spots
B. Spots gray, elongate, spores bacillar, 2.3-4.6 X 1-1-4 j«
P. owensii Sprague (on Panicum the spots were
variable, sometimes vague or brown)
BB. Spots vague
C. On dead leaves, spores bacillar, 5-9 X 1-1.6 ii
P. anthoxella Sprague
CC. Spores ovate, 4.8-9.6,X 3:6 /J.
P. panici Young
CCC. Spores 3-4 X 0:4-0.7 /^
P. minutaspora Sprague
BBB. Spots cream-colored to pale yellow, emarginate, spores oval to
elliptic, 6.5-8.0 X 3-4 t^
P. avenophila Tehon and Daniels
BBBB. Spots brown, sometimes paler later ' !
• C. Spots brown for long periods, spores capsular, guttulate, ll-l5„X
3.6-4.5 n I • '
P. healdii Sprague
CC. Spots fading later
D. Spores fusoid, 7.5-10 X 3-4 ii
P. miyakei Sydow I
DD. Spores cylindrical-elliptical, 6.0-7.4 X 1.6-2.8 /x
P. helenae Sprague
Phyllosticta anthoxella Sprague—Leaf Mold

Spots vague or without definition, pycnidia scattered but with a
tendency to be in rows parallel to the leaf veins, erumpent, black
minute, 35-50 /A, ostiole with brown marginal cells, 10-12 "X 4-5 /*,
pycnospores bacillar-shaped, hyaline with 2-3 small guttulae in most
spores, aseptate, exuded in long chains j n water mounts, 5-9 X 1-1.6 [i.
f
On: Anthoxanthum odoratum L., Oreg.

/
P. anthoxella (Fig. 19,A) occurs 0n dead basal leaves associated
with Colletotrichum graminicola (Ces.) Wils. and Titae sp. This spe-
0
^ " ^
FIG. 19.—Pycnospores'of Phyllosticta spp.: A, P. anthoxellaon-Anthoxanthum

odoratum, Triangle Lake, Oregon (type); B, P. sorghina on Setaria viridis, Lang-
don, N. Dak.; C, P. rogleri on Digitaria sanguinalis, Albia, lowa-(type); D, P.
healdii on Panicum huachucae, Valentine, Nebr. (type); E, P. m.inutaspora on
Muhlenbergia filijormis, Idaho (type); F, P. helenae on Glyceria occidentalis,
Priest Lake, Idaho (type).
/
cies may be a saprophyte. I t has never been reported since the type
specimen was collected at Triangle Lake, Oregon, in May, 1938
(O.S.C. 264).
Reference: Sprague (1941,6).
FUNGI IMPERPECTI—SPHAEROPSIDALES 193
Phyllosticta avenophila Tehon and Daniels—Leaf Spot

Spots diaphyllous, very extensive, one half cm. to several cm. long,
half as wide or fully as wide as the leaf, cream-colored to pale yellow,
unlimited, emarginate. Pycnidia numerous, on the sides of the veins,
in closely clustered rows which are widely scattered over the spot,
spherical to slightly applanate, somewhat rostrate, occupying the
mesophyll, protruding epiphyllously, darkly membranaceous to carbo-
naceous, 70-100 /* diam., ostiole round, 10-15 jw, wide; rostrum 15-20 ju,
high, 12-20 /x broad, completely erumpent. Spores hyaline, continuous,
mostly appearing biguttulate, oval to elliptic, 6.5-8 X 3-4 fi.
On: Avena sativa L., 111., Miss.
The writer has identified a leaf spot on oats sent from Mississippi
as this species. If the oats fungus is an autonomous species it is either
very scarce or, more likely, so obscure as to pass unnoticed by most
observers.
Reference: Tehon and Daniels (1927, pp. 118-119).
Phyllosticta glumarum (.Ell. and Tracy) Miyake—•

Brown-Bordered Leaf Spot and Glume Spot
Phoma glumarum Ell. and Tracy
Pycnidia erumpent-superficial, black, 90-120 fi, ostiolate; pycno-
spores ellipsoid, nonseptate, hyaline to faintly smoky, 3-4 X 2.0-2.5 /i.
On: Oryza sativa L., Ark.-, La., Miss., Tex.—Uniola latifolia Michx., Miss.—
U. paniculata L., Miss.
This was collected on rice at Starkville, Mississippi, by Tracy.
Miyake (1910) transferred it to Phyllosticta. The literature is some-
what confused because Phyllosticta glumarum Sacc. was described in
1916 (Saccardo, 1916) which was six years after the paper by Miyake.
The spores of the 1916 species were 4.8-6 X 1.8-2.0 /x, and were said
to differ from Ellisând Tracy's species in the longer, strictly hyaline
spores and the pale bordered spots. Considering the number of species
of Phyllosticta or Phoma described on rice it would be surprising if
Saccardo's species was not a synonym of another species. It appears
to be distinct from the earlier named Phyllosticta glumarum and prob-
ably from P. miurai Miyake (1910) which has spores 3-4 X 1-1-5 i^.
P. miyakei Sydow (P. japonica Miyake) has larger spores, 7.5-10 X
3-4 ij.. Ling (1948) reported P. oryzae Hari from China. P . oryzaecola
Hara has spores 5-7 X 3-4 ^ (Hara, 1930, p. 158). P. oryzae is said to
be a synonym of P. oryzaecola (Hara, 1930) and to have a perithecial
stage called Phaeospora oryzae Miyake with ascospores 16-23 X 4-5 /^
(Hara, 1930, p. 158).
j j
There is also Phoma oryzae Cooke and MaSsee with ovoid spores,
3 X 2 / * , collected on rice straw at Calcutta. P. riecatrix v. Thuem. was
reported on culms and necrotic sheaths of rice from northern Italy
(cf. Sylloge Fungorum V. 10, p. 185). It had hyaline, eguttulate, ovate
to ellipsoid spores 10-12 X 6-8 /*. ]\
Range: United States, China, Japan, Philippines, Uganda, and other
tropical regions.
References: Ellis and Tracy (1888); Ideta (1926, V. 2, p. 642); Miyake
(1910).
Phyllosticta healdii Sprague—^Leaf Spot

Spots covering most of the surface of infected leaves, diffuse, pale
brown, of one uniform color without margin; pycnidia black, more or
less grouped in lines, erumpent to nearly superficial, ellipsoidal, mostly
definitely longer than broad, 111-156 X-80-115 fi, eventually ostiolate,
ostiole small, spores in mounts usually .need to be crushed from the
pycnidia, less often emerging through /the ostiole; pycnospores hya-
line, capsular to bacillar-shaped, 1-septate but many spores have slight
constriction or contraction at the centers; most spores with a prominent
inclusion at each end with sometimes one or more smaller inclusions
elsewhere, 11-15 X 3.6-4.5 /x.
On: Panicum huachucae Ashe, Nebr.
This fungus is characterized by the numerous black, linearly ar-
ranged pycnidia on diffuse brown lesions and the short-cylindric,
"blocky" spores (Fig. 19,D) with their prominent guttulae at each end.
The fungus is perennial and is found on current season living foliage
and on the dead growth from last year. This fungus has much larger
spores than P. panici or P. sorghina. The spores of P. healdii are some-
what different in shape from those of P. rdgleri (Fig. 19,C).
Greene (1942) reported abundant ostiolate pycnidia on Panicum
scribnerianum Nash containing bacillary spores. This ^spedes could
have been the microspore stage of a species of Septoria.
Reference: Sprague (1948,o).
Phyllosticta helenae Sprague—Spot

Spots brown, fading to straw-color, pycnidia elliptical in outline,
between the veins, much longer than wide, flattened, erumpent, fragile,
parenchymatousj light golden brown, ostiol^feT" 180-230 X 88-100 /i
diam.; pycnospores exuded in cirrhi, cylindric'al-elliptical, both ends
rounded uniformly, with small guttulae at each end, contents hyaline,
6.0-7.4 X 1-6-2.8 /z.
FUNGI IMPERPECTI—SPHAEROPSIDALES 195
On: Glyceria occidentalis (Piper) J. C. Nels., Ida.
This fungus is associated with other fungi on dried basal leaves of
this grass growing in wet beach sand or shallow water in the Priest
Lake area of Idaho. The type is A.S. 20,507. The fungus has relatively
long, narrow, thin-walled pycnidia and the spores are cylindrical but
also somewhat elliptical in outline (Fig, 19,F) with a small oil drop
at each end. The fungus is associated with a yellow spored aseptate
fungus with spores up to 12 X 3 ;a long borne in brown globose pyc-
nidia. This immature material appears to be a phase of an associated
species of Hendersonia. In addition microspores of Septoria avenae
Frank occur on the same leaves. These spores are 4-5 X 1.1-1.5 jx.
Range: Idaho in United States.
Phyllosticta minutaspora Sprague—-Yellow Leaf

Infected leaves pale yellow, dying, pycnidia in black narrow lines,
gregarious, somewhat or thinly surrounded by stromatic tissue, small,
black, irregularly ostiolate, carbonaceous, globose, 40-80 /j. diam. Pyc-
nospores bacteria-like, hyaline, 3-4 X 0.5 /x. (microspores) (Fig. 19,E).
On: Muhlenhergia filiformis (Thurb.) Rydb., Ida.
The lower leaves of current season foliage turn yellow after inva-
sion. The pycnidia are somewhat embedded in fungus tissue, which
gives a black appearance to the lesions. However, this tissue is inter-
rupted and more or less Phoma-like.
Phyllosticta miyakei Sydow—On Rice Leaves and Glumes

P. japonica Miyake
Spots effuse, brown, then becoming bleached; pycnidia globose to
ellipsoid, immersed, black, 75-100 /j. diam., 55-90 /j. high; sporophores
small; pycnpspores fusoid, biguttulate, 7.5-10 X 3-4 û,.
On: Oryza sativa L., S. Car., Tex.
Fields reported this fungus from the United States (19451 • As men-
tioned under P. glumarum, above, this species has larger spores than
the latter. Ideta (1926) indicated that Sydow created a new name for
the fungus which was first described as P. japonica Miyake (nee v.
Thuem.; nee Fautr.).
Range: United States, Japan.
References: Fields (1945); Miyake (1910); Saccardo (Sylloge v. 22, p
864).
'i
Phyllosticta owensii Sprague—Gray' Leaf Streak
Leaf spots elongate, gray, pycnidia sparse^ globose, superficially
erumpent, black, tardily ostiolate, 70-100 /J. diam.; pyenospores numer-
ous, bacillar, nonseptate, hyaline, 2.3-4.6 X l.Q-!l.4 ix,.
O n : Dactylis glomerata L., Oreg.—Panicum capillare L., Wash.
The type is a Phoma-like saprophyte or weak parasite on the dead,
living, or salt-spray-injured leaves near Waldport, Oregon. It is asso-
ciated with Scolecotrichum graminis Fckl. A collection in a comparable
area at Long Beach, Washington (A.S. 20,510), on stunted plants of
Panicum capillare has gray to vague spots on the leaf tips. The tiny
spores are identical with the type on Dactylis.
Reference: Sprague (1941,6; 1950,a).
Phyllosticta panici E. ôung—Leaf Spot

Spots indefinite, diffuse, pycnidia few, often in clusters; epiphyllous,
dark brown to black, spherical, 72^144 /* diam.; mycelium a brown
mass of septate hyphae; pyenospores hyaline, ovate, 4.8-9.6 ><^ 3.6 ju,.
On: Panicum maximum Jacq., Fla.—P. virgatum L., Okla.
Weiss (1945) questioned whether this was distinct from P. sorghina'
and according to Bourne (1934) it is a synonym of the latter. The ma-
terial from Oklahoma which we assigned to P. panici was possibly dif-
ferent from P. sorghina. ,
Range: United States, Puerto Rico. j
References: Bourne (1934); Weiss (1945) ;| Young, E. (1915).
. Phyllosticta rogleri Sprague—White Leaf Spot

Leaf spots small, elongate, white, pycnidia scattered,~"yellow brown,
globose, erumpent, thin-walled, composed of polyhedral cells, ostiolate
(6-12 IX.), small, 50-100 ix; pyenospores ellipsoid, nonseptate, hyaline,
3-6 guttulate, 11-15 X 2.5-4.5 /^ (Fig. 19,C)\
On: Digitaria sanguinalis (L.) Scop., Iowa. /
This species causes spots on living leaves. I t is known only from
the type area, near Albia, Iowa. Weiss, however, also lists Idaho (1945)
which, perhaps, is a typographical error.
Phyllosticta sorghina Sacc.—Leaf Spot

Lesions on living leaves irregular, mostly subcircular, 2-10 mm.
diam., pale straw to isabelline with narrow, purple borders; pycnidia
45-150 [1. diam., black, subglobose, walls fuliginous, strongly erumpent,
spores nonsept.ate, hyaline, eguttulate, exuding in cirrhi when placed
in water, ellipsoid with rounded ends, 3.5-7 X 1.7-3.4 n (Fig. 19,B).
On: Panicum dichotomiflorum Michx., Fla..—Saccharum officinarum L., Fla.,
Hawaii, La.—Setaria verticillata (L.) Beauv., Colo.—<S. viridis (L.) Beauv.,
Manit., N. Dak.—Sorghum halepense (L.) Pers., Fla., Tex.—S. vulgare
Pars., Ark., Fla., Ga., 111., Kans., La., Miss., Mo., Okla., Tex., Va.—S. vid-^
gave var. sudanense (Piper) Hitchc, Md., Va.—Tricholaena rosae Nees,
Wash., in plots.
The synonymy of P. sorghina Sacc. as determined to this date is
as follows:
Name Host Locale Citation
Phyllosticta sacchari Speg.. . -. Saccharum Argentina Spegazzini (1896)
P. setariae Ferr Setaria Italy Ferraris (1902)
P. glumarum-sorghi P. Henn.. Sorghum Congo Hennings (1907)
P. glumdrum-setariae P. Henn. Setaria Congo Ibid.
P. phari Speg Pharus Argentina Spegazzini (1911)
P. penicillariae Speg Pennisetum Senegal Spegazzini (1915)
P. hawaiiensis Caum Saccharum Hawaii (1919); J. P.
Caum (1919):
Martin (1938)
Phoma insidiosa Tassi Sorghum Abyssinia Tassi (1898)
The fungus occurs on a wide range of warm-temperature hosts in

many regions and for that reason its synonymy is considerable. It is
common on Setaria viridis in North Dakota and adjacent Manitoba
and is well known on sugar cane and sorghum in warm parts of the
world (cf. Fawcett, 1922).
Unamuno (1931) described P. halepensis on dry leaves of Sorghum
halepense from Spain: The amphigenous, punctiform, globose or ellip-
soid, black, pycnidia measure 62.8-88.5 X 68.5-74.5 /J. diam. and the
spores are hyaline, oblong to ellipsoid, Macrophoma-like, straight or
curved, 2-3-septate,. 8;5-12 X 3.2-3.8 ,i.
Another species of Phyllosticta on sugar cane, similar to, if not
identical with, P. saccharicola P. Henn., is the pycnidial stage of Lepto-
sphaeria sacchari V. Breda de Haan. The latter is a common secondary
spot associated with Helminthosporium sacchari (V. Breda de Haan)
Butler, the cause of the eye spot (ring spot) in cane leaves (Bourne,
1934). P. saccharicola P. Henn. was described with spores 10-30 X
3.0-3.5 /*. Bourne, incidentally, considered P. sorghina a saprophyte
associated with H. ocellum Paris.
Range: United States, Abyssinia, Argentina, Brazil, Canada, Congo, Hawaii,
India, Italy, Portugal, Puerto Rico, Senegal, South America, Tanganyika,
U.S.S'.R.
i\
References: Bourne (1934); Ferraris (1902); Hennings (1907); Koch and
Rumbold (1921); Martin (1938); Parris (1940); Rumbold and TisdaJe
(1921,a,6); Spegazzini (1896, 1911, 1915)^; Sprague (1941,6); Stevenson
(1926); Tassi (1898).
Phyllosticta spp.—Undetermined
Weiss (1944, 1945) listed a number of collections of Phyllosticta
undetermined as to species in his check list. W e also have a number
of fragments which have not been placed in definite species. T h e v a r i -
ous hosts, locales, and citations are as_follows:
Host Locale Citation

Andropogon furcatus Muhl North Dakota Sprague, notes
Bromus sp. {secalinus) Georgia Weiss (1944)
Calamovilfa longifolia (Hook.) Scribn North Dakota Sprague, notes
Cenchrus bifloriis (Hook.) Scribn Florida Weiss (1945)
C. ince-tus M. A. Curtis ., .'. Wisconsin Greene (1948)
Cortaderia selloana (Schult.) Asohers. and Graebh. Kentucky Weiss (1945)
Digitaria violascens Lk : . . Kentucky Ibid.
Festuca subulata Trin ,. . . . Idaho, Oregon Sprague, notes
Paspalum dilatatum Poir .•. . . . South Carolina Weiss (1945)
Schizachne purpurascens (Torr.) Swallen. .y. Minnesota Sprague, notes
Zea mays L Florida, Texas Weiss (1945)
Among those collections reported b y the writer t h e only material

which has been seen in any q u a n t i t y , other t h a n a small fragment, is ,
the fungus on Festuca subulata which has been collected several times
in the woods of western Oregon and along the south fork of the Clear-
water River, I d a h o . Greene (1948) reported spores 5-8 X 3 /u,, ellipsoid
or short-cylindric, straight or slightly curved on' Cenchrus.
Pyrenochaeta terrestris (Hansen) I Gorenz, Walker, and

Larson—Pink Root
Producing a pink, slow rot of rootlets and mojie extensive sapro-
phytic decay of m a t u r e or dead roots in later season, forming a slow-
growing, mounded, gray growth with rosy or magenta colors in the
medium (potato dextrose a g a r ) . Pycnidia subglobose, ostiolate, papil-
late, dark brown to black, with scattered Setae about the ostiole, setae
light to d a r k brown, 1- to 5-septate, 8-120 ix., pycnidia carbonaceous,.
170-350 fi, sometimes gregarious. Pycnospores continuous, hyaline, ob-
long-ovoid, 3.7-5.8 X 1.8-2.3 /x, biguttulate, sessile in the pycnidium,
escaping as a gelatinous mass through ruptures, rarely as a gelatinous
cirrus through the ostiole. Mycelium septate, hyaline, guttulate, 1.0-
4.5 /x, frequently anastomosing. ~^^^
O n : Agropyron ciliare (Trin.) Franch, N. Dak.—A. cristatum (L.) Gaertn.,

Mont., N. Dak., S. Dak., Wyo.—A. desertorum (Fisch.) Schult., Mont.,
Nv Dak., S. Dak.,'Wyo.—A. elongatum (Host.) Beauv.,'N. Dak.—A.
intermedium (Host.) Beauv., N. Dak.-—A. michnoi Roshev, N. Dak.,
S. Dak.—A. mongolicum Keng, N. Dak.—A. pendulinum (Nevski) ined.
(fide Swallen), N. Dak.—A. repens (L.) Beauv., Minn., Nebr., N. Dak.,
S. Dak.—A. semicostatum (Steud.) Nees, N. Dak.—A. sibiricum (Willd.)
Beauv., N. Dak., Wyo.—A. smithii Rydb., Mont., Nebr., N. Dak., S. Dak.,
Wyo.—A. trachycaulum (Lk.) Malta, N. Dak.—Andropogon furcatus
Muhl., N. Dak.—A. hallii Hack., N. Dak.—Arthraxon hispidus var.
cryptatherus (Hack.) Honda,'N. Dak.—Avena sativa L., Calif., Minn.,
Mont., Nebr., N. Dak., S. Dak., Wyo.—Bouteloua curtipendula (Michx.)
Torr., N. Dak., S. Dak.—5. gracilis (H. B. K.) Lag., Mont., N. Dak.,
S. Dak.—Bromvs arvensis L., N. Dak.—B. carinatus Hook, and Am.,
N. Dak.—B. erectus Huds.^ N. Dak.—B. inermis Leyss, Nebr., N. Dak.,
S. Dak.—Calamovilfa longifolia (Hook.) Scribn., N. Dak.—Cenchriis
pauciflorus Benth., N. Dak.—Cynodon dactylon (L.) Pars., N. Dak.—
- Dactylis glomerata L., N. Dak.—Distichlis striata (Torr.) Rydb., S. Dak.
—Echinochloa crusgalli (L.) Beauv., N. Dak., S. Dak.—E. crusgalli var.
frumentacea (Roxb.) Wight, N. Dak.—Elymus canadensis L., N. Dak.—
E. dahuricus Turcz., N. Dak.—E. interruptus Buckl., N. Dak.—E. juncem
Fisch., N. Dak.—E. sibiricus L., N. Dak.—Festuca octofiora Walt., N.
Dak.—F. rubra L., N. Dak.—Hordeum brevisubulatum (Trin.) Lk., N.
Dak.—H. distichon L., Minn., Nebr., N. Dak., S. Dak.—H. vulgare L.,
Iowa., Minn., Mont., Nebr., N. Dak., S. Dak., Wyo.—Koeleria cristata
(L.) Pers., N. Dak.—Muhlenbergia squarrosa (Trin.) Rydb., N. Dak.—
Oryzopsis hymenoides (Roem. and Schult.) Rick., N. Dak., S. Dak.—
Panicum capillare L., Mont., N. Dak.—P. miliaceum L., N. Dak., S. Dak.
—P. virgatum L., N. Dak.—^Phleum alpinum L., N. Dak.—P. pratense
L., N. Dak., S. Dak.—Poa compressa L., N. Dak.—P. pratensis L., N.
Dak.—P. secunda Presl, S. Dak.—Saccharum officinarum L., La.—
Schedonnardus paniculatus (Nutt.) Trel., Mont., N. Dak.—Setaria italica
(L.) Beauv., Iowa, N. Dak.—S. lutescens (Weigel) F. T. Hubb, N. Dak.
-—S. viridis (L.) Beauv., N. Dak.—Sorghum vulgare Pers., Iowa, Nebr.,
N. Dak., S. Dak.—S. vulgare var. sudanense (Piper) Hitchc, N. Dak.—
Stipa comata Trin. and Rupr., Mont.,,S. Dak.—S. viridula Trin., N. Dak.
—S. williamsii Scribn., N. Dak.—Triticum aestivum L., Mont., N. Dak.—
T. dicoccum Schrank, N. Dak.—T. durum Desf., N. Dak.—T. spelta L.,
N. Dak.—T. timopheevii Zhuk., N. Dak.—Zea mays L., Colo., N. Dak.,
S. Dak., Wise.
Hansen (1929)^ found that this species, the cause of pink root in
onions, was common in soil and he suggested that it was probably
parasitic, or at least present, on hosts other than onions. Kreutzer
(1941) showed that it could cause injury ,to cereals under., artificial
'conditions. It has since been determined that P. terrestris occurs on
many cereals and grass hosts in nature as had been indicated by
Kreutzer's work (Carvajal, 1945; Sprague in Fischer, et a l , 1942;
Johann, 1943; Sprague, 1944,a; 1946,e). The array of susceptible hosts
is more impressive, however, than the actual damage done to them by
P. terrestris. It causes a very mild seed rot and slight root rot (Sprague,
1944,a) which is accompanied by various pink, rosy, or purple dis-
colorations of the smaller roots or crown parts adjacent to the seed in
young plants. The pink color seen on most roots of 1 maturing cereals,
however, is due to the saprophytic invasion of Fusarium oxysporum
(Schl.) em. Snyder and Hans.
Gorenz, Walker, and Larson found that the pycnidia of the fungus
produced setae about the ostiole. They therefore transferred the fungus
to Pyrenochaeta (1948). |
A few isolates of this species readily fruit on potato dextrose agar
at room temperature, or in the refrigerator, but most of the isolates
remain sterile under these conditions. Fruiting is obtained with more
ease on sterilized plant parts under common storage conditions.
References: Carvajal (1945); Gorenz, Walker, and Larson (1948); Hansen
(1929); Johann (1943); Kreutzer (1941); Sprague (1944,a; 1946,e).
Scaphidium boutelouae Clements—Narrow Brown Spot

Spots brown to gray, linear, emarginate or sometimes brown-bor-
dered with limited fawn-colored centers,'1-4 X 0.4-1.0 mm. diam.
Pycnidia small, more or less gregarious, black, carbonaceous, innate-
erumpent, linear to subglobose or subanastomosed, opening irregu-
larly or somewhat hysteroid, 60-100 /x, diam., sporophores simple, hya-
line 10 X 2 ja, spores yellow-brown or subhyaline, 1-septate, curved
fusiform-obclavate, 19-30 X 2.8-5.3 [i., commonly 24-29 X 3.7-5.0 /i.
On: Bouteloua curtipendula (Michx.) Torr., Ariz., N. Mex.—B. hirsuta Lag.,
Nebr.
Material sent by W. G. Solheim from Santa Catalina Mts., Arizona
(Rocky Mtn. Herb. 2447), and Silver City, New M ^ i c o (Rocky Mtn.
Herb. 2390), have curved yellow-brown spores (Fig.
20) in poorly formed pyciiidia which answer the
description of Scaphidium.. We are not certain
whether Scaphidium belongs in the Exaipulaceae or
the Leptostromataceae but Clements placed- it in
the first mentioned. —
The material from New Mexico appears to be
the result of mild parasitic action. At least some of
the linear lesions have a defiiîte border and ap-
FIG. 20.—Pyc- parently developed while the leaf was still alive.
nospores of Sca- The. gray streaked areas which are the most com-
phidium boute- mon phase had apparently developed as the leaf
louae from Boute- was dying. The material from Arizona has the linear
loua curtipendula,
Ariz. brown-bordered lesions surrounded by'living green
tissue. ' '' ' '
References: Clements (1901); Saccardo (Sylloge, 1906, v. 18).
7[^^
-FUNGI IMPERFECTI—SPHAEROPSIDALES 201
Selenophoma Maire.
Lunospora Frandsen
This genus is characterized by spherical, compact pycnidia bear-
ing sickle-shaped (falcate) hyaline spores (Maire, 1906). Frandsen's
Lunospora (1943) was described during World War I I when communi-
cation with other workers was largely cut off. His genus does not
appear to be needed (Sprague and A. G. Johnson, 1940, 1945, 1947).
If Pseudoseptoria Speg. (1911, pp. 388-389, Fig. 51) is the same as
Selenophoma Maire, Spegazzini certainly misdescribed the fungus. He
discussed a fungus with superficial pycnidia.
A key to the known species of Selenophoma is given here.
KEY TO THE KNOWN SPECIES OP SELENOPHOMA ON GRAMINEAB
A. Spores acute tipped, boomerang, or arcuate ,

B. Spores arcuate, typically small, 10-6 X 1.1-1.8 /i
S. everhartii (Sacc. and Syd.) Sprague and
A. G. Johnson
BB. Spores lunate to boomerang-shaped, typically 18-30 X 2-4 /»
S. donacis (Pass.) Sprague and A. G. Johnson
BBB. Spores lunate to subarcuate, typically 12-24 X 1.5-3.0 /*
S. donacis var. stomaticola (Baueml.) Sprague
and A. G. Johnson
BBBB. Spores lunate, typically 17-25 X 2.0-3,2 M, on Bromus
S. bromigena (Sacc.) Sprague and A. G. John-
son
AA. Spores obtuse, subrenifprm-falcate
(S. obtusa Sprague and A. G. Johnson
Selenophoma bromigena (Sacc.) Sprague and A. G. Johnson—

Leaf Spot
Spots on leaves gray, circular to irregular with narrow brown bor-
ders, pycnidia spherical to subglobose, light golden brown, ostiolate,
subepidermal, 50-150ju. diam., pycnophores prominent, truncate to
flask-shaped, 3-5 X 2.5-3.6 /j.; pycnospores regularly lunate, rarely
boomerang-shaped, 17-25 X 2.0-3.2 fi (21-33 X 3.0-4.3 n in one collec-
tion on B. ciliatus from Climax, Colorado, A.S. 20,525).
On: Bromus carinatus Hook, and Am., Calif., N. Dak., Wash., Wyo.—
B. ciliatus L., Colo.—B. inermis Leyss, Alta., Ariz., B. C, Manit., Minn.,
Mont., Ida., Nebr., N. Dak., Sask., S. Dak., Utah, Wash., Wise, Wyo —
B. purgans L.j Colo.
The common host, smooth brome, is severely damaged by this
fungus which forms gray spots on the leaves in early spring. I t is espe-
cially prevalent in the Red River Valley in North Dakota and Minne-
sota but occurs westward to Pullman, Washington. I t is not uncommon
/I
on smooth brome growing in the Idaho prairies and we have collected
it in the vicinity of Logan, Utah, and as far south âs Grand Canyon,
Arizona (Sprague, Fischer, and Meiners, 1948).
This species of Selenophoma seems'to be largely confined to smooth
brome but during recent years it has become more common on B. cari-
natus than was the case in earlier years. Allison (1939;a) found that
Minnesota isolates did not attack any host except B. inermis. The re-
cent increase in host range may be associated with the development
of a new race or races. We found B. carinatus scatteringly infected to
as far south as Tioga Pass, California. The fungus was found on
FIG. 21.—Pycnospores of: A, Selenophoma donacis on Elymus flavescens,

Hanford, Wash.; B, S. bromigena on B. inermis, Craigmont, Idaho; C, S. obtusa
on Sitanion hystrix, Mt. Shasta, Calif.
B. ciliatus L. at Climax, Colorado, in August, 1948. The spores were

large as is noted in the description above.
S. bromigena differs from S. donacis somewhat vaguely in that the
spores of S. bromigena are regularly lunate (Fig. 21,B), sharp pointed
and rarely boomerang-shaped. It has somewhat/larger spores than
S. donacis var. stomaticola and, considering! all angles, the writer and
A. G. Johnson concluded that it should be left as an autonomous
species, at least until a great deal more study can be made with the
group. ^
Phlyctaena bromi 'Clements and Clements (Crypt. FT Coloraden-
sium) appears to be a saprophytic phase of S. bromigena. The'pycnidia
are subsuperficial and connected by coarse creeping mycelia. The
packet containing this material is undated -and the fungus so poorly
described as to have little status as a good species. Certainly there is
little reason to propose a new combination for this fungus. Since not-
ing Clements' material we have collected typical material of S. bromi-
gena on B. ciliatus in Colorado, as above noted.
Range: United States, Canada, Hungary (Krypt. ]5xs,_PaIatino No. 1465).
References: Allison (1939,a,6; 1945); Barley. ^9|1),; Saccardo (1915);
Sprague and A. G. Johnson (1940, 1945,1947,1950).
Selenophoma donacis (Pass.) Sprague and A. G. Johnson—

Eye Spot
Septoria donacis Pass. ^
S. oxyspora Penz. and Sacc.
Lunospora oxyspora (Penz. and Sacc.) Frandsen
Spots more or less elliptical, straw-colored with purple, red, or
brown borders (eye spots) which may fade later, on c«lms, sheaths,
and leaves; pycnidia erumpent, small to subprominent, brown, globose,
ostiolate, 40-150 ju, sometimes up to 145 ;u, diam., peridia of pycnidia
composed of coarse compacted polygonal cells which produce cuspidate
pycnophores, from the inner pycnidial wall, composed of hyaline bulb-
ous initials, walls 5-12.5 ^ thick, pycnophores 2.5-5 X 2-3 jn; pycno-
spores are stoutly falcate to boomerang-shaped, 18-35 X 2.0-4.5 ;«,.
On: Agropyron albicans Soribn. and Sm., N. Dak.—A. cristatum (L.)
Gaertn., Wyo.—A. dasystachyum (Hook.) Scribn., Oreg.—A. inerme
(Seribn. and Sm.) Rydb., Colo., Wash.-—A. pendxdinum (Nevski) Swallen,
N. Dak.—A. repens (L.) Beauv., N. Dak., Wash.—4. riparium Scribn.
and Sm., N. Dak., Wyo.—A. smithii Rydb., Ariz., Mont., Nebr., N. Dak.,
S. Dak.,.Wyo.—A. spicatum (Pursh) Scribn. and Sm., Colo., Nev., N. Dak.,
Utah, Wash., Wyo.—A. subsecundum (Lk.) Hitchc, Mont., Wyo.—A.
trachycaulum (Lk.) Malts, Mont., N.- Dak., Sask.,^ Wyo.—Arundo donax
L., Calif.—Avena sativa L., N. Dak.—Elymus aristatus Merr. (hybrid),
\Wash.—E. canadensis L., N. Dak.—E. condensatus Presl, Cahf., Mont.,
Nev., N. Dak., Oreg., Wash.—E. flavescens Scribn. and Sm., Wash.—
E. giganteus Vahl, Oreg., Wash.—E. junceus Fisch, N. Dak.—E. macounii
Vasey (hybrid), Sask.—E. villosus Aluhl., N. Dak.—E. virginicus L.,
N. Dak.—Panicum virgatum L., Kans., N. Dak., Wise.—P. wilcoxianum
Vasey, Nebr.—Phalaris arundinacea L., N. Dak., Wash.—Phragmites
communis Trin., IN. Dak.—Poa alpina L., Colo., Mont.—P. ampla Merr.,
Ida., Mont., Oreg., Wash., Wyo.—P. cenisia Allescher, Greenland—P. com-
pressa L., Cole, N. Dak.—P. cusickii Vasey, Oreg.—P. epilis Scribn.,
Calif., Wyo.^P. glauca Vahl, Greenland—P. gracillima Vasey, Oreg., Utah
—P. juncifolia Scribn., Oreg., Utah, Wash.—P. nevadensis Vasey, Oreg.,.
Wash.—P. palustris L., Colo.—Secale cereals L., Minn., N. Dak.—
Stipa Columbiana Macoun, Wash.—S. comata Trin. and Rupr., Mont.,
Nebr., N. Dak., S. Dak:,'Wyo.—S. lemmonii (Vasey) Scribn., Wash.—
S. richardsonii Lk., Wyo.—S. spartea Trin., N. Dak.—S. viridula Trin.,
Mont., N. Dak., S. Dak.—Triticum aestivum L., Ida., Wash.
This fungus is frequently called Septoria oxyspora, less often Sep-

toria donacis, an earlier name. While it was described on Arundo donax
L., a giant reed, it occurs also on Phalaris arundinacea in some abun-
dance and rarely on Phragmites in the area surveyed. The spores of
these hosts and on Elymus (Fig. 21,A) average around 20-21 X 3 /t
and are more or less boomerang-shaped. Spores of this species on
Panicum are narrower, one collection, for instance, being 21.5 X 2.2 y..
Perhaps this should be recognized as a distinct variety.
S. donacis on Stipa and Poa has stubbier spores than on Phalaris
and Arundo, averaging 18.3 X 2.8 p. on Stipa. That on Phragmites has
thicker spores, 16-18 X 3-4 /t but is typically boomerang-shaped
(Sprague and A. G. Johnson, 1947). Most of the spores in European
material from Phragmites are 1-septate &nd are usually called Sep-
toria curva Karsten (1887). Petrak (1940) plaped it in Selenophoma
curva (Karst.) Petrak. Frandsen called it Lunospora curva (1943).
Allescher (1901, p. 916) called it Rhahdospora curva, while Sprague
and A. G. Johnson considered it an overwintered 1-septate phase of
S.donacis (1947). Like Migula (1921) and Diedicke (19],5) these men
also found that current season material was nonsepta,te.
FIG. 22.—Pycnospores of Selenophoma donacis on: A, Secale cereale, Vem-

dale, Minn.; B, Triticum aestivum, from leaf fragments intercepted in bulk
wheat at San Pedro, Calif., originating in Australia in 1944; C, on wheat, Mos-
cow Mts., Idaho.
Most of the lesions caused by S. donacis are straw-colored, with

tawny, fuscous, or lilac-tinted borders, and the pycnidiaâre small and
relatively obscure. However, the spots often fade, leaving the pycnidia
in unspotted tissue on the culms, sheaths, and Reaves.
S. donacis on rye {Secale cereale) also needs study as soon as viable
material is available. The pycnidia are prominent but not large, 95-111
X 72-85 [x., and the spores are 12-18 X 2-3.5 /x (Fig. 22,A), mean size
14.5 X 2.4 fi. in one collection, 16 X 2.7 /x in another. This fungus has
been found in an area' north of Jamestown, North Dakota, a,nd also in
Minnesota. The symptoms may be confused with those caused by bac-
terial leaf spot and therefore the disease on rye may be more wide-
spread than is known at the present. It has no relation to Septoria
falcispora DeraidoYa (1926) (nonBubak).
S. donacis on wheat was recognized for the' first time in the United
States in the vicinity of Pullman, Washington, in 1948 on winter
wheat {Triticum aestivum) (Fig. 22,C). It had been collected at Pull-
man in 1915 but not determined. When it first appeared in mid-March,
1948, the spots were emarginate, pinkish buff and irregularly elliptical
(Fig. 23). In a later development in early JuflCthe more or less circu-
lar spots were narrowly margined with a darker brown or fuscous than
the ashy center of the spots. In the younger lesions of those seen in
March, the centers, of the lesions were umber color froni the aggre-
•;m?^-

gated pycnidia. When viewed with a hand lens the pycnidia seemed
to be grouped in an elliptical area with the surrounding pinkish-buff
portion abruptly differentiated from the living green leaves. In later
material the fungus could be distinguished from Septoria tritici by the
shape of the lesions, those of S. tritici being somewhat elongated, those
of Selenophoma more or less circular. The spores were lunate, 16-21 X
FIG. 23.—Leafspots caused by Selenophoma donacis on wheat (Golden vari-

ety) near Moscow Mts., Idaho.
2.2-3.5 /A. The fungus produces a colony on potato dextrose agar which
is soon reddish brown to terra cotta, is convoluted with irregular mar-
gins, and produces terra cotta pigment in the medium. This fungus does
not resemble the common races at Pullman and the source of this
sudden appearance is not known. Cross inoculation trials in the green-
house were disappointing in that only scattered, sterile spots formed
on wheat and Poa pratensis and nothing on smooth brome, mountain
brome {Bromus carinatus), Dactylis, oats, barley, and Poa compressa
when all were inoculated with abundant fresh material from wheat and
incubated in the greenhouse. The time was late in the spring, condi-
tions were not favorable, and opportunity for further work was not
available.
S. donacis on wheat was found from the (foothills of the Thatuna
Hills near Moscow, Idaho, west to .Central Bridge, Washington, and
north to Rosalia, Washington. It was serioiis enough on the flag leaf
of wheat in the boot to draw the attention of Orville Vogel in his
varietal trials at Pullman, Washington .(U.S.DJA. and W.S.C. cooper-
ating) . Notes taken by the writer showed wide .range .of susceptibility
to the fungus. Some varieties including Rex land Orfed were very
susceptible, while others such as Comanche, Kharkov, and Hymar x
Elgin* (F4 composite) were highly resistant (Sprague, 1949,c). In the
preliminary report we placed this under S. donacis var. stomaticola but
in a later recent revision of the entire Selenophoma collections we'de-
cided that it should be placed in the species proper.
On the basis of single spore cultures there appear to be several dis-
tinct strains, groups, or races of S. donacis, besides those on wheat and
rye. The common race on Arundo and Phalaris (see also Lunqspora
baldingerae Frandsen) forms a Congo pink to testaceous rose mycelia.
A race on Panicum virgatum has a similar appearance in culture, but
the spores are recognizable by their narrower width. The race on
E. fiavescens which was collected in the sand hills south of Hanford,
Washington, before the atomic age, produced an ivory-colored growth
which was very different in macroscopic appearance from the Congo
pink of the other isolates. No isolations were made from the other
hosts. Weiss (1944) listed S. donacis on bamboo from California. This
may have been Arundo which is a common host.
Range: United States, Czechoslovakia, Denmark, Finland, France, Italy,

Latvia, Poland, Spain.
References: Diedicke (1915); Frandsen (1943); Grove (1935); Karsten
(1887); Migula (1921); Passerini (1879);, Penzi'g (1884); Petrak (1929,
1940); Sprague (1949,c); Sprague and A.I G. Jbhnson (1940, 1945, 1947,
1950). I
Selenophoma donacis var. stomaticola (Baueml.)

Sprague a,nd A. G. Johnson—Eye Spot ^'"~^-~,^
Phyllosticta stomaticola Baueml. - " '
Septoria culmiflda Lind ~
Spots on leaves, sheaths, and culms generally elliptical, white to-
tawny, with purple, red, or lavender borders, later fading] pycnidia
small, 40-150 X 40-110 fi, golden brown to b/own or sometimes black,
globose, sometimes somewhat longer than broad especially in winter
material,, ostiolate, erumpent, composed of coarse polyhedral cells;
pycnophores cuspidate, relatively prominent, arising irom globose or
somewhat cubical hyaline initials; spores faJcate, septate, variable,
iO-20 X 1-3 /x, less often as much as 25 ix longr ''
On-:,[Ârrhenatherum elatius (L.) Beauv., Ida., Mont.—Calamovilfa longi-
folia (Hook.) Scribn., N. Dak., S, Dak.—Dactylis glomerata L., Ida., Oreg.,
Wash.—Danthonia californica BoL, Oreg.—D. spicata (L.) Beauv., N. Y.—
Elymus caput-medusae L., Wash.—Festuca elatior L., Calif.—F. ida-
hoensis Elmer, Wash.—F. kingii (S. Wats.) Cassidy, Mont., Utah, Wyo.—
F. octoflora Walt., Nebr.—F. ovina L., Wash.—Hordeum brevisubulatum
(Trin.) Lk., N. Dak.-—H. vulgare L., Oreg., Sask.—Koeleria cristata (L.)
Pers., Ida., Mont., N. Dak., S. Dak., Wash., Wyo.—Melica harfordii Bol.,
Calif.—Oryzopsis hymenoides (Roem. and Schult.) Rick., Cahf., Mont.,
N. Dak., Oreg., Utah, Wyo.—Phleum pratense L., Alaska, Colo., IST. Dak.,
Sask., Wash.—Poa cenisia AUesch., Greenland—P. nervosa (Hook.)
Vasey, Mont., Wyo.—P. pratensis L., Colo., Mont., N. Dak., Wash., Wyo.
—P. secunda Presl, Mont., Nev., N. Dak., Oreg., S. Dak., Wash., Wyo.—
Puccinellia nuttalliana (Schult.) Hitchc, N. Dak., S. Dak.—Sitanion hys--
trix (Nutt.) J. G. Sm., N. Mex., Oreg., Wash.—Sporobolus asper (Michx.)
Kunth, Iowa, Mont., N. Dak.—S. clandestinus (Spreng) Hitchc, Okla.—
S. cryptandrus (Torr.) A. Gray, Ida., N. Dak., S. Dak., Wyo.—S. giganteus
Nash, Ariz.—S. heterolepis A. Gray, Wise.—Stipa lemmonii (Vasey)
Scribn., Wash.
This complex, which some would divide into as many as ten spe-
cies, was assigned to the shorter, nari'ower-spored variety of S. donacis
only after long and detailed study. What we have called groups
(Sprague and A. G. Johnson, 1950) cannot be justifiably separated
into species because so many of them grade from one group type to
another. For instance, in some groups the spores equal or approach
those of the slender form of S. donacis on Panicum, while in others,
"•'as those on Sporobolus, the spores are small and close to those of
S. everhartii. The group on Danthonia has evenly lunate spores with
softly tipped ends. It is, nevertheless, difficult to segregate these into
species on any satisfactory basis. Petrak (1929) used Selenophoma
drabae (Fckl.) Petrak to cover these groups and this name has con-
siderable merit. He reported it as the common species of the genus
on sedges, grasses, and crucifers, especially in the arctic. Recently he
reported it on Stipa imbricans from China (1947,a). Our reason for
not accepting S. drabae is that it appears to be too inclusive to be
practical. To many it will seem to be stretching a species to the break-
ing point to recognize a maze, such as variety stomaticola.
The symptoms induced by this variety are of two kinds: (a) light-
colored spots with pale, tawny, red, or lilac borders, or (b) no spots
at all with the minute pycnidia scattered over straw-colored areas. On
Calamovilfa, in the Dakotas, the spots are particularly pale, while in
Danthonia californica Bol. in Oregon the spots are a "frog eye" type
with a prominent red or maroon border.
Pure culture studies, plus examination of a large number of collec-
tions on western grasses and cereals, have shown that var. stomaticola
is divisible into 10 major groups as follows:
Group 1, on Danthonia californica, is closest to Septoria lunata
Grove (1935) with the regularly lunate spores with soft-pointed tips.
It produces a light gray, crustose colony with scant mycelium and
opalescent grayish white spore droplets. The medium was tinted rosy-
!\
vmaceous. The fungus, in inoculation trials, was restricted to Dan-
thonia and would not attack other hostsôf yar. stomaticola. This one
group then, if not the others, could be termed a race, in, its recognized
sense. |
Group 2, on Dactylis glomerata and Koelena] cristata from Pull-
man, Washington. The isolates from these two hosts are identical, each
producing a lavender-tinted colony. This '"race" is becoming Very
prevalent in the Palouse region of Washington arid Ida-ho and is one
of the leaf spots to which the plant breeders are endeavoring to find
resistance. There is considerable difference in the amount of spotting
on various selections and strains of orchard grass in the plots at Pull-
man. In some clones the disease is so severe as greatly to reduce seed
set and the eye spots occur on all the leaves and even on the heads.
Group 3, on Festuca idahoensis from Lyle, Washington, has the ap-
pearance of group 2 in pure culture but the spores are narrower and
resemble those of the next race.
Group 4, on Poa pratensis from Quad Creek, Montana, is akin to
S. donacis on Poa spp. but the burnt-orange, later brown, colonies plus
somewhat boomerang-shaped but smaller spores indicate that this is
at least a localized variant.
Group 5, on Arrhenatherum elatiics, is on the type host for Phyl-
losticta stomaticola. An isolate from Bozeman, Montana, shows less
tendency to produce lavender tints in the substrate than group 2, and
in addition it forms dirty, tan, mucose, conidial masses in early stages
of growth. I
Group 6, on Phleum pratense, is distinct from group 1 in host range
but its relation to the others is undetermined. The fungus was de-
scribed by Lind (1907) as Septoria culmifida. However, it is morpho-
logically indistinguishable from most of the coniplex. Further study
may show that this group is unnecessary. I
Group 7, oh Sporoholus asper, has not !been cultured either and
little is known of its relationship to the grouij. Since it has some boom-
erang-shaped spores it somewhat resembles S. donacis^^ but the spores
are narrower. We have separated it into a group becauseljrspore char-
acteristics. _
Group 8, on Festuca kingii from the mountains of the West..It tends ^
to merge with S. everhartii. I t differs to some-extent from other groups
in having relatively prominent pycnidia.
Group 9, on Deschampsia spp., is somewhat similar to S. everhartii
but the spores range longer. It needs further study. /
Group 10, on Hordeum spp., appears to represent a weak phase of
this complex. I t also needs further study. ' ^
A considerable number of the known hosts of^his variety have not
been classified into groups. The entire problem,4n fact, has scarcely
been touched. Were the groups of more serious importance they would
warrant a great deal of study because they are comparatively unknown
to the plant pathologist. Since they appear to be increasing in impor-
tance they should probably be investigated further. Such work is in
progress.
An extensive list of synonyms for var. stomaticola is given by
Sprague and A. G. Johnson (1947, 1950).
Range: United States, Alaska, Algeria, Canada, China, Czechoslovakia, Den-
mark, Eire, England, France, Germany, Greenland, Hungary, Libia, Nor-
way, Spain, U.S.S.R., Wales.
References: Bauemler (1890); Bisby, et al. (1938); Bubak (1909); Demi-
dova (1926); Frandsen (1943); Grove (1935); Jorstad (1924); Lind
(1907, 1910); Lobik (1928); McKay (1946); Petrak (1929, 1947);
Rostrup (1888); Sprague (1946,e; 1949); Sprague and A. G. Johnson
(1940, 1945, 1947, 1950); Trotter (1916).
Selenophoma everhartii (Sacc. and Syd.) Sprague and

A. G. Johnson—Eye Spot
Septoria everhartii Sacc. and Syd.
S. calamagrostidis Ell. and Ev. [non (Lib.) Sacc]
Lesions on leaves, culms and sheaths pale with light purple or red-
dish borders, soon fading to uniform straw color, pycnidia small, black,
to creosote brown, ostiolate, globose, often 50-80 /j,, less often as large
as 100 fj. on such small grasses as Agrostis, but 80-120 X 75-80 Ju, on
Calamagrostis and coarser grasses; pycnophores few, cuspidate, arising
N from hyaline initial cells; pycnospores short, sharply arcuate, 10-15 X
1.0-1.5(2.2) /t, on Bouteloua 10-15 X 1-3-2 /x.
On: Agrostis diegoensis Vasey, Calif.—A. scabra Willd., N. Dak., S. Dak.—
Aristida longiseta Steud., N. Dak.—A. oligantha Michx., Iowa—Bouteloua
curtipendula (Michx.) Torn, N. Dak.—Calamagrostis canadensis (Michx.)
Beauv., Alaska, Sask., Wise.—C. inexpansa A. Gray, Greenland—C.
koelerioides Vasey, Wyo.—C. montanensis Scribn., N. Dak.—C. rubescens
Buckl., Wash.—C. scribneri Beal, Wyo.—Deschampsia atropurpurea
(Wahl.) Scheele, Mont., Wash., Wyo.—D. caespitosa (L.) Beauv., Mont.,
Wyo.—D. danthonioides (Trin.) Munro, Ida., Wash.—D. elongata (Hook.)
Munro, Wash.—Festuca octoflora Walt., S. Dak.—F. ovina var. brachy-
phylla (Schult.) Piper, Calif., Mont., Wyo.—7^. rubra L., Mont., Wash.—
Muhlenbergia cuspidata (Torr.) Rydb., N. Dak.—M. racemosa (Michx.)
B. S. P., N. Dak.—Puccinellia nuttalliava (Schult.) Hitchc, N. Dak.—
Sphenopholis obtusata (Michx.) Scribn., N. Dak.—Trisetum spicatum (L.)
Richt., Colo., Mont., Wyo.—T. subspicatum Beauv., Greenland—T. wolfii
Vasey, Wyo.
S. everhartii is, in part at least,'an assembly of related fungi placed
for convenience in this one species. With our knowledge of the group
so incomplete it is deemed unwise to segregate this group further. In
general, we distinguish S. everhartii from S. donacis var. stomaticola
by the small size of the spores and their arcuate shape and sharp
points. Those on the smaller-leaved Calamagrostis spp. and on Agrostis
are typicai'while those on alpine fescues and on Trisetum are at least
'I
very similar to those on Agrostis. But there are; others which may be
different from S. everhartii but which we have included in that species
at this time. For instance, collections on B'outelqua have small, some-
what boomerang-shaped spores which are slighfly thicker than those
on the hosts previously mentioned. The material Jon Muhlenbergia and
on Aristida may also be different. 'i
Pure culture isolations from a number of hosts do not clarify the
situation with reference to the group. The fungi from every host, with
few exceptions, are different in appearance and single spore isolates
from the same collection sometimes differ T^ne from another. Isolates *
from Agrostis scabra, North Dakota, have gray or rose-colored,
leathery mycelia. Isolates from Deschampsia atropurpurea have leath-
ery, moist, gray to isabelline colonies. Those on D. danthonioides have
leathery to mucose, finally subcottony colonies which are burnt orange
in color mixed with patches of pink and black, a weird color combina-
tion. Isolates from Bouteloua form off-white crustose colonies. An iso-
late from Trisetum spicatum has moist,/leathery colonies which are
gray and creosote-coloxed with a pale lavender vmderlayer. An isolate
from Muhlenbergia racemosa obtained in extreme western North
Dakota formed a black carbonaceous growth. This isolate, when incu-
bated on sterilized smooth brome grass culms in an uhheated barn over
winter at Mandan, North Dakota, produced small, black perithecia
measuring 80-90 /x diam. and belonging to the Mycosphaerellaceae. A
few contained fascicled asci with eight ellipsoidal ascospores 7-9 X 2.0-
2-6 fjL. As reported previously (Sprague, 1946,e), it could not be de-
termined whether these were slightly immature and only biguttulate
(Ascospora) or somewhat immature'and ultimately 1-septate [Myco-
sphaerella). The spores could not be germinated. By using common
storage conditions, particularly in regions having mild moist winters,
it is probable that the ascigerous stage of jthis. form could readily be
obtained, particularly if the organic medium were sterilized with pro-
pylene oxide. A great deal more study is needed with this group.
The type of Septoria nebulosa Rostr. (1888) is apparently referable
to S. donacis but material on Calamagrostis inexpansU-&nd__Trisetuni
subspicatum Beauv., referred to by Rostrup, appears to be Sêverhartii.
To show how interrelated these species are the reader is referred
to Selenophoma calamagrostidis Moesz and Smarods which was de-
scribed as appearing on very coarse grass, C. epigeios (L.) Roth (Moesz,
1941). This fungus has spores 15-25 X 2.5-4-0<V and is typical in all
ways of S. donacis. We therefore have S. donacis on a coarse species
of Calamagrostis and S. everhartii on the smaller species of the same
genus. The intertangled relationships of all of these become increas-
ingly obvious as further study is made of them^There is a definite cor-
relation between the size of the pycnosporesS^nd the abundance of
nutrient available in the host, that is, in general, S. donacis tends to be
on the coarse grasses and canes, its variety on the pasture grasses, and
S. everhartii on the smaller grasses. The latter is the most common
species on alpine and subalpine plants and is abundant on the dry culms
of grasses in the plains country in late summer.
Range: United States, Canada, Greenland, U.S.S.R.
References: Ellis and Everhart (1898); Rostrup (1888); Saccardo and
Sydow (in Sylloge v. 16, p. 973); Sprague (1946,e); Sprague and A. G.
Johnson (1945, 1950).
I Selenophoma obtusa Sprague and A. G. Johnson—Eye Spot

Spots tawny, margin fuscous to pale lavender, irregular to subcircu-
lar, pycnidia globose, black, 40-150 X 40-138 ^x.; pycnophores cuspidate,
prominent, 3-7 X 2.0-3.5 /,i; pycnospores curved, ends obtuse to some--'
what acute, 13-17 X 2.5-4.2 /i.
On: Agro-pyron inerme (Scribn. and Sm.) Rydb., Wash.—A. spicatum
(Pursh) Scribn. and Sm., Wash.—A. subsecundum var. andinum
(Scribn. and Sm.) Hitchc, Colo.—Elymus condensatus Presl, Mont., N.
Dak., Oreg., Wash.—E. glaucus Buckl., Calif., Oreg., Wash., Wyo.—Poa
arida Vasey, N. Dak.—P. secunda Presl, Oreg.—Sitanion hansenii (Scribn.)
J. G. Sm., Calif., Wash.—S. hystrix (Nutt.) J. G. Sm., Calif., Mont.,
Utah—Stipa coronata Thurb., Cahf.—S. richardsonii Lk., Mont.
S. obtusa is a short-spored variant of the S. donacis complex with
typically curved spores but with blunt ends (Fig. 21,C). They are
somewhat sausage-shaped except that the ends taper to some extent.
There are, however, intermediate forms. Pure cultures isolated from
E. glaucus collected on Mt. Shasta, California, showed prominent black
pycnidial aggregates scattered in salt and pepper pattern over a light
isabelline to very pale flesh background. While the spore exudate is
pale lavender, the rosy or lavender color of the substrate which is so
prominent in some races of S. donacis var. stomaticola is nearly absent
in this case. A culture isolated from Agropyron inerme at Washtucna,
Washington, developed tan, later brown, conidial masses which became
carbonaceous and eventually leathery. This race was practically identi-
cal in appearance to isolates of S. donacis var. stomaticola from Poa
ampla at Pullman, Washington, and from P. epilis, Big Horn Moun-
tains, Wyoming. This collection belongs, perhaps, with S. donacis var.
stomaticola. Further study is needed because our species is not too well
differentiated from S. donacis, except in the type material on Sitanion.
References: Sprague and A. G. Johnson (1945, 1950).
Septoria' Fr. em. Sacc.

Many of the species 'included here were the subject of a detailed
study (Sprague, 1944,d). Additional data (Sprague, 1946,e, 1948,a,c,
etc.) have been added. In addition to the western material this text
includes a certain number of species which occur outside the western
states.
Septoria Fr. em. Sacc. is a genus with wiell-formed; unilocular,
ostiolate, pycnidia-bearing scolecospores whichj are typically at least
ten times as long as wide. They are usually 0-, 1-, 3-, or 7-septate
although variations occur. The pycnospores are hyaliAe or rarely faintly
yellow. The brown or yellow-brown group belongs in Phaeoseptoria
Speg. (Sprague, 1943,b). Septoria merges with Stagonospora Sacc.
which has broadened spores, typically less than ten times as long as
broad.
The genus Septoria has emerged from obscure beginnings whij^h
have been well presented by various writers. Most recently Rogers
(1949) called for retention of the genus as discussed by Fries. Rogers
did not offer a method for segregating from Septoria the group with
poorly formed pycnidia which was first described under Phloeospora
(Phleospora). The first species in the genus with good, well-formed
pycnidia was Septoria rosae Desm. which Fries did discuss (1829). We
prefer to accept Phleospora as a separate genus and to refer all the
true pycnidial forms, generally recognized as Septoria, to the almost
universally recognized emendation by Saccardo with S. rosae as type.
Whether the genus.be credited to Fries em. Saccardo, Saccardo, or even
Fries does not matter too much to ihe writer if the species are left in
the genera to which they are now assigned.
KEY TO T H E SPECIES OF SEPTORIA ON GRAMIJSTEAE I N NORTH AMERICA
A. Pycnospores 0- to 2-septate, less often 3-septate I

B. Pycnidia prominent, as much as 180-250 f. diam.
C. Spores filiform, 36-50 X 1.4-1.6 i>.
Septoria poliomela Sydow
CC. Spores obclavate-filiform, 33-65 X 1.2-2.5 //
S. bromi Sacc. |
CCC. Spores obclavate, 28-50 X 2.5-4.5 M
S. infuscans (Ell. and Ev.) Sprague
BB. Pycnidia intermediate, 100-180 M in diam.
C. Spores filiform,^ 2-septate, 40-56 X 1.8-2.4 /* - ^ ^
)S. elymi Ell. and Ev. f. eZymina^CDavis) Sprague
(Sprague, 1948,c) _
CC. Spores elongate-fusiform, 26-38 X 2-2.5 i^
S. spartinae (Trel.) Sprague
CCC. Spores elongate-fusiform, 24-42 X 2,5-4 t^, pycnidia sometimes
less than 100 /* diam. / /
S. calamovilfae Pettak
CCCC. Spores bacillar, 1-septate; 3-septate in one collection from
Mandan, N. Dak.
S. oudemansii Sacc. /
CCCCC. Spores filiform, faintly septate, 5Qi20-X 1-1.5 f^
S. cenchrina J. "J. BaVis "
BBB. Pycnidia small, obscure, 50-120 /^ diam.
C. Pycnidia typically 50-80 ii, spores 0- to 1-septate
S. triseti Speg. em. Sprague '
FUNGI IMPERFEGTI—SPHAEROPSIDALES 213
CC. Pycnidia typically 80-120 M, spores 0- to 2-septate
S. tenella Cke. and Ellis
AA. Spores typically more than 2-septate
B. Pycnospores usually less than 50 /* long
C. Spores averaging more than 2 /* wide
D. Spores bacillar
E. Spores seldom more than 30 M long
S. nodorum Berk.
EE. Spores frequently more than 30 M long
F. Produces a pale-colored leaf spot, spores 3-septate, 25-50 X
2.5-3.5 li, on rye
S. secalis Prill, and Del.
FF. Produces pale-colored spots, spores 1- to 3-septate, 37-62 X
2.6-3.4 /*, on Stipa and Agrostis
S. secalis var. stipae Sprague
FFF. Produces dark brown leaf spots, spores 3-septate, 25-50 X
2.7-4.1 /*, on Lolium
S. loligena Sprague
FFFF. Produces a brown leaf spot, spores 3-septate, 22-38 X 2.8-
3.8 /J., on Melica
S. melicae Pass.
FFFFF. Produces a pale, brown-bordered leaf spot; spores 3- to
4-septate, 25-45 X 3-4 /», on Avena '
S. avenae Frank {S. avenae f. sp. triticea
T.. Johnson occurs on wheat and the spots
are less well defined or the pycnidia are
borne in vague stramineous areas)
DD. Spores narrowly obclavate, apex more or less pointed
E. Pycnidia 100-140 M, on Sporobolus
S. andropogonis f. sporobicola Sprague
EE. Pycnidia 75-100 n, on Andropogon
S. andropogonis J. J. Davis
EEE. Pycnidia 130-160 /i, on Sphenopholis
S. andropogonis
CC. Spores less than" 2 /J. wide
D. Pycnidia golden-brown, ellipsoid
E. Pycnophores 4-7 fi long, spores 35-40 X 1.3-1.6 t^
S. macropoda Pass.
EE. Pycnophores 3-4 M long, spores 30-50 X 1.2-2.1 ii
S. elymi Ell. and Ev.
DD. Pycnidia creosote-brown, subglobose to ellipsoid, spores 14-46 X
1.3-2.0 f^
S. passerinii Sacc.
BB. Spores frequently more than 50 /« long
C Spores narrowly filiform, about 1 M wide
S. stipina Died.
CC. Spores more than 1 fi wide
D. Spores often more than 2.7 /J. wide
E. Spores averaging less than or approximately 65 n long
F. Spores strictly hyaline
/j
G. Spores 2- to 4-septate, stiffly curved, obclayate-filiform,
pycnophores 4-9 i" long (in part)
S.infuscans (Ell. and Ev.) Sprague
GG. Spores mostly 3-septate, somewhat whiplike, obclavate-
filiform, pycnophores 5-11 i« long ,
iS. bromi var. phalaricola Sprague
GGG. Spores 3-septate, obclavate-cylindric to narrower
S. agropyrina Lob.
GGGG. Spores 3- to 5-septate, narrowly obclavate, up 'to 73 /x
long, on Stipa comata
S. andropogonis i. sporobolicola Sprague
FF. Spores somewhat chlorinous
S. mississippiensis Sprague
EE. Spores averaging more than 65 M long
F. Pycnidia prominent, as much as 320 t^ diam.
G. Spores sometimes over 90 X 5 /« diam.
H. Spores mostly 60-95 X 3.6-5.6 M
S. digitarivoxa Sprague
HH. Spores mostly 60-91 X'5.3-6.6 ii
Stagonospora foUicola (Bres.) Bubak
GG. Spores smaller
H. Spores seldom more than 5-septate, lance-shape, obclavate
Septoria-jacidella Sprague '
HH. Spores 3- to 7-septate, obclavate, apex elongated
S. arctica Berk, and Curtis
FF. Pycnidia smaller, commonly less than 150 f- diam.
G. Pycnophores narrowly ampuUiform, spores long-filiform
S. pacifica Sprague '
GG. Pycnophores cylindrical, spores obclavate-filiform
iS. munroae Ell. and Barth.
DD. Spores less than 2.7 ii wide but more than 1 ii wide
E. Pycnidia brown, cells compact, frequently elongated
F. Pycnophores awl-shaped, 2-5 /* long
G. Spores very narrowly obclavkte-filiform
H. On Agrostis exarata
S. calamagrostidis (Lib.) Sacc.
HH. On Koeleria cristata '^ ^
JS. calamagrostidis'f. koeleriae (Coc^ and ,Mor.)
Sprague —
GG. Spores filiform, less often very narrowly obclavate-filiform.
H. Spores typically sinuous
S. calamagrostidis on Agrostis palustris
HH. Spores usually slightly curved, but less often sinuous
S. calamagrostidis on Trisetum spp.
FF. Pycnophores short, papillate, 2-3 /J. long
S. macropoda Pass, on Pba howellii
EE. Pycnidia golden-brown, cells moderately compact, variable in
shape, not uniformly elongated • ^ - --
F. Spores seldom over L7 ti wide
S. macropoda var. septulata (Gonz. Frag.)
Sprague
FUNGI IMPERFECTI—SPHAEEOPSIDALES 215
FF. Some spores more than 1.7 M wide
G. Spores never as much as 8- to 9-septate
H. Pycnophores 2 to 6 /« long
S. macropoda var. grandis Sprague
HH. Pycnophores 4 to 13 M long
I. Spores typically sinuous
S. tritici f. avenae (Desm.) Sprague
II. Spores variously curved, less often sinuous
J. Microspores present, macrospores 25-90 X 1.4-2.6 M on
wheat
S. tritici Rob. in Desm.
JJ. Microspores not known
K. Pycnospores as much as 105 /^ long on Holcus
S. tritici f. hold Sprague
KK. Pycnospores as much as 85 M long, on Lolium
S. tritici var. lolicola Sprague and A. G. John-
son
GG. Spores 3- to 9-septate, up to 95 /i long
JS. andropogonis var. sorghastri H. C. Greene
and Sprague
GGG. Spores usually 5-septate, up to 60 n long
S. quinqueseptata Sprague
Septoria agropyrina Lobik—Leaf Spot and Blotch

Pycnidia brown, ostiolate, somewhat flattened, 80-100 X 60-150 /x,
spores coarsely obclavate to subcylindric, 3-septate, hyaline, apically
pointed, 30-65 X 2.5-4.0 /^.
On: Agropyron caninum (L.) Beauv., Wash.—A. inerme (Scribn. and Sm.)
Rydb., Wash.—A. pertenue (G. A. Meyer) Nevski, N. Dak.—A. repens
(L.) Beauv., Minn., N. Dak., S. Dak., Wash., Wise.—A. semicostatum
(Steud.) Nees, N. Dak.—4. smithii Rydb., Mont., Nebr., N. Dak., S. Dak.,
Wyo.—A. trachycaulum {Ik.) Malte, Mont., Nebr., N. Dak., S. Dak.,
Wyo.—Elymus canadensis L., N. Dak., S. Dak.—E. condensatus Presl,
N. Dak.—E. giganteus Vahl, N. Dak.—E. glaucus BuckL, Oreg., Wash.-—
E. junceus Fisch., N. Dak.—E. macounii Vasey (natural hybrid), N. Dak.
—E. triticoides Buck!., Calif.—E. virginicus L., Miss., N. Dak.
This species is a very common but relatively unimportant com-
ponent of the leaf-spot complex on Elymus and Agropyron spp. in the
Great Plains. Its range, however, extends west into Oregon and Wash-
ington. It apparently occurs widely in Asia and adjacent Europe. It is
usually found as a saprophyte-on the tips of dead or nearly dead leaves
but some collections show' it associated with a brown or fuscous blotch.
It is sometimes difficult to distinguish this fungus from a slender phase
of Septoria infuscans on the one hand, and from Stagonospora arenaria
or even Septoria avenae f. sp. triticea T. Johnson on the other. Its
apically pointed spores (Fig. 24) distinguish it from St. arenaria, while
its development on potato dextrose agar (it forms a cottony buff
colony) contrasts sharply with the scanty pellicular growth of S. in-
fuscans. While typical S. infuscans does not appear to extend east-of
Letter, Wyoming, S. agropyrina does occui: with St. qrenaria west to
the coast (see also Fig. 43>,E,F,G,H,I,J). j
Septoria agropyrina is readily segregated from most material of
S. elymi, as the latter has much smaller spores and produces a yeasty
colony on potato dextrose agar. There is, how-
ever, a phase of S. elymi that is possibly the
same as S. elymicola Died., which may gome-
times be intermediate between typical S. elymi
and the more slender-spored collections of S.
agropyrina. This fungus, which is a phase of
S. elymi, recently assigned to S. elymi f. elymina
(Davis) Sprague (1948,c), occurs on Elymus
and Agropyron in Michigan, North Dakota,
Iowa, Utah, and Wisconsin.
Septoria agropyrina is further briefly dis-
cussed in connection with associated forms men-
tioned in this text. It should be pointed out that
critical comparison with European types is
needed, something which has" been impossible
to accomplish under recent, and to some extent,
current conditions.
24. —• Pycno-
of Septoria Septoria agropyrina Unam. (1930) has non-
agropyrina on Ely- septate spores 46.5-71.5 X 3.0-3.5 jn and is ap-
mus glaucus, Orleans, parently very distinct from Lobik's specimen.
Oreg.
Lobik and Unamuno have several fungi bearing
the same name whose descriptions were published within two years of
each other. /
Range: United States, U.S.S.R.
References: Lobik (1928); Sprague {19i4,d', 1946,e).
I
Septoria aîdropogonis J. J. Davis—Leaf-Sp^
Causing narrow elongated areas of a reddish yellow color some-
times becoming sordid; pycnidia epiphyllous, subseriate or scattered,
dark brown, depressed-globose, 75-100 /i; Spores hyaline, straight or
slightly curved, more acute at the apex and slightly thickened at the
base, typically stiff and not sinuous, becoming 2- to 4-septate, 30-
51 X 2-4 ,i..
On: Andropogon jurcatus Muhl., Kans., Nebr., Okla.,-S. Dak., Wise.—A.
scoparius Michx., Nebr.—Bouteloua curtipendula (Michx-f) Torr., S. Dak.
—Sphenopholis obtusata (Michx.) Scribn., 'N^âk.-^Sporobolus hetero-
lepis A. Gray, Wise.
This fungus, and its various subdivisions, is common in the Great
riaifas region. The species proper on Andropogon jurcatus Muhl. has
been collected in Kansas (Lefebvre, see Sprague, 1943,c), Oklahoma
(Preston), and Wisconsin which is the type area (Davis, 1915,a), and
recently in Nebraska and adjacent South Dakota (Sprague, field notes).
FIG. 25.—Pycnospores of: A, Septoria andropogonis f. sporobolicola on

Sporobolis heterolepis, N. Dak.; B, S. andropogonis on Andropogon furcatus,
Kansas; C, ;S. mississippiensis on Muhlenbergia mexicana, N. Dak.; D, S. andro-
pogonis i. sporobolicola on Stipa comata, N. Dak.; H, S. andropogonis on
Sphenopholis obtusata, N. Dak. (From Mycologia.)
Gray elongate lesions are formed with moderately prominent pyc-

nidia scattered in the spots or more or less seriately arranged. The
spores are characteristically obclavate with tapering blunt bases (Fig.
2b,B) and narrowly sharpened apices (Sprague, 1943,c, Fig. l,b). They
are usually 3-septate, 30-51 X 2.0-3.9 ii and the contents are hyaline to
yellowish.
As far as we know, this fungus has never been cultured and nothing
has been done to determine strains or races on the|widely^different hosts
which have been encountered. • ' <
One collection on Sphenopholis obtusatd frorn Devil's Lake, North
Dakota, appears to be S. andropogonis (Fig. 25,H) rather than S.
quinqueseptata Sprague, which also occurs on this host
Septoria fontii Gonz. Frag., described (1923)| from Spain on An-
dropogon should be compared with S. andropogonis. While the spores
are 20-30 X 3-5-4 /n, the illustration strongly suggests Davis' species.
References: Davis (1915,a); Sprague (1943,c).
Septoria andropogonis var. sorghastri H. C. Greene and Sprague—

Leaf Spot
As in the species but the spores are 3- to 9-septate, hyaline, 56-
100 X 1.5-2.4 /A, filiform at the base, tapering to a narrow, yet rounded,
apex. '
On: Sorghastrum nutans (L.) Nash, Nebr.,/Wisc. ^
This differs from the species in its much longer and multiseptate
spores. Except for their extreme length they are fundamentally the
same type of. spore as the species and on a genus closely related to
Andropogon. The first collection of this fungus on Sorghastrum nutans
(L.) Nash was made by the Rev. J. M. Bates many years ago at Long
Pine, Nebraska, but the specimen was undetermined until Greene and
the writer compared notes with some recently collected Wisconsin
material and concluded that the fungus was only an unusual phase of
S. andropogonis. I
Range: United States. j
References: Greene (1944,6); Sprague (1944;d; 1946,e).
Septoria 'andropogonis forma sporobolicola Sprague—Leaf Spot

and Blotch ~~~^ —
Spots oval to subelongate, elliptical, small, 1-3 mm., scattered in
living, vigorous leaves; white to straw-colored with narrow but well
defined, anthocyanin red borders; pycnidia 2 to several in each spot,
strongly erumpent, almost superficial, pale brown, thin-walled, some-
what elongate but depressed-globose, ostiolate, 120-150 X 100-140 /»;
pycnospores hyaline but becoming yellow in saprophytic stage, con-
tents with a few small, not prominent, guttulae more or less adjacent
to the cross walls, basal cell tapering, blunt, pr^subtruncate, spores
stiffly curved or nearly straight, obclavate,-afiex^ sharp pointed, 45-
70 X 2.4-3.4 /x, typically about 50 X 2.6 /x in spot material, broader in
saprophytic material, 3-septate.
On: Sporobolus heterolepis A. Gray, N. Dak., Wise.—Stipa comata Trin. and
Rupr., Mont., N. Dak.—S. viridula Trin., N. Dak., S. Dak.
The type of this occurs as a nonstriate, circular spot on Sporobolus
heterolepis but similar material has been found on Stipa comata at
Mandan, North Dakota. The fungus on the latter host causes a blotch
at first, later a light-colored spot. The spores of the type on Sporobolus
(Fig. 25,A) are slightly narrower than those of Septoria andropogonis
proper, while those on Stipa are somewhat longer (40-73 X 2.4-3.4 jn),
but also proportionately slightly narrower (Fig. 25,D). This character,
plus the somewhat larger, light-colored pycnidia in f. sporobolicola
readily segregates the form from the species. Except for these differ-
ences, they all belong in one species. At present f. sporobolicola is con-
fined to the Agrostideae of the Gramineae while the species proper and
its variety sorghastri occur on the Andropogoneae except for the col-
lection on Sphenopholis of the Avenae. The possibly related species, S.
quiUqueseptata also occurs on Sphenopholis, while another related, but
we believe distinct species, S. mississippiensis, occurs on Muhlenbergia
of the Agrostideae. I t would appear that this group has adapted itself
to a considerable range of tribes under the favorable conditions of as-
sociation in the plains country.
Forma sporobolicola was recently identified also on Stipa viridula
Trin. from South Dakota and in great quantities at Amidon, North
Dakota. It is probably much more common on this host than these
few collections indicate as much of the material on Stipa is not fruiting
and we have frequently collected similar unidentifiable material.
References: Sprague (1943,c; 1944,c?).
Septoria arctica Berk, and Curt.—Leaf Spot

Spots elongated, ill-defined, brown to dull gray, merging with the
green surrounding tissue, on leaves, sheaths, glumes, and pedicels,
pycnidia prominent, strongly erumpent, black, tardily ostiolate, globose,
with firm peridium, semiloculated, walls 8-12 fx. thick, dark creosote-
brown, composed of tightly packed corky cells with large, irregular,
polyhedral ones included in some folds of the walls; pycnophores stout,
broadly to narrowly pyriform; pycnospores exuded in thick masses,
hyaline, obclavate-scolecosporous with tapering, rounded bases and
sharply pointed ends, 2- to 7-septate, 60-90 X 2.8-4.6 ju,, straight to
stiffly curved.
On: Calamagrostis nutkaensis (Presl) Steud., Alaska, Oreg.—Dupontia
fischeri R. Br., Bering Straits (U.S.S.R.).
The type of this species was collected on Dupontia fischeri on an
island off the Siberian coast in the Bering Straits, many years ago
V
(Berkeley and Curtis, 1860). It also occurs on Calamagrostis near Sea-
side, Oregon, and apparently in southern.Alaska and northern Europe
(Sprague, 1944,d). The" spots are ill-defined brown
to dull gray lesions; pycnidial are large (140-250 ix)
(Fig. 48,A''). The spores in 'the type specimen are
2- to 3-septate or- more, 60-80 X 2.8-3.6 jn, straight
to stiffly curved, and somewhat resemble spores of
S. jaculella Sprague on Bromus spp. The spores
on C nutkaensis are 3- to 7-septate, 70-89 X 3.5-
4.6 /J. (Fig. 26). The fungus is probably common
in coastal Alaska and it appears to be distinctly
parasitic.
Range: United States, Siberia.
References: Berkeley and Curtis (1860); Sprague
(1944,d; 1946,e).
Septoria arechayaletae Wint.—Leaf Spot

Spots tawny to straw-colored, vague; pycnidia
sparse, subgregarious, totally immersed, globose,
about 100 fx diani.; pycnospores "filiform, obscurely
septate, hyaline, 50-115 X 1-0-1.5 ja.
On: Panicum capillare L., Va.
Most of the material on Panicum in the United
States has gregarious, black, cupulate, collapsed
FIG. 26.—Pyc- pycnidia which appears to be S. tandilensis. The one
nospores of Sep-
toria arclica on collection of David Fairchild's from Virginia is
Calamagrostis nut- closer to S. arechavaletae (Sprague, 1946,a).
kaensis, Clatsop
Co., Oreg. Range: United States, j^rgentina.
References: Sprague (1946,a); Winter (1887).
Septoria avenae Frank^Speckled Blotch of OaTsând, Grasses

Ascigerous stage is Leptosphaeria avenaria G. F. Weber
Stagonospora arrhenatheri Smith and Ramsb. "
Spots small, circular to elongate, elliptical, yellowish to light brown,
mottled, eventually fading, leaving the pycnidia to produce a speckled
lesion; pycnidia more or less scattered or in rows, subepidermal, globose
to subglobose, 90-150 [i diam., average 120 /n, wall smooth (Fig. 27,B),
brown to black, ostiole round to oval, 20-30 /x diam.; pycnospores bacil-
lar, 3-septate, hyaline, 25-45 X 3-4 jn; perithecia globose to subglobose,
60-130 /i diam., walls smooth, black, thin,_ osftole 12-20 [x diam.; asci
narrowly clavate with rounded tips, hyaline, thin-walled, 30-100 X 10-
18 fi; ascospores fusoid, straight or slightly curved, ends obtuse,
rounded, 3-septate, constricted at the septa, especially the central
septum, second cell from t h e top usually swollen, light yellow to slightly
olivaceous, 23-48 X 4.5-6.0 /*.
O n : Agrostis exarata Trin., Oreg.—Arrhenatherum elatius (L.) Beauv.,

Penn., Wash.—Avena sativa L., Alta., B. C , Iowa, Ky., Manit., Mass.,
Minn., N. B., N. Dak., N . S., Ont., P. E. Is., Que., Wise—Calamagrostis
canadensis (Michx.) Beauv. (spores 21-28X3.8-4.8 M), Mont.—Festuca
elatoir L., Utah—Glyceria elata (Nash) Hitchc, Wyo.—G. grandis S.
Wats., Wyo.—G. occidentalis (Piper) J. C. Nels., Ida.—G. pauciflora Presl,
Ida., Wash.—G. striata (Lam.) Hitchc, Wyo.—Oryzopsis hymenoides
(Roem. and Schult.) Rick., Mont.—Schizachne purpurascens (Torr.) Swal-
len, Nebr.—Scolochloa festucacea Lk., N . Dak., S. Dak.
FIG. 27.—Septoria avenae (except C and F ) : A, pycnospores on Arrhenatherum

elatius, State College, Pa.; B, cross section of pycnidium from same source as A;
C, S. melicae on Melica scabrosa, N. Dak.; D, pycnospores on A. elatius, Engl.;
E, pycnospores on Avena sativa; F, pycnospores of Stagonospora arenaria on
Dactylis glomerata, near Crescent Lake, Wash., showing somewhat coarser spores
than those of S. avenae (F. Columb. 4771), Canada. (From Mycologia.)
/ '
Since this species is well known as the pycriidial 3tage of Lepto-
sphaeria avenaria possibly the taxonomyxof thjC form genus is not
especially important. In an earlier paper we included St. arrhenatheri
under Stagonospora arenaria Sacc. (Sprague, 1941,c). Further study
shows that St. arrhenatheri (Fig. 27,A,D) is only B. avenae (Fig. 21,E;
28,B) on another host, and this host is very simila'r to Avena. A collec-
tion on Oryzopsis belongs here (Fig. 58,F).
A common purple blotch or spot on the common marsh grass
Scolochloa festucacea seems to be S. avenae. The uniformly'cylindrical'
spores are 27-31 X 2.4-3.0 /* (Sprague, 1948,a). We also found a dark
purple spot on meadow fescue in Logan Canyon, Utah. The spores
were mostly 1-septate, rarely 2- to 3-septate, cylindrical, hyaline, 16.8-
27.2 X 3.1-4.7 IX (Sprague, 1948,c). We also found abundant material
of S. avenae on Glyceria paucifiora in a shaded area in the Chelan Na-
tional Forest, Washington. The spots were gray, elliptical to striate,
later fading to straw color and with surrounding yellow areas, or the
lesions were nearly buff when mature. The spores were truly cylindrical
and hence distinct from those of S. glijcericola which have pointed
apices. They measured 35-46 X 3.4-4.1 ju. Similar material was found
at Priest Lake, Idaho, with spores 37-45 X 3.5-4.2 ja, and in addition
spermatia-like microspores were present, 4-5 X 1.1-1-5 ju,. Another col-
lection on G. elata at Teton Pass, Wyoming, had spores 28-34 X 3.0-
4.4 fi and some of the spores were somewhat pointed but too wide for
S. glycericola (A.S. 17,091). ;
S. avenae was found on Agrostis exarata Trin. near Elsie, Oregon.
The 3-septate spores were 33-39 X 3.2-3.8 ^ (Sprague, 1949,a).
Range: United States, Argentina, Australia, Canada, Denmark,- England,
France, Germany, Norway, Poland, South Africa, U.S.S.R.
References: Drayton (1926); Frandsen (1943); Johnson, T. (1947); Smith
and Ramsbottom (1916); Sprague (1941,c; 1944,rf; 1946,e; 1948,a,c;
1949,o; 1950,a); Weber (1922,a). I
Septoria avenae Frank f. sp. triticea T. Johnson—Blotch

Lesions on leaves ovate, frequently coalescing, straw color or buff,
central portion bearing pycnidia, often gray. Pycnidia sometimes fol-
lowed in late summer by perithecia. Perithecia at first subepidermal,
later erumpent, black by reflected light, browti by transmitted light,
globose to subglobose, 100-220 /A diam., surface cells polyhedral, often
pentagonal; ostiole circular, 14-20 /* diam., asci clavate, straight or
slightly curved, hyaline, 40-80 X 8-11 /A, 8-spore^,_ascospores biseriately
arranged, fusoid, straight or curved, 3-septat^7^frequently constricted
at the septa, light yellow, (16)-19-25-(28) X 4-6 ju.; paraphyses fila-
mentous, septate, slightly longer than the asci, hyaline.
FUNGI IMPERFECTI-SPHAEROPSIDALES 223
Pycnidia at first subepidermal, later erumpent, scattered, spherical
or slightly elongate, black by reflected light, golden brown by trans-
mitted light, (80)-90-140-(210) [i diam., surface cells polyhedral; walls
composed of three or four layers of brown cells, 7-11 ft, thick at base and
11-14 )i thick at top of pycnidium; ostiole circular or oval, 10-15 /x.
diam.; pycnospores hyaline to faintly chlorinous, thin-walled, cylindri-
cal, straight or slightly sinuate, obtuse or rounded at the ends, 3-septate
or rarely 4-septate, (18)-26-42-(53) X (2.3)-2.8-3.5-(4.2) y,. Spores
are generally shorter and more slender than in Septoria avenae proper.
On: Agropyron smithii Rydb., Mont., N. Dak.—A. trachycaulum (Lk.)
Malte, N. Dak.—Elymus antarcticus Hook, f., Wash.—E. excelsus Turez,
N. Dak.—E. glaucus Buckl., Ida., Oreg.—E. junceus Fisch., N. Dak.—E.
sibiricus L., N. Dak.—Hordeum murinum L., Oreg.—fZ^. vulgare L., Manit.,
Sask.—Triticum aestivum L., Alta., Manit., Maritime Provinces, N. Dak.,
Ont., Sask., Wash.—T. dicoccum Schrank, N. Dak.—T. durum Desf.,
Manit., N. Dak.
T. Johnson (1947) has found that the species of Septoria on wheat,

and less often barley, whose cylindrical spores are larger (Fig. 28,C)
than those of S. nodorum is morphologically very similar to S. avenae
and to Stagonospora arenaria. He found that a perithecial stage was
also virtually identical with Leptosphaeria avenaria. The spores of the
pycnidial stage averaged slightly smaller on wheat than they did on oats
but there was considerable variation as indicated by the measurements
given. This species causes a pale spot on leaves of wheat and all indi-
cations are that the -fungus is not nearly as actively parasitic as S.
nodorum. Johnson found the fungus on grasses, and he found that
isolates from these grasses, which he did not specify, could attack
wheat. He obtained negative infection results on Canada wild rye, tall
meadow oats grass, orchard grass, and reed canary grass with the
fungus from wheat. While both he and the writer have inferred that
Septoria avenae and Stagonospora arenaria are very closely related,
neither has stated that they are identical. It is still not possible to
prove this point. As will be pointed out later under St. arenaria, the
spores of the material which we have left in the last mentioned
(Sprague, 1948,c) are somewhat larger than those that Johnson reports
on wheat. They are as large or larger than those of S. avenae on oats.
In addition many of the cultures have the appearance of those illus-
trated by Johnson for S. avenae on oats. The genetic coniiectioh with
the Leptosphaeria complex has not been proved for most of these col-
lections and the negative results which Johnson obtained in his inocu-
lation studies suggests that the time is not right for placing St. arenaria
under the Leptosphaeria avenaria group. We have, however, in check-
ing over a considerable number of specimens, removed some of these
from St. arenaria and placed them in Septoria avenae f. sp. triticea
because the spores are small enough to justify •iiiclusion there. This has
been anticipated for some years but we awaited the article by Johnson.
As he pointed out (1947) Bisby and the writer have both assigned'some
of these smaller forms to S. nodorum (cf. Sydow, 1940) and it is these
which we now know have genetic connection with a Leptosphaeria.
A collection of Septoria on Puccinellia nuttalliana is close to S.
avenae (Fig. 28,A).
FIG. 28.—Pycnospores of: A, Septoria^sp. (probably S. avenae) on Puccinellia

nuttalliana, Crystal Springs, N. Dak., compared with B, S. avenae on oats
(B.P.I. 80,888), Mandan, N. Dak.; C, S. avenae f. sp. triticea on wheat, Langdon,
N. Dak.
Range: United States, Canada. '

References: T. Johnson (1947); Sprague (1948,a, pp. 185-186;c).
Septoria brevispora Ellis and J. J. Davis—Leaf Spot

Spots sordid white, with brown margins; pycnidia dark brown, flat-
tened, composed of very small, crushed .cells, short papillae serving
as pycnophores; spores hyaline, clavate,; 3-septate, as much as 15-
30 X 2.5-4.0 /A. , --.
On: Bromus ciliatus L., Wise. '_
This species (Davis, 1903) is recognized mainly because there is
not sufficient information, except the type description, to do otherwise.
It is questioned if it is a distinct species. Sectioned material proved that
the fungus was different from S. hromi and S. jaculella (Sprague,
1944,d) (Fig. 37,N). Later Davis (1929) reported material on a coarse
grass, possibly B. ciliatus, with spores 15-30 X 2.5-4.0 /*. It is suspected
that this is not the same as the type and is like Stagonospora bromi
"Smith and Ramsb. St. hromi, however, has $rown' or purple brown
lesions instead of the sordid white ones of Davis' later collections
'(1929). We also compared this fungus with the spores found in the
type of S. bromivora (Fig. 37,M) (Sprague, 1944,d). S. brevispora may
on further study prove to be either S. nodorum or St. bromi.
References: Davis (1903, 1929); Sprague (1944,d, p. 122, Fig. 18^n).
Septoria bromi Sacc.—^Leaf Spot

Spots elongate to elliptical, indefinite to brown, leaves often yellow,
drying to brown; pyenidia black-brown, spherical to flattened (Fig.
48,J), outer pycnidial wall composed of closely intertwined cells, 4 to
10 /A thick; pycnophores narrowly ampulliform to subcylindric, 5,r
7 X 1.5-2.0 /x; pycnospores characteristically whip-like, narrowly fili-
form-clavulate, usually 2-septate, 33-65 X 1.2-2.5 /*.
On: Bromus ciliatus L., N. Dak., N. W. Terr.—B. commutatus Schrad., Ida.,
,Okla., Oreg., Wise.—B. inermis Leyss, Alta., Ida., Iowa, Manit., Mont.,
N. Dak., Oreg., Wash., Wise.—B. japonicus Thunb., Ida., Nebr., Utah,
Wash., W. Va., Wye—5. latiglumis (Shear) Hitchc, Wise. (Fig. 37,0)—
B. mollis L., Oreg., Wash.—B. purgans L., Okla.—B. racemosus L., Oreg.—
B. secalinm L., Ida., 111., Ind., Iowa, Ky., Mo., N. Car., Ohio, Okla., Oreg.,
Tenn., Tex., Wise.—B. texensis (Shear) Hitchc, Okla. Preston (1945)
also listed B. carinatus as a host of S. bromi from Oklahoma.
The spores are whip-like (Fig. 29), not javelin-like as in S. jaculella
(Fig. 37,A-L). S. bromi is common in regions where moisture is suffi-
ciently prevalent to permit this leaf spot to thrive. I t is present in
prairie regions, and in the late fall or during open winter weather in
some areas such as the Palouse region of Washington and Idaho, this
fungus develops extensively on necrotic leaves.
Range: United States, Austria, Chile, Denmark, Italy, Siberia.
References: Saccardo (1878); Spegazzini (1910); Sprague (1944,cf;
1946,e); Weber (1923).
Septoria bromi var. phalaricola Sprague—Buff Spot

Lesions on basal leaves elliptical, buff, as much as 1 cm. diam., or
involving entire leaf, pyenidia brown, globose to subglobose, 150-
i90 X 204-256 /j. ostiolate; walls thin, 6 [i, consisting of one to two
intertwined, light brown, parenchymatous layers of oblong cells from
which hyaline hyphae arise terminating in cylindrical t o narrowly subu-
late pycnophores, 4-11 X 1.5-2.2 jw,; pycnospores narrowly obclavate-
filiform, somewhat whip-like, commonly 3-septate, 34-66 X 1.4-2.9 /t
(Fig.29,£>).
On: Phalaris arundinacea L., Oreg.
This species may be distinguished from certain other pycnidium-
bearing leaf spots on reed canary grass. A key is presented here.
FIG. 29.—Septpria bromi: A, pycnospores from Bromus japonicus, Romney,

W. Va.; B, pycnospores from B. secalinics near Alsea, Oregon (O.S.C. 8299);
FVKGl IMPERFECTI—SPHAEROPSIDALES 227
KEY TO PYCNIDIAL FORMS ON PHALARIS
A. Spots pale with narrow fuscous or tinted border, spores nonseptate, falcate
Selenophoma donacis (Pass.) Sprague and A.
G. Johnson
AA. Spots buff, yellow, or obsolete
B. Spots pale or absent, spores 0- to 1-septate, falcate, from Argentina
Septoria macrostoma Speg. (Sprague, 1944,d,
Fig. 19,c)
BB. Spots buff
C. Spores 1-septate, hyaline, 20-25 X 1.3 M, from Italy
Septoria phalaridis Cocc. and Mor.
CC. Spores 3-septate, hyaline, obclavate-filiform, from Oregon and-^
probably Italy
Septoria bromi var. phalaricola
BBB. Spots tawny blotches, spores 38-71X4.5-6.0 ii (when mature),
3- to 8-septate (3-septate phase is 25-40 X 2.7-4.5 M) from the Middle
West and the Great Plains and in Europe
Stagonospora folikola (Bres.) Bub.
Septoria bromi var. phalaridis Trail (1887,£)) is referred to Phaeo-

septoria phalaridis (Trail) Sprague (1943,6), a saprophyte. S. bromi
var. alopecuri Karst. (1884,a) is similar to Hendersonia crastophila
(Sprague, 1943,c).
, Var. phalaricola was collected at Astoria, Oregon, in row plots at
the Agricultural Experiment Station near Young's Bay. It has not been
found since.
References: Cocconi and Morini (1884); Saccardo (1878); Sprague
(1943,6; 19U,d; 1946,e).
Septoria calamagrostidis (Lib.) Sacc.—Leaf Spot

Ascochyta calamagrostidis Lib.
• Septoria agrostidis Frandsen (1943)
Lesions pale green, gray to straw-colored, usually on the tips of the
leaf blades, pycnidia obscure, scattered, strongly flattened, subepi-
dermal, dark brown, appearing black in spots, 50-140 X 50-180 fi. diam.,
50-85 f). deep, walls thin, of overlapped hyphae; pycnophores small,
C, cross section of pycnidial wall from B. latiglumis, Bruce, Wise; D, cross sec-
tion of pycnidial wall of var. 'phalaricola on Phalaris arundinacea, Astoria, Oreg.;
E, cross section of pycnidial wall, from JS. secalinus, Temple, Texas; F, pycno-
spores from B. mollis, Oregon; G, pycnospores from B. commutatus, Lane Co.,
Oreg.; H, pycnospores on B. mollis, near Stevenson, Wash.; I, pycnospores and
attached pycnophores from B. mollis, Linn County, Oreg.; J, cross section of
pycnidial wall on B. mollis, Bellfountain, Oreg.; K, pycnospores from B. lati-
glumis (B. incanus), Wise; L, cross section of pycnidial wall, on B. secalinus,
near Alsea,. Oregon (O.S.C. 8300); M, cross section of pycnidial wall, on B. se-
calinus, near Corvallis, Oreg. (O.S.C. 10,402); N, pycnospores from B. mollis,
Austria. (From Oregon State College Bot. Monogr. 6, fig. 17.)
FIG. 30.—Septoria calamagrostidis: A, pycnospores from Agrostis pahtstris,

South ¥ork, Alsea River, Oreg., May 7, 1938 (O^S.C. 189); B, pycnospores from
blunt, bottle-shaped, seldom more than 3.5 X 1-5 /t (Fig. SO,C,D,I,K);
pycnospores filiform, distinctly 0- to 5-, mostly 3-septate, curved and
often sinuous, 25-73 (40-65) X 1.0-2.0 fj. (Fig. ZQ,A,B,E,F,G,H,J).
On: Agrostis diegoensis Vasey, Oreg.—A. exarata Trin., Alaska, Oreg.—A.
palustris Huds., Oreg., Wash.—A. rossea Vasey, Wyo.—A. scabra Willd.,
Alaska, Wise.—Trisetum canescens Buckl., Ida., Oreg.—T. cernuum Trin.,
Oreg.
There are two species of Septoria on Agrostis spp. in the Pacific
Northwest. The species above discussed is common on A. palustris
(Seaside Bent). S. triseti is equally common on A. tenuis (Astoria
Bent). S. triseti has smaller, more globose pycnidia and very different,
short, few-septate spores.
The common race of S. calamagrostidis which is that on A. palustris
is numbered 1 (Fig. 30,A,D,G,I). A race on A. exarata is race 2 (Fig.
30,H) while collections on Trisetum (Fig. ZO,B,F,J) are called race 3.
The spores of race 3 are usually stiffer and straighter than those of
races 1 and 2. Such species as S. caballeroi Gonz. Frag. (1914), S.
caballeroi var. panicei Gonz. Frag. (1917) and S. graminum f. triseti-
loeflingianii Caballero (1928) are synonyms of S. calamagrostidis,
race 3.
. It appears to us that S. agrostidis,Fva,ndsen (1943), which has spores
30-55 X 1.0-1.5 ju, is indistinguishable from S. calamagrostidis.
Range: United States, Belgium, Denmark, France, Greenland, Italy, North
Africa, Spain, U.S.S.R.
References: Anderson, J. P. (1920); Caballero (1928); Coccini and Morini
(1884); Frandsen (1943); Gonzalez Fragoso (1914, 1917); Saccardo,
Peck, and Trelease (1904); Sprague (1938,rf; 1944,d; 1946,e; 1949,a).
Septoria calamagrostidis f. koeleriae (Cocc. and Mor.) Sprague—

Blotch
S. koeleriae Cocc. and Mor.
Spots definite, circular to subcircular, fuscous to vague, brown dead
leaf tips; pycnidia prominent, ellipsoid, often flattened at the top,
black-brown, composed of compacted cells, ostiolate, 80-140 X 70-120
/J.; Dycnophores awl-shaped, 2-5 X 1.6-2.0 /*; pycnospores needle-like,
0-2-septate, 40-60 X 1.1-2.0 /i.
Trvietum canescens, Silver Creek Falls Park, Oreg. (O.S.C. 271); C, cross section
of pyenidial wail, on T. canescens, Crater Lake area, Oreg. (O.S.C. 8498); D,
cross section of pyenidial wall, on A. palustris, Corvallis, Oreg. (O.S.C. 8490);
E, pycnospores from A. diegoensis, near Kellogg, Oreg. (O.S.C. 419); F, pycno-
spores from T. cernuum, Bergsvik Creek, Oreg. (O.S.C. 198); G, pycnospores from
A. palustris, Corvallis, Oreg.; H, pycnospores from A. exarata. Silver Creek Falls
Park, Oreg. (O.S.C. 273); I, cross section of pyenidial wall from A. palustris,
Corvallis, Oreg. (O.S.C. 53); J, pycnospores from T. canescens, Sutherlin, Oreg.;
K, cross section of pyenidial wall from A. diegoensis. (From Oregon State Col-
lege Bot. Monogr. 6, fig. 14.)
On: Koeleria cristata' (L.) Pers., Ariz':, Colo., Ida.j IST. Dak., Oreg., Wash.,
Wyo.
- i •
This fungus is similar to S. calamagrostidis. The compacted pyc-
nidial wall is somewhat distinctive (Fig. 48,C) j and the needle-like
spores are different in some ways from racfes 1 and 2 but are more like
race 3 on Trisetum. Host range trials show that f. \koeleriae is confined
to June grass. S. koeleriae var. koeleriae (-) vallesianae Unam. was de-
scribed on K. vallesiana from Spain (Unamuno, 1941) and S. koeleriae
var. macrocarpa Rayss from Palestine (Rayss, 1941).
Range: United States, Africa, Asia, Denmark, Italy, Spain. » •
References: Bubak (1909); Cocconi and Morini (1883); Frandsen (1943);
Sprague (1944,d; 1948,a).
Septoria calamovilfae Petrak—Leaf Spot

Spots white or pale gray on the leaves' and sheaths, pycnidia 70-
130 n diam., ostiolate, wall 7-12 fi thick, pycnospores smoky-hyaline,
spindle-shape, 0- to 2- sometimes 3-septate, 24-42 X 2.5-4.0 /i.
I
On: Calamovilfa.longifolia (Hook.) Scribn., Mont.,
N. Dak., S. Cak., Wyo.
This fungus is not -uncommon in the north-
ern Great Plains of the western United States
and possibly occurs north into Saskatchewan.
It is distinct from any other species, of Septoria
in the area. It is apparently not very parasitic
but lesions are frequently surrounded by living
tissue. No investigation been made of this
species other than the original taxonomic study
FIG. 31. —Pycno- by Petrak and some cursory field observations
spores of Septoria by the writer (Fig. 31).
calamoviljae. on Cal-
amovilfa longifolia,
near Valley City, N. Range: United States. ^''~~~-^-.__ .
Dak. References: Petrak (1925, p. 86)j_ Sprague
(1944,rf; 1946,e; unpublished field notes).
Septoria carricerae Fairman—Leaf Spot

Leaf spots somewhat elongate, white above, brown beneath, pycnidia
small, epiphyllous, sparse, black, smooth, ostiolate, immersed, then
erumpent; jaycnospores filiform, straight to curved, apices acute, com-
monly 3-septate, hyaline, 30-35 X 1.5-2.0 /*. ^_ //
On: Oplismenus hirtellus (L.) Beauv., Pueblo Viejb,iVI'exico.
This fungus has not been seen by the writer and the only report
available is that of Fairman's type description (1913). The fungus was
collected in June, 1911, by the Rev. H. 0. Morton on a grass called
Carricera.
Range: Mexico.
Reference: Fairman (1913).
Septoria cenchrina J. J. Davis—Leaf Spot

Spots linear, gray, variable in size, often confluent, pycnidia am-
phigenous, brown-black, globose to depressed-globose to elliptical with
a thick wall which is often defective distally, 60-115 /^ diam.; spores
filiform, faintly 3- to 7-septate, straight or more or less curved, hya--"'
line, 30-100 (-125) X 1-5-3.0 /*.
On: Cenchrus myosuroides H. B. K., Tex.—C. pauciflorus Benth., Minn.,
N. Dak., Wise.
A collection made on August 14, 1946, at Mandan, North Dakota,
produced obscure, relatively pale pycnidia on yellowed leaves without
appreciable spot formation although some lesions were delimited by the
linearly arranged veins and stomata. Pure cultures on potato dextrose
agar produced light, cream-colored, juvenile colonies which became
carbonaceous, black, mounded growths with minute, distinct, dirty pink
spore masses exuding. The spores in pure culture were typically long
'and straight-sided without appreciable difference in dimension from
one end to the other. They were faintly 3- to 7-, mostly 5- to 7-septate
under critical illumination. The spores appeared opaque. Some were as
long as 125 X 2.4 /x. Davis (1924) spoke of the fact that the spores were
often so long that they extruded through the ostiole.
References: Davis (1924, p. 296); Sprague (1946,e; 1948,a).
Septoria cynodontis Fuckel—Leaf Spot

Pycnidia minute, seriately arranged in a black, thin, stroma, ostiole
small, papillate; pycnoipores filiform, hyaline, 50-65 X 1-5-2.0 /*.
On: Cynodon dactylon (L.) Pers., Tex.
This is listed from Texas (Weiss, 1945) but the fungus'has probably
not been critically studied. If this,is a true Septoria it is worthy of
detailed study because it was.named in 1863 and may involve synonymy
of later described fungi. It should be compared with the group on
Panicum, for; instance.
Range: United States, Germany, Italy, Persia, Spain.
References: Fuckel (1863; Symb. 1869, p. 389).
Septoria digitarivora Sprague.—Leaf Spot

Spots elliptical, finally diffuse, brown or fuscous, later isabelline in
the centers, pycnidia scattered, few to several to a lesion, seldom gre-
garious or two or three may be grouped, brown', parenchymatous, erum-
pent, subglobose to subellipsoidal, 110-128 )x, ostiole siiaall; pycnospores
clavate-filiform, apex acuminate, base sub-
obtuse, 3- to 7- often 5- to 7-septate, con-
tents somewhat coarse, hyaline, with yellow
or chlorine inclusions, 45-95 X 3.6-5.6 fx,, btit
mature spores are mostly 70-95 X 3.6-4.5 ju.
Mean spore size 84 X 4 ju, ratio of length to
width is 21 to 1.
On: Digitaria sanguinalis (L.) Scop., W. Va.
This species is known definitely only from
the type locality at Mineral Wells, West
Virginia, where it/ was collected by A. D.
Hopkins in 1943. '
Seymour (1929) reported S. graminum
Desm. on Digitaria sanguinalis and this is
probably from New Jersey (Ellis, N.A. Fungi
750) and Alabama (Weiss, 1944). Weiss
questioned the identity of S. graminum. Un-
derwood and Earle (1897) report a Duggar
collection as S. graminum on D. sanguinalis
from Alabama. In his routine discussion of
Septoria, Martin (1887) discusses the speci-
men from New Jersey (Elhs, N.A.F. 750) as
having hyaline, linear, flexuous, nonseptate
32. — Pycno- spores, 55-75 X 1-1-3;/A in pycnidia measur-
of Septoria ing 150 /t diameter, ilt appears likely that
tarivora on Digi-
taria sanguinalis, W. the New Jersey specimen, at4east, is very
Va. (type). different from the coarse-spored 'STliigitari-
vora (Fig. 32). Martin's description indicates
that N.A.F. 750 is closer to S. calamagrostidis than to S. graminum,
which has somewhat shorter spores (cf. Sprague", 1938,d).
Range: United States. Reference: S'prague (1946,a)'.
Septoria elymi Ell. and Ev.—Speckled Leaf Blotch '

S. agropyri Ell. and Ev.
/
Lesions pale gray to tan or fuscous with inoflerately prominent
pycnidia which are numerous enough to give a mottled effect to the
lesions; pycnidia flattened, ostiolate, brown, thin-walled, 95-200 X 60-
160 ju,; pycnophores ampulliform to subulate, 3.5 X 1.0-1.5 ;«;,pycno-
spores 0- to 3-septate, filiform bacillar, hyaline, 25-50 X 1.2-2.1 /j,.
On: Agropyron inerme (Scribn. and Sm.) Rydb., Oreg.—A. repens (L.)
Beauv., Alta., Iowa, Ky., Mich., Minn., Mont., Nebr., N. Dak., Ohio,
Oreg., Penn., S. Dak., Wash., Wise.—A. smithii Rydb., Mich., Nebr.,
N. Dak., S. Dak., Utah, Wise—A. spicatum (Pursh) Scribn. and Sm.,
Oreg., Wash., Wyo.—A. subsecundum (Lk.) Hitchc, Manit., Mont., N.
Dak.—A. trachycaulum (Lk.) Malte, Manit., Mont., Nebr., N. Dak.,
Sask., S. Dak., Wyo.—Elymits canadensis L., Minn., N. Dak., Okla.,
Ont., S. Dak., Wise.—E. canadensis var. rohustus (Scribn. and Sm.)
Mack, and Bush, Iowa, Mich., Wise.—E. glaucus BuckL, Mich., Oreg.,
Wash.—E. triticoides Buck!., Calif.—E. virginicus L., Wise.
S. elymi (synonym, S. agropyri) occurs over a large portion of the
United States where it is one of the most common spots on Agropyron
and Elymus spp. growing in thick or ungrazed areas or at the edge of
somewhat shaded patches adjacent to woodlands or thickets. It is read-
ily distinguished from the other species of Septoria on these hosts by its
relatively narrow, short spores (Fig. 33,A^,C,D,E,F,G,H). There is
one form-of it on Elymus canadensis rohustus in Iowa, Michigan, and
Wisconsin which has 2-septate, larger spores, 34-56 X 1.8-2.4 /t [S.
elymi f. elymina (J. J. Davis) Sprague]. The same form occurs on A.
smithii at times (Sprague, 1948,c). Greene recently found this form in
Wisconsin again (Fig. 33,7).
In pure culture S. elymi produces mucose colonies on potato dex-
trose agar. The following key based, in the main, on pure culture char-
acters, will aid in distinguishing S. elymi from associated pycnidial
forms on Elymus and Agropyron.
KEY TO SPECIES OF SEPTORIA ON AGROPYRON AND ELYMUS
A. Produces a mucose or slowly growing, noncottony colony in juvenile

growth on 2 per cent potato dextrose agar
B. Produces pycnidia in culture
C. Colony very slow growing, pale caenstone colored, later becoming
black, exuding white spore masses
S. pacifica Sprague
CC. Similar to S. pacifica but colony pelleted, gray color
S. infuscans (Ell. and Ev.) Sprague
BB. Produces mucose masses of conidia in juvenile growth
S.'elymi Ell. and Ev.
AA. Produces a buff or cottony, whitish growth in juvenile condition
B. Spores uniformly cylindrical, up to 60 M long
C. Spores mostly 26-42 X 2.8-3.5 M .
S. avenae f. sp. triticea T. Johnson
CC. Spores commonly 30-45 X 3.5-4.3 /*
Stagonospora arenaria Sacc.
BB. Spores uniformly cylindrical or slightly fusoid, mostly less than 30 M
long Septoria nodorum Berk.
BBB. Spores tapering abruptly at apex
S. agropyrina Lobik
/;
FIG. 33.—Seploria elymi: A, pycnospores on E.^glaucus var. jepsonii, Alsea

Mts., Oreg.; B, pycnospores with fragment of pycnidial wall and pyonophores
attached, on Agropyron repens, Madison, Wise; C, cf'oss section of pycnidial
wall on Agropyron spicatum near High Prairie, Wash.; D, pycnospores from
pure culture, isolated from A. inerme, Rowena, Oreg.; E, cross section of pycnid-
ial wall on Elymus glaucus, High Prairie, Wash.; F, pycnospores froin A. ciliare,
Chekiang, China; G, pycnospores from type of S. agropyri; H, ditto, type of
S. elymi; I, pycnospores of &. elymi i. elymina, on A. smithi^, Logan, Utah.
(Taken in part from Oregon State College Bot. Monogr^~and from Myoologia.)
Range: United States, Central Asia, China, France, Germany.
References: Davis (1924); Diedicke (1912,a); Ellis and Everhart (1892,a,
p. 132; 1893,a, p. 163); Petrak (1947); Sprague (1944,d; 1948,c); Weber
(1923).
Septoria glycericola Sprague—Buff Leaf Spot

Spots prominent, buff, semiorbicular, becoming paler in the center,
pyenidia scattered, golden brown, subglobose, slightly flattened, walls
6 to 7 /A thick, composed of rectangular cells which finally collapse to
a corky parenchymatous structure, pyenidia 90-140 fi diam., up to
90 /* tall; pycnophores obscure, absent or they consist of short papillae;
pycnospores narrowly cylindrical, somewhat pointed to blunt at the
apex, rounded at the base but definitely blunter than at the apex,
hyaline, 1- to 3-septate, 20-34 X 1.9-3.4 /*, mean size 25-26 X 2.8 /it.
On: Glyceria canadensis (Michx.) Trin., Mass.—G. elata (Nash) Hitchc,
Oreg.—G. grandis S. Wats., Colo., Mont., N. Dak.—G. pauciflora Presl,
Calif., Colo., Wash.—G. striata (Lam.) Hitchc, Ky., Oreg., Wash.
When this fungus was first described it was readily distinguished
from S. nodorum Berk, by the proportionately narrower spores with
their pointed ends (Fig. 34,^). Since collecting additional material'of
•f\
FIG. 34.—Pycnospores of: A, Septoria glycericola on Glyceria canadensis,

Dennis, Mass.; B, S. nodorum on Hystrix palula, Cotton Lake, Minn. (From
Myoologia.) - ' "
this and S. avenae, it becomes apparent that the problem is to distin-

guish between the latter and S. glycericola. S. avenae has more uni-
formly cylindrical, slightly broader spores and the lesions are some-
times darker. The fungi on GZi/ceria should be studied further in pure
culture and in overwintered cultures to determine perithecial affinities.
S. avenae appears to be the common form on Glyceria in the moun-
tains of the interior and S. glycericola is more prevalent nearer the
coasts.
Summer material of S. glycericola on plants growing in exposed
JIG, 35.—Pycuospores of Septoria infuscans; A, from Elymus triiicoides,

Warwick, "Wash. (O.S.C. 8378); B, same, with longer spores; C, from E. con-
areas has small black pycnidia (60-100 /x) and spores as small as
10-24 X 1.3-1.9 t,..
Reference; Sprague (1948,a).
Septoria infuscans (Ell. and Ev.) Sprague—Brown Leaf Blotch

Cylindrosporium infuscans Ell. and Ev.
Lesions fuscous, brown to gray, somewhat vague on their margins,
sometimes fading to white, especially in linear lesions or becoming
stramineous; pycnidia covered-cupuliform, subglobose (Fig. 48,P), -
elliptical in cross section, 70-110 X 75-200 jm, ostiolate; macropycno-
phores 3-8 X 1.5-3.5 /*, ampulliform, sometimes 1-septate; micropycno-
phores closely packed, narrowed, 8-10 X 1.0-1.5 ju,; macropycnospores
blunt at the base, tapering to a stubby point, stoutly clavate, 1-septate,
28-50-(60) X 2.1-4.5 fx. on Elymus condensatus, or spores narrowly
clavulate, stiffly curved to nearly straight, 1- to 4-septate, 40-75 X 1.7-
2.7 /I on E. triticoides, and on E. condensatus at times; micropycno-
spores nonseptate, small, 5-7 X 1.0-1.5 /t or smaller (4-7 X 0.7-1.5 /x),
narrowly ovate, short cylindric, straight or often curved.
On: Agro-pyron spicatum (Pursh) Scribn. and Sm., Oreg.—Elymus con-

^ densatus Presl, Calif., Colo., Ida., Mont., Oreg., Utah, Wash., Wyo.—
E. condensatus var. pubens Piper, Calif., Oreg.—E. glaucus Buckl, Oreg.,
Wash.—E. triticoides Buckl., Calif., Oreg., Wash.
There are twb kinds of macrospores involved in this species, not to
mention the spermatia-like microspores (Fig. 35, 36). I t was decided
after detailed study that the aberrant, coarse 1-septate macrospore on
E. condensatus (Fig. 35,^4), which unfortunately is the type, is the same
as the several-septate, somewhat more slender phase found widely dis-
tributed in the West (Fig. 35,F,G,I). The latter resembles the later-
described S. elymi-europaei Jaap (1907) (Fig. 43,C) (Fig. 48,11). The
reader is referred to a detailed discussion of this fungus in an earlier
article (Sprague, 1944,d/pp. 66-75). This species should prove to be an
interesting one for those studying variation in cultures from single
spores.
densatus, Yellowstone Natl. Park; D, from E. condensatus, near Warwick, Wash.

(O.S.C. 8392); E, microspores and malformed macrospores in pycnidia from
E. condensatus, near Biggs, Greg.; F, from E. triticoides. More, Greg.; G, same
host, Davis, Calif.; H, from pure culture isolated from E. condensatus, near
Biggs, Greg.; I, from E. triticoides, Klamath Falls, Greg. (O.S.C. 10,308); J,
micro- and macrospores and attached pycnophores from E. triticoides near
Wasco, Oreg. (G.S.C. 269); K, microspores from E. condensatus, Warwick,
Wash. (O.S.C. 8392); L, from E. condensatus, Logan, Utah (B.P.I. 80,048); M,
from E. condensatus, Kalispel, Mont., in linear faded white spots. (From Ore-
gon State College Bot. Monogr. 6, fig. 8.)
FIG. 36.—Septoria injuscans on Elymus condensatus, cross section of pycnid-

ial wall: A, inverted cross section of pycnidium (O.S.C. 8392) showing thicken-
ing formed inside wall adjacent to ostiolum (to right) developed during micro-
spore formation, normal wall and pycnophore development on left; B, showing
normal macropycnophore development at base of a pycnidium; C, showing de-
velopment of micropycnophores and microspores; D, macropycnophores from
Yellowstone Natl. Park. (From Oregon State College Bot. Monogr. 6.)
/
I t should be mentioned in passing t h a t this fungus is n o t a Cylindro-
sporium b u t has a good pycnidium (Fig. 48,-K).
R a n g e : United States and possibly across Asi^ and Europe to the moun-
tains of Switzerland. -. ^
References: Cooke, W. B. (1940,a, No. 13); Ellis and ^ v e r h a r t ( 1902);
Jaap (1907); Sprague (1940,c; 1944,d). '_
Septoria jaculella Sprague—Western Brome Speckle
Lesions vague, chlorotic, gray to isabelline 'patches on t h e leaves

with t h e prominent black pycnidia forming in t h e still-green tissue of
the leaf, mârgins yellow to indefinite; infected p a r t s eventually die
(these areas are sometimes long streaks involving half of the entire leaf
w i d t h ) ; pycnidia carbonaceous and suberumpent-(Fig./48,A,G), walls
4 to 20 ju, thick, or even thinner t h a n 4 /<, in s6me specimens, ostioles
prominent; pycnophores variable, from prominent bottle-shaped struc-
tures t o short nipple-like ones, mostly 4.8-7.0 X 1.6-2.4 /A or absent
(Fig. 37,C,D,E,F,G,H); pycnospores lance-like, straight, obclavate-
filiform, hyaline (diamond blue), 2- to 5-septate, 45-90 X 1.9-4.0 ju
(55-75 X 2.1-3.0 /j.); micropycnospores bacillar, 2-7 X 0.4-0.6 [i (Fig.
37,L).
On: Bromus carinatus Hook, and Arn., Colo., Ida., Oreg., Wash.—B. ciliat-us
L., Ariz., Mont.—B. laevipes Shear, Calif.—B. rigidus Roth, Oreg., Wash.
—B. tectorum L., Wash.
This fungus appears to be almost an obligate parasite. I t lies in
close association with the living host cells on vigorously growing leaves.
They die only after a considerable length of time, often some time after
the fungus has produced spores from pycnidia growing adjacent t o '
living cells. The fungus has resisted all attempts to induce it to grow
in pure culture (Sprague, 1944,d). The spores germinate, sending out
irregular, narrow mycelium which dies after the nutrient from the
spore has been exhausted. The stiffly lance-like spores are characteristic
of this_ species (Fig. 37,A,B,I,J,K).
This species was described in 1940 but was collected, inadvertently,
by David Douglas 108 years before that date in "Nova California,"
probably in the vicinity of Roseberg, Oregon. It is therefore a native,
if not an invader in prehistoric times from Asia. While this fungus is
common only west of the Cascade Mountains it occurs east to Colo-
rado, Arizona, and Montana.
References: Cooke (1940,6, No. 114); Sprague (1940,d; 1944,^; 1949,a).
Septoria loligena Sprague—Brown Leaf Spot

Spots fuscous to deep chocolate brown, somewhat paler in the
center, with borders surrounded by lighter areas. Pycnidia light golden
brown to brown, walls thin, subglobose, not prominent, 80-120 /x diam.;
pycnospores bacillar, clear hyaline, 0- to 3-septate, 25-50 X 2.7-4.2 n.
On: Lolium multiflorum Lam., Calif.
S. loligena, S. avefiae, S. melicae and possibly S. secalis and even
Stagonospora arenaria compose a group which probably sooner or later
should be placed under one species. As mentioned earlier, study of pos-
sible perithecial stages is needed beyond what we already know about
this group. (S. loligena was segregated- because of the inadequate knowl-
edge of the whole series and beca:use it appeared to represent a distinct
species when compared with the other species on Lolium and associated
grasses.
Range: United States, France.
References: Castagne (1851); Sprague (1944,rf; 1946,e).
FIG. 37.—ArL, Septoria jaculella; A, pycnospores from Bromus^ carinatus,

Pacific Grovej,Calif.; B, pycnospores from B. carinatus, Fox Valley; Oreg.; C,
crofs section of'pycnidial wall, same collection ^s just mentioned; D, cross sec-
FUNGI IMPERFECTI-SPHAEROPSIDALES 241
Septoria macropoda P a s s - -Leaf Blotch

S. annua Ell. and Ev.
S. poae-annuae Bres.
Spots elliptical, at first mottled green, later brown or straw-colored,

finally bleached, off-white; pycnidia subepidermal, slightly erumpent,
mostly flattened and appressed to the upper leaf epidermis, sometimes
subglobose, walls thin (5-11 fi), 60-135 X 66-160 fi diam., 50-80 ti high;
pycnophores subulate to subcylindric (Fig. 38,B,C,E,F), 4-7 X 1.5-2.5
[i; pycnospores filiform, 30-40 X 1.0-1.5 [i (Fig. 38,A,D).
FIG. 38.—Septoria macropoda: A, pycnospores from type of S. annua; B, cross

section of pycnidial wall on Poa annua, Black Earth, Wise.; C, cross section
through very thin pycnidial wall on P. annua, Corvallis, Oreg. (O.S.C. 10,354);
D, pycnospores from P. kelloggii, Union Creek, Oreg.; E, cross section of pyc-
nidial wall and pycnospores on P. annua, Corvallis, showing thicker walls; F,
cross section of another pycnidium with thin walls, Corvallis, Oreg., but some-
what intermediate between- C and E. (From Oregon State College Bot. Monogr.
6,fig.4.)
tion of pycnidial wall, on B. carinatus, Astoria, Oreg., with resinous products

between pycnidial wall and host cells; E, cross section of pj^cnidial wall, on
B. carinatus. Fossil, Oreg.; F, cross section of pycnidial wall on B. carinatus,
Corvalhs, Oreg. The wall in this early spring material is composed of large
cells, while the pycnophores are poorly developed and scarcely differentiated
from the hyphal layer. In some cases the spores grow directly from the pyc-
nidial wall; G, cross section of pycnidial wall from B. rigidus (O.S.C. 8070); H,
cross section of pycnidial wall from B. tectorum, Lyle, Wash.; I, pycnospores
from type; J, pycnospores from B. ciliatus, Bradshaw Mts., Ariz.; K, pycnospores
from B. rigidus, near Lyle, Wash. (O.S.C. 8070); L, microspores from B. cari-
natus. Fox Valley, Oreg.; M, pycnospores found in type of S. bromivora; N,
cross section of pycnidial wall of S. brevispora (type); O, Septoria bromi on B.
latiglumis (B. altissimus), J. J. Davis, Bruce, Wise. (From Oregon State College
Bot. Monogr. 6, fig. 18.)
On: Poa annua L., Oreg., Wash., Wise.—P. kelloggii :Vasey, Oreg.
S. macropoda produces a slow-growing mucose, flesh pink, later
carbonaceous colony on 2 per cent potato-dextrose agar. It is widely
distributed.
Range: United States, China, Germany, Morocco.
References: Bresadola (1900); Cocconi (1896-97); I Ellis and Everhart
(1900); Gilman and Archer (1929); Sprague (1944,d).
Septoria macropoda var. grandis Sprague—Leaf Spot

Lesions deep brown to gray, later fading, pycnidia often large but
averaging 65-190 /* diam., 40-90 /^ tall, pycnidia subepidermal, brown,
somewhat flattened at the apex, wall 7-14 /x thick, mostly 9-11 //.; pycno-
phores ampulliform, subulate, or subcylindric, 3-6 X 1.3-2.3 /i; spores
filiform to very narrowly filiform-clavate, 1- to 3-septate, 50-70 X 1-4-
2.4 [i in winter, 40-60 X 1.4-1.9 /x in spring. ,
On: Poa ampla Merr., AVash.—P. arida Vasey; S. Dak.—P. canbyi (Scribn.)
Piper, Mont., S. Dak., Wash.—P. cusickii /Vasey, Mont., Oreg.—P. gracil-
lima Vasey, Wash.—P. nervosa (Hook.) Vasey, Mont.—P. nevadensis
Vasey, Calif., Oreg.—P. pratensis L., Wyo.—P. scabrella.[Thnrh.) Benth.,
Wash.—P. secunda Presl, Ida., Mont., N.
Dak., Oreg., S. Dak., Wash., Wyo.—P.
vaseyochloa Scribn., Wash.
This variety is apparently native oii
the western prairies and range lands. The
spores are larger than those of var. sep-
tulata and are sometimes curved or sinu-
ous from crowding in the pycnidium. This
variety approaches S. tritici in size of
spore and general morphology of the pyc-
nidium (Fig. 39, 48,B). At least three
races of this variety are known (Sprague,
1944,d). • ---^_^
One specimen from Wyoming~on Poa
EiG. 39.—Septoria mac- pratensis appears to be var. grandis rather
ropoda var. grandis, pycno- than var. septulata, the common one on this
spores on Poa secunda, near grass.
Warwick, Wash., and cross
section of pycnidial wall on Range: United States./ '
Poa ampla, Lyle, Wash.
References: Sprague (1942,6; lMi,d; 1946,e).
Septoria macropoda var. septulata (Gonz. Frag.) Sprague—

Leaf Blotch ^— /
Lesions elliptical to elongate, dark gray to brown, up to 3 mm. long,
margins red, maroon, or yellow, pycnidia obscure, at least until the
lesions fade to straw color; pycnidia subepidermal, slightly erumpent.
FUNGI IMPERFECT!—SPHAEROPSIDALES 243
variable, from somewhat flattened globose to true globose, compact,
wall 5-9 /t thick, dark brown, 60-180 X 70-240 /*, commonly about 160
fj. diam., 50-95 m tall; pycnophores elongate, flask-shaped to nearly fili-
form, 3-6 X 1.5-2.0 /x, spores filiform, sharp pointed (needle-like), cross
walls obscure, typically the spores measure 40-60 X 1.3-1.7 /A.
On: Poa compressa L., Ida., N. Dak., Oreg., S. Dak.—P. nervosa (Hook.)
Vasey, Wash.—P. pratensis L., Ida., Iowa, Ky., Mich., Minn., Mont.,
Nebr., N. Dak., Ohio, Oreg., Penn., S. Dak., Tenn., Va., Wash., Wise,
Wye.
This variety is readily recognized by the needle-like spores which
are longer and stiffer appearing than those of Poa annua. It appears to'^
be largely restricted to P. pratensis but we sometimes have encountered
it on P. compressa and once on P. nervosa. The material on P. com-
pressa from near Rainier, Oregon, was weakly parasitic and repre-
sented a distinct race from the common one on P. pratensis. The spot
on bluegrasses is common during rainy weather in the summer and iaJl,
and sometimes in open winters in mild climates. It is especially com-
mon on ungrazed pastures or along roadsides at the edge of thickets.
Range:-United States, Spain, probably widely distributed.
References: Gonzalez Fragoso (1920, 1926); Sprague (1944,c;; 1946,e);
Weber (1923).
\^
Septoria melicae Pass.—Leaf Blotch
Spots elliptical, white with brown margins; pycnidia yellow brown,
subglobose, 80-120 ft diam.; pycnospores bacillar, 22-38 X 2.8-3.8 /*,
1- to 3-septate.
On: Melica scabrosa Trin., N. Dak. (in plots).
This fungus is very similar to S. avenue Frank. The only other
collection on Melica which might be said to resemble the Mandan col-
lection is one oniW". califomica Scribn. collected by W. B. Cooke on Mt.
Shasta, California. This was assigned to S. nodorum Berk. (Sprague,
1948,c). The spores in-this material were 1-septate, subcylindrical,
17-25x2.8-3.5^1 (Fig. 27,C).
Guyot (1946) described S. allorgei on Melica ciliata from France
with small pycnidia and cylindrical somewhat tapering 3-septate. spores,
10-23 X 2.5-3.0 ft.
Range: United States, Denmark, Italy.
References: Frandsen (1943); Sprague (1946,a,e; 1948,c).
Septoria mississippiensis Sprague—Leaf Spot

Spots orbicular to elongate, buff with brown margins, pycnidia few,
subgregarious, subcarbonaceous-parenchymatous, erumpent, ostiolate.
black brown, 100-113 /t; pycnospores hyaline to. yellowish, clavulate-
filiform, apices subacute, bases obtuse; 3-septate, guttulate, 35-
65 X 2.1-3.1;«, (Fig. 25,C). ~ '
I
On: Muhlenbergia asperifolia (Nees and Mey.) Par., H. Dak.—M. mexicana
(L.) Trin., Minn. (type). ''
This distinct, but apparently rare species, causes well defined buff
spots with prominent brown borders on the sheaths and leaves.
References: Sprague (1943,c; 1946,e).
Septoria munroae Ell. and Barth.—

Leaf Spot
Pycnidia epiphyllous, seriately arranged,
punctiform, flattened-cupuliform to subspheri-
cal, 95-100 /x wide, 72-108 /x, tall, walls light
golden brown, 11-14 /A thick, hyphae 2 /x thick,
ramifying into adjacent host tissue, pycno-
phores cylindrical, 5-9.5 X 1.5-2.1 /j.; pycno-
spores, 80-110 X 2.5-3.0, hyaline, oblong-flli-
form-clavulate, 1- to 3-septate (Pig. 40).
On: Munroa squarrosa (Nutt.) Terr., Colo., Kans.,
Okla. ,
The pycnidia are prominent -against the
faded straw-colored leaves of the host in the
dried material which we have seen. The host
FIG. 40. — Septoria range of this species has'not been studied.
munroae, cross section of
pycnidial wall, Yuma, Range: United States.
Colo. (From Oregon
State College Bot. References: Ellis and Bartholomew (1902);
Monogr. 6.) Sprague (1944,d; 1946,e).
Septoria nodorum (Berk.) Berk.—Glume Blotch_

Depazea nodorum Berk.
Septoria glumarum Pass.
Phoma hennebergii Kuehn
Hendersonia nodorum (Berk.) Petra;k '
Discolored to brown lesions on the glumes, leaves, and culms; pyc-
nidia scattered, sometimes seriately arranged, spherical, flattened, or
elongate, subepidermal, 160-210 /j. diam., wall thin, soft, and paren-
chymatous, at first light brown, later becoming^'Sark brown or black,
ostiolate; spores short, cylindrical, 0- to 3-septate, 15-32 X 2-4 ;«,.
On,: Aqropyron re-pens (L.) Beauv., N. Dak.—A. smithii Rydb., N. Dak.—
A. spicatum (Pursh) Scribn. and Sm., Wash.—A. trachycaulum (Lk.)
Malte, N. Dak.—Cinna latifolia (Trev.) Griseb., Minn.—Deschampsia
atropurpurea (Wahl.) Soheele, Wash.—D. elongata (Hook.) Munro, Wash.
—Elymus canadensis L., Manit., Mich.—E. excelsus Turcz., N. Dak.—
E. giganteus Vahl, Manit.—E. glaucus Buokl., Wash.—E. junceus Fisch.,
N. Dak.—E. sibiricus L., N. Dak.—Festuca obtusa Spreng., Wise.—
F. subulata Trin., Wash.—Glyceria striata (Lam.) Hitchc, Oreg., Wash.—
Hordeum jubatum L., Alta., Manit., N. Dak.—H. nodosum L., Alta., Ida.
—H. vulgare L., Manit.—Hystrix patida Moench, Minn.—Melica bulbosa
Geyer, Utah—M. californica Scribn., Calif.—Poa palustrxs L., S. Dak.—
P. pratensis L., Oreg.—Secale cereale L., Ark., Ind., Md., Ohio, Wise.—
Stipa viridula Trin., N. Dak.—Triticum aestivum L., Alta., Ark., Del.,
Ga., 111., Ind., Iowa, Kans., Ky., Manit., Md., Minn., Mo., Mont., Nebr.,
N. B., N. J., N. Y., N. Car., N. Dak., N. S., Ohio, Okla., Ont., Penn.,
P. E. Is., Que., Sask., S. Dak., Tenn., Va., Wash., W. Va., Wise—r. dicoc-
cum Schrank, N. Dak.—T. spelta L., Minn., Oreg.
S. nodorum has relatively small spores (Fig. 41) which are short-
cylindric or sometimes slightly wedge-shaped. They are definitely
smaller than those of S. avenae. Cer-
tain material on Agropyron, Elymu,s,
and Triticum with spores 25-45 X
2.5-4.3 ju, has been discussed under
S. avenae f. sp. triticea and will not
be considered here.
S. nodorum is now generally
known as a species of Septoria al-
FIG. 41.—Pycnospores of Sep-
though the spores are short-cylin- toria nodorum.: A, on Triticum
drical and technically belong close spelta, Alsea Valley, Greg.; B, on
to Stagonospora. Recently Petrak Elymus virginicus, Lansing, Mich.
(1947,b) placed this fungus in Hen-
dersonia. In order to justify placing this species in the genus Hender-
sonia he must have referred Hendersonia back to the pre-Saccardian
concept.
S. nodorum is apparently more prevalent on wild grasses than
earlier information had indicated (Fig. 34,B). For instance, in 1947
the writer found a common blight or brown leaf scald of a woodland
species, Festuca subulata Trin., in the Chelan National Forest, Wash-
ington, which seems to be only a form of S. nodorum (Sprague, 1948,c).
The spores were 21-30 X 3.1-4.8 /*, which is larger than most collections
of this species but probably too small for S. avenae. He found typical
spores of S. nodorum on Elymus glaucus Buckl. during the same year
and material on Melica californica assignable to the species. A speci-
men on Hordeum nodosum in Idaho had bacillar-shaped, 1-septate
spores 15-23.6 X 2.4-3.0 ju. wHich we also place in S. nodorum (1948,c).
Material on Festuca elatibr had spores approximately the same size as
those of S. nodorum on F. subulata but the material from the former,
collected in Utah, was placed in S. avenae because of the cylindrical
aspect of the spores. S. nodorum is rare in the West because the humid-
ity in summer is low in most of the wheat-growing regions. Glume
/'
blotch on wheat was not found in Washington until !l947 in the western
part of the state. - -
Stagonospora graminicola Ishiyama (1916) was 'described on wheat.
We have not obtained a description of .this species and do not know
whether it is similar to S. nodorum or not.
Range: United States, Argentina, Australia, Austria, Cahada, China, Czecho-
slovakia, Denmark, England, France, Germany, Italy, Kenya, Nether-
lands, Norway, Portugal, Spain, Sweden, Switzerland, Tanganyika,
Uruguay.
References: Alfaro (1940); Bensaude (1929); Berkeley (1845); Berlese
and Voglino (1886); Bockmann (1932); Diedicke (1915); Frandsen
(1943); Grove (1916, 1935); Kuehn (1878); Luchetti (1939),; Machacek
(1945); Marchionatto (1941); Passerini (1877); Petrak (1947,6);
Petrak and Sydow (1925); Rosella (1929); Rosen (1921; 1947,6);
Sprague (1944,d; 1946,e; 1948,a,c; 1950,a).
Septoria oryzae Catt.-Ôn Rice

Pycnidia innate, prominent, numerous,'black, aggregated, ostiolate,
between veins of the host in seried arrangement; spores small, cylin-
drical, straight to curved, subhyaline, S^septate, 21 X S-fi. •
On: Oryza sativa L., Fla.
This species has been reported from Florida. A form of it, S. oryzae
var. braziliensis Speg. (1908) has yellow 2- to 5-septate spores 20-
30 X 3 /x. Phaeoseptoria oryzae Miyake (1910) has flavid, 4- to 6-sep-'
.tate spores 30-45 X 2.5-3.0 /J,. Perhaps these are all related. Since it has
been our experience that the yellow- or brown-spored Phaeoseptoria
and Hendersonia are dubiously parasitic it is likely that S. oryzae is
not an important parasite on this important host.
A number of species of Septoria are described on rice. S. poae (Lib.)
Catt. (Cattaneo, 1879) has strongly curved Ifiliform spores 50 ju. long.
»S. curvula Miyake (1910) has 5- to 8-septate curved spores SO-
SO X 2.5-3.0 IX. S. longispora Miyake (1910) which bectsmes S. miyakei
Sacc. and Trav. (Syll. Fung. V. 22, pp. 1120-1121) hasônseptate
spores with rounded or truncated apices, 30-40 X 2.5-3.2 n, which some-
what belies the "longispora" name. Septoria oryzaeicola Hara is listed
as new on rice from Japan (Shirai, 1927).
Range: United States, Brazil, Italy. '
References: Cattaneo (1879); Miyake (1910); âccardo (Sylloge V. 3,
p. 562; V. 22, p. 1120); Shirai (1927); Spegazzini (1908). '
/
Septoria oudemansii Sacc.—LeafrSpot /
Lesions light purple, finally straw-colored, mostly confined to dis-
tal portions of leaves, obscure; pycnidia yellow brown, subglobose,
somewhat erumpent, outer wall composed of a yellow brown layer 6-7 î
thick, surrounding a hyaline layer 4-4.5 jn thick, all cells of wall with
solid contents (Fig. 48,7); pycnophores bulbous,-4-5 X 1-8-2.2 /<.; ma-
crospores cylindrical but slightly thicker at the usual central septum,
other spores tapered to nearly lanceolate, contents bright hyaline with
small uniform oil drops adjacent to the septum, 1-septate macrospores
12-24 X 1.7-2.8 IX, 3-septate material at Mandan, North Dakota, with
spores 35-55 X 3-4 jx mingled with spores 11-18 X 2.5 /*. Microspores
sometimes present, ranging in size from those as small as 4 X 2 /x to
ones nearly as large as the macrospores.
On: Poa ampla Merr., Mont., N. Dak.—P. arachnifera Torr., N. Dak., in
plots—P. canbyi (Scribn.) Piper, Mont., N.'Dak.—P. compressa L., N.
Dak., Oreg.—P. juncifolia Soribn., N. Dak., in plots—P. palmtris L.,
Colo., N. Dak.—P. pratensis L., Calif., Colo., Ida., Iowa, Ky., Mich.,
Minn., Mont., Nebr., N. Dak., Oreg., Utah, Wash., Wyo.—P. reflexa
Vasey and Scribn., Colo.—P. secunda Presl, Ida., N. Dak., Wash., Wyo.
This fungus is scattered widely but is seldom serious. It was preva-
lent at Mandan, North. Dakota, on native and introduced Poa spp,,
however. In western Oregon and Washington it occurs during the rainy
periods in winter and early spring. Griffith (1943) indicated that the
disease- .was important on leaves of P. pra-
tensis in Kentucky during the cool wet
season. Seymour (1929) lists S. oudemansii
on P. /lexuoso var. elongata I^lytt and on
Hierochloa alpina (Sw.) Roem. and Schult.,
probably from northeast North America. The
first record from this country appears to be
a collection made by the writer in Klickitat
County, Washington, in February, 1935, al- of FIG. 42.—Pyonospores
Septoria oudemansii
though he had the fungus in pure culture as on : A, Poa compressa,
early as 1932 but did not determine its iden- Wild Rice, N. Dak.; B,
tity until later. In 1935 he reported it on P. pratensis, Ft. Lewis,
Wash.
P. pratensis in two places in western Oregon
as Ascophyta sp. (Sprague, 1935,c) and in 1942 corrected this to Sep-
toria oudemansii (Sprague, 1942,a,6). He also found it on material sent
to him from Michigan lor determination (Hardison, 1942). Except for
the 3-septate material from Mandan, above mentioned, this species is
close to Ascochyta (Fig. 42).
Range: United States, Canada, Denmark, Greenland, Netherlandsr'
References: AUescher (1901); Fischer, Sprague, Johnson, and Hardison
(1942); Griffith (1943); Hardison'(1942); Sprague (1935,c; 1942,a,M;
1946,e; 1949,o).
Septoria pacifica Sprague—Leaf Blotch

Lesions fuscous, soon straw-colored; pycnidia prominent in seried
arrangement on the faded lesions, pycnidia nearly spherical to some-
EiG.. 43.—Sepiom elymi-europaei, S. pacifica and S. agropyrina compared;

A, liyctiogpores of S. pacifica from Elymus mollis, near Newport, Or'eg. (O.S.C.
what flask-shaped, erumpent but deep-seated, tardily ostiolate, walls
heavy, parenchymatous to pseudocorky, 110-150 ix; pycnophores nar-
rowly ampulliform, hyaline; pycnospores strongly 1- to 8-septate, fili-
form, 68-110 X 1.6-3.2 II (winter), 60-85 X 1.5-2.5 /* (spring).
On: Elymus condensatus var. pubens Piper, Calif.—E. mollis Trin., Oreg.
S. pacifica has distinctly longer and slightly narrower spores (Fig.
4S,A,B) than S. elymi-europaei Jaap and the pycnidial walls are
thicker (Fig. 43,D). As mentioned in another publication (1944,d,
p. 79), S. -pacifica is distinct from S. ammophilae H. and P. Sydow and
S. arenaria Rostr. The last mentioned has extremely narrow spores,
60-100 X 0.5-1.0 /i. Frandsen illustrated and discussed S. arenaria
(1943, Fig. 3,/; pp. 57, 60, 61). The spores in his drawings are nonsep-
tate.
In comparing S. pacifica with S. agropyrina it may be noted that
the latter has distinctly broader spores than the former (Fig.
A3,A,B,E,F,G,H,I,J).
S. pacifica appears to be confined to the coastal area. It is related
to the S. injuscans complex. In culture it produces pale caenstone-
colored, pellet-shaped colonies which finally become covered with black
carbonaceous hyphae. ,
References: Sprague (1944,d; 1946,e).
\^
Septoria passerinii Sacc..—Barley Leaf Speckle
' S. murina Pass, (non v. Thuem.), 1879
Lesions indefinite, yellowed, finally fuscous; pycnidia subepider-
mal, creosote brown, mostly subglobose (Fig. 48,-D), scarcely erumpent,
90-150 IX, diam. and up to 100 [i deep, walls relatively thin, 5 to 10 /*,
composed of strongly pigmented cells consisting of an outer layer of
crushed, intertwined, oblong cells and an inner layer of polyhedral
cells; pycnophores are somewhat filiform to subulate, 3.4-5 X 1.2-1.8 p.,
mostly on the lower hemisphere of the pycnidium (Fig. 44,5,C); ma-
crospores broadly filiform, obtusely pointed at both ends, rarely curved,
1- to 3-septate, 23-46 X 1.7-3.0 ft (Fig. 44,A), microspores 3-6 X 0.3-
0.6 p., bacteria-like (Fig. 48,M).
On: Hordeum brevisuhulatum (Trin.) Lk., N. Dak.—H. distichon li-., Wyo.
—H. jubatum L., Iowa, Minn., N. Dak., S. Dak., Wise.—H. nodosum L.,
36); B, pycnospores of S. pacifica' from E. mollis, near Waldport, Oreg. (O.S.C.

421); C, pycnospores of S. elymi-europaei from Elymus europaeus, Bavaria; D,
cross section of pycnidial wall of S. pacifica on E. mollis, Waldport, Oreg.; E,
pycnospores of S. agropyrina from E. virginicus, Starkville, Miss.; F, pycno-
spores of S. agropyrina from Agropyron smithii, Lincoln, Nebr.; G, S. agropy-
rina from Agropyron smithii, Forsythe, Mont.; H, S. agropyrina from A. repens,
Vermillion, S. Dak.; I, S. agropyrina from A. trachycaulum, Tagus, N. Dak.; J,
S. agropyrina from Elymus canadensis. (From Oregon State College Bot.
Monogr. 6.)
Mont.—H. vulgare L., Alta., Ark., B. C, Ida., lUl, Ind., Iowa, Manit.,
Mich., Minn., N. Car., N. Dak., Ohio,-Okla., Ôntj., Oreg., Penn., Que.,
Sask., S. Car., S. Dak., Va., W. Va., Wise.—Hystrixl patula Moench, Ind.,
Minn.—Sitanion hystrix (Nutt.) J. G. Sm., Oreg.
This species is not important on barley in the Ear West but is some-
times common on the cultivated host in the Middle West (Weber,
1923) and in Canada. S. passerinii is, however, very common on H.
jubatum in the northern Great Plains. It was recently shown that S.
FIG. 44.—Septoria passerinii: A, pycnospores, including, microspores on Hor-

deum nodosum, Lewisburg, Greg.; B, cross section of pycnidial wall on ditto; C,
same collection but very thin-walled pycnidium; D, type of S. microspora,
Indiana; E, conidia on potato-dextrose agar isolated from Sitanion hystrix, Mt.
Hood, Oreg.; F[ on Hystrix patula. Cotton Lake, Minn. (From Mycologia, in
part.) ^
passerinii occurs on other grasses such as Sitanion hystrix (Sprague,
1944,d) (Fig. 44,E; Fig. 48,S) and Hystrix patula (Ellis, 1893;
Sprague, 1948,a) (Fig. 44,F). Microsporous stages of this species are
especially confusing (Sprague, 1944,d; 1946,e; 1948,a) (Fig. 44,D).
Range: United States, Australia, Bulgaria, Canada, Cyprus, Denmark, Libya,
Romania.
References: Ellis (1893); Frandsen (1943); Johnson, A. G. (1916); Pas-
serini (1879); Saccardo and Trotter (1913); Sprague (1944,rf; 1946,e;
1948,a); Weber (1923).
Septoria pertusa Heald and Wolf—^Leaf Spot

Amphigenous, spots large, elongated parallel to the veins, 1-2 cm.
long, center of lesion brown which is surrounded by a yellowish margin
or zone which pales into the green of the leaf; by confluence entire leaf
may become dry and yellowish brown; pycnidia innate, flask-shaped,
fuscous, protruding by a short papilla, spores hyaline, straight to
curved, som'ewhat clavulate, abundant, 60-75 X 3 ju.
On: Sorghum halepeme L., Tex.
This leaf spot on Johnson grass was described by Heald and Wolf
from Texas (1911). A¥eiss gives only Texas in his list (1945), indi-'
eating that the fungus is either restricted in its range, rare, or little
observed. The writer failed to find 'the fungus in the Rio Grande Val-
ley, New Mexico, in 1947. The host was abundant along irrigation
ditches but the season was very dry.
Bever (1949) found Septoria sp. prevalent on broomcorn {Sorghum
vulgare) in Illinois in July, 1949. Lesions measured 0.5-2.0 X 0.125-1.0
inches. The lesions were brownish gray with a yellowish halo. Pycno-
spores were 3- to 7-septate. Koehler and Woodworth (1949) also re-
port the blight from Illinois and mention that it had been confused
with Helminthosporium turcicum. Unofficial recent reports indicate that
Ramulispora was involved in the Illinois problem.
Range: United States, China.
References: Heald andjôlf (1911); Porter (1926).
Septoria poliomela Sydow—Leaf Spot

Lesions indefinite, tan to straw-colored, mostly on the lower leaves,
pycnidia prominent, brown, somewhat gregarious, often coalesced,
strongly erumpent,''ostioles circular, bordered by brown oxidized cells,
parenchymatous, somewhat loosely woven, 40-160 X 140-200 fx,; pycno-
phores, 3.2-4.2 X 0.9-1.1 /A (Fig. 45,D), spores clear hyaline, strongly
curved, 36-50x1-4-1.6-2.0 /x, 1- to 3-septate (Fig. 45,A,C), curved,
somewhat needle-like, tapering towards each end or slightly blunt at
one end.
252 DISEASES OF CEREALS AND GRASSfiS
On: Deschampsia caespitosa (L.) Beauv., Oreg.—Di danthonioides (Trin.)
Munro, Oreg., Wash. .
A few collections of this species were made west of the Cascade
Mountains or in foothills in Oregon and Washirigton. While the ma-
terial on D. caespitosa is somewhat different fromlthat oh D. danthoni-
oides the fungus on both hosts was referred by jSpragu'e (1944,d) to
S. poliomela (Sydow, 1937). This species was described from Madeira
FIG. 45.—Septoria poliomela: A, pycnospores from Deschampsia danthoni-

oides under oaks and firs, near Peoria, Oreg. (O.S.C. 8491); B, pycnospores from
Aira caryophyllea (type); C, pycnospores from D. danthonioides, High Prairie,
Wash.; D, cross section of pycnidial wall on D. danthonioides, near High Prairie,
Wash., under ponderosa pines (O.S.C. 8130). (From Oregon State College Bot.
Monogr. 6.) i
material collected on Aira caryophyllea. The spores were mostly 20-30

X 1.5-2.2 n (Fig. 4:5,B), borne in prominent pycnidia, 80-130 jj, which
occur in straw-colored'lesions. ^~'~~'^_
Isolations obtained from D. danthonioides produced a white to'pale
cream color, slow-growing, mounded, yeasty colony on potato dextrose
agar which soon became covered with shoTt-nap, velvety, gray my-
celium, similar to S. elymi (Sprague, 1944,d).
Range: United States, Madeira Is., Morooco (cf. Maire and Werner, 1938),
References: Maire and Werner (1938); Sprague (1942,a,6; 194;i,d; 1946,e);
Svdow (1937).
^- ^ /
Septoria quinqueseptata Spragu^^Leaf Spot
Pycnidia few, somewhat in lines, widely spaced, erumpent, globose
or sUbglobose, brown, black about the ostiole which is small and late
to open; pycnospores filiform, blunt at the base, subacute at the apex,
5-septate, hyaline, straight or stifHy curved, 45-58 X 1-9-2.4 /x.
On: Koeleria cristata (L.) Pers., N. Dak., Wash.—Sphenopholis obtusata
(Michx.) Scribn., N. Dak.
The type of this species has 5-septate spores (Fig. 46,A) which is
an unusual number to expect in a genus which normally has 1-3-7 cross
walls. ;
FIG. 46.—Pycnospores of: A, Septoria quinqueseptata, type; B, S. quinque-

septata isolated from Koeleria cristata (a) compared with S. calamagrostidis f.
koeleriae (b); C, S. quinqueseptata on Koeleria, Mandan, N. Dak. (From My-
cologia.)
This species is not" well known. The type is a fragment, while the
collections on Koeleria (1948,a) appear to be occasional developments
on this host (Fig. 46,B,C), which is usually attacked by S. calama-
grostidis f. koeleriae. Perhaps these collections represent variants of
*S. andropogonis; but at the time that S. quinqueseptata was described,
and before the polymorphic nature of S. andropogonis was apparent,
it was very distinct from known forms.
References: Sprague (1944,d; 1946,a,e; 1948,a).
Septoria secalis Prill, and Del.—Leaf Blotch

Phoma secalina Jancz.
Septoria s^caliha (Jancz.) Sacc.
Leaf lesions yellow brown, pycnidia brown, walls composed of
layers of light brown, rectangular cells which give rise to creeping
hyphae terminating in prominent subulate to nearly fusiform pycno-
phores, 5.5-8 X 1.9-2.2 y, (Fig. 47,B); pycnospores hyaline, 3-septate,
narrowly cylindrical, 25-50 X 2.0-3.5 /i, mostly 35 'x 2.7 ii, microspores
mostly bacillar-shaped, 10 X 0.5 /x. -- , ,
On: Secale cereale L., Alabama, Alta., D. C, III, Iowa, Minn., N. Car.,
N. Dak., Que., Sask., S. Dak., Tenn., Tex., Va., W. yk.. Wise.
The disease is sometimes prevalent in the rye-grpwing areas of the
Middle West and in the northern Great Plains, but it seldom does any
appreciable damage because it is confined mainly to basal culm leaves
in late season. It has not yet been found in the Far West, probably
because rye is usually sown on semiarid lands in this region.
Range: United States, France, Germany, Poland.
References: Janczewski (1894); Sprague (1944,d; 1946,e); Weber (1923).
Septoria secalis var. stipae Sprague—Leaf Spot

Lesions at first dark brown, circular, soon^pale bufi' in center, finally
white to grayish white and elongate; pycnidia prominent, black, glo-
bose, 130-190 ju, ostiolate, erumpent; pycnbspores sometimes straight
but usually slightly and stiffly curved, broadly filiform with roundeH
and only slightly tapering ends, 3-septate with 2, 4, of several small
oil drops on either side of the cross walls, rarely slightly constricted
at the septa, 37-54 X 2.5-3.1 /x, (Fig. 4:7,A,C,D), oozing out from the
pycnidia in large cirri which retain their identity for some time in
water. ' |
On: Agrostis hallii Vasey, Oreg.—A. scabra Willd., N. Dak.—Stipa Colum-
biana var. nelsonii (Scribn.) Hitchc, Wash.—S. viridula Trin., N. Dak.,
S. Dak.—S. mlliamsii Scribn., N. Dak. /
The variety has longer spores (Fig. 47,A; \50,M} than the species,
except some summer material from North Dakota which has 1- to
3-septate spores 22-40 X 2.6-3.7 fx. (Fig. 47,Z)).'Microspores 5-10 X 0.5-
0.7 /t occur (Sprague, 1946,a). This variety is readilyîstinguished
from S. adropbgonis f. spoYoboKcola by the bacillar shape of~lhe-former
(Fig. 25,D). This form of S. andropogonis occurs in abundance on
Stipa comata at Mandan, North Dakota.
S. secalis var. stipae belongs in the S. avenae complex but at present
it is recognized. ,
References: Sprague (1941,6; 1944,rf; 1946,a,e; 1949,a, p. 499).
o ' /
Septoria spartinae (Trel.) Sprague—Leaf Spot
Ascochyta spartinae Trel. " - ' '
Lesions on living leaves elliptical to striate, straw to buff color,
with ofctcure, tardily erumpent pycnidia 100-140 /x wide and 70-90 fi
A n
i
i
FIG. 47.—Septoria secalis: A, pycnospores of var. stipae on Siipa viridula,
type; B,ôross section of pyonidial wall on Secale cereale, Pottawattamie County,
Iowa; C, pycnospores of var. stipae on Agrostis hallii, Polk County, Oreg.; D,
pycnospores of var. stipae on Stipa wiUiamsii, Mandan, N. Dak. Summer /ma-
terial. (
•deep, smaller on Beckmannia, strongly immersed, black, heavy-walled,
darker adjacent to the ostiole; pycnophores prominent, cuspidate to
narrowly pyriform, arising from hyaline pycnophore initials along the
floor of the pycnidium, 4-7 X 2.5-3.7 /J.; spores mostly 2-septate, nar-
rowly elongate-fusiform, tapering to a relatively sharp apex and a
more gradually blunted or truncated base, 26-38 X 2.0-2.5 [JL, ranging
to 3.0 ;n (in type).
On: Beckmannia syzigachne (Steud.) Fern., N. Dak.—Spartina gracilis
Trin., Utah—S. pectinata Lk., S. Dak., Wise.
The pycnidia are somewhat "Rhabdospora-like," that is, dark and
heavy-walled, because they are buried in the silicified, thick leaves of
the host. The fungus has never been collected in any great quantity
and does not seem to be important. Collections on Beckmannia from
two places in North Dakota appear to be saprophytic material of
S. spartinae. The pycnidia were dark, sunken, 60-90 /* diam. containing
hyaline (yellow, only in mass), 1- to 2-septate spores, 35-44 X~2-9-3.3 /x.
References: Davis (1919,6)^ Sprague (1944,d; 1946,a,e).
Septoria stipina Died.—^Leaf Spot

Spots small, linear, brown-margined, center gray to white, 2 mm.
long, 0.5 mm. wide; pycnidia epiphyllous, minute, light brown, 90-
FIG. 48.—Septoria spp., cross sections of pycnidia from prepared slides stained
in Ehrlich's haematoxylon and photographed. {Continued on next page.)
120 ju, ostiolate, context indistinct; spores filiform, curved to flexuous,
hyaline; 25-40 X 0.5 [i.
On: Stipa Columbiana var. nelsonii (Scribn.) Hitchc, Oreg.
The above description is taken from that of the type. This species
was described on Stipa sp. from Kashmir (Sydow, Sydow, and Butler,
1916, p. 214). The only collection which has been made in the United
States occurred at Long Creek, Oregon, in the Blue Mountains. At-
tempts to find further material in 1947 in this area failed because we
could find no Stipa sp. in the vicinity. The material obtained by W. E.
Lawrence some 30 years ago consisted of some fragments which the
writer removed from a phanerogamic herbarium specimen at Oregon
State College. Lesions were confined to leaves which were dead or
necrotic at the time of collecting. The pycnidia were prominent, dark
brown, erumpent, ostiolate, finally collapsed, 100-120 X 150-200 /u,,
composed of compacted creosote-brown crushed cells. The spores are
straight, very narrow, 1- to 4- (mostly 3-) septate, 39-63 X 0.8-1.1 ja
(average 52 X 0.95 /x) (Fig. 50,0). The Oregon material could, per-
haps, be aberrant material of S. andropogonis f. sporobolicola but it
would be difficult to reconcile the utterly filiform spores of S. stipina
with the distinctly broader ones of S. andropogonis. It is also hard to
understand why this fungus (S. stipina) has not been found again. We
have examined a large number of collections of Stipa from 15 western
'^.- states without finding any material approaching S. stipina and in the
Far West have not found any Septoria material on Stipa except one
collection of S. secalis var. stipae Sprague (1949,a, p. 499).
Range: United States, Kashmir.

References: Sprague (1944,d; 1946,e); Sydow, Sydow, and Butler (1916).
Septoria tandilensis Speg.—Black Spot

Spots none to diffuse but numerous black pycnidia form a promi-
nent black spot; pycnidia somewhat lenticular, black, cupulate, finally
(See opposite.)
All X170 except M which is X600: A, S. jaculella on Bromus rigidus, near
Coburg, Oreg., showing chambered and ill-formed pycnidial walls in this collec-
tion; B, S. passerinii on Sitanion hystrix, Mt. Hood Natl. For., Oregon; C, S.
calamagrostidis f. koeleriae on Koeleria cristata, CorvalHs, Oreg.; D, 8. pas-
serinii on Hordeum vulgare, Corvallis, Oreg.; E, S. macropoda var. grandis on
Poa nervosa, Missoula, Mont.; F, S. tritici f. avenae on oats, Eugene, Oreg.;
G, S. jaculella on Bromus laevipes,' Calif.; H, S. elymi-europaei, Obersdorf,
Bavaria; I, S. oudemansii on Poa compressa, Corvallis, Oreg.; J, S. bromi on
Bromus commutatus, Lane County, Oreg.; K, S. pacifica on Elymus mollis,
Newport, Oreg.; L, S. triseti on Agrostis tenuis, Comstock, Oreg.; M, S. pas-
serinii on Hordeum distichon, Kiev area, Russia showing basal portion of pyc-
nidium with microspores (X600); N, S. arctica on Calamagrostis nutkaensis,
Delmoor Bog, Oreg.; O, S. tritici f. hold on Holcus lanatus, Corvallis, Oreg.;
P, S. injuscans on Elymus glaucus (O.S.C. 8444); Q, small pycnidium of S. tritici
on wheat, Linn Co., Oreg., contains microspores. (From Oregon State College
Bot. Monogr. 6, Plate 2.)
FIG. , 49.—Sepioria tenella on Festuca spp. (except I, which is f!fiyllosticto.

sp.): A, pycnospores from type of S. festucina ^= S. tenella); B, pycnospores
collapsed, 100-120 ix, closely gregarious, seriated, often filled in winter
with prosenchymatous material; spores filiform 50-80 X 1.3-1.6, less
often up to 111 X 2.7 /*.
On; Panicum boreale Nash, Wise.—P. capillare L., N. Dak.—P. implicatum
Scribn., Wise.—P. praecocius Hitchc. and Chase, Wise.—P. scribnerianum
Nash, Wise—P. spp., PJa., Ga., N. Car., Oida., S. Car., Tex.
This species is distinguished by the numerous black pycnidia which
stand out in relief against the straw-colored or brown dead leaf ma-
terial of fall or winter collections. Greene (1944,a,c) reported a num-
ber of additional hosts from Wisconsin, and it is likely that the host
range as now known is somewhat incomplete.
Range: United States, Argentina.
References: Greene '(19440,0); Spegazzini (1902); Sprague (1946,a;
1948,0).
Septoria tenella Cke. and Ell.—Leaf Blotch

Spots none or small, vague gray to brown lesions on culms, sheaths
and leaves of maturing or living plants, pycnidia creosote-brown, 60-
120 X 95-130 fi, more or less elliptical in cross section, composed of
tightly crushed, oblong to polyhedral cells averaging 0.5 to 3 /i'wide;
pycnophores small, cylindrical to very narrowly subulate, averaging
4 /i long, 1 fi wide; spores exti'emêly variable, filiform to short bacillar,
5-70 X 0.8-2.1 ,1, commonly 25-45 X 1-0-1.5 /x.
On: Festuca dertonensis (All.) Aschers. and Graebn., Greg.—F. elatior L.,
Wash.—F. idahoensis Elmer, Greg., Wash., Wyo.—F. mairei St. Yves,
Greg., in plots—F. octoflora Walt., Colo., Ida., III., Kans., Mont., Nebr.,
N. J., N. Dak., S. Dak., Wyo.—F. ovina L., Wash., Wyo.—F. rubra L.,
Greg., Wash.—F. rubra var. commutata Gaud., Greg.
Spores on Festuca dertonensis from western Oregon vary from 28-
41 X 0.8-1.2 ja to 7-17 X 1.0-1.5 /u, and even 9-12 X 1.4-1.7 fi in adja-
cent pycnidia (Fig. 4:9,F,G,J), while on F. rubra var. commutata all
spore types are intermingled and found to some extent in the same
pycnidium (Fig. J:9,D,i?,K). It is the most common, and ^.widespread
material on F. octoflora Walt, from Idaho to Minnesota, with spores
0- to 2-septate, 44-67 X 1.6-2.1 ,* (Fig. 49,B). In May, 1941, this fun-
from F. octoflora, N. Dak. 618; C, conidia from pure culture isolated from
F. rubra, Bay City, Oreg.; D, pycnospores from F. rubra var. commutata, near
Alpine, Oreg.; E, cross section of .pycnidial wall on F. octojiora, Stockton, Kans.;
F, Ascochyta-\\ke spores of_/S. tenella from F. dertonensis. Fish Hatchery at
Roaring River, Oreg.; G, Septoria-like spores from Roaring River collection;
H, pycnospores from spore exudate from pure culture obtained from F. rubra,
Alpine. Oreg.; I, pycnospores of Phyllosticta sp. on F. subulata, Silver Creek
Falls, Oreg.; J, intermediate spore stage on F. dertonensis. Roaring River, Oreg.;
(compare Fig. 49, F, G, and J ) ; K, cross section of pycnidial wall from Alpine,
Oreg., material of F. rubra var. commutata; L, pycnospores of S. festucae (=8.
tenella) on F. gigantea, F. Pol. Ex 232; M, pycnospores from F. rubra. Bay City,
Oreg. (From Oregon State College Bot. Monogr. 6.)
gus caused a speckled lesion on the basal leaves and culms of almost
100 per cent of this grass over vast areasîn-the Dakotas. At that time
it was by far the most abundant parasite on any grass in the region
(see also Fig. 4:9,A,E). , "I
Material on F. rubra commutata from Oregon appears to be a
variant similar to Septoria jestucae Died. (Fig. 49,L). However, the
material on F. dertonensis could be assigned to S. tenella, S. jestucae,
and Ascochyta sp., depending on the particular pycnidium examined
(Fig. 4Q,F,GJ). The material from the Great Plains is more typically
and uniformly like S. tenella. I t is no doubt racially distinct from the
polymorphic West Coast material. S. tenella also occurs on F. ovina in
the Yellowstone National Park and on F. idahoensis in both Wyoming
and Oregon. The Oregon material (W.S.C. Coll. Ser. 3840) was obtained
at an elevation of 4,994 ft. (conveniently located at a Geodetic Survey
mark) in the Malheur National Forest. It had stiff filiform spores with
some of the bases swollen and they were 2.7 /i wide. However, some of
the spores were partially germinated, and the width of the spore is
attributed to this partial germination—a condition which is not unusual
when the material has been left in seed packets or coin envelopes for
several days without sufficient dehydration. The above-mentioned
Oregon material from the forested interior seemed to be more nearly
like the coastal material on F. rubra (Fig. 49,C,D,F,G,J,M) as the
spores were variable in length, from 15 to 45 /*. F. idahoensis, inciden-
tally, is rarely parasitized by this species; at least, out of several hun-
dred specimens examined of this grass only the two collections of this
fungus have been made. Apparently moisture is unable to cling to the
narrow, somewhat glaucous leaves sufficiently long to permit spore
germination and host infection. /
In pure culture, the isolates from F. rubra commutata produced a
slimy, cocoa brown mass of conidia, which contained many Ascochyta-
like spores (Fig. 49,H) as well as filiform ones plus many variants and
freaks. Later the conidial masses, through germination^f the conidia,
produced a somewhat catbonized surface over the colonyrT?he isola-
tions from F. octoflora obtained from Grassy Butte, NorthJDakota,
were different in that mucose, mounded, flesh pink colonies similar to
S. tritici were formed.
A collection of a species of Phyllosticta on F.,subulata had spores
4.5-11 X 1.4-2.0 jn and was assigned to no definite species (Fig.'49,/).
It was considered as being close to Septoria tenella but possibly was
different (Sprague, 1944,d). This fungus was mentioned under Phyllos-
ticta spp. earlier in this text. Recently the same fungus was noted on
F. subulata collected on the South Fork Clearwai^'lîver, Idaho.
Range: United States, probably Europe and Asia.
References: Gooke and Ellis (1879); Diedicke (1912,a); Sprague (1944,d);
Tehon and Daniels (1927).
FUNGI IMPBRFECTI—SPHAEROPSIDALES 261
Septoria triseti Speg., em. Sprague—Leaf Mottle

Lesions on the leaves indeterminate, sublinear to irregular, or on
leaf tips, cinerous; pycnidia not prominent, black brown, ostiolate, sub-
stomatal, globose to flattened at the ostiolar end where it is appressed
against the leaf epidermis, 40-100 /A (mostly 40-80 /x) in diam., formed
of irregular, oblong-polyhedral, parenchymatous cells, in strands,
which converge in radii toward the ostiole, brown outer cells two layers
thick (3 ju.) with 2 or 3 inner hyaline layers {3 to 5 fi thick) giving rise
to narrowly bulbous-cylindric pycnophores, 2.5-3.5 [i. Pycnospores fili-
form to subbacillar or sometimes narrowly fusiform, straight, bent or
less often moderately curved, 0- to 1-septate, 16-43 (18-35) X 0.8-2.0
(1.3-1.7) p,.
On: Agrostis alba L., Oreg., Wash.—A. castellana Boess. and Rent., Oreg.,
. in plots—A. exarata Trin. var. ampla (Hitchc.) Hitchc, Oreg. (Fig. 5Q,E)
—A. stolonifera L., Oreg.—A. tenuis Sibth., Oreg., Wash.
S. triseti is associated with S. calamagrostidis on Agrostis spp. in
Oregon, but it is readily distinguished from the latter by the much
smaller spores and pycnidia (Fig. 48,L) and from its somewhat dis-
tinct host range. Spegazzini (1887) described this species from south-
ern Argentina material collected on both Trisetum phleoides (Vill.)
Trin. [Koeleria phleoides (Vill.) Pers.] and Agrostis magellanica Lam.
with spores 20-30 X 1-1.3 /J, borne in pycnidia 70-80 jn. The type has
apparently disappeared. Sprague (1944,d) emended the description to
include only Agrostis, considering that the material on Koeleria was
probably S. calamagrostidis f. koeleriae.
Material on A. alba (Fig. 50,D) is racially distinct from that on
A. tenuis (Fig. 50,A,B,C,F,G,H,I,J,K,L). That on the latter is termed
race 1, that on A. alba, race 2. S. phleina Baudys and Picb. is somewhat
similar to S. triseti (Fig. 50,iV).
Range: United States, Argentina.
References: Spegazzini (1887, p. 164); Sprague (1944,d; 1946,e).
Septoria'tritici Rob. in Desm.—Speckled Leaf Blotch

Lesions' straw-colored, appearing speckled due to the^numerous
pycnidia (Fig. 52,C); pycnidia golden brown, substomatal, ostiolate,
somewhat flattened, ellipsoid, 60-200 fi, mostly 100-150 /*, walls thin,
composed of 2 to 5 outer layers of golden brown, oblong, box-shaped,
thin-walled cells; then inside that 1 to 4 intermediate layers of poly-
hedral, compacted cells and inside this a hyaline layer; pycnophores
narrowly ampuUiform (Fig. 51,B,C,D,F,G,L), 5-13 X 1-3-3 fi; spores
of two kinds (Fig. 51,A), macrospores obclavate-filiform, 35-98 X
1.4-2.8 /x (Fig. 51,H,I,J,M); microspores bacteria-like, curved or
hooked, aseptate, 5-9 X 0.3-1 fi (Fig. 51,K,N).
FIG. 50.—A"L, Septoria triseti: A, conidia from young culture isolated from
Agrostis palustris X A. tenuis near Newport, Oreg. (O.S.C. 16); B, conidia from
staling pure culture, isolated from A. tenuis; C^ pycnospores from A. tenuis,
On: Secale cereale L., Wash.—Triticum aestivum L., Alta., B. C, Calif., D. C,
Ga., Ida., 111., Ind., Iowa, Kans., Ky., Manit., Mich., Minn., Mo., Mont.,
Nebr., N. Y., N. Car., Ohio, Okla., Ont., Oreg., Penn., Que., Sask., Tenn.,
Wash., W. Va., Wise.—T. dicoccum var. farrum Bayle, Oreg.—T. spelta L.,
Oreg.—T. turgidum L., Oreg.
S. tritici is an important parasite on winter wheat in the moister
parts of Oregon, Washington, and northern California and sometimes
it is serious in the middle western and eastern states. It is less common
east of the Cascade Mountains in Washington and Oregon during most
seasons, in fact was rarely reported until the unprecedented season of
1948. In that year (Sprague, 1948,(i; 1949,a) of prolonged precipitation,
leaf blotch appeared in spring in considerable quantity on the basal
leaves of winter rosettes and later as linear spots on the leaves of
winter wheat in June. When a survey was made in early June most
of the spots on leaves proved to be the small circular ones of Seleno-
phoma, but two weeks later these had been partially replaced by the
longer, less distinct spots of S. tritici. It was found in all areas in
eastern Washington except in the drier parts in the central portion of
the state. The fungus was also common in adjacent Idaho where it
had been reported only once previously (Schade, 1936). In the usually
humid parts of Oregon and Washington winter rosettes of wheat are
100 per cent covered with the salt-and-pepper lesions of S. tritici and
this injury has a depressing effect on the host, even though the fungus
-often disappears entirely when warm, drier weather comes in spring
and summer.
It has been shown by Weber (1922,6) that S. graminum does not
occur on wheat in Wisconsin while the writer reported that S. tritici
was the only filiform-spored species present on wheat in the United
States (1944,d) and that S. graminum (1938,d) was apparently limited
to Brachypodium spp. and does not, as far as known, occur in this
country. Most of the species that the writer assigned to synonymy are
obvious synonyms, but one, S. briosiana Morini (1886) is less appar-
ent. In this form microspores (9-11 X 0.5-0.75 fi) are borne on more
or less abortive macrospores (32-40 X 1.5-2.0 ju). (See also Fig. 48,Q.)
Septoria tritici produces mucose, flesh-color colonies, on potato-
dextrose agar, which later become carbonaceous, finally subcottony or,
on staling, cottony.
Corvallis, Oreg. (O.S.C. 80); D, pycnospores from A. alba, Ft. Steilacoom, Wash.;
E, pycnospores from A. exarata var. ampla, Lewisburg, Greg.; F, pycnospores
from A. tenuis, Young's Bay, Clatsop Co., Greg.; G, pycnospores from A. tenuis,
Alsea, Greg.; H, pycnospores.from A., tenuis, Tillamook Co., Greg.; I, conidia
from young materia! in pure culture isolated from A. tenuis, Greg.; J, cross sec-
tion of pycnidial wall froin A. tenuis, Benton Co., Greg.; K, pycnospores from
A. tenuis, Benton Co., Greg.; L, cross section of pycnidial wall from same collec-
tion, just mentioned; M, pycnospores of Septoria secalis vsi,r. stipae on Aarostis
hallii, Polk Co., Greg.; N, pycnospores of S. phleina Baudys and Picb. on Phleum
arenarium from Croatia; O, pycnospores of S. stipina on Stipa columbiana var.
nekonii. Long Creek, Greg. (G.S.C. 10,761). (From Oregon State College Bot.
Monogr. 6, fig. 15.)
DISEASES OE CEREALS AND GRASSES
FIG. 51.—Septoria tritici (except E and G): A, microspore and, maerospore

stage on wheat, 40 miles south of Davis, CaliJ,; B, cross section of pycnidial
The information on varietal resistance in wheats to speckled blotch
is somewhat scanty, although the subject is under investigation in the
United States at present, it is understood. Savulescu (1930) reported
some differences in susceptibility of wheat varieties to S. tritici in
Romania. Bremer, et al. (1948) found Elbistan and Eskischir free from
attack in Turkey. Sprague (1944,d) reported that the earlier maturing
varieties appeared to be the most susceptible. These observations were
substantiated in the field in 1948 in the Palouse region, although it is
possible that the difference is due to maturity at the time of maximum
infection rather than any strong resistance on the part of the late
varieties. As mentioned in our 1944 study (1944,(i) C. 0 . Johnston had
noted that most of the soft red winter wheat varieties in southeastern
Kansas had more spots than the hard red winter wheat varieties.
Range: United States, Algeria, Argentina, Australia, Austria, Czechoslo-
•vakia, Denmark, France, Germany, Great Britain, Hungary, India, Iran,
Italy, Morocco, Netherlands, New Zealand, Romania, Spain, Turkey,
U.S.S.R., Uruguay.
References: Baudys and Picbauer (1924); Beach (1919); Bremer, et al.
(1948); Cormier (1932); Desmazieres (1842, 1843, 1847); Grove (1916,
1935); Luthra, et al. (1937, 1938); Maokie (1929,6); Marchionatto
(1941); Morini (1886); Rivera (1923); Rosella (1929); Rosen (1947,6);
Saccardo (1913,0,6); Savulescu (1930); Sprague (1938,d; 1944,d;
1950,a); Unamuno (1930); Weber (1922,6).
Septoria tritici f. avenae (Desm.) Sprague—Speckled Leaf Blotch

Lesions on leaves and sheaths gray to straw-colored, appearing
speckled due to prominent pycnidia (Fig. 52,A,B), irregular to rec-
tangular, pycriidia as in type but peridium slightly looser (Fig. 48,f);
pycnophores ampuUiform to subcylindric, 4-7 X 1.3-1.6 /*, developing
from hyaline, somewhat bulbous initials (Fig. 51,E); spores slightly
narrower than on wheat but readily recognized by their sinuous ap-
pearance, 35-120 X 1.6-3.5 /i (averaging 65-80 X 2.1-2.3 tx) [Fig.
52,2;, ( a ) ] .
On: Avena byzantina K. Koch, Oreg., Wash.—A. fatua L., Oreg., Wash.—•
A. sativa L., Oreg., Wash.
wall on wheat, Warwick, Wash. (O.S.C. 134); C, cross section of pycnidial wall
on wheat, Denmark; D, cross section of pycnidial wall, on wheat, Pendleton,
Oreg. (O.S.C. 10,362); E, cross section of pycnidial wall of Seploria tritici f.
avenae on Avena sativa, Astoria, Oreg. (O.S C. 133); F, cross section of pycnid-
ial wall, on spring-sown wheat, Benton County, Oreg.; G, cross section of
pycnidium from type of S. neglecta; H, pycnospores from spring-sown wheat,
Benton County, Oreg.; I, pycnospores from winter wheat collected in Decem-
ber, Benton County, Oreg.; J, pycnospores and K, microspores from Khapli
emmer, Benton County, Oreg.; L, cross section of pycnidial wall on wheat.
Friend, Oreg.; M, pycnospores from wheat, Rabat, Morocco; N, microspores,
young macrospores and pycnophores from cross section of pycnidium on wheat,
East Farm, Linn County, Oreg. (From Oregon State College Bot. Mongr. 6.)
OOA
DISEASES OF CEREALS AND GRASSES ^
FIG. 52.—Septoria tritici and S. indict f. atênae. A, forma avenae on Red

Rustproof oats, Alse Valley, Oreg., showing lesions on the leaves; B, enlarged
portion of lesions on red oats X3'^; C, S. Irilici on wheat, enlarged view showing
pycnidia, X3%; D, young culture of f. avenae on potato-dextrose agar; E, (a)
pycnospore of S. trilici avenae from Red Rustproof oats X375 and (6) pycno-
spores of S. avenae, Puslinch, Ont. (F. Columb. 4471) X375. (From Phyto-
pathology.)
This form is common on wild oats and at times on winter oats in

western Oregon and Washington during open wet winter weather.
Frandsen (1943) recently described S. sativa on oats. It is the same
fungus as the one in Oregon and Washington. On the basis of present-
day species concept in this country it is questioned if the form on oats
warrants complete separation from the form on wheat. In culture they
are, in addition (Fig. 52,D), very similar.
S. avenae—barbatae Gonz. Frag. (1921) has filiform-clavulate
spores 60-75 X 3.0-3.2 /jt. As described and illustrated it is distinct.
Range: United States, Argentina, Denmark, France, Morocco, New Zea-
land, Poland.
References: Desmazieres (1847); Frandsen (1943); Sprague (1934,a;
1944,rf; 1946,e).
Septoria tritici f. holci Sprague—Speckled Leaf Blotch

Lesions occur on the lower leaves of the plant, mostly indefinitely-
bordered, irregular, coalescing to cover all or large portions of the
leaf. Lesions at first are yellow green to vinaceous buff to vinaceous
fawn or fawn, later becoming Saccardo umber in the center of the
lesions. Pycnidia moderately prominent, scattered or grouped in rows
parallel to the veins, erumpent but deep seated, visible from both
surfaces, hypophyllous, sometimes distinctly flattened, often subglo-
bose to globose (Fig. 48,0), ostiolate, dark brown to black brown;
pycnidial wall 10-12 /x thick, outer portion composed of dark, crushed,
oblong, polyhedral cells surrounding intertwined, lighter polyhedral
cells (about 2 ju diam.) producing hyaline pycnophore initials; pycno-
phores are also hyaline, ampuUiform to nearly cuspidate, 3.5-5.7 X
1.4-2.1 /x (Fig. 53,JD); pycnospores exuded in very short, pale, flesh-
cotored cirri or guttulae, straight or somewhat curved, less often
strongly curved, except at distal cells, scolecosporous, slightly broad-
ened at base, crystal-hyaline contents, cross walls obscure in many
collections, 1- to 4- (0- to 5-) septate, 30-80 X 1-4-1.8 fi (spring col-
lections), 35-105 X 1.4-2.5 ^ (winter collections) (Fig. 53,E), rarely
narrower. ,,
On: Holcus lanatus L., Oreg., Wash.
This fungus is very common in late winter and early spring. Cross
inoculation trials indicate that it is confined to Holcus. S. holci Pass, is
probably S. nodorum.
Range: United States, Scotland.
Reference: Sprague (1944,d).
Septoria tritici var. lolicola Sprague and A. G. Johnson—

Speckled Leaf Blotch
Lesions indefinite on the leaves, sheaths, and glumes, at first green,
later yellow, grayish brown, brown or Saccardo umber to isabelline,
sometimes reddish, in- old lesions, white in the center; pycnidia moder-
ately prominent, scattered or in rows parallel to the leaf veins, usually
globose to subglobose, sometimes ellipsoid, 80-150 X 90-180 n, mostly
120-150 X 120-150 IX, ostiole 10-25 X 10-30 /j. diam., peridia as much
as 15 fi thick, pale to deeper amber brown, smooth, outer cells sub-
cubical to oblong, inner cells interwoven, polyhedral, giving rise to
pycnophores, 3-6 X 1.5-2.-3 fi, slightly pyriform to subcylindric, ends
acute with blunt terminal (Fig. 53,B); pycnospores numerous, hyaline,
with small oil drops, 0- to 5- (mostly 1- to 3-) septate, slightly enlarged
at the base, and tapering to a rounded base, distal cells tapering to
subwhip-like, slightly to strongly curved, 31-85 X 1.3-2.8 fi (Fig. 53,.4),
in Oregon collections on leaves in winter, or 22-30 X 1.1-1.7 /t in sum-
FIG. 53.—A, Pycnospores of S. tritici var. lolicola, East* Corvallis, Oreg.; B, C,

cross section of pycnidial walls of same species; D, cros^ section of pycnospores
of S. tritici hold on Holcus lanatus, Blodgett, Oreg.; E, pycnospores of S. tritici
hold, Corvallis, Oreg. :
mer material in Oregon, or 23-52 X 1.2-2.6 fi on the glumes in Des-

mazieres Plant Cryptogames de France No. 2169, in pgî.
On: Lolium multiflorum Lam., Oreg.—L. perenne L., Oreg. . ""
In early years this variety and two species, Bhabdospora lolii Cast.
and S. graminum var. lolii Mont, were all confused. R. lolii is, as men-
tioned elsewhere, assigned to S. loligena Sprague (1944,d) while S.
graminum var. lolii is a species of Ascochyta known as A. desmazieri
Cav. (1944,d). Frandsen (1943) described Sepioria cavarae based on
A. desmazieri Cav. and a collection of Septoria made by Rostrup.
Frandsen's description and illustration show that his S. cavarae is the
same as our S. tritici var. lolicola. Frandsen, we-believe, erred in con-
sidering S. graminum var. lolii and A. desmazieri as synonyms of
S. cavarae as they represent Ascochyta phases, either of a distinct
species or possibly of S. loligena (1944,d) but not of S. tritici yar. loli-
cola. We feel that the filiform material on Lolium is adequately recog-
nized as a variety of S. tritici. Because of the ambiguity in Frandsen's
description we prefer not to reduce his species to a variety of S. tritici
but to retain the later described but nonambiguous 5. tritici var. lolicola.
Range: United States, Denmark, France.
References: Castagne (1851); Cavara (1893); Desmazieres (1851);
Frandsen (1943); Sprague (1944,rf); Westendorp (1857).
Stagonospora Sacc.
The pycnidia of this genus are globose to flattened, golden brown,
ostiolate and bear cylindrical to fusiform (broadened, not filiform)^
multiseptate spores on blunted to short cylindrical pycnophores. The
genus occurs on leaves, culms, and sheaths. Most of the species are
parasitic although some of those on coarse grasses, such as on Phrag-
mites, are saprophytes. Most of those which are saprophytic have
coarse spores ranging from hyaline to yellow, therefore approaching
Hendersonia Sacc, which has brown spores.
We present a key to the species of Stagonospora which have been
identified in the United States and Canada on Gramineae.
KEY TO SPECIES OP STAGONOSPORA ON GRAMINEAE I N T H E

UNITED STATES AND CANADA
A. Spores mostly over 8 i^- wide (24-45 X 8-11 M)

Stagonospora simpUcior Sacc. and Br.
AA. Spores narrower (but some up to 9 M at times)
B. Spores somewhat boat-shaped, 20-45 X 6-9 M
C. Spores mostly 3- to 4-septate
D. In vague lesions on necrotic leaves
St. subseriata Desm.
DD. In purple lesions on living leaves
St. maculata (Grove) Sprague
CC. Spores 5- to 7-septate
St. graminum Sacc. and Scalia
CCC. Spores 2- to 3-septate, 24-28 X 7-9 M
St. paspali Atk.
BB. Spores more or less cylindrical
C. Spores somewhat fusiform
D. Spores 22-30 X 4.6-6.4 /i
St. glycericola Sprague
DD. Spores 15-30 X 2-3.2 ii
E. Spores somewhat truncated at base
.,St. agrostidis f. angusta Sprague
EB. Spores more fusiform than truncated
St. bromi A. L. Smith and Ramsb.
DDD. Spores 30-50 X 3 ,«, 7-septate
St. intermixta (Cooke) Sacc.
DDDD. Spores 12 X 4 M
St. ischaemi Sacc.
CC. Spores definitely cylindrical ' |
D. Spores 25-60 X 2.5-5 A ~" - - I
St. arenaria Sacc. ^ '
DD. Spores 30-80 X 4-9 /x, in buff spots i
St. foliicola (Bres.) Bubak
DDD. Spores 35-55 X 4.5-5.0 ti, in vaguer lesions
St. vexatula Sacc. I
DDDD. Spores 18-22 X 3-5 ic*
iSt. graminella Sacc. (saprophyte)
DDDDD. Spores 40-47 X 6-9 /*
St. spartinicola Sprague
CCC. Spores subclavate or cylindrical, 13-33 X 5-7 ;»
. St. brachyelytri Greene
The genus Stagonospora is separable from Septoria with some diffi-
culty, because on grasses there is a group of fungi, some placed in
Septoria, some in Stagonospora, that have spores about 30-40'60 ix
long X 3-4-6 pi, wide and therefore are both filiform and cylindrical,
depending on your point of view. Some workers consider Stagonospora
as having coarse spores similar in all respects to Hendersonia, except
for color, and similar also to Hendersonia in being mainly weak para-
sites or saprophytes (Kirschstein, 1939). There are exceptions to this
classification that makes its value questionable. In general, the writer
uses the length-wddth ratio of 10 to 1 as the division point between cy-
lindrical and filiform. Since the lengths of the spores of many of these
species range from about 9 to 1 to 13 to 1, they become difficult to
classify. In such cases they are left in the genus to which they have
been customarily referred—e.g., Septoria nodorum, Stagonospora are-
naria, and Septoria avenae.
Stagonospora agrostidis f. angusta Sprague—Leaf Blotch

Spots on leaves and sheaths fuscous, straw-colored or none, pyc-
nidia globose, golden, parenchymatous, ogtiolate, 120-280 /*; pycno-
spores fusiform, subapute, straight to somewhat curved, bases sub-
obtuse to subtruncate, nonconstricted, 3-
septate, hyaline, conspicuously guttulate
(4-8 oil drops), 15-27 X 2.2-3.2 /JL (averag-
ing 22.3 X 3 ja):
On: Stipa rohmta .Scribn., Colo.—S. viridula
Trin., N. Dak. '
This is an exceedingly obscure fungus
Stagonosvora agrostidis f. ^îch is often a secoridary parasite and
angusta, N. Dak. saprophyte on Sj-gro- viridula m western
North Dakota "and-it'has also been found
on rusted plants of St. robusta in Colorado. The spores are typically
fusoid-cylindrical, blunt at the base, tapering sharply toward the apex
(Fig. >54). Their typical shape and the guttulate nature of the spores
FUNGI IMPERFECTI—SPHAEROPSIDALES 271'
readily distinguish them from Septoria nodorum and summer material
of S. secalis var. stipae (Sprague, 1946,a). The fungus is considered a
form of Si. agrostidis H. and P. Sydow (1900) which has fusoid curved,
subfalcate, 3-septate spores, 24 X 4 /x.
Stagonospora arenaria Sacc.—Purple Brown Blotch

Lesions dark purple, purple black to deep brown, later variously
mottled, fading to buff or straw color later; pycnidia scattered, not
prominent, golden brown to sepia, subglobose, erumpent, ostiolate, thin-
walled, 50-160 (240) ju, diam.; pycnophores short, botuliform or sub-
cylindric, spores cylindrical or less often, subfusiform, 3- (1- to 4-)
septate, 25-60 X 2.5-5.0 /t, often 30-45 X 3.5-4.3 /^.
On: Agropyron ciliare (Trin.) Franch, N. Dak.—A. cristatum (L.) Gaertn.,
N. Dak.—A. desertorum (Fisch.) Schult., N. Dak.—A. repens (L.) Beauv.,
Oreg,—A. smithii Eydb,, N. Dak,—A. spicatum (Pursh) Scribn. and
Sm., N. Dak.—A. trachycaulum {Ik.) Malta, Mich., N. Dak.—A. tricho-
phorum {Ik.) Richt., N. Dak.—Bouteloua curtipendula (Michx.) Torr.,
Okla.—Cinna latifolia (Trev.) Griseb., Minn., Oreg.—Dactylis glomerata
L., Ky., Md., Mich., Ohio, Oreg., Wash.—Elymm canadensis L., Ky., Mich.,
N. Dak.—E. dahuricus Turcz., N. Dak.—E. glaucus Buckl., Oreg., Wash.
—E. mollis Trin., Alaska—E. pseudoagropyron (Griseb.) Trin., N. Dak.
—E. triticoides Buckl., Calif.—E. virginicus L., Mich.—Oryzopsis race-
mosa (J. E. Smith) Ricker, Wise.
As shown by the key under Septoria elymi, this fungus has only
slightly larger spores (Fig. 55) than S. avenae f. sp. triticea T. John-
son. It has been used somewhat as a catch-all for those forms with
narrowly cylindrical spores (Fig. 27,E) which slightly exceed the size
of Septoria avenae f. sp. triticea. It appears to be especially common
on Agropyron and Elymus spp. It was originally described on E. arena-
rius from northern Europe with spores 30-35 X 3.5-4.0 /x (Saccardo,
1878, p. 124). The material from Sweden has spores of about the size
of those of S. avenae f. sp. triticea. Grove (1935) gave the spores as 1-
to 5- (or even 7-) septate instead of 3-septate as given by Saccardo,
both workers basing their data on the type.
Until more information is available St. arenaria is maintained as a
'distinct species (Sprague, 1948,c) from Septoria avenae f. sp. triticea,
the pycnidial stage of Leptsphaeria avenaria f. sp. triticea T. Johnson
(1947).
Septoria dactylinaGonz. Frag, on Dactylis (1921) has spores 36 X
4 fi and probably belongs somewhere in this group.
Range: United States, Canada, Sweden.
References: Johnson, T. (1947); Saccardo (1878, p. 124); Sprague (1941,c;
1944,d; 1946,a,e; 1948,c).
FIG. 55.—Stagonospora arenaria. A, pycnospores on Elymu^glaucits; B, ditto,

Tangent, Oreg.; C, ditto oii Dactylis glomerata, Ohio; D, ditto^"Wallace Bridge,
Greg. ~
Stagonospora brachyelytri Greene—^Leaf Spot

Spots pale brown with narrow, dark brown/margins, mostly elon-
gate, 5 - 1 5 x 1 - 5 mm., following the venation;'pycnidia brown, sug-
globose, 100-150 ;u, diam., mostly 100-110 ii, indistinctly ostiolate; pyc-
nospores hyaline, granular, subclavate or cylindrical, often slightly
curved, distinctly 1- to 3-septate, 13-33 X 5-7 /*.__ /
- -^—',
On: Brachyelytrum erectum (Schreb.) Beauv., Wise."
Shaw, Fisk,- and Bryan collected topotype material at Devil's Lake,
Wisconsin, on July 19, 1941. This has dark brown, linear or marginal.
later tawny or fuscous spots. The pycnidia are golden brown, flattened
globose, thin-walled and contain somewhat cylindrical, hyaline spores
with more or less blunted bases and rounded to slightly tapering apices
or sometimes the spores are semifusiform, 21-31 X 3.8-4.9 /x in a mount
in KOH made by Shaw. This specimen is filed, in part, in the Herbar-
ium of the Department of Plant Pathology, State College of Wash-
ington, Pullman.
Stagonospora rubi-albida Dearness and House was described, in
English, on B. erectum in 1940 (Circ. N. Y. State Mus. 24, p. 48) with
3-septate eylindric-allantoid spores 18-26 X 4-5 /i..
Reference: Greene (1946,6).
Stagonospora bromi A. L. Smith and Ramsb.—

Purple Brown Blotch
Spots amphigenous, elongate, finally confluent, purple brown (Fig.
56), center finally paler, pycnidia globose-lenticular, 130-175 /A diam..
~Ts*«i^^î
FIG. 56.—Leafspot on Bronius ciliaius caused by Stagonospora bromi, Roose-

velt Natl. For., Colo.
context parenchymatous, spores fusiform, to subcylindric, straight to

curved, 3-septate, nonconstricted (15)-17-24-(42) X 2.7-3.6-(4.0) [i
(Fig. 58,C,D).
On: Bromus carinatus Hook, and Arn., Ida., Mich., N. Dak.—B. catharticus
Vahl, Ky.—B. ciliatus L., Colo., Wise.—B. frondosus (Shear) Stand, and
Wootn., Colo.—B. inermis Leyss, Ky., Wise.—B. kalmii A. Gray, Wise.
—B. purgans L., Wise.—B. vulgaris (Hook.) Shear, Ida., Minn.
This fungus, originally described on Bromus ramosus Huds. from
England (Smith and Ramsbottom, 1914) appears to be a relatively im-
portant parasite of brome grass where it occurs in the Middle West
and adjacent areas in eastern North Dakota. Its relation to Septoria
avenae, Ascochyta sorghi, and Septoria nodorum needs study. One
specimen from Idaho (A.S. 17,214) had spores 15-42 X 2.7-4.0 /i.
Range: United States, Scotland.
References: Grove (1935); Hardison (1942); Smith and Ramsbottom
(1914); Sprague (1946,e; 1949,a).
/'
Stagonospora curvula Sacc, Bomm. and R'ouss.—Leaf Spot
Pycnidia innate, subglobose, 300-450 ^ dJam.,^ reddish black brown,
densely aggregated to sometimes confluent, (Joverpd with the gray epi-
dermis of the leaf, tardily ostiolate; spores linear, obtuse, curved, at
first 1-septate then 3-septate, 12-20 X 3 /i, yelloyish.
On: Panicura capillarioides Vasey, Tex.
Altstatt (1942) listed this fungus from Texas. We have not seen
this particular collection, nor are we familiar with the species. St. cur-
vula was described from dry grass culms in Belgium (Bommer and
Rousseau, 1890).
Range: United States, Belgium.
References: Altstatt (1942); Bommer and Rousseau (1890); Sacoardo
(Sylloge V. 10, p. 337).
Stagonospora foliicola (Bres.) Biib^k—Tawny Blotch

St. ^Jexata ">fw:. ôlucola B\:es.i
Cylindrosporium phalaridis Sacc. and Dearn.
Lesions elliptical, brown to vina(5eous, finally wine-colored, tawny
to buff, sometimes covering entire leaf, margins indefinite, pycnidia
epiphyllous, globose-applanate, between the leaf veins, gray to light
brown, 100-270 fx. diam., parenchymatous, ostiolate; pycnophores papil-
liform; pycnospores stoutly obclavulate with a tapering, finally blunt
base and an acute apex, 3- to 8-septate, sometimes, b u t less o f t ^ not,
constricted at the septa, 38-90 X 4.5-6.0 p., type material up to 9.5 [i
wide and 6- to 12-septate; most of the materiahin the United States
is immature with fusiform-cylindric to irregularly cylindric spores
with blunt bases, 30-35 X 3.4-3.9 ^ or 25-45 X 2.7-4.1 /..
On: Glyceria data (Nash) Hitchc, Qolo.-^Phalaris arundinacea L., Colo.,
Idaho, Iowa, Ky., Minn., Mont., N. Dak.,iOnt., S. Dak., Wise—P. cali-
fornica Hook, and Arn., Calif. ^--^..^^^
As mentioned, much of the material in the United States is imma-
ture. Such specimens have fusiform-cylindVic to irregularly cylindric
spores with blunt bases, 30-35 X 3.4-3.9 ^, sometimes 25-41 X 2.7-4.1 ^.
Until abundant material was obtained at Mandan, North Dakota, col-
lections (Fig. 57,A) were hesitatingly assignefd to Stagonospora are-
naria Sacc. (Sprague, 1941,c) while Davis placed these in Stagono-
spora intermixta (Cooke) Sacc. (1942). However when mature ma-
terial was finally obtained it was obviously distinct from these forms.
St. ioliicola is abundant in parts of nortberpJEurop/. Bubak (1915)
raised the fungus to specific rank. St. vexata,Jrora which it was re-
moved, has large coarse spores. Another variety of this fungus, St.
vexata var. baldingerae Sacc, is only a saprophytic phase of St. folii-
cola. Cylindrosporium phalaridis (F. Columb. 5011; Sacoardo, 1915,
FIG. 57.- -A, Pycnospores of Stagonospora }oliicola on Phalaris arundinacea.

Mandan, N. Dak.; B, pycnospores of St. ischaemi on Andropogon jurcatus, Okla.,
X2,000.
p. 125) is an insect-eaten collection of the same species. True pycnidia

which escaped the insects occur in the type. The insects had eaten the
ostiolar ends of the pycnidia reducing them to acervuli!
St. foliicola produces a slow-growing, white cottony growth with
pink color in the substratum. In comparison with St. arenaria it is
slow-growing. No spores were obtained in pure culture on melilotus
culms held at common storage conditions (Sprague, 194d,a).
The disease caused by this fungus was extremely prevalent in late
summer at Mandan, North Dakota, during years of abundant precipi-
tation. The fungus was found there on upland seedings in plots and in
native stands in the Missouri River bottomlands. I t was common in
marshes in Minnesota and South Dakota. Some distinct differences in
the susceptibility of clones of reed canary grass were noted in upland
plots at the Northern Great Plains Field Station (Sprague, field notes).
The differences were striking enough to warrant consideration in select-
ing for increase.
St. foliicola was found once, probably an accidental occurrence,
on Glyceria in Colorado (Sprague, 1949,a). Except for a strong brown
border around the spot, the fungus and the disease were identical with
the material on Phalaris.
/1 .
Range: United States, Canada, Czechoslovakia, France, Italy, Latvia.
References: Bubak (1915); Davis (1915,6,-.p. 98;' 1942); Saccardo (1915,
p. 125); Sprague (1941,c; 1946,a,e; 1949,a).' '
„l
Stagonospora glycericola Sprague—Leaf Rot
Spots none to straw color, vague at best; pyenidia globose, erum-
pent, nongregarious, golden brown in transmitted light, 90-140 /t diam.;
pycnospores subcylindrical to subfusiform, constricted to noncon-
stricted, apex and base subobtuse, context hyaline, globulate, 1- to
3-septate, 22-30 X 4.6-6.4 fi.
On: Glyceria grandis S. Wats., N. Dak.—G. striata (Lam.) Hitchc, Minn.
St. glycericola differs from St. subseriata (Desm.) Sacc. in its dis-
tinctly less broadened spores. St. glycericola is similar to St. brachy-
podii Died, but the latter has darker pyenidia and yellow spores and,
as Saccardo says, is close to Rendersonia-culmicola Sacc. (Sylloge V. 5,
p. S67). Comparing St. gl-ycericolo, with. St. hrachyelytri the spores of
the two species are somewhat similar /with the exception that the
species on Glyceria has wider spores with coarse globulate contents.
St. glycericola is a saprophyte, p.t least the material which we
studied did not show very much evidence that the lesions had devel-
oped in living tissue.
Reference: Sprague (1948,o).
Stagonospora graminum Sacc. and Scalia—Leaf Rot

Pyenidia gregarious on dead leaves, subcutaneous, globose to
slightly ellipsoid, ostiolate, 275-380 X 160-200 /t, context parenchyma-
tous, chestnut brown; pycnospores oblong,' 5- to 7-septate, noncon-
stricted at the septa, 23-28 X 6.5-8.0 /j., yellow hyaline.
On: Deschampsia atropurpurea (Wahl) Scheele, Alaska. ^"^ --,^__
This species is another of those saprophytic forms which invade
frosted or otherwise necrotic tissue. It is probably very close to St. sub-
seriata (Desm.) Sacc. in its morphology. It 'will probably be found in
the higher mountains of the western United States.
Range: Alaska. ,
Reference: Saccardo, Peck, and Trelease (1904).
/
Stagonospora intermixta (Cooke) S a c c . ^ ^ a f and Culm Rot
Pyenidia thin-walled, size not given, spores fusoid-elongate, hyaline
or tinted, 50 X 3 ;a (on Milium multifiorum Cav., in type), 30-40 X 3 /*
(Greene, 1944,a on Agrostis alba L.), 7-septate.
On: Agrostis alba L., Wise.—Cinna arundinacea L., Wise.—C. latifolia
(Trev.) Griseb., Wise.—Poa arachnifera Torr., Miss.
Except for the fact that the spores in this group are 7-septate, this
species could probably be ignored in this text. Possibly St. intermixta
and St. arenaria are not distinct. In Davis' final text (1942) he deleted
A. alba from the list of hosts of St. intermixta but Greene (1944,a)
found abundant material on this host at Eagleville, Wisconsin. The
pycnidial walls were not well defined in this material, the spores, a s '
mentioned, were 30-40 X 3 /* and 7-septate. Davis (1942) also listed
this fungus on the two species of Cinna and on Phalaris but the last
mentioned was transferred to St. foliicola. The collection on Cinna
arundinacea was discussed by Davis (1915) as follows: "I have not
seen an authentic specimen of this species. The specimens which I have
referred here have depressed-globose pycnidia, 40-60 fi in diameter with
a round apical pore which is surrounded by a thick black ring. The
long fusoid sporules are 7-septate, 40-60 X 3.5-5.0 jx." The small size
oi the pycnidia must he a typographical error for 140-160 fi. Davis
(1924) collected material on C. latifolia with 7-septate spores as long
as 90 X 4 ;n. The pycnidia were pale and thin-walled, except around
the ostiole. Phyllachora occurred on the leaves also.
Range: United States, England.

References: Cooke (187S); Davis (1915, 1924, 1942); Greene (1944,a);
Saccardo (Sylloge, 1884, V. 3, p. 454).
Stagonospora ischaemi Sacc.—Leaf Spot

Lesions narrow, tawny, pycnidia grouped in lines, erumpent, globose,
papillate, ostiolate, parenchymatous, brown; pycnospores cylindrical to
oblong, at first 4-guttulate, 1-septate, finally 3-septate, nonconstricted
at the septa, hyaline, 12-18 X 2.8-4.1 ,j..
On: Andropogon furcdtm Muhl., Okla.—A. hallii Hack., Okla.
Preston listed this fungus as present on two species of Andropogon
in Oklahoma (1945). The writer has seen some of this material col-
lected by K. Starr Chester and forwarded by W. W. Ray. The spores
are spindle-shaped, flattened on one side, subhyaline, somewhat con-
stricted at the 3 septa, 15-18 X 2.8-4.1 /x, with usually a pair of small
oil drops on each side of the cross walls (Fig. 57,5).
The spindle-shaped Ascochyta material on A. furcatus (cf. A.
brachypodii) may be a phase of this fungus but it is doubtful if this
is true because of the comparatively large size of the Ascochyta stage.
Preston also lists Sorghastrum nutans (L.) Nash (1945) as a host
for Stagonospora sp. in Oklahoma.
References: Preston (1945); Saecardo (1878; Sylloge, 1884, V. 3, p. 454).
/i
Stagonospora maculata (Grove) Sprague—[Purple Leaf Spot
Stagonospora subseriata yar. maculata Groye
Lesions blackish brown to purple, scattered to] confluent, pycnidia
in lines in the blotches, globose, golden brown, | |)stiolate, 130-200 jm
diam.; pyenophores cylindrical, 4-6x1-8-2.2 n ([Fig. 59,C); pycno-
spores fusiform, 3- to 4-septate, yellow with coarse contents, strongly
constricted at the septa, bases and apices tapering but eventual tip
blunted at the base, sharper at the apex, 27-40 X 4.8-6.5 {i, mean size
3 3 x 5 . 5 j a (Fig. 58,B;59,B).
On: Dactylis glomerata L., Md., Penn.
The writer raised this fungus to species rank because it differed in

morphology from St. subseriata (Fig. 5S,B,E; 59,D,E) as well as in its
very distinct parasitic nature and its production of characteristic purple
blotches on orchard grass leaves. /
This fungus is of some economic importance in Pennsylvania and
is recognized as one of the major diseases of orchard grass in the area.
Range: United States, England. ^
References: Grove (1935); Sprague (1941,6; 1948,a).
Stagonospora maritima Sydow—On Manisuris

1
Pycnidia frequently sparse, not in spots, hypophyllous, globose,
black, 100-130 /t diam., ostiole 15-20 /x diam., context of pycnidia paren-
chymatous, fuliginous; pycnophore short, pycnospores oblong, some-
what irregular at times, ends obtuse, at first radltiguttulate, finally
2-septate, hyaline, 16-20 X 4.0-6.5 /x, central 'cell somewhat longer than
other ones. '
On: Manisuris spp., Tex.

This species was originally described on sedge from Germany (cf.
Saccardo, Sylloge, 1931, v. 25, p. 365, or Sydow Myc. Germ. Fasc. XXI
no. 1025 listed in Ann. Myc. 9, p. 537, 1911). Weiss (1945) listed this
species on Manisuris spp. from Texas. We hav(; not found the source
of this report and are not familiar with the s|)ecies. Manisuris is a
genus with 5 species listed in Hitchcock (1935) and belongs in the
Andropogoneae. It is suspected that the fungus on Manisuris is another
saprophyte referable, perhaps, to an earlier name than St. -maritima.
Range: United States, Germany. --^_ ^

References: Saccardo (Sylloge, 1931, v. 25, p. 365); Weiss (1945).
FIG. 58.—Pycnospores of: A, Stagonospora paspnli on Paspalum sp., N. Car.;

B, St.^mdculata on Dactylis glomerata, Laurel, Md.; C, St. bromi on Bromus
carinatus, Ann Arbor, Mich.; D, St. bromi on Bromus cilialus, Lake Itasca,
Minn.; E, St. subseriata on Poa pratensis, near Bemedji, Minn.; F, Septoria
avenae or St. arenaria on Oryzopsis hymenoides, Terry, Mont.
Stagonospora paspali Atk.—Leaf Spot

Spots elliptical to irregular, off-white to buff with purple margins,
sometimes confluent, pycnidia applanate-globose, spores hyaline, 2-sep-
tate, finally 3-septate, oblong-elliptical, ends tardily rounded, 24-
28X7-9/X (Eig. 58,A).
On: Paspalum dilatatum Poir, Ga., N. Car., Tex.—P. floridanum var,
glabratum Englm.,-JS[. Car.—P. laeve Michx., Alabama—Trioda flava
(L.) Smyth, Va.
This species is common along the Atlantic and Gulf Coasts from
North Carolina to Texas. The fungus appears to be parasitic.
Reference: Atkinson (1897, p. 33). . _
Stagonospora simplicior Sacc. and Berl.—Leaf Spot

Spots elliptical, pycnidia densely seriate, subglobose, ostiolate, black,
80-200 fi.; pycnophores short cylindric, 8 x 5 - 5 / ^ ; pycnospores oblong-
fusoid, straight or slightly bent, moderately, slightly, or scarcely con-
stricted at the septa; 3-septate with coarse contents, somewhat taper-
ing toward the broad ends, 24-44 X 9-111^-
On: Andropogon jurcatus Muhl., N. 'D&k.—Sorghastrum nutans (L.) Nash,
Wise.—Stiâ columbiana Macoun, Wyo".—S.- colun^biana var. nelsordi
(Scribn.) Hitchc, Calif., Utah—S. lemmonii (Vasey) Scribn., Oreg.—S. let-
termanii Vasey, Colo.—S. occidentalis Thurb., Oieg.—^S. vindtda-Trin., N.
Dak.—S. williamsii Scribn., Wyo.—Stiporyzo'psis bloomeri (Boland) John-
son, Calif.
i
Material sent to us by Greene from Wisconsin appears to be para-
sitic. St. simplicior Sacc. and Br., has coarse spores, 32-38 X 8-9 /x, in
the type on Phragmites communis Trin. in France (Sylloge, v. 10, p.
336). Greene (in manuscript) has also found material on Panicum spp.
which is apparently parasitic. Most of the Wisconsin material appears
to be parasitic and probably this fungus is of some economic im-
portance. Some North Dakota material was saprophytic such as a col-
lection on Calamovilja longifolia (Hook.) Scribn. from Mandan, North
Dakota, which had faintly yellow spores 28-35 X 8-9 fi (Fig. 59,A)
borne on blunt pycnophores in somewhat gregarious, black, globose,
ostiolate pycnidia, 100-140 /x, diam. This was an early autumn invader
of dead tissue, the lesions being vague gray or stramineous blotches.
We have also seen material, probably ,of the variety andropogonis
Sacc. on Andropogon jurcatus at Mandan. The pycnophores were short
cylindric and the spores 37-44 X 10-11 /*. Greene (1946,6) reported
spores on Andropogon from Wisconsin measuring 40-55 X 8-13 /*.
Alpine material on Festuca ovina var. hrachyphylla has 1-septate,
finally 3-septate, spores 28-35 X 7.4-10 ju, more or less cylindrical while
a specimen of Melica bulbosa Geyer from Waterton Lakes National'
Park, Alberta, has spores 26.8-36 X 9.8-12.8 ju,. This one occurs with
Septoria nodorum. These alpine collections are saprophytic phases of
St. simplicior proper, although the one from Waterton Lakes is some-
what parasitic. Material on Stipa is, however, parasitic (Sprague,
1950,a). j
Range: United States, France. |
References:'Greene (1944,6; 1946,6; 1949); Saccardo (1916; Sylloge, 1892,
V. 10, p. 336, and 1921,'v. 25, p. 366); Sprague (1946,a,eT^95Q,a|.
Stagonospora spartinicola Sprague—Streak Spot

Spots none to straw-colored, drab or faintly buff to light tan,
elongate, sometimes very narrow, 60 X 1-5 mm/ diam.; pycnidia flat-
tened globose, sometimes elliptical in outline, immersed with wide
(80 ix) ostiole protruding through a slight rupture in the host'epidermis,
parenchymatous, the wall thin, pale brown to nearly hyaline toward
the apex, 150-280 //,; pycnospores cylindrical, constricted at the septa,
ends rounded, one end more so than other, .centents coarse including
one large chlorine guttula in each cell, spores usually immature in
summer and fall collections, 3-4-5-septate, 40-47 X 6-9 jx.
On: Spartma pectinatatLk., N. Dak. (type),^Wisc.
FIG. 59.—A, Pycnospoies of Stagonospora simplicior var. andropogonis on

Calamovilja longijolia, N. Dak.; B, pycnospores of St. subseriata on Elymus
mollis, Manhattan, Oreg.; C, cross section of pycnidial wall of Stagonospora
. maculata on Dactylis glomerata from State College, Pa.; D, pycnospores of St.
•subseriata on Deschampsia caespitosa, Lyndhurst, Hampshire, Engl.; E, pycno-
spores of St. maculata on Dactylis glomerata, State College, Pa.
We have a fragment of this fungus collected at Wilton, North

Dakota, September 14, 1941 (B.P.I. 80,798), and similar material sent
by H. C. Greene in 1949 from Madison, Wisconsin. Weiss (1945) re-
ported Stagonospora sp. on Spartina from Maryland.
While related to the St. vezatula (not to be confused with St.
vexata) material this fungus is recognizable as distinct because of its
wider, coarser appearing spores (Sprague, 1950,a). The material sent
by Greene shows some evidence of being the result of parasitic action.
The reader is referred to the complex discussed under St. vexatula
for comparable fungi.- Determination of any of these saprophytic and
mildly parasitic species of Stagonospora is difficult and morale-
destroying.
Reference: Sprague (1950,a, probably will appear in Greene's series).
Stagonospora subseriata (Desm.) Sacc.—Leaf Rot

Hendersonia subseriata Desm.
Pycnidia erumpent, somewhat gregarious, small, black to dark
golden brown, globose, ostiolate, spores boat-shaped, 20-45 X 6-8 /*,
3- to 4-septate, constricted or nonconstricted, hyaline.
/ : •
On: Danthonia intermedia Vasey, Colo.—Elymus', \nollis Trin., Oreg.—

Phleum pratense L., Minn.—Poa pratensis L., ._Minn.
This has boat-shaped spores, sometimes but not always constricted
at the septa (Fig. 59,B,D). We have found it in,'a few places in the
western United States on dead grasses. It was originally described on
Molinia (Enodium) caerulae (L.) Moench in Belgium and Germany.
Petrak discussed it on Dactylis (1927) but he may have been dealing
with St. maculata.
A collection on Danthonia from Colorado is associated with a
species of Leptosphaeria (A. S. 20,071).
Range: United States, Belgium, Germany.
References: Petrak (1927); Saccardo (Sylloge, 1884, V. 3, p. 454); Sprague
(1941,6; 1948,a).
Stagonospora vexatula Sacc. and Other Saprophytic Species

Description of Si. vexatula:
Pycnidia gregarious in fuscous spots, Surrounded by a buff or yel-
lowish halo, pycnidia innate-erumpent, globose but slightly elongated
into a semicone-shaped or mammiform-âpex, about 200 ft diam., some-
times smaller but reported up to 500 /* diam., black, firm; spores cylin-
drical, ends rounded, 35-38 X 4.5-5.0 )x. or longer (to 55 /A) , 3- to 7-sep-
tate, hyaline.
On: Deschampsia caespitosa (L.) Beauv., Ida.—Phragmites communis Trin.,)
N.J.
A considerable number of species of Stagonospora occur on dead
culms, leaves and sheaths of grasses, especially on Joarse marsh grasses
such as Phragmites communis. In his check jlist, Weiss (1945) writes
about Stagonospora graminella Sacc. from North Dakota as follows:
"The Hendersonia-Stagonospora complex on Phragmites can not be
resolved without the study of cultures. Sonle reports of Hendersonia
arundinacea .should perhaps be referred to S. vexata SStcv-Qjie or more
of these appear to be the conidial stage of Leptosphaeria arundinacea
(Sow. ex Fr.) Sacc." Weiss also lists St. phragmites Ell. aifd Ev. on
leaves from Indiana, associated with Mycosphaerella sp. W. B. Cooke
sent material of Phragmites to us from New Jersey which was affected
with St. vexatula Sacc. (Sprague, 1950,a). The/fungus was an early
invader of necrotic or dead tissue in the fall o'f the year. The spots
consisted of small fuscous, more or less linear lesions surrounded by a
buff halo. The spores were hyaline, stoutly obclavate, frequently curved,
25-53 X 3.0-4.8 [i, mostly 1- to 3-septate in brown-pycni/ia, 140-180 [i.
The type of this species was described w i t h - s i ^ e s 35-38 X 4.5-5.0 /j.
(Saccardo, Sylloge v. 3, p. 454) and the spores were 5- to 7-guttulate.
Petrak gave the spores as 18-56 X 3.5-5.0 /x, (1927) in very large pyc-
nidia, 400-500 [i diam., while Saccardo gave the measurements as about
200 [X. diam. The material from New Jersey, as above mentioned, had
pycnidia smaller than those reported by Saccardo. The New Jersey
fungus had some resemblance to St. foliicola. Perhaps detailed study
will show that St. foliicola and St. vexatula are developing phases of
St. vexata from which the first mentioned was removed by Bubak
(1915).
A number of other species of Stagonospora described on Phragmites
should be mentioned at this time. They all appear to differ from St.
vexatula. One of these is St. elegans (Berk.) Sacc. and Trav. and it has
spores with prominent guttulae, 60-90 X 9-10 jx, resting on a gelatinous
substratum in the coarse pycnidium. On the basis of the description
(Sylloge V. 20, p. 878) this species and St. dolosa Sacc. and Roum.
(Sylloge, 1884, v. 3, p. 455; Petrak, 1927), which Grove says is a
synonym of St. elegans (1935, pp. 356-357), are distinct from St.
vexatula. St. hysterioides (Karst.) Sacc. (Karsten, 1884,6) and the
similar St. hydrophila Br. and Har. (Sylloge v. 10, p. 336) appear to
have too short and proportionately thick spores ever to develop into
St. vexatula. St. simplicior, which we have discussed, has thick fusiform
or roughly cylindrical spores. St. neglecta (West.) Sacc. (Sylloge, 1884,
V. 3, p?455) has fusoid, 3-septate spores. St. graminella (subsp.) Iqphi-
oides Sacc. has 4-nucleate spores, 15-16 X, 3 /* (Sylloge, 1884, v. '3, p.
454), while St. mucipara Sacc. (Sylloge, 1899, v. 14, p. 964) may, per-
\ haps, be a young phase of St. vexatula, as the 3-septate spores are
25-30 X 3.2 IX.. They are tinted yellow, however. Material on Agrostis
oregonensis Vasey from Idaho County, Idaho, had yellowish hyaline
cylindrical 3-septate guttulate spores 18-22 X 3-5 ju,. This was assigned
to St. graminella Sacc. Another species, St. arundinis (Lev.) Sacc, is
not available for comparison and the description is too vague to de-
termine even approximate relations. The spots are said to be yellow,
fuscous or none, and the spores hyaline and septate. Originally de-
scribed as Dothidea arundinis Lev. (Leveille, 1842), it probably repre-
sents a smaller species than St. vexatula. Brenckle (1918) listed St.
graminea Sacc. on Phragmites from Kulm, North Dakota, probably a
typographical error from St. graminella Sacc. The latter is listed by
Fairman on Sporobolus auriculatus Vasey from New Mexico (1918, p.
259). Zeman (1921) reported it on Phalaris bulbosa from Argentina.
St. phragmiticola Hansford was described on Phragmites communis
from Uganda in 1945 (Proc. Linn. Soc. 157).
Consideration of the complex on Phragmites involves not only some
16 entities in Stagonospora but a number in Septoria as well. Septoria
arundinacea (Sylloge, 1884, v. 3, p. 564) causes oblong, amphigenous,
sordid, ochre spots with fuscous borders. The pycnidia are globose-
lenticular, 140 ju, diam. with bacillar spores, 60-70 X 5-6 ju,, 6- to 7-sep-
tate. On the basis of the description Cooke's material from New Jersey
differs in not having bacillar spores, in having the spots fuscous, the
284 • DISEASES OF CEREALS AND GRASSES
margins yellow, and the pycnidia globose. Septona littoralis Sacc. is
' • ' . .
closer to Stagonospora vexatula. It has 3-septate spores, 50-65 X 3.5-

4.0 [I, which are cylindrical, straight to curved with rounded apex and
a subtruncate base borne on ampulliform pycnophores (Sylloge, 1884,
V. 3, p. 565). Gonzalez Fragoso (1920) described) Septoria littoralis f.
culmicola with spores 55 X 4 jn. This form is even closer to St. vexatula.
Septoria paludosa Kabat and Bubak (Sylloge, 1906, v. 18, p. 395) on
Phragmites is associated with a leaf spot. The lesions are oblong linear,
delimited, gray brown; the pycnidia 150 jj. diam.; the numerous spores
are exuded in rose-colored cirri and are bacillar, 8- to. 12-septate, 40-
70 X 4.0-4.5 fx. Septoria phragmites Sacc. (Sylloge, 1884, v. 3, p. 564)
has spores 20-30 X 1.5-2.0 ju, and is distinct from St. vexatula. Septoria
arundinis (Mont.) Sacc. (Ascospora arundinis Mont.) has oblong-
linear, 3-septate, hyaline spores 20 X 3 iX. Since critical study with
type material plus pure culture work is needed before this group can
be reduced to a working number of species they are simply permitted
to pass briefly in review here in relation to/the material on Phragmites.
The material in question (W. B. Cooke, 17,079) is left in the species
to which it seems morphologically related, St. vexatula.
In the course of routine examination of leaf specimens we have
noted fragmentary material from time to time of a considerable num-
ber of Stagonospora forms on dead or dying grass in the fall of the
year, especially in western North Dakota following frosts. Many of
these collections were not recorded because of the scantiness of the
material and because they were obviously of no economic importance.
A few collections were kept and camera lucida drawings made of sonie
of them. A specimen on Deschampsia caespitosa (L.) Beauv. from the
vicinity of Corvallis, Oregon (O.S.C. 51), bore, among other fungi, a
Stagonospora-like fungus with the pycnidia sur^rounded by a prosen-
chymatous matrix. The spores were short cj^lindric, somewhat rounded,
and slightly tapering, with heavy walls, yellow with a few nearly brown,
20-24 X 6.7-8.0 ix. Possibly this fungus belongs in Sclerostagonospora
V. Hoehn. (1917). Petrak (1925, p. 4), however, does_not see very much
difference between this' genus and Stagonospora. Clements-and Shear
also give Sclerostagonospora secondary status (1931). Recently col-
lected spots of Stagonospora on D. caespitosa from Plummer, Idaho
(C.S. 3969), have cylindrical spores 30-40 X 4.5-5.5 /^ borne in vague
spots on necrotic leaves. The spores were 3-septate but immature.
These are assigned to St. vexatula. The fungusfis, in this case', weakly
parasitic. /
Preston (1945) reported Stagonospora sp. on Axonopv^ compressus
(Swartz) Beauv. from Oklahoma. We have seen material of Stagono-
spora on this host from Poplarville, Mississippi^with 5-septate spores
20.8-25.6 X 6.4-9.6 [i. It was distinct from St.'paspali]
References-: For this miscellaneous group the reader is referred to the
various volumes of Sylloge Fungorum and to Grove's text (1935).
MELANCONIALES
The Melanconiales, holding an intermediate position, morphologi-
cally, between the Sphaeropsidales and the Moniliales, are somewhat
more difficult to classify than the other two orders of the Fungi Imper-
fecta Grove (1935) placed the Sphaeropsidales and the Melanconiales
in the Coelomycetes, the leaf spot fungi. Some genera of the Sphaerop-
sidales have poorly formed apical portions of the pycnidium so that
it resembles an acervulus. But at the other extreme the acervuli may
have no walls at all and are composed only of stromatic masses of
mycelia which become thick enough to break the epidermis of the host,
forming pustules.
A key to the few representatives of the Melanconiales on Gramineae
is as follows:
A. Acervuli setose Colletotrichum
AA. Acervuli not setose ;
B. Spores elongated
C Spores filiform, 0- to several-septate Cylindrosporium
\ CC. Spores broadly obclavate or fusoid Septogloeum
BB. Spores somewhat oblong '. Gloeosporium
Colletotrichum falcatum Went—See Physalospora tucumanensis

Speg. for details of this organism
C. falcatum (C. lineola Corda var. halepense Heald and Wolf) has
been combined with C. graminicola (Ces.) G. W. Wilson, but the com-
paratively recent finding of the perithecial stage of the southern an-
thracnose, fungus has thrown doubt on whether it is desirable to in-
clude both C. falcatum and C. graminicola under P. tucumanensis. If
the two are distinct it is probable that C. falcatum extends north into
the range of C. graminicola. C. falcatum has been reported as far north
as Minnesota. We have seen material of anthracnose on sudan grass at
Lincoln, Nebraska, that seems to answer the description of C. falcatum.
A study of a brief tabular summary of the morphology of 'these two
species will disclose their similarity:
Size of Setae
Name Size of Spores Shape of Spores and Septations
C. falcatum. . . 20-27 •>< 4-5 n falcate, acute pointed up to 75 n; 1-
to 2-septate
C. graminicola. 18-26 X 3-4 11 falcate or somewhat 60-120 M; 1- to
spindleform 2-septate
Reference: See Physalospora tucumanensis Speg.
, 285
/i
CoUetotrichum graminicola (Ces.) Wils.—Anthracnose
C. cereale Manns ^"" I '
Acervuli dark brown or black, elongate; setae few or many, dark
brown or black, 1- to 2-septate, 60-120 jx. long, and 6-8 fn thick at the
base; conidiophores very short, 6-12 X 1-2 /*; conidia falcate, spindle-
or boat-shaped, 2- to several-guttulate, 18-26 X 3-4 /x.
On: Agropyron desertorum (Fisch.) Sclmlt., Minn., N. Dak.—A. elongatum
(Host.) Beauv., Okla.—A. intermedium (Host.) Beauv., Okla.—A. repens
(L.) Beauv., Minn., Nebr., N. Dak., Penn., Tex.—A. smithii Rydb., Okla.
—A. trachycaulum (Lk.) Malta, N. Dak.—Agrostis alba L., Ida., Iowa,
Minn., Ohio, Wise.—A. exarata Trin., Wash.—A. palustris JInds., Wash.—
A. scabra Willd., Colo.—Alopecurus aequalis Sobol, Minn.—A. pratensis
L., Minn.—Ammophila breviligulata Fernald, Wise.—Andropogpn furcatiis
Muhl., Colo., Nebr., N. Dak.—A. hallii Hack., Miss.—A. scoparius Miehx.,
Okla.—Anthaenantia rufa (Ell.) Sehult., Fla.—Anthoxanthum odoratum
L., Oreg.—Aristida dichotoma Michx., Okla.—Arrhenatherum elatius (L.)
Beauv., Minn., Ohio, Penn.—Avena Jatud L., N. Dak.—A. hookeri Seribn.,
N. Dak.—A. sativa L., Alta., Ark., 111., Iowa, Kans., Ky.,. Mo., N. J.,
N. Y., N. Dak., Ohio, Okla., Ont., Oreg., Penn., P. E. Is., Que., Sask.
(Weiss lists the range as "general")—Beckmannia ^syzigachne (Steud.)
Fern., Minn., Nebr., N. Dak.—Brachypodium mucronatum, Willk., Md.—
Bromus carinatus Hook, and Am., Okla.—B. catharticus Vahl, Tex.—B.
inermis Leyss, Nebr., Que.-—B. latiglumis Hitehc., Wise.—B. purgans L.,
Iowa—B. secalinus L., N. J., Ohio, Penn., Tex.—Buckloe dactyldides
(Nutt.) Englm., Nebr.—Calamagrostis canadensis (Miehx.) Beauv., Ida.,
Minn., Mont., N. Dak., Ohio, Wise.—C. inexpansa A. Gray, Mont., N.
Dak., Wash.—C. purpurascens R. Br., Mont.—C. rubescens BuckL, Mont.,
Oreg., Wash.—Calamovilfa gigantea (Nutt.) Scr'ibn. and Merr., Okla.—
C. longijolia (Hook.) Seribn., Wise.—Cendhrus'pauciflorus Benth., Nebr.
—C. tribuloides L., Del.—Chloris verticilldta Nutt., Okla.—Cynodon dac-
tylon (L.) Pers., Okla.—Dactylis glomerdta L., Ky., Md., Minn., Miss.,
N. J., N. Car., Ohio, Okla., Oreg., Va.—Danthonia californica BoL, Ida.,
Wash.-^jDi'arrAena flmericana Beauv., Okla.—Digitaria filiformis (L.)
Koel., Okla.—Z). sanguinalis (L.) Scop., N. J., N. Dak., Gl^S^—Êchinochloa
crusgalli (L.) Beauv., Ga., Kans., La., Miss., Mo., Ohio,'D.kla., W. Va.,
Wise.—Eleusine indica (L.) Gaertn., Okla.—Elymus canadensis L., Wise.—
E. junceus Fisch., Okla.—E. virginicus -L., Okla.—Eragrostis curvula
(Schrad.) Nees, Okla.—iJ. trichodes (Nutt.) Wood, Okla.—Eremochloa
ophiuroides (Munro) Hack., Fla.—Festuca elhtior L., Ky., N. Y., Okla.,
Penn.—F. megalura Nutt., Wash.—F. octoflora Walt., N. Dak.—F. ovina
L., Minn.—-F. rubra L., B. C, Minn., Penn., Va., Wash.—F. subulata
Trin., Ida.—F. versuta Beal, Okla.—Glyceria striata (Lam.) Hitche.,
Wise.—Hierochloa odorata (L.) Beauv., Wisc^TrzHolcm lanatus L., Ind.,
Md., Penn., Wash.—Hordeum bidbosum -IxfTDkla.—i7. pusillum Nutt.,
Kans.—H. vulgare L., Alabama, Ga., Ind., Iowa, Md., Minn., N. J., N. Y.,
N. Car., Okla., Penn., S. Car., Tex., Va.—Leptoloma cognatum (Sehult.)
Chase, Okla.—Lolium multiflorum Lam., Oreg., Wash.—L. perenne L.,
FUNGI IMPERFECTI—MELANCONIALES 287
Oreg., Penn.—Melica nitens (Scribn.) Nutt., Okla.—Milium efjusum L.,
Mieh.—Muhlenbergia capillaris (Lam.) Trin., Okla.—M. mexicana (L.)
Trin., Okla.—M. racemosa (Michx.) B. S. P., Okla.—M. schreberi Gmel.,
Okla.—Oryzopsis micrantha (Trin. and Rupr.) Tliurb., N. Dak.—Panicum
boscii var. molle (Vasey) Hitchc. and Chase, Okla.—P. capillare L.,
Minn.—P. hemitomon Schult., Fla.—P. implicatum Scribn., Wise.—P.
tennesseense Ashe, Nebr.—P. virgatum L., Alabama, Kans., La., Minn.,
Wise.—P. sp., Iowa—Paspalum dilatatum Poir, Fla., Ga.—P. stramineum
Nash, Okla., Wise.—Phalaris tuberosa var. stenoptera (Haek.) Hitchc.,
Okla.—Phleum pratense L., 111., Ind., Minn., Mo., N. J., Ohio, Okla.,
Penn., Sask.—Poa ampla Merr., Okla.—P. canbyi (Scribn.) Piper, Okla.,
S. Dak.—P. compressa L., Minn., Okla.—P. palustris L., Minn.—P.
pratensis L., Minn., N. J., N. Y., N. Dak., Ohio, Penn., Que., S. Dak.,
Tex., W. Va.—Schizachne purpurascens (Torr.) Swallen, Minn., Wise—
Secale cereale L., Ark., 111., Ind., Iowa, Ky., Md., Mich., Minn., Miss.,
Mo., N. J., N. Y., N. Car., N. Dak., Ohio, Okla., Penn., S. Car., Tenn.,
. Tex., Va.—Sorghastrum nutans (L.) Nash, Kans., Okla., Wise.—Sorghum
halepense (L.) Pers., confused with C. falcatum so that the range on this
host is uncertain, possibly N. Car. to Ind. and Iowa—S. vulgare var.
sudanense (Piper) Hitchc, Ga., Mo., Nebr., N. Dak., Va., Wise, probably
widespread but confused at present with C. falcatum—Sphenopholis ob-
tusata (Michx.) Scribn., N. Dak.—Sporobolus asper (Michx.) Kunth,
Okla.—S. clandestinus (Spreng.) Hitchc, Okla.—S. heterolepis A: Gray,
N. Dak., Okla.—Triodia albescens Vasey, Okla.—T. elongata (Buokl.)
Scribn., Okla.—r. flava (L.)'Smyth, Alabama—Tripsacum dactyloides L.,
N. Y., Okla., S. Car., Va.—Trisetum canescens Buekl., Wash.—T. cernuum
Trin., Oreg.—T. flavescens (L.) Beauv., Mich.—Triticum aestivum L.,
Alta., 111., Iowa, Mo., N. J., N. Dak., Ohio, Okla., Que., Sask.—T. dicoc-
cum Desf., N. J.—Zea mays L., Conn., Ill, Ind., La., Md., N. J., N. Car.,
Ohio, Penn., S. Car., Va.
As indicated, the host range of this fungus is very great. The symp-
toms on many of these hosts are obscure, often appearing only as a
light mold on the leaves. Bolley (1903) reported Colletotrichum, on the
roots of wheat from North Dakota but this fungus was apparently
Gloeosporium bolleyi Sprague (1948,6). C. graminicola has, however,
been reported on the roots &f cereals in Canada (Sanford, 1935) and
in Germany (Winter, 1940,d).
Lobik (1933) described C. zeae on spots caused by Ascochyta zeina.
The former produced a mycelium on which arose cylindrical, hyaline
conidiophores and dark brown septate setaC The spores were hyaline,
short clavate or oblong ovate, 8.4-18.2 X 4.9-5.6 ix.
Padwick and Henry ,(1933) listed Colletotrichum sp. as isolated
from the roots of Agropyron repens in Alberta. C. graminicola is com-
mon on above-ground parts of grasses in the area and no doubt
Gloeosporium bolleyi is also present. The latter is common from North
Dakota to Washington and into Saskatchewan and probably occurs in
Alberta.
/I
C. graminicola produces mycelium in pure culture without mucose
development of spore masses. Setae are sparingly produced in culture.
Bruehl (1948) reported that a group of isolates of C. graminicola on
Sorghum spp. were more tolerant to higher temperatures than isolates
from cool-temperature cereals and grasses,- Seedling blight on sorghum
developed well at any temperature between 16° G and 32° C.
Range: United States, Australia, Burma, Canada, China, France, Germany,
Gold Coast, India, Italy, Peru, Tanganyika, Uganda.
References: Boening and Wallner (1936,&); BoUey (1913); Bruehl (1948);
Chowdhury (1936); Ellis and Halsted (1888); Jennings-(1890); Koehler
(1943); Rosen (1947,c); Sanford (1935); Schwarze (1917); Selby and
Manns (1909); Wilson, G. W. (1914); Winter (1940,d). (See also U.S.D.A.
Tech. Bull. 1005, just received.)
Cylindrosporium bambusae Miyake and Hara—On Culms

Spots on the culm are at first elliptical to lanceolate or irregular,
later confluent and sometimes occupying the entire inter node; gray
with a dark brown margin; the pustules elliptical, black, not prominent,
0.8 mm. long, bearing subepidermal acervuli, 200-250 n diam.; conidio-
phores straight, simple, 4 - 5 x 1 /t;-conidia filiform, ends rounded,
curved, or flexuous, 15-17 X 1 z^-
On: Bamboo, Ga.
Weiss (1944) listed this species on bamboo from Georgia. It was
described on the culms of Phyllostachys bambusoides Sieb. and Zucc.
from Kyoto, Japan, in 1912 (cf. Saccardo, Sylloge, 1931, v. 25, pp.
623-624). /
Range: United States, Japan. J
Reference: Saccardo (Sylloge, 1931, v. 25). ,
Cylindrosporium calamagrostidis Ell. and Ev.—Leaf Spot

Spots linear, white, purple-margined; acervuli epiphyllouS", seriate,
150-200 ju, diam.; conidia filiform, curved, 40-72 X 1-5-2.5-/x, multi-
nucleate. ^,
On: Calamagrostis canadensis L., Wise.—Muhlenbergia filiformis (Thurb.)
Rydb., Utah. ( '
This species was collected in 1893 by J. J. Davis. The western ma-
terial on Muhlenbergia (C.S. 3685) appears to be^the same fungus
(Sprague, 1948,c). The spots on the tiny yellowed leaves were pale
brown with faintly darker borders. They boie^live-brown, seriately
arranged acervuli, 80-170 X 50-80 /*. These had definite walls but were
too ill-formed to be considered as eroded pycnidia. The spores were
hjralihe, guttulate, curved, filiform with slightly swollen bases. Their
general shape was whip-like and they measured 50-72 X 1.7-2.5 ;u,
sufficiently close to C. calamagrostidis to permit including in this
species (Fig. 60).
Moesz (1941) reported Cylindrosporium
arundinaceum on Calamagrostis arundinacea
from Latvia. This has not been seen and the
only available abstract (R.A.M. 25:474)
states only that it caused a small foliar
spot.
References: Ellis and Everhart (1893,6, p.
460); Moesz (1941); Sprague (1948,c).
Cylindrosporium glyceriae Ell. and Ev.—

Leaf Spot
Acervuli amphigenous, innate, 100-150 fx FIG. 60.—Conidia of
Cylindrosporium calama-
in elongated white spots with purple margins, grostidis on Muhlenbergia
0.4-0.6 X 1.5-2.0 cm., conidia cylindrical, 15- filiformis, Uinta Natl. For.,
30 X 2.5-3.0 IX, 3-septate, hyaline. Utah (C.S. 3685).
On: Glyceria canadensis (Michx.) Trin., Wise.—G. striata (Lam.) Hitchc,

Wise.
"^ This material was collected by Davis at Racine, Wisconsin. He re-
ported later that a collection made at Athelstane, Wisconsin, on G.
canadensis had longer spores than the type with an occasional one
being much longer. He gave no spore measurements, however (1915,
p. 99). I have not seen material of this fungus and have not compared
it with other species.
References: Davis (1915, p. 99); Ellis and Everhart (1893,6, pp. 459-460).
Gloeosporium bolleyi Sprague—Root Necrosis

On necrotic roots. Pseudopionnotes copious in pure culture, at first
mucose, pink to carrot red,' finally carbonaceous and black, mycelium
in soil gray, cottony; conidia numerous in culture, hyaline, sub-bean-
shaped to oblong, eguttulate at first, finally guttulate, 4-9 X J--7-4.0 fx,
mostly 5-8 X 1.9-2.5 ix.
On: Agropyron albicans Scribn. and Sm., N. Dak.—A. angustiglume Nevski,
N. Dak.—A. cristatum (L.) Gaertn., Mont., N. Dak., S. Dak., Wyo.—
A. desertqrum (Fisch.) Schult., N. Dak., S. Dak.—A. elongatum (Host.)
Beauv., N. Dak.—A. griffithsii Scribn. and Sm., Mont.—A. intermedium
(Host.) Beauv., N. Dak.—A. michnoi Roshev, N. Dak.—A. mongolicum
Keng, N. Dak.—A. repens (L.) Beauv., Minn., N. Dak., S. Dak.—A. ripa-
rium Scribn. and Sm., Wyo.—A, semicostatum (Steud.) Nees, N. Dak.—A.
/!
sibiricum (Willd.) Beauv., Mont., N. Dak., S. Dak.—A. smithii Rydb.,
Mont., Nebr., N. Dak., S. Dak., Wyo.^-r4 spicdtum (Pursh) Scribn. and
Sm., Wyo.—A. subsecundum (Lk.) Hitchc.,''Wyo.-'—^.trac/ij/cawZMm (Lk.)
Malte, Mont., Nebr., N. Dak., S. Dak., Wyo.—r4. trichophorum (Lk.)
Richt., N. Dak.—Agrostis alba L., N. Dak., S: JDak., Wyo.—A. scabra
Willd., N. Dak., S. Dak.—Andropogon jurcatus, Muhl., Nebr., N. Dak.,
S. Dak.—A. hallii Hack., Nebr., S. Dak.—A. scpparius Michx., Mont.,
Minn., Nebr.—Aristida longiseta Steud., Mont., Nebr.—Avena byzantina
K. Koch, Minn., N. Dak., S. Dak.—A. fatna L., N. Dak., Wash., Wyo.—
A. sativa L., Minn., Nebr., N. Dak., S. Dak., Wyo.—Bouteloua gracilis
(H. B. K.) Lag., Mont., Nebr., N. Dak., S. Dak., Wyo.—Bromus anomalus
Rupr., Mont.—B. arvensis L., N. Dak.—B. carinatus Hook, and Arn.,
N. Dak., Wyo.—B. erectus Huds., N. Dak.—5. inermis Leyss, Minn.,
N. Dak., S. Dak., Wyo.—B. japonicus Thunb., Mont., S. Dak., Wash.,
Wyo.—B. tectorum L., Mont., Nebr., N. Dak., S. Dak., Wyo.—Buchloe
dactyloides (Nutt.) Englm., N. Dak.—Calamagrostis epigejos (L.) Roth,
N. Dak.—C. montanensis Scribn., S. Dak.—Calamovilfa gigantea (Nutt.)
Scribn. and Merr., Nebr., S. Dak.—C. {ongifolia (Hook.) Scribn., Mont.,
Nebr., N. Dak., S. Dak., Wyo.—Cenchrus pauciflorus Benth., Nebr., N.
Dak., S. Dak.—Cynodon dactylon (L.)^Pers., N. Dak.—Dactylis glome-
rata L., Md., Mont.—Distichlis stricta (Torr.) Rydb., N. Dak., S. Dak.—
Echinochloa crusgalli (L.) Beauv., Wyo.—E. crusgalli var. frumentacea
(Roxb.) Wight, N. Dak.—Elymus ambiguus Vasey and Scribn., N. Dak.—
E. antarcticus Hook, f., N. Dak.—E. canadensis L., Nebr., N. Dak., S.
Dak., Wyo.—E. condensatus Presl, N. Dak., Wyo.—E. dahuricus Turcz.,
N. Dak.—E. dasystachys Trin., N. Dak.—E. excelsus Turcz., N. Dak.—
E. giganteus Vahl, N. Dak.—E. glaucus Buckl. N. Dsik.—E. interruptus
Buckl.,'N. Dak.—E. junceus Fisch., N. Dak.—E. macounii Vasey {Hordeum
jubatum x Agropyron trachycaulum), N. Dak.—B. pseudoagropyron
(Griseb.) Trin., N. Dak.—E. salsuginosus Turcz., N. Dak.—E. sibiricus L.,
N. Dak.—Eragrostis curvula (Schrad.) Nees, N/ Dak.—Festuca elatior
L., N. Dak.—F. elatior var. arundinacea (Schreb.) Wimm., Mont., Wyo.
—F. octoflora Walt., Nebr., N. Dak., S. Dak,—F. ovina L., Wyo.—F.
pacifica Piper, Wash.—F. rubra L., N. Dak.—Hilaria mutica (Buckl.)
Benth., N. Dak.—Hordeum brevisvbvlatum (Trin.) Lk., N. Dak.—H.
distichon L., Minn., Mont., Nebr., N. Dak!, S. Dak.^Wyo.—H. jubatum
L., Mont., N. Dak., S. Dak.—H. nodosum L., Mont^^'Br-Dak^î?. pusil-
lum Nutt., N, Dak., S. Dak., Wyo.—ff. vulgare K, Minn., Mont., Nebr.,
N. Dak., S. Dak., Wash., Wyo.—Koeleria cristata (L.) Pers., S. Dak.,
Wyo.—Melica scabrosa Trin., N. Dak.—If. virgata, N. Dak.—Muhlen-
bergia japonica Steud., N. Dak.—M. racemosa (Michx.) B. S. P., Minn.,
N. Dak.—Oryzopsis hymenoides (Roem. and Schult.) Rick., N. Dak.—
Panicum capillare L., N. Dak.—P. miliaceum Ll, N. Dak., S. Dak., Wash.
—P. tennesseense Ashe, Nebr.—P. virgatum L., N. Dak.—Phalaris arundi-
nacea L.,. Minn., N. Dak.—Phleum alpinum L., N. Dak.—P. phleoides
(L.) Karst., N. Dak.—P. pratense L., Minn., Nebr., I^ Dak., S. Dak.,
Wyo.—Poa bulbosa L., N. Dak.—P. canbyi ^^îbn.) 'Piper, N. Dak.—
P. compressa L., Minn., N. Dak., S. Dak".-^P; 'glaucifolia Scribn. and
Will., N. Dak.—P. interior Rydb., Wyo.—P. nevadensis Vasey, N. Dak.—
P, palustris L., Minn., Mont., N. Dak.—P. pratensis L., Minn., Nebr.,
N, Dak., Wash., Wyo.—P. secunda Presl, N. Dak.—Schedonnardus panicu-
latus (Nutt.) Trel., Nebr., S. Dak.—Secale cereale L., Minn., Nebr.,
N. Dak., S. Dak.—S. cereale X S. montanum Guss., N. Dak,—Setaria
italica (L.) Beauv., Minn., N. Dak.—S. lutescens (Weigel) F. T. Hubb,
N. Dak., S. Dak.—S. viridis (L.) Beauv., Nebr., N. Dak., S. Dak.—
Sitanion hystrix (Nutt.) J. G. Sm., Wyo.—Sorghastrum nutans (L.) Nash,
N. Dak.—Sorghum halepense (L.) Pers., N. Dak.—S. vulgare Pers., N.
Dak.—S. vulgare var. sudanense (Piper) Hitclic, N. Dak.—Sporobolus
cryptandrus (Torr.) A. Gray, N. Dak.—S. neglectus Nash, N. Dak.—
Stipa baicalensis Roshev, N. Dak.—S. columbiana Macoun, N. Dak.—
S. comata Trin. and Rupr., N. Dak., S. Dak., Wyo.—S. spartea Trin.,
Nebr.—S. viridula Trin., N_. Dak., S. Dak., Wyo.—Triticum aestivum
L., Calif., Manit., Minn., Mont., Nebr., N. Dak., Oreg., S. Dak., Wash.,
Wyo.—T. dicoccum Schrank, N. Dak.—T. durum Desf., N. Dak., S. Dak;
—T. spelta L., N. Dak.—T. timopheevi Zhuk., N. Dak.—Zea mays L.,
CaUf., Minn., N. Dak., S. Dak.
The host range of this species, 120 at present, is very great and is
perliaps more or less unlimited on the northern grasses and cereals.
c? o
FIG. 61.—Conidia of Gloeosporium bolleyi from pure cultures on potato-dex-

trose agar: A, from culture 3679 B-5, pink strain, isolated from Hordeum brevi-
subulatum, under snow, Mandan, N. Dak., Jan. 17, 1946; B, from culture 49 A-2,
pink strain, isolated from Agropyron cristatum, Huron, S. Dak.; C, from culture
5 B-1, intermediate strain, from A. cristatum, Mandan, N. Dak.; D, from cul-
ture 3703 B-3, carrot red strain, from Elymus canadensis, Mandan, N. Dak.
(From Phytopathology.)
When I first encountered this fungus on necrotic cereal roots, I

assumed that it was-a form of CoUetotrichum graminicola, as had a
number of earlier workers. However, after referring to the work by
Selby and Manns (1909) it was obvious that the North Dakota ma-
terial was a species-of Gloeosporium (Fig. 61) with only the most
superficial resemblance to CoUetotrichum. The early inoculation work
of ours in all but one case proved negative which resulted in dropping
work with the fungus (1944,6) until more presaging problems had been
studied. Later tests disclosed, to our surprise, that some of the isolates
caused a severe seed rot and root'necrosis on a number of hosts. The
fungus, therefore, must be-recognized as a parasite of Gramineae even
though many strains or isolates seem to be nonparasitic.
G. bolleyi is readily isolated on potato-dextrose agar from water
agar plates of water-washed roots and crowns. The isolates grow slowly
at first but they eventually cover test tube slants with yeasty masses
of conidia. The isolates which form pink spore masses first produce
pale cream-colored colonies which soon become f^right pink or carrot
red masses of conidia and which eventually gernainate to form black,
smooth or wrinkled, carbonaceous colonies. On sterilized sand plus 0.85
per cent bran and 0.5 per cent dextrose the colonies are gray, cottony,
and are composed of true mycelium. This species constituted about
4 per cent of 22,832 pure cultures of fungi isolated from Gramineae
roots up to 1944. In 1944 this species averaged 6.7 per cent of the total
and in 1945 it composed 14 per cent of the fungi isolated. During a
period in midwinter with the temperature below zero F., material
collected yielded 26 per cent of G. bolleyi from dormant tissue. In these
isolates 96 belonged to the strains forming pink conidial masses and
52 to those developing carrot red ones (Sprague, 1946,/).
G. bolleyi is most prevalent on roots of grasses and cereals growing
in crowded conditions in poor soil, especially sandy soils deficient in
nitrogen.
Range: "United States, Canada. ^ -
References: Bolley (1913); Sprague (1944 6,c; 1946,/; 1948,6).
Gloeosporium graminum Rostr.—Ânthracnose

Acervuli numerous, brown, conidia irregularly oblong, 11-14 X 4-
6 /A.
On: Poa pratemis L., Oreg.

Two collections of a fungus which answers the brief description of
G. graminum were obtained some years ago in Benton County, Oregon,
in a humid region along a small creek. Their status remains uncertain
but the size of the spores excludes them and Rostrup's fungus from G.
bolleyi. G. graminum was described on LoliUm multiflorum from Den-
mark. Possibly these forms should be compared with the macroconidial
stage of Phialea temulenta Prill, and Del. whose spores are 11-21 X 3.3-
6.0 ;a. , ' --
Rothers (1928) reports a wilt on Lolium perenne, caused by
Gloeosporium "graminis." The original of this article is not available
and it is not possible to say whether this is believed to be the same as
Rostrup's fungus.
Range: United States, Denmark, U.S.S.R. j* '
References: Rostrup (1892); Rothers (1928); Sprague (1937,e; 1946,e;
1948,6).
/
Gloeosporium meinersii Sprague-êaf Spot
Spots on leaves elongate, 0.5-2.0 X 1-3 mm., pale buff to tawny,
emarginate or with a narrow brown border, center of spots finally paler;
conidia numerous, hyaline, nonseptate subphaseoliform to oblong, gut-
tulate, 9-12.5 X 3.0-3.8 /i (on Phleum), sometimes 10-14 X 3-4 ^ (on
Poa).
On: Phleum pratense L., Wash.—Poa alpina L., Colo.
Jack P. Meiners and G. W. Fischer collected this fungus in western
Washington. We found comparable material on Poa alpina at 11,700 ft.
elevation above Monarch Pass, Colorado. This material had tawny
lesions which occupied much of the surface of basal leaves. It was
named G. meinersiivar. alpina Sprague (1949,a).
Reference: Sprague (1949,ffl).
Melanconium iliau Lyon—See its ascomycetous stage Gnomonia

iliau Lyon
• Melanconium sacchari Massee apud Speg.—Rind Disease

Browning of surface followed by souring and reddening of interior,
numerous small acervuli, 250-500 /x diam.; gregarious beneath the epi-
dermis with a pore, spore horns prominent, black; conidiophores hya-
line, 7-10 X 3 /x; conidia cylindrical with rounded ends, 10-15 X 3.5-4.0
ju,, straight or curved, olivaceous, 1- to 2-guttulate (Fig. 62).
On: Saccharum officinarum L., widely distributed where the crop is grown.
\
In their trials Faris and Allison (1927) found that this fungus was-
nonparasitic. Nowell (1923) indicated that the species may be a wound
parasite on weakened plants. Abbott (1932) reported
that the species is much less serious as a seed rot than
Colletotrichum jalcatum. Cook, M. T. (1932,b) believed
that this fungus is only a weak parasite. Abbott (1932)
mentioned that this fungus was first described as
Strumella sacchari Cooke in 1890. Stru7nella belongs in
the Tubercularieae of the Moniliales. The fungus is some- pm g2 _
times placed in Pleocyta. Seaver, Chardon, and Tore Conidia of
(1932) listed M. sacchari as a synonym of Pleocyta sac- Melanco-
, . . T^ , T^- -- J J a mum sac-
chan from Puerto Kico. chari.
Averna-Sacca (1933) found no evidence that M.
sacchari and Ceratostomella paradoxa were related. Bremer, G. (1926)
obtained evidence that Nigrospora panici Zimm. was a form of M.
sacchari.
M. arundinaceum Ell. and Ev. appears to be saprophytic on Arundi-
naria in Louisiana (1897, p:290). The acervuli were 1.0-1.75 X 0.75-1.0
mm. diam. and the conidia were globose, 15-20 n, or elliptical, 18-
22 X 13-16 ix, on short conidiophores.
Range: United States, Argentina, Australia, Brazil, British Guiana, Cuba,
Dominican Republic, Egypt, Hawaii, India, Java, Kenya, Mauritius,
Natal, Peru, Puerto Rico, S. Africa, Tanganyika, West Indies.
/'
References: Averna-Sacca (1933); Bijl (1921); Earis and Allison (1927);
Fawcett (1922); Johnston, J. R. (1917); Massee (:1895); Nowell (1923);
Saccardo (Sylloge, 1899, v. 14, p. 1019). '^ ' ^
Septogloeum bartholomaei (Pk.) Wr.-J-Leaf Spot

Fusarium bartholomaei Pk. |
Trichofusarium bartholomaei (Pk.) Sacc.
Sporodochium compact, subglobose, oblong, or pulvinate, arranged*
in series, whitish, long covered by the elevated epidermis; conidiophores
slender, erect, straight, crowded, 40-60 X 1-2 /J.; coriidia slender, curved,
5-6 nucleate, continuous, hyaline, terminating at each end in a very
thin bristle-like point, the point or bristle being 10-12 /x, long. On lower
leaf surface.
On: Sorghastrum nutans (L.) Nash, Kans.
This fungus has not been seen by the writer and he is accepting the
diagnosis of Wollenweber. Saccardo placed it in the Trichojusarium
(Sylloge, 1913, v. 22), while Peck had/originally considered it as be-
longing to the subgenus Fusamen oi, Fusarium (1909, p. 157). Peck
emphasized that the fungus was readily distinguished by the pointed
spores and the long covered acervuli. The fungus differs from Spermo-
spora in the presence of true acervuli, in the fact that the spores are
bristle-like at each end while Spermospora avenae has a cilium-like
appendage at the lower end which is produced obliquely, and finally,
because the spores of the Kansas species are much narrower and non-
septate. S. avenae grows during cool winter or early spring weather
while S. bartholomaei was found in September oi/maturing leaves.
Range: United States. |
References: Peck (1909); Saccardo (Sj'Uoge, i913, v. 22).
I'
Septogloeum oxysporum Sacc, Bomm. and Rous^.^~B.lptch and
Char Spot - ^
Lesions at first tawny, yellow margined, circular but soon elliptical
to elongate, becoming covered with dull bfack, charcoal-like streaks,
2- to 5-mm. wide and often several times as long, frequently sharp-
pointed at each end; spores borne in obscure acervuli or in pycnidia
usually in the center of the lesion or about its periphery, 80-160 jx,
diam., ostiolate; conidiophores hyaline, subcuspidate, or nearly globu-
lar in some cases; spores yellow to subhyaline, fusoid/often flattened
slightly on one side, subtruncate at the base;^t5pering to an obliquely
pointed apex, 0- to 3-septate, but mostly 2-septate, 20-38 X 2.5-6.0 /x.
Associated are perithecia in some cases, these are often filled with
prosenchymatous material, globose to slightly flattened, as much as
250 ju. diam., asci when formed ace short-fascicled, no paraphyses and
no mature spores seen.
On: Agropyron repens (L.) Beauv., N. Dak.—A. spicatum (Pursh) Scribn.
and Sm., Wyo.—A. trachycaulum (Lk.) Malte, Mont., N. Dak., S. Dak.—
Agrostis alba L., Colo., Utah, Wash., Wyo.—A. exarata Trin., Wyo.—
A. hallii Vasey, Oreg.—A. scabra Willd., Oreg.—Arrhenatherum elatius
(L.) Beauv., Oreg., Wash.—Bromus carinatus Hook, and Am., Colo.—
B. ciliatus L., Colo.—Calamagrostis canadensis (Michx.) Beauv., Alta.,
Mont., Utah—C. inexpayisa k. Gray, N. Dak.—C. rubescens Buckl., Ida.
—Elymus condensatus Presl, Cahf., Colo., Ida., Mont., Oreg., Utah, Wash.,,
Wyo.—E. glaucus Buckl, Alta., Ida., Utah, Wash., Wyo.—Glyceria elata
(Nash) Hitchc, Wash.—Muhlenbergia asperifolia (Nees and Mey.)
Parodi, N. Dak.
A number of years ago the writer discussed this fungus, or fungi,
in-some detail (1941,a). Typical blotch material on Arrhenatherum
elatius from Oregon has been determined as true S. oxysporum. On the
other hosts such as Agrostis, Agropyron, and Elymus the blotch stage
becomes slowly overrun by black stromatic mycelia which form the
char spot symptoms. Some pycnidia occur in this material and they
contain spores (Fig. 63,S) which are similar to those produced in the
earlier-formed spot blotches. In still other cases immature perithecial
material has been found. Thi^ is probably similar to Dothidella aris-
tidae which we are keeping distinct from S. oxysporum at this time.
Earlier (1941,a) we illustrated pycnospores of a &eptogloeum-Y\k&
fungus in char spot material in North Dakota (Fig. 63,F). Perhaps we
were correct in considering this close to, if not identical with, Septo-
gloeum. But not only must future research consider both Dothidella
spp. and Sepipgloeum, but the genus Davisiella, especially D. elymina
(J. J. Davis) Petrak (1924, pp. 133-134). This group was discussed by
Greene in 1942 (pp. 86-87). Davisiella is based on Cytodiplospora
elymina and has been-discussed in the Sphaeropsidales in this text. The
writer agrees with Greene that probably more than one fungus is in-
volved in the whole Middle Western and Western complex. In this
publication, on the basis of very incomplete information, we consider
that most of the Middle Western material belongs under Davisiella
which is probably a hemiparasite on tar spot and char spot. The ma-
terial on Arrhenatherum is true Septogloeum {Fig. 63,B) as above men-
tioned (Fig. 63,A), the char spot on Distichlis in the Far West is
Dothidella aristidae. The char- spot on Agrostis, Calamagrostis, Agro-
pyron, and Elymus is Septogloeum but it contains pycnidia at times
and frequently developing, but never mature, perithecia. It is possible
that the material on Agrostis and Calamagrostis is different from that
on Agroptjron and Elymus (Fig. 6S,C,D,E). The material, for instance,
which Fischer, Meiners, and the writer collected on C. rubescens at
FIG. 63.—Septogloeum oxysporum Sacc, Bomm. and Rouss.: A, conidia from

Agrostis hallii, Corvallis, Oreg. One of the spores has a oonidiophore still at-
tached and broken away from the parent stroma; B, conidia from Arrhena-
therum elatius, Corvallis, Oreg. (O.S.C. 23); C, conidia ancl pycnospores from
Elymus glaucus, Logan Canyon, Utah (B.P.I. 80,003); D, /{lonidia from Elymus
condensaius, Hooper, Wash.; B, pycnospores fromj Agropyron spicatum, Mt.
Wagoner, Lincoln County, Wyo.; F, pycnospores from Distichlis spicata, Man-
dan, N. Dak. (From Northwest Science.)
Smith Ferry, Idaho, in 1947 (Sprague, Fischer, and Meiriers, 1948),

and again in 1948, showfed typical circular to elliptical'"gray_ black
blotches or spots with white masses of conidia piled in the centner of the
lesions (see also Fig. 64). On the other hand, the sometimes abundant
material on Elymus condensatus is more likely-to be a black char spot
which often covers most of the leaf surface of plants growing in small
clearings in pine forests or in unfavorable habitat^ where the humidity
is somewhat higher than normal for this semidesert plant. These lesions
seldom show conidia but pycnidia are sometimes found which contain
the Septogloeuni-like spores. Immature perithecia are usually present
on black, well-developed material but we have never found any that
had mature ascospores. We heard reports that mature' material was
collected in Logan Canyon, Utah, some years ago but search there in
1947 and again in 1948 did not disclose any that could be identified.
FIG. 64.—Lesion caused by Scplnglocum, oxysporum on Calamagrostis cana-

densis, Grand Mesa, Colo. T h e black stromatic mass is covered in the center
of the lesion with a lighter mass of conidia.
Some abundant material collected north of Burns, Oregon, in the

Malheur National Forest was the nearest to mature of any seen. It
might be added that D. aristidae seldom produces mature perithecia.
The dry summers of the Far West apparently discourage the develop-
ment of the ascigerous stage.
Recently we saw conidial material on Elymus glaucus from Rainier
National Park, Washington, which closely resembles that of Septo-
gloeum on Arrhenatherum. This tends to link the groups on Agrostidae
with those on Hordeae.
Range: United States, Belgium, Canada, Germany, and in northern Europe
in general.
References: Bommer and Rousseau (1890); Greene (1942); Sprague
(1941,a; 1946,e). .,-- -
Septogloeum spartinae (Ell. and Ev.) Wr.—Spot

Fusarium spartinae Ell. and Ev.
Sporodochia pale-yellow, floccose; conidia oblong-elliptical to
oblong-fusoid, 1- to 3-septate, 12-15 X 3-4 /x.
On: Spartina leiantha Benth., Cahf.
The type is from Pacific Grove, California, which is also the type
locality of Phyllachora serialis Ell. and Ev., reputedly on the same
l \ •
host, but more probably on Elymus triticoides Buckl. instead. I am
rather dubious about the autonomy of this species but have not seen
it. Seymour (1929) lists the fungus on Spartina alternifolia Loisel.
References: Ellis and Everhart (1902, p. 14); Saccardo (Sylloge, 1906, v.
18, pp. 674-675); WoUenweber and Reinking (1935, i). 336).
MONILIALES
This order contains those forms of the Fungi Imperfecti which pro-
duce their conidia on free conidiophores which are not contained in any
fruiting body. Fundamentally the spores are produced in the open on
either loose scattered mycelium, or distinct conidiophores which are
sometimes intricately branched, or they are aggregated on sporodochial
pads or such structures as coremia which are bundles of vertical grow-
ing fertile hyphae. Some important scalds, blasts; molds, and root rots
are caused by members of this order.
The key to the included genera of Monihales is as follows:
A. Spores hyaline
B. Spores borne sessile or on short conidiophores
C. Spores in slimy matrix, elongate to filiform Gloeocercospora
CC. Spores not as above
D. Spores cylindrical, boat-shaped Mastigosporimn
DD. Spores apicallj' elongated (subulate) sometimes with basal cilia
Spermospora
. DDD. Spores 1-septate, wedge-shaped with oblique apical beak, or
cylindrical without beak Rhynchosporium
DDDD. Spores globose, small Sporotrichum
BB. Spores on evident conidiophores
C. Conidiophores branched
D. Conidia in heads
E. Conidia not in chains Cephalosporium
EE. Coiiidia in chains on penicilhately branched conidiophores
F. Conidia enclosed in mucus Gliodadium
FF. Conidia not enclosed in mucus Penicillium
DD. Conidia not iii heads but often in brightly colored sessile masses
(sporodochia) Fnsarium.
CC. Conidiophores simple
D. Spores nonseptate, ovoid Ovularia
DD. Spores somewhat elongated or septate
E. Some of spores branched Ramulispora
EE. Spores not branched
F. Spores fusoid, attenuated below .' Ellisiella
FF. Spores fusoid but not,attenuated below
. Ramularia (graminicola)
FFF. Spores filiform Cercosporella
AA. Spores colored
B. Spores borne on very short and obscure conidiophores
C. On leaves, spores 15 /* or less in length, ellipsoidal Coniosporium
CC. On young Claviceps sclerotia, spores are 3-chambered glomerules
Cerebella
299
CCC. Spore masses in flower parts, spores small,! 4.6 ^ diam., causes a
"false smut" ^ , ..., ^., Ustilaginoidea
BB. Spores borne in evident conidiophores ^ '
C. Conidiophores short but evident, simple ^
D. Causing extensive black streaks and death ofi leaves (on Phragmites)
Napicladium
DD. Conidiophores arising from less extensive flat black pads
E. Spores echinulate, globose, multiseptate Epicoccum
EE. Spores smooth
F. Spores arising from an elongate black pad, spores globose to
pyrif orm . . . . : . . Hadrotrichum
F F . Spores arising from a small flattened pad, spores flattened-
globose, opaque Nigrospora
GC. Conidiophores elongate or sometimes branched
D. Spores nonseptate
E. Arising from a sporodochium, spores lenticular with a narrow
hyaline rim around the periphery .Papularia
EE. Spores otherwise •. •
F. Spores smooth, on irregularly' branched prominent, brown
conidiophores / Botrytis
F F . Spores globose, muriculate, when mature on stout black
conidiophores ^ Periconia
FFF. Spores echinulate, variable, on slender brown conidiophores
Cladosporium
(in part)
DD. Spores septate
E. Spores most frequently 1-septate, broadened j
F. Spores echinulate, variable on slender brown conidiophores
Cladosporium
(in part, see also Fusicladium)
FF. Spores ampulliform on fascicled conidiophores. .Scolecotrichum
EE. Spores most frequently more than' 1-septate
F. Spores filiform , Cercospora
FF. Spores otherwise .
G. Spores muriform 1
H. Conidia not in chains, conidiophores decumbent, spores
typically equidimensioned Sfemphyllium
HH. Conidia in chains —Alternaria
GG. Spores otherwise
H. Spores more or less cylindrical
I. Spores echinulate Heterosporium
II. Spores smooth [. Helminth'osporium
HH. Spores otherwise
I. Spores helicoid .Helicoceras
II. Spores otherwise ,
J. Spores curved or bent with one-of the central several
cells distinctly larger than -the^ther cells . .Curvularia
J J . Spores pyriform Piricularia
(see also Dactylaria)
FUNGI IMPERFECTI—MONILIALES 301
Alternaria tenuis auct. sensu Wiltshire—Black Mold

Conidiophores olive brown, septate, simple or branched, length
variable, 3-6 /J. diam. Conidia light olive brown to dark brown, smooth
or warty, usually with three to five cross walls and with longitudinal
ones in the second and third cells, obclavate with rather short beaks
and borne in long chains (16)-20-50-(70) X (7)-10-16-(20) fi, beaks
somewhat longer on plant parts in nature. Produces abundant cottony
or tufted ashy gray to dark gray, greenish to dark olivaceous mycelium
on potato-dextrose agar, somewhat tufted. Sporulation in pure culture
variable, sometimes slow, usually abundant.
On: Saprophytic on many plants including the Gramineae on dead parts,
especially in late season following rains, on shocked grain or hay or on
stubble, causing a black mold; on seeds of such moldy grain and from the
roots of dead matured plants.
Most of the saprophytic black molds referred to Alternaria spp. on
grass and cereals are now placed in Alternaria tenuis. This species is
generally recognized but it has no type specimen and no legal status
even though it is relatively certain of being accepted permanently as a
good species. The work by Elliott (1917), Mason (1928), Wiltshire
(1933) and Groves and Skolko (1944,b) all clearly indicate thai; this
species should be conserved. A. tenuis Nees was pre-Fresian and is the
^.same as Toruta altemata Fries. Corda illustrated A. tenuis (1840) but
his "A. tenuis" is not like A. tenuis Nees. Mason (1928) defined A.
tenuis sensu Corda as one "whose spores are obclavate, borne in long
chains, and the majority of whose spores have 3 to 5 cross septa and,
especially in culture, fall within the limits 20-50 X 10-14 ^." This is
the fungus which most of us know as Alternaria tenuis.
The seed mold of wheat is called "black germ" in Siberia, "punta-
tura" in Italy, "moucheture" in Morocco, and such names as "kernel
smudge," "black point" (in part Helminthosporium sativum), seed
blight, or "durum blight" in Canada and the United States. A mela-
nistic (black skin pigment) browning of wheat lemmas is termed
"Alternaria blotch" by Johnson and Hagborg (1942).
Almost all careful research on Alternaria tenuis shows that it is a
saprophytic fungus on Gramineae. It is, however, economically im-
portant because of the unattractive gray flecking it imparts to maca-
roni flour made from North Dakota durum wheat (Brentzel, 1941).
There is a strong tendency for buyers to dock the price of any wheat
which shows black point.
E. C. Johnson worked with A. tenuis in earlier years and he came
to the conclusion that the fungus was not parasitic (1914). The writer,
working with A. tenuis which he isolated from necrotic roots of Grami-
neae was of the same opinion (1944,6). Grintescu (1923) also obtained
negative results with A. tenuis.
Other species of Alternaria have been reported associated, with
cereals. Bockmann (1933) included A. peglionii Ci:(rzi and A. circinans
(Berk, and Curt.) Bolle as species present on cereals. All of the isolates
of Alternaria from North Dakota Gramineae were assigned to A. teriuis
by David R. Sumstine (Sprague, 1946,e). They varied considerably in
morphology. |
Because A. tenuis is often associated with parasitic species, such as
Helminthosporium sativum Pam. King and Bakke, moldy wheat seed
should be treated with New Improved Ceresan or Ceresari-M at recom-'
mended rates.
A. tenuis is sometimes associated with Stemphyllium botryosum but
the sarcinaeform spores of the latter are readily distinguished. Macro-
sporium as a genus seems to have faded from the scene. At one time
Alternaria and Macrosporium were segregated on the basis of whether
the conidia were borne in chains or singly. Elliott (1917) and others
pointed out that the shape of the spore, whether clavate or sarcinae-
form, was more diagnostic. Alternaria, being pre-Fresian, required ac-
cepting the lesser used Macrosporium to include the forms with clavate-
shaped spores. Recent workers have preferred to accept Alternaria and
reject Macrosporium (Wiltshire, 1933; Groves and Skolko, 1944,&).
Stemphyllium with sarcinaeform spores borne singly on somewhat lax
conidiophores partly replaces Macrosporium.
Range: World-wide.
References: Bockmann (1933); Bolley (1913); Brentzel (1941); Christen-
sen and Stakman (1935); Curzi (1926); Elliott (1917); Grintescu (1923)';
Groves and Skolko (1944,a,6); Hagborg (1936); Hanson and Christensen
(1943); Henry (1924); Johnson, E. C. (1914); Johnson, T., and Hag-
borg (1942); Machacek and Greaney (1938); Mason (1928); Miege
(1930); Peyronel (1926,a); Rosella (1930,a,5); S^llans (1931); Sprague
(1944,6); Wiltshire (1933, 1938); Ziling (19*32).
Botrytis cinerea Pers. ex auct^-^Gray Mold

Hyphae: slender, coiistricted at septa, gregarious, simple-or^parsely
branched, erect, cinerous, conidia globose, obovate, ellipsoid^ variable,
nearly colorless or very pale brown, collected in somewhat globose
heads on short straight-side branches, 7-14 X 4-8 [i.
On: Avena sativa L., N. B., N. S., P. E. Is., Wash.—Bromus mollis L.,
Wash.—Poa secunda Presl, Wash.—Triticum aestivum L., N., 13., N. S.,
P. E. Is.
Gray mold sometimes develops on cereals growing during cool moist
periods- in the maritime provinces of Canada, especially on the matur-
ing or ripened heads and lower leaves. While S.'xiwerea is an important
mold on ornamental plants in cool coastal climates its role on the grass
family is a,pparently not important.
One isolate mad^ by Neil Allan MacLean from Poa'secunda at
Pullman, Washington, appears to be B. cinerea, which is a variable
species. In his studies MacLean found that this isolate could attack
certain bulbous ornamental plants (1948).
The species 5 . cinerea dates to Persoon but is unofficially conserved
by many workers.
Peck described B. uredinicola (1909) on Panicum virgatum from
Stockton, Kansas, but the fungus occurred on rust pustules and ap-
parently was a hyperparasite or secondary organism. The description
reads more like a species of Sporotrichum than Botrytis. The snowy
white tufts had short conidiophores bearing hyaline globose spores 8-10
/J, diam.
L. Stakman (1923) reported B. sp. from the roots of rye seedlings in
Minnesota. The mycelium was said to resemble that of B. compada
(Pat.) Sacc. (see Sylloge v. 10, p. 537). The fungus produced on the
roots a slight brown discoloration with vertical black lines, up to 1 mm.
in length, scattered over the upper half. A cross section through one of
these lines showed a wedge of discolored tissue, wider at the epidermis
and gradually narrowing through the cortex. The mycelium was mostly
intracellular, hyaline at first and then dark green. The walls between
severely attacked host cells were frequently broken down and the inter-
stices filled with numerous vertical, sporulating conidiophores. /
B. elegantula Cooke (Grev. 12, p. 27) is reported on "grass" from
South Carolina.
V
Range: General on many hosts in cool temperate areas, occasional on
Gramineae, especially necrotic plant parts. Ling (1948) listed it on
wheat from Szechwan Province, China.
References: Conners (1929-42); Conners and Savile (1943-46); MacLean,
Neil Allen (Thesis, in manuscript, 1948); Stakman, L. (1923).
Centrospora bromi (Sprague) Newhall—See Ramulispora bromi

(Sprague) comb. nov.
Cephalosporium acremonium Corda sensu Fresenius—^Black

Bundle Disease
Hyphae creeping, conidiophores short, erect, not apically swollen,
continuous, conidia held together in slightly gelatinous heads averaging
6-10 /A diam., spores spherical or ovate, hyaline or slightly colored,
4X1/^.
On: Zea mays L., Ark., Calif.,- Conn., Del., 111., Ind., Iowa, Kans., Ky., Md.,
Mich., Minn., Miss., Mo., Nebr., N. J., N. Y., N. Car., N. Dak., Ohio,
Penn., S. Car., S. Dak'., Tex., Wise.
The original description by Corda (1839) did not include spore,
measurements and the identity of the fungus he dealt with is not clear.
However, according to Reddy and Holbert (1924), his illustration re-
sembled C. acremonium f. major Penzig. Corda reported the fungus on
various substrata, including other fungi, while Fresenius was the first
to report it on corn (1863).
In somewhat earlier work than that of the past dozen years, C.
acremonium was reported as associated at times With a stunting of corn
in the Middle Western United States. Symptoms jinclude black fibro-
vascular bundles accompanied by development of nubbins in the plant,
multiple ears at the node, barrenness and purple discoloration of the
plant. The fungus is seed- and seedling-borne. The symptoms' were well
presented by Reddy and Holbert (1924).
Today the disease is not considered of very much importance by
many workers; in fact, some believe that the fungus is a saprophyte
growing on resinous products of decomposition in the vascular system
of corn, especially in certain inbred corn strains (Harris, 1936). Harris
has personally told the writer that all his evidence indicated that the
fungus was virtually saprophytic.
C. sacchari has been reported on corn- (Manns and Adams, 1921)
but this species has larger spores than C. acremonium and the spores
often become 1- to 3-septate (Butler and Hafiz Kahn, 1913). Reddy
and Holbert (1924) found that the isolates obtained by Manns and
Adams (1921,a,b) were C. acremonium. Bijl (1921) listed C. sacchari
from South Africa on sugar cane. Khanna and Chacravarti (1949) re-
ported that wilt ( C sacchari) from Bihar reduced the sugar content
of cane. This might be an angle to consider in the com plant grown for
special purposes. ;
At this point it might be important to mention in some detail a
locally serious disease of wheat in Japan caused by C. gramineUm
Nisikado and Ikata. Because of increased communication with Japan
there is always the chance that this fungusi will /appear in the United
States. C. gramineum (Nisikado, Matsunioto, and Yamauti, 1934)
causes a clear-cut leaf stripe disease of wheat. In the early stages dur-
ing late February and early March the disease resembles the yellow
type of wheat mosaic. A month later the lesions somewhat resemble
barley stripe. One or two, rarely as many as four, contiliuOus, yellow
brown stripes form on the leaf blades, sheaths, and culms.-By the end
of May the symptoms are acute. This phase is similar to take-all,
stunting, and white heads. The vascular bundles in the diseased areas
turn yellowish brown and the spiral and pitte^ vessels, in particular,
contain hyphae and conidia. Other cereals and grasses are attacked.
The conidiophores are simple, 5-20 X 1-5-4.0 ;a, bearing at their apices
capitate agglomerations of hyaline, long-elliptical to ovoid, continuous,
usually biguttulate conidia, 5-10 X 1-5-3.0 fi. The fuiigus is readily
cultured and infections are easily obtained..Ikâ, Kasai, Yosida, and
Yokota (1936) reported that this species was so adaptable to changes
in acidity that the spores passed through the alimentary tracts of cattle
without loss of viability. '
FUNGI IMPERFECT!—MONILIALES 305
Ikata and Kawai (1937) reported that the minimum, optimum, and
maximum temperatures for growth and conidial germination of C.
gramineum were approximately 5° C , 20° C , and 30° C , respectively.
Nisikado and Higuti (1938) determined that C. gramineum and C.
acremonium were distinct species.
Range: Of C. acremonium: United States, Australia, Czechoslovakia, Ger-
many.
References: Branstetter (1922); Corda (1839); Fresenius (1863); Harris
(1936); Koehler (1942); Koehler and Holbert (1938); Manns and
Adams (1921,a,6); Reddy and Holbert (1924).
Cercospora spp.
This genus is represented by a number of species which cause leaf
spots on various grasses. Atkinson (1892, 1897) described several of
these on southern grasses, especially from Alabama. Solheim (1929)
published a monograph on the genus while Chupp has issued (1937) a
list of described species and has assembled extensive unpublished data.
The published data on the species on Gramineae, except for the original
descriptions, is limited. We spent considerable time attempting to as-
semble a usable key for this work. The fungi can be divided to some
extent into two groups, depending on whether the lesions are linear
or broadened. However, juvenile spots were not classifiable. Some
• species have emarginate spots; others have red, brown, or yellow tissue
around the lesion. The shape of the spore is distinctive in some species
but many are, to me, indistinguishable. The geniscar on the conidio-
phores aids in distinguishing other species. The arrangement or group-
ing of the conidiophores appears to be recognized as diagnostic. Some
species have the conidiophores clustered in short, thick fascicles. Others
have long lax ones, while some species have few or even single stalks.
The mycelium at the base of the conidiophore fascicles is used to dis-
tinguish some species. The stroma or basal mycelia may be abundant
or scanty.
Solheim and Stevens (1931) listed 38 sections of Cercospora. Sol-
heim (1929) had published a monograph with over 100 species included.
A key to the numerous sections might be used in developing a key to
the species on grasses.
Johnson and Valleau (1949) indicate by cross inoculation studies
that many described species of Cercospora are referable to C. apii Fres.
The latter has spores about 50-80X 4 jn. While the problem was too
great for them to do much more than outline it, there are strong indi-
cations that many species that are well known in literature on many
hosts will not be left with even the identity of a racial status. In the
following pages we have mentioned some species that appear to fall
within the limits of C. apii. More and more we need to compare our
grass fungi with those on other host families.
306 DISEASES OF CEREALS AND GRASSES'
l\
Cercospora agrostidis Atk.-—Leaf Spot
Spots amphigenous, broadly elliptical, very 'light brown in center
with broad border of dull red brown, 3-5 mm. Ipng. Hyphae amphige-
nous, loosely fasciculate tufts irregularly scattered and few in a spot,
bright reddish brown, septate, nearly straight to kubflexuous and spar-
ingly toothed near apex, 40-60 X 2.4-3.1 ,1. Conidia hyaline, 1- to
7-septate, terete, straight or little curved, 10-60 X 2.5-3.5 /*.
On: Agrostis perennans (Walt.) Tuckerm., Alabama—A. scabra Willd., Ida.
—A. sp., Alabama—Sphenopholis obtusata (Michx.) Scribn., N. Dak.
Chupp identified material on the last mentioned host which we sent
from Mandan, North Dakota.
References: Atkinson (1892). See also Johnson and Valleau (1949).
Cercospora asprellae Ell. and Gall. nom. nud.—Leaf Spot

Apparently undescribed, specimen in mycological collections.
Bureau of Plant Industry, Beltsville^ Maryland.
On: Hystrix patula Moench, 111.
Reference: Weiss (1945, v. 29, n. 9, p. 227).
I
Cercospora boutelouae Chupp and Greene—Leaf Spot
Spots narrowly elliptic to elongate, 0.5-4 mm/in length, pale brown
to almost black, immarginate or with a yellowish halo; fruiting mostly
hypophyllous, seriate; stromata lacking qr only a few brown cells;
fascicles 2-14 spreading stalks; conidiophores pale to medium brown,
slightly paler and narrower near the liip, sparingly septate, not
branched, 0-1 abruptly geniculate, straight to curved-jJipsubtruncate,
20-100 X 3.0-5.5 /x; conidia hyaline to pale brown, obclavate', mildly
curved, indistinctly multiseptate, base long obconically truncate, tip
subobtuse, 20-80 X 3.5-5.0 /t.
On: Bouteloua curtipendula (Michx.) Torr., Okla., Wise, (type)—B. gracilis
(H. B. K.) Lag., Colo. /
Known from the type area at Madison, Wisconsin (Greene, 1944,6),
from Oklahoma (Preston, 1947) and from Buckhorn Creek in the
Roosevelt National Forest, Colorado. The latter collection contains
spores that are pale brown. ^/^ ^
This fungus has spores falling within the rangeôf C. apii Fres.
References: Greene (1944,6, p. 579); Preston (1947); Sprague' (1950,a).
Cercospora bromi Sprague—See Ramulispora bromi (Sprague)

comb. nov.
Cercospora caespitosa Ell. and Ev.—Leaf Spot

Spots hypophyllous, brown, 0.2-0.3 mm. diam., conidiophores densely
caespitose, brown, 3- to 5-septate, undulate, geniculate toward the top,
70-100 X 4 ;a; conidia cylindrical-fusoid, curved, hyaline; 20-35 X 3.0-
3.5 /t.
On: Chloris petraea Swartz, Miss. /
Reference: Ellis and Everhart (1891).
Cercospora echinochloae J. J. Davis—Leaf Spot

Spots elongate-linear, reddish brown, becoming arid in the center;
conidiophores hypophyllous in small tufts, brownish tinted, continuous,
straight or bent, entire or denticulate, or oblique at the apex, 10-
26 X C5-5.0 jx, conidia hyaline, straight or curved, cylindrical to/ ob-
clavate-cylindrical, distinctly 1- to 7- (mostly 2- to 3-) septate, 23-
53 X 3-4 ix.
On: Echinochloa crusgalli (L.) Beauv., Wise.
This species was collected in late summer in 1913 at Devil's Lake,
Wisconsin. Apparently there are no published reports of the occurrence
of the fungus elsewhere. Material which I collected on E. crusgalli at
Mandan, North Dakota, was referred to C. sorghi by Dr. Chupp.
,Re£erence: Davis, J. J. (1915,6, p. 106).
Cercospora festucae Hardison—Leaf Spot

(But see Johnson and Valleau, 1949)
Leaf spots oval to elongate, 0.5-4.0 mm., gray to purplish border;
stromata none or a few brown cells; fascicles 2-8 divergent stalks;
conidiophores near the base are pale to medium olivaceous brown, paler
and sometimes narrower near the tip, sparingly septate, rarely genicu-
late, almost straight, not branched, rounded to subtruncate tip, 50-
800 X 3.5-5.0 iJ.; conidia hyaline, acicular, curved or undulate, indis-
tinctly multiseptate, base truncate, tip acute, 40-300 X 2-4 fi.
On: Festuca elatior var. arundinacea L., Ky., Oreg.
Preston (1945) listed Cercospora sp. on F. elatior from Oklahoma.
This may be the same species which Hardison described (1945,6).
Hardison reported that this fungus caused only a mild leaf spot on
tall fescue (Fig. 65). He mentioned (in personal correspondence) that
he had noted a few spots on fescue in Oregon.
,•1
'•J !l
t I
ii
m
i i
I
I'
1
IV HI 'i^
l-p^
jSl: lUf raK
Hit
FIG. 65.—Lesions caused by Cercospora festucae (C. apii) on Festuca elatior,

from type, Xl- (Photo by John Hardison.)
Johnson and Valleau (1949) place this under C. apii. They did not
inoculate celery with C. festucae although morphologically the latter
is close to C. apii, C. beticola, and others which Johnson and Valleau
considered identical. C festucae caused eye spots on beans, cabbage,
and beets but not on petunia or canteloupe.
Reference: Hardison (1945,6).
Cercospora fusimaculans Atk.—Leaf Spot

Spots amphigenous, light brown bordered by dark brown, broadly
fusoid or elliptical, 3-4 mm. long, frequently confluent; conidiophores
epiphyllous, fasciculate, olive-reddish-brown, straight, sub-geniculate
or nodulose, sparingly denticulate towards the apex, septate, 50-
100 X 4.0-4.5 IJL; conidia hyaline, 3- to 4-septate, 25-40 X 2.0 /t.
On: Echinochloa crusgalli (L.) Beauv., Towa, Wise.—Leptoloma cognatum
(Schult.) Chase, Wise.—Panicum dichotomiflorum Michx., Iowa—P.
dichotomum L., Alabama—P. latifolium L., Wise.—P. leibergii (Vasey)
Scribn., Wise.—P. maximum Jacq., Canal Zone—P. pacificum Hitche.
. and Chase, Oreg.—P. perlongum Nash, Wise.—P. praecocius Hitche. and
Chase, Wise.—P. scribnerianum Nash, Wise.—P. virgatum L., N. Dak.,
Wise.—P. xalapense H. B. K., Okla.
The~light brown blotches are obscure and probably account for the
limited hosts and geographic range reported for this species. It is 'un-
doubtedly on many species of Panicum, within most of the states
abounded approximately by the Scattered reports which are listed above.
C panici J. J. Davis (1919,6) may be & synonym.
Range: United States and Canal Zone.
References: Atkinson (1892); Greene (1944,o); Underwood and Earle
(1897).
Cercospora longipes Butler—Brown Spot

Spots at first narrow, oval, about % inch long, reddish; brown
center and yellow border form later; mature spots are as large as %
inch diam.; oval, deep brown rings with straw-colored centers; conidio-
phores caespitose, flexuose, brown, geniculate to denticulate, 100-
200 X 4 ju, conidia obclavate, sometimes attenuate, straight to curved,
hyaline, 4-6 septate, 40-80 X 5 /A.
On: Saccharum ojficinarum L., Alabama, Fla., La.
Matsumoto and Yamamoto (1934) state that this species caused an
extensive streaking of sugar cane leaves in Taiwan (Formosa). The
spores in their material were hyaline to subhyaline, 2- to 15-septate,
29-153 X 2.5-3.7 fi, commonly 62-91 /«. long. They say it is close to
Cercosporina imperatae Sydow, but this species has spores 50-65 X
5-6 /i.
Range: United States, Brazil, Cuba, India, Lesser Antilles, Puerto Rico,
S. Africa, Tanganyika, Uganda, Venezuela.
/i
References: Bourne (1933); Butler (1906); Matsiimoto and Yamamoto
(1934); NoweU (1923). '
"""" - - I
Cercospora muhlenbergiae Atk.—Leaf Snot and Stripe
'1
Amphigenous, fascicles few, scattered over the leaf, dense at the
base but conidiophores becoming flexuous above and nodulose, brown,
septate, 70-140 X 3.5-4.0 n; conidia hyaline, straight, narrowly lanceo-
late, 1- to 5-septate, 25-35 X 4-5 /x.
On: Muhlenbergia tnexicaud (L.) Trin., Alabama—M. Tcicetnosct (Michx.)
B. S. P., N. Dak., Wise.—M. schreberi Gmel., Alabama—M. silvatica
Terr., Wise.—M. sp., Iowa.
This species causes a leaf spot and olivaceous stripe mold. We have
seen material in the Missouri River Valley near Wilton, North Dakota,
which showed marked injury. Nagel (1934) used this fungus, among
others, in his studies on sporulation under artificial culture conditions.
References: Atkinson (1897, p. 46); Nagel (1934).
Cercospora oryzae Miyake—^Narrow Brown Spot, Glume Spot

Leaf lesions linear, 3 to 5 mm. long, usually up to 1 or 1.5 mm.
wide, central part dark brown, fading toward margin, also spots on
sheaths, peduncles, and glumes, conidiophores solitary or 2-3, in groups
arising from stomata, 3- to multi-septate, 88-140 X 4-5 /*; conidia
cylindrical to subclavate, 3- to 10-septate, 20-60 X 5 /x.
On: Oryza sativa L., Alabama, Ark., La., Tex. /
C. oryzae produces narrower and lighter brown lesions than those
caused by Helminthosporium oryzae or Piricularia grisea (Tullis,
1937). i
Ryker reported 5 races and variants of G. oryzae on rice in Louisi-
ana, differentiating them by their relative ability to attack Blue Rose,
Blue Rose 41, Fortuna, Caloro, Calusa, Zenith, Delitus, andôuthern
Red rice (1943). Another variety, Rexoro, previously resistant, proved
susceptible to a new race in 1944 and 1945 (Chilton, 1945; Chilton and
Tullis, 1946) in both Texas and Louisiana. This race 6 was not serious
on most of the commercial varieties but the fa^t that it was parasitic
on the important variety Rexoro added to the difficulty of breeding for
resistance. Chilton and Tullis (1946) noted that out of 703 foreign
varieties and selections 555 were resistant to race 6, 109 moderately
resistant, and only 16 susceptible, while 23 appeared tb be segregat-
ing. Recently Ryker reported race 7 (1947)._3Fhis race differed from
race 4, which could only attack Fortuna, in that race 7 also attacked
Blue Rose; He reported that race 4 had not developed extensively but
race 7 appeared to be increasing. He also reported that Rexoro had be-
come heavily diseased since 1944 with a widespread epiphytotic of race
6. Nira, the only remaining resistant variety, showed some infection
in 1945 and some fields were severely infested in 1946. Tests with this
material disclosed still another race, 8. Of the six differential hosts used
at this time race 8 attacked Nira and Fortuna readily but Blue Rose,
Blue Rose 41, Caloro, and Rexoro were resistant. Ryker and Cowart
(1948) discussed the breeding program for combating this disease and
at that time they reported additional races 9 and 10. Some years ago
Ryker and Jodon (1940) had reported that resistance to C. oryzae in
crosses was due to a single dominant factor.
Ganguly (1946,a) reported that Bengal material of C. oryzae pro-
duced the spores in chains.
Range: United States, Brazil, Burma, China, Colombia, Cuba, Dominican
Republic, East Indies, India, Japan, Philippines, Puerto Rico, Venezuela.
References: Chilton (1945); Chilton and TuUis (1946); Ganguly (1946,a);
Metcalf (1906); Miyake (1910); Ryker (1943, 1947); Ryker and Chil-
ton (1942); Ryker and Cowart (1948); Ryker and Jodon (1940);
Sawada (1921, 1922). Tullis and Altstatt (1944).
^ Cercospora panici J. J. Davis—Leaf Spot

Cercos-porina panici (Davis) Sacc. /
Spots fusoid, ferruginous, central portion sordid white, 2-4 X 1-2
mm. Conidiophores amphigenous, caepitose, fuliginous, straight or more
or less flexviose and nodulose, 30-40 X-3 JA; conidia hyaline, cylindrical,
straight or curved, catenulate (?), 30-40 X 2-3 /*.
On: Panicum boscii Poir, Wise.—P. latijolium L., Wise.
Davis found this species in three localities in Wisconsin but did not
report C. fusimdculans Atk. from the state. Greene (1944,a) stated that
Chupp considers this Wisconsin species a synonym of C. fusimaculans.
Apparently the difference in length of the conidiophores in the two
species has not been found to be a consistent character.
Range: United States, Dominican Republic.
References: Davis, J. J-.-(1919,6; 1942); Greene (1944,a).
Cercospora paspali Ray—Leaf Spot

Leaf spots oval to linear, the largest attaining 12 X 5 mm., dark
brown to almost black, frequently with narrow tan centers; fruiting
amphigenous; stromata a few cells to 30 /* diam.; dark brown, fascicles
2-15 divergent stalks; conidiophores moderately brown but paler and
distinctly attenuated toward the tips, 2-5 septate, sparingly geniculate,
straight to tortuous, not branched, medium spore scar at the narrow,
subtruncate tip, 10-80 X 3.0-4.5 /x; conidia hyaline, narrowly acicular
to almost cylindric, straight to slightly curved or undulate, indistinctly
/'
multiseptate, truncate to obconically truncate at'the bases, subacute to
subobtuse at the apices, 30-140 X 1.2-4.0./X;
On: Paspalum stramineum Nash, Okla.
The type was collected at Perkins, Oklahoma, August 26, 1942, and
is filed at Cornell University as No. 33,134. '
Range: United States, possibly Uganda.
Reference: Ray (1944).
Cercospora scolecotrichoides Atk.—Leaf Spot

Fascicles of conidiophores densely aggregated, black, punctiform,
conidiophores brown, fiexuose, nodulose, sometimes ramose, 100-200 X
4-6 jx, septate; conidia formed terminally or laterally, obclavate, fuligi-
nous, abruptly attenuated, 3-septate, 30-50 X 8-10 H-
On: Arundinaria tecta (Walt.) Muhl., Alabama:
This has not been seen. From the description alone it could be
related to Scolecotrichum graminis Fckl., as is mentioned by Atkinson
(1897).
Range: United States. ^
Reference: Atkinson (1897, p. 46).
Cercospora seminalis Ell. and Ev.—Leaf and Glume Spot,

False Smut '
In spikelets, forming a compact olivaceous mass enclosed by the
spines of the involucre, mycelium penetrating the seed which it replaces
with a mass of conidia; conidiophores between tips of enclosing palea
dark olive, long-branched, obliquely truncate above, septate below,
conidia acrogenous, subhyaline, obclavate, granulose, 3- to 5-septate,
80-110 X 6-7 /t. '
On: Buchloe dactyloides (Nutt.) Englm., Colo., Kans., NebPt,J5^Dak., Okla.,
S. Dak., Tex., Wisp. Weiss (1945) listed Cynodon dactylon (l7:)~Pers. as
a host from Texas. —
False smut is an important disease because it materially reduces
seed yield and the resulting shortage of seed has retarded regrassing
programs where buffalo grass is the most importa-nt grass used. Buffalo
grass seed is usually presoaked before planting. Whether viable conidia
of the fungus have any detrimental effect on seedlings is not known,
but no isolations have been made that indicate that such is the case.
Some studies were made at Mandan, North Dakota, by the writer.
The fungus is sometimes moderately serious in. small, experimental plots
there although this region is beyond the main northern limits where
buffalo grass, is important. Dusting trials with fermate, basic copper
sulfate, and other materials in the spring of the year were inconclusive
because of the relatively scanty infection available in the year in which
the trials were conducted.
Reference: Ellis and Everhart (1888,a).
Cercospora seriata Atk.—Leaf Spot

Spots amphigenous, cinereous with definite brown border margined
with indefinite yellow, irregularly oblong, sometimes confluent. Hyphae
epiphyllous, fasciculate, faint reddish brown, in age darker, flexuous
and toothed, 20-50 X 4 ju, tufts in parallel rows. Conidia hyaline, nearly
cylindrical, straight or curved, faintly 2- to 6-septate, 30-70 X 3-3.5 ;u.
On: Sporobolus asper (Michx.) Kunth., Alabama—S. clandestinus (Spreng.)
Hitchc, Okla.
References: Atkinson (1892); Underwood and Earle (1897).
Cercospora setariae Atk.—Leaf Spot

Spots amphigenous, dark with indefinite pale border, elliptical;
hyphae epiphyllous, dull reddish brown, fasciculate, sometimes fasci-
cles very dense, others with the stalks divergent, sometimes conidio-
'^"phores branched near the base, septate, with a few small guttulae, scars
small, giving denticulate appearance near apex, 50-100 X 4.5-5.0 ju,;
conidia hyaline, 1- to several-septate, cylindrical to obclavate, straight
to curved, 20-150 X 4-5 fi.
On: Setaria lutescens (Weigel) F. T. Hubb, Alabama, Iowa, Ky., N. Y.,
N. Dak., Wise.—Sporobolus cryptandrus (Torr.) A. Gray, Wise.
Reference: Atkinson (1892); Greene (1950).
«^
Cercospora setaricola Tehon and Daniels—Leaf Spot
Foliicolous, spots at first small, broadly oval, dark brown, lying
between the larger veins, 0.5-1 mm. long, becoming elongate-elliptic
with a cinereous, nonfriable center and an unraised, sharply delimited,
brown border; sometimes confluent. Fasciculae strictly hypophyllous,
erumpent in rows through the interveinular stomata, not numerous or
crowded, composed of 50 or more conidiophores which arise from a
dark, compact, oval, flat tubercle measuring 15-17 X 30-36 jx. Conidio-
phores straight, erect, continuous, or 1-septate, 17-40 X 3-4 /j,, trans-
lucent, light olivaceous, tapering to a definitely conic apex, with small
but marked internal geniscars. Spores straight or sometimes curved,
acicular, hyaline, 4- to 12-septate, 30-90 X 3-4 [i; geniscar small but
marked, internal.
/',
On: Setaria geniculata (Lam.) Beauv., Okla.—S. liit^scens (Weigel) F. T.
Hubb, 111., Mich., N. Dak., Okla., Wise.
Tehon and Daniels (1927) furnished the key here presented to dis-
tinguish this fungus from others on Setaria.
KEY TO SPECIES OP CERCOSPORA ON SETARIA
Conidiophores long, 50-120 M-

Spores 20-150 X 4-5 /i^
C. setariae
Spores 45 X 1.5 j"
C. striaeformis (Africa)
Conidiophores short, 17-40 /x
C. setaricola
Tehon and Daniels say that C. setaricola is essentially a species of

Scolecotrichum except for the scolecospores. Along this line Scoleco-
trichum is essentially a species of CercospQra except for the ampulli-
form 1- or 2-septate spores. Horsfall (1930) has proposed Cercospora
graminis (Fckl.) Horsfall.
Reference: Tehon and Daniels (1927).
Cercospora sorghi Ell. and Ev,—Leaf Spot

Lesions dark purple on sorghum (Fig. 66) or brown on corn (f.'
maydis, see Sylloge v. 10, p. 656) elongate in stripes several-centimeters
in length, spots later becoming dead and dry, conidiophores amphige-
nous in minute scattered tufts on the dead leaf partfe, few, brown, trun-
cated above and laterally subdentate, 60-80 X 4-6 fx, continuous or
sparingly septate below; conidia slender, faintly 3 or more septate,
70-80 X 3 /x, hyaline. (Ramakrishnan, 1931,j gave 30-132 X 3-8 /x, 1-
to 12-septate.) I
On: Echinochloa crusgallh (L.) Beauv., N. Dak.—Sorghumr-haU'pense (L.)
Pers., Alabama, La., Miss., Nebr., Okla., S. Dak., Tenn., Tex.—5. vul-
gare Pers., Alabama, Fla., Ga., La., Miss., N. Car., S. Car., S. Dak., Va.—
Zea mays La., Alabama, Fla., La., Miss., S. Car., Tenn., Va., W. Va.
Material of a Cercospora on old leaves of Echinochloa found along
the Missouri river bottoms near Mandan, North Dakota, in September,
1944, was determined as C. sorghi by Dr. Chupp. This material had
acicular to obclavate conidia, almost straight, multiseptate, 30-120 X
2-4 fi with amphigenous conidiophores 25-85 X 4-5 /x.'
AVallace and Wallace (1945) report a species-of Cercospora from
Tanganyika on sorghum which was "other t-halnrG. sorghi" (R.A.M.
24:442).
Possibly C. sorghi would fall under the interpretation of C. apii
given by Johnson and Valleau (1949) but the spores appear to'be longer
I'K.. (j(i.- Ci iiuxpiirn sorghi on Hodo sorghuiu, Meridian, Miss. Xl-5- (Photo
by C. L. Lefcbvre, U.S.D.A.)
and wider in some material. The robust size of the host might account
for these differences.
Range: United States, Burma, Cliina, Gold Coast, India, Italy, Peru, Philip-
pines, Uganda.
References: Ellis and Everhart (1887,o, p. 15); Raniakrishnan (1931).
Cercospora tessellata Atk.—Leaf Spot

Spots indefinitely bordered above, usually narrowly oblong, nearly
black below with a bluish tinge caused by numerous black tufts and
bluish cast of tissues affected; conidiophores hypophyllous, densely
fasciculate, fuliginous, short, 10-12 X 2.5-3.0 jn, denticulate, tufts in
longitudinal and usually transverse rows giving a checkered appearance
to the group, 10-12 X 2.5-3.0 ju,; conidia hyaline, slender, terete, curved,
septate, 50-90 X 2.0-2.5 /x.
On: Dactyloctenium aegyptium (L.) Richt., Alabama.
There appears to be no recently published information on this
fungus.
Reference: Atkinson (1897, p. 27).
Cercospora vaginae Krueger—Red Leaf-Sheath Spot

Spots bright red, irregular patches visible from both sides of the
leaf sheaths; by confluence the spots are sometimes large; mycelium
effuse, olive brown, septate; conidiophores arising from the mycelium,
erect, simple, solitary, 80 X 3 ix; conidia subclavate or subcylindric,
hyaline, pleurogenous, 0- to 3-septate, 12.5-37 X 3.0-6.5 /*.
On: Saccharum officinarum L., Fla., Hawaii, La.
An illustration by Roldan (1938) shows narrowly subclavate spores.
Matsumoto and Yamamoto (1934) illustrate C. kopei Krueger as hav-
ing spores which are somewhat broader than C. vaginae. They _give
spore measurements of 26-55 X 4.3-5.7 ji. C. kopei causes a "yellow
spot" or "rotfleckenkrankheit." The type of C. kopei has subfusoid
spores 20-50 X 5-8/i. -•
C. vaginae is widely reported as the cause of a sheath spot on sugar
cane but does not seem to be serious in most cases.
Range: United States, Barbados, Brazil, British Guiana, China, Cuba, For-
mosa, Hawaii, Jamaica, Japan, Java, Mauritius, Peru, Philippines, Puerto
Rico, St. Lucia Is., West Indies in general.
References: Kiryu (1938); Krueger (1890); Matsumoto and Yamamoto
(1934); Roldan (1938); Stevens, F. L. (1925).
Cercospora zeae-maydis Tehon and Daniels—Leaf Spot

Spots brown or tan, evident above and below, 0.5 X 1-2 cm. or con-
fluent and then much more extensive, not limited or ^sardered. Fasci-
culae amphigenous, abundant, scattered, low spreading, long-oval, aris-
ing from nearly closed stomata. Conidiophores lax but ascending, 3- to
8-septate, olivaceous to brown, bearing a single apical geniscar, 70-90
X 4 /i. Spores hyaline, distinctly obclavate, 4- to 10-septate, 50-85 X
5-9 /x.
On: Zea mays h.,J.lL, Ky., Tenn.
Tehon and Daniels (1925) state that this species is distinctive in
that the conidiophores bear a single apical spore_^scar. During the re-
cent War Plant Disease Survey the same spe€iffs,was found in three
states and is of some economic importance (Miller and Wood, 1947).
The type material (111. Ace. 4276) was collected August 29, 1924, indi-
cating from the season of the year that the fungus was probably an
active parasite and not a late autumn saprophyte. Tehon and Daniels
illustrated a spore and conidiophore in their article (1925).
Range: United States, South America.
References: Miller and Wood (1947); Tehon and Daniels (1925).
Cercosporella spp.
The species of Cercosporella found on Gramineae may be distin-
guished by the key which follows.
KEY TO SPECIES OF CERCOSPORELLA ON GRAMINEAE
A. Attacks base of culm causing foot rot, spores 30-80 X 1.5-3.5 i^

Cercosporella herpotrichoides Fron
AA. Attacks leaves causing leaf spot or scald
B.,Causes an eye spot on leaves of Poa praterms, spores 45-90 X 3.6-5.0 /*
C. poagena Sprague
BB. Causes a scald or non-eye-spot type of leaf injury
C. hold Sprague
Cercosporella herpotrichoides Fron—Strawbreaker or

Cercosporella Foot Rot
Fawn color to white, broad'bordered, elongate, less often circular
spots up to 3 cm. in length, occurring on outer leaf sheaths at base of
culms, later penetrating into the interior of stem in spring of year or
earlier, causing light-colored, later brown to black charred-appearing
lesions at the ground level, rarely higher; margins of lesions dark
brown to golden brown; tissues of lesion firm and brittle, later sunken,
culms eventually falling.
Mycelia septate, of two sorts: (1) vegetative, yellow to dark brown,
linear-celled; (2) stromatic, medium- to very heavy-walled, frequently
consisting of polygonal cells forming charred masses on the outside of
stems and sheaths or occurring in cells in the interior of attacked culms
(Fig. 67,A,B,C,D). Conidiophores simple or slightly branched, some-
times swollen at the base and elongated, produced from macrophyphae
subicula. Spores usually produced in spring of the year, rarely as early
as December (in 1934) on spots or lesions, frequently in pairs, often
singly, somewhat curved, obclavate, 2- to several-celled, (mostly 5-7),
variable, 30-80 X 1-5-3.5 /x, mostly 40-60 /A long. On cornmeal in pure
culture sporulation occurs in cool fall or winter weather; spores are
produced in pseudopionnotes, coremia, and in loosely formed sporo-
dochial pads; spores strongly obclavate on this medium, blunt, some-
times pointed at the apex, cells sharply constricted, somewhat doliform,
20-45 X 1.5-3.5 [x (Fig. 68); on potato dextrose agar the spores usually
germinate from terminal cells but sometimes from any cell, and pro-
duce smoky-gray, mounded, compact colonies.
FIG. 67.—Cercosporella herpotrichoides, cross section/through lesions at base

of wheat plants: A, young infection in outer leaf sheath showing direct pene-
tration of the outer cells following the formation 'of a stroma. Note the slight
constriction of the hyphae in passing through the J cell walls of the host and the
tenaency to expand on reaching the inner walls, X675; B, detail of older infec-
tion in leaf sheath, showing increased septation and hyphal development. Shows
detail of the passage of the fungus through the thickened cgll walls, X675; C,
detail of very young infection in leaf sheaths in March, showing^penetration
through stomatal opening. Note strands passing through stomatal (scY, fungi in
guard cell (gc) and stromatic mass completely filling substomatal cavity (ssc),
X190; D, portion of a section at base of a young wheat tiller 7 weeks after the
wheat has been inoculated with C herpotrichoides^ at sowing time in the green-
house. The fungus has penetrated through the leaf sheaths (a,b,c) and is about
to attack the inner sheath, d. Note the fungus strortjata, the bulbous enlarge-
ments on some of the hyphae within the cells, and tjhe thickenings of the cell
walls in certain of the infected areas, especially beneath stromata. This section
happens to be taken so that no well-developed veins are shown. ,>><500. (From
U. S. Dept. of Agr.)
' /
On.: Aegilops cylindrica Host., Oreg.—A. ovata L.,'^reg.—A. triuncialis L.,
Oreg.—Agropyron cristatum (L.) Gaertn., Wash.-Â. inerme (Scribn. and
Sm.) Rydb., Wash.—A. riparium Scribn. and Sm., Wash.—Avena sativa
•^L., Oreg., Wash.—Bromus carinatus Hook, and Arn., I d a . ^ B . inermis
Leyss, Ida.—B. japonicus Thunb., Ida.—B. sterilis L., Ida.—B. tectorum
L., Oreg., Ida., Wash.—Hordeum vulgare L., Ida., Oreg., Wash.—Koeleria
cristata (L.) Pers., Wash.—Poa secunda Presl, Wash.—Secale cereale L.,
Ida., Oreg., Wash.—Sitanion hystrix (Nutt.) J. G. Sm., Ida.—Triticum
aestivum L,, Ida., Oreg., Wash.—T. dicoccum Schrank, Oreg., Wash.—
T. durum Desf., Oreg., Wash.-—T. monococcum L., Oreg., Wash.—T.
spelta L., Oreg., Wash.
In the United States the strawbreaker foot rot is restricted to prairie
areas in the winter wheat region of the Columbia Basin where the
precipitation in the infested places is 14 to 25 inches and the climate
FIG. 68.—Conidia of Cercosporella herpotrichoides from culms of wheat.

Klickitat County, Wash.
is not. severe. During the past decade the disease seems to have spread
to some extent but still lies within the same ecological areas as it did
during the years in which it was intensively studied (Sprague,
1931,a,£)).
In other parts of the world strawbreaker occurs in areas close to
the ocean in cool temperate zones. The Columbia Basin area has one
of the few inland infestations and even there the weather is tempered
by the prevailing winds from the not-too-distant western part of
Oregon and Washington. C. herpotrichoides is greatly'favored by open
wet winters which induce excessive growth in -winter wheat or winter
barley. In the United States it has never been found on spring-sown
grain but Glynne (1946) reported the disease on] spring-sown barley
in southern Scotland and to a lesser extent in sp;uthern England. In
Germany, Bockmann (1934) found that the fungus produced spores in
winter. They could germinate at 0° C. and infestation, occurred at
15° C.
This fungus causes a true foot rot as contrasted with a root rot. The
injury occurs at the base of the culm. Fawn-colored to white lesions
with brown borders form at the ground line in early spring. These are
an eye-spot type of lesion. Later stromatic mycelium covers the base
of the culms with black charred-appearing hyphae. The culms may
break over at this point. This is called strawbreaking in Europe and
this seems to be a logical name. Rhizoctonia solani Kuehn also some-
times causes strawbreaking but the outer, lesions often have longer,
sharper apical and basal portions, hence the name "sharp eye spot."
Wheat fields which are seriously infested with C. herpotrichoides
have characteristically pale green leaves and the heads are smaller than
normal. Later so much of the grain may be broken over and tangled
together that harvesting is difficult and costly. Fallen grain was ex-
tremely prevalent in eastern Washington in 1948 and snarling and
twisting in falling was largely caused by Cercosporella. Overly tall
grain may lodge but it tends to fall in one direction while strawbreaker
grain tends to fall in criss-cross or jack-straw arrangement. ;
In the years since intensive study was made of this disease some
of the recommendations for control have become obsolete, but for the
main part these suggestions for control of the fungus are still applicable
(Sprague, 1949,b). Orfed, Rex, and Hymar wheal; and Olympia barley
are preferred varieties while short wheats are preferred to taller ones.
In a very bad season, such as 1948, even the short-straw wheats, such
as Elgin, broke over with strawbreaker. i
While strawbreaker,has appeared in places where«-it,had not been
found in earlier years it has also disappeared, because ol^'change in
cropping practices and because of soil erosion, from some places where
it was once serious. In the High Prairie region of Klickitat County,
Washington, the top soil has largely washed away and with it the
foot rot fungus which does not thrive in the subsoil (Sprague, 1937,c;
1949,d). An extensive acreage of alfalfa has replaced much of the land
formerly seeded to wheat in Klickitat County and this has solved their
foot rot problem. Perennial legumes could well be seeded in parts of
the Palouse where it is obvious the top soil is _fast disa'^ppearing. The
spasmodic foot rot problem there is vanishing by^he saine route.
Fertilizers have not been widely used to combat strawbreaker.
Calcium cyanamid in pellet form, 100 lbs. per acre, had some merit on
High Prairie in earlier years to permit wheat to recover. Ammonium
phosfate (16-20-0) in pellet form is suggested for the same reason
where legumes in rotation are not immediately available.
In earlier times in France sundry materials were applied to the soil
to combat root rots. Foex (1926) and others tried sulfuric acid on
various plots during January to March but this drastic method is
seldom mentioned today. Rabate (1927) reported that sulfuric acid
was effective only if the root-rot fungi had not penetrated deeper than
the first sheath of the young plants. No doubt dusting with some of the
modern organics in late winter would be an effective but highly expen-
sive means of control.
The host range of this fungus is not extensive and when found on
grass weeds the infection has obviously come from the heavily infected
cereal to the adjacent less severely diseased grasses. In 1948 brome
grasses growing in or on the edge of diseased wheat fields in Latah
County, Idaho', were heavily infected. No doubt intensive search would
have revealed other unreported hosts in diseased fields during this year
of heavy loss.
Range: United States, Australia (S. Australia), Belgium, Denmark, Eng-
land, France, Germany, Italy, Netherlands, New Zealand, Norway, Scot-
land, Sweden, S. Africa.
References: Adam (1940); Bockmann (1934); Detroux (1946,a,6); ,Foex
and Rosella (1930; 1931,a,6; 1933; 1934); Fron (1912); Glynne (1936;
1939,0,6; 1942); Glynne, Dion, and Weil (1945); Gorter (1941); Heald
(Foot rot of wheat. Wash. Bui. 155, 167, 175, 180, 187, in 1920 to 1924);
Moritz and Bockmann (1933); Oort (1936); Parisot (1926); Rabate
(1927); Roemer et al. (1938); Saxby (1943); Sprague (1931,a,6;
1934,6,c; 1936,o; 1937,c; 1939,a,d; 1949,6,d); Sprague and Fellows (1934).
(See also Glynne: J. Minist. Agric. 56:510-14. 1950.)
Cercosporella holci Sprague—^Bufl Scald

Spots tan, yellow bordered to dark buff or tawny, elliptical or ex-
tensive scald-like lesions on any or all leaves of a plant in mid-winter
to late winter and early summer. Mycelia hyaline or faintly chlorinous,
intracellular and intercellular, 0.5-1.2 ju diam., septate, branched; stro-
matic mycelia loose or nearly absent except subhyaline aggregates in
the stomatal and adjacent internal and external leaf parts. Conidio-
phores frequently but little differentiated from the fruiting stroma,
or short erect, simple or once branched, 5-20 X 1-2-2.5 /x, hyaline or
faintly chlorinous, most common near or in the stomata, sterile hyphae
and/or developing spores .10-31 X 1.5-2.0 /x mingled with the fruiting
hyphae. Conidia hyaline, straight to slightly curved, rarely double
refiexed, attached at the larger, elliptically rounded base, slender, ob-
clavate-filiform, 1- to 9-septate, distal and tapering, rounded, contents
homogenous with small globules in the larger cells, 40-105 X 1-5-3.0 /x.
On: Holcus lanatus L., Oreg., Wash.
In severe cases of infection C. hold causes >& j condition similar to
scalding or snow injury. The fungus is favored! by cold, wet weather
and is very common in winter and early spring in' western Oregon and
Washington. Conidia are produced in great numbers in winter (Fig. 69)
but in the spring they may be entirely absent. Suqh late spring lesions
were called nonparasitic by workers who had not fouiid the fungus
fruiting on the grass in winter. That they had not found, the fungus
FIG. 69.—Conidia of Cercosporella hold: A, from type collection; B, another

collection sliowing longer spores. (From Mycologia.)
fruiting was due solely to the fact that they had not ventured out in
the wet and often raw isveather of western Oregon in'-mid-winter. The
fungus is exceedingly abundant in the Willamette Valley, Ore'gon. This
fungus is probably one of the reasons why velvet grass is not a satis-
factory crop for pastures in western Oregon.
Reference: Sprague (1937,6, Fig. 1).
Cercosporella poagena Sprague—Cercosporella Eye Spot

, Lesions light brown with a distinctive straw-colored center and a
surrounding border or halo of yellow; lesions-^fff first small, circular,
later enlarging to nearly the width of the leaf or elongating longitu-
dinally on the leaf for some distance; spots finally becoming dull brown
and eventually fading to straw color. Conidiophores hyaline, obscure.
arising from hyaline, thinly formed aggregating hyphae; conidia hya-
line, elongate, broadly filiform to obclavate, bases blunt, tapering grad-
ually, apices either abruptly acuminate or attenuate at tips, 4- to
7-septate (mostly 4-septate), 45-90 X 3.6-4.6 ix
in the type and 57-90 X 4-5 /x in material from
East Corvallis (Fig. 70).
On: Poa pratensis L., Oreg.
This eye-spot fungus appeared somewhat
mysteriously at Forest Grove and East Corvallis,
Oregon, in 1939. It was collected again in 1940
but the writer has not had an opportunity to
search for it further in the Willamette Valley.
C. poagena is readily distinguished by its yellow-
bordered eye-spot lesions. The spot caused by
Septoria macropoda var. septulata are ashy gray
with reddish borders in some cases while the
lesions caused by Helminthosporium vagans are
purple black or dark brown, also with reddish
borders.
Reference: Sprague (1942,c).
Cerebella andropogonis Ces.—False Smut

or Blackhead
FIG. 70.—Conidia
In the ovaries, often on young ergot, of Cercosporella po-
agena on Poa praten-
masses globose, 5-6 mm. diam., enveloping the sis, type. (From Phy-
glumes, dark or black throughout, glomerules topathology.)
subglobose, usually composed of 3, "spores,"
smooth or slightly roughened, brown, 8-10 (i diam., "spores" ovoid,
6-8 X 5-6 [i.
On: Andropogon furcatus Muhl., Okla.—A. ischaemum L., Okla.—A. sac-
charoides Swartz, Okla.—A. scopdrius Michx., Okla.—A. ternarius Michx.,
Okla.—Anthaenantia-rufa (Ell.) Schult., Miss.—Axonopus affinis Chase,
Fla., Ga., La., Md., N. Car., S. Car., Va.—A. compressus (Swartz) Beauv.,
La.—Brachiaria extensa Chase, Md.—Heteropogon contortus (L.) Beauv.,
Alabama—Panicum virgatum L., Miss.—Paspalum dilatatum Voir, Ala-
bama, La.—P. floridanum Michx., Fla., Ga.—Sorghastrum nutans (L.)
Nash, Alabama, Ark., Kans., Miss., Okla., Tex.—Tricholaena rosea Nees,
Fla.
There have been a number of species described on ergotty material
of grasses from the southern United States. We have included the host
range of most of these under the earliest name that is well known. The
descriptions of most species of Cerebella on grassed are similar. Those
described include C. paspali Cke. and Massee, C. panici Tracy and
. /'!
Earle (1899), and C. spartinae Ell. and Ev. whicH occurs on Distichlis
spicata in Mississippi. . ,
This species belongs in the Tuberculariaceae and is not a smut.
Range: United States, Italy, widely scattered. ,
References: Tracy and Earle (1899); Weiss (1943, p. 193). '
I
Cladosporium herbarum Lk.—Saprophytic Mold
See Mycosphaerella tulasnei (Jancz.) Rothers, its ascigerous stage
Coniosporium Arundinis Sacc.—Mold

See Papularia sphaerosperma (Pers.) v. Hoehnel
Coniosporium shiraianum (Sydow) Bubak—Mold

See Papularia sphaerosperma (Pers.) v. Hohnel
Curvularia spp.
/
Curvularia Boedijn (1933) has 3- to 5-celled, brown, fusiform
spores typically curved or bent, with one or two of the central cells
somewhat enlarged. The spores are universal molds on countless kinds
of plants and on debris in the soil. Because their classification is still
likely to be changed by future investigations, the segregation into
species in this manual is somewhat tentative. It is based on Groves
and Skolko (1945). Their segregation of C. geniculata (Tracy and'
Earle) Boed. from C. inaequalis (Shear) Boed. is apparently justified
but difficult for the routine worker to follow. We have tended to lump
most of this group under C. geniculata. j
I '
KEY TO T H E NORTH AMERICAN SPECIES OP CURVULARIA ON GRAMINEAE
A. Spores sometimes with more than 3 septa
B. Spores averaging 11-14 /* wide
C. geniculata^,,^
BE. Spores averaging 1^-16 M wide
C. inaequalis
AA. Spores 3-septate
B. Spores usually less than 12 /"• wide
C. lunata (Wakker) Boed.
BB. Spores wider, up to 15 /t, olive brown
C. trifolii (Kauff.) Boed.
Curvularia geniculata (Tracy and Earle) Boed.T^Mold

Helminthosporium geniculatum Tracy/Sn^ Earle
Conidiophores brown, septate, simple, geniculate at the tips, varia-
ble in length, (3)-4-5 /x diam., spores brown, unequally ventricose-
fusiform, jiiostly 4-septate, the third cell from the base sliglitly larger
and darker, and the end cells nearly hyaline, more or less curved to
nearly straight, mostly not curved, (21)-28-36-(42) X 11-14-(15) /i.
Spores in pure culture (malt extract agar) (21)-28-40-(50) X (10)-11-
14-(15) [I.
On: Eragrostis rachitricha (type host, apparently not listed in Hitchcock,
1935), Miss., from heads. Isolated from the roots and seeds of all com-
mercially grown cereals and from about 100 species of native and exotic
grasses in Ida., Manit., Minn., Mont., Nebr., N. Dak., Ont., Sask., S. Dak.,
Wash., Wyo. Since most of this material represents saprophytic molds
FIG. 71.—Conidia of Curvularia geniculata isolated on potato-dextrose agar

from Agropyron trachycaulum, Mandan, N. Dak., culture 365 B-3.
it is not deemed necessary to list them here. The complete host range
on grasses and cereals is given in a manuscript by Sprague and Fischer
filed at Washington State College.
Groves and Skolko (1945) isolated this species only from cabbage,
flax, and pea seed. They recognize it as distinct from C. inaequalis be-
cause the third cell of the spores of the latter gives them a broader
aspect than those of C. geniculata. The spores of C. geniculata are
usually under 15 /* wide (Fig. 71), those of C. inaequalis as much as
16 ij. wide. The writer, who has examined hundreds of mounts of this
group from material isolated from the roots of Gramineae, admits some
confusion in his mind. Before the appearance of the article by Groves
and Skolko (1945) he had placed all these in C. geniculata. Most of
this material on potato-dextrose agar had spores 24-46 X 8-14 /*. This
material resembled an Australian species which Hynes (1935) reported
as C. ramosa (Bainier) Boed. The spores from one of Hynes's collec-
tions averaged 32.6 X 13 jn and those of another 35.3 X 13.3 /i. Accord-
ing to the method used by Groves and Skolko both of these collections
7 i
would belong in C. geniculata, but they place C. rdmosa as a synonym
of C. inaequalis. Hynes (1935) found that there'were eleven races of
C. ramosa associated with wheat in Australia.' H^ believed that this
species, or races of it, were more virulent than either H. sativum or
H. tetramera McK. (? Curvularia spicijera'^. Thei forms of C. ramosa
differed considerably, one from another, in pure culture. Hynes believed
that Helmintkosporium-M of Henry (1924) was the'same as C. ramosa.
Chesters (1948) claims that H. biforme Mason and Hughes is a new
name for C. ramosa.
In the northern Great Plains the writer (1944) found that most
isolates of C. geniculata were nonparasitic on seedlings of many grasses
and cereals. In a few cases, however, isolates were parasitic on certain
grasses, but scarcely so on cereals. One culture 367 B-6, isolated from
seedlings of Agropyron sibiricum in mid-April, caused injury to roots
of A. cristatum and moderate injury to Sudan grass. Most isolates
caused some injury, mostly pre-emergence killing, to proso millet
{Panicum miliaceum) and Bouteloua gracilis when added to the soil
at seeding time. In general, however, the fungus is a saprophyte. Since
some isolates are parasitic the fungus deserves watching. Since plant
breeders are upsetting the natural equilibrium by development of re-
sistant varieties of plants other strain^- of fungi may gain ascendance
as did H. victoriae.
In culture C. geniculata produces a luxuriant dark gray to nearly
black, more or less stringy growth on potato-dextrose agar. Other iso-
lates are velvety black, similar to some isolates of H. sativum, and.
such isolates produce quantities of spores. '
Since C. geniculata is so often associated with Helminthosporium
sativum on moldy grain and grass seed it is usually wise to treat such
seed with a good disinfectant such as New Improved Ceresan or Cere-
san-M. Most of our trials have not shown jvery much benefit from
treatment of range grass seeds except those infected with H. sativum.
Slooff, Thung, and Reitsma (1947) lepovtl Curvularia andropogonis
(Zimm.) Boed. {Napicladium andropogonis^ on sereh [Andropogon
(Cymbopogon) nardus] ,from Java. The elongated lesionsjisually ex-
ceeded 20 X 0.5 cm. with grayish necrotic centers and violet-^reddish
margins mostly at the apices of the leaves. The spores were dark brown,
triseptate, 36-60 X 14-25 [i, on conidophores 36-41 X 16-21 [i. Varietal
resistance was noted.
Wernham and Kirby (1941) reported a "going-out" or "rnelting-
out" of Metropolitan bent and Poa annua by either Helminthosporium
or Curvularia, or both, during hot weather in Pennsylvania. This con-
dition was referred to somewhat indirectly in later reports but little
additional information is available at this writing. /
/ ^ '
Range: United States, Australia, Canada, Mexico.-"^ - '
References: Bainier (1908); Boedijn (1933); Groves and Skolko (1945);
Henry il924); Hynes (1935; 1937,c); Lefobvre and Johnson (1941);
Mason (1928); Sprague (1944,6; 1946,e); Wernham and Kirby (1941).
Curvularia inaequalis (Shear) Boed.-—Mold

Helminthosporium inaequale Shear
Conidiophores brown, septate, simple or sometimes branched, genic-
ulate at the tip, variable in length, 4-6 /A diam. Spores 3- to 5-celled,
ventricose fusiform, mostly 4-septate, the third cell especially enlarged,
27-35 X 11-16 jx; in pure culture (malt extract agar) the spores are
(24)-28-40-(60) X (10)-11-16 [i, sometimes 5-septate. Differs from
C. geniculata in being slightly broader in proportion to length with the
ends more obtusely rounded, this difference being mostly confined to
the third cell and, as a result, the spores are more strongly curved than
in C. geniculata. -^
On: The type is from the pulp of diseased cranberries from New Jersey
(Shear, 1907, p. 307). Groves and Skolko (1945) isolated it from
Dactylis glomerata L., Festuca elatior L., F. rubra L., Secale cereale L.
and Zea mays L. from 111., Manit., Ont., and Quebec. Also isolated from
Gramineae in Manit., Minn., Mont., Nebr., N. Dak., Sask., S. Dak., Wash.,
and Wyo. but not always distinguished from C. geniculata (Sprague iso-
lation records).
As we understand these forms the majority of the material in the
northern Great Plains is C. geniculata. In the collection (A. S. series)
series at Washington State College are a number of dried cultures of
these forms for reference.
References: Bainier (1908); Boedijn (1933); Groves and Skolko (1945);
Hynes (1935, 1937,c); Kirby (1941); Mason (1928); Shear (1907, p.
307); Sprague (1944,6; 1946,e); Wernliam and Kirby (1941).
Curvularia lunata (Wakker) Boed.—Mold

Acrothecium lunatum Wakker
Conidiophores are pale brown, septate, simple or branched, genicu-
late at the tip, variable in length, 3-5 /x diam.; conidia pale brown,
3-septate, the third cell from the base is larger and darker colored than
the others, unequally ventricose-fusiform, more or less curved to nearly
straight, 19-30 X 8-12 fx (up to 16 /x. according to Boedijn), on malt
agar spores are 19-32 X 9-15.5 jn, commonly 22-28 X 10-13 /j..
On: Many hosts in warm climates, especially well known as black mold of
rice heads.
Whether this mold is very actively parasitic is still open to ques-
tion. I t appears to be of some economic importance because it is an
almost omnipresent mold of rice grain. Some workers have found this
species to be parasitic. Cralley and Tullis (1937), Matsuura (1927),
and Boedijn (1933) report that the fungus causes a seedling blight of
rice. Martyn (1936) said that C. lunata was present in the brown dis-
/
, i
colored palea of flowering heads of rice in British' Guiana. He was able
to cause the condition by inoculating with spores of C. lunata and in-
cubating under humid conditions. Contrary^ to 'tHese claims Bunting
(1927) was unable to find that the fungus was conclusively'parasitic
on maize in the Gold Coast of Africa. i
Nigam (1936), in India, isolated C. lunata ivom red spots on sor-
ghum leaves and found that the fungus produced) on 2 per cent rice
agar, alternate zones of pink and white mycelium, pink in daylight and
white in darkness. In continuous darkness the colonies were a dense
bottle-green color.
Sanitary measures to inhibit these molds, particularly in warmer
climates, would appear to be difficult. However, such places as rice
straw stacks (Martin, 1939) may be sources of abnormally heavy in-
fection and can be eliminated. Martin and Altstatt (1940) also recom-
mended burning all weeds and volunteer rice around fields, keeping late
seedings away from early ones so that spores are not spread during
threshing of early crops, and keeping the grain and shocks dry to avoid
heating.
C. lunata proper is not common in the northern Great Plains, if it
occurs at all, and seems to be, in general, confined to the warmer cli-
mates. It may occur farther north in^the eastern United States (Le-
febvre and Johnson, 1941). Wernham and Kirby (1941), as mentioned
previously, also reported Curvularia sp. as associated with a "going-
out" or "melting-out" disease of Agrostis palustris and Poa annua in
Pennsylvania. This fungus grew best at 30° C. and was inhibited if
the temperature dropped below 25° C. While this fungus will need to
be compared critically with the two previously mentioned species its
affinity with warm temperatures would suggest comparing it first with
C. lunata. /
Range: World-wide in the warm temperate and;tropical regions.
References: Bainier (1908); Boedijn (1933); Bunting (1927, 1928);
Cralley and TuUis (1937); Groves and Skplko (1945); Lefebvre and
Johnson (1941); Martin (1939); Martin and Altstatt r 1940); Martyn
(1936); Mason (1928); Matsuura (1927); Nigam (1936); TuUis. (-1936);
Wernham and Kirby (1941). '_
Curvularia trifolii (Kauf.) B6ed.—Mold

Brachysporium trifolii Kauf.
Conidiophores brown,, septate, simple, geniculate at the tip, 5-6 /u,
diam.; spores brown to olive brown, 3-septate, the third cell much
larger and darker than the others, unequally ventricose-fusiform,
mostly strongly curved, sometimes forked or triangular lobed, 25-35-
(38) X 11-15 iJ.; in pure culture on malt extract agar the spores are
(20)-25-35-(38) X (10)-12-16-(19) /..
On; Ecfmochloa crusgalli (L.) Beauv., Minn., N. Dak., S. Dak. '
This fungus, which has been isolated only from this one species of
grass, has in the opinion of the writer olive green or olive brown rather
than brown spores. Our material resembles the illustration by Groves
and Skolko (1945, plate VI) in that the spores are very plump, almost
triangular, even more pronounced in this tendency than those of the
related C. lunata (Fig. 72,A).
C. trifolii was originally described by Kauffman on white clover
(Bonar, 1920) and studied further by Bonar (1924). Groves and Skolko
FIG. 72.—A, Conidia of Curvularia trifolii isolated on potato-dextrose agar

from Echinochloa crusgalli, Dickinson, N. Dak., culture 210 B-2; B, conidia of
Brachycladium spicijerum isolated on potato-dextrose agar from Agropyron
dasystachyum, Mandan, N. Dak., culture 141-A-3.
found it on soybeans, peas, cucumbers and pumpkins. They apparently

did not isolate this species from grasses although Machacek has ob-
tained it in Manitoba (personal letter) and he determined the writer's
isolates from Echinochloa.
G. cymbopogi (C. W. Dodge) Groves and Skolko occurs in Guate-
mala on citronella and lemon grass (Dodge, 1942). It belongs in the
"geniculata" group and is probably not distinct from C. geniculata.
C. jalcata (Tehon) Boed. occurs on Setaria in Puerto Rico (Tehon,
1919) and has spores 35 X 10 ;u but which Groves and Skolko remeas-
ured and recorded as (22)-24-28-(31) X (10)-12-13-(14) p..
C. andropogonis (Zimm.) Boed. was originally described as Napic-
ladium with spores 40 X 20 ;a but Boedijn (1933) gave the spore size
as 38-66 X 13-23 ix and stated that the basal cell bore a small hyaline
appendage.
C. penniseti (Mitra) Boed. has spores 30.4-41.8 X 17.1-20 /* and
was compared critically with C. lunata by Mitra (1941).
Range: Of C trifolii: United States, Canada. ' |
References: Bonar (1920, 1924); Groves-and Skolko (1945); Sprague
(1946,e). " '
Dactylaria graminum (Schw.) Sacc.—îLeaf Spot

Dactylium graminum Schw. j
Conidiophores sparse, simple gray, erect, conidia 3-4 digitate, con-
colorous.
On: Leersia oryzoides (L.) Swartz, Penn.
This fungus is not determinable from the description. Perhaps the
digitate nature of the spores refers to the whorls of conidiophore
branches such as occur in Dactylium.
Dactylaria leersia Sawada (1916 and Tanaka, 1920) which S. Ito
(Bot. Mag. Tokyo 32:307-308, 1918) placed in Piricularia has been
carefully described. The fungus is very sinailar to Piricularia oryzae.
The conidiophores were fasciculate or solitary, simple, 2- to 3-septate,
48-88 X 4-5 /A. The conidia were 20-35 x'7-10 /i, short conic to elon-
gate conic.
References: gaccardo (1886, Sylloge v. 4, p. 195); Sawada (1916);
Schweinitz (1832); Tanaka (1920).
EUisiella caudata (Pk.) Sacc.^—Leaf Spot on Old Leaves

Colletotrichum caudatum Pk.
Tufts erumpent, oblong or sublinear, black, minute, 250-330 /i long,
125 [I wide. Sterile hyphae erect, cuspidate, rather rigid, continuous, or
often distinctly septate, 100-180 X 7 jn, dark fuliginous, subbulbose at
base. Conidiophores at the base of the hyphaie, subpyriform, subobtuse
at the apex, 2-3 spored, 15-20 X 6 ;«,, very pale olivaceous. Conidia
fusoid, slightly curved, 28 X 5-6 fi, hyaline or yellowiBb,,nucleate, at-
tenuated below into a slender, curved base or pedicel, 25-30'X 1 /^
(from Journ. Mycology 1:105, 1885). ~
On: Agropyron trachycaulum (Lk.) Malte, Okla.—Andropogon annvlatus
Forsk., Ga.—A. furcatus Muhl., Okla., Wise—A. halii Hack., Okla:—A.
scoparius Michx., Okla., Wise.—A. ternarius Michx., Okla.-—Aristida cur-
tisii (A. Gray) Nasli, Okla.—Eragrostis secundiflora Presl, Okla.—E. spec-
tabilis (Pursh) Steud., Okla.—Glyceria paucijiora Presl, Wyo.—Koeleria
cristata (L.) Pers., Okla.—Manisuris cylindrica (Michx.) Kuntze, Okla.—
Setaria. lutescens (Weigel) F. T. Hubb, Okla.—Sorghastri^m nutans (L.)
Nash, N. J., Okla.—Sporobolus heterolepis A,.-Gray, Wise.
A description of this species is given in English in the Journal of
Mycology (v. 1, p. 105), probably by the editor. He states: "We agree
with the opinion expressed by Professor P ^ k in the 35th Report of the
New York State Museum, page 139, that this genus is not suificiently
distinct from Colletotrichum." Saccardo described the genus EUisiella
as follows: "Hyphae sterile, erect, simple, fuscous. Conidia fusoid, with
a long, curved beak above." The description in the Journal of Mycology
mentions the curved base but not the curved beak above. Peck (1884)
states: "If scarcely differs from Colletotrichum, except in having spores
longer pointed." Clements and Shear placed this fungus in the Mo-
niliales.
References: Anon. (J. Myc. v. 1, p. 105, 1885,o); Peck (1884); Saccardo
(Sylloge, 1886, v. 4, p. 315).
Epicoccum spp.—Molds
Species of Epicoccum, generally referred to E. neglectum Desm.,
occur on ripening heads of grasses and cereals in wet weather. The
material which we have seen in the western United States seems to be
saprophytic secondary or tertiary black molds on dead plant parts.
Epicoccum is readily recognized by its short black conidiophores aris-
ing from shallow sporodochial pads and bearing globose, somewhat
echinulate, multiseptate dark spores.
There is a report by Ito and Iwadare (1934) on two species of
Epicoccum which caused a red blotch disease of harvested rice grains
and*which were associated with a complete loss of germinative capacity
of the grain at Hokkaido, Japan. Besides E. neglectum they discussed
E. oryzae Ito and Iwadare. It has branched, septate, olivaceous hyphae,
3.7-6.2 ju diam. producing globose or subglobose, black, punctiform
sporodochia, 45-210 m diam.; and yellow to olivaceous conidiophores
2.5-7.5 IX. long, bearing globose to subglobose or pyriform, granular-
verrucose, olivaceous conidia, 9.9-23.1 X 6.6-16.5 /i, consisting of 1 to 5
cells. These men were unable to infect healthy leaves or stems of
wheat, oats, corn, beans or buckwheat, all of which were saprophytic
hosts for E. neglectum. Properly dried rice bundles were not infected
by the red blotch disease.
Range: Of the various species—world-wide.
References: See Saccardo Syll. Fung., various volumes; Ito and Iwadare
(1934).
Fusarium spp.
The number of species of Fusarium that occur on Gramineae is not
great but the artificial system of classification, still in vogue, in part,
has so multiplied the number of these subdivisions as to represent a
considerable array of names. The recent studies by Snyder and Hansen
(1940, 1945, 1948) which have only been outlined in some cases, indi-
cate that single spore cultures, repeatedly made, yield facsimiles of
variously described species in the Roseum group. I t is their belief
that a number of species are unnecessary, such as F. graminearum,
F. culmorum, F. equiseti, F. scirpi^ F. 'sporotrichioides, and F. ave-
naceum which are variants of the pink and buff types all lumped under
F. roseum. While the evidence which they advance Seems highly discon-
certing to those who cling religiously to the Wollenweber system, we
hesitate to use their whole system until some of the details have been
ironed out. We do, however, prefer to use the all-inclusive F. oxysporum
Schlecht. em. Snyder and Hansen and, for all practical purposes, not
explore the multiplicity of forms that are involved in this species which
is of comparatively little consequence as a parasite of cereals.
A key to the species of Fusarium which occur on Gramineae in the
United States and Canada as grown on 2 per cent potato-dextrose agar
in diffuse light at 70° F. is here presented.
KEY TO SPECIES OP TUSARIUM ON GRAMINEAE
A. Typically producing red, pink, or wine-colored mycelium. Sometimes color

is scanty or absent in group with pip-shaped {Sporotrichum) microscopes
B. Producing at least a few iSporotric/iMm-like microspores
F. poae (Pk.) Wr.
BB. No Sporotrichum-like microspores produced
[The next four species are variants of F. roseum of Snyder and Hansen
(1945, 1948)]
C. Colony typically rosy pink; spore masses bright carrot color, often
absent; macrospores somewhat acuminate at distal end, 5-septate
spores average 45 X 4.1 fi, 7-septate average 55 X 4.5 /* (the common
"pink" Fusarium on the roots of plants in the semiarid country) '
F. acuminatum Ell. and Ev;
CC. Colony white and rose or pink; spore masses carrot red; spores
very narrow, 3-septate, average 34 X 3.1 f^, 5-septate average 50 X 3.7
M, common on heads of Gramineae in moister regions
F. avenaceum (Fr.) Sacc.
CCC. Colonies bright carmine; spore masses orange red, often profuse;
spores stout, not acuminate, 3-septate, average 29 X 6.1 n, 5-septate
38 X 6.6 iJ- and 7-septate 46 X 7 M, usually strongly parasitic
I F. culmorum (W. G. Sm7)-Sacc.
CCCC. Colonies carmine or darker; conidial masses brighr"orange;
spores longer than in F. culmorum, proportionately narrower; 3-septate
average 41 X 4.3 /*, 5-septate average 51 X 4.9 M, 7-septate 73 X 5.4 M,
strongly parasitic
F. graminearum pchw.
CCCCC. Colonies white and bright rose or pale salmon; spores small,
scarcely heeled, 1- to 3-septate, 1-septate average 16 X 2.8 ^, 3-septate
25 X 3 M, thrives at or near freezing temperatures
F. nivale (Fr.) Ces. .
/
AA. Typically producing a peach-colored, lateryi¥liff to nearly brown
myceli^mi; macrospores typically acuminate on distal end, macroconidia
3- to 7-Heptate; common as a saprophyte on grass roots in early to mid-
spring
F. equiseti^(Cda.) Sacc.
AAA. Typically producing white or bluish mycelia
B. Microspores produced in chains, growth generally white on potato-dex-
trose agar
F. moniliforme Sheldon
BB. Microspores not produced in chains
C. Colonies white, soon lavender, blue, purplish or less often pink or
isabelline tinted in the depths of the medium; mostly aseptate micro-
spores, 7-10 X 2.1-3 ft; terminal and intercalary chlamydospores
numerous; very common on roots of grasses in the Plains country and
in the Far West in late spring and summer
F. oxysporum Schl. em. Snyd. and Hans.
CC. Colonies becoming intensely blue in the depth of the medium;
spore masses dirty cream to brownish; macrospores long cylindrical,'^
fusiform, 3-septate, 28-42 X 4.1-6.2 /*, 5-septate mostly 42-51 X 5.0-6.3
fi; saprophytic on Gramineae and not often abundant
F. solani (Mart.) Appel and Wr.
The above key may be of some use to students. It represents a con-

siderable reduction in the number of species once recognized on Gra-
mineae. It seems to us that the number, if reduced much further,
might eliminate one evil and create another. If we lump such sapro-
phytes as F. equiêti in with such parasites as F. graminearum we cer-
tainly must segregate the components with species subdivisions or
racial groups of some category or other. We have followed Gordon
(1933, 1939, and 1946) and at this time feel that the system is practi-
cal, at least. The final decision on the group awaits publications based
on study of the entire life cycles of these species or forms. Until workers
have accepted any other system of classification we prefer to use the
hybrid system above designated. This will be somewhat more thor-
oughly discussed in connection with the various species of Fusarium
which are parasitic on Gramineae. Our studies of this group have been
from the viewpoint of a field-laboratory pathologist.
References to Fusarium as a Genus: Atanasoff (1923); Bennett (1938);
Carrera (1939, 1940); De Haan (1937); Dickinson (1932); Forster and
CroU (1941); Gordon (1933, 1939, 1946); Gordon and Sprague (1941);
Leonian (1929); Millrr, J. J. (1946,a,6); Padwick (1941); Raillo (1935,
1936); Sherbakoff (1947); Simmonds (1941); Snyder and Hansen (1940,
1945, 1948); WoUenweber and Reinking (1935); Wollenweber, Sherbakoff,
Reinking, Johann, and Bailey (1925).
Fusarium acuminatum Ell. and Ev.—Secondary Root Rot

F. scirpi var. acuminatum (Ell. and Ev.) Wr.
Mycelium white to buff with carmine-pink substratum, firm, felty,
cottony to fluffy, conidia in sporodochia orange in mass, sickle shaped
but definitely narrowed at the distal end, with a foot-shaped basal cell,
0- to 9-septate, mostly 3- to 7-septate, 19-114 X 2.0-6.5 /t, commonly
334 DISEASES OF CEREALS AND-GRASSES
' \ ' .
30-70 X 2.5-5.0 jx] chlamydospores 12(6-16) /^ thick, globose to ovoid,
intercalary. ' ~' ^ - _|
On: Aegilops cylindrica Host., Oreg., Wash.—A. ovata L., Oreg.—A. tri-
uncialis L., Oreg.—Agropyron angustiglume'NevsM\N. Dak.f—4. caninum
(L.) Beauv., Wash,—A. ciliare (Trin.) Franch, N. Dak".—A. cristatum
(L.) Gaertn., Ida., Minn., Mont., Nebr., N. Dak., Oreg., S. Dak., Wash.,
Wyo.—A. desertorum (Fisch.) Schult., Mont., Nebr., N. Dak., Oreg.,
S. Dak., Wyo.—A. elongatum (Host.) Beauv., Nebr., N. Dak.—A. grif-
fithsii Scribn. and Sm., Mont.—A. inerme (Scribn. and Sm.) Rydb., Nev.,
Oreg., Wash.—A. intermedium (Host.) Beauv., Minn., Nebr., N. Dak.,
S. Dak., Wash.—A. michnoi Roshev, Ida., N. Dak., S. Dak., Wash.—A.
mongolicum Keng, N. Dak.—A. pubicalum var. pilipes (Keng) ined.,
fide Swallen, N. Dak.—A. repens (L.) Beauv., Minn., Mont., Nebr.,
N. Dak., Oreg., S. Dak., Wyo.—A. riparium Scribn. and Sm., N. Dak.,
Wash., Wyo.—A. sibiricum (Willd.) Beauv., Minn., Mont., Nebr., N.
Dak., Oreg., S. Dak., Wash., Wyo.—4. smithii Rydb., Colo., Minn., Mont.,
Nebr., N. IDak., S. Dak., Wyo.—A. spicatum (Pursh) Scribn. and Sm.,
Ida., Oreg., Wash.—A. subsecundum (Lk.) Hitchc, N. Dak., Wyo.—A.
trachycaulum (Lk.) Malte, Mont., Nebr., N. Dak., S. Dak., Wyo.—A.
trichophorum (Lk.) Richt., N. 'Da.k.—;Agrostis alba L., Minn., Nebr.,
N. Dak., Wyo.—A. canina L., Oreg.—A. palustris Huds., Oreg., Wash.—
A. scabra Willd., S. Dak.—Andropogon jurcatus Muhl", Nebr., N. Dak.,
S. Dak., Wyo.—A. hallii Hack., N. Dak., S. Dak.—A. scoparius Michx.,
Mont., Nebr., S. Dak.—Aristida longiseta Steud., Nebr., Wyo.—Bouteloua
curtipendula (Michx.) Torr., N. Dak., S. Dak., Wyo.—B. gracilis (H. B.
K.) Lag., Mont., Nebr., N. Dak., S. Dak., Wash., Wyo.—Bromus arvensis
L., N. Dak.—B. carinatus Hook, and Arn., Mont., N. Dak., Oreg., Wash.,
Wyo.—B. commutatus Schrad., Oreg.—B. erectus Huds., NT Dak., Wash.
—B. inermis Leyss, Ida., Minn., N. Dak., S. Dak., Wyo.—B. japonicus
Thunb., Ida., Minn., Mont., N. Dak., S. Dak., Wash., Wyo.—5. mollis
L., Ida., Oreg., Wash.—B. pumpellianns Scribn., AVyo.—B. tectorum L.,
Ida., Mont., Nebr., N. Dak., Oreg., S. Dak., Wash., Wyo.—BuchloS
dactyloides (Nutt.) Englm., Nebr., N. Dakl—Calamagrostis montanensis
Scribn., S. Dak.—Calamovilja longifolia (Hook.) Scribn., N. Dak., S. Dak.
—Cenchms pauciflorus Benth., N. Dak., S. Dak.—Chloris virgata Swartz,
N. Dak.—Dactylis glomerata L., Minn., Mont., N. Dak7pS^-Dak^ Wash.—
Danthonia pilosa, N. Dak.—Deschampsia elongata (Hook.) Munro, Wyo.—
Digitaria sanguinalis (L.) Scop., N. Dak.—Distichlis striata (Torr.) Rydb.,
Mont., N. Dak., S. Dak., Wyo.—Echinochloa crusgalli (L.) Beauv., Minn.,
N. Dak., S. Dak., Wyo.—E. crusgalli var. jrumentacea (Roxb.) Wight,
N. Dak.—Elymus ambiguus Vasey and Scribn.,. N. Dak.—E. canadensis
L., Minn., Mont., Nebr., N. Dak., S. Dak., Wash., Wyo.—E. canadensis
var. robustus (Scribn. and Sm.) Macken. and Bush, N. Dak.—.£^. dahuricus
Turcz., N. Dak.—E. dasystachys Trin., N. Dak.—E. excelsus Turcz.,
N. Dak.—E. giganteus Vahl., N. Dak.—E. glaucus Buckl., N. Dak.—S.
interruptus Buckl., N. Dak.—E. junceus Fisch^^JSfebr., ,N. Dak.—E. ma-
counii Vasey (Hordeum jubatum x Agropyron trachycaulum), N. Dak.,
* Wyo.—E. sibiricus L., N. Dak.—E. triticoides Buckl., N. Dak.—Eragrostis
curvula (Schrad.) Nees, N. Dak.—E. pilosa (L.) Beauv., N. Dak.—E.
trichodiis (Nutt.) Nash, N. Dak.—Festuca dertonensis (All.) Aschers.
and Graebn., Oreg., Wash.—F. elatior L., Oreg., N. Dak., Wyo.—F. elatior
var. arundinacea (Schreb.) Wimm., N. Dak., Oreg.—F. megalura Nutt.,
Oreg.—F. myuros L., Oreg., Wash.—F. octoflora Walt., Ida., N. Dak., S.
Dak., Wash., Wyo.—F. ovina L;, N. Dak.—F. rubra L., N. Dak., Oreg.—F.
rubra var. commutata Gaud., Minn., N. Dak., Oreg.—Hilaria jamesii
(Torr.) Benth., N. I)a.k.—H. mutica (Buckl.) Benth., N. Dak.—Hordeum
brevisubulatum (Trin.) Lk., N. Dak.—H. bulbosum L., N. Dak.—H.
distichon L., General in Ida., Minn., Mont., N. Dak., Oreg., S. Dak.,
Wash., Wye—F. jubatum L., Minn., N. Dak., Oreg., S. Dak., Wyo.—
H. pusillum Nutt., Mont., S. Dak.—H. vulgare L., Alta., Calif., Colo.,
Ida., Iowa, Manit., Minn., Mont., Nebr., N. Dak., Oreg., Sask., S. Dak.,
Wash., Wise, Wyo.—Koeleria cristata (L.) Pers., Mont., N. Dak., Oreg.,
S. Dak., Wash., 'Wyo.—Lolium multiflorum Lam., N. Dak., Oreg., Wash.—
L. remotum Schrank, Wash.—Melica virgata, N. Dak.—Muhlenbergia
pungens Thurb., Nebr.—M. racemosa (Michx.) B. S. P., Minn., N. Dak.—
M. squarrosa (Trin.) Rydb., N. Dak.—Oryzopsis hymenoides (Roem.
and Schult.) Rick., N. Dak., S. Dak., Wash., Wyo.—Panicum capillare
L.; Mont., N. Dak., S. Dak.—P. dichotomiflorum Michx., Nebr.—P.
miliaceum L., Nebr., N. Dak., S. Dak., Wash.—P. perlongum Nash, Nebr.—
P. subvillosum Ashe, Minn.—P. virgatum L., Nebr., N. Dak., S. Dak.,
Wyo.—Phalaris arundinacea L., Minn.—P. tuberosa L., N. Dak.—Phleum
alpinum L., N. Dak.—P. phleoides (L.) Karst., N. Dak.—P. pratense L.,
Minn., Mont., N. Dak., S. Dak., Wash., Wyo.—Pkragmites communis
Trin., N. Dak.—Poa alpina L., Mont., Wyo.—P. ampla Merr., Mont.,
N. Dak., Wash.—P. annua L., N. Dak., Oreg., Wash.—P. arida Vasey,
N. Dak.^P. bulbosa L., Ida., N^ Dak., Wash.—P. canbyi (Scribn.) Piper,
N. Dak., S. Dak., Wyo.^P. compressa L., Minn., N. Dak., Oreg., Wyo.—
P. cusickii Vasey, Wyo.—P. glaucifolia Scribn. and Wils., N. Dak.—P.
interior Rydb., Wyo.—P. nevadensis Vasey, N. Dak.—P. palustris L.,
Minn., Mont., N. Dak., S. Dak.—P. pratensis L., Calif., Ida., Minn., Nebr.,
N. Dak., Oreg., S. Dak., Wash., Wyo.—P. scabrella (Thurb.) Benth., N.
Dak., Oreg., Wash.—P. secunda. Presl, Mont., N. Dak., Oreg., S. Dak.,
Wash., Wyo.—Schedonnardus paniculatus (Nutt.) Trel., Mont., Nebr.,
N. Dak., S. Dak., Wyo.—Secale cereale L., Calif., Colo., Ida., Iowa, Minn.,
Mont., Nebr., N. Dak., Oreg., S. Dak., Wash.—Setaria italica (L.) Beauv.,
Iowa, N. Dak.—S. lutescens (Weigel) F. T. Hubb, N. Dak,, S. Dak.—
S. viridis (L.) Beauv., Minn., Mont., Nebr., N. Dak., S. Dak., Wyo.—
Sitanion hystrix (Nutt.) J. G. Sm., Ida., Mont., Oreg., Wash., Wyo.—
Sorghum vulgare PersTJ'lowa, Minn., Nebr., N. Dak., S. Dak.—S. vulgare
var. sudanense (Piper) Hitchc, Kans., N. Dak.—Spartina pectinata Lk.,
N. Dak.—Sporobolus asper (Michx.) Kunth., Mont.—S. cryptandrus
(Torr.) A. Gray, Ida., N. Dak., Wyo.—Stipa columbiana. Macoun, N.
Dak.—S. columbiana var. nelsonii (Scribn.) Hitchc, Wash.—S. comata
Trin. and Rupr., N. Dak., S. Dak.,'Wash., Wyo.—S. sibirica (L.) Lam.,
N. Dak.—S. viridula Trin., N. Dak., S. Dak., Wyo.—Trisetum cernuum
Trin., Wash.—Triticum- aestivum L., general—T. dicoccum Schrank, N.
Dak., Oreg., Wash.—T. durum Desf., N. Dak., Oreg., Wash.—T. monococ-
cum L., Oreg., Wash.—T. spelta L., N. Dak., Oreg., Wash.—T. timophe-
evii Zhuk., N. Dak.—Zea mays L., Ida., Iowa, Minn., Mont., Nebr.,
N. Dak., Oreg., S. Dak., Wash., Wyo.
/
This common pink mold causes yel'low-browiii lesions on the roots
of some grasses and cereals but is usuallY associajted with other fungi
without any specific symptoms. It is a common saprophyte on practi-
cally all Gramineae in the western United States but causes some seed
rot in spring and fall on small-seed crops. It is only a weak parasite on
wheat and barley and is apparently nonparasitic'on oats. It is part of
the complex which Snyder and Hansen place in tl\e species F. roseuni
with several others (1945).
This species differs from F. equiseti in the rose-pink, firm colony
that is produced on potato-dextrose agar. The mycelium tends to be
coarse and breaks up into characteristic stiff fragments in slide mounts.
It often produces no spores, although some isolates form carrot-red
masses of macrospores which are slightly smaller than those of F.
equiseti. Five-septate macrospores average 45 X 4.1 /*, and 7-septate
55 X 4.3 /JL. The curved spores have the typical Gibbosum shape,
narrow at the apex. This fungus is readily distinguished from F. cul-
morum and F. graminearum by its firm- felty growth with rose-pink
coloration instead of the carmine and yellow buff looser mycelium
of the two scab-forming species. At Pullman, Washington, however,
we noted a greater tendency for the fungus to produce forms approach-
ing these species than we had at Mandan, North Dakota.
F. acuminatum apparently is favored by cool, wet weather in March
and April when it develops on the roots of overwintering perennial
grasses and on young seedlings and sprouting seeds. It was isolated
from frozen grass sod in subzero weather in January, 1946 at Mandan,
North Dakota (Sprague, 1946,/). I t is a very common saprophyte fol-
lowing Cercosporella herpotrichoides in the Pacific Northwest, and is
one of the components of common root rot and dry-land root rot. It
also occurs in acid soils in the peat area of norjthern Minnesota and
westward, and is common in soil of relatively high alkalinity. Cultures
of this fungus isolated from red soil in Wyortiing were particularly pale
in color and evidently were a local race. Sometimes other cultures from
various areas have shown marked buff colorations, but these colorations
are frequently induced by bacterial associates in the^cultures. The
writer's inoculation trials (1944,b) have shown considerable variation
in racial traits among the isolates used in the pathogenicity tests. Some
were mildly parasitic but many were essentially saprophytes.
Range: General. /
References: Appel (1924); Fellows (1925-1944);'Gordon (1939); Gordon
and Sprague (1941); Sprague (1944,&; 1946,/); Wollenweber and
Reinking (1935).
/
Fusarium avenaceum (Fr.) Sacc.—M«ld'and Stem Rot
Mycelium white with carmine tones, cottony, substratum often red
or dark amber, conidia scattered or in dull orange masses, very narrow,
FUNGI IMPERPECTI—MONILIALES 337
curved, with narrow foot cell, 0- to 7-septate, mostly 3- to 5-septate,
18-74 X 2.7-4.4 /., mostly 35-65 X 3.0-4.4 ;a.
On: Avena sativa L., Calif., N. B., N. S., Ohio, P. E. Is., Que.—Bromus
pumpellianus Scribn., Wyo.—Dactylis glomerata L., Oreg.—Elymus
glaucus Buokl., Wash.—Festuca rubra var. commutata Gaud., Oreg.—
Hordeum vulgare L., Manit., N. B., N. S., P. E. Is., Que., Wise.—Secale
cereale L., Ind., Ohio—Triticum aestivum L., B. C, Calif., Manit., N. B.,
N. S., Ont., Que. F. avenue also reported as a shoot Wight of Agrostis alba
in Quebec.
This fungus apparently prefers marine or moist climates. It some-
times occurs as a parasite on rust spores on grasses in the western
United States during rainy periods. It is a common mold on grass and
cereal seeds in eastern Canada. The writer has seen it on a stem canker
of Dactylis glomerata in the Cascade Mountains, Oregon. It was not
found on any root-rot material in North Dakota, although it occurs
not far north of there in Canada (Gordon, 1933). It was isolated from
the roots of Bromus pumpellianus Scribn. collected at 7,200 feet eleva-
tion in the Big Horn Mountains, Wyoming, in June, 1946. Johnston
and Greaney (1942) present evidence to indicate that it is a weak para-
site. Snyder and Hansen (1945) would include it under F. roseum.
This narrow-spored variant or form, or what-have-you, apparently
occurs in California on the common small grains, possibly in the coastal
area. The fungus in its "wild" state is certainly readily recognized by
its narrow macrospores.
Range: General.
References: Appel (1924); Bennett (1928,6); Dickson, Johann, and Wine-
land (1921); Gordon (1933); Gorlenko (1936); Johnston and Greaney
(1942); Krampe (1926); Snyder and Hansen (1945); Wollenweber and
Reinking (1935, Fig. 2, p. 52).
Fusarium culmorum (W. G. Sm.) Sacc.—Root Rot, Head Mold, Scab

Mycelium loosely cottony, carmine red with buff or off-white my-
celium scattered through it; sporodochial masses golden to ochre, later
darker, conidia thick, curved, bluntly pointed apex, slightly more taper-
ing at base with prominent foot, often 5-septate but 3- to 9-septate
spores are present, mostly 30-60 X 4.5-6.5 /J,.
On: Agropyron cristatum (L.) Gaertn., Oreg., Wash.—A. desertorum
(Fisch.) Schult., Oreg.—A. elongatum (Host.) Beauv. X Triticum aestivum
L., Wash.—A. inerme (Scribn. and Sm.) Rydb., Ida., Oreg., Wash.—A.
sibiricum (Willd.) Beauv., .N. Dak., Oreg.—A. 'smithii Rydb., Minn.—A.
trachycaulum (Lk.) Malte, Manit.—Agrostis palustris Huds., Minn.,
Oreg., Wash.—A. tenuis Sibth., Minn., Oreg., Wash.—Avena hyzantina
K. Koch, Calif., Ida., Mont., Oreg., Wash.—A. fatua L., Ida., Oreg., Wash.
—A. sativa L., Alta., B. C, Cahf., Ida., 111., Ind., Manit., Mich., Mont.,
Ohio, Oreg., Sask., Wash., Wise.—Bromus carinatvs Hook, and Arn., Ida.
—B. inermis Leyss, Ida.—B. mollis L., Ida.—B. tectorum L., Ida., Oreg.,
/I
Wash.—Dactylis glbmerata L., Wash.—Echinochloa crusgalli (L.) Beauv.,
Minn., N. Dak., S. Dak.—E. crasgdK-(L.) Beauv. var. frumentacea
(Roxb.) Wight, N. Bak.—Elymus condensatus PresI, Ida.—Festuca rubra
var. commutata Gaud., Wash.—Hordeum distichdn-{L., Alta., Calif., Oreg.,
Sask., Wash.—H. jubatum L., Wash.—JE?. mlgareiL., Alta., B. C, Calif.,
Ida., Iowa, Manit., Minn., Mo., Nebr., N. B., N. S.j Ont., Oreg., P. E. Is.,
Sask., S. Dak., Wash.—Lolium multiflorum. Lam., Oreg., Wash.—L.
perenne L., Oreg.—Poa annua L., Oreg., Wash.—P. juncijolia Scribn.,-
Oreg.—P. pratensis L., Wash.—Secale cereale L., Ark., Calif., Conn.,
Manit., Md., N. J., Oreg., Penn., P. E. Is., Sask., Va., Wash.—Tn'ticMW
aestivum L., Alta., Calif., Conn., Ida., Manit., Minn., Mont., N. B., Nebr.,
N. Dak., N. S., Oreg., P. E. Is., Que., S. Dak., Wash., .general in the
Middle West and eastward—T. dicoccum Schrank, Oreg., Wash.—T.
durum Desf., Manit., Oreg., Sask., Wash.—T. monococcum L., Wash.—
T. spelta L., Oreg., Wash.—Zea mays L., Calif., Oreg.
Fusarium culmorum causes pre-emergence killing, stunting, and
brown root rot of cereals and grasses. The bases of some plants, such
as oats, are sometimes covered with pint/spore masses. It may cause
pink-head mold or scab, and sometimes develops a small red-bordered
sheath and leaf spot on wheat in the early season, as noted in the
Palouse area of Washington. While the fungus is scarce in the northern
Great Plains, it is very common from central Montana westward. In
the coastal region, it and crown rust {Puccinia coronata Corda) com-
bine to make the growing of oats a somewhat hazardous commercial
venture. The fungus is also important east of the Great Plains. In the
Pacific Northwest it is an associate of Cercosporella herpotrichoides
and of Helminthosporium sativum (dry-land root rot). Ledingham
(1942) found that F. culmorum was not inhibited by H. sativum but
that the latter was inhibited by F. culmorum. There is increasing evi-
dence that the various components of common root rot are not entirely
compatible. I
Fusarium culmorum root rot is one of the causes of red leaf in oats
in Oregon and Washington (Sprague, 1939'b). This symptom is par-
ticularly apparent in the Willamette Valley and -in_^the vicinity of
Moro, Oregon. Inoculation trials show that isolates of tKi^fungus from
North Dakota and Oregon are destructive parasites, far mo're parasitic
than F. acuminatum.
The name F. culmorum has been a catch-all for any pink- or
carmine-tinted fungus in the United States. Very often F. acuminatum
has been called F. culmorum, and the name applied to fungi which are
assignable to F. equiseti and F. oxysporum. This accounts for the
tendency to consider F. culmorum as a quasi-parasite of slight im-
portance in some regions. Quite to the contrary, it is in many areas
essentially as parasitic as F. graminearum, and-where it occurs in
quantity it is to be reckoned with as a maôr^ problem, as it is in
Canada and in the Pacific Northwest. However, in parts of Canada
and. in California some isolates of F. culmorum are said to be weakly
parasitic. '
Range: United States, Argentina, Canada, England, Germany, Kenya,
Netherlands, Norway, Peru, Switzerland, Uganda, U.S.S.R.
References: Anliker (1935); Appel (1924); Baltzer (1930); Bennett
(1928,6); Blair (1936); Broadfoot (1933,6; 1934,6); Doyer (1921);
Foster and Henry (1937); Greaney (1938); Krampe (1926); Leding-
ham (1942); Machacek (1936); Machacek and Greaney (1936); Mar-
chionatto (1935); Rose (1924); Sadasivan (1939); Samuel and Greaney
(1937); Sanford (1946); Schmidt and Feistritzer (1933); Sliands, H. L.
(1944); Shen (1940); Simmonds (1928, 1939,a); Slykhuis (1947);
Sprague (1944,6); Sprague and Fellows (1934); Stakman, L. (1923);
Tyner (1940, 1941); Vosbein (1935); Walker (1941).
Fusarium equiseti (Corda) Sacc.—Secondary Root Rot

F. scirpi Lamb, and Fautr.
Mycelium peach colored when young, becoming buff or light brown,
rarely with scattering of rosy mycelium, cottony, microconidia scat-
tered through mycelium, macroconidia in ochre-colored sporodochia,
spores sickle-shaped, definitely narrowed at the apical cell, 0-8, mostly
5-celled in the macroconidia, mostly 35-55 X 2.5-5.0 fi.
On: Saprophytic in virtually all cereals and grasses in the northern Great
' Plains and adjacent areas, mildly parasitic on Bouteloua gracilis (H. B. K.)
Lagerh. in Mont., Nebr., N. Dak, S. Dak., and on Hordeum vulgare L.
and Triticum aestivum L. in Calif, and Manit.
F. equiseti represents about 50 per cent of all Fusarium spp. iso-
lated from roots of Gramineae in April to June in the northern Great
Plains and is exceeded in numbers during the summer only by F.
oxysporum.
It first produces a peach color or apricot tinted, cottony colony that
later turns buff or medium brown. Microspores and later macrospores
are produced in considerable numbers when grown in north light on
potato-dextrose agar. Putty-colored masses of macrospores, often with-
out pseudopionnates, are formed later. All our pathogenicity trials at
Mandan, North Dakota, indicated that local isolates were nonparasitic
on cereals and some grasses. Mild seed rot apparently developed on
Bouteloua gracilis (H.B.K.) Lag. during germination of seedlings
(Sprague, 1944,6). Johnston and Greaney (1942) found that F. equiseti
was moderately parasitic on older cereal plants in Manitoba. F. equi-
seti is considered a weak parasite in California. Bennett (1932), how-
ever, found that English strains of Fusarium scirpi var. pallens F. T.
Bennett were mildly parasitic on wheat and barley. All evidence, there-
fore, indicates that F. ,equiseti is of slight economic importance as a
parasite of Gramineae.
Range: General in the north temperate zone, at least.
References: Anliker (1935); Bennett (1932; 1935,6); Gordon and Sprague
(1941); Johnston and Greaney (1942); Sprague (1943,a; 1944,6).
l\
Fusarium graminearum Schw.-rzScab, Seedling Blight
Mycelium carmine red, loosely cottony to somewhat cobwebby,
conidia in pionnotes, rarely in sporodochia, ochr^ to bright orange in
mass, conidia longer than F. culmorum, oxhorn-shaped,: commonly 3-
to 5-septate but ranging from 1- to 9-septate, mostly 40-80 X 4.0-6.5 /*.;
chlamydospores scarce.
Perithecial stage (see Journ. Myc. 3:2.1887): Perithecia gregarious,
confluent-caespitose and concrescent, coriaceo-membranaceous, verru-
cose, at length flaccid, plicate ovoid, subcontracted at the base, bluish,
200-370 X 170-220 jn, papillate; asci oblong-lanceolate, acuminate
above, contracted at the base into a short, thick stipe, 8-spored, 60-
76 X 10-12 ix; sporidia uniseriate or subseriate, fusiform, curved or
straight, subacute, 3-septate, but scarcely constricted, nearly hyaline,
18-24 X 4-5 /*. Ellis and Everhart had difficulty in separating this
species {G. saubinetii) from G. pulicharis Er.
On: Agropyron cristatum (L.) Gaertn., N., Dak.—A. subsecundum (Lk.)
Hitchc, N. Dak.—Agrostis sp. (P. I. 131,533), N. Dak.—Avena sativa L.,
111., Ind., Iowa, Mich., N. B., N. Dak., Ohio, Okla., P. E. Is., Tex., Wise—
Bromus tectorum L., S. Dak.—Cenchrus echinatus L., Fla.-—Cortaderia
selloana (Schult.) Aschers. and Graebn., Calif.—Echinochloa criisgalli
(L.) Beauv., Minn., N. Dak., S. Dak.—Hordeum distichon L., same
range as H. vidgare—H. nodosum L., S. Dak.—H. vulgare L., Calif., Ill,
Ind., Iowa, Kans., Ky., Manit., Md., Mich., Minn., Mo., Nebr., N. Y.,
N. Car., N. Dak., N. S., Ohio, Okla., Penn., P. E. Is., Que., S. Car.,
S. Dak., Va., W. Va.—Lolium perenne L., Pa.—Poa annua L., Wise.—
P. pratensis L., Wise, Wyo.—Secale cereale L., Calif., Iowa, Md., Minn.,
Mo., N. Y., N. Car., Okla., Penn., Ya.—Setaria lutescens (Weigel) F. T.
Hubb, Ind., Ill, Iowa, Wise,—Sorghum halepense (L.) Pers., Ga.—
Triticum aestivum L., Alaska, Alta., Calif.,) Ga., 111., Ind., Iowa, Kans.,
Ky., Manit., Md., Mieh., Minn., Mo., Nebr., N. B., N. Y., N. Car.,
N. Dak., Ohio, Okla., Penn., P. E. Is., Que.,; S. Car., S. Dak., Tenn., Va.,
Wise.—T. dicoccum Schrank, Que., S. Dak.—T. durum Desf., Manit.,
Minn., N. Dak., S. Dak.—T. spelta L., Minn.—Zea -mauslj., Ark., Conn.,
Del., Ill; Ind., Iowa, Ky., Md., Mass., Minn., Mo., NebriT-N-.^J., N. Y.,
N. Car., N. Dak., Ohio, Okla., Ont., Penn., S. Dak., Wise. '_
This well-known cause of scab in cereals has been the subject of

intensive study in the Middle West. An outline of recent studies is
givenby Dickson (1939, 1947). /
F. graminearum is especially prevalent in southern Minnesota
(Hanson, 1939; 1944,b) and is important westward into eastern South
Dakota and the Red River Valley in extreme eastern, North Dakota on
spring wheat and barley. I t is isolated only occasionall/ from roots of
grasses in North Dakota west of the region .wiiere,it occurs on cereals.
It occurs in California as part of the F. roseum complex. Mourasch-
kinski (1924) reported that it was rare in Siberia. Conners (1942) in-
dicated that it was. rare in Manitoba. Palm (1932) found that the
fungus was important as a cause of seedling blight in Guatemala but
not as a cob rot.
Fetch (1936) found that Gibberella saubinetii (Mont.) Sacc. was
not the same as Gibbera saubinetii Mont, (in Sylloge Crypt., p. 252,
1856). He found that Sphaeria cyanogena Desm. was earlier and the
same as Montague's fungus. He proposed Gibberella zeae (Schw.)
Fetch.
Range: United States, Argentina, Australia, Belgium, Canada, China, Eire,
England, Finland, Germany, Guatemala, India, Japan, Kenya, Nether-
lands, S. Africa, Tanganyika.
References: Adams (1921); Anderson (1948,a,6); Appel (1924); Bennett
(1930); Blanchard and Carrera (1933); Christensen and Kernkamp
(1936); Christensen and Stakman (1927); Conners (1942); Dickson
(1923; 1939; 1947); Dickson et al. (1923); Dickson, Johann, and
Wineland (1921); Doyer (1921); Dufrenoy and Fremont (1931)
Eide (1935); Goddard (1939); Gorter (1943); Hanson (1939; 1944,6)
Hayes, Johnson, and Stakman (1933); Hemmi, Seto, and Ikeya (1931)
Henry (1931,a); Ho and Melhus (1940); Hopkins (1922); Hoyman
(1941); Ikeya (1933); Immer and Christensen (1943); Ireland (1928);
Janova (1936); Johnson and Dickson (1921); Kasai (1923); Koehler,
Dickson, and Holbert (1924); Mclndoe (1931); McKay (1943); McKay
and Loughnane (1945); Maclnnes (1922); Maelnnes and Fogelman
(1923):; Manns and Adams (1921,6); Manns and Phillips (1924); Mar-
chionetto (1935); Melhus (1944); Falm (1932); Pearson (1931); Fetch
' (1936); Scott (1927); Senn (1932); Shands, H. L. (1944); Shands,
R. G. (1937, 1939); Sprague (1944,6; 1946,e); Tanja (1933); Taylor and
Maher (1931); Titus and Godfrey (1934); Tu (1929; 1930); UUstrup
(1935); Young, H. C, Jr. (1943).
Fusarium moniliforme Sheldon—Secondary Root Rot

Ferithecial stage is Gibberella fujikuroi (Saw.) Wr.
or G. moniliformis (Sh.) Wineland, the former
preferred by Wollenweber and Reinkihg (1935).
See Nisikado (1931). See also G. fujikuroi (Saw.)
Ito in Ito and Kimura (1931).
Mycelium white, cottony, sometimes tinted with gold or rose, micro-
conidia produced in chains, except in var. subglutinans in which spores
are borne singly or in false heads, 5-12 X 2-3 fi, ellipticaLto ovate,
mostly aseptate, macroconidia slightly curved, stiff, sporodochia or
pionnotes later produce bright masses, tinted isabelline to milky orange,
1- to 5-septate, 3-septate spores dominating, mostly 15-60 X 2.4-4.9 /j.,
3-septate averaging 40 'X 3.8 /i.
Perithecia superficial on corn stalks, globose, brown, 270-350 X 240-
300 (JL, asci long, narrow, subclavate 75-100 X 10-16 j«., 4, 6, or 8 asco-
spores which are straight, constricted at the septa, short elliptical,
mostly 1- (rarely 2-, 3-) septate, 12-17 X 4.5-7.0 /t (1-septate spores).
Var. subglutinans differs from the species in that the 8-spore asci
are more common than the 4- to 6-spore~asci-ahd the 2- to 3-septate
ascospores are fewer in number (Edwards, 1935). •
1
On: Agropyron sibiricum (Willd.) Beauv., N. Dak.—A. smithii Rydb.,
Nebr.—Avena sativa L., N. Dak.—Calamovilfa lorigifolia (Hook.) Scribn.,
Nebr.—Festuca elatior L., Wash.—Hordeum vulgare L., Minn., N. Dak.—•
Oryza sativa L., Tex.-—Oryzopsis hymenoides (Roem. and Schult.) Rick.,
N. Dak.—0. micrantha (Trin. and Rupr.) Thurb., N. Dak.—Panicum
miliaceum L., N. Dak., Wash.—P. perlongum Nash, Nebr.—Saccharum
ojficinarum L., Alabama, Hawaii, La., Miss.—Setaria lutescens (Weigel)
F. T. Hubb, Minn., N. Dak.—S. viridis (L.) Beauv., N. Dak.—Sorghum
vulgare Pers., Calif., Kans., Nebr., N. Dak., Tex.—S. vulgare var.
sudanense (Piper) Hitchc, N. Dak.—Triticum aestivum L., N. Y.—T.
durum Desf., N. Dak,—Zea mays L., Alabama, Ariz., Ark., Calif., Colo.,
Conn., Del., Fla., Ga., 111., Ind., Iowa, Kans., Ky., La., Manit., Md.,
Mass., Mich., Minn., Miss., Mo., Nebr., N., Y., N. Car., N. Dak., Ohio,
Okla., Ont., P. E. Is., Que., S. Car., S. Dak'., Tenn., Tex., Va., Wash., W.
Va., Wise, Wyo. '
Top rot or "pokkah-boeng" [F. monilijorm,e) as it is known in Java
(Bolle, 1927, 1928, 1934) is a common disease of sugar cane in the
tropics. Priode reported pokkah-boen^disease of maize and sorghum in
Cuba (1933). Top rot (undetermined, possibly bacterial) is reported by
Altstatt (1926) from Texas. Hemmi and Seto (1928) reported that
the bakanae disease was featured by development of abnormally long
and slender, finally pale green, rice seedlings. Kurosawa (cf. R.A.M.
7:668) had reported earlier that the abnormal seedling growth was cor-
related with toxic excretions by the causal organism. Hemmi and Seto
(1926) confirmed these results by obtaining Elongation in plants
watered with extract from pure cultures of Fi moniliforme. The bakanae
disease of rice is featured by tall, slender, yellowish-green plants accom-
panied by basal culm rot, pink seed, and resulting seedling infection
from diseased seed (Ito and Kimura, 1931).!Kreibohn (1947) discussed
control of the disease, i;ecently discovered in TucumaB-,^^gentina. Seto
(1932) and Kurosawa (1931, cf. also Trans. Nat. Hist. Soc."~Formosa,
p. 87, 1926; see R.A.M. 7:668) found that the abnormal lengthening of
diseased seedlings could be induced by liquor extract from G. monili-
formis but only between 20° C.-35° C. Shimada (1932) found that
wheat, barley, corn, azuki beans, and soybeaijs grew abnorrnally tall
when watered with an alcohol extract from G. 'moniliformis.
Nisikado and Matsumoto (1933,b) working with 66 strains of G.
fujikuroi from rice and 5 strains of G. moniliformis'horn wheat, using
corn as a test plant, noted that the isolates frorg^rice caused overgro^vth
while those from wheat did not. As based'^n a-review available
\ (R.A.M. 12:589-590, see Nisikado and Matsumoto, 1933,a) they dis-
•tinguished the two as separate entities. Kurosawa (1932) further en-
larged his studies on^ this phenomenon, reporting that potassium is the
essential element in the production of the elongating substance. He
reported on its properties.
In 1933 Seto published several papers on the bakanae disease. In
one paper (1933,a) he philosophized on the reason why some plants
grew tall while others were stunted. In another paper (1933,6) he re-
ported that the bakanae disease was confined to rice in moist ground;
rice on dry ground when invaded was stunted. In still another paper
Seto (1933,c) reported that while the fungus grew best at 25° C , the
temperature most suitable for seedling growth and infection was 35° C.
"Bakane" seedlings developed to some extent at 25° C. but not at all
at 20° C. At 40° C. the seedlings were retarded and the development
of diseased ones was suppressed. The effect on the disease would thus
appear to be due to the action of the soil temperature on the seedlings
rather than on the fungus.
While this is a common mold on corn ears in the South and Middle
West, it is somewhat less abundant in the northern Great Plains,
although Edwards (1940,a) found it to be common on corn seed from
Nebraska, and Hoppe (1940) reported it as abundant even west to
Colorado. Boyle (1944) reported that Ray isolated it from corn stalks
from~Pasco, Washington. This species has been isolated by the writer
from the roots of corn, sorghums, millets, and a number of grasses
during the summer months in North Dakota and adjacent areas. Our
inoculation results indicate that it is a weak seedling parasite on a few
small-seeded grasses such as blue grama.
Recently Edwards (1941,a), working with F. moniliforme and F.
moniliforme var. subglutinans Wr. and Reinking, found that, because
the corn seed contained sufficient nutrient for independent seedling
growth for 10 to 12 days, soil fertility did not influence the action of
the parasite on the plant during this critical period.
Range: United States, Africa, Argentina, Asia, Australia, Barbados, Brazil,
British Guiana, Burma, Cameroons, Canada, China, Colombia, Costa
Rica, Cuba, Dutch East Indies, England, Formosa, Germany, Gold
Coast, Guatemala, Hawaii, Honduras, Hungary, India, Indochina, Italy,
Jamaica, Japan, Java, Kenya, Manchuria, Mexico, New Zealand, Peru,
Philippines, Puerto Rico, Reunion Is., Spain, Sumatra, Switzerland,
Svria, Tanganyika, Trinidad, Turkey, U.S.S.R.
References: BoUe (1927, 1928, 1934); Boyle (1944); Butler (1947);
Dickson (1940); Edwards (1935; 1940,a,6; 1941,a,&); Hemmi (1937);
Hemmi and Seto (1928); Hemmi, Seto, and Ikeya (1931); Ho (1941);
Hoppe (1940); Ireland (1928); Ito and Kimura (1931); Koehler (1942);
Koehler and Dungan (-1940); Koehler and Holbert (1938); Kreibohm
(1947); Kurosawa ,,(1931; 1932); Leonian (1932); Limber (1927);
McLaughlin (1946); Manns and Adams (1921,6); Manns and Phillips
(1924); Marchionatto (1935); Neil and Brien (1935); Nisikado (1931);
Nisikado and Matsumoto (1933); Porter (1927); Priode (1933); Seto
(1928; 1932; 1933,a,6,c); Sheldon (1904); Shimada (1932); Sprague
1944,6); Tochinai and Ishizuka (1934); Ullstrup (1936; 1943); Vorhees
344 DISEASES OF CEREALS AND GRASSES •
„, I
(1933,o; 1934,0); WoUenweber and Reinking (1925; 1935); Wineland
(1924); Yabuta and Hayashi (1939; 1940). •> ^ \
Fusarium nivale (Fr.) Ces.—Snow Mold .

Perithecial stage is Calonectria graminicola (Berk, andlir.) Wr.
Mycelium somewhat cobwebby or stringy when young, faintly pale
salmon or flesh color to pale rose, producing conidia loosely in the
mycelium or in orange to paler colored spore masses; most of the
spores in younger cultures are nonseptate to 3-septate, often without
an evident heel, 0-septate spores averaging 9.6 X 2.4 ju,, measuring
8-12 X 2.0-2.8 [i; 1-septate are 13-18 X 2.4-3.0 /t, 3-septate are 19-
27 X 2.8-3.8 n, and 4- to 7-septate, 19-30 X 2.5-4.0 /^.
Perithecia attached to a wrinkled stromatic substrate, biscuit shaped
or oval in outline, 80-300 fi., mostly 125-200 y. diam., plectenchymator.s.
Spores in one or sometimes 2 rows, 8 in each' elongate ascus, spindle-
form, straight or curved, smooth, 1- to 3-septate, 15.2 X 3.2, mostly
13-18 X 3.0-3.4 f, (10-20 X 2-4 /.). /
/
On: Agropyron cristatum (L.) Gaertn., Ida., Wash.—A. dasystachyum
(Hook.) Scribn., Wash.—A. desertoncfn (Fisch.) Schult., Wash.—A.
elongatum, (Host.) Beauv., Wash.—A. elongatum X Triticum aestivum
L., Wash.—A. inerme (Scribn. and Sm.) Rydb., Oreg., Wash.—A. inter-
medium (Host.) Beauv., Ida., Wash.—A. michnoi Roshev, Wash.—A.
repens (L.) Beauv., Ida., Wash.—A. ripariwn Scribn. and Sm., Wyo.—•
A. semicostatum (Steud.) Nees, Wash,—A. sibiricum (Willd.) Beauv.,
Ida., Mont., Oreg., Wash.—A. smithii Rydb., Wash.—A. spicatum
(Pursh) Scribn. and Sm., Oreg., Wash.—A. subsecundum (Lk.) Hitchc,
Ida., Wash.—A. trachycaidum (Lk.) Malte, Ida,, Wash.—A. trichophorum
(Lk.) Richt., Ida., Wash.—Agrostis alba L., Oreg., Wash.—A. canina L.,
Mich., Minn., Oreg.—A. exarata Trin., Oreg.—JA. palustris Huds., B. C,
Mich., Minn., Ont., Oreg., Wash.—A. tenuis Sibth., Mich., Minn., Ont.,
Oreg., Wash.—Avena byzantina K. Koch, N.'J.—A. sativa L., N. J.—
Bromus carinatus Hook, and Arn., Ida., Wash.^—B. commutatus Schrad.,
Oreg., Wash.—B. erectus Huds., Wash.—B. inermis Leyss7~'Ida.,^Wash.—
B. japonicus Thunb., Wash,—B. mollis L., Ida., Wash.—B. purgans L.,
Wash., in plots—B. racemosus L., Wash.—B. rigidus Roth, Oreg., Wash.
-^B. se.calinus L., Wash.—B. tectorum L., Ida., Oreg., Wash.—Calama-
grostis breweri Thurb., Cahf.—C. canadensis (Michx.) Beauv., Minn,—
Dactylis glomerata L., Ida., Wash.—Elymus canadensis L., Wash.—E.
glaucus Buckl., Wash.—E. macounii Vasey (hybrid). Wash.—E. sibiricus
L., Wash.—Festuca dertonensis (All.) Aschers. and Graebn., Oreg., Wash.
—F. elatior L., Ida., Wash.—F. elatior var. arundinacea (Schreb.) Wimm.,
Ida.-—F. idahoensis Elmer, Ida., Oreg., Wash.—F. megalura.'Nntt., Oreg,,
Wash.—F. myuros L., Oreg.—F. ovina L., Ida.—Fy.~rM.bra L., Alta., Ida.,
Mich., Minn,, N. Y., Oreg., Wash., Wise.—Fr^rjibra var. commutata
Gaud., Ida., Oreg., Wash,—Holcus lanatus L,, Mich., Minn., Oreg.,
Wash., W'isc.—Hordeum brevisubulatum (Trin.) Lk., Wash.—H. bulbosum
L., Wa^h,—R. jubatfim L., Wash.—H. murinum L., Oreg., Wash,—H.
nodosum L., Oreg., Wash.—H. vulgare L., Minn., Oreg., Wash.—Hystrix
patula Moench, Wash., in plots—Koeleria cristata (L.) Pers., Ida.
Oreg., Wash.—Lolium multiflorum Lam., Oreg., Wash.—L. perenne L.
Oreg., Wash.—Oryzopsis hymenoides (Roem. and Schult.) Ricker, Calif.
—Phleum pratense L., Wash.—Poa ampla Merr., Ida., Oreg., Wash.—
P. annua L., Ida., Oreg., Wash.—P. arctica R. Br., Wyo.—P. bulbosa L.,
Ida., Wash.—P. cusickii Vasey, Oreg.—P. leptocoma Trin., Oreg.—P.
palustris L., Oreg., Wash.—P. pratensis L., Ida., Mich., Minn., Oreg.,
Wash.—P. secunda Presl, Oreg., Wash.—Secale cereale L., Minn., N. Dak.,
Wash.—S. cereale x S. montanum, Wash.—Sitanion hystrix (Nutt.) J. G.
Sm., Calif., Wash.—S. jubatum J. G. Sm., Wash.—Stipa columbiana
Macoun, Calif.—iS. columbiana var. nelsonii (Scribn.) Hitchc, Wash.-—
S. comata Trin. and Rupr., Calif., Wash.—S. lemmonii (Vasey) Scribn.,
Wash.—Stiporyzopsis bloomeri (Boland) Johnson {Oryzopsis hymenoides
X Stipa occidentalis), Calif.—Triticum aestivum L., Calif., Ida., Oreg.,
Wash.
The pink dead leaves of plants affected with pink snow mold are
conspicuous just after the snow leaves the winter wheat in eastern
Washington during certain years. The characteristic spores are pro-
duced in great quantities. Most of them are 1-septate, small, more or
less heelless. Sometimes red-bordered, fawn-colored spots are formed
on new foliage in early spring. After growth starts in the spring, how-
ever, the mold rapidly disappears. Pink snow mold is associated, with
Typhula spp. in parts of Oregon and Idaho in areas where snow lies
deep. No doubt pink snow mold is the cause of late winter killing of
cereals in the Palouse, at times, which has been indiscriminately at-
tributed solely to low temperature. Blodgett (1946) has indicated this
point in his discussion of winter injury in the eastern counties of
Idaho. In the spring of 1948 snow mold {F. nivale) was prevalent in
the Palouse region of Washington and adjacent Idaho on winter wheat
and many species of grasses. I t formed great mats of dead pink leaves.
Some plants were killed outright and others severely set back although
most of the grasses finally recovered after .warm weather set in
(Sprague and Meiners, 1948).
In the humid parts of Oregon and Washington pink snow mold is
important on lawns in the winter, causing dead snow-mold areas. Snow-
mold injury in lawns occurs in Minnesota and eastward to the At-
lantic. Possibly some of the injury formerly attributed to this fungus
in the eastern United States is due to Typhula (Wernham, 1941).
Pink snow mold occurs on grasses in the higher mountain country
from Canada south into the eastern part of California (Sprague,
Fischer, and Meiners, 1948)...Haenseler (1948) reported it on winter
oats in New Jersey on Wintok and Traveler oats but not on Forkadeer,
Lee, Pioneer, Stanton, Winter Turf, and C.I. 4316.
Eleneff (1926) emphasizes the importance of inclement weather,
poor drainage, weedy, poorly-tilled fields as factors predisposing fall-
seeded plants to attack by F. nivale. Besides this organism Eleneff says
that Sderotinia graminearum Eleneff is the cause of killing of winter
/I . .
wheat and Eye in north and central Russia. The Sclerotinia attacks
many grasses. — ^ - ,
Bennett (1933,t>) controlled F.nivale by regular application to golf
greens of a combination of Bordeaux mixture and Malachite green at
the rate of 3 ^ gallons per 150 sq. yards. He applied the liquid twice
weekly in April and once a week from May to September. Malachite
green was inhibitory to the fungus at a strength of 1 to 20,000.
In Europe when pink mold is said to be seed-borne, seed treatment
is practiced [cf. Lindfors (1924); Esmarch (1924)]. While pink snow
mold is common in northern Europe it was reported as rare in Siberia
by Mouraschkinski (1924), but he mentioned that he gave winter
grains only cursory examination. F. nivale is not common in North
Dakota but we have seen it at Mandan on winter rye in very early
spring and it was isolated occasionally from necrotic perennial grasses
in the Northern Great Plains.
Roussakov (1925) believed that smothering due to shortage of
oxygen aided destruction of the plants by pink mold.
In Austria, Pichler (1933) stated seed -treatment was not always
effective because of soil infection which >vas an important factor in
Austria. He recommended good clean plump seed, properly treated, as
important. Locally adapted hardy varieties and varieties deficient in
dextrose were needed. Late sowing, relatively wide spacing, shallow
cultivation, thorough tillage, and good drainage are important. Potash
and phosphoric acid should be applied in the autumn but nitrogen
should not be given (unless seriously deficient) until after the snow i
melts in the spring. Where the snow persists it should be broken up
by driving over the field or making holes with sticks, and the melting
accelerated by strewing ashes. In 1943 his suggesticins were similar.
Range: United States, Austria, Belgium, Canada,! Czechoslovakia, Denmark,
Finland, France, Germany, Great Britain, Hungary, Netherlands, Nor-
way, Poland, Romania, Sweden, Switzerland, U.S.S.R., including Siberia.
References: Andren (1939; 1940); Anliker (11935); Atanasoff (1923);
Bennett (1933,a,6); Blodgett (1946); Broadfoot (1936r-lfl38); Dahl
(1934); Dennis (1943); Doyer (1921); Ekstrand (1938,a); Elener(1926);
Fries (1825); Hiltner and Ihssen (1911); Krampe (1926) ;-Lindfors
(1920; 1924); Markevich (1939); Monteith and Dahl (1932); Nilsson
(1940); Peuser (1931); Pilcher (1933; 1943; 1946; 1947); Pokorny
(1865); Roussakov (1925); Scliaffnit, E. (1913); Schaffnit and Volk
(1926); Snyder and Hansen (1945); Sprague (194'2,a,6; 1946,e; 1947,6);
Sprague and Meiners (1948); Tyson (1936,6); Wernham (1941); Wol-
lenweber and Reinking (1935).
/
Fusarium oxysporum (Schl.) em. Snyder and/Hansen( Secondary
Root Rot '' - "
Mycelium white with lavender, purple, to nearly sepia, rarely rosy
colors in \y^ substratum, microspores scattered, numerous, oft'en giving
powdery appearance to the cultures, not in chains, aseptate, 8-10 X 2.1-
3.0 /x, macrospores infrequent in cultures, somewhat stiffly oxhorn
shaped, not large, mostly 3- to 5-septate, 20-50 X 3.0-4.5 ft. in sporo-
dochia, seldom in pionnotes, chlamydospores numerous, terminal and
intercalary, globose, smooth, or roughened.
On: Saprophytic in dead pink roots of many cereals and grasses. Possibly
weakly parasitic on: Boutelous curtipendula (Miclix.) Torr., N. Dak.,
S. Dak., Wyo.—B. gracilis (H. B. K.) Lag., Mont., Nebr., N. Dak., S.
Dak., Wyo.—Bromus tectorum L., S. Dak.—Melica scabrosa Trin., N.
Dak.—Oryzopsis hymenoides (Roem. and Schult.) Rick., N. Dak.—0.
micrantha (Trin. and Rupr.) Thurb., N. Dak.-—Panicum miliaceum L.,
Minn., N. Dak., S. Dak.—Seiam lutescens (Weigel) F. T. Hubb, N. Dak.
—S. viridis (L.) Beauv., Iowa, Minn., Mont., Nebr., N. Dak., S. Dak.,
Wyo.—Sorghum vulgare Pers., N. Dak., S. Dak.—S. vulgare var. su-
danense (Piper) Hitchc, N. Dak.—Zea mays L., Iowa, Minn., Mont.,
Nebr., N. Dak., Oreg., S. Dak., Wyo.
During the summer months this species leads all others in number
of isolates obtained from grass roots. However, it is saprophytic in
most cases, giving a pink or bluish color to dead roots of maturing or
mature plants. I t is especially common on oats, wheat, and barley. In
the greenhouse under cool conditions the writer obtained moderate seed
rot in artificial inoculation trials in some grasses, but on cereals and on
most cool-temperature grasses the fungus was harmless (1944,6). John-
" ston and Greaney obtained the same results on cereals (1942). Sim-
monds and Ledingham (1937) also found that the cereals were not
attacked, except oats which was only mildly susceptible. F. oxysporum
produces a white cottony growth with characteristic lavender tints in
the substratum. Sometimes the tints are dark purple to sepia, rarely
reddish. Most of the spores formed in pure culture are microspores.
Range: General.
References: Anliker (1935); Gordon (1933); Gordon and Sprague (1941);
Johnston and Greaney (1942); Simmonds and Ledingham (1937);
Snyder and Hansen (1940).
Fusarium poae (Pk.) Wr.—Silver Top, Secondary Root Rot

F. sporotrichioides Sherb.
Mycelium cobwebby, carmine rose or buff, powdery with citron
shape, 0- to 1-septate microspores {Sporotrichum), 6 - 1 4 x 4 - 7 /x;
macrospores narrowly sickle shape, pointed at both ends, foot cell not
prominent, 3- to 5-septate, 18-36 X 3.0-4.2 /x.
On: Parasitic potentialities not great, possibly weakly parasitic on roots of
some Gramineae and known to cause seed rot in the soil in some species.
Avena byzantina K. Koch, Oreg.—A. sativa L., Ont., Oreg., Que.—
Bovteloua curtipendula (Michx.) Torr., N. Dak., S. Dak.—B. gracilis
(H. B. K.) Lag., Nebr., N. Dak., S. Dak.—Bromus inermis Leyss, Minn.,
/j
Nebr., N. Dak., S. Dak.—Dactyhs glomerata L., N.,Y., Term.—Elymus
junceus Fisch., N. Dak.—Festuca octoflora Walt., Wash.—F. rubra L.,
N. Dak., Penn.—F. rubra var. commutata ^ Ga.u&., 'Perm.—Hordeum
jubatum L., N. Dak.—ff. vulgare L., Manit., N. B., N. Dak., N. S., Ont.,
P. E. Is., Que.—Muhlenbergia racemos'a (Michx.)i B. S. P., Minn.^—
Panicum capillare L., N. Dak.—P. miliaceum L;,' N. T)ak.—Phleum
pratense L., Ind., Iowa, Minn., N. Y.—Poa palustris JL., Minn., S. Dak.,
Wyo.—P. pratensis L., Iowa, Minn., Nebr., N. Y., N. Dak., Wash., Wise.
—Setaria lutescens (Weigel) F. T. Hubb, N. Dak.—S. viridis (L.) Beauv.,
N. Dak.—Triticum aestivum L., B. C, Manit., N. B., N. Dak., N. S., Ont.,
P. E. Is., Que., S. Dak., Wash.—T. durum Desf., N. Dak.—Zea mays
L., Wyo.
Gordon (in conversation) has doubted the specificity of F. sporo-
trichioides. The writer has felt that the difference between this species
and F. poae was slight.
F. poae has the characteristic citron-shaped or pip-shaped inicro-
spores which are produced in carmine red,-cobwebby cultures. Some-
times the red color is missing but the pip-shaped spores remain to
identify the species. /
F. poae was once described as Sporotrichum poae Pk. from a "silvery
top" disease of the panicles of Poa pratensis. The heads turn white.and
die prematurely. They are covered with a pale buff growth of F. poae
which is associated in some instances with a species of mite {Pedicu-
lopsis graminum Renter). Heald (1908) found a similar association of
Fusarium and mite in bud rot of carnation, while Stewart and Hodg-
kins (1908) reported it on carnation and blue grass. The writer found
a similar blast of oat heads but no mites were present. Wollenweber
determined an Oregon isolate from this material asyF. poae (Sprague,
1937,(i). The culture on potato-dextrose agar was buff-colored and pro-
duced many pip-shaped Sporotrichum-type spores. The fungus ap-
peared to be a secondary parasite and was usually found on sterile
spikelets. " |,
In addition to the molds on cereal and grass heads, F. poae was
sometimes isolated in early spring from the roots of a number of
grasses in the northern Great Plains and Washington State:-During
the summer months of July and August F. poae is seldom found on
the roots of Gramineae but reappears as a pin"k mold or false scab on
wheats and barley, at or near harvest time. It is a/ minor component of
durum blight and appears to be saprophytic in' these instances. Its
chief importance lies in the fact that wheat is sometimes docked for
scabby grain when this grain may be only carrying the relatively harm-
less false scab organism, F. poae. /
Inoculation trials show that isolates of F.marium poae are com-
paratively weak parasites or saprophytes. A tew strains cause some
seed rot in blue grama and millets. F. poae is, however, a relatively
minor mold on Gramineae. >
Range: United States, Argentina, Canada, Italy.
References: Gordon (1933); Gordon and Sprague (1941); Heald (1908);
Marchionatto (1935); Peyronel (1926); Sprague {1937,d; 1944,6;
1946,e); Stewart and Hodgkins (1908); Wollenweber and Reinking (1935).
Fusicladium alopecuri Ell. and Ev.—^Mold

Spots gray, conidiophore tufts compact, grayish, thickly scattered
over the partly dead tips of the leaves; hyphae smoky-hyaline, con-
tinuous, subtruncate and more or less shouldered and toothed above
but otherwise nearly straight; conidia clavate-oblong, olivaceous, gran-
ular, continuous or with faint indications of a medial septum, 20-
35 X 7-10 IX.
On: Alopecurus genicvlatus L., Mo.
, This occurs on the apices of dying leaves at Columbia, Missouri
("Montana" in Saccardo Sylloge Fung. v. 10, p. 599). This species has
not been seen but the writer suspects that it may be young material of
Scolecotrichum graminis Fckl. which is common on this host. Ellis and
Everhart' (1888,b, p. 53) state that F. alopecuri is closely allied to F.
fasciculatum C. and E. but the tufts are more compact, the hyphae
thicker and straighter, and the conidia larger.
References: Ellis and Everhart (1888,6, p. 53); Maneval (1937).
Fusicladium destruens Peck—Spot

Hyphae short, 20-30 ju, fascicled, continuous to 1- to 2-septate,
colored; conidia acrogenous, continuous to finally 1-septate, subcatenu-
late, ellipsoid to oblong, colored, 7-20 X 5-7 /x.
On: Avena sativa L., N. Y.
This species also has not been seen but Peck's illustrations (PI. 3,
Figs. 19-22) strongly suggests Cladosporiuni herbarum. It was said to
occur in living foliage-of oats (Peck, 1890). He (1890, p. 30) writes as
follows: "In the affected plants the apical part of the leaf first shows
symptoms of disease. The tissues die and the color changes to rusty
red or dead brown. This change goes on till the whole leaf is involved.
Soon the minute and inconspicuous tufts of the fungus appear. In the
southern part of St. Lawrence County . . . scarcely a field of oats was
free from this disease. So prevalent was it, that the general color of
the fields was changed thereby, and it was the opinion of the owners
that their oats were 'rusting' badly."
Reference: Peck (1890).
'1
Gloeocercospora sorghi D. Bain andÊdg.—Zonate Leaf Spot,
Copper Spot " ~ |
Vegetative hyphae septate, iiyaline; branchina; mycelial pad be-
tween guard cells and above stomatal aperture; conidiophores hyaline,
septate, simple or branched, short, 5 to 10 ju,; conidia hyaline, elongate
to filiform, of variable length, the longer ones tapering, 20-195 X 1-4-
3.2 fjL, average 82.5 X 2.4 fx, borne in a slimy matrix,,salmon color in
mass; sclerotia 0.1-0.2 mm. diam., lenticular to spherical, black, oc-
curring inside the necrotic tissue of the host, abundant. •
On: Agrostis canina L., Penn., R. I.—A. palustris Huds., Penn., R. I.—A.
tenuis Sibth., Penn., R. I.—Saccharum ojficinarum L., Miss.—Sorghum
halepense (L.) Pers., La., Miss.—S. vulgare Pers., Fla., La., Miss., Tex.—S.
vulgare var. sudanense (Piper) Hitchc, La., Va.—Zea mays L., La., Miss.
Shear examined this material and suggested that it belonged in a
new genus close to Cercosporella but differing in that the spores are
borne in a slimy matrix. It was placed in the Moniliales because the
fruiting pad or sporodochium is in no sense an acervulus (Bain and
Edgerton, 1942, 1943). I examined this material a number of years ago
and also was of the opinion that it^was different from any species
which had been described on Gramineae up to that time.
Mature spots on sorghum vary from dark red to light brown, de-
pending on the host. They form large semicircular or irregular lesions
with a tendency towards zonation. These zonations start in'June and
become more definite later. Minute spherical to lenticular pads or
sclerotia appear in the necrotic areas. The conidia are borne in a
pinkish- to salmon-tinted, slimy matrix. The fungus grows readily in
pure culture. i /
Apparently the same fungus attacks bent grasses causing small
copper-tinted spots on lawns. The spots are irregular, coalescent, and
differ in hue from the bleached straw color of the otherwise similar
dollar spot [Sclerotinia^ When the leaves 'are wetr\vith dew, pink or
reddish gelatinous fungus pustules the size of pinheadsâre noted
(Wernham and Kirby, 1943). Some of the newer turf fungicides con-
trol the disease moderately well. Keil (1946) used a spray at the rate
of 10 gallons per 1,000 square feet applied oh July 26 and 31. He used
Puraturf (phenyl mercuri triethanol lactate) ,at a dilution of 1 to
10,000; Puratized 177 (phenyl amino cadmium lactate) at a dilution
of 1 to 5,000; and Zerlate (zinc dimethyl dithiocarbamate) at a dilu-
tion of 1.5 lbs. in 100 gallons. Protection lasted over a month, and in
the case of Puratized 177 the plots showed only 1 per cent disease after
58 days, whereas the plots which were untreaieSFaveraged 24 per cent
injury. The standard treatments failed to. control this disease and the
plot treated with Phygon, used at the rate of 1 lb. per 100 gallons,
had 34 "snr cent injury.
Range: United States, Tanganyika, Venezuela.
References: Bain (1941,a; 1944); Bain and Edgerton (1942, 1943); Cic-
carone (1949); Howard and Keil (1947); Lefebvre (1947); Wallace,
G. B. (1945); Wallace, G. B., and Wallace, Maud (1945); Wernham and
Kirby (1943).
Gliocladium spp.—Root Necrosis

Gliocladiuin Corda has erect septate conidiophores, penicillately
branched above, branches and branchlets septate, appressed, forming a
solitary gelatinous head; conidia unicellular, borne upon the tips of
branchlets and held together by mucilaginous substances in a dense
mass (Thorn, 1930, p. 498).
Various species of Gliocladium have been reported on the roots of
Gramineae. L. Stakman (1923) isolated Gliocladium sp. from Minne-
sota soil. The fungus caused a brown discoloration of the shortened
roots of infected plants. The irregular hyaline hyphae wound spirally
around the walls of the epidermal and cortical cells and sent haustoria
into the lumina, eventually causing the disintegration of the contents.
Young branches and root tips were readily infected. The fungus was
said to-resemble G. viride. The species is listed by Thom (1930) in the
floccose-green series and with conidia 6-7 X 3 ju.
Sanford reported G. roseuni (Lk.) Bainier on oats from Alberta
(1933) and Simmonds and Ledingham (1937) reported it from wheat
in Saskatchewan. The latter workers .found that it was slightly para-
sitic on wheat, but not on oats or barley. One trial at Mandan, North
Dakota, with an isolate of Gliocladium in unsterilized soil was negative
on all cereals and grasses tested (Sprague, 1944,6). G. roseum is dis-
cussed by Thom (1930, pp. 504-505).
Range: Scattered in the United States and Canada on Gramineae, general
on some other hosts as an associate in the rhizosphere.
References: Sanford (1933); Simmonds and Ledingham (1937); Sprague
(1944,&); Stakman, t ; (1923); Thom (1930).
Hadrotrichum lineare Peck—Mold

Conidiophores short, thick, stump-like, borne on black elongated
pads arranged in rows. Conidia acrogenous, egg shaped, elongate or
pear shaped, colored, 16-27 X 11-13 /M. _
On: Calamagrostis canadensis (Michx.) Beauv., N. Y.
There does not appear to be any good evidence that this fungus is
parasitic. The common species on grasses seems to be H. phragmitis
Fuckel. This species is discussed in Rabenhorst (Lindau, 1907, pp. 683,
684). The spores of H. phragmitis are globose. It has not been widely
reported in North America although Seymour (1929) lists it on
Phragmites. H. lineare is also reported on this host from Manitoba but
/!
this may not be correct. V. Hoehnel (1923) said that H. phragmitis, as
recognized by Saccardo, is the same "as~ Napicladium^ arundinaceum
(Cda.) Sacc. It is also said to be the same as TV. laxum Bubak and
allied to Cycloconium (v. Hoehnel, 1923).
Range: United States, possibly Canada, Germany.
References: Lindau (in Rabenhorst, 1907, v. 8, pp. 1682-684); Peck (1885,
p. 101, PL 1, Figs. 4-6).
Helicoceras cryzae Linder and Tullis—Reddish Brown Sheath Rot

Vegetative mycelium creeping, hyaline to hght fuscous, septate,
branched, 1.5-5.4 fi diam. Conidiophores subhyaline to hyaline, smooth,
simple or short-branched at the inflated apices, of varying length, 1.8-
5.4 /x thick, enlarging terminally to 5.4-7.4 /x. Conidia curved or some-
what helically coiled, light fuscous, multiseptate, constricted at the
septa, the cells longer than wide, mostly pf equal diameter, echinulate,
the basal cells abruptly rounded, 64-90 X 5.4-9.0 fi.
On: Oryza sativa L., Ark., La., Tex.
H. oryzae was isolated from kernel rot in the United States and it
causes a sheath rot of rice plants in the field (Tullis, 1936). Tullis
stated that Helicoma echinosporum Ito and Sasaki apparently is
morphologically identical with H. onjzae (Ito, 1932).
Range: United States, China, Japan. I
References: Ito (1932); Linder (1931); Tullis (1936).
Helminthosporium /
This important genus, as it occurs on Gramineae, was well pre-
sented over 25 years, ago by Drechsler (1923) although his work was
limited to the eastern part of the United States, outside most of the
vast grass range country. His work antedates the recent period of
intensive grassland agronomic research which is well summarized in the
United States Department of Agriculture Yearbook for 1948. As a side
line of recent investigations by the writer, some information has been
obtained on various members of the genus Helminthosporium to con-
vince him that the group needs a modern presentation. In this manual
he has relied heavily on Drechsler with additio'ns and information avail-
able from other sources. A considerable increase in host range is noted,
based partly on root-rot investigations. A critical study, above sug-
gested, would eliminate a goodly number of species which we have
been forced to retain and it might add a few/not recognized.
Drechsler's lengthy descriptions have-been-slightly abridged in a
few instances, but as far as space permits they have been included as
given. In the great majority of cases the descriptions of fungi which
we have dealt with in preparing this manual have been too meager.
Heltninthosporium avenae Eidam—Leaf Blotch of Oats

On coleoptile and roots of seedlings, causing longitudinal lesions
with stunting and blighting later in July or August or on second
growth at other times; first appears as minute brown flecks which
become longitudinal to elliptical light spots with anthocyanin or purple
borders with red or yellow halos beyond the outer edge of the dark
border; mycelium thin-walled, hyaline, later darker, 5-7 ju, diam.;
conidiophores appear singly or in groups; stout, cylindrical, multisep-
tate, fuliginous, 150-200 X 8-12 p.; spores olivaceous to yellow brown,
cylindrical to slightly swollen in the middle, rounded at the ends., 4- to
6-septate, 80-110 X 15-16 ju. Sometimes distorted pycnidia bearing
spermatia-like spores are formed in culture, spores 2-3 X 2 ju; peri-
thecia formed in culture have been called Pleospora avenae Schaffnit
and Rathschlag, while Pyrenophora avenae Ito and Kuribayashi was
described on straw. Pleospora avenae Schaffnit and Rathschlag: Peri-
thecia nearly spherical, black, without setae, 118-340 /x, asci sac-
shaped, 192 X 29 n, paraphyses present, 160-180 /x; spores amber yel-
low to brown, 5-8 times transversely and vertically septate, oblong,
slightly constricted at the cross walls, 33 X 14 jn.
' Pyrenophora avenae Ito and Kuribayashi: On half-rotten straw of
common oats and Avena jatua L. Perithecia at first subepidermal, then
erumpent, semiglobose when young, later flask-shaped or conical with
a short paraboloid or cylindrical ostiolar beak; the blackish brown
walls bear conidiophores and setae; perithecia 300-600 jx tall, 450-800 fx.
long diam., 350-700 fi short diam. The fasciculate, clavate, or cylindri-
cal hyaline asci are often slightly curved, rounded at the apex, shortly
stipitate at the base, 250-350-400 X 35-45 [i, containing 2-8, mostly 8,
pale yellowish or yellowish brown ellipsoidal or oval ascospores, with
3 to 6 (mostly 5) transverse septa, sometimes accompanied by 1 to 4
longitudinal septa, markedly constricted at the septa, measuring 50-
75 X 17.5-30 y..
On: Arrhenatherum elatim (L.) Beauv., CaUf.—Avena barbata Brot., Calif.

—A. byzantina K. Koch, Calif., Ga., Ky., Oreg.—A. jatua L., Oreg.—A.
sativa L., Alaska.Alta^, Ark., B. C, Calif., Conn., Fla., Ga., Ida., 111., Ind.,
Iowa, Ky., La., Me., Manit., Minn., Mc, Mont., Nebr., N. B., N. Y., N.
Dak., N. S., Ohio, Okla., Ont., Oreg., Penn., P. E. Is., Que., Sask., S. Dak.,
Wash., Wise, Wyo.—Koeleria cristata (L.) Pers., S. Dak., Wash.
The perithecial stage oi^ this species has been widely studied.
Drechsler (1923) reported a perithecial form at Madison, Wisconsin,
in 1919 and 1920 with either immature Pleospora or Pyrenophora, asso-
ciated with saprophytic conidial forms. The fungi described in Ger-
many and in Japan have been mentioned in the description above.
Dennis (1933) rejected the Japanese species but later (1935) reported
it from Scotland. Weiss also listed the conidial stage as associated with
the Japanese perithecial stage. Drechsler (1934) preferred Pyrenophora
for the perithecial forms associated with Helminthosporium, forms, but
Wehmeyer put them back in Pleospora "(1949)-. j
This fungus was originally described by Briosi and Cavara in 1889
(I fungi par delle piante colt, od utili no. 80) as H. teres f. avenae-
sativae. Drechsler (1923) also found that H. teres and H. avenae were
morphologically similar, if not identical. Denni^ mentioned that the
two species were similar (1933). Eidam described H. avenae from
Silesia (1891) and first noted that the species would not attack barley.
Ravn (1900) arrived at the same conclusion. He proposed H. avenae
(Br. and Cav.) Ravn, deriving his combination, without justification,
from H. teres f. avenae-sativa. Cooke (1889) described H. avenaceum
Curtis, Herb., a fungus found on straw in the United States (oat
straw, according to Saccardo in a discussion by Drechsler, 1923). The
conidia were cylindrical to fusoid, pale honey color, 75-85 X 15' /*.
Harkness (1885) found Helminthosporium on oats at San Francisco
in April. Atkinson reported H. avenaceum from Mississippi, collected
June 26, 1891 (1897). Apparently Drechsler hesitated to consider this
saprophytic material the same as H. avenae Eidam (1923, p. 664),
particularly as he mentioned having seen two species of Helmintho-
sporium on old oat straw. None of these seems to resemble the darker-
spored H. victoriae Meehan and Murphy.
Harvey (1895) reported a Helminthosporium on oats from Maine
and this was assigned t,o H. inconspicuum var. brittanicum. He listed
the spores as 40-80 X 15 /i, 1- to 5-septate. His illustration of an ellip-
soidal spore indicates, according to Drechsler, no close relation to either
H. inconspicuum, H. teres, or H. avenae (1923). Perhaps Harvey's
material (1895) was an aberrant form of H. sativum which .occurs on
oats, especially in marine climates. /
In summer, in the northern Great Plains, -ihe spots on oat leaves
may be poorly-defined brown areas merging into yellowish red or
orange lesions. In winter or in cool weather, in Oregon, they are more
frequently definite spots of red or purjjle anthocyanin shades sur-
rounded ;by halos of, yellow. As they become oldejj, the spots fade to
straw color in the center. Dennis (1933) reported irT-Scotland pre-
emergence killing in seedlings from seed-borne mycelia or spores. Sur-
vivors showed leaf stippling. Later spotting developed, often secondarily
in late season, particularly on second-growth oats.
H. avenae requires cool, moist growing conditions such as prevail in
the winter, early spring, and fall in the coast region of Oregon, Wash-
ington, and in parts of the British Isles. The optimum temperature for
the fungus is 20° C , the maximum, 30° C.
In Oregon, H. avenae was isolated from the bases of diseased plants
but appeared to have washed down the culm^smd developed as an asso-
ciate of Ophiobolus graminis, Fusarium culmoruin,Aphanomyces camp-
tostijlus and Pythium spp., which thrived in the highly acid soil of the
coaf' al region where the diseased condition prevailed (Sprague, 1938,a).
H. avenae is somewhat lighter gray ii),pure culture than most isolates
of H. sativum, and it tends to form bracketed growths in test tubes
which are somewhat distinct from any developed by H. sativum.
While this fungus causes a minor leaf spot in many regions, it is
very serious in Scotland and Ireland and moderately important in
western Oregon and parts of eastern Canada. Seed treatment with New
Improved Ceresan (or Geresan-M) at % ounce per bushel is very
necessary wherever the disease is serious. Varietal resistance has not
been observed in Scotland although the varieties Sovereign and Ascot
appeared to be especially susceptible. In Oregon winter-hardy oats,
such as Support or Gray Winter, are more tolerant to the disease than
less winter-hardy oats such as Red Rustproof or Kanota of the red oats
group {A. hyzantina). Wild oats {A. jatua) sometimes carries the dis-
ease. It has been found on tall oats grass and on Koeleria. Sampson
and Davies (1923) reported H. avenae on 41 varieties of spring-sown
oats in Wales. The varieties Record, Yellow Naesgaards, Leader, and
GoMfinder were the most severely affected while Algerian sterilis was
"immune."
Recently, in a survey of Helminthosporium spp. on oats in Georgia,
Miller, J .H. (1947) reported a general scorch of oats in November and
December, 1946. Most of the injury in the early stages, at least, was
due to H. avenae although H. victoriae was present and accountable
for necrosis in culms later in the season.
Range: United States, Argentina, Australia, Austria, Belgium, Canada,
China, Czechoslovakia, Denmark, England, France, Germany, India, Ire-
land, Italy, Madagascar, Netherlands, Scotland, Tanganyika, Turkey,
U.S.S.R., Wales.
References: Atkinson (1897); Cooke (1889); Dennis (1933, 1935);
Drechsler (1923); Eidam (1891); Harkness (1885); Harvey (1895);
Ito (1930); Miller (1947); Muskett (1937, 1938, 1944); O'Brien and
Dennis (1933); Rathschlag (1930); Ravn (1900); Smith and Putterill
(1932); Sprague (1938,a; 1946,e); Turner (1931); Turner and Millard
(1931).
Helminthosporium bromi Drechsler—Leaf Spot

Perithecial stage is Pyrenophora bromi (Died.) Drechsl. or Pleos-
pora trichostoma (Fr.) Ges. and de Not. (Wehmeyer, 1949).
Lesions in early spring at first minute, dark brown or black with a
yellowish halo, appearing on the first leaves, later elongating,-becoming
as much as 2 X 6 mm. in size and, through coalescence, may cover most
of a leaf surface which becomes yellow, withers, and dies. Conidio-
phores singly or in pairs, 100-150 X 7-10 /x, rarely 250 /A long, 2- to
6-septate, geniculatioris not pronounced; conidia cylindrical, hght
brown, 1- to 10-septate, 45-265 X 14-26 /*, basal cell hemi-ellipsoidal
in shape, hilum of conidia an inconspicuous scar included entirely
within the contour of the peripheral wall which is thin, germinating
from any cells, usually no more than two. Perithecia may form as early
as the latter part of July, 0.3-0.4 mm. with- an osti'olar beak extending
0.1 mm. above the surface, some sterile bristles forming around the
ostiole; asci develop the following spring or as e^rly as February or
March, completing formation in April, asci,170-215 X 40-50 /^ but en-
larging to as much as 300 X 65 /* by discharge time; ascospores dis-
tichously arranged, light brown, 45-72 X 20-30 n, • 3-sept'ate, longi-
tudinal septa when present in middle segments.
On: Bromus carinatus Hook, and Arn., Ida., Ky., N. Dak., Wash.—B. ciliatus
L., Wise, Wyo.—B. inermis Leyss, Alta., 111., Ind., Manit., Md., Minn.,
N. Y., N. Dak., Ohio, Penn., Wise—5. japonicus Thunb., Mont., S. Dak.,
Wise, Wyo.—B. pumpellianus Scribn., Alta.—B. rigidus Roth, Oreg.—
B. secalinus L., Cahf.—B. tectorum L., Wash., Wise.
This species is favored by cool weather with temperatures ranging
from 2° C. to I C C , and becomes less conspicuous as the season ad-
vances. It is very common on B. inermis, in North Dakota, particu-
larly in the Red River Valley of the North (Weniger, 1932). It occurs
also, more or less incidentally, on the roots of this grass in the northern
Great Plains and scatteringly westward. It is probably common east
of Wisconsin and Kentucky where it has been reported.
The perithecial stage, Pyrenophora hromi (or Pleospora trichos-
toma, cf. "Wehmeyer, 1949) is well known (Drechsler, 1923). Chamber-
lain and Allison (1944, 1945) recently reported that P. hromi was an
important parasite of brome in "Wisconsin during cool, wet, spring
weather. The perithecia are considered to be more important in over-
wintering the fungus than the conidial stage, as the latter does not
live over winter in the area. Varietal resistance was noted.
Range: United States, AustraUa (N. S. W.), Canacja, Germany.
References: Chamberlain and Allison (19,^14, 1945); Diedioke (1902);
Drechsler (1923, PI. 8, 9); Wehmeyer (1^49); Weniger (1932).'
Helminthosporiutri buchloes (Ell. and Ev.) Lefebvre and A. G.

Johnson—Mold "~~~—^^
H. inconspicuum (Cke. and Ell.) var. buchloes Ell.'^nd Barth.
(in Herb.)
H. inconspicuum (Cke. and Ell.) var. buchloes Ell. and Ev.
(nom. nud.)
Spots on leaves at first yellow to reddish 'yellow, later bleaching to
straw color with, dark blotches on both sides of leaf as/stromata and
conidiophores are produced. Conidiophores arising singly or in groups
of *2 to 20 or more, the larger groups being_ produced from brown
stromata between the fibrovascular bundleg^'Tonidiophores olivaceous-
brown, lighter toward apices, 2- to 9-septate, 5-8 [i diam. at basal sep-
tum, and 60-120 ju, long; conidia fuliginous, darker when mature, rarely
subhyaline when very immature, 2- to 9-septate, straight or curved,
occasionally tending to be somewhat sinuous, tapering toward the
rounded apex, basal cell rounded; with hilum inconspicuous, germinat-
ing by two polar germ tubes, 27-86 X 8-11 ja (mean 51.8 X 9.7 [i).
On: Bouteloua curtipendula (Michx.) Torr., Kans., Nebr., N. Dak., S. Dak.,
Wise, Wyo.~B. gracilis (H. B. K.) Lag., Kans., Mont., Nebr., N. Dak.,
S. Dak., Wyo.—B. hirsuta Lag., Wise.—Buchloes dactyloides (Nutt.)
Englm., Kans., Mont., Nebr., N. Dak., Okla., S. Dak., Tex., Wise.
This differs from H. inconspicuum in the very much smaller, darker
spores which are fusiform and thick-walled.
I have seen this fungus many times during the course of studies on
.root and crown diseases of cereals and grasses in the Plains region.
What may be the same fungus occurs widely on blue grama grasses and
we have added these to the host range.
This fungus was undescribed until 1949 (Lefebvre and A. G. John-
son). It was issued in 1895, as a nom. nudum (cf. Lefebvre and John-
son, 1949).
Range: United States, possibly on Bouteloua into Canada.
References: Lefebvre and A. G. Johnson (1949); Pammel, King, and
Bakke (1910); Sprague (1946,e); Wenger (1943).
Helminthosporium californicum Mackie and Paxton—Spot Blotch

This species is referred to H. sativum P.K.B. as suggested in an
'.earlier article (Sprague, 1946,e'). The difference, if any, from the last-
mentioned species is probably due to the influence of the salt-laden air,
especially fogs from the nearby ocean.
Helminthosporium carbonum IJJlstrup—Charred Ear Mold and

Leaf Spot
RACE I.—Lesions small, pale gi-een or yellowish in early infections.
Fully developed lesions show zonate pattern, dry light brown centers,
pale to purphsh-brown margins. Lesions range in size from hardly
visible spots to those 5 X 20 mm. Fruiting of fungus abundant on leaf
sheaths and husks under humid conditions. Ears infected through tips,
shanks, or directly through the husks. Diseased ears covered with black
mycelium imparting a charred appearance.
RACE II.—Lesions in early stages not unlike those caused by Race I.
Fully developed lesions elongated, irregular, and up to 3-20 mm. in
size; chocolate brown, zonate patterns less distinct; not highly special-
ized in parasitism toward, inbred lines of corn; ears more frequently
attacked than leaves, symptoms on ears as in Race I.
Conidia 25-100 X 7-18 fi (mean size 62.6 X 13.2 ^), widest in the
center and tapering slightly towards rounded ends, straight or slightly
curved, dark olivaceous brown, 2- to 12-septate (average 7) with hilum
rather inconspicuous, and germinating by two polar germ tubes. Coni-
diophores arising singly or in small groups froin' the stomata, 90-
230 X 5-7 ft, dark olivaceous brown, bearing one tojseveral conidia.
On: Zea mays L., 111., Ind., Iowa, Kv., Ohio, Penn., Tenii., Wise.
' 'I
The dark, relatively straight, stiff, "cigar-shaped" conidia of this
species distinguish it from the regularly curved, almost lunate spores
of H. maydis (1941,a; 1944).
Two races, which differ in symptoms caused, were recognized by
Ullstrup (1944). Race II is, in general, not nearly so virulent nor so
highly specialized in parasitism as Race I. Race I was originally found
on a dent corn inbred line Pr. Later two additional lines, K.G.I, and
K44, were found susceptible. H. carbonum seems to be a case, as with
H. victoriae, of an obscure species arising to prominence as a result of
man's attempt to direct nature into producing varieties and hybrids
different from those found in nature.
Ullstrup (1941,6) reported that susceptibility to Race I (under H.
maydis) of H. carbonum was inherited as a monogeneric recessive
factor. Additional studies were reported, later (Ullstrup and Brunson,
1947). Robles (1949) reported on H. "Z" which had longer spores than
H. carbonum and differed in pathogeiiicity.
References: Ullstrup (1941,a,6; 1944); Ullstrup and Brunson (1947).
Helminthosporium catenarium Drechsl.—Leaf Blight

Tips, margins, or larger areas of tissue prematurely killed, without
definite spotting or coloration; conidiophores/brown or olivaceous,
usually emerging singly or in pairs from the stomata, 60-200 X 5-8 ft,
producing the first spore at a distance of 25-60 /x from base and suc-
cessive spores at intervals of 15-30 fi, the point of attachment marked
by scars at the apices of pronounced geniculations. Conidia 1- to 10-sep-
tate, the septa sometimes associated with slight consltictions or irregu-
larities in the contour of the thin peripheral wall; subhyaline to light
yellowish, 30-200 X 14-18 ft, the shorter ones usually straight, widest
at the basal or second segment, tapering uniformly to approximately
half the maximum diameter at tip; the longer ones often perceptibly
crooked, irregular in diameter, frequently shcjiwing both a basal hilum
and an apical scar, the apical scar marking the attachment of a sec-
ondary spore and usually associated with a peculiar modification of the
distal portion of the terminal segment, consisting'in the prolongation
of the latter at a uniform diameter representing the minimum of the
sporr. Germination by production of 1 or' 2-lateral germ tubes from
basal .-egment, and a single lateral or oblique tube from terminal seg-
ment. Contour of basal end hemi-ellipsoidal, of distal end hemispheri-
cal; lillum and apical scar not protruding.
On: Cinna arundinacea L., N. Y.—Sorghum vulgare var. sudanense (Piper)
Hitchc, Va. (as reported by Lefebvre and Johnson, 1941).
On potato-dextrose agar the aerial mycelium is white or dirty yel-
lowish, present as dense erect tufts 2 to 5 mm. high at point of inocula-
tion and as small flecks scattered sparsely over the surface. Conidial
fructifications arising on the expanded terminations of hyphae consist
of a series of successively proliferated spores that may be either ses-
sile or separated by interecalary, narrower sporophoric segments
(Drechsler, 1923).
Drechsler (1923) collected drought-like lesions on Cinna in Sep-
tember, 1920, on Long Island, N. Y. The spores have some of the
appearance of H. dictyoides and to some extent of H. gramineum. The
larger ones resemble H. tritici-repentis but the latter does not shov.^
an apical hilum or apical attenuated prolongation. In addition H.
catenarium does not have the peculiar modification of the basal seg-
ment typical of H. tritici-repentis.
Atkinson (1897) reported H. turcicum on Cinna arundinacea from
Alabama but this may have been H. catenarium. These appear to be
the only reports of Helminthosporium on Cinna.
Helminthosporium cyclops Drechsl.—Leaf Spot

Causes small, dark brown spots, not abundant. Conidiophores oliva-
ceous, 3- to 7-septate, 100-250 X 7-8 fx, producing first spore from 80
to 160 /x from base and successive spores at the apices of moderately
pronounced geniculations. Conidia dark olivaceous except'in restricted
subhyaline regions at apex and base; straight or rarely slightly curved,
cylindrical or tapering gently toward the bluntly rounded ends, the
shorter ones often ellipsoid; measuring usually 45-110 X 12-17 jn, 4- to
12-septate, usually not constricted, peripheral wall thick except in the
subhyaline regions from which the two polar germ tubes are produced,
one at the apex, the other immediately surrounding the unusually large
and conspicuous hilum included within the basal contour.
On: Danthonia californica Boland, Oreg.—D. spicata (L.) Beauv., Me.,
Mass.—Poa pratensis L., Wash.—Triticum aestivum L., Wash.
This fungus produces brown spots from 0.5-2.0 mm. diam. on living
leaves of Danthonia. The fungus'resembles H. sativum and H. vagans
but Drechsler considered-//, cyclops distinct because of the more pro-
nounced attenuation of the peripheral wall at the base and apex. In
addition the basal end of the conidia of H. cyclops is distinguished by
a conspicuous hilum, the largest seen by Drechsler. The fungus was
named H. cyclops because of the one-eye aspect of this hilum.
H. cyclops produces, on potato-dextrose agar, moderately profuse
gray aerial mycelium which is interspersed with a considerable number
of subspherical, dark superficial sclerotia wliich could be undeveloped
peritheci^ (Drechsler, 1923). Compound upright conidial stick-like
structures as tall as 0.3-0.5 mm. are formed in culture in Beyerinck's
agar. These bodies are composed of hard whitfe pseudoparenchyma with
a black mottled surface, and they bear numerous | radially oriented
sporophores on the part above the somewhat expanded sterile base.
We found this fungus in abundance on Danthonia in the white,
Dayton clay pasture area east of Corvallis, Oregon. Strangely enough
we identified the same fungus on overwintered wheat seeds collected
near Pullman, Washington, by Conley Lowther. Further search by
Lowther, Goheen, and Shaw (1950) indicated that the fungus was not
uncommon on weathered seeds of wheat. The black, twisted, root-like
appendages were 1-3 mm. long. They could be mistaken for moldy
rootlets of a partially germinated seedling. However, they proved to be
fungus tissue. Lowther found that the horns produced many hyaline,
septate, radially oriented conidiophores which formed singly, or in
pairs, on forks, cigar-shaped, very dark brown conidia with a promi-
nent hilum. It was my opinion that morphologically the fungus was
virtually identical with H. cyclops. I ^had seen these ,moldy seeds in
earlier years but had never noted that they were covered with this
Helminthosporium. The fungus is not readily recognizable as such until
placed in a moist chamber and spores are produced. In pure culture it
developed the strange stick-like stromatic coremia which Drechsler
had illustrated. Mary Hafercamp of The Washington State College!
Seed Laboratory found the fungus producing horns from seeds of Poa
pratensis in 1949 (C. G. Shaw, notes).
Range: United States. i /

References: Drechsler (1923); Lowther, Goheen, and Shaw (1950, in manu-
script); Sprague (1946,e). ,'
!•
Helminthosporium cynodontis Marig.—^Mold
Spots olive to fuliginous, conidio'phores dark brown, single or in

pairs, 2- to 5-septate, 50-150 X 4-6 fx; conidia elongate, straight or
more often curved, tapering slightly towards the abruptly rounded
ends; subhyaline to fuliginous, never brown or dark olivaceous, 3- to
9-septate, nonconstricted, germinating by production of two polar germ
tubes, 27-80 X 11-14 fi.
On: Cynodon dactylon L,,' Cahf., Fla., Ga., N. Gar., .Okla., Tex., Va.—
, Eleusine indica (L.) Gaertn., Fla.—Muhlenbergia mexicnna (L.) Trin
N'\Y., Va. _ ^/^ '
Thi^ fungus is common on Cynodon in the southern United States,
especially along the seacoasts north to Washington, D. C , and San
Franciscb, Calif. It sometimes disfigures lawns but is not actively para-
i
sitic. Nattrass (1939) reported it as the cause of brown patch of
Cynodon lawns in Kenya.
H. cynodontis produces a moderate quantity of light gray, fluffy
mycelium, especially at some distance from the point at which the
culture was planted. Drechsler did not obtain spores in pure culture.
Range: United States, Kenya, northern Italy.
References: Drechsler (1923); Marignoni (1909); Nattrass (1939).
Helminthosporium dematioideum Bubak and Wrob.—Mold

On withered leaves without distinct lesions or coloration, sporo-
phores singly or in pairs, subtorulose, brown, usually short, 25-60 >,
rarely 175 fi long, 5-6 ju. wide; conidia yellowish, 2- to 6-septate, 18-
48 X 14-18 /i, cylindrical-oblong, broader in some at the rounded apex
than at the pointed base, proximal septum frequently constricted,
small dark hilum contained within basal contour, germination from
basal cell.
On: Agrostis alba L., Conn., N. Y.—A. perennans (Walt.) Tuckerm., Conn.,
N. Y.—Anthoxanthum odoratum L., D. C, N. Y., Oreg., Wash.
b e a d weathered leaves are abundantly covered with the fruiting
stalks of this species. The fungus has not been studied to any 'extent
and it is not known whether it is parasitic or not. It is not uncommon
along the seacoast of northwestern Oregon on sweet vernal grass. The
type area is Galicia (Bubak, 1916).
Range: United States, U.S.S.R.
References: Bubak (1916); Drechsler (1923).
Helminthosporium dictyoides Drechsl.—Net Blotch of Fescue

Causing a net blotch in which infected leaves show irregular brown-
ish areas marked with dark longitudinal and transverse streaks form-
ing a delicate reticulate pattern, leaves later withering progressively
from' the tips. Sporophores dark brown or olivaceous; emerging singly
or in groups of 2 to 6 from stomata or between epidermal cells; 3- to
6-septate, 70-150'><^6-8 /^ diam., spore attachment points marked by
moderately to strongly pronounced geniculations. Conidia subhyaline
and nearly colorless at first, finally yellow, typically straight, some-
what larger at base, 14-17 ji* wide, tapering uniformly and perceptibly
to apical segment, the latter in long spores frequently not exceeding
8 to 9 /x diam., in short ones usually of greater diameter; more rarely
approximately cylindrical or short ellipsoidal. Length 23-115 ft., usually
50-70 fi, 1- to 7-septate, typically 3- to 5-septate, rarely constricted at
septa except at times at basal septum; contour of basal cell hemi-
spherical, or apical end hemispherical or hemi-ellipsoidal; peripheral
wall of spore uniformly thin, and entirely including the hilum within its
contour, germinating mostly from end segments.
On: Festuca elatior L., CoBn., D. C, Ida., Iowa, Me., iMd., Mass., Mich.,
N. y., Ont., Va., Wash., Wise.—F. elatior var. ari^ndinacea (Schreb.)
Wimm., Wash.—F. occidentalis Hook., Wash.—F. pacifica Piper, Wash.—
F. rubra \a.r/commutata Gaud., Oreg.
Drechsler (1922,a) reported this species as the mosf!' important
parasite of meadow fescue (F. elatior) in the eastern United States,
where it is so common that it serves as a ready means of identification
of the host in its vegetative stage. Tall fescue {F. elatior var. arundi-
nacea) appears to be resistant. It is not common in the West—at least
it has not been found in any quantity as yet.
References: Drechsler (1922,o; 1923, pp. 677-679, PI. 11); Sprague
(1946,e).
Helminthosporium erythrospilum Drechsler—Spot

Spot on leaves straw colored with a reddish-brown border and a
water-soaked area surrounding, numerous; more or less overlapping to
form a pseudozonate to halo spot appearance of variable width, 0.5-
0.8 X 0.5-2.5 mm., sometimes more expensive; often tfie withering of
the leaves resembles drought injury. Conidiophores erect, brown or
fuliginous, simple or occasionally somewhat branched, emerging singly
or less often in pairs or groups, mostly from between epidermal cells
but sometimes from stomata, 75-275 X 6-9 ;«,, 1- to IG-septate. Conidia;
yellowish when mature, sometimes light olivaceous; straight, rarely
curved, mostly nearly cylindrical, ends abruptly rounded off, 2- to
10-septate, sometimes, but more often not, constri'cted, 25-105 X 8-16
/i, averaging 65 X 12-3 /x, basal hilum nonprotruding, germinating from
any or all cells. i
On: Agrostis alba L., Conn., D. C, Ind,, Md., Mich., N. Y., Ohio, Va., Wise.
—A. canina L., Va.—A. palustris Huds., Va.—A. perennans (Walt.)
Tuckerm., N. Y.—A. t'enuis Sibth., Va. ""^--^
During wet weather the russet-bordered lesions are readily recog-
nizable but local discoloration is less evident in dry weather, when
infected leaves often wither in a manner suggestive of drought injury.
Drechsler distinguished this species from H. gramineum, H. dictyoides,
H. siccans, and H. avenae by its smaller diameter and closer septations
of its conidia; from H. arundinis by its conidia, being abruptly rounded
at both ends, and by the position of the hilum within the contour of
the basal segment. H. arundinis is discussed by Nisikado (1928,a).
Ito (1,930) placed this species in Drechslera/Mhich is a genus, now
little lised, based on Nisikado's subgenus Cylindfo-Helminthosporium
(1928^). Kisikado divided Helminthosporium into this division and
another, Eu-Helminthosporium. The former was distinguished by
cylindrical conidia germinating from any><!ells and having Pyrenophora
FUNGI IMPBRFECTI—MONILIALES 363
ascigerous stages (P. bronii, P. tritici-repentis, P. teres). Eu-Helmintho-
sporium has fusiform conidia germinating from the poles and having
Ophiobolus ascigerous stages (but see Drechsler 1934,6 on Cochlio-
bolus).
This fungus produces submerged sclerotia in pure culture that are
more numerous and much smaller than those produced under similar
conditions by H. bromi. The aerial mycelium is rapid-growing, not
profuse, often' zonate and light gray or steel gray in color. I t resembles
that of H. siccans from Loliuni but the latter, perhaps, is less inclined
to form zonations. H. erythrospilum differs distinctly from H. dictyoides
in pure culture as the latter produces, on cornmeal agar, a more
abundant, almost white growth with tufted or somewhat columnar
mycelium.
This obscure fungus apparently is common on redtop and seaside
bent in the Middle West and central Atlantic States. Colonial bent {A.
tenuis) and A. canina appear to be less susceptible to the parasite.
References: Drechsler (1925,c; 1935).
Helminthosporium giganteum Heald and Wolf—Zonate Eye Spot

Spots numerous, yellowish or pale straw-colored, narrow brown
border,, 0.5-1 X 1-4 mm., often confluent; conidiophores brown, multi-
septate, base somewhat inflated, 200-400 X 9-12 jn; conidia cylindrical,
5-septate, pale brown, extremely large, 300-315 X 15-21 fx.. s
On: Agropyron repens L., Md., Mo., Tex., Va,—Cynodon dactylon (L.)
Pers., Fla., Mo., N. Car., Okla., Tex.—Echinochloa crusgalli (L.) Beauv.,
Mo.—Eleusine indica (L.) Gaertn., Fla., Md., Mo.—Elymus virginicus
L., Md., Penn.—Eragrostis cilianensis (All.) Lk., Md., Va.—E. pectinacea
(Michx.) Nees, Md., Va.—Leersia virginica Willd., Md.—-Muhlenbergia
mexicana (L.) Trin., Md.—M. schreberi Gmel., Md.—Panicum dandesti-
num L., Md.—P. dichotomiflorum Michx., Mo.—Pennisetum alopecuroides
(L.) Spreng., Md.—Phalaris arundinacea var. picta L., Md.—Phleum
pratense L., Md.—Poa compressa L., Md., Va.—P. pratensis L., Md.
The zonate eye spot is readily recognized and the huge spores cannot
be mistaken for any other species. I t is primarily a disease of the
southern United States but its range extends north to Maryland and
Pennsylvania. It is especially common on Bermuda grass.
References: Drechsler (1923; 1928,c; 1929,6); Heald and Wolf (1911).
Helminthosporium gramineum Rab.—^Barley Stripe

Causes a striping of leaves and death of plants, sporophores in clus-
ters of 3 to 5 (1 to 6), yellowish, 1- to 5-septate. Spores subhyaline to
yellowish brown when fully mature, straight or very slightly curved;
/I
subcylindrical, but frequently widest in the basal portion and tapering
more or less toward the apex, 1- to 7-septafe, 50-100 X 14-20 /x; periph-
eral wall thin, rarely constricted at the septa, germinating from any or
all cells.
On: Hordeum vidgare L., Alta., Ark., B. C, Calif., Colo., Conn., Ga., Ida.,
111., Ind., Iowa, Kans., Ky., Manit., Md., Mich., jilinn.. Mo., Mont.,
Nebr., N. B., N. Y., N. Dak., N. S., Ohio, Okla., Ont., Penn., P. E. Is.,
Que., Sask., S. Dak., Va., Wash., W. Va., Wise., Wyo. According to
Drechsler it occurs in almost every state in the Union where barley is
grown to any extent (1923).
The symptoms of this important disease are detailed by' Drechsler
(1923). Previous to heading time, the leaves become streaked with yel-
low bands that sometimes traverse the entire length of the leaf. The
streaks later become yellowish brown or brown and the plants die pre-
maturely. The leaves eventually become ragged or shredded. Heads
may be completely sterile and the undeveloped kernels more or less
brown.
Barley stripe is not important west of the Red River Valley, North
Dakota, except in certain areas in California and scattered spots in
western Oregon, and sometimes in thCPalouse region 'of Washington
and Idaho. It was serious in a few small fields near Rifle, Colorado, It
is more important in the Middle "West and east to the Atlantic coast.
However, in the better barley-growing areas the disease is now far less
ij:nportant than formerly because it readily yields to control by seed;
treatment with organic mercurials..
The seed infection is favored by high humidity at and following
flowering time. The disease may be carried over ,as mycelium pene-
trating the glumes into the seed coat, or it may be transmitted by
conidia on the seed, or wind-borne. Infection of the seedling occurs at
sprouting time and is favored by cold, wet soil. AVinkleman (1929)
discussed grain seedling infection methods' Fuchs (1930) used the
vacuum method to inoculate seeds. Christensen and-Graham (1934)
found that H. gramineum was polymorphic and multiracial. They-found
about 125 races showing some morphological differences -in certain
cases and many variations in cultural characters. About 20 races could
be distinguished by their relative virulence on 16 varieties of barley.
Shands, H. L. (1934) found that H. gramineum was somewhat
adaptable to varying temperature conditions. Bferley will grow at cool
temperatures (12° C.-16° C.) as will H. gramineum, but theiatter also
developed well at cooler temperatures and also at 20? C.
The following varieties have been more or less resistant to stripe:
Glahron, Spartan, Trebi, and Wisconsin barbleSfT, -
Pyrenophora graminea S. Ito and Kuribayashi occurs in Japan and
is reputed to be the perithecial stage of H. gramineum, but Weiss states
that it has not been reported from the United States (1945)'.
H. gramineum usually does not sporulate in pure culture although
under common storage conditions in northern California it was made
to fruit readily and abundantly (Houston and Oswald, 1946).
Range: United States, Argentina, Australia, Canada, China, Cyprus, Den-
mark, England, France, Germany, India, Ireland, Japan, Morocco,
Netherlands, Norway, Peru, S. Africa, Sweden, U.S.S.R., world-wide.
References: Aberg (1945); Anon. (1921); Arny (1945,a,6); Christensen
and Graham (1934); Dickson (1939, 1947) ;,DrechsleT (1923, pp. 650-
656); Fuchs (1930); Graham (1935); Haan (1926); Hayes and Stakman
(1921); Houston and Oswald (1946); Isenbeck (1930); Johnson, T.
(1925); Leukel, Dickson, and Johnson (1926, 1933); Majdrakoff (1932);
' Nisikado (1928); Paxton (1922); Reddy and Burnett (1930);'Roemer,
et al. (1938); Shands, H. L. (1934); Shands, R. G. (1939); Shands and
Arny (1944); Smith, N'. J. G. (1929); Smith and PutteriU (1932); Smith
and Rattray (1930); Stelzner (1934); Suneson and Santoni (1943);
Tapke (1946); Verhoeven (1921); Vogt (1923); Weniger (1932);
. -Winkleman (1929).
• Helminthosporium hadrotrichoides Ell. and Ev.—Mold

H. leucostylum Drechsl.
In white spots on withering or withered leaves which eventually
shred; conidiophores subhyaline to light fuliginous, emerging from
stomata singly or in groups pi 2 t o . 6 ; measuring usually 35-100 X
5-6 /x, 2- to 8-septate, geniculations very pronounced; spores dark
olivaceous, measuring 15-67 X 11-17 /x, averaging 50 X 15 /.i, 1- to 6-
septate, typically 3- to 5-septate, not constricted, narrowdy ovoid, wid-
est about one half the distance above the base, proximal portion ex-
hibiting a paraboloid contour, the distal portion tapering uniformly to
the narrow apex, then abruptly rounded off, more rarely and atypically
subcylindrical, exospore thick except at extreme apex and at hilum.
On: Elemine indica (L.) Gaertn., Alabama, D. C, Kans., Ky., La., Md.,
Wise.—Eragrostis cilianensis (All.) Lk., Del., Md., Mont., N. Dak., Va.
Drechsler distinguished his H. leucostylum from H. hadrotrichoides
because of the "smoky brown" color of the latter and in its having
"the apex swollen so as to form a knob-like head of a pestle, 8-12 /A
in diam." Lefebvre however has determined material on Eragrostis as
belonging to H. hadrotrichoides and it is our belief that he considers
H. leucostylum the same as H. hadrotrichoides- (see also Greene, 1944,d,
p. 261).
Material on Eragrostis cilianensis from Miles City, Montana, col-
lected by the writer and determined by Lefebvre as H. hadrotrichoides,
had medium gray, linear streaks followed by withered, sometimes
shredded leaves or vague spots on dead leaves. The writer has isolated
the fungus from the roots of this grass which had been growing in
sandy soil at Mandan, North Dakota.
366 DISEASES OP CEREALS AND GRA^ÊS
Drechsler also determined H. rostratum on E, citianensis but this
species is readily distinguished by the beaked elongated; apical portion
of the spores. Mitra and Mehta (1934,fo) found that H. leucostylmn
grew best on nutrient media at pH 6.7. These authoBs studied both this
species and H. nodulosum on Eleusine spp. in Indiai {1934,a,6).
Nisikado (1928,6) described Ophiobolus kusanoi on Eragrostis
major as the perithecial stage of a species of Helminthospqrium which
he said resembled H. eragrostidis P. Henn. The conidia were short,
obclavate, 3- or 4-septate, dark olive, averaging 45 X 15 /*. The fungus
could attack both Eragrostis major and Eleusine indica.
Range: United States, India, S. Africa, Uganda.
References: Drechsler (1923, pp. 710-711," PI. 23); Ellis and Everhart
(1888,6); Mitra (1931,c); Mitra and Mehta (1934,a,&); Nisikado
(1928,6); Smith and Putterill (1933).
Helminthosporium halodes Drechsl.—Mold

Causing poorly defined, bluish, discolored areas, particularly on the
leaf sheath immediately below its junctiire with the blade, and from
brown roots of plants. Conidiophores arising from discolored spots after
death of the host, singly or in groups of two, geniculate towards the
apex, 60-150 X 4-7 /x., 1- to 5-septate; conidia on type host (Distichlis)
straight or curved, brownish yellow, and segments lighter, fusiform,
apical portion tapering to a blunted, somewhat hemispherical terminal,
basal cell, pointed or acuminate with prominent protruding hilum, 20-
105 X 10-14 ft, 1- to 12-septate, scarcely constricted at the-septa, spores
on artificial media olivaceous, except for hyaline end cells, fusiform,
definitely smaller than H. sativum. I
On: Agropyron repens (L.) Beauv., N. Dak.—Distichlis spicata (L.) Greene,
N. Y.—Muhlenbergia japonica Steud., N. Dak.—Panicum miliaceum L.,
N. Dak.—Schedonnardns paniculatus (Niitt.) Trel., N. Dak.—Setaria
italica (L.) Beauv., N. Dak.—S. viridis (L'.) Beauv., Minn., N. Dak., S.
Dak.—Triticum aestivum L., Sask. (glume discoloration).
H. halodes appears to be fairly common as a saprophytic mold and
sometimes causes secondary seed rot or possibly root rot in Agropyron
repens (L.) Beauv. and other grasses in the'northern Great Pl&ins. The
species is not recogiiized as being associated with any particular root
rot symptom, and does not appear to be at alf prevalent in any series
of isolations from grass roots.
In culture ,the dark spores are somewhat longer and less "kneed"
than those of Curvularia geniculata although some isolates are con-
fusable, yfhen immature, with the 4-septate phase of,this species.
Mitra, (1931,6, p. 287) described H. h'atodes var. tritici Mitra on
wheat with decayed roots in India. The conidia were cylindrical or
elliptical with the distal end abruptly rounded and the proximal end
tapering somewhat towards an acute base with prominent, protruding
hilum. The spores were widest at their middle, straight or slightly
curved, 2- to 9-septate (average 6), light yellowish brown to smoky
brown, the two end cells being slightly lighter, 23-73 X 13-20 /t, mean
50 X 16.5 [i. The description resembles some of the material isolated
from roots of various grasses at Mandan, North Dakota.
Most of the isolates of H. halodes which I worked with at Mandan
were saprophytes but some cultures caused mild seed rots (Sprague,
1944,6). The pure cultures on potato-dextrose agar resemble the dark
olive to gray cultures of many isolates of H. sativum or of Curvularia
•geniculata on the same medium.
Range: United States, India, Italy.
References: Drechsler (1923, pp. 707-709, PI. 22, 23); Mitra (1931,&);'
Sprague (1944,6).
Helminthosporium leersii Atk.—Leaf Spot

Spots irregularly oblong, amphigenous, at first dark brown, then
dirty white with a dark brown border. Hyphae amphigenous, brown,
irregularly nodulose or flexuose, 200-350 X 4-6 /x. Conidia slightly
curved) -5- to 9-septate, elliptical, faintly fuliginous, 50-95 X 11-14 /x.
On: Leersia virginica Willd., Alabama, Conn., D. C, Md.
The fungus first appears as minute brown spots not over 1 mm. in
length. Later the spots enlarge by formation of additional brown tissue
which merges with the green of the leaf. Later the brown center fades
to gray which contrasts sharply with the margin as an eye spot type of
infection. The leaves eventually wither and the fungus fructifications
follow.
On artificial media this species forms a dense gray, slow-growing
mycelium. Drechsler reports that the imbedded hyphae at the edge of
the colony produce short swollen elements which are intricately
branched.
Preston (1945) reports H. leersii on Bromus inermis from Okla-
homa.
References: Atkinson (1897); Drechsler (1923).
Helminthosporium maydis Nisik. and Miy. See its perithecial stage,

Cochliobolus heterdstrophus Drechsl.
Helminthosporium micropus Drechsl.—Leaf Blight

Attacking leaf blades of young plants, killing foliar tissues in elon-
gated areas, usually varying in length from 5 to 30 mm., the affected
parts having at first a scalded aspect and later becoming dry and
shriveled. On the foliage of older plants, affected regions more re-
stricted in extent, elliptical, usually with brownish margins, bring
about death of leaf by more gradual progressive withering.'
Conidiophores on withered leaves emerging singly or in groups of
2 or 3, approximately 70-165 X 4-6 [i, moderately geniculate; conidia
slightly tapering, except smaller spores which arej somewhat elliptical,
apex usually rounded off by hemispherical or hemi-ellipsoidal contour;
basal segment approximately obconical, tapering uniformly to a width
of 2 to 3 /i, then produced as a very short prolongation terminating in
hilum. Peripheral wall moderately thin, but visibly double-contoured
except in small circular spot at apex and about b ^ a l modification im-
mediately adjacent to hilum, where it becomes very thin and appears
single-contoured. Germinating by polar tubes from each of the thin-
walled regions, 28-92 X 10-18 ix, 3- to 9-septate, nonconstricted.
On: Paspalum bosdanum Flugge, Fla.
Drechsler reported (1923) that this fungus caused a quick flaccid
wilting followed by drying and death of the leaves or plant. The conidia
can be readily distinguished from those' of any other Helminthosporium
by the combination of their being light colored as iuv H. tritici-repentis
and being more or less curved, and because they germinate by pro-
ducing two polar tubes. H. micropus does iiot taper towards the ends
as much as H. turcicum. A final distinguishing feature is the short
cylindrical prolongation at the conidial base which terminates in the
flat hilum. '
Drechsler reports that the violent wilting induced by this species is
unique among the ones which he studied (1923), although he induced
similar quick decline by inoculating corn, seedylmgs with quantities of
conidia of H. sativum. j
Drechsler compared his species to two others described on Paspalum
and found that the species in question was distinct (1923). H. peni-
cillosum Speg. is a saprophyte described' from Ar^ntina on decaying
culms of P. platensis and on Andropogon sp. (Spegazzini,_^1899). This
had large conidiophores 150-600 X 10-15 ju, and spores 60;^5 X 10-12 fi.
Miles (1917) described H. mayaguezense on P. conjugatum in Puerto
Rico with conidiophores 300-500 X 18-22> and conidia 135-155 X 35-
45 /x but only 3- to 4-septate.
Weiss listed Helminthosporium sp. on P. urvillei Steud., P. notatum,
Flugge and on P. paniculatum L., all from Florida (1945). He also
listed H. rostratum Drechsl. on P. bosdanum Flugge from Georgia.
Helminthosporium monoceras Drechsl.—Blotch

H. crusgalli Nisikado and Miyake
Forms dark brown or chocolate spot blotches, 0.3-1.0 X 1-3 mm.,
nonzonate, foliage eventually withered and larger spots less distinct.
Conidiophores appearing after the death of the affected tissues, thick-
walled, emerging mainly from stomata in groups of 2 or 3,120-325 X 6-
9 fx, dark brown or olivaceous except at the extreme tip which is nearly
.hyaline,,usually 3- to 7-septate, moderately geniculate. Conidia yellow-
. ish when young, becoming dark olivaceous when mature, same color
as H. sativum, 40-150 X 15-22 /j., typically straight or showing a slight
crescentic or sigmoid curve, yet not infrequently exhibiting more pro-
nounced bends; widest at the middle segment unless modified by a
median constriction, tapering gradually towards the tip to one third
or one half of its maximum width, then bluntly rounded off; tapering
towards the base to approximately one sixth of the median diameter,
the contours then curving gently into the protruding hilum, 3- to 10-
septate, seldom constricted. The peripheral wall at maturity thick as
in H. sativum or H. vagans except at the apex and about a subhyaline
narrow zone at the proximal end near the hilum. Germination occurs
from these two polar thin areas.
On: Echinochloa crusgalli (L.) Beauv., N. Y., Wise. Has been reported as
isolated from the roots of Panicum miliaceum L. and Setaria viridis
(L.) Beauv. from N. Dak. by the writer but perhaps these isolations
represented off-types of H. turcicum or an undescribed species.
The long-curved or geniculate, sharp-pointed conidia are character-
istic of this species. The fungus has, when mature, much darker spores
than has H. turcicum.
Schweinfurth and von Thuemen (1878) list H. flexuosum Corda
{Brachysporium flexuosum (Cda.) Saec.) on the culms and inflorescence
of E. crusgalli in Egypt: This may possibly be the same as Curvularia
trifolii (Kauf.) Boed.
An undescribed species of Helminthosporium is listed from the
vicinity of Sydney, Australia, on E. crusgalli (Noble, et al., 1934).
Range: United States, China, Japan.
References: Dickinson (1932); Drechsler (1923, pp. 704-707, PL 20, 21);
Greene (1949,c); Ling (1947); Nisikado and Miyake (1925).
Helminthosporium nodulosum Berk, and Curtis—Blight

Spots on leaves at first oval and light brown, later elongating
parallel to the axis of the leaf, becoming deep brown, lesions as much
as 1 cm. long and 1 mm. wide or somewhat larger through coalescence;
leaves which are severely injured wither prematurely and the seed yield
is reduced. Leaf sheaths and heads may become infected. Fungus forms
a small woolly, grayish brown mass in. center of! each spot, composed
of sporophores arising from internal mycelium (through the stomata
and also, though less often, directly across the epidermal cells. Conidio-
phores erect, unbranched, often much curved or with prfcjminent genicu-
lations, dark brown, paler at the distal portion, with numerous septa,
80-250 X 5-7 ft. Conidia produced singly at apex, geniscars formed,
conidia straight or curved, somewhat fusiform and flattened on the
incurved side, obtuse but gradually tapering, clear brown, relatively
heavy-walled, 4- to 10-septate, 50-100 X 12-17 jx..
OQ»C Eleusine indica (L.) Gaertn., Miss., S. C.
Butler (1918) gives the longest description of this species and illus-
trates the spots, conidiophores, and spores. Possibly the fungus he dis-
cusses on Eleusine coracana Gaertn. from India is distinct from the
meagerly described H. nodulosum, but we are using Butler's-description
at this time. Mitra (1931,c) reported results of studies in which cross
inoculations showed that H. nodulosum could infect corn, Italian
millet, and Panicum fnimentaceum but not wheat, oats, or barley. He
found that the species attacked all parts of the plant at all stages of
its growth. McRae (1932) found that the temperature range for infec-
tion by H. nodulosum was 10° C. to 37° C. with an optimum of 30° C -
32° C. This species grows best in nutrient agar with pH 7.1 (Mitra
and Mehta, 1934,6). Castellani (1938) described the symptoms on E.
coracana from Eritrea and reported 4- to 8-septate conidia, 48 X 10 /«
on conidiophores 105 X 5 /x. Thomas reported that H. nodwiosum'at-
tacked both E. coracana and wheat at Madras and even grew at a
temperature of 36.5° C. (1939).
Range: United States, China, India, Italian E.'Africa (Eritrea), Philip-
pines, j
References: Butler (1918); Castellani (1938); McRae (1932); Mitra
(1931,c); Mitra and Mehta (1934,6); Thomas (1939).
Helminthosporium oryzae V. Breda de Haan—^rGwn Spot

H. oryzae Miyake and Hori —
Cochliobolus miyabeanus (Ito and Kuribayashi)
fide Dickson is perithecial stage
Spots in center dry and stramineous to giây (Tullis) surrounded by
a brown or reddish brown margin, longitudinally elongated, 0.5 to 3
mm. diam. Conidiophores arising from underside of the leaf or from
dark olivaceous mats on the glumes, 150-600 X 4-8 î, those on the
glxirues arising from stout prostrate hyphae^,8^15 ju, diam.; conidia large,
fuliginous, fusiform, acrogenous, 5- to" 8-septate, 16-90 X 11-17 jx,
sometimes 170 /i long and then containing as many as 13 septa, these
larger ones usually formed from mycelial mats on the glumes. Spores
slightly curved, widest at the middleôr somewhat below the middle.
the distal portion tapering towards the hemispherical apex where its
width is half the median width, the proximal portion tapering toward
the base which is similarly rounded off although less so than apex.
Spore wall when fully mature fuliginous or brownish and moderately
thin, germinated by polar germ tubes. Perithecia globose, pseudo-
parenchymatous, black with an ostiolar beak. Asci cylindrical to long
fusiform, slightly curved, containing 4 to 6 filiform ascospores in a
close spiral (helix), the spores longer than asci and up to 468 /^ long
when uncoiled after emergence from the ascus, 6-9 /t diam.
On: Oryza sativa L., Ark., Calif., Fla., La., Miss., Tex.
This causes brown spotting of the leaves, injury to the neck of the/
culms, the panicles, and glumes, and also causes a seedling blight.
Nisikado and Miyake (1922, see R.A.M. 2:230-231) discussed the
symptoms. When present on the grain it causes a discoloring which is
called "pecky rice" (Tullis, 1935). This fungus produces the same
general group of disease symptoms on rice that H. sativum causes on
upland grains.
Von Thuemen discussed H. macro carpum Greville as a saprophytic
mold on dead rice plants. According to Drechsler (1923) the sporo-
phores of H: macrocarpum are 15 to 20 /i wide which equals or exceeds
the width of the conidia. Apparently, according to Drechsler, the iden-
tity of this material which von Thuemen had was not clear, although
••JVei considered it the same as H. oryzae. It appears to be necessary to
reject this earlier name'as ambiguous and take up H. oryzae described
later by Van Breda de Haan (1900). In 1901 Miyabe and Hori de-
scribed H. oryzae Miyabe and Hori (cf. Hori, 1922) which is consid-
ered the same as H. oryzae Van Breda de Haan. This species has been
widely reported from the tropics (Wei, 1936). In Russian Central
Asia, Loukyanovitch et al. state that H. oryzae spreads to rice from
barley (1931).
Nisikado and Miyake (1920) found that spores of H. oryzae were
killed by immersion in water at a temperature of 51° C. for ten minutes
while air-dry rice seed was not injured by immersion for 10 to 15
minutes in water at a temperature of 54° C. to 55° C. They recom-
mend treatment of rice' seed in water for 10 minutes at 52° C. or for
five minutes at 54° C. after presoaking for 24 hours at room tempera-
ture. However H. oryzae overwinters in the field and cannot be con-
trolled by seed treatment, according to Adair (1941). These results
were reported by Cralley and Tullis from Arkansas earlier (1937).
Adair went on to state that the-only satisfactory control of this fungus
appeared to be through breeding of resistant varieties. He found that
resistance in rice was • a recessive character. Others have reported
varietal differences in reaction to H. oryzae. Lin (1936) found three
varieties of rice very resistant to H. oryzae while Ocfemia (1924,a),
Tullis (1935), and Suzuki (1935) reported various degrees of resistance
among rice' varieties to H. oryzae. Sherf, Page, Tullis, and Morgan
(1947) stated, however, that no varietal resistance to brown spot was
demonstrated in their trials. They determined tjiat high temperatures
did not predispose rice plants to infection, but .high humidity not ac-
companied by cleansing rains did increase iniection. The optimum
temperature range for infection was 20° C.-25'^. C. Ocfemia (1924,5)
gave 16° C.-24° C. as optimum, 16° C.-36° C. as extreme, and 20° C -
30° C. as the extreme range. They found that rice was susceptible to
leaf infections at all stages of development. Tullis found that in re-
sistant varieties of rice the invading fungus was hemmed in by forma-
tion of intercellular deposits, the chemical nature of which was not
determined (1935). Baldacci (1947) found that early-season tempera-
tures prevented infection (below 18° C ) .
Control is made further difficult by the fact that rotation of crops
is scarcely practical because of the longevity of the mycelium and of
the spores in the field. Such workers as Ito (1932), Lin (1936), Suzuki
(1930), or Wei (1936) found that fungus material remained viable for
1 to 3 years under field conditions.
The perithecial stage of H. oryzad was found in pure culture and
named Ophiobolus miyabeanus Ito aiid Kuribayashi (1927). Drechsler
suggested placing this species in -Cochliobolus because its very long
filamentous ascospores were spirally coiled in the asci. Thomas (1940)
had difficulty inducing H. oryzae to sporulate on potato-dextrose agar
but it sporulated more readily on corn meal-sand media and on potato-
dextrose agar plus tannic acid. Its host range was considerable.
Range: United States, Afghanistan, Belgian Congo, Brazil, Burma, Ceylon,
China, Colombia, Egypt, French Guiana, Gold Coast, India, Indochina,
Italy, Jamaica, Japan, Java, Malaya, Peru,/Philippines, Puerto Rico,
Sierra Leone, Sumatra, Tanganyika,) Togoland, Trinidad, Turkey,
Uganda, U.S.S.R. |
References: Adair (1941); Baldacci (1947; 1947-48,a,&); Baldacci and
Ciferri (1936); Cralley and Tulhs (1937); Drechsler (1923; 1934,^);
Ganguly-41946,6); Haan (1900); Hemmi (1937^ Hemmi and Nojima
(1931); Hemmi and Suzuki (1931); Hemmi and YokQgi^(1928); Hori
(1922); Ito (1932); Ito and Kuribayashi (1927); Katsura (1937); Lin
(1936); Motte (1947); Nisikado (1923; 1927,c); Nisikado and Miyake
(1920; 1922); Ocfemia (1924,a,&); . Padmanabhan, Choudhry, and
Ganguly (1948); Page, Sherf, and Morgan (1947); Sherf, Page, Tullis,
and Morgan (1947); Sundararaman (1922); Suzuki (1930; 1935);
Tanaka (1920); Thomas (1940); Tisdale (1922); Tochinai and Sakamoto
(1937); Tullis (1935); Wei (1936).
Helminthosporium poae Bâudys-rMold

Spots on the leaves yellowish with brown margins; conidia 36-73 fi
long, 2r to 6-septate. Somewhat more fusiform than cylindrical.
O n : Poa secunda Presl, Greg., Wash.-^P. trivialis L., Greg.
Scattered and scanty material seen at Astoria, Oregon, on P. triv-
ialis, and on P. secunda in the Malheur National Forest in eastern
Oregon and near Bingen, Washington, does not seem to fit the descrip-
tion of H. vagans. The description of H. poae Baudys (1915-16) from
Bohemia is too meager to be at all certain that our material is this
species. Until such fragments are better known they can be assigned to
this species.
Range: United States, Czechoslovakia.
References: Baudys (1915-16); Drechsler (1923, p. 688); Sprague (1937,e;
194:2,a,b).
Helminthosporium ravenelii Curt, and Berk.—Head Mold

H. crustaceum P. Hennings
H. hojjfmannii Berk, and Curt.
H. tonkinense Karst. and Roum. (1890)
Affected floral parts covered with a velvety or spongy brownish
olive, later black-crusted mass of hyphae; conidiophores branching, in
masses light fuliginous to light yellow, torulose and variable in diam-^
eter, 5 to 10 /* and up to 500 jn long, septate at intervals of 20 to 40 /x;
spores borne in great quantities at the apices and geniculations of the
conidiophores, slightly sigmoid, rounded at both ends, the apical
slightly broader as the two basal cells taper more than the apex,
22-78 X 12-19 fi, 1- to 5- (usually 3- to 4-) septate, septations rarely
constricted.
On: Sporobolus cryptandrus (Torr.) A. Gray, Wise.—S. elongatus Dur. and
Schinz, Hawaii—S. indicus (L.) R. Br., Alabama, Fla., Ga., La., Miss.,
N. C, S. C, Tex.—S. poiretii (Roem. and Schult.) Hitchc, Fla., Ga., La.
The mold is so common in the southern United States on the host
S. indicus that this grass is known as smut grass.
Drechsler (1923) summarized the early history of this fungus which
was first described from Carolina material.
Range: United States, Argentina, Australia, Bermuda, Brazil, China, Co-
lombia, Costa Rica, Cuba, Hawaii, Java, Mexico, New Zealand, Philip-
pines, Uruguay.
References: Curtis (1848); Drechsler (1923); Hennings (1902); Karsten
and Roumeguere (1890).
Helminthosporium rostratum Drechsl.—Leaf Spot

Lesions small, 2-5 X 1-2 mm., vein delimited, straw-colored with
slightly browned edges and terminal yellowing; conidiophores dark
olivaceous, emerging singly or in groups of 2 to 5, the swollen bases
more or less united, 4 0 - 1 8 0 x 6 - 8 fi, 1- to 6-septate; conidia when
mature dark olivaceous, straight or less frequently somewhat curved,
FiQ. 73.—IIelviinlhosporium roslratum on Pennisetum glaucum, Tifton, Ga.,

X360. (Photo furnished by A. G. Johnson from Young, Lefebvro and Johnson,
U.S.D.A.)
often short, widest at or somewhat below the middle, tapering moder-

ately or more markedly toward both ends, the hemispherical apex
abruptly romided off; the basal end somewhat more acute, often ex-
hibiting a rounded conical contour but often strongly rostrate at the
tip, many spores almost oblanceolate, hilum protruding and prominent,
8- to 15-septate but elliptical spores only 3- to 9-septate, basal septum
and distal septum often thicker and more prominent than others,
peripheral wall thick, spores 32-184 X 14-22 fi.
On: Eragrostis cilianensis (All.) Lk., D. C, Wise.—E. pectinacea (Michx.)
Nees, Wise.—E. spectabilis (Pursh) Steud., Wise.—Paspalum boscianum
Flugge, Ga.—Pennisetum glaunim (L.) R. Br., Ga.—Sorghum vulgare
Pers., Fla.—S. vulgare v;ir. smlanense (Piper) Hitchc, Fla.—Zea mays
L., Ga., Miss.
Recently H. rostratum was found by Young, Lefebvre, and A. G.
Johnson (1947) on various warm-temperature-loving hosts (Fig. 73)
including corn and sorghum in the southeastern states. Their excellent
photomicrograph illustrates typical rostrate spores of the species. Bunt-
ing (1927,1928) reported a similar fungus on corn from the Gold Coast
in Africa.
Range: United States, Gold Coast.
References: Bunting (1927, 1928); Drechsler (1923); Lefebvre and John-
son, H..W. (1941); Young, Lefebvre, and Johnson, A. G. (1947).
Helminthosporium sacchari (Van Breda de Haan) Butler;—Eye,

Spot (Florida Ring Spot)
H. ocellum Faris
Causes an eye spot the center of which, when mature, is antique
brbnze, finally straw color or Spanish raisin color surrounded by a
yellow halo; spots somewhat elongate, often with sharp ends; entire
leaf may wither; spots sometimes on sheaths and culms, as much as
11 cm. long; conidiophores with bulbous bases, 70-278 (av. 30-160
ju.) X-3.7-9.25 }!., 3- to 8-septate; spores brown, curved, fusiform, taper-
ing to bluntly pointed ends, 3- to 9-septate, 24-93 X 11-17 /x.
On: Cymho'pogon citratus (D. C.) Stapf., Fla.—Pennisetum glaucum (L.)
R. Br., Hawaii—P. purpureum Schumacli., Fla., Ga., Hawaii—Saccharum
officinarum L., Alabama, Fla., Hawaii, La.
.Weiss (1945) credits this fungus to Cercospora sacchari Van Breda
de Haan which is the same as H. sacchari Butler. In 1934 Bourne com-
pared the species to H. ocellum Faris (1928,a) who claimed that his
species, H. ocellum, was distinct from H. sacchari and H. stenospilum
Drechsl. Parris (1942) reviewed the history of these forms showing
that Butler in his revision of Cercospora sacchari Van Breda de Haan
had given smaller measurements than actually occurred in H. sacchari.
Parris (1942) concluded that H. ocellum Faris was, therefore, a syno-
nym of H. sacchari. Parris (1942) detailed H. sacchari as the cause of
eye spot of Napier grass [Pennisetum purpureum) in Hawaii. In a foot-
note Parris reported that some isolates produced spores which closely
resembled H. stenospilum. McRae (1933) also found a "mutant" of
H. sacchari which resembled H. stenospilum. Eye spot disease of
Napier grass also occurs in Florida (Vorhees, 1938) as does eye spot
of lemon grass (Cymbopogon) (Bourne, 1941).
H. sacchari is widely distributed and one of the important diseases
of sugar cane. Bourne (1934) determined that the associated fungus,
Leptosphaeria sacchari Van Breda de Haan, was only a saprophyte,
thus refuting observations made by various workers including Van
Breda de Haan (1892,a). Bourne also found that the associated Phyl-
losticta sorghina Sacc. was "saprophytic" but was capable of increas-
ing the size of the spot and "inducing a target spot effect." Another
species of Phyllosticta, P. saccharicola Hennings, was found to be the
conidial stage of the saprophytic Leptosphaeria sdcchari, above men-
tioned. P. sacchari Speg, is a synonym of P. sorghina. These various
species differed in pure culture, one from another i(pourne, 1934).
Matsumoto and Yamamoto (1934) illustrated in color what they
determined as eye spot {H. ocellum) but which they somewhat
obliquely later concluded (supplement) was probably brown stripe
{H. stenospilum).
Lee and Martin (1928) recommend avoiding application of nitrog-
enous fertilizer on canes in Hawaii except from April to July inclusive.
Phosphoric acid at the rate of 400 lbs. per acre tended to reduce in-
fection. Martin (1933) further reported that the application of nitrog-
enous fertilizers in late fall appeared to favor development of H.
sacchari.
Range: United States, Argentina, Barbados, • Borneo, Brazil, Canal Zone,
Colombia, Cuba, Egypt, Fiji, Gold Coast„'Hawaii, India, Jamaica, Java,
Lesser Antilles, Madagascar, Malaya, ^ Mauritius, Peru, _ Philippines,
Puerto Rico, Queensland, Santo Domingo, Singapore, S. Africa, Uganda,
Venezuela.
References: Averna-Sacca (1916); Bell (1929); Bijl (1921); Bourne
(1934, 1941); Butler (1918); Butler and Hafiz (1913); Caum (1921);
Cobb (1909); Cook, M. T. (1924,a,&; 1926; 1931,a,fe); Faris (1928,6);
Haan (1892,5); Halma and Fawcett (1925); Johnston and Stevenson
(1917); Krueger (1890); Lee (1926, 1929); Lee and Martin (1926);
McRae (1933); Martin (1938); NoUa (1949); North (1923); Nowell
(1923); Parris (1942); Parris and Ripperton (1941); Eriode (19^1);
Ripperton (1940);^Vorhees (1938).
Helminthosporium sativum P. K. B.—Spot Blotch, Root Rot, Seedling

Blight, Black Point and Dryland (Common) Root Rot.
Cochliobolus (Ophiobolus) sativus (Ito and Kuribayashi) Drechsler
fide Dickson is the perithecial stage which is seldom found
Causes a dark brown seedling rot of the coleoptile region, a spot
blotch on the leaves, a mold of the leaves, spikes and seed, a root rot
or crown rot of maturing plants; conidiophores brown, emerging from
the stomata or between epidermal cells after death of the host tissue,
singly or in fascicles of 2 or 3,110-150 X 6-8 /*, up to 8-septate. Grenicu-
lations well defined, 5 or 6; spores curved, tapering evenly toward each
rounded end, or spores irregularly, thickly boomerang-shaped, bent
elliptical ovate to pip-shaped, rarely bilobed or triangular in outline,
3- to 10-septate, 60-120 X 15-20 fi, more variable yon wheat heads,
dark olive brown with a thick brittle episp9j£;~hilum conspicuous but
situated within the contour of rounded basal'end.
On: Agropyron buonapartis (Spreng.) Dur. and Schinz., ,N. Dak.—.4.
ciliare (Trin.) Franch, N. Dak.—A. cristatum (L.) Gaertn., Md., Minn.,
Mont., Nebr., N. Dak., Ont., Oreg., ^. Dak., Wash,, Wyo,—A. dasys-
tachyum (Hook.) Soribn., N. Dak.—A. desertorum (Fisch.) Schult., N.
Dak., S. Dak.—A. elongatum (Host.) Beauv., Nebr., N. Dak.—A. inter-
medium (Host.) Beauv., N. Dak., S. Dak.—A. michnoi Roshev, N.
Dak., S. Dak.—A. mongolicum Keng, N. Dak.—A. pendulinum (Nevski)
ined. (fide Swallen), N. Dak.—A. repens (L.) Beauv., Alta., Iowa, Minn.,
Nebr., N. Dak., S. Dak., Wise.—A. riparium Scribn. and Sm., N. Dak.—
A. semicostatum (Steud.) Nees, N. Dak., Wash.—A. sibiricum (Willd.)
Beauv., Minn., Nebr., N. Dak., Oreg., S. Dak., Wyo.—A. smithii Rydb.,
Colo., Minn., Mont., Nebr., N. Dak., S. Dak., Wyo.—A. subsecundum
(Lk.) Hitchc, Alta., N. Dak.—A. trachycaulum (Lk.) Richt., Alta., Mont.,
Nebr., N. Dak., S. Dak., Wyo.—A. trichophorum (Lk.) Richt., N. Dak.—
Agrostis alba L., Wyo.—Ammophila arenaria (L.) Lk., N. Dak.—Andro-
pogon hallii Hack., N. Dak.—Arthraxon hispidus (Hack.) Honda, N. Dak.
—Avena byzantina K. Koch, Ga., N. Dak., Oreg.—A. fatua L., Minn., N.
Dak., S. Dak.—A. sativa L., Alta., Colo., Ga., Iowa, Minn., N . B., N. Dak.,
Okla., Oreg., Que., Sask., S. Dak.—Bouteloua curtipendula (Michx.)
Torr., N. Dak., S. Dak.—B. gracilis (H. B. K.) Lag., N. Dak., S. Dak.—
'Bromus arvensis L., N. Dak.—B. carinatus Hook, and Arn., N. Dak.,
Wash.—B. catharticus Vahl, N . Dak.—B. erectus Huds., N . Dak.—B.
inermis Leyss, Alta., Iowa, Minn., Nebr., N. Dak., S. Dak., Wyo.—B.
japonicus Thunb., Calif.—B. madritensis L., N. Dak.—B. tectorum L.,
Nebr., N. Dak., S. Dak.—Calambvilfa longijolia (Hook.) Scribn., N.
Dak.—Cenchrus paucifiorus Benth, N. Dak.—Chloris verticillata ,Nutt.,
Minn.—Dactylis glomerata L., Minn., N. Dak., S. Dak., Wash.—Digitaria
sanguinalis (L.) Scop., Minn., N. Dak.—Echinochloa crusgalli (L.) Beauv.,
Minn., N. Dak., Wyo.—Elymus canadensis L., Minn., Nebr., N. Dak.,
S. Dak., Wise.—E. canadensis var. robustus (Scribn. and Sm.) Mackenz.
and Bush, N. Dak.—E. dahuricus Turcz, N. Dak.—E. dasystachys Trin.,
N. Dak.—E. interruptus Buckl., N. Dak.—E. junceus Fisch., N. Dak.,
S. Dak.—E. macounii Vasey {Hordeum jubatum X Agropyron trachy-
caulum), N. Dak.—E. sibiricus L., N. Dak.—E. virginicus L., Minn.-—•
Eragrostis cilianensis (All.) Lk., N. Dak.—E. curvula (Schrad.) Nees, N .
Dak.—E. pilosa (L.) Beauv., N. Dak.—Festuca elatior L., Iowa, N. Dak.
—F. octoflora Walt., N. Dak.—F. rubra L., N. Dak., Va.—Hordeum brevi-
subulatum (Trin.) Lk., N. Dak.—H. bulbosun L., N. Dak.—H. distichon
L., Calif., Colo., Iowa, Manit., Minn., Mont., Nebr., N. Dak., S. Dak.,
Utah, Wash., Wyo.—H. jubatum L., Minn., N. Dak., S. Dak.—H. murinum
L., Oreg., Wash.—il. vulgare L., Alta., Ark., B. C , Calif., Colo., Conn.,
Ga., Ida., 111., Ind.,- Iowa, Kans., Ky., Manit., Md., Mass., Mich., Minn.,
Mo., Mont., Nebr., N. B., N. Y., N. C , N. Dak., Ohio, Okla., Ont., Oreg.,
Penn., P. E. Is., Que., Sask., S. C , S. Dak., Tenn., Tex., Va., Wash.,
W. Va., Wise, Wyo.—Hystrix patula Moench, Minn.—Koeleria cristata
(L.) Pers., N. Dak.—Lolium perenne L., Minn.—Muhlenbergia foliosa
(Roem. and Schult.) Trin., Wise—M. racemosa (Michx.) B. S. P., N. Dak.
—Oryzopsis hymenoides (Rbem. and Schult.) Rick., N. Dak., S. Dak.,
Wash.—Fanicum capillare L., Minn., N. Dak., S. Dak., Wise, Wyo.—
P. implicatum Scribn., Wise.—P. miliaceum L., Minn., N. Dak.—P. perlon-
gum Nash, Nebr.—P. virgatum L., Nebr.—Phalaris arundinacea L., Md.,
Wise.—P. tuberosa L., Oreg.—Phleum pratense L., N. Dak., Wash.—Poa
bulbosa L., N. Dak.—P. compressa L., N. Dak., Oreg., S. Dak., Wash.—
P. juncifolia Scribn., N. Dak.—P. palustris L., Minn., N. Dak.—P. pratensis
'I
L., N. Dak., S. Dak.—P. secunda Presl, N.,Pak., Oreg.—Secale cereale L.,
Alta., Calif., Colo., Ida., 111., Ind., Manit., Mich., Miip., MoBt., Nebr., N.
Dak., Ohio, Oreg., Perm., Sask., S. Dak., Utah, Wash., Wise, Wyo.—Setaria
italica (L.) Beauv., Minn., N. Dak.—S. lutescens (Weigel) F. T.Habb, N.
Dak.—S. viridis (L.) Beauv., Iowa, Minn., N. DakJ, S. Dak.—Sorghum
vulgare Pers., Minn., N. Dak.—S. vulgare var. sudanense (Piper) Hitchc,
N. Dak.—Stipa baicalensis Roshev, N. Dak,—S. comata Trin. and Rupr.,
N. Dak.—S. pulchra Hitchc, N. Dak.—S. sibirica (L.) Lam., N. Dak.—iS.
spartea Trin., N. Dak.—S. viridula Trin., N. Dak., S. Dak.—Triticum
aestivum L., general in the United States and Canada—T. dicoccum
Schrank, Minn., N. $)ak.—T. durum Desf., Manit., Minn., N. Dak., Sask.,
S. Dak.—T. monococcum L., Minn.—T. polonicwn L., Minn.—T. spelta
L,, Minn., N. Dak., Oreg.—T. timopheevii Zhuk., N. Dak.—T. turgidum
L., Minn.—Zea mays L., Iowa, Manit., Minn., N. Dak., S. Dak,
H. sativum, which was first described from Iowa, is now known
throughout great areas of the temperate zones where wheat and barley
grow.
Helminthosporium sativum occurs on many hosts, but it is most
important as a root rot and kernel smudge or blight of wheat and
barley. I t has been reported on dozens of species of grasses as well as
on oats, corn, and wheat relatives. In^the northern Great Plains, it has
been isolated from the roots of 79 species of Gramineae. It is especially
common on Elymus, Agropyron, Bromus, and Hordeum. It is less
abundant on species of Poa, which are resistant to some extent. The
host range as established by seedling and seed inoculation trials in the
greenhouse is very extensive. '
As mentioned, H. sativum is world-wide in distribution in the tem-
perate zone. It occurs in very acid soil (pH 4.8) in a region of very
heavy rainfall (70-100 inches yearly) in Oregoiyand is, paradoxically,
omnipresent in the arid and semiarid parts of the western United States.
While it is an important parasite of wheat( and barley, it takes second
place as a parasite of grass roots when compared with Pythium graihini-
cola. Its importance on grasses may have been somewhat exaggerated
in the past. However, because it is so universally distributed, it cannot
be dismissed as a minor parasite, even on grasses. • -
As a component of dryland root rot of fall-sown wheat, it is not
considered as important as it is on spring grain. Its effect is, neverthe-
less, important in spite of its obscure nature. In the Columbia Basin
of Oregon and Washington it is now recognized as an increasingly seri-
ous disease of bearded winter wheats. ^
Helminthosporium sativum has been studied in great detail by a
number of workers. Christensen has shown that it possesses a large
number of physiological races; also, it is g^gcally recognized as being
readilj' influenced by environment. Changes in light or moisture, or the
presence or absence of antagonists, can change a single isolate to almost
unrecognizably different subcultures. While H. sativum is able to thrive
in cool and moist weather, it often causes greatest injury in drought
periods that follow. The diseased plants have weakened root systems
and are unable to supply the plant with water and nutrients as readily
as do healthy plants with larger root systems. Vladimirsky (1937)
found that root rot was worse in soil with 30 per cent moisture at
tillering than in soil with 60 per cent moisture. It is one of the most
common components of common root rot in grasses in the plains of the
central United States and Canada (Padwick and Henry, 1933).
The seedling phase of this disease is characterized by a slow dark
rot of the coleoptile region with injury spreading later to the cortex and
small feeder roots. The seedlings of small grasses may be wiped out
early or even killed before emergence (pre-emergence rot), but the in-,
jury is less spectacular than that caused by Pythium graminicola. Such
hosts aa barley, which shows stunting or killing of lower leaves, very
often recover later with slight apparent ill effects if growing conditions
are optimum for the host.
- After the seedling phases have been passed, leaf spotting may occur.
This is a brown blotch rather than a definite spot or stripe. Abundant
spotting in late season on barley or wheat speeds up premature ripen-
ing of the grain. The fruiting bodies of the fungus may give the ma-
turing plants a blotchy or olivaceous cast. Barley seed and glumes show
"dark brown stains that are accompanied by shriveling of the kernels.
The basal portions of kernels are more likely to be discolored. Black
point of wheat and other grains is partly due to H. sativum (Evans,
Nevada S., 1921), and most of the injury that occurs at seeding time
in the northern Great Plains from seed-borne parasites is due to this
fungus.
In winter wheat, injury due to Helminthosporium sativum is vague,
particularly on wheat grown in the semiarid regions. The more or less
recognized term, "dryland root rot" (or foot rot), is used to designate
a particularly vague type of injury in which the plants show pro-
nounced stunting in more or less circular to irregular areas in the fields.
The crowns of these plants, when bisected with a sharp knife, uniformly
show a conical area of brown, prematurely dead, or necrotic tissue.
These areas of greatest injury in the field are frequently associated
with shallow soil, hardpan under-layers, or other conditions unfavor-
able for the growth of the plant. In the dry, winter wheat regions, oc-
casional white heads (sterile heads) develop in winter wheat showing
dryland root rot. These are particularly common in eastern Oregon and
Washington and are a phase of premature killing of plants by the
omnipresent H. sativum.
H. sativum was not described until 1910 (Pammel, King, and Bakke,
1910) which is surprising'because of its great prevalence. H. sorokinia-
num Sacc, however, was mentioned in 1891 (Sorokin, 1891) and de-
scribed by Saccardo (Sylloge 1892, v. 10, pp. 415-416) as a fungus on
the spikes of wheat and rye from Russia. The spores were described as
acrogenous, ovate-fusoid, 80-100 X 30 /x, reddish brown, 5- to 10-sep-
tate. Drechsler (1923) felt that this species was not proved the same
• (
as H. sativum because of the reddish brown color of the spores. While

this difference is merely that of personal opinion' as to color standards,
H. sativum cannot be reduced to synonymy until the types are com-
pared. This Saccardian species H. sorokinianum, as far as I know, is
in the same state as other Saccardian species—^the type is not available.
Since H. sativum is now so well established in literkture it would ap-
pear unwise to take up the Russian fungus. However, it is used,
especially in Russia. Mourashkinski (1924) considered H. sativum a
synonym.
H. sativum produces a velvety layer of gray to olive black mycelium
on potato-dextrose agar. The spores are light olive brown at first, often
ovate (pip-shaped), 1-septate to subglobose, but soon the spores formed
become fusoid, ellipsoid, wedge-shaped, or bent and are dark brown
with the characteristic brittle wall. Drechsler (lj923) and F. L. Stevens
(1922) show excellent illustrations. The worker who deals with H.
sativum becomes well aware that this species is extremely variable in
pure culture. Isolates which limit themselves to the production of a few
pip-shaped spores are especially confusing'.
Arny (1948) found that the difference between susceptibility and
resistance to H. sativum in barley appeared to be due to a single-factor
pair, with susceptible dominant. He found no detectable association of
spot blotch reaction with lemma and pericarp color (black or white),
awn barbing, rachilla hair length, or reaction to stem rust. Vladimirsky
(1937) reported that the spring wheat varieties Palestine, Tunis, and
Morocco and the winter wheat type ferrugineum showed a moderate
degree of resistance in the field.
Burnett, Murphy, Reddy, and Johnson (1947) report a selection of
Wisconsin 38 barley with unusual resistance to M. sativum; Mars was
the most resistant variety and Montcalm the most susceptible. Inci-
dentally in observations made in North E)akota I also found Mars
somewhat tolerant to common root rot and at Langdon, North Dakota,
during one season Montcalm was somewhat resistant. Velvon also
showed tolerance to cbmmon root rot in central and~.^western North
Dakota (Sprague, field notes). . "~
Christensen and Schneider (1948) found that a monosporous cul-
ture of Helminthosporium sativum that ^had been passed through
Marquis wheat for ten successive generations was relatively stable
with the frequency of mutation only 1:2900. /
Ito and Kuribayashi described Ophiobolits sativus (Kuribayashi,
1929) as- the perithecial stage of H. sativum. It is inferred that
Drechsler considered this as having a helicoid type of arrangement of
the ascospores in the asci and hence belonged in Cochliobolus (1934).
Dickson has credited Drechsler with this.Gombination, which should
perhaps best be called Cochliobolus sativus (Ito and Kuribayashi)
Drechfel. in Dickson (1947). As Dickson pointed out Dastur proposed
C. tritiQi (1942) and this appears to be a synonym of C. sativus.
Brittlebank and Adam (1924) reported,a conidial form identical to
Podosporiella verticillata O'Gara in Victoria (Australia). Pycnidia and
perithecia developed in nutrient agar. T h e peritliecial stage was called
Pleosphaeria semeniperda. T h e perithecia were slightly stalked, large,
680 X 500 [i, bristled, and bore conidia on their outer surface. T h e
fungus was parasitic on oats and wheat. Since this fungus is an earlier
name t h a n 0. sativus it should be investigated.
R a n g e : United States, Australia, Bulgaria, Burma, Canada, China, England,
Germany, Gold Coast, India, Italy, Japan, Kenya, Mexico, Morocco,
Norway, Peru, S. Africa, Uganda, U.S.S.R.
R e f e r e n c e s : Andrews (1943); Arny (1928); Beckwith (1911, 1912);
Blanchard and Carrera (1933); Bolley (1913); Boening and Wallner
(1934,a); Brentzel (1941, 1944); Brittlebank and Adam (1924); Broad-
foot (1933,6; 1934,0,6); Broadfoot and Tyner (1938,a,6); Burnett,
Murphy, Reddy, and Johnson (1947); Christensen (1922, 1925, 1926,
1936); Christensen and Davies (1937, 1940); Christensen and Schneider
(1948); Christensen and Stakman (1935); Dastur (1942); Dickson
"(1946, 1947); Dosdah (1923); Dosdall and Christensen (1923); Drechsler
. (1923, pp. 690-704, PL 17-19); Evans (1921); Greaney (1946); Greaney
and Bailey (1927); Greaney and Machacek (1933, 1942); Greaney,
Machacek, and Johnston (1938); Greaney and Wallace (1943); Gries
(1943); Griffee (1925); Hamblin (1922); Hanson and Christensen (1940,
1943); Hayes, Stakman, Griffee, and Christensen (1923); Henry (1924;
1931,a,6; 1935); Hynes (1923; 1935; 1937,c; 1938); Johnson, 'E. C.
(1914); Kuribayashi (1929); Ledingham (1942); Machacek (1936, 1943);
Machacek and Greaney (1941); McKinney (1923, 1925); Mead
(1942,0,6); Miller (1947); Mitra (1931,a,6); Mitra and Bose (1935);
Mourashkinski (1924); O'Gara (1915); Pammel, King, and Bakke (1910);
Paxton (1933); Pinck and AUison (1944); Richardson (1942); Robles
(1949)"; Roemer, et al. (1938); Russell (1943); Russell and Ledingham
(1941-); Russell and Sallans (1940); Saccardo (Sylloge, 1892, v. 10, p.
415-416); Sallans (1931,a; 1933; 1940,a,6; 1947; 1948); Sanford (1939,
1946); Sanford and Broadfoot, 1934); Sanford and Cormack (1940);
Sewell and Melchers (1924); Simmonds (1939, a,6; 1941; 1947); Sim-
monds, Russell and Sallans (1935); Simmonds and Sallans (1930, 1933,
.1946); Smith, N. J. G. (1929); Smith and Rattray (1930); Sorokin
(1891); Sprague (1938,a; 1941,d; 1942,a,6; 1943,a,d; 1944,6; 1946,c,d,e);
.Sprague and Fischer (1948); Stakman, L. (1920); Stevens, F. L. (1919,
1922); Stover (1922); Tropova (1940); Tyner (1940, 1941); Tyner and
Broadfoot (1943); Valleau (1935); Weniger (1932); Ziling (1932).
H e l m i n t h o s p o r i u m s e t a r i a e Saw.—Spot Blotch and Secondary

Root Rot
H. setariae Lind
Cochlioboliis setariae (Ito and Kuribayashi) Drechsl. fide D i c k -
son, the ascigerous stage
Conidia dark olivaceous when m a t u r e , sometimes lighter colored in

both end cells, fusiform obclavate or ellipsoidal, mostly somewhat
curved, slightly broader a t or somewhat below the middle, tapering
''(
toward both ends, 5- to 10-septate, 40-120 X ,10-18 /u.,- wall fragile.
Spores germinating by the production of two pola|- germ tubes.
Perithecia dark brownish, pseudoparenchymatous, flask-shaped,
with ostiolar beak; bodies globose or short ellipsoidal, 240-500 X 220-
315 ju; beaks well developed, paraboloid o'r cylindmcal, 60-125 X 50-
110 n- Asci numerous, fusiform, straight or slightly curved, widest
somewhat below the middle, rounded at the apex, shortly stipitate at
the base, hyaline, thin-walled, 130-150 X 22-32 ^, with 1 to 8 asco-
spores. Ascospores filiform, wavy, obtusely pointed at both ends, 5- to'
9-septate, coiled in a close helix, 200-315 X 6-7 ja, hyaline or light olive
colored in low magnification.
On: Setaria italica (L.) Beauv., Md.—5. lutescens (Weigel) F. T. Hubb,
Iowa—S. viridis (L.) Beauv., Iowa, Nebr., N. Dak., S. Dak.
The disease of Italian millet and other Setaria spp. was first ob-
served by K. Yoshina in 1906 in Japan and was attributed to a species
of Helminthosporium somewhat related to'H. turcicum Pass. K. Sawada
(1912) of Formosa described the symptoms of the disease and the diag- •
nosis of the causal fungus which he named. A detailed account of this
fungus was reported by Nisikado (1928,a) and he noted that H. setariae
Lind (1913) was the same fungus.
Ito and Kuribayashi (1931; Ito, 1930) reported that affected leaves
were brought into the laboratory in the fall of 1927 and the diseased
pieces incubated on rice-culm decoction agar in petri dishes at a tem-
perature of 25° C. Perithecia formed in about four weeks. Conidia wpre
obtained from germinating ascospores and typical lesions produced by
artificial inoculation. When seeds of Italian millet were smeared arti-
ficially with the conidia and sown in the soil, the germinating seedlings
died from a brown stem rot and abundanti conidia formed on the dead
plants. They determined that the perithecia belonged in Ophiobolus
and were similar to 0. sativus. They segregated it from that species
because of the narrower width of the conidia which otherwise were also
related in general morphology (Ito and Kuribayashi,W31). The fungus
is assigned (Dickson, 1947) to Cochliobolus seiartae~~(Ito-and Kuri-
bayashi) Drechsler in Dickson (cf. Dreehsler, 1934,b)_The present
trend toward consolidation of species would probably indicate that
this belongs under H. sativum.
In the United States this species is sonêtimes isolated from the
roots of Setaria viridis and, less frequently, S. italica in North Dakota.
Spotting of the leaves is common in Japan (1931) but it was not
noticed In North Dakota until late in the season, where it was mixed
with typical H. sativwn. The conidia are narrower in //. setariae on
North Dakota material than are the associated spores of H. sativum,
but the difference is not great. " - '
Median isolated the fungus from Setaria spp. in Iowa (letter sent
to writer) although it appeared to be less common than other forms on
this host. This was also the case at Mandan, North Dakota, where I
PUNGI IMPERFECTI—MONILIALES 383
found that much of the material obtained from Setaria and Panicum
was a species, of Helminthosporium somewhat resembhng H. turcicum.
Inoculation trials at Mandan (Sprague, 1946,e) showed that H.
setariae, when added as a sand-bran inoculum to electrically sterilized
soil, was somewhat pathogenic to Setaria spp., Echinochloa crusgalli,
Panicum miliaceum, and Bouteloua gracilis, but was less parasitic
on wheat and crested wheatgrass and only moderately parasitic on
Sorghum vulgare.
The fungus is not important in this country at present. A. G. John-
son first determined that H. setariae was present in the United States
(Haehseler, 1941).
Huang (1935) reported H. setariae on corn in China.
Range: United States, China, Denmark, Formosa, Japan, Korea.
References: Haenseler (1941); Huang (1935); Ito (1930); Ito and Kuri-
bayashi (1931); Lind (1913); Nisikado (1928,a); Padwick and Henry
(1933); Sawada (1912); Sprague (1946,e).
Helminthosporium siccans Drechsl.—Brown Blight

Producing dark brown, typically numerous, elongated spots, or
bhghting starting from the leaf tips; conidiophores olivaceous, emerg-
ing singly, less often in pairs or threes, 50-300 X 7-9 n, 1- to 9-septate;
conidia subhyaline or light fuliginous when young, later becoming yel-
low, brownish or brownish olivaceous, never dark olivaceous like H.
sativum; when fully mature provided with a moderately thick
peripheral wall, typically straight or curved, subcylindrical or tapering
slightly or more markedly toward apex, the distal segment often not
exceeding 10 JJ. diam. or rarely greater in diameter at the distal end
than at the base, 35-130 X 14-20 /x, hilum moderately conspicuous but
included within contours of the peripheral wall.
On: Lolium multifioriim Lam., Calif., D. C, Ky., Md., Va., Wash.—L.
perenne L., Md., Ohio, Ont., Oreg., Va., Wash.
The spores of this fungus^resemble those of H. gramineum. It is
very common durijigjwiriter and early spring in western Oregon and
has been found at Pullman, Washington. It is particularly prominent
on uncut lawns. There the leaves in certain areas are nearly all killed
back to their bases. The spots are numerous, dark brown, 0.2-1.0 X 0.1-
0.3 mm. and so abundant that much of the leaf surface may be covered
with a pseudoreticulate scorching, or browning.
H. siccans is one of the most important diseases of L. multifiorum
in the eastern United States (Drechsler, 1923, pp. 679-682, PL 12).
Diedicke (1903) reported a species of Helminthosporium on L. perenne.
H. siccans also occurs in Scotland (Dennis and Foister, 1942, p. 274)
and England (Sampson and Western, 1940).
Dovaston (1948) described Pyrenophora lolii from a pure culture
obtained from Helminthosporium sp., apparently H. siccans. The peri-
thecia were irregular, 300-1000 ;», diam. with a short;, conical beak, when
mature. One to fifty flexuous, tapering multiseptate, elongate setae
occur externally. The 8-spored asci measured 1721230 X 27-41 ^. The
spores were pale brown, 49-67 X 16-22 ju., usually with 5 transverse and.
2(0-4) longitudinal septa. Since he discussed bothlspot and qft blotch
symptoms there may be some question in, his mind regarding the
identity of H. siccans with the symptom complex he encountered. His
fungus produced indefinite net blotch lesions some four or five days
after inoculation of Italian ryegrass. The evidence appears rather
strong that H. siccans and P. lolii are identical. In addition, on refer-
ring to Wehmeyer's recent paper on Pleospora (1949) it appears ve,ry
clear to me that Pyrenophora lolii closely answers the description of
Pleospora trichostoma. This throws H. siccans, if such Dovaston (1948)
was dealing with, in with H. avenae, H. graniineum, H. bromi, and H.
tritici-repentis. This is a problem for a mycologist who sympathizes
with the everyday needs of the routine plant pathologist. Some sub-
dividing is called for if the lumping under one perithecial stage is
accepted.
Range: United States, Canada, Englarfd, Germany, Scotland, Wales.

References: Dennis and Foister (1942); Dovaston (1948); Dreehsler
(1923); Sampson and Western (1940); Sprague (1946,e).
Helminthosporium sigmoideum Cav. See Leptosphaeria

salvinii Catt.
Helminthosporium sigmoideum var. irregulare Cralley and

Tullis—Stem R'ot '
Hyphae white to olivaceous, septate, profusely branched, 2-5 û
diam., aerial hyphae usually scanty, ,submerged mycelium dark.
Numerous appressoria produced at times. Scleroiâ very numerous,
irregular in outline, black, surface rough, 90-119 X 26J-342,|it on leaves,
sheaths, and culms. Conidiophores dark brown, septate,'_erect, simple,
75-200 X 4-5 yu, conidia borne singly on sharp-pointed sterigmata, fusi-
form, typically 3-septate, frequently with germ tubes 2 or 3 times the
length of the spore on spores still attached to the conidiophore, inter-
calary cells densely granular, terminal cells less granular/'than inter-
calary cells, 41-58 X 9-12 ft.
On: Oryza sativa L., Ark., La., Tex.
/
This variety apparently has no connectj6n'~with Lepiosp/iaem sal-
vinv,, and differs from the sclerotial stage" of the species in having
smaller sclerotia with characteristic irregular outlines. In comparison
with the regular circular outlines of the sclerotia of L. salvinii, those of
var. irregulare appear aborted and ill-iormed (Cralley and Tullis, 1935,
• FUNGI IMPERFECT!—MONILIALES 385
Fig. 2). The conidia of var. irregulare are very similar morphologically
to those of L. salvinii. In pure culture the spores of var. irregulare are
frequently,nonseptate or tardily septate and often the cultures obtained
from germinating conidia produce sclerotia first and conidia later,
which is atypical for L. salvinii. In addition the variety is not as ac-
tively parasitic as L. salvinii, although it is by no means a saprophyte.
Nakata (1934) considered the irregular sclerotia as being the same
as those described by Cattaneo but this is not true as shown by Cralley
and Tullis (1935). Nakata described the spherical sclerotia as Hel-
minthosporium sigmoideum var. microsphaeroides Nakata which be-
comes a synonym of L. salvinii.
Range: United States, Ceylon, Japan, Philippines.
References: Cralley and'^Tullis (1934, 1935); Nakata (1934); Park and
Bertus (1934); Sakurai (1917)..
Helminthosporium sorghicola Lefebvre and Sherwin—Target Spot

Producing well-defined spots, mostly in the southern United States.
Spots small, tan or reddish purple, depending on host pigment—^tan on
Tift Sudan grass, reddish purple on common Sudan grass; older lesions
usually show target or zonate pattern, with light centers, surrounded
by narrow darker, then wider lighter bands in succession. Lesions range
iu size from barely visible to"l X 15 mm.;'these may coalesce to" involve
the whole leaf. Lesions at first round to elliptical, becoming more
elongated by delimitation caused by leaf veins. Fruiting of fungus on
leaves and glumes sparse under field conditions, abundant under moist
chamber conditions.
Conidiophores-arising singly or in groups of 2 to 4, usually from
s'tomata, or occasionally singly between ordinary epidermal cells;
typically simple, occasionally branching; dark, olivaceous brown; meas-
uring 5.5 to 10.5 /Li in diameter and 115 to 700 /x in length, sometimes
longer under very moist conditions; secondary conidiophores common,
especially under moist conditions, shorter, 5 to 7 /A diam. and 25 to
200 fi long. Conidia on Tift Sudan grass leaves in moist chamber golden
yellow to light olivaceous, 20-105 X 8.5-20.6 /x, mean 59.2 X 14.1 in,
usually curved, widest near the middle, tapering slightly toward
rounded ends; 2- to 8-septate, average 5.1; peripheral wall thin; hilum
moderately broad, not conspicuous; germinating by production of two
polar germ tubes.
On: Sorghum halepense (L.) Pars., Ga., Tex.—S. vulgare Pars., La., N. C.—
iS. vulgare var. sudanense. {Viper) Hitchc, Fla., Ga.—S. vulgare X S. vul-
gare var. sudanense (hybrid), Ga.
Lefebvre and Sherwin (1948) described this fungus as new. I t
closely matches Helminthosporium m,aydis Nisikado and Miyake. The
sorghum material has spores which are less strongly curved, smaller,
and have fewer septa than the corn fungus. The average number of
' i
septa in the conidia of the sorghum fungus is 5.1 as compared with 7.7
for the corn. ^ " j.
On cornmeal agar and potato-dextrose agar the fungus produces a
compact mass of whitish, soon light gray, or grayish mycelium,
finally grayish to olive to slate olive. Small whitish upright .tufts of
hyphae are sometimes formed within or on the border of the mat.
In their study of fungi on Sorghum spp., Lefebvre and Sherwin
(1948) examined the type of H. sorghi Schw, and could find nqftiing
except a species of CoUetotrichum on this material- and on cotype
material. H. cookei Sacc. {H. sorghi Cooke) did, not appear to be a
species of Helminthosporimn. They found that H. sudanensis Gonz.
Frag, and Cif. was similar to Curvularia geniculata (Tracy and Earle)
Boed.
This fungus should be studied for its relationship to the Cochliobolus
group.
Range: Southeastern United States. ':
References: Lefebvre (1944); Lefebvre and Sherwin (1948).
Helminthosporium stenacrum Drechsl.—Leaf Mold

On withered leaves, sporophores dark olivaceous, emerging singly or
in pairs, 80-250 X 7-10 //., 3- to 10-septate, geniculations well defined,
spores subhyaline to yellowish when fully mature, 53'135 X 15-23 /x,
subcylindrical with hemispherical or hemi-ellipsoidal ends, sometimes
widest below the middle and tapering moderately toward the ends; the
apical portion sometimes produced into a somewhat narrowed distal,
prolongation; 1- to 11-septate, nonconstricted or barely so; peripheral
wall of spore thin and including the dark hilum within its contour.
Spores subhyaline to yellowish when fully mature; 15 to 23 by
53 to 135 /*; subcylindrical with hemispherical or hemi-ellipsoidal ends,
or widest somewhat below the middle and tapering moderately toward
the ends; the apicaL portion sometimes produced iijto a somewhat nar-
rowed distal portion; 1- to 11-septate, the septa narâssociated with
constrictions, or marked by barely perceptible constrictions; the
peripheral wall thin and including the dark hilum within its contour.
Germinating by the production of germ tubes from several or all seg-
ments, the intermediate segments proliferating usually not more than
one tube, the "end segments occasionally giving rise to two or even three
tubes. -
On: Agrostis alba L., D. C, Minn., Mont., N. C.—A. oregofiensis Vasey, Ida.—
A. palustris L., Conn.—A. scabra Willd., .Cpjo., Ida.—A. tenuis Sibth.,
Conn., Oreg.—Eleusine indica (L.) Gaertff.,3Id. -,
This species is probably not as actively parasitic as H. erythro-
spilum Drechsl., in fact little appears to be known as to its importance
as a parasite. , '
H. stenacrum will usually not sporulate on nutrient agar at room
temperature although small atypical spores form on water agar.
Helminthosporium stenospilum Drechsl.—Brown Stripe

Very narrow linear brown streaks, becoming more extensive later;
conidia dark olivaceous with a thick peripheral wall, 3- to 12-septate,
average 7- to 8-septate, 40-128 X 12-22 ^ (average 84 X 17 ;».).
On: Saccharum officinarum L., Fla., Ga,, Hawaii, La.
An ascigerous stage has been developed in Hawaii and called
Ophiobolus stenospilus Carpenter (Martin, 1931). T. Matsumoto and
Yamamoto (1936) report this from Japan as Cochliobolus stenospilus
but according to Weiss (1945) neither of these names has been formally
presented. The latter is widely used, however.
Martin (1933) found that severe infection was associated with
available-phosphate deficiency. Application of phosphoric acid was
followed by a marked decrease in infection.
Matsumoto and Yamamoto (1934, plus appendix) illustrate H.
stenospilum, calling it H. ocellum but correcting this error in an ap-
pendix to their Formosan publication.
Range: United States, Brazil, Cuba, Formosa, Guam, Japan, Puerto Rico,
S. Africa, Venezuela.
•References: Drechsler (1928,a); Faris (1928,a); Martin (1931, 1933);
Matsumoto and Yamamoto (1934, plus appendix; 1936); NoUa (1949);
- Weiss (1945).
Helminthosporium teres Sacc.—Net Blotch

Pyrenophora teres (Died.) Drechsl. is ascigerous stage. (But see
recent paper by Wehmeyer, 1949, on Pleospora trichostoma (Fr.) Ces.
and de Not.)
Spots when mature composed of a dark brown network in a diffuse
brown spot surrounded by a narrow yellowish zone, conidiophores
brown or olivaceous, emerging from the stomata or between the epi-
dermal cells singly or in groups of 2 or 3y 120-200 X 7-9 ji (swollen
base is broader), stouter than H,. gramineum and H. sativum; septa-
tions in conidiophores 15-60-/* apart, averaging about 35 /x; conidia
subhyaline when young, becoming greenish fuliginous or yellowish when
older, cylindrical, thin walled constriction of basal cell somewhat pro-
nounced, proximal contour of basal cell nearly hemispherical, apical
cell similar, sometimes tapering slightly toward apex, 1- to 10-septate,
30-175 X 15-22 fj. diam., hilum contained in contour of outer wall.
. Pycnidia sometimes produced (Ravn, 1900; Smith and Putterill, 1932).
;I
Perithecia up to 500 /* diam., flattened, nearly hemispherical in out-
line, ostiolar beak short or missing, setae few' or m'oderate in number,
scattered toward the base, dark olivaceous taperiiig toward their apex
to % basal size; conidiophores often produced in large numbers on the
perithecia; asci often ill-formed, subcylindrical, the proximal portion-
tapering toward the short stipitate base and the wall of the apical end
modified by a ring-like thickening, 220-250 X 30-36 /j.; ascospoftes 8,
arranged distichously, light brown, with 3 transverse septa associated
with perceptible and often pronounced constrictions, longitudinal con-
strictions confined to one or both middle segments, contents more or
less granular and vacuolate, 52-60 X 18-22 /x.
On: Hordeum distichon L., Calif., 111., Mich.,' Minn., Wise.—H. murinum
L., Calif.—H. vulgare L., Alta., Ark., B. C , Calif., Conn., Del., Ga., Ida.,
111., Ind., Iowa, Kans., Ky., Manit., Md., Mass., Minn., Mo., Nebr., N. B.,
N. J., N. Y., N. C, N. Dak., N. S., Ohio, Okla., Ont., Oreg., Penn.,
P. E. Is., Que., R. I., Sask., S. C, Va., Wash., W. Va., Wise.
This fungus is instrumental in causing a typical reticulate blotching
on barley leaves in the Middle West and eastern states where it is
sometimes common but less often as'serious as congeneric diseases of
cereals. The only other fungus producing comparable symptoms is H.
dictyoides on Festuca elatior.
H. teres was confused with stripe {H. gramineum) in early work,
and, in fact, Saccardo's original description was not clear. Drechsler.'s
discussion (1923), while profusely wordy, clarifies the_status of H.
teres. The work by A. G^ Johnson (1914) who proposed the name net
blotch was one of the first attempts in this country to distinguish the
barley helminthosporioses and it is this work wh/ch forms the basis for
our present ability to distinguish these forms macroscopically in the
field in most instances. ;
Ravn (1900) found that the disease is carried by infected seed.
Infection of the primary leaf at germination time^wta-kes place at a
temperature of 10° C. to 15° C. in the field but at 2l)'^C.'-Ravn ob-
tained no infection. Drechsler (1923) found that much of the infection
in Wisconsin came from stubble and residue. He also indicated that
the perithecia were important in the overwintering of the disease just
as Chamberlain and Allison reported recently for P. bronii (19,44,1945).
On a rich organic medium such as potato-dextrose agar a profuse
white aerial growth results, consisting of fluffy mycelium and of more
or less compact erect columnar masses. These, according to Drechsler
(1923), represent incipient stages of sclerotia,_and ^rom Ravn's ob-
servations (1900) and those of A. G. JohnsdnT (1914), are developing
phases oi perithecia.
Range: United States, Australia, China, Cyprus, Denmark, Germany, Italy,
Mesopotamia, Netherlands, Peru, Poland, S. Africa, Tunis, U.S.S.R.
References: Atanasoff and A. G. Johnson (1920); Boening and Wallner
(1934,6); Drechsler (1923, pp. 656-663, PI. 2, 3); Geschele (1928); Ito
and Kuribayashi (1931); Johnson, A. G. (1914); Ravn (1900); Sac-
cardo (F. ital. del. 1877/86); Smith and Putterill (1932); Wehmeyer
(1949).
Helminthosporium tetramera McKinney—Root Rot and Mold

Conidiophores dark olivaceous to brown, simple or compound, with
septa 5 to 50 /i apart and with the conidia produced at irregular inter-
vals. Conidia chiefly 4-celled, borne in clusters of 2 to 3 to 50 or more,
dark olivaceous to brown, usually rather symmetrical in shape, taper-
ing toward the rounded ends, 20-41 X 10.2-20.4 /*, frequently 30-
34; X 10.2-13.6 fi.
On: Triticum aestivum L., Tex.
The writer has isolated a similar fungus from cereals and grasses in
North Dakota, South Dakota, Montana, Wyoming, and Nebraska. The
spores on potato-dextrose agar are dark olive smoky, relatively thin-
walled, regularly elongate, elliptical, 3-septate, 23-31 X 8-12 /i borne
on simple or sparsely branched conidiophores. A number of attempts
to produce root-rot lesions or even seed rot on various Gramineae with
this fungus have been unsuccessful. The local fungus appears to be
Brachycladium spiciferum according to a letter received from J. E.
Machacek in 1941. B. spiciferum was transferred to Curvularia spicifera
(Bainier) Boed., but without much justification as the regular elliptical
spores are seldom Curvularia-\ike. The fungus has small spores but
could be placed in Helminthosporium or Brc.chysporium, according to
Machacek. He believes that B. spiciferum differs from H. tetramera.
The spores of the former may be smaller than in H. tetramera, and the
nonparasitic nature of the former may indicate specific differences be-
tween the two. In our case we question if H. tetramera proper has been
found in the northern Greats Plains.
Brachycladium spiciferum is usually isolated from overwintering
roots and crowns of perennial grasses and from roots and sprouting
seeds of grasses and-cereals in the early season and late fall. It pro-
duces a luxuriant dark gray-black colony on potato-dextrose agar.
Spores (Fig. 12,B) are produced in moderate quantities but seldom in
large numbers. McKinney (1925,a) stated that H. tetramera produced
long, simple, or branched sclerotia composed of hard white pseudo-
parenchyma with an outer black layer or rind. Many hyphal strands
developed from these rinds, but no conidia were found on them. How-
ever, McKinney did have conidia in pure culture, as he stated earlier,
and there seemed to be little if any change in spore characters when
the fungus was cultured artificially. Since B. spiciferum produces a
readily growing gray-black cottony colony that resembles Alternaria
tenuis Nees and shows none of the sclerotial development of McKinney's
390 DISEASES OF CEREAI^ AND GRASSES .
•' !
cultures, the two forms are possibly distinct. Machacek mentioned in

a letter to the writer (May 2, 1941) that he had seen material on oats
and barley from eastern Canada that is closer to H. tetramera than
B. spiciferum although, as he points out, Hynes ,01937) has referred
Australian material to C. spicifera while Mason has referred it to H.
tetramera. The tendency during recent years appears to be to lump the
two species together. While no critical taxonomic report is available,
the group had been under study by Machacek. Crosier and Weimer
(1940), who also had the aid of Machacek, concluded that C. spicifera
was a saprophyte, although they did obtain mild .seedling injury on
Poa pratensis, P. compressa, P. trivialis, and Agrostis alba [A. palus-
tris). Machacek points out in his letter to the writer that in Crosier
and Weimer's paper the legends on the illustrations of C. spicifera and
C. geniculata are transposed as is obvious on reference to this paper.
In 1937 (Anon.), B. spiaiferwn was reported as an associate of R.
solani in a brown patch of lawns in New South Wales. Bensaude (1930)
obtained consistently negative pathogenicitJy results on wheat in Portu-
guese East Africa with a fungus also referred to H. tetramera. Thomas
(1939) obtained H. tetramera from rice and with it failed to obtain
infection on rice leaves, stems, andxheads nor did .it infect wheat,
barley, sorghum, Pennisetum, or Eleusine. Some further study with the
Texas material is indicated as being desirable.
Range: United States, Canada, India, perhaps elsewhere.
References: Anon. (1937); Bensaude (1930); Crosier and Weimer (1940);
Hynes (1935, 1937); McKinney (1925); Thomas (1939).
Helminthosporium triseptatum DrechsK—Leaf Mold

Conidiophores scattered on withering (leaves, singly or in pairs,
dark olivaceous, 200-400 X 6-8 /J., usually 6- to 11-septate, proliferation
of spore at apex associated with conspicuous localized thickening of
conidiophore wall resembling a fan or wing, actually a band or ring,
around conidiophore, these thickenings occurring with ct)nstnctions be-
tween, give upper portions of sporophore a more or less moniliform con-
tour. Spores dark olivaceous; ellipsoidal or short cylindrical with hemi-
spherical ends sometimes tapering more ~ or less toward basal end;
regularly 2- to 3-septate, not constricted, wall ûnusually thick although
somewhat thinner at the distal end and at the basal end diminishing
markedly in thickness toward the hilum; the hilum not projecting be-
yond contour of proximal end, 35-50 X 15-21 /* germinating at or near
hilum. /
/^- /
On: Agrostis alba L., N. Y.—A. exarata Trin1,_0reg.—Dactylis glomerata
L, N. Y.—Holcus lanatus L., Mich., N, Y., Oreg., Wash.
This is a saprophytic or weakly parasitic species causing a mold on
necrotic leaves, especially of Holcus lanatus in fog-frequented areas of
the Pacific Northwest and New York State. The fungus is readily dis-
tinguished by the thickenings on the upper part of the conidiophores.
They resemble wings or fans of growth but are ring-hke thickenings
with thin (constricted) areas separating them.
References: Drechsler (1923); Sprague (1946,e).
Helminthosporium tritici-repentis Died.—Leaf Bhght

(Ascigerous stage, Pyrenophora tritici-repentis (Died.) Drechsl. or
Pleospora trichostoma (Fr.) Ces. and de Not., cf. Wehmeyer, 1949.)
Vague to brown spots on the leaves followed by leaf withering and''
•abundant production of conidiophores which are dark olivaceous, some-
,what swollen at the base, 80-220 X 7-9 /x; conidia typically subhyaline,
straight-cylindrical, 1- to 9-septate, 45-175 X 12-21 /x diam., slightly
con^ricted in the contour of the thin peripheral spore wall. Basal spore
segment tapering to a rounded conejShape.
Perithecia produced in great abundance on dead culms of Agropyron
repens, in particular, and sometimes on leaves. Perithecia black, form-
ing in .autumn, maturing in the spring from April to June (southern
Wisconsin), basal portion in leaf material subspherical 200-350 ft diam.,
tapering apically into a short, well-defined beak, in stem material peri-
thecia often irregular, beak obscure; short dark brown setae around
ostiole, straight or fluxuose, sometimes septate or branched, sometimes
conidiophores produced in great numbers from upper half of peri-
thecium often checking further ascigerous growth, conidiophores often
suppressed when culms are covered with litter, perithecium 200-350 fx
diam., asci clavate, flattened on inner side, stipitate at base curving
toward center of base of ascus with small cap at apex of membrane,
spore distichously arranged, 8 in an ascus,. brown, 45-70 X 18-28 /<.,
3-septate transversely and 1- to 2-septate longitudinally from median
cells, constricted.
On: Agropyron intermedium (Host.) Beauv., N. Dak.—A. repens (L.)
Beauv., Manit., Mich., N. Y., N. Dak., Oreg., Wise.—A. smithii Rydb.,
Minn., Mont.,_ .N-Dak., S. Dak., Wyo.—A. subsecundum, (Lk.) Hitchc,
N. Dak.—Elymus canadensis L., Manit.—E. glaucus BuckL, Wash,—E.
hirsutus Presl, Oreg.—E. junceus Fisch., N. Dak.—Hordeum jubatum
L., Iowa—Secale cereale L., Minn.—Triticum aestivum L., Manit., Que.,
Sask.—T. dicoccwn Schrank, N. Dak.—T. durum Desf., Manit., Que.,
Sask.
Drechsler (1923, pp. 667-670, PI. 5, 6) described the symptoms on
Agropyron repens as being vague, manifested in a gradual fading and
necrosis of the leaves, which finally become gray and heavily covered
with the conidiophores and later perithecia, particularly on dead culms.
In Oregon, definite brown to fuliginous spots and streaks occur on the
leaves of A. repens and these are followed later by general necrosis. On
l\
E. hirsutus Presl the dark brown spots were abundant on all leaves of
plants growing along the Santiam River in Oregon- jn May.
In the northern Great Plains saprophytic deyelopment is some-
times great on wheat leaves during rainy periods in later season; how-
ever, little evidence has been obtained that t'he fungus is actively para-
sitic on this host in that region. On wheat the fungus is usually asso-
ciated with Septoria nodorum, S. avenae f. sp. triticea, and other fungi.
Fungi similar to H. tritici-repentis are occasionally isolated from the
roots of cereals and grasses in the northern Great Plains. Isolates from
the roots of crested wheatgrass cause pre-emergenqe injury in crested
wheatgrass, blue grama, proso millet, Italian millet, Japanese millet
(Echinochloa), and very slight root injury to wheat (Sprague notes).
Conners (19th Ann. Rpt. Can. PI. Dis. Survey, pp. 12-14, issued
1940) gave a full report of this fungus in Canada. He was inclined to
include this fungus and H. tritici-vulgaris Nis. in one species. Machacek
reported the fungus as abundant on durum and common wheats in
Manitoba (cf. Conners, 1939).
Drechsler (1923) reviewed the early history of H. tritici-repentis
which is somewhat confused.
Wehmeyer (1949) recently placed the perithecial stage of this
fungus under Pleospora trichostoma together with that of H. bromi,
H. avenae, and others. A restudy of the conidial forms is in order.
Range: United States, Canada, Germany, India, Japan.
References: Barrus (1942); Conners (1938, 1939); Diedicke (1902, 1903;);
Drechsler (1923); Johnson, A. G. (1942); McRae (1932);; Mitra (1934);
Nisikado-(1928,a); Noack (1905); Wehmeyer (1949).
, /
Helminthosporium tritici-vulgaris Nisikado—Yellow Spot
Infecting leaf blades and sheaths. Spots fusiform, with somewhat
irregular concentric zones, and yellowish halos; at first light yellowish
brown and 1 X 0.5 mm., later becoming somewha?t~-.darker, and 5-
10 X 2-4 mm.; the infected leaves dying prematurely, and"the wither-
ing proceeding from the tip toward the base. ' ~
Conidiophores arising from the infectecl tissues, usually singly and
not in groups of more than two; simple and rigid, dark olive, becoming
lighter toward the apex, 90-400 X 6-8 ju (meta 238.9 X 5.6 /*) 1- to
13-septate, the basal swollen cell being 10-15 /x wide, bearing 1-3
conidia. -
Conidia yellowish brown or light yellowish brown, thin-walled,
cylindrical, generally straight and somewhat^ curved/to one side, not
or slightly constricted at the septum; •2gf.S-183>< 8.9-21.9 /x (av.
118 X 17.3 ju.), 0- to 10-septate; the basal cell protruding to a rounded
conical end; germinating from the polar cells as well as the inter-
mediate cells.
On: Triticum aestivum L., Kans., Md., Nebr., N. Y., N. C, Penn., S. C,
Va., W. Va., possibly same as reports from Manit., Que., Sask.
A. G. Johnson (1942) identified material from Maryland and New
York (Barrus, 1942) as being caused by this species.
Range: United States, Canada, Japan.
References: Barrus (1942); .Conners (1939); Johnson, A. G. (1942); Nisi-
kado (1928,c; 1929); Raabe (1937).
Helminthosporium turcicum Pass.—Leaf Blight

H. inconspicuum Cooke and Ellis
Blighting of leaves to a general gray color followed by a grayish
green efflorescence of conidiophores which emerge in groups of 2 to 6
through the stomata. Conidiophores olivaceous up to 260 /A long, 6-9 ju.
wide, 2- to 4-septate; spores straight or slightly curved, widest near
the middle and tapering strongly toward each end, basal portion tapers
toward hilum in cone-like arrangement with basal end tending to be
rounded off, apical end rounded off but entire shape is approximately
fusoid, 45-132 X 15-25 /*, 1- to 8-septate, peripheral wall thin, sub-
hyaline, light fuliginous, later moderately fuliginous, greenish yellow,
yellowish brown, or pale olive; hilum of spore strongly protruding,
germinating by production of two polar germ tubes, one from each end,
rarely from other cells.
On: Sorghum halepense (L.) Pars., Alabama, Fla., Ga., La., Miss., N. C,
Ohio, S. C, Tex.—S. vulgare Pars., Ariz., Fla., Ga., Ill, Md., Mo., N. Y.,
N. C, Penn., S..C., Tex., Va., W. Va., Wise.—S. vulgare var. sudanense
(Piper) Hitchc, Ga., 111., La., Md., Mo., N. Mex., N. Dak., Ohio, Okla.,
S. Dak., Tex., Wise.—Zea mays L., Calif., Conn., Del., Ga., 111., Ind., Iowa,
Kans., Ky., Me., Md., Mass., Mex., Mich., Minn., Mo., N. J., N. Y.,
N. Dak., Ohio, Oreg., Penn., R. I., Tenn., Va., W. Va., Wise.
This parasite appears on the leaves in mid-late summer causing
extensive scald lesions. I have seen it on leaves of corn in the coast
region of Oregon where it was so extensive as to resemble early frost
damage. While it attacks late in the season it is, in most cases, ag-
gressively parasitic under cool wet conditions.
The fungus fruits well on potato-dextrose agar, forming a sub-
stantial mat of grayish black aerial mycelium.
Fungi referable to this species are not infrequently isolated from
the roots of various grass plants'in the' northern Great Plains. The
identity of a number of comparable isolates remains in doubt, however.
Sherbakoff and Mayer (1937) reported a black ear rot of corn in
Tennessee as due to H. turcicum but ho doubt this case represented
H. carhonum (Ullstrup, 1944). Chilton (1940) has showed that H.
turcicum is seed-borne on sudan grass and is potentially destructive.
The disease is said to be serious in the tropics (Mitra, 1923; Reinking,
394 DISEASES OF CEREALS AND GRASSES .
'I
1918) and it interferes with seed production in the'United States, par-
ticularly on sweet corn in the eastern United States!
Elliott and Jenkins (1946) set up six classes-of infection ratings
ranging from very slight to very heavy and .by field inoculations tested
the leaf blight ratings on 200 inbred lines of dent corn. Line NC 34
was outstandingly resistant. In general, the resistance of resistant lines
was transmitted to their hybrids.
Lefeybre and Sherwin (1945) found distinct racial differences in
isolates from common sudan grass, Atlas sorghum, Johnson grass, and
corn. Two cultures from sudan grass and one from sorghum were patho-
genic on these hosts but not on corn. Four out of six isolates from corn
attacked sorghum while all six attacked corn. An isolate from Johnson
grass was restricted to that host. These 10 isolates represented 4 races.
Indications are that corn readily carries the disease to sorghum; it is
transmitted less often from sorghum to corn.
Sanitary measures that reduce the anaount oi inoculum help to con-
trol the disease. Ducomet (1903) recommended wider spacing of rows
to increase aeration. Seed treatment with various reliable chemicals
kills seed-borne spores.
Range: United States, Brazil, China,-Êgypt, France, 'Guatemala, India,
Japan, Ken3'a, Madagascar, Mexico, N. South Wales, N. Caucasus, Peru,
Philippines, Puerto Rico, Queensland, Romania, Sierra Leone, S, Africa,
Tanganyika, Turkey, Uganda, U.S.S.R.
References: Campi (1939); Chilton (1940); Drechsler (1923, pp. 712-718);
Ducomet (1903); EUett (1943); Elliott and Jenkins (1945, 1946);
Gentner (1942); Lefebvre and Sherwin (1945); Mitra (1923); Nisikado
(1927,a); Nisikado and Miyake (1926,a,6); Reinking (1918); Robles
(1949); Sherbakoff and Mayer (1937); Ullstrupt (1941,fl; 1943, 1944);
Valleau (1935); Zhavoronkov (1915). | /
Helminthosporium vagans Drechsl.-^Bluegrass Purple Spot

Formitig purple black to reddish brown spots,^-8 X 0.5-3 mm.',
later becoming lighter in the center and then diffuser-brQ\vn as the
lesions progress toward the base of the plant. Conidiophores emerging
from stomata or between epidermal cells of tissue soon after death of
host cells, singly or in pairs, simple, sometimes branched, dark oliva-
ceous; 50-280 X 8-10 p, 1- to 10-septate, geniculate toward apex. Coni-
dia dark olivaceous when fully mature; cylindrical or slightly tapering
toward the hemispherical ends; 25-130 X 17-23 fi, 1- to 10-septate
(usually 5- to 8-septate), the septa not associated with constrictions
in the peripheral wall; the latter always thick and including the dark
hilum within the contour; germinating by the-production of 3 to 11
germ tubes indiscriminately from end and'^middle segments, a single
gerra^ tube usually being produced from several or all segments.
On: Poa arida Vasey, N. Dak.—P. bulbosa L., Wash.—P. compressa L.,
Ida., N. Dak., Oreg., Wash.—P. prateyisis L., Alta., Ariz., B. C , Calif.,
Conn., D. C, Ida., III., Ind., Me., Md., Mass., Mich., Minn., Mont., Nebr.,
N. J., N. Y., N. Dak., Ohio, Oreg., Penn., S. Dak., Utah, Va., Wash., Wise,
Wyo.—P. secunda Presl, Ida., Oreg., Wash. The Canadian Plant Disease
Survey reports this species on Koeleria cristata.
This spot disease is not important in the West, except possibly at
times in foggy or rainy coastal regions, but is considered important in
the Middle Atlantic States on Kentucky bluegrass pastures. Since the
fungus is favored by rainy weather it develops most actively during
late winter and early spring in the Pacific Northwest and during the
summer and fall in the northern Great Plains. At Mandan, North
Dakota, it caused moderate injury during September and October.
Horsfall (1930) found that copper lime dust inhibited this fungus:
Very little information is available on commercial control of this spot
by use of chemicals.
Range: United States, England.
References: Drechsler (1922,6; 1923, pp. 686-688, PL 15); Horsfall (1930);
Sampson and Western (1941); Sprague (1942,c; 1946,e).
Helminthosporium victoriae Meehan and Murphy—Blight

Causes a necrosis and reddening of seedling leaves, later basal ,stem
and root rot, striping of leaves, and strawbreaking; conidiophores fulig-
inous to dark olivaceous, slighj^ly curved, rounded at the base; conidia
widest near the center, tapering to a rounded tip, 40-130 X H-25 /^
(av.' 70 X 15 /i), 4- to 11-septate (average 8), walls moderately thin.
On: Agropyron cristatum (L.) Gaertn., Iowa—Avena byzantina K. Koch,
Ark., Ga., Iowa, Ky., Minn., Mo., N. Dak.—A. sativa L., Ark., Fla., Ga.,
Ida., Iowa, Ky., Manit., Minn., Mo., Nebr., N. B., N. Y., N. Dak., Ont.,
Que., S. Dak., Tex. (19 states, but not specified)—Chloris gayana Kunth,
Iowa—Dactylis glomerata L., Iowa—Hordeum vulgare L., Iowa—Paspahim
notatum Fluegge, Iowa—Phleum pratense L., Iowa, N. Dak.—Setaria
viridis (L.) Beauv.,-Iowa—Sorghum vulgare L., Iowa. Also incidental on
Soja max in Iowa.
This recently described species has suddenly appeared in epiphy-
totic form in the oat-growing areas of the Middle West, especially in
the Upper Mississippi Valley. Unfortunately it appears to be especially
aggressive on a large number of promising and also commercially ac-
cepted rust-resistant types of oats involving crosses with the variety
Victoria (Murphy and Meehan, 1946). This chain of events resulted
in a serious setback to cooperative investigations in charge of the
plant breeders. The fungus, according to Meehan and Murphy
(1946,a,6), is remarkably parasitic on oats in the vicinity of Ames,
Iowa, causing a devastating blight in field nurseries. Even a liquor
extract from the mycelium can cause death of seedlings. In fact (1947),
they state that H. victoriae is actually a weak parasite which uses a
toxic secretion to break down the resistance of the host.
396 DISEASES OF CEREALS AND GRASSES '
;' i
Because the fungus did not appear to^ fit tlie' description of any
known graminicolous Helminthosporium, the.Iow£( workers described
it as new. They found that seed from widely scattered parts of the
country was infected with the fungus. 'The writer sent Meehan pure
culture material from Mandan, North Dakota, but none of the forms
which he isolated belonged to H. victoriae except! possibly one from
roots of timothy growing in red soil in a field (Field-P) at the North-
ern Great Plains Field Station. Meehan did obtain the fungus from
oat seed grown at Fargo, North Dakota, however. Perhaps the disease
will prove less serious in the arid west but may be destructive in some
of the coastal areas where root rots and rust already make oat-growing
hazardous.
The host range at Ames, Iowa, is relatively extensive but it is in-
ferred that the locally prevalent inoculum on oats may account for the
milder attacks on other Gramineae.
The writer has not found the disease .in oats in the Palouse region
of Washington, and while he has heard that it has been reported from
the state, he has not seen material of it nor has the Department of
Plant Pathology, Washington State Colliege, any specimens which have
been mailed in by growers. Extension pathologist, M. R. Harris, has
no record of it as of this writing. The writer has not had opportunity
to search for the,disease in the western pari of Washington and Oregon,
however. The last report of the disease in the state proved to be crown
rust.
Hansing, Fellows, Johnston, and Clapp (1948) recommend, for
Kansas, seeding the following varieties of oats: Cherokee,'Nemaha,
and Clinton. Crop rotation practices affected losses, according to these
workers. In eastern Kansas, when susceptible oAt varieties were pre-
ceded by various crops, the losses were as follows: after susceptible
oats the loss in susceptible oats the secondiyear was 38 per cent; after
corn the loss was 27 per cent; after clover and meadow, 26 per cent;
after soy beans, 24 per cent; after wheat, l8 per cent, and after resist-
ant oats only 4 per cent. Hamilton and Broadfoot X'WitT) recommend
seed treatment with an organic mercury whenever varieties such as
Beacon, Garry, Vicland, and others with Victoria parentage are sown.
H. victoriae produces a dark gray or v gray black cottony growth,
which is not particularly distinguishable from isolates of H. sativum
and H. turcicum, among others. /
Miller, J. H. (1947) reported that an early winter scorching of oats
in Georgia was caused more by H. avenae than by H. victoriae al-
though the latter was found in 16 of 32 counties in the state. He men-
tioned that H. victoriae in the seedling stage^cpuld not be distinguished
with certainty except through cultures, biit'in May such fields showed
much lodging and necrosis of lower nodes and root system. No such
symptoms were found in plants affected with H. avenae^ All the at-
tacked' oat varieties contained Victoriar oat genes.
References: Elliott, E. S. (1948); Hamilton and Broadfoot (1947); King-
solver (1947); Litzenberger (1949); Litzenberger and Murphy (1947);
McLaughlin (1948); Meehan and Murphy (1946,a,6; 1947); MOler, J. H.
(1947); Murphy and Meehan (1946); Rosen (1947,a,c); Whitehead and
Dickson (1948).
Helminthosporium zeicola Stout—Corn Stalks

Caulicolous; occurring in a dark olivaceous effuse patch at and
below the first node above the uppermost roots. Sporophores super-
ficial, chocolate brown to black olivaceous, arising singly or in groups^
of two to four, usually about 6.5 /x diam., but ranging from 5.5-7.7 X
160 fi, to more than 500 /A long; up to 15-septate, or more, at intervals
of 8.5-45 /x, the spores produced at or between the septa at the apices
of well-marked geniculations; the first spore produced at 75-165 n or
more from the base and successive spores at intervals of 6.5-45 /*, the
basalend of the basal cell of the sporophore swollen as into a bulb
usually about twice the diameter of the base of the sporophore. Spores
concolorous with the sporophores to dilute-olivaceous, narrow-ellipsoid
to subcylindrical, widest near the middle or basal part, often tapering
considerably to the rounded ends, straight to slightly curved, rairely
irregular or abruptly bent, 3- to 11-septate, rarely constricted at the
septa, the hilum external and' sometimes obscure, 33-115 X 10-17 /x,
germination in the water by end cells within several hours. Type is
111. Nat. Hist. Survey Ace. No. 19, 884.
Ullstrup (1944) concluded that his H. carbonum was distinct from
H. zeicola. Little is known regarding H. zeicola other than the original
description by Stout (1930).
References: Stout (1930); Ullstrup (1941,o; 1944).
Helminthosporium spp.
It seems desirable to list some of the cases where the species name
of Helminthosporium disease fungi have not been determined. This
may, perhaps, point out unsolved problems or aid in determining the
geographical range of these fungi. In most cases the undetermined
fungi will later prove to be. referable to already well-known forms but
some of them are probably undescribed. The following are some of the
cases known to the writer:
On: 'Agropyron cristatum (L.) Gaertn., a thin-walled cylindrical spored form
somewhat Mke H. tritici-repentis but possibly distinct, from necrotic roots
at Winner, S. Dak. Possibly some of the material'we assigned to H.
398 DISEASES OF CEREALS AND GR/^SSES
tritici-repentis, which was isolated from othei Species of wheatgrass
roots, belongs here also. These latter, however, appeared to be typical for
the species even though in an unusual habitat.—Agrostis palustris Huds.
by T. Brooks from Arlington, Y&.—Alopecurus pmtensis L., a blight was
reported on this host by Weniger (Brentzel, 19p). This may be H.
sativum. I have never isolated a species of Helminthosporium from this
host.—Axonopus afjinis Chase, collected on this host by C. L. Lefebvre at
Ft. Acres, Tex.—Beckmannia syzigachne (Steud.) Fern., from West
Fargo, Bismarck, and Carrington, N. Dak., and near Moorcroft, Wyo.,
a species resembling H. catenarium Drechsl. The fungus is usually found
in ditches and is mingled with various molds, Septoria spartinae and
Physoderma sp.—Brachiaria dictyoneurum (Fig. 'and de Not.) Stapf,
collected by G. E. Ritchey, Gainesville, Fla.—Brachypodium pinnatum
Beauv., C. L. Lefebvre, Beltsville, Md.—Bromus inermis Leyss, possibly
aberrant material of H. bromi on the roots of this plant at Fargo and
McCanna, N. Dak., and at Winner, S. Dak,, was isolated.;—Bromus cathar-
ticus Vahl, from Fla. and Ga.—Calamagrosiis canadensis (Michx.) Beauv.,
leaf spot from Minn.—Cenchrus biflorus' Roxb., Gainesville, Fla.—Chloris
gay ana Kunth, Gainesville, Fla.—Cortaderia selloana (Schult.) Aschers and
Graebn., collected by Howard Johnson 'from Tifton, Ga.—Cynodon dac-
tylon (L.) Pers., from Fla. and Ga.—Ddctylis glomerata L., leaf spot from
Woodland, Wash., and from Minn., perhaps near H. catenarium. Some of
the material from Woodland, Wash., was sent to Drechsler about 16 years
ago, the remainder filed at Oregon State College.—Deschampsia danthonio-
ides (Trin.) Munro from Ford, Ida.—D. elongata (Hook.) Munro from
Camel's Hump Pass, Mont.—Digitaria sanguinalis (L.) Scop, a leaf spot
from Minn. (Plant Dis. Reptr.).—Erianthus hostii Griseb, by G. E. Ritchey,
Fla.—Eriochloa polysta.chya H. B. K. by G. E. Ritchey, Belle Glade, Fla.
—Festuca ovina L., C. L, Lefebvre collected this material at Beltsville,
Md.—F. rubra L., Va.—Gastridium ventricosum (Gouan) Schinz and
Thell, a fragment collected in the southern part di the Willamette Valley,
Greg. The small circular spots were white|with a faint vinaceous margin
(Sprague, 1M6,e) .—Hierochloa odorata (L.) Beauv., a trace of leaf rot
was found near San Francisco, Cahf,—Muhlenbergia asperifolia (Nees
and Mey.) Parodi, a leaf blotch near Ft. Ijotten, N. Dak.—Panicum capil-
lare L., Leonard, Ida., Iowa, Minn., and N. Dak., a. leaf spot. This fungus
shows some traits of H. turcicum but is, perhaps, disfinet^Similar mate-
rial on P. flexile (Gatt.) Scribn. at Tokio, N. Dak., was isolated. Some of
the isolations from P. miliaceum which the writer has assigned to H. turci-
cum may be the same as the undetermiijed material on P. capillare.-—
P. maximum Jacq., Fla.—P. purpurascens Raddi, Fla.—Paspalum almum
Chase, by C. L. Lefebvre, Ft. Acres, Tex,—P. notatum Fluegge, Fla.—
P. paniculatum L. and P., Fla.—P. urvilUi Steud., Fla.—Pennisetwrn
orientale (L.) Rich., Ga. The collections on Paspalum und .Pennisetum are
probably H. rostratum—Phalaris arundinacea L., N. C.—Phleum pratense
L., a leaf rot near Niawa, Minn. Perhaps this was H./dictyoides but the
.symptoms were obscured by associatedyôlds.—Redfieldia flexuosa
(Thurb.) Vasey, causing a necrosis of rhizomes'south of Martin, S. Dak.,
in dunes near the Nebraska line. The spores are small, brown, somewhat
capsular shaped. The fungus produces a gray, somewhat stringy growth
on potato-dextrose agar. The fungus^ may be distinct.—Schedonnardus
paniculatus (Nutt.) Trel. Isolated from roots at Long Pine, Nebr. This
is the same general form as the undetermined material on Panicum and
Setaria from Iowa, Nebr., N. Dak., and S. Dak. The spores are some-
what like H. monoceras but tend to have thicker walls. This material
has all been isolated fronl necrotic roots. It produces a nondistinctive
dark gray cottony growth on potato-dextrose agar.—Sorghum vulgare
Pers. and var. sudanense harbor the same fungus as above mentioned
but it is not common on this host.—Sporobolus neglectus Nash was af-
fected with a species of Helminthosporium causing a leaf spot.—Steno-
taphrum secundatum (Walt.) Kuntze by Lefebvre in La.—Stipa wil-
liamsii Scribn. also was parasitized at Pullman, Wash., by an undeter-
mined species of Helminthosporium, possibly H. sativum.—Zea mays L.,
a fungus comparable to H. monoceras, mentioned above on Panicum, etc.,
is rarely isolated from the roots of corn in the northern Great Plains.
In pure culture it somewhat resembles H. sativum although the mycelium
may have a stringy aspect in some sectors.—Zizania aquatica L. harbors
a Helminthosporium leaf spot in Minn.
• The above annotated records are largely taken from Lefebvre and
Johnson, H. W. (1941) and from notes by the writer (Sprague, 1942,a;
1943,a; 1944,b; 1946,e; 1947,6; Sprague and Fischer, 1948).
*The following citations are taken from the check list by Weiss
(1943, 1944, 1945) eliminating, as far as possible, all citations which
have been assignable to known species or to the above miscellaneous
list: Agropyron spicatum (Pursh) Scribn. and Sm., Mich.—A. sub-
secundum (Lk.) Hitchc, Minn.—Axonopus affinis Chase, Tex.—"Bam-
boo,"' Fla.—Beckmannia syzigachne (Steud.) Fern., Mich.—Buchloe
dactyloides (Nutt.) Englm., Ky., Tex. (but see H. buchloes)—Elymus
condensatus Presl, Mich.—Cynodon dactylon (L.) Pers., Okla., Tex.—
Dactylis glomerata L., Mass., Minn., Pa.—Digitaria sanguinalis (L.)
Scop., Minn.—Heteropogon melanocarpus (L.) Benth., Fla.—Hilaria
belangeri (Steud.) Nash, Tex.—Panicum miliaceum L., N. J., Tex.—•
Phleum pratense L., Conn., Mo., [N. Car., collected by R. P. Korf on
•mycology foray (Mycologia 41:215, 1949)]—Poa ampla Merr., Mich.
—P. nevadensis Vasey, Mich.—Setaria geniculata (Lam.) Beauv.,
Tex.—S. italica (L.) Beauv., Mo.—S. macrostachya H. B. K., Tex.—
Stipa robusta Scribn., Ky.
Preston Hated Helminthosporium spp. on the following from Okla-
homa (1945): Andropogon saccharoides Swartz; Cynodon dactylon
(L.) Pers.; Sorghum vulgare var. sudanense (Piper) Hitchc; Triticum
aestivum L.; Uniola latifolia Michx.; Zea mays L.
References: Lefebvre and Johnson, H. W. (1941); Preston (1945);
Sprague (1942,a; 1943,a; 1944,6; 1946,e; 1947,&); Sprague and Fischer
(1948); Weiss (1943, 1944, 1945).
Heterosporium avenae Oud.—Leaf Mold

Spots scattered, elongate or in series, fuscous or often vague stra-
mineous or discolored areas; conidiophores somewhat fascicled or
'!
grouped, up to 200 ju, tall, 6 j«. thick, simple, nodulose-geniculate, pale
brown, conidia at first nonseptate, later 1- to several-septate, elliptical
to oblong or cylindrical, ends rounded, muriculate io granulate-asperu-
late, light brown, 18-35 X 9-14 ju,. 1
On: Agropyron cristatum (L.) Gaertn., Sask.—Avena sativa L., Oreg., Sask.,
Wash.—Danthonia parryi Scribn., Calif.—Elymus cbndensatus Presl, Ida.,
Oreg., Wash., Wyo.—Hordeuni vulgare L., Manit., Mont., Wash.—'
Koeleria cristata (L.) Pars., Wash.—Phalaris arundinacea L., Nebr.—Poa
gracillima Vasey, Utah—P. nevadensis Vasey, Oreg.—Triticum aestivum
L., Wash., Wyo.
The collections which have been placed under this species are not
identical by any means. They represent a group of spiny-spored fungi
which have too large spores for Cladosporium herbarum and which
seem to belong to Heterosporium. They have coarser conidiophores
than C. herbarum but not all the collections have the relatively long
conidiophores cited by Oudemanns for H: avenae. Because type ma-
terial has not been available we have preferred to follow the trend and
place them in E. avenae. They should be' compared with H. maculatum
Klotzsch. which was described on Typha and Sparganium in 1832.
R. G. Atkinson (Conners and Savile, 1944) reported this species ten-
tatively on oats from Ottawa, Ontario, collected on July 10. The spots
were nearly white with a narrow orange-yellow margin, irregularly
oval to rectangular, 4-37 X 1-5-6.0 mm. The spores were finely echinu-
late, olive brown, mostly 2-celled, 9.3-16.3 X 4.6-6.5 p.; a few 4-celled,
17-24.8 X 6.2 JU,; conidiophores 55.8-96 X 3.1-4.6 fx. Lindau (Raben-
horst, 1910, V. 9, pp. 75-76) gives spores 25-28 X 12 fx-tor H. macu-
latum. The material on Elymus condensatus which often has short
stout conidiophores should be compared with HJmaculatum and prob-
ably with H. hordei Bubak. H. secalis Dippenaar (1931) from S. Africa
has larger spores, 42.5-59.5 X 6.8-9.0 /* which are 1- to 2-septate and
borne in elongated, straw-colored blotches.
We have seen material of Heterospbrium on Bromus carinatus
Hook, and Arn., Oregon, Lolium multifiorum Lam^j.. Qregon, and L.
perenne L., Oregon, which was not assigned to a species. - ~
H. avenae is a common winter saprophyte and it may possibly be
weakly pa,rasitic.
Range: United States, Canada, Netherlands, widely distributed in northern
climates but seldom reported. [ '
References: Dippenaar (1931); Lindau (Rabenhorst, 1910, v. 9); Oude-
manns (1898); Saccardo (Sylloge v. 16, p. 1065); Solheim (1934); Sprague
(1946,e). ^
Heterosporium phlei Gregorj^^—Eye Spot

Leaf spots small, scattered, oval, 0.5-1 X 1-4 mm., diam., rarely
larger, center of spot light mauve with a narrow madder'violet or dark
•\{h3-
FUNGI IMPERFECT!—MONILIALBS 401
nigrosin violet margin fading to brown or sometimes intensifying to
black; intervening tissue becomes yellow and severely infected leaves
may die. Conidiophores arising from mycelium or stomata, simple,
usually septate, primary portion reaching 325 X 6-9 jn; spores few,
cylindrical, echinulate, pale to dark brown, not constricted at the septa,
13-57 X 6-14 [J, averaging 23 X 9 ju, 1- to 6-celled, chiefly 1-septate.
On: Phleum alpinum L., Calif., Colo.—P. phleoides (L.) Karst., Wash.—P.
pratens'e L., Alta., Ariz., B. C, Calif., Conn., Ida., 111., Ind., Ky., Manit.,
Mich., Minn., Mo., Nev., N. B., N. Y., N. W. Terr., Ohio, Ont., Oreg.,
Penn., P. E. Is., Que., Wash., Wise, Wyo.
Jacques (1941) stated that Heterosporium phlei is possibly merely
a physiologic race of some earlier-described species, such as H. avenae
Oud., H. graminis McAlp., H. hordei Bub., or H. phragmitis Sacc.
Jacques (1941) found that the fungus grew rapidly on potato-
dextrose agar. The colony was bordered by a white margin, surround-
ing the olive green center, in the outer portion of which four or five
zones of different shades of green could be recognized. A certain amount
of white aerial mycelium gave the center a grayish tinge. The medium
showed a reddish or purplish coloration for some distance around the
margin of the colony.
In the original description, Gregory (1919) reported that the
fungus grew slowly in pure culture and did not sporulate until a thallus
•of considerable size was formed. Jacques (1941), working with this
strain,' found that the fungus produced spores abundantly in pure cul-
ture after only 4-5 days when the thallus was only 12 mm. diam. .
Heterosporium phlei occurs on timothy in the Far West to a limited
extent, particularly in prairie areas in shaded canyons (Idaho, Utah,
Arizona) and along the coast where the rainfall is moderate to heavy.
It is more prevalent in Minnesota and appears to be important at times
in New York and in the eastern states in general. The spores are able
to germinate at from 3° C. to 33° C , with the optimum at 24° C. Since
the disease has been noted in February, March, July, and August, the
fungus is evidently adaptable to a wide range of temperature. Horsfall
(1930) found that relatively satisfactory control could be obtained
with Kolodust and somewhat less control with 300-mesh dusting sulfur.
Range: United States, Canada, Europe, Japan.
References: Gregory (1919); Horsfall (1930, Figs. 18, 19); Jacques (1941).
Mastigosporium Riess ap. Fres.

This actively parasitic group has broa,dly cylindrical, hyaline spores
borne on short stout conidiophores. A key to the species on Gramineae
follows.
si.
402 DISEASES OF CEREALS AND GRASSES.

.' I
KEY TO THE SPECIES OF MASTIGOSEORITJM ON GRAMINEAE
V - j
A. Spores with apical appendages
M. album Rless ap. Fres'.Unot in U.,S.)
AA. Spores muticate, 3-septate , 1
B. Spores 25-32 X 4.5-5.9 n \
M. cylindricum Sprague '
BB. Spores 29-90 X 9-17 /i
M. rubricosum (Dearn. and Earth.) Nannf.
AAA. Spores muticate, 3- to 7-septate
M. deschampsiae Jorstd. '(1947) (not in U. S.)
Most keys in texts include only Mastigosporium album which is
readily traceable by its simple or forked, elongate, hyaline apical ap-
pendages, and for anyone not familiar with the large clear hyaline,
3-septate, cylindrical spores of M. rubricosum^ its determination is
likely to be difficult with any available keys.
/
Mastigosporium cylindricum/ Sprague—Brown Spot
Spots brown, elliptical to elonga1;e, finally confluent and mottled,
commonest along the sides and tips of the leaves. Mycelia mostly
endophytic, somewhat coalesced beneath the upper leaf surface, coarse,
hyaline or slightly tinted, producing short, stout conidiophores which
produce the spores by expansion of the distal portion and eventual
abscission. Spores straight-sided, slightly or scarcely constricted at the
septa, cylindrical with rounded, blunt ends, typically capsular, hyaline,
3-septate, 25-32 X 4.5-5.9 /x. / .
On: Bromus vulgaris (Hook.) Shear, Oreg. | '
This very distinct species is known only from a restricted forested
area in the Coast Range of Oregon, on Bromus vulgaris. No further
collections have been made since the writer found it in wooded areas
in the Uinpqua Valley, Oregon, in February and"TVIar^, 1939. The
type is O.S.C. 488 but 0:S.C. 482-487 are from the type-area.
Reference: Sprague (1940,6, Fig. 1,6).
/
Mastigosporium rubricosum (Dearn. and Barth.) Nannf.—
Eye Spot
Fusoma rubricosa Dearn. and Barth. ap. Dearn.
Mastigosporium rubricosum (Dearn. and Barth.) Sprague
Fusidium agrostidis Rostr. (nom-subiuidurh) ^
Mastigosporium album var. calvum Ell.'and Davis ap. Davis
M. calvum (Ell. and Davis) Sprague
M. album var. muticum Sacc.
Amastigosporium graminicola Bon.—Montev.
Forms dark purple-brown flecks on Dactylis which enlarge, become
elliptical with ashy gray to fawn-colored centers, 1-8 mm. long, vein
restricted; later lesions show various shades of gray, ashy or light
fawn color with more or less purple, red, or ochre borders (Fig. 7i,A).
On Agrostis the lesions are broader, elliptical, light brown with fawn
centers becoming an eye spot or frog-eye spot with a broad red, or red
and yellow margin. On Calamagrostis and Trisetum same as on Agros-
tis or small brown flecks, 1- to 3-mm. diam. with small gray centers,
or are sometimes surrounded by fawn or pale buff tissues. On Phleum
the spots are fawn color with vinaceous borders, that is, an eye spot.
Mycelia in host epidermal and subepidermal, endophytic, or some-
what ectophytic, septate, coarse, branched, hyaline; conidiophores
brief, single or in small groups, short, stipitate, continuous; conidia
clear hyaline, elliptical-navicular, constricted at the 3 septa, 29-60 X
9-17 ^.
On: Agrostis alba L., Oreg., Wash., Wye.—A. canina L., Oreg.—A. exarata
Trin., Oreg.—A. palustris Huds., Oreg., Wash.—A. stolonifera L., Oreg.—
A. tenuis Sibth., Oreg., Wash.—A. verticillata VilL, Oreg.—Calamagrostis
canadensis (Michx.) Beauv., Ida., Mont., Utah, Wash., Wise, Wyo.—
G...canadensis var. scabra (Presl) Hitchc, Alaska, Alta., B. C, Mont.—
Dactylis glomerata L., B. C, Oreg., Wash.—Phleum pratense L., Ida.,
Wash.—Trisetum cernuum Trin., Oreg.
The crystal-white, thick,' muticate spores with the boat-shaped
ends (Fig. 74,B) are confusing to the uninitiated who are unable to
liey this muticate form in available texts. The writer segregated this
fungus from M. album which has forked hyaline appendages on the
apex of the spore (1938,6). The earliest acceptable name proved to be
M. rubri'cosum based on Fusoma ruhricosum Dearn. and Barth. de-
scribed from material on Calamagrostis from Glacier National Park
(Dearness, 1917). M. ruhricosum is common in Europe (Lundell and
Nannfeldt, August, 1939) and in the humid parts of Oregon, Washing-
ton (Sprague, 1938, January, 1940), British Columbia, and Alaska it
is an important spot of orchard grass, redtop, creeping bent, and in
the mountains is coinmon on Calamagrostis. In Oregon the fungus on
A. alba was found to be a race distinct from that on orchard grass
(Sprague, 1938,6; 1940,6). There are also differences in susceptibility
of several species of Agrostis to the fungus (Sprague, 1938,6).
Jlf. rubricosum requires rainy or foggy weather. The spores are com-
paratively fragile and readily break down under dry or adverse con-
ditions. The fungus develops most luxuriantly during open winter
weather in Oregon and Washington and during rainy or foggy weather
throughout the growing season. In Oregon it occurs commonly on lawns
in late winter. In the Yellowstone National Park and Glacier National
Park the fungus is prolific during wet weather in August, especially on
grass growing in shaded creek valleys.
FIG. 74.—Mastigosporium rubricosvm: A, eye spot lesions on leaf of Daclylis

glomerata, Oregon; B, conidiuni fiom Daclylis, X2000.
M. rubricosiim produces a slow-growing, leatliery mat on potato-

dextrose agar. The surface becomes somewhat folded and mounded
and pebbly or roughened by the formation of small partially embedded,
stromatic or sclerotic mycelia that resemble pycnidia. The color ranges
from tan to fawn with a pale amber substrate. Normal-appearing
conidia were produced on potato-dextrose agar after several weeks in
an electric refrigerator. After growing several months at this tem-
perature the colonies became orange brown and the sclerotia increased
in number, frequently forming rings in the substratum around the
original point of transfer.
A culture of M. album, the species with forked appendages, from
Alopecurus pratensis from Aberystwyth, Wales, had the same general
yellow-brown color of M. rubricosum but produced masses of conidia
in glistening orange-brown mounds on the surface of the colony. After
transfer to potato-dextrose agar the sporulation continued but was at
first putty-colored, later darker. Strangely enough this species has
never been found in North America although it is very common in
similar areas in northern Europe (1938,6).
Inoculation trials showed that M. rubricosum from orchard grass
readily attacked that host but not Agrostis, Avena, Deschampsia
caespitosa, Holcus, Lolium, and Poa pratensis. An isolate from redtop
was mildly parasitic on A. palustris, strongly so on 4 . tenuis, weakly so
on orchard grass, and negative on the others. M. album would attack
only Alopecurus, Brachypodium, and Deschampsia (Sprague, 1938,6).
Recently J0rstad described M. deschampsiae (1947) with muticate,
4- to 7-septate spores, 40-68 X 15.5-20 /A. Deschampsia is a host for
M. album and' collections are available (1938,b) but apparently it is
not a host for M. rubricosum. Some spores of M. deschampsiae resemble
pointed spores of M. album without their apical appendages, but others
have the rounded boat shape of M. rubricosum except for their multi-
septations.
Spores of M. rubricosum develop from expansion and elaboration
by hyphal tips accompanied by two nuclear divisions to produce 4 cells.
Germination is from the apical end, producing one or two stout germ
tubes which soon branch repeatedly.
In 1947 two collections of M. rubricosum were obtained on timothy,
one in, a meadow at Laird Park, Idaho, and another on a mist-covered
cliff from Snoqualmie Falls, Washington. Whether these represent a
new-race or had escaped observation because of resemblance to Hetero-
sporium spot is not certain. J0rstad reported M. rubricosum from
Norway on timothy (1945). English material of M. album used by the
writer (1938,b) would not infect timothy in Oregon, but unfortunately
no trials with M. rubricosum were run on this host at the time. Further
study is therefore needed.
Range: ' United States, Canada, England, France, Germany, Norway,
Sweden, U.S.S.R., Wales.
References: Bondarzeva-Monteverde (1921); Davis, J. J. (1903); Dear-
ness (1917); Dennis and Foister (1942, p. 276); Jones (in Conners, of.
19th Ann. Rpt. Canad. PI. Dis. Survey, 1939); J0rstad (1945, 1947):
Lundell and Nannfeldt (1939, p. 32); Sampson and Western (1941);
Sprague (1938,6; 1940,6);-Wakefield (1918).
Napicladium arundinaceum (Cda.) Sacc.—Black Leaf Mold

Helminth asp orium arundinaceum Corda
Scolecotrichum roumegueri Cav. (cf. V. Hoehnel, 1923)
Causing long-olive-velvety black streaks or extensive areas on
leaves, often accompanied by drying, yellowing, and death of leaves
or shoots, conidiophores short, fascicled, rounded or thickened at the
base, 1- to 2-septate, 50-60 X 8 /*, olivaceous; conidia obclavate, 40-
45 X*18 /*, mostly biseptate, pale olive.
On: Phragmites communis Trin., Manit., Minn., Nebr., N. Dak., S. Dak.
Large streaks or coalesced areas coyered with black mold form on
the leaves, progressing from the tip downward. Yellow or necrotic areas
may be extensive. Either the fungus is systemic or it produces toxic
material which kills the host. The invaded leaves often fire, turning
straw-colored or yellow.
* J
There are comparatively few specimens available of this fungus
but it is relatively abundant in the lakes fegionj of 'Minnesota and
adjacent areas. It is nearly always present on reeds along the shores
of Devil's Lake, North Dakota. '
Range: United States, Austria, Belgium, Czechoslovakia, England, France,
Germany, Hungary, Italy.
References: Corda (1839, v. 3, p. 10, Fig. 25); Saccardo (Sylloge, v. 4, p.
482); Sprague (1946,e).
Nigrospora oryzae (Berk, aijd Br.) Fetch—Dry Rot of Corn

Monptospora oryzae Berk, and Broome
Conidiophores erect; stiffly flexuose, apices briefly branched, spores
subglobose to definitely flattened, 12-15 ju. diam., sometimes 11-12 /i
vertical diam., black, opaque. , •
On: Saccharum officinarum L., general—Zea.mays L., Conn., 111., Ind., Iowa,
Kans., Manit., Mass., Miss., Mo., N. Mex., Ohio, general in central and
eastern states.
The reader is referred to Mason (1927) for a discussion of this
species and also of N. sphaerica. The latter has slightly larger spores
than N. oryzae. The two have been described under a considerable
number of species, most of them in different genera. Possibly the com-
monest synonym is Basisporium gallarum Molliard (1902) which
Mason placed under iV. oryzae. In addition to the article by Mason it
is noted that Saccardo (Sylloge, v. 25, p. 775, 1930) gave a list of
synonyms for iV. oryzae and N. sphaerica. j
The disease condition caused by this fungus is confined to a sapro-
phytic or weakly parasitic dry rot of the cob in late season or at har-
vest time. The cobs tend to shred at the base or shank and sometimes
the kernels are shriveled. Infected ears show black spores around the
area where the ear was broken away from the shankôr at the base of
kernels. Yellow kernels are often bleached or discolored^yellow brown.
The fungus may attack injured seed and rot it in the soil if the corn
seed is planted at a time when the soil is cold and wet. Alfaro (1946)
reported wheat jointly injured by Pediculopsis mite and A^. oryzae in
Spain. •• (
Range: United States, Australia, Barbados, El Salvador, Gold Coast, India,
Italy, Kenya, N. Caucasia, Palestine, Peru, Romania, Spain, Tanganyika,
Uganda, .W. Siberia. ^
References: Alfaro (1946); Berkeley and Broome (1875); Butler (1947);
Purrell (1925,b); Koehler (1942); Mas,ccnrCl927)'; Molliard (1902);
Fetch (1924); Reddy (1933); Savulescu and Rayss (193i; 1932); Standen
(1944, 1945).
FUNGI IMPERrECTI—MONILIALES 407
Nigrospora sphaerica (Sacc.) Mason—Dry Rot of Corn

Conidiophore short, grouped and arising from a small stroma;
spores flattened globose, 16-18-(20) ju, black opaque.
On: Avena sativa L., Sask.—Sorghum vulgare Pars., Okla.—Stenotaphrum
secundatum (Walt.) Kuntze, Fla.—Triticuni aestivum L. Sask.—Zea mays
L., in the, central and eastern United States and south to Okla. Reported
as an ear rot and stalk rot in Ontario.
This is said to be more common in the Middle West than N. oryzae
but often the two species are not distinguished or for practical purposes
are referred to one species, usually JV. oryzae. The available descrip-
tions which can be relied on are meager, as indicated above. N. sphae-
rica has larger spores than N. oryzae. Mason also lists A^. sacchari
(Speg.) Mason which has still larger spores, 20-22 X 15-18 [i. This is
derived from Glenospora sacchari Speg. (1896).
N. sphaerica was originally described as Trichosporium sphaericum
Sacc. (1882) from material sent by Ellis from New Jersey (N.A. Fungi
968, on Zea). Coniosporium gecevi Bubak (Bubak and Kosaroff, 1912)
was referred to JV. sphaerica by Mason (1927) as was Hadrotrichum
arundinaceum Cooke and Massee.
Range: United States, Bulgaria, Burma, England, Gold Coast, India,
Tanganyika, Uganda.
References: Arzberger (1913); Bubak and Kosaroff (1912); Butler
(1913,6); Cooke (1887); Mason (1927); Massee (1893); Miyake (1910);
••Saccardo (Micheha, 1882); Spegazzini (1896).
Ovularia hordei (Cav.) Sprague—Leaf Spot

Ophiocladium hordei Cav.
Straw-colored to buff, later pale brown spots, amphigenous, elon-
gate, striate, often 10r30 X 1-2 mm., finally more or less confluent and
covering or killing most of the leaf surface. Spots emarginate or sur-
rounded by a diffuse yellow buff area, finally the lesions covered with
a chalky fruiting material of the fungus. Conidiophores in compact
hyaline fascicles in neat rows between the veins of the leaf, emerging
from stomata and arising from abundant yellow to mostly hyaline
compacted or serpentinous conidiophores, 20-40 X 3-4 /i, tortuous,
twisted, or remarkably serpentinous, particularly at the apex. Spores
produced near the apex on the side of the conidiophores and on break-
ing off, a slight hilum is left at the base of the elliptical to elongate-
ovate spores. Spores hyaline, nonseptate with an evident wall which
is faintly roughened, 11-15.5 X 6.0-7.8 /*.
On: Festuca kingii (S. Wats.) Cassidy, Colo., Utah—Glyceria grandis
(S. Wats.) Wise.—Lolium multiflorum Lam., B. C.—L. perenne L., B. C,
Oreg.—Phalaris arundinacea L., Ida., Mont., N. Dak., S. Dak., Wise,
Wyo.
J0rstad (1930) mentioned that Ophiocladium hordei produced dark
spots, bearing a white or pale pink' coating produced by the conidio-
phores on barley leaves. While workers in this country have assigned
the material on Phalaris to 0. hordei it is surprisiiig that barley has not
been found parasitized by the fungus in the United States. The fungus
on Phalaris is common along creeks in the Black Hills of South Dakota
and Wyoming and has also been found as far west as Priest Lake,
FIG. 75.—Fascicle of conidiophores and attached conidia of Ovularia hordei

on Phalaris arundinacea. (From Mycologia.) I
Idaho (Sp'rague, 1950;b). Davis (1932, 1942) and Gfeene41942, p. .89)

have found it in Wisconsin where it is apparently not rare o"n Phalaris.
We have assigned this species to Ovularia because" the genus
Ophiocladium, does not appear to be fundamentally distinct from Ovu-
laria. Cavara described 0. hordei with spores only 6-8 X 4.5 {x, but with
the characteristic conidiophores and spores ['with evident -walls. We
found spores 11-15 X 6-7 /x. Another species on Phalaris, 0. baldingerae
Eliasson-which seems to belong in 0. hordei, has intermediate spores
9-12 X 6-7 IX. These measurements are very close to 0. pulchella var.
agropyri Davis, 9 - 1 2 x 6 - 9 ju, (1919,6). Davis, later apparently dis-
carded this variety (1942). It appears difficult,- in-some instances, to
segre<.fate 0. hordei from 0. pusilla (pulchella) on the basis of spore
size. The tortuous aspect of the conidiophores of 0. hordei plus the
evident spore wall (Fig. 75) is of considerable help, however. Some
material on Festuca kingii (Sprague, 1948,c) from Logan, Utah, is less
helpful. The conidiophores on this host are only moderately serpen-
tinous to nearly straight but 30-40 X 3.5 /t and hence similar in size
to the Phalaris material. The spores are even larger, 16-22 X 9.5-10.5 fj.
and, in addition, are definitely more echinulate than the faintly muri-
culate ones on Phalaris. It was not believed that they should be placed
in Ramulaspera, however. 0. lolii Volk. is assigned to 0. hordei, but
see discussion by Sprague (1948,c) regarding this group.
Range: United States, Canada, Italy, Norway,
References: Cavara (1893); Davis (1932, 1942); Greene (1942); J0rstad
(1930); Saccardo (Sylloge 1895, v. 11, p. 587; 1906, v. 18, p. 531);
-Sprague (1946,o; 1948,c).
Ovularia lolii Volkart—Eye Spot of Lolium

See also 0. hordei (Cav.) Sprague
The conidiophores of this species were up to 80 /x long but other-
wise it seemed assignable to 0. hordei (Sprague, 1948,c). It may be a
developing phase of Ramulaspera holci-lanati (Cav.) Lindau (cf. Ovu-
laria holci-lanati Cav.). This species in addition to its definitely spiny
spores has conidiophores up to 170 [t, long (Lindau in Rabenhorst
Krypt. Fl. 1907). 0. pulchella var. lolii-italici Ferr. may also belong
in this complex. We have collected poor material on Lolium marschallii
Stev. in Oregon which we assigned at the time to 0. pulchella because
the spores were smaller than 0. lolii Volk. (Sprague, 1935,c). We have
left this in 0. pusilla {pulchella) but perhaps this too needs restudy.
This group should be critically studied in an area where fresh viable
material is available.
Marchionatto (1947) reported 0. lolii from Argentina. It produced
elongated (2- to 3-mm.) discrete, brown lesions with yellow, shaded
margins. The caespitose, cylindrical, erect hyaline conidiophores were
60-80 X 4-5 [I and the spores were oval or subpyriform, 18-19 X H -
13/..
Range: Argentina, Europe, possibly in Oregon.
References: Lindau (in Rabenhorst, 1907); Marchionatto (1947); Sprague
(1935,c; 1948,c).
Ovularia pusilla (Ung.) Sacc. and D. Sacc.—Leaf Spot

0. pulchella (Ces.) Sacc.
Spots elliptical to round, somewhat elongate at times, on fescue
grasses, spots brown or tan on the stem, sheath, or leaves; on Agrostis,
Lolium, and Holcus spots are ashy with red borders (eye spot); on
brome the spots are minute, 1-2 mm., round ashy with brown borders
(round eye spot). Conidiophores hyaline, fascicled, 4 0 - 6 5 x 2 - 3 /j,;
conidia acro-pleurogenous, crystal hyaline, few, ovoid, 8-13 X 6-10 /..
On: Agropyron cristatum (L.) Gaertn., Colo.—A. repehs (L.) Beauv., Ont.—
A. trachycaulum (Lk.) Malte, Wise.—Agrostis alba'L., Oreg., Utah, Wise.
-A. humilis Vasey, Wyo.—A. oregonensis Vasgy', Mont.—A. palmtris
Huds., Oreg.—A. tenuis Sibth., Utah^-AZopecwrus alpinus J. E. Smith,
Mont.—Arrhenatherum elatius (L.) Beauv., Ore •Bromus carinatus
(Hook.) Am., Ida., Wash.—B. inermis Leyss, Ida'., Wise—Calamagrostis
canadensis (Miehx.) Beauv., Mont.—Elymus glaiicus Buckl,, B. C.—
Festuca idahoensis Elmer, Ida.—F. megalura Nutt., Oreg.—F. myuros L.,
Oreg.—Glycerin data (Nash) Hitchc., Wash.—G. erecta Hitchc., Calif.—
Holcus lanatus L., Oreg.—Lolium marschallii Stev., Oreg.—Poa ampla
Merr., Ida., Wyo.—P. longiligula Scribn. and Wils., Wyo.—Trisetum spica-
turn (L.) Rieht., Wyo. .
,0n most hosts this widely distributed species is usually limited to
a few small scattered spots. In addition to the fact that the spots are
scanty, the fascicles of conidiophores are difficult to see, even with a
low-power binocular, and the spores are few. The crystal white, egg-
shaped spores which average 8-12 ^ long- are so typical, however, that
the presence of only one or two is a clue that 0. pusilla is present.
While most of the material, as stated/has few lesions on the leaves,
species of annual fescue growing in thick stands in sandy soil or in
open wooded areas in the Willamette-Valley, Oregon; sometimes have
a considerable percentage of the plants infected. The infection on
Bromus inermis at Lawyer Canyon State Park, Idaho, had relatively
numerous spots of a characteristic round eye-spot appearance.
As pointed out recently (Sprague, 1948,c), 0 . pusilla is apparently
an earlier name than 0. pulchella, dating back to Ramularia pusilla
Ung. described in 1833 (linger, 1833). Saccardo and D. Saccardo (Syl-
loge, v.-18, p. 531, 1906) state that the spore dimensions of 0. pusilla
as based on an illustration by Unger (1833, Fig. 10 of Table II) are
8-10 X 5-6 f*. I
0. pulchella (Ces.) Sacc. (Sylloge, 1886, v. 4, p. 145), which we
call a synonym of 0. pusilla, was originally described as Ramularia
pulchella Cesati (1853) and its description later amplified by Fresenius
(1863) as causing a purple leaf spot of Dactylis glomeratn.-Srhis host,
strangely enough, has not been noted in the Pacific North.west where
the species is common.
Davis described 0. pulchella var. agropyri on A. trachycaulum from
Wisconsin (Daias, 1919,5) but in his final work he placed this under
the species 0. pulchella without differentiating it. As we have pointed
out under 0. hordei, the spore measurements of his variety are close to
those of 0. hordei (Davis gave the spores as 9-12 X 6-9 fx.). While we
have listed this material under 0. pusilla it should be-' critically com-
pared with the two species. We have seen the-fype -'of Davis' fungus
but that was a number of years ago before we had the opportunity of
seeing the several western collections which display a divergent mor-
phology.
Weiss (1945) listed Ovularia sp., on Panicwm hemitomon Schult.,
perhaps a Sporotrichum.
Range: United States, Austria, probably Canada, Germany, Italy.
References: Cesati (1853); Davis (1919,6; 1942); Fresenius (1863); Sac-
cardo (Sylloge, 1886, v. 4, p. 145; 1906, v. 18, p. 531); Sprague (1948,c;
1949,a); Unger (1833).
Papularia sphaerosperma (Pers.) von Hoehnel—Mold

Fruiting surface forms under the epidermis which is forced aside,
becoming somewhat elongated sporodochia, 1-2 X 0.5-0.75 mm. diam.,
conidiophores variable, in culture 10-30 X 4-6 /x; conidia 1-celled, len-
ticular, black with a narrow hyaline rim around the periphery; spores
7-12 /t diam. in face view, 7-12 X 3.5-6 fi side view.
On.: Arundinaria gigantea (Walt.) Chapm., Gulf States—Arundo donax L.,
Calif.—Bambusa sp., Calif., Fla., Ga., Tex.—Cortaderia selloana (Schult.) ,
Aschers. and Graebn., Calif.—Cynodon dactylon (L.) Pars., Miss.—Sac-
charum ojficinarum L., La.
Mason (1933) gives the most useful discussion of this group which
is represented by the species Papularia sphaerosperma. Coniosporium
Link has multiseptate spores and Coniosporium Sacc. must be referred
to Papularia. The genus has the characteristic lens-shaped spores which
appear strongly flattened when viewed from the side. The sphaero-
sperma group is widely distributed in the tropics on a large number of
hosts. C. shiraianum (Sydow) Bubak has been reported from this coun-
try but this appears to be just another synonym of P. sphaerosperma.
It is doubted if this group is very parasitic, if at all. All the forms listed
by Weiss (1943, 1945) are listed together here. No doubt the saprophy-
tic host range of this species is far greater than given.
Range: General in the^tropics, and in Europe in some places.
References: Lindau (in Rabenhorst, 1907); Mason (1933); Sydow (1899).
There are numerous other citations which are listed in part by Mason,
Saccardo (Sylloge), and Lindau.
Penicillium expansum (Lk.) em. Thorn—Blue Mold

Colonies on gelatin, potato, or bean agar, green, becoming gray
green and slowly brown after several weeks (especially when exposed
to light), with concentric zones tufted with short, loose coremium-like
aggregations of conidiophores, not over 1 to 2 mm. in height except in
old cultures containing sugar, broadly spreading with broad white
margin in growing colonies; reverse somewhat brown, conidiophores
either very short lateral branches of aerial hyphae or very long (1 mm.
or more), arising singly or grouped with others to form coremia; peni-
cilli consist of 1 to 3 main branches bearing verticils of branchlets
supporting crowded whorls of sterigmata with cohidial chains totalling
130 to 200 X 50-60 fi. at base in cultures withou|. sugar. With sugar
continuing for some weeks to produce great numbers of conidia which
come to form masses perhaps 1 mm. iii thickness! sterigmata 8-10 X
2-3 /A; conidia elliptical to globose 2 X 3.3 n atlfirst, later 3-3.4 ju,
green, homogenous persisting in chains when mounted; colonies begin
to liquefy gelatin slowly after about ten days and continue until it is
completely liquefied. The fungus grows readily and rapidly upon all
common media.
On: Agropyron cristatum (L.) Gaertn., N. Dak., Oreg., S. Dak., Wash.—
Avena sativa L., Colo., Ida., Iowa, Minn., Mont., Nebr., N. Dak., Oreg.,
S. Dak., Wash., Wye.—Hordeum vulgare L., Ida., III., Iowa, Minn., Mo.,
Nebr., N. Dak., Oreg., S. Dak., Wash., Wise, Wyo.—Panicum miliaceum
L., Minn., Mont., N. Dak., S. Dak.—Saccharum officinarum L., Hawaii
(on seed canes); La.—Sorghum vulgare Pers., Colo., III., Ind., Iowa,
Kans., Mo., Mont., Nebr., N. Dak., S., Dak., Wash., Wise.—Triticum
aestivum L., Ida., 111., Ind., Iowa, Minn.,' Mo,, Mont., N. Y., N. Dak.,
Ohio, Oreg., S. Dak., Wash., Wise—T. d;u.rum Desf., N. Dak., S. Dak.—
T. spelta Desf., N. Dak., Oreg., Wash.—r. timopheevii Zhuk., N. Dak.—
Zea mays L., general and in Ontario.
This, and probably a considerable number of other species of Peni-
cillium, occurs on the seeds of many plants, of which the above are but
some of those which appear to be definitely damaged by this sapro-
phyte. The starchy endosperm is reduced in amount, resulting in weak-
ened seedlings. It is possible that by-products of saprophytic decay kill
young seedlings in the same manner reported under P. oxalicum where
oxalic acid kills the plant's tissue in advance of the actually sapro-
phytic organism (perthophyte). /
Most of the injury caused by blue molH occurs in storage on seed
which has been exposed to moisture or has. been improperly cured and
has sweated in storage. In the field most of the injury occurs on planted
seed in the soil during prolonged rainy periods in April or May (and
sometimes' even in June) in the northern Great Plains,^()r earlier in the
Pacific Northwest. Semeniuk, Nagel, and Oilman (1947r>eported on
the development of mold on stored yellow dent shelled corn in large
steel bins with a capacity of 1,000 to 2,740 bushels. The early invaders
were Penicillium politans of the blue-eye fungi group with association
of Aspergillus spp. Gentner (1925) listed various molds on oats, in-
cluding P. glaucum, but stated that invasion probably followed frit fly
(Oscinis frit) damage.
Milner, Christensen, and Geddes (1947) report on "sick" or germ-
damaged wheat seed. Penicillium sp. caused 20 per cent of the storage
damage in wheat from Montana, North D^Kota, South Dakota, and
Minjlesota but Aspergillus glaucus was involved in 60 per cent of the
loss.
Seed treatment (New Improved Semesan Jr.) will help reduce the
endosperm-destroying molds. Proper curing of corn seed is necessary.
FUNGI IMPERFECT!—MONILIALES_ 413
In the drier parts of the northern Great Plains, seed molds usually are
less important and if clean local seed is used, seed treatment of field
corn does not often show any benefit. However, sweet corn and pop-
corn usually show definite benefit from seed treatment in this area.
The slurry technique using Arasan-M has shown promise in reducing
seed rot.
For a critical taxonomic study of the blue molds the reader will, of
course, wish to refer to Thorn (1930).
Range: General.
References: Diachun (1939); Hurd (1921); Leukel and Martin (1943);
Marchionatto (1942); Noble (1924); Noll (1936); Rands (1926);
Thorn (1930).
Penicillium oxalicum Currie and Thom—Green Mold Necrosis

_A soil-inhabiting species producing oxalic acid rapidly in sugar
media. Colonies ivy-green, velvety, broadly spreading, with reverse
pale yellow, and agar uncolored or only slightly colored; conidiophores
up to 200 [1. by 3.3-5.4 ju, producing a verticil of 2 to 3 metulae, verticils
of closely parallel sterigmata 10-14 X 2.5-3.5 z^; conidia cylindrical
then elliptical, uneven in size from 3 X 2 /A when first formed to a
maximum of 5 X 3.5 ^ when ripe and becoming subglobose, 7-10 j«.
diam. in germination, in long chains packed in close columns. In
favorable media producing great masses of conidia which break off
readily. Variants are discussed by Thom (1930).
On: Zea mays L., D. C, 111., Ind., Iowa, Ky., Ohio, Va., Wise.
This species invades the seed embryo, causing yellowing of the
leaves of the seedling and a gradual dying and drying of infected
seedlings before the sixth leaf has unfolded (Johann, 1928; Johann,
Holbert, and Dickson, 1931). The dead plants, which remain stiffly
upright, show little or no decay of the rool.s. The fungus is essentially
a saprophyte and would technically be called a perthophyte. The in-
jury is said to be due apparently to oxalic acid, which kills the plant
tissue in advance of the fungus.
Thom (1930) verified the identity of P. puberulum Bainier which
was isolated from rotting corn at Lincoln, Nebraska, by F. D. Heald.
He gave an emended description of this fungus (1930, p. 272).
Range: United States, Manchuria.
References: Currie and Thom (1915); Diachun (1939); Johann (1928);
Johann, Holbert, and Dickson (1931); Thom (1930).
Periconia circinata (Mangin) Sacc.—Milo Root Rot

Aspergillus circinatus L. Mangin
Conidiophores filiform, fuliginous, gregarious, septate, erect, 150-
300 fi long, expanded somewhat at the top which is obtuse and 1-2-lobed,
conidia in short chains in dense black heads, yoiing spores smooth but
mature ones minutely muriculate to "spiny,. spherical, fuliginous to
black, 15-27 ft diam., muriculations 0.5 /x long.
On: Sorghum vulgare Pers., Md., Minn.,'Mont., N, âk'., S. Dak., Wyo. In
addition to the reports on sorghum the writer has isolated a species of
Periconia with coarser spores from the following plants: Agropyron cris-
tatum (L.) Gaertn., Mont., N. Dak.—A. inerme (Scribn.'and Sm.) Rydb.,
Wash.—A. intermedium (Host.) Beauv., N. Dak.—A. michnoi Roshev,
N. Dak.—A. semicostatum (Steud.) Nees, N. Dak.—A. trachycaulum
(Lk.) Make, N. Dak.—Agrostis cernua, N. Dak.—Andropogon hallii
Hack., N. Dak.—A. scoparius Michx., Minn., S. Dak.—Arthraxon hispidus.
var. cryptatherus (Hack.) Honda, N. Dak.—Avena fatua L., Wash.—
A. nuda L., N. Dak.—A. sativa L., Minn., N. TiAkr—Bromus inermis
Leyss, N. Dak..—Calamovilfa longifolia (Hook.) Scribn., Mont.—Dac-
tylis glomerata L., N. Dak.—Elymus dahuricus Turcz., N. Dak.—E.
juncetis Fisch., N. Dak.—E. mlsuginosus Turcz., N. Dak.—Eragrostis
cilianensis (All.) Lk., N. Dak.—Festuca. elatior L., N. Dak.—F. octofiora
Walt., S. Dak.—Hordeum brevisuhulatum (Trin.) Lk., N. Dak.—H. vul-
gare L., N. Dak.—Koeleria cristata (L.) Pers., N. Dak.—Panicum milia-
ceum L., N. Dak.—Phleum pratense L., Mont., N. Dak.—Poa juncifolia
Scribn., N. Dak.—Setaria italica (L.) Beauv., N. Dak.—S. viridis (L.)
Beauv., N. Dak.—Sorghastrum nutans (L.) Nash, "N. Dak.—Sorghum
halepense (L.) Pers., N. Dak.—^S. vulgare var. sudanense (Piper) Sitchc,
N. Dak.—Triticum aestivum L., N. Dak.—T. dicoccum Schrank, N. Dak.
—T. durum Desf., N. Dak.—Zea mays L., N. Dak. The fungus was also
isolated from Medicago sativa at Mandan, N. Dak.
Leukel and Pollack (1947) first reported evidence that P. circinata

was the cause of the disease of mile sorghum which was originally
thought to be caused by Pythium arrhenomanes (cf. P. graminicola).
Still more recently Leukel (1948) and iLeukel and Johnson, A. G.
(1948) further substantiate the findings.'
P. circinata was described on wheat;in France (Mangin, 1899,6).
It was originally considered to be one of the soil-borne black molds
with no parasitic ability. Stevenson and Imle (194^reported a locally
severe leaf spot caused by P. heveae Stevenson and Tiiile-as a parasite
of rubber plants in Costa Rica and Mexico. A number_of species of
Periconia have been described from decayed roots of plants but no
evidence of their pathogenicity is available. I ran one series of soil-
borne inoculations with an isolate of P. sp. on cool-temperature grasses
and cereals in a cool greenhouse at Mandan, North Dakota, a number
of years ago but at the temperature prevailing the isolates were non-
parasitic to the seeds and sprouting seedlings in the flats. As shown by
the above list Periconia sp. was isolated from the roots of a number of
Gramineae in the northern Great Plains and-in adjacent regions during
the years 1940-46 (Sprague, unpublished reports). This fungus, which
Leukel has found distinct from P. circinata, was rarely isolated as early
as May, sometimes in June, more often in July and August into Sep-
tember. The plants which harbored the fungus included such different
hosts as proso millet, oats, wheat, and sorghum. The fungus was never
found in any great abundance in the area and in most years it averaged
from 0.3 to 1.9 per cent of the season's total. In August, 1944, it was
isolated 75 times out of 520 pure cultures of all fungi obtained that
month. In one case it was isolated several times from a red root condi-
tion in Freed sorghum at Mandan. But in general our isolations were
scattered, often one pure culture from a collection mixed in with other
fupgi. In 1942 we isolated it twice at Pullman, Washington, during
cool weather in May. In 1947 no pure cultures were obtained of the
species at Pullman but no isolations were made in June, July, and
August from likely material so that its absence is not significant.
Zeman (1939) reported Periconia pycnospora Fres. on Phalaris
bulbosa from Argentina. Glynne (1939,c, p. 210) reported P. circinata
on wheat roots from England but no further data were reported.
Range: United States, France.
References: Glynne (1939,c); Leukel (1948); Leukel and Johnson (1948);
Leukel and Pollack (1947); Mangin (1899,6); Quinby and Karper (1949).
Piricularia grisea (Cke.) Sacc.—Blast, Gray Spot

Spots gray or dirty yellow, ashy with darker brown or purple or
water-soaked borders, elliptical to nearly round, 1 to 5 mm. diam.,
conidiophores gray or tinted, septate, basal cell somewhat swollen,
simple or sparingly branched, penetrating through stomata in clusters
of 2 to 5, geniculate 60-120 X 4-5 /x; conidia single, fuscous, terminal
in scorpioid cymes, ovate, 2-septate with the apical cell cone-shaped or
slightly rostrate, conidia broadest at lower septum, base with slight
hilum-like terminal where it breaks from the stalk, 18-22 X 9-11 jn.
On: Agrostis stolonifera L., Tex.—Cynodon dactylon (L.) Pars., Okla.—
Digitaria sanguinalis^ (L.) Scop., general in the south and north to at
least Mo., Okla., Wise, on this host and probably indistinguishable from
P. oryzae—D. serotina (Walt.) Michx., Fla,—Echinochloa colonum (L.)
Lk., Okla., Tex.—Eragrostis lugens Nees, Okla.—Hystrix patula Moench,
'Wise.—Leersia oryzoides (L.) Swartz, Nebr., Wise.—L. virginica Willd.,
Nebr., Wise.—Muhlenbergia racemosa (Miehx.) B. S. P., Nebr.—Panicum
miliaceum L., Del., 111., Minn., Ohio—P. texanum Buckl., Tex.—Paspalum
plicatulum Miehx., Alabama—P. stramineum^ Nash, Okla.—Poa pratensis
L., Nebr.—Scolochloa festucacea (Willd.) Ek., N. Dak.—Setaria italica
(L.) Beauv., Alabama, Conn., Del., Fla., Ga., Iowa, Md., Mass., Minn.,
Mo., Nebr., N. J., N. C, N'. Dak., R. I., S. C, Tex., Va., Wise.—S.
lutescens (Weigel) F. T. Hubb, Conn., Del., Ind., Kv., Md., Mass., Mieh.,
Nebr., N. J., N. Y., Ohio, Okla., Penn., R. I., Vt., Va., W. Va., Wise.—
S. viridis (L.) Beauv., 111., Ind., Iowa, Kans., Minn., Mo., Nebr., N. Dak.,
Okla., W. Va., Wise, (on Setaria has been reported as P. setariae Nisikado)
—Sorghum vulgare var. sudanense (Piper) Hitchc., N. Dak.—Stenotapk-
rum secundatum (Walt.) Kuntze, Fla., Tex.
" ' i
P. grisea is common on Setana millets in tlie; United States and is
a minor to sometimes destructive leaf spot on a large number of hosts.
When severe the spotting becomes a blighting or blasting of the foliage.
This species is not readily distinguishable from he later described P.
oryzae and probably all should be known as P. grisea. Bremer et al.
(1948) were of the same opinion. Altson (1926) ,i for instance, used P.
grisea for the disease on rice in British Guiana. However, most of the
rather voluminous literature is found under P. 'oryzae on tropical
plants, mostly rice. For that reason the writer has retained P. oryzae
in this text.
Early collections in Nebraska include a considerable number of
hosts of P. grisea which do not occur elsewhere. This fungus must either
be adapted to this area or else was prevalent during the'years in which
Bessey.and Webber were actively collecting there. These collections
should, perhaps, be verified by further study.
Schwarze (1917) illustrated P. grisea on an undetermined grass,
possibly a millet. '
Range: United States, Australia, British Guiana, Japan.
References: Heald and Wolf (1912)'; Pammel, Weems, and Scribner
(1901); Schwarze (1917, p. 120, Figs. 735, 736); Tanaka (1920).
Piricularia oryzae Briosi and Cavara—Rice Blast, Rotten Neck

Spots oblong, arid, fuscous to ashy gray, conidiophore terete to
terete-subulate, basal cell somewhat swollen, simple or sparingly
branched, emerging through stomata in small clusters, geniculate.
60-120 X 4-5 /x; conidia obclavate, apex attenuate, base truncate with
slight hilum where it breaks from the conidionhore, 2-septate, fuscous,
diaphanous, 20-22 X 10-12 /x.. I
On: Oryza sativa L., Ark., Hawaii, La., Tex.
This fungus has been intensively studied by oriental workers,
especially the Japanese, who determined etiology^^host range, and
racial differences in great detail. (Cf. Abe, 1931; 1933,^6 ;-1935; 1936;
1937; Konishi, 1933, etc.) -
In Italy the disease is called "brusgre" or blast. Farneti (1921)
attributed the blast to Piricularia oryzae under which he lumped P.
grisea, Helminthosporiwn oryzae, H. macôcarpum, H. sigmoideum,
Cladosporium sp., and Hormodendron sp. in one chaotically polymor-
phic species (cf. R.A.M. 1:344, 1922). The introduction of resistant or
adapted varieties of rice saved the Italian rice industry from further
heavy loss after about 1909. , -.^ /
Recently Anderson, Henry, and Tullis'('i'§47) published data on the
factors which affect the spread, persistence, and infectivity of P. oryzae
in this country. These workers determined the relative susceptibility
of 19 varieties of rice. The variety ^ u t t e was moderately resistant.
Such varieties as Acadia, Onsen, Asahi, and Blue Rose were very sus-
ceptible. They found that barley, corn, rye, and wheat could be in-
fected with P. oryzae but sorghum and oats were not attacked. Lesions
appeared after 5 to 7 days and conidia formed on such leaves after 24
hours of incubation in a moist chamber.
They found that rice plants were most susceptible to P. oryzae in
the seedling, early tillering, and heading stages. Maximum infection
could be obtained by exposing inoculated plants to continuous wetting
for 16 to 24 hours at temperatures' between 24° C. and 28° C. High
humidity and the presence of free moisture on the leaves were essential
for secondary spread. Conidia live for a week on dry plants but ap-
parently die in irrigation water by the end of 24 hours.
Ryker (1938) reported that farmers had observed that P. oryzae
was mQre severe on newly reclaimed marsh land than in succeeding
crops.
.Anderson and Henry (1946) used a spreader-sticker combination of
0.05 per cent sodium oleate and 0.25 per cent gelatin in a conidial
suspension of P. oryzae employed to inoculate rice leaves. It facilitated
infection appreciably, raising the infection from 0.2 to 0.8 in the water
controls and from 6.5 to 11.1 per cent in those employing the spreader
stickers.
Ideta (1926) lists this fungus as Dactylaria oryzae (Cav.) Sawada.
Range: tfnited States, Argentina, Brazil, British Guiana, Bulgaria, Burma,
Ceylon, China, Colombia, Dominican Republic, Fiji, Formosa, Gold
Coast, Greece, Hawaii, India, Indochina, Italy, Japan, Java, Kenya,
Korea, Martyn, Philippines, Puerto Rico, Romania, Sierra Leone,
Tanganyika, Turkey, Uganda, Uzbekistan, Venezuela.
References: Abe (1931; 1933,a,6; 1935; 1936; 1937); Anderson and Henry
(1946); Anderson, Henry, and TuUis (1947); Aoki (1935); Baldacci and
Ciferri (1936); Bremer, et al. (1948); Farneti (1921); Hemmi (1930;
1937); Hemmi and Abe (1931); Hemmi and Imura (1939); Henry, B. W.,
and Andersen (1948); Ideta (1926, V. 2, p. 865); Imura (1938); Konishi
(1933); Liver, Leal, and Brewer (1947); Nakatomi (1927); Nisakado
(1927,&); Nose (1933); Sakamoto (1940); Sawada (1937); Sueda (1928);
Sundararaman (1921); Suzuki (1930, 1933, 1934, 1935, 1937); Tanaka
(1920); Thomas (1931); Tochinai and Komiya (1939); Tochinai and
Shimamura (1932);'Yoshii (1936, 1937).
Piricularia parasitica E. and E.—On Tar Spot

Gray white, densely caespitose; conidiophores erect, simple to
branched, septate, hyaline, 70-110 X 3-4 /j.; conidia 15-22 X 5-7 /*,
obclavate-ovoid, hyaline, finally 2- to 3-septate.
On: Elymus virginicus L., Wise.—Hystrix patula Moench., Wise.—Muhlen-
bergia foliosa (Roem. and Schult.) Trin., Wise.
Davis (1926) reports that this fungus occurs only on Elymus and
Hystrix in association with spots of Phyllachora graminis (Pers.) Fckl.
.' i
Greene (1944,a) found it on Phyllachorayulgata Theiss. and Sydow on
Muhlenbergia. We have seen material of this fu;ngus and it does not
resemble any species seen in the Far West.
Range: United States. '
References: Davis, J. J. (1926); Ellis and Everhart (1893, proc. Phil.,
p. 462). '
Ramularia graminicola Pk.—Leaf Spot

Spots small, not numerous, irregular to subelliptical, pale, sur-
rounded by a broad, brown, indeterminate margin; conidiophores am-
phigenous short, hyaline, 12.5-25 ^ long; conidia subcylindric to fusoid,
sometimes acuminate, sometimes pointed, at each end, finally falsely
1-septate, hyaline, 25-35 /x long.
On: Poa palustris L., N. Y.
There is no information on this species except that of the original
description. I t should be studied to determine if it is distinct or a
synonym of a better known species. I t is surprising that, if this is dis-
tinct, it has not been reported elsewhere. Peck's illustration (PI. 4, Figs.
1-3) suggests a species of Fusarium.
Ramularia cornijormis (Ell. and TracyJ Sherb. has been reported
on spikelets of Tripsacu7n dactyloides L. from Mississippi.
Range: United States." •
Reference: Peck (1891, pp. 26-27, PL 4, Figs. 1-3).
Ramulispora bromi (Sprague) comb. nov.—Blotch

Cercospora bromi Sprague |
Ansatospora bromi (Sprague)l Sprague
Centrospora bromi (Sprague)iNewhall
. ' . . •
Spots gray to sor.did, elliptical or striate with-broader margins dark

brown to umber. Vegetative mycelia variable, hyalme~to-subhyaline to
fuliginous, hyphal cells variable in size and shape, 'rectangular to
angular, becoming subspherical or bulbous, stroma dark olive to dull
black, filling epidermal cells with polygonal and irregular cells and
spreading from there to outer surface of the host through stomata.^
Conidiophores variable in length, sometimes obscure, siniple or once
branched, single or fascicled to form coremium-like aggregates, fuligi-
nous to chlorine-colored. Conidia hyaline, borne singly or in pairs,
acrogenously or subacrogenously, obovate-fusifbrro, slightly curved,
1- to 4-septate, slightly constricted at septa,-ibase blunt with prominent
scar or hilum, 15-45 X 2.5-5 n (mostly 25^43 X 3-4.5 ii); secondary
conidia produced singly on from 25 to 80 per cent of the conidia,
usually from the second, or less commonly from the first, basal cell,
deflected at 45 degrees usually toward the leaf base, slender, fusiform,
3-15 X 0.8-2 IX. (mostly 10-12 X 1.5-1.8 /n), (Fig. 76) nonseptate, semi-
deciduous.
*
On: Bromus rigidus Roth, Oreg.—B. vulgaris (Hook.) Shear, Oreg.
Originally described as Cercospora bromi, this was later transferred
to Ansatospora (Sprague, 1946,a); but Newhall later found that his
genus Ansatospora was a later synonym of Centrospora (1946). He
FIG. 76.—Conidia of Ramulispora bromi from type of (Cercospora) bromi.

(From Mycologia.)
casually made the combination Centrospora bromi Sprague but with

the rider, "if it belongs here rather than in the genus Ramulispora."
It would appear to be reasonable to have this fungus known as Centro-
spora bromi (Sprague) Newhall but on comparing with Ramulispora
sorghi (Ell. and Ev.) Olive and Lefebvre (1946) it appears to show
even closer relationship to this genus and therefore Ramulispora bromi
(Sprague) comb. nov. is proposed for this considerably kicked-around
fungus.
This fungus was found three times, once in eastern Oregon in a
semiarid region, once in a •vacant lot at Corvallis, and once on B. vul-
garis in a wooded area in Yamhill County, Oregon. The material from
Corvallis had gray, striate lesions which were superficial and anthrac-
nose-like on the green plant parts. While very superficial on the silici-
fled stem the fungus nevertheless apparently caused considerable stunt-
ing, particularly where the attack had reached thfc heads. The collection
on B. vulgaris was scanty and the fungus was suspected, in this case, of
being a hyperparasite on undetermined fungi in tne gray spots. Possibly
this collection is distinct from the type of C. hromi but seemed to be
morphologically similar to it.
References: Newhall (1946); Sprague (1937,6; 1942,a; 1946,o).
Ramulispora sorghi (Ell. and Ev.) Olive and Lefebvre—

Sooty Blotch
Septorella sorghi Ell. and Ev.
Ramulispora andropogonis Miura
Titaeospora andropogonis (Miura) Tai ,
Spots elongate-elliptical, with straw-colored centers, surrounded by
reddish purple to tan borders according to the variety of the host.
Sclerotia amphigenous, gregarious, superficial on straw-colored center
of lesions, subglobose, coarsely tuberculate, subcarbonaceous, 53-170 fx.;
sporodochia amphigenous, develop from subepidermal stromata, becom-
ing erumpent through stomata, conidiophores fasciculate, 10-35 X 2-3
ju,; conidia filiform, with 1 to 3 branches, 5-53 X 1.1-2.5 y., hyaline,
curved, tapering toward apex, 38-86.3 X 1-9-3.0 jn, 3- to 8-septate.
On: Sorghum halepense (L.) Pars., Alabama, Miss., N. C.—S. halepenseX
S. vulgare, Miss.—S. vulgare Pers., Alabama, Ark., Fla.7 Ga., La., Miss.,
Okla.—S. vulgare var. sudanense (Piper) Hitchc, Fla., Ga., Tex., Va.—
S. vulgare X S. vulgare var. sudanense, Ga. /
1 '
Ellis and Everhart (1903) mistook the sporodoehium-like pad of
this fungus for a pycnidium and thereby^ placed it. in Septorella. Miura
(1920) created the new genus Ramidispora with cylindrical, flexuose,
branched, septate spores and resembling Cylindrosp^riwm in other re-
spects. He described E. andropogonis with spores 36-100'^X~2-4 /*. Olive,
Lefebvre, and Sherwin (1946) aided by recent work by-Bain (1941,t);
1945) and Bain and Edgerton (1943), as well as notes by Larsh (1944,
1945) assigned this fungus to B. sorgHi, making the necessary new
combination. They believed the considerably discrepancy in spore size
between the American and Manchurian collections to be of little sig-
nificance because the species is greatly influenced by environment. Tai
(1932), who placed Miura's species in Titaeospora, gave spores 47.6-
106.9 X 2.04-3.06 /*. ^ /
Artificial inoculations indicated that 16-18 days were required for
formation of maturing lesions. These take their sooty appearance from
the numerous small, black, tuberculate sclerotia which form on the leaf.
in pure culture the fungus grows very slowly, producing raised
tuberculate black masses, on which *he conidia appear in light pink.
gelatinous aggregates. Olive, Lefebvre, and Sherwin (1946) transfer
Bamulispora to the Tuberculariales of the Fungi Imperfecti.
Gloeocercospora sorghi (Bain and Edgerton, 1943) differs from
Bamulispora in having nonbranching spores borne on a small stomatal
pad in a slimy matrix. Copper spot on bent grasses has been given as
caused by Gloeocercospora and by Ramulispora, which indicates that
the two genera have some resemblance to each other.
Range: United States, China, India, Manchuria.
References: Bain (1941,6; 1945); Bain and Edgerton (1943); Ellis and
Everhart (1903); Larsh (1944, 1945); Mehta and Bose (1946); Miura
(1920); Olive, Lefebvre, and Sherwin (1946); Tai (1932).
Rhynchosporium spp.
The spores are produced from a very flat obscure pad but not on a
true sporodochium (Caldwell, 1937). A key to the two species on
Gramineae follows.
KEY TO SPECIES OF RHYNCHOSPORIUM SPP. ON GRAMINEAE
Conidia apically obliquely beaked R. secalis (Oud.) J. J. Davis

Conidia cylindrical R. orthosporum Caldwell (Fig. ,77,4)
Rhynchosporium orthosporum Caldwell—Scald or White Leaf Stripe

• On leaves, spots 2.5-3.0 cm. long, lenticular, coalescing to form
irregular lesions, at first water-soaked, dark olivaceous, later becoming
FIG. 77.—A, Rhynchosporium orthosporum conidia from Calamagrostis cana-

densis, Wyo.; B, R. sp., conidia from Elymus glaucus, Linn County, Oreg.
gray surrounded by a brown margin, or on Dactylis fading to off-white;

conidia 14.4-20 X 2.3-4.7 /n, erect, cylindrical, medianly septate, formed
directly on cells of superficial stroma, more or less covering leaf spots.
On: Agropyron subsecundum (Lk.) Hitchc, Ida., Utah—Agrostis alba L.,
Oreg.—Alopecurus pratensis L., Oreg.—Calamagrostis canadensis (Michx.)
Beauv., S. Dak., Wyo.—Dactylis glomerata L., Calif., Ida., Mont., Utah,
Wash., Wise, Wyo.—Elymus glaucus Buckl., Oreg.—Lolium multiflorum
Lam., Oreg.—L. perenne L., Oreg.
' i
This species is very destructive to Alopecurus in Benton County,
Oregon, and is occasional on grasses in the"^hills ^nd canyons east to
Wisconsin. While typical B. orthosporuni differs distinctly from R.
secalis, there are collections that are'more or Uss intermediate be-
tween the two species. Both species occur on Lolium :in Oregon. A
collection on Agropyron subsecundum from Bountiful Canyon, Utah,
is hesitatingly assigned to R. orthosporum.. I t has spores which vary
from typical for the species to blocky capsular ones as short as 7 X 4.2
fji. A few tend to form oblique apices as in" R. secalis. Material on
Dactylis from Kendrick, Idaho, also was a variant. It had a few spores
that were 2- to 3-septate and up to 31 ji long.
Some material on Elymus glaucus, restricted to an area'in Linn and
Marion Counties, Oregon, may be an undescribed species or a varietj'
with somewhat distinct morphology (Fig. 77,B).
Drechsler's report of R. secalis (1921) from Wisconsin, on Dactylis,
is probably referable to R. orthosporum. •
References: Caldwell (1937); Drechsler (1921); Sprague (1942,o; 1946,e;
1950,a).
Rhynchosporium secalis (Oud.) J. J. Davis—Scald

Ramularia hordei McAlpine
Lesions at first dark bluish gray, water-soaked in appearance, later
the collapsed tissue becomes light gray with a darker brown margin
(Pig. 78), sometimes irregularly zonate, frequently elongate with
pointed terminals, often completely killing scalded leaves. Mycelium
penetrating by small appressoria and germ tube directly through the
epidermis; mycelium 0.6-3.0 fi diam., grow'th sparse in the interior of
the leaf, subcuticular at first but superficial stroma which are several
cells thick formed on the spot; conidia formed as direct outgrowths of
stroma, no evident conidiophores, conidia numerous^Jiyaline, allantoid
with a short oblique apical beak, medianly septate, 11-22 X-2:3-5.4 fi.
On: Agropyron dasystachyum (Hook.) Scribn., Oreg., WasH.—A. elmeri
Scribn., Wash.—A. repens (L.) Beauv., B. C, Calif., Colo., Ida., Minn.^
N. Dak., Oreg., Wash., Wise.—A. semicostatum (Steud.) Nees, Wash.—A.
subsecundum (Lk.) Hitchc, Wash.—A. trachycaulum (Lk.) Malte, Colo.,
Ida., Wash., Wye.—Agrostis alba L., Oreg.—Bbuteloua gracilis (H. B. K.)
Lag., Nebr.—Bromus carinatus Hook, and Arn., Ida., Wash.—B. jron-
dosus (Shear) Stand, and Wootn., Wyo.—B. inermis Leyss, Alta., Calif.,
Manit., Oreg., Wise.—B. pumpelliamis Scribn., Wyo.^5. secalinm L.,
Calif.—Danthonia (?) spicata (L.) Beauv., Qr.eg7—Elymus canadensis L.,
Wash., Wise.—E. glaucus Buckl., Ida., Ore'g., Wash., Wyo.^JS'. innovatus
Beil, S. Dak.—E. junceus Fisch., Was-h.—E. triticoides Buck!., Wash.—
E. virginicus L., Ind.—Hordeum distichon L., Calif., 111., Ind., Mich., Ohio,
Oreg., Wash., Wise, Wyo.—H. jubatum L., Oreg., S. Dak., Wise.—H.
murinum L., Calif., Oreg.—H. vulgare L., Alabama, Alta., B. C, Calif.,
Ida., 111., Ind., Iowa, Mich., Mo., Ohio, Okla., Oreg., Sask., Tex., Wash.,
Wise.—Lolium multiflorum Lam., Oreg.—L. perenne L., Oreg.—Phalaris
arundinacea L., Alta., Ida., Mont., N. Dak., S: Dak., Wyo.—Secale cereale
L., Alta, Calif., 111., Sask., Utah, Wash., Wise.—S. montanum Guss., Wash.
In his detailed study, Caldwell (1937) recognized six highly special-
ized physiologic races, which were distinguished by their ability to
attack one of six hosts: rye, barley, quack grass, smooth brome, Canada
wild rye, or Hordeum jubatum. None of these except the race from
Canada wild rye could attack a host species of a genus other than that
from which it was isolated. The race from Canada wild rye could attack
Agropyron trachycaulum, which is more or less to be expected as fungi
attacking the genus Agropyron are likely to attack members of the
genus Elymus, there being little, if any, fundamental genetic difference
between the two genera. Were all the additional hosts found since Cald-
well's day to be tested at this time, several unrecognized strains would
be found.
The collections on Phalaris arundinacea are less strongly beaked
than most of the other collections (see B.P.I. 80, 456) and this material
may represent another race. The spores are 13-18 X 2.3-2.8 /x. It should
be added that recently collected material by G. W. Fischer from Water-
ton National Park, Alberta, has spores which are definitely interme-
diate between R. orthosporum and R. secalis.
Caldwell (1937) found that R. secalis penetrated directly through
the epidermis of the leaf, although the host sometimes formed re-
sistance wall material around penetrating hyphae. These hyphae or
pegs arose from small appressoria. The fungus produced scant mycelia
in the interior but after collapse of the outer host tissue it formed a
thin but extensive pad from which masses of conidia arose directly.
This trait of forming masses of conidia apparently out of nowhere has
confused workers who have encountered the fungus for the first time or
are not familiar with its development. Rodriguez-Vallejo (1948) found
that Mars, Tregal, Manchuria, Bolivia, Svansata, and Texan (C.I.
5127) were not infected in 1947 trials at Chapingo, Mexico. A number
of varieties were' susceptible. Riddle and Suneson (1948) recently
reported varieties of barley including backcross-derived C.I. 7189
(Turk X Atlas'') and Atlas 46 (C.I. 7223). Sarasola and Campi (1947)
report comparable studies from Argentina.
Range: United States, Argentina, Australia, Belgium, Canada, Denmark,
England, France, Germany, Mexico, Netherlands, New Zealand, Peru,
S. Africa, Tanganyika, Tunis, Turkey, U.S.S.R.
References: Bartels (1928); Brooks (1928); Caldwell (1937); Davis
(1919,6); Drechsler (1921); Frank (1897); Heinson (1901); Johnson
and Mackie (1920); Mackie (1929,a); Oudemans (1897); Riddle and
Suneson (1948); Rodriguez-Vallejo (1948); Sarasola and Campi (1947);
Smith, N. J. G. (1937); Sprague (1946,e).
FIG. 78.—Lesions caused by Rhyncliosporium secalis on Agropyron repens,

near Priest River, Idaho.
Scolecotrichum graminis Fckl.—Leaf Streak

Producing gray, brown, or ashy streaks on leaves; conidiophores
fasciculate on the upper surface, resembling a seried arrangement of
black dots; olive brown, unbranched, irregular or slightly geniculate,
30-50 X 4-5 n; conidia 35-45 X 8-10 ju,, elongate, bottle-shaped, base
broadly rounded with prominent hilum, 1-, less often 2-septate.
On; Agropyron caninum (L.) Beauv., N. Dak.—A. cristatum (L.) Gaertn.,
i
N. Dak. (rare)—A. dasystachyum (Hook.) Scribn., N. Dak., Oreg., Wash.,
Wyo.—A. grifjithsii Scribn. and Sm., S. Dak.—A. inerme (Scribn. and Sm.)
Rydb., Ida., Oreg., Utah, Wash., Wyo.—A. repens (L.) Beauv., Ida.,
Minn., Nebr,, Utah, Wash., Wise.—A. riparium Scribn. and Sm., N. Dak.,
Wyo.—il. smithii Rydb., Nebr., N. Dak., Oreg., S. Dak., Wyo.—^.
spicatum (Pursh) Scribn. and Sm., Colo., Ida., Mont., Nev., Oreg., Utah,
Wash., Wyo.—A. subsecundum (Lk.) Hitchc, Alta., Colo., Ida., N. Dak.,
Utah—A. trachycaulum (Lk.) Malte, Alta., Ariz., Colo., Ida., Minn.,
Mont., Nebr., Nev., N. Dak., Oreg., S. Dak., Utah, Wash., Wyo.—
Agrostis alba L., Colo., Ida., Minn., Mont., Nebr., N. Dak., Ont., Oreg.,
Que., S. Dak., Utah, Wash., Wise, Wyo.—A. castellana Boiss. and Rent.,
Oreg.—A. exarata Trin., Ida., Mont., Oreg.—A. hallii Vasey, Oreg.—A.
oregonensis Vasey, Ida., Mont.—A. rossae Vasey, Wash.—A. scabra Willd.,
CaUf., Colo., N. Dak., Oreg., S. Dak., Utah—A. scabra var. geminata
(Trin.) Swallen, Wyo.—Alopecurus aequalis Sobol, Oreg., Wash.—A.
carolinianus Walt., La., Okla.—A. geniculatus L., Ida., Iowa, Okla., Oreg.,
Wash., Wise.—A. pratensis L., N. Dak., Oreg.—Arrhenatherum elatius
(L.) Beauv., B. C, Calif., Ida., Ky., Md., Miss., Mo., Mont., N. Y., Oreg.,
Penn., Wash.—Arundinaria gigantea (Walt.) Chapm., Alabama—Avena
sativa L., 111., N. Y.—Beckmannia syzigachne (Steud.) Fern., Ida., Mont.,
Nebr., N. Dak., Oreg., S. Dak., Wash., Wyo.—Bromus arenarius Labill,
Oreg.—B. carinatus Hook, and Arn., Cahf., Colo., Ida., Mont., N. Y.,
Oreg., S. Dak., Utah, Wash., Wyo.—5. catharticus Vahl, N. Y., Wash.—
B. ciliatns L., Alta., Colo., Wyo.—B. frondosus (Shear) Stand, and
Wootn., Colo.—B. inermis Leyss, Alta., Ariz., Ida., Oreg.—B. laevipes
Shear, Mont.—B. mollis L., Wash.—B. orcuttianus Vasey, Oreg.—B.
pacificus Shear, Oreg.-—B. purgans L., Colo.-—B. sitchensis Trin., Wash.—
B. vulgaris (Hook.) Shear, Ida.—Calamagrostis canadensis (Michx.)
Beauv., Ida., Mont., N. Dak., S. Dak., Wash., Wise.—C. canadensis var.
scabra (Presl) Hitchc., Alt^-, Wash.—C. inexpansa A. Gray, Wash.—
Cinna arundinacea L., Wise.—C. latifolia (Trev.) Griseb., Ida.—Cynosorus
cristatus L., B. C", N. Y.—Dactylis glomerata L., B. C, Colo., Ida., Iowa,
Ky., La., Md., Mo., N. Y., Okla., Ont., Oreg., Utah, Vt., Va., Wash.,
Wise, said to be general in the eastern United States—Danthonia cali-
fornica BoL, Oreg.—Deschampsia atropurpurea (Wahl.) Scheele, Wash.-—•
D. caespitosa (L.) Beauv., Oreg., Wash.—D. danthonioides (Trin.) Munro,
Ida., Oreg.^—D. elongata (Hook.) Munro, Ida., Mont., Oreg., Wash.—
Digitaria sanguinalis (L.) Scop., Va.—Elymus antarcticus Hook, f., N.
Dak.—E. aristatus Merr., Oreg.—E. canadensis L., Ida., Iowa, Kans.,
Minn., Mont., Nebr.", N. Y., N. Dak., S. Dak., Utah, Wash., Wise, Wyo.—
E. condensatus Presl, Alta., Colo., Ida., Mont., Utah, Wash.—E. flavescens
Scribn. and Sm., Mont., Wash.—E. glaucus Buckl., Alta., B. C, Calif.,
Colo., Ida., Kans., Oreg., Wash., Wyo.—E. giganteus Vahl, Manit.—E.
innovatus Heal, B. C.—E. mdcounii Vasey {Hordeum jubatum x Agro-
pyron trachycaulum), Colo., N. Dak., Wyo.—E. mollis Trin., Alaska—
E. sibiricus L., Wash.—E. triticoides Buckl., Calif., Oreg.—E. virginicus L.,
Kans.—Eragrostis secundiflora Presl, Okla.—Festuca elatior L., Kans.—
F. rubra L., Ont., Wise.—Glyceria borealis (Nash) Batchelder, Wise—
G. elata (Nash) Hitchc, Cahf., Wash.—G. fluitans (L.) R. Br., Mich.,
N. Y., Oreg., Wise—G. grandis S. Wats., Ida., Mont., N. Dak., Oreg.,
Wise—G. leptostachya Buckl., Wash.—G. pauciflora Presl, Calif., Wyo.—
G. septentrionalis Hitchc, N. Y., Oreg., Wise—G. striata (Lam.) Hitchc,
Oreg.—Hordeum jubatum L., Colo., Ida., Minn., Mont., Nebr., Nev., N.
Dak., Oreg., S. Dak., Utah, Wash., Wise, Wyo.—H. jubatum var.
'I
caespitosum (Scribn.) Hitchc, Colo., Nev^ Wash.—H. nodosum L., Alta.,
Ida., Kans., Mont., Oreg., Wash., Wyo.—H. nodosurif, var. boreale (Scribn.
and Sm.) Hitchc, Oreg.—H. pusillum Nutt., Kans., Mo., Mont., Okla.,
S. Dak.—H. vulgare L., Iowa, Ohio, Okla., Sask., Utah—Hystrix patula
Moench, Wise.—Koeleria cristata (L.) Pers.; Mont!,|Nebr.,iN. Dak., Wyo.
—Leersia oryzoides (L.) Swartz, N. Y.—Loliim multifiorum Lam., Oreg.
(rare)—L. perenne L., Oreg.—Melica bulbosa Geyer, Oreg.—M. geyeri
Munro, Wyo.—M. smithii (Porter) Vasey, Ida.—M. spectabilis Scribn.,
Wyo.—Melica subvlata (Griseb.) Scribn., Ida.—Milium effusum L., N. Y.,
Wise.—Muhlenbergia filiformis (Thurb.) Rydb., Ida.—M. mexicana (L.)
Trin., Iowa—M. racemosa (Michx.) B. S. P., Wise.—M. silvatica Torr.,
Wise.—Oryzopsis hymenoides (Roem. and Schult.) Rick.,-Mont., Utah,
Wash., Wyo.—Phleum alpinum L., Colo., Wyo.—P. pratense L., Alta.,
B. C, Colo., Ida., Iowa, Ky., Manit., Md., Minn., Mo., Mont., N. Dak.,
N. S., Ont., Oreg., P. E. Is., Sask., S. Dak., Utah, Vt., Wash., Wise,
Wyo., said to be general in the eastern United States—Pkragmites com-
munis Trin,, Wise.—Poa alpina L., Wyo.—P. ampla Merr., Mont,, Wash.,
Wyo.—P. arida Vasey, Mont., N. Dak.—P. canbyi (Scribn.) Piper, Colo.,
Mont., N. Dak., S. Dak., Wash., Wyo.—^. compressa L., Colo., Ida., Iowa,
Md., Minn., Mo., Mont., Nebr., N. Dak., Oreg., Penn., S. Dak., Wash.,
Wise, Wyo., general in the eastern United States—P. epilis Scribn., Wyo.
—P. gracillima Vasey, Ida.—P. interior Vasey, Ind:, N. Y., Wyo.—P.
juncifolia Scribn., Oreg.—P. longiligula Scribn. and Wils., Wyo.-—P. ner-
vosa (Hook.) Vasey, Oreg., Wyo.—P. nevadensis Vasey, Colo., Mich., N,
Dak., Oreg., Wash., Wyo.—P. palustris L., Minn., Oreg., Wise.—P.
pratensis L., Alaska, Alta., B. C, Iowa, Manit., Minn., Mont., N. Y.,
Ont., Wise.—P. secunda Presl, Ariz., N. Dak.—P. stenantha Trin., Mich
—P. trivialis L., N. Y.—Secale cereale L., Oreg., S. C.-^S. cereale x S.
. montanum, Wash.^S. montanum Guss., Wash.—Sitanion hansenii
(Scribn.) J. G. Sm., Calif., \Yash.—S: hystrix ONutt.) J. G. Sm., Ariz.,
Mont., Nev., N. Mex., Oreg., Wyo.—S. jubaturp, J. G. Sm., Oreg., Utah,
Wash.—Spartina gracilis Trin., Oreg.—Stipa Columbiana Macoun, Ida.,
Mont., Oreg., Utah, Wash., Wyo.—S. Columbiana var. nelsonii (Scribn.)
Hitchc, Calif., Mont., Wash.—^S. comaia) Trin. and Rupr., Colo., Mont.,
Nebr., Oreg., S. Dak., Utah, Wyo.—S. coronata Thurb., Calif.—S.
elmeri Piper and Brodie, Wash.—S. lemmonii (Vasey^Scribn., Ida., Oreg.,
Wash.—S. lettermanii Vasey, Oreg., Utah, Wash., Wyo>=-)S. occidentdis
Thurb., Calif., Oreg., Wash.—S. speciosa Trin. and Riipr., Calif.—S.
thurberiana Piper, Nev., Oreg., Wash., Wyo.—S. viridula Trin., Ida.,
Mont., Utah—S. vnlliamsii Scribn., Ida., Wyo.—Trisetum canescens BuckL,
Ida., Mont.—T. flavescens (L.) Beauv., N. Y.—T. montanum Vasey, •
Mont. / /
i
The_^ shape, color, and extent of the lesions caused by this fungus
vary with the age of the plant and with the host species (Fig. 79). In
its most common condition it occurs as elongate grayybrown-ochraceous
•to stramineous streaks on the leaves wjtlj'.^niline/to sulphine yellow
borders in which the black clusters of conidiophores appear as minute
dot& in seried arrangement. Younger stages show water-soaked, circu-
lar to elliptical lesions which are deep olive gray in the morning when
dew-wet, and dull gray when dry. These spots become brown, purple
FIG. 79.—Lesions caused by Scolecoirichum qraminis on: A, Beckmannia

syzigachne, Pot latch, Idaho; B, Bromus carinalus, Pullman, Wash.
to ochre with gray centers, with a tendency to form streaks as the

leaves slowly die.
This fungus is readily distinguished by the j^roininent black, densely
fasciculate conidiophorcs arranged in parallel rows as black dots along
the surface. Without a hand lens they are sometimes mistaken for
pycnidia. Less frequently they may be plentiful enough to give a sooty
mold aspect to the underside of the leaves. The spores are Indian-club-
I
shaped, i.e., fusoid-obclavate, 35-45 X 8-10 /A, 1- or sometimes 2-sep-
tate, olive brown.
The correct name for this fungus has never been settled to the
satisfaction of all workers, although most of them use the old name
which we employ and which Fuckel proposed in 1863. Horsfall (1930)
suggested that all available evidence indicated that Cercospora graminis
that V. Hoehnel (1923) considered Scolecotrichum Kze. as probably
invalid because of the indefinite nature of S: virescens Kze. the type
species. Until the matter is settled the writer uses the old name.
Synonyms for the fungus include: S. compressii,m Alleschr. (Hedw.
35(2) :34. 1896); Cercospora graminicola Tracy and Earle (Torr. Bot.
Club 22:179. 1895); Passalora hordei Otth. (v. Hoehnel, 1923); P.
graminis (Fckl.) v. Hoehn. (1923); P. punctiformis Otth. (v. Hoehnel,
1923); P. dactylina Pass. (Lindau, 1907 in Rabenhorst, Krypt. Flora
V. 8, pp. 793-799); S. graminis var. nanum Sacc. (in v. Hoehn., 1923);
Napicladium gramineum Pk. (Sprague, 1942,c) and probably Cerco-
spora poae Baudys and Picb. (1924).
V. Hoehnel stated that S. graminis (1923) had a perfect stage called
Carlia recutita (Fr.) v. Hoehn., sometimes known as Sphaeria recutita
Fr. and Metasphaeria recutita (Fr.) Sacc. Carlia is the same as Myco-
sphaerella (R.A.M. 3, p. 102) with a Passalora conidial stage. Today
this fungus would probably be known as Mycosphaerella recutita (Fr.).
We have examined many collections of Scolecotrichum graminis asso-
ciated with various ascomycetous fungi' but have never noticed any
that appeared to be consistently associated with it. This point needs
study. -^
The known host range of S. graminis is great. Since Horsfall listed
28 hosts for the entire world in 1930 the known host range of this fungus
has greatly increased. An extensive collection of this species is now
available at Oregon State College, AVashington State College, and in
the Mycological Collections, U.S.D.A., Beltsville, Maryland. S. gram-
inis is particularly common on the Pacific coast, but is abundant in
the Plains country as well. In a trip made south from Pullman, Wash-
ington, with G. W. Fischer and Jack P. Meiners in June, 1947, a very
dry year, the fungus was abundant in all but the sagebrush area of
Idaho, was prevalent in the canyons and hills but not in the plains of
Utah, and disappeared in Arizona, New 'Mexico, and southern Cali-
fornia, except for scant quantities in the Kaibab National Forest, Ari-
zona. In California it was absent from San Diego noi^th-to.about Bridge-
port, California. It is scarce in the alkali valleys of Nevada but ex-
ceedingly abundant in eastern Oregon in forested areas visited by re-
cent rains. East of the Rockies the fungus-is one of the most important
leaf spots of timothy, orchard grass, blue grass, tall meadow oats grass,
redtop, and native Stipa and Poa grasses. Spmetimes it is less serious
than it appears to be because the fungus sporulates profusely in late
season and develops saprophytically on many hosts on which it is only
mildly parasitic. It is sometimes secondary to other fungi and to bac-
teria. It is not important on cereals but spmetimes occurs on barley,
oats^, arid rye. It is sometimes moderately prevalent in localized areas
on Secale in "shoe string" valleys in coastal'Oregon and once was found
on 40 per cent of the leaves of winter rye in early May near Alsea,
Oregon (Sprague, 1938,c), and on another occasion it was found on rye
heads in the same general area. Guarch (1941) reported S. graminis
var. brachypoda Speg. on rye in Uruguay and Jankowska (1929) listed
it from Poland on rye, and also said that it caused slight injury to
wheat. Shitakova-Roussakova (1939) reported the disease as widely
distributed and of moderate importance on rye in the Soviet Union. She
found some varietal resistance. Landaluze (1941) reported S. graminis
as a secondary parasite of rye in a hilly region in Spain where the soil
was pH 5.4. This condition is similar to that encountered near Alsea,
Oregon, where the soil is highly acid (pH 4.9-5.5) and also very de-
ficient in nitrogen and phosphorus. Probably the injury on rye is not
representative of very active parasitism.
A tabulation of about 300 collections of S. graminis shows that this
fungus may be found in any month of the year, but in Oregon its great-
est activity was from March to July with a secondary development in
the fall after the rains started again. Injury is greatest on older plants
in the spring. In the northern Great Plains, the fungus developed in
May and June but profusely fruiting material was collected from May
until late September.
This fungus needs critical study to determine its physiological
status. Morphological variants occur (as on Poa ampla Merr. in Klicki-
tat County, Washington) but whether they are anything more than
variations due to host and humidity is not known.
Information on the control of this disease is scanty. Certainly grass
weeds, such as Hordeum jubatum and Beckmannia syzigachne, aid in
spreading the fungus. Burning dead grass reduces the inoculum.
Range: United States, Argentina, Austria, Canada, China, England, Ger-
many, Poland, Spain, U.S.S.R., Uruguay, Wales.
References: Fischer, Sprague, Johnson, and Hardison (1942); Fuckel
(1863); Guarch (1941); Horsfall (1930); Jankowska (1929); Johnson,
A. G., and Hungerford (1917); Krause (1909); Landaluze (1941); Nils-
son-Enle (1908);, Pammel, Weems, and Scribner (1901); Saccardo
(1886); Sampson and Western (1941); Shitakova-Roussakova (1939);
Sprague (1938,c; 1942,c; 1946,e; 1949,o); Trelease (1887).
Scolecotrichum maculicola Ell. and Kell.—Leaf Spot

Spots amphigenous, narrowly elliptical, about 5-8 X 2 mm., dirty
white with a dark border, hyphae hypophyllous, subundulate, continu-
ous or with a faint septum near the base, subfuliginous, about 40 X 4-5
fjL, growing in dense, spreading, olivaceous, seriate tufts, forming a single
continuous line along the middle of each spot; conidia terminal, gran-
ular, continuous, ovate, ovate-elliptical or oblong, subhyaline, 20-
22 X 8-11 [i.
On: Arundo donax L., Okla.—Phragmites communis Trin., Iowa, Kans.,
Nebr., N. Dak., Okla., Oreg., Wise.
430 DISEASES OF CEREALS AND G M S S E S
Both the spores and conidiophores of this fungus are considerably-
smaller than those of S. graminis. Von B'oehnel placed this fungus in
Cladosporium, near C. phragmitis Opiz.
This species is relatively common in the plains region. It is abundant
in one locality on Young's Bay, near Astoria, Oregon.
Range: United States, Austria. '
Reference: Ellis and Kellerman (1887).
Spermospora avenae (Sprague and A. G. Johnson) comb. nov.—Red

Leather Leaf
Pseudodiscosia avenae Sprague and A. G. Johnson
Mycelium branched, septate, hyaline to faintly tinged with yellow,
intracellular or intercellular in the leaf, aggregating at or near the
surface, more frequently the upper surface, producing blunt or rarely
sharp-pointed, 1- to 3-celled (mostly, 1-celled) conidiophores 5-
12 X 0.8-2.0 fi. Conidium with one cilium at each end. Body of the
spore narrowly fusiform, slightly curve,d, hyaline or faintly tinged with
yellow, 2- to 3-septate, 10-42 X 2-4, > . Conidia are borne singly or
sometimes in pairs, acrogenously or subacrogenously. Bases of spores
rounded or blunt wdth a rod-shaped or sometimes sharp-pointed ap-
pendage or cilium, 1-16 X 0.3-1.3 /x, attached obliquely near the hilum.
The apical appendage or cilium is usually longer than the one at the
base and usually merges very graduallj' with the body of the spore.
On: Arrhenatherum elatius (L.) Beauv., Ida., Wash.—Avena byzantina K.
Koch, Oreg., Wash.—A. sativa L., Oreg., Wash.
In the original discussion of this fungus (Sprague and A. G. John-
son, 1936) it was pointed out that the spores were borne on very short
conidiophores compacted together on a poorly developed stroma in the
epidermis. There was no apparent rupture of the leaf epidermis as in
typical acervuli, and in assigning this to the Melanconiales the de-
scribers were guided by expediency in their uncertaiijty as to where
this fungus belonged. There was no species in the Moniliffceae nor in
the Tuberculariae, both in the Moniliales, to which this^fungus could
have been assigned. The nearest genus, morphologically, was Pseudo-
discosia in the Melanconiales. Since Spermospora as revised (Sprague,-
1949,a) can now logically be used as a depository for this, fungus it is
now removed from its dubious place in the Melanconiales and placed
in Spermospora in the Moniliales where it appears to be better assigned.
In western Oregon and Washington entire 'fields of winter oats
sometimes show brilliant red color in February, in other cases the
disease is confined to areas in the fields.' In-these areas all but the in-
nefmo'st leaves of a plant may be affected. These lesions cause a leath-
ery, fawn to red or brown appearance. An excess of red anthocyanin
pigmentation in the uninjured portiojis of the diseased leaves gives the
reddish brown color to affected fields. The diseased plants are stunted,
compact, and rigid. The leaves are slightly rolled. The disease where it
occurs is of some importance, although oats recover to a considerable
extent as soon as warm weather arrives in the spring. Red leather leaf
is almost unrecognized as a definite disease, even at the several experi-
ment stations in the area where the trouble is so prevalent. The symp-
toms are confused with red-leaf symptoms in general (Sprague, 1939,6).
Many specimens are sent to the experiment stations during early spring.
While this fungus is very largely confined to the mild, humid,
coastal parts of the Pacific Northwest it was found recently in the
interior at Pullman, Washington, on the closely related genus Arrhena-
therum in February, 1947 and at Moscow, Idaho, in January, 1948.
Bremer (1943) also found the fungus on oats in Turkey where it ap-
peared to be favored by cool late winter weather. Bremer found the
species on a wild grass, perhaps Avena sterilis L., growing in the
mountains. He considered that it was probably native.
Range: United States, Turkey.

References: Bremer, H. (1943); Sprague (1934,e; 1935,6; 1936,6; 1938,c;
1939,6; 1947,6); Sprague and A. G. Johnson (1936).
Spermospora subulata (Sprague) Sprague—Blast

Cercosporella subulata Sprague
Lesions extensive, elongate, sometimes diffused as a scald over large
portions of the leaves, sordid dull straw color to olive buff and isabelline
bordered by darker tones of umber to buffy brown, in severe cases
causing a dark brown or umber brghting of the entire plant which in
the case of bulbous species of Melica is readily broken off at the base.
Vegetative mycelia hyaline to strongly chlorine, septate, branched, 1-2
ju, diam., stromatic mycelia olivaceous to dull black, aggregated at or
near the surface of the leaf from which are produced short, hyaline to
strongly chlorinous conidiophores, 5-20 X 1.5-2.0 JM. Conidia hyaline,
produced in large numbers, borne blunt, broad base down, subulate to
subulate-filiform, tapering to a whip-like distal portion which is some-
times straight, usually curved or fiexuous, rarely reflexed. Conidia some-
times 1-septate, usually 2-septate with basal septum at broadest por-
tion of subulate base, sometimes lightly constricted at second septum
from base, which usually is at basal end of whip-like distal portion,
sometimes with an oblique basal cilium, 10 X 0.4 jj. on Agrostis alba
and Calamagrostis rubescens (Sprague, 1949,a on f. ciliata). In old
material conidia frequently break at second septum. Conidia 20-35-
50-(80) X 2.5-5.4-(8.4) fi, mostly 28-40 X 2.8-3.5 /x. Material on Fes-
tuca rubra has 3-septate spores 33-55 X 3.3-4.3 ju.; on Deschampsia
spores faintly yellowish, 2-3 septate, 33-47 X 3.0-4.5 ju,; on Calama-
grostis rubescens spores 35-50 X 4.5-5.4 î, but swollen material (in
humid container) as much as 50-80 X 7.0-8.4 fl. P'ype is O.S.C. 10,669
on Melica subulata. ., i
On: Agrostis alba L., Wash. (f. ciliata on this host)—Bromus vulgaris
(Hook.) Shear, Mont.—Calamagrostis rubescem BuckL (also f. ciliata)
Wash.—C. scribneri Beal, Wyo.—Deschampsia •caespitosa (L.) Beauv.,
Ida., Mont., Wyo.—Festuca rubra L., Oreg., Wa^h.—F. rubra var. com-
mutata Gaud., Wash.—Melica bulbosa Geyer, Alta.-, Wyo.—M. smithii
(Porter) Vasey, Ida.—M. spectabilis Scribn., Wyo.—M. .subulata
(Griseb.) Scribn., Oreg.—Stipa lettermanii Vasey, Alta.—Trisetum spica-
tum (L.) Richt., Mont.
The fungus was originally found on a herbarium specimen in the
phanerogamic collections at Oregon State College from material of
Melica subulata collected in forested country in eastern Oregon. The
fungus has since been found scatteringly over a wide area in the moun-
tains and grass areas of the West on an increasing number of hosts. In
some areas in the moimtains of western Wyoming the fungus caused a
severe blight of Melica spp. in 1948.
Because of their sperm-like shape (Fig. 80) the spores of this fungus
were placed in a new genus which was described as follows: "SPERMO-
SPORA Sprague-conidia subulate ta-subulate filiform, apices elongate-
fusiform, appearing appendage-like, hyaline, borne on evident conidio-
phores in spots" (Sprague, 1937,b). Later .after basal cilia were found
in two collections the genus was emended as follows: "SPEBMOSPOEA
Sprague emend.—Conidia subulate to subulate-filiform, hyaline, sep-
tate, apical cell appearing appendage-like, sometimes with an oblique-
basal cilium, borne superficially on evident conidiophores in spots"
(Sprague, 1949,a). S. subulata f. ciliata Sprague (1949,a) was described
as being the same as the species but with some'of the spores having the
oblique-basal cilium 10 X 0.4 jn. |
Spermospora is distinguished from Cercosporella and from Mastigo-
sporium as follows: i '
)
KEY TO SPECIES SPERMOSPORA AS DISTINGUISHED PROM CERCOSPORELLA
A. Conidia more or less broadly filiform, hyaline - ^

Cercosporellar
AA. Conidia subulate or narrow subulate With distal cell elongated into an
attenuated, appendage-like terminal, sometimes with obUque-basal cihum-
^Spermospora /
AAA. Conidia broadened, cylindrical or boat-shaped, sometimes (European)
with apical appendages
Masiigosporium
The three above cited genera actually have very little in common
and all are distinct in many ways from the others.
An isolate from red fescue on potato-dextrose agar produced fawn-
', colored, slow-growing, felty mycelium which produced a few natural-
FIG. 80.—Conidia of Spermospora subulata: A, from Melica subulata, Natl.

For., Oreg.; B, ditto on Festuca rubra, Granger, Oreg.; C, ditto on Deschampsia
caespitosa, Beartooth-Passr Mont.
appearing spores at 40° F. after several weeks in the refrigerator witli-

out light. Macrospores were 35-50 X 3.4-4.6- /x and microspores, not
seen in nature, were 4-6 X 1-5-2.0 fi,.
References: Sprague (1937,6, Fig. 2,a; 1948,a,c; 1949,a; 1950,a).
Sporotrichum columbiense Sprague—White Stipple

Lesions on leaves obscure, straw-colored or eventually tawny,
curled, often without detail and then resembling drought-injured plants,
434 DISEASES OF CEREALS AND GHASSES
/j
stromata minute, vein delimited and resembling nearly white stippling
which resembles machine stitching, firm, riot cottony, 4O7I5O X 1.5-24
n; mycelium compacted, sometimes crushed but when distinguishable
the hyphae are 2-5 /J. diam.; branched, septate;the conidiophores are
3-5 IX diam. and there are hyaline, nonseptate,conidia of two sorts:
microspores 2-4.5 X 0.8-1.5 ^ of various shapes Tom globose to cylin-
drical, and macrospores 3-8 X 2.5-6 /*, mostly globose to elliptical
(Fig.81,A).
O0Q f
VoO.
FIG. 81.—A, Spore forms associated with siroma of Sporotrichum coluinbiense
on Agropyron inerme, Wash.; B, conidia. of Sporotrichum peribebiiyense on
Setaria lutescens, Okla. ,
On: Agropyron inerme (Scribn. arid Sm.) Rydb., 'Wash.—A. spicatum

(Pursh) Scribn. and Sm., Mont., Wash.—A. subsecundum (Lk.) Hitchc,
Ida.
This very little imderstood species was described after considerable
study and placed in Sporotrichum. The fungus is too obscure to be
visible without aid of the microscope. Diseased plants show dried,
curled, sometimes puckered leaves with only the vaguest of lesions.
This fungus should' be studied to determine if/it is, perhaps, systemic.
Range: United States. j
Reference: Sprague (1947,a). 1
I
I
Sporotrichum peribebuyense Speg.-^-Jjcaf Spot
Spots hypophyllous, linear, sparse, pulvinate, -cottony-woolly,
1-5 X 0.5-1.0 mm., white to pale rose color, hyphae densely and intri-
cately ramose, 5.0-5.5 ju diam., septate; -sterigmata papilliform, sparse,
hyaline; conidia globose, guttulate, hyaline, 6-8 [x. (8-11.2 /x in Okla-
homa collection). [ /
On: Setaria lutescens (Weigel) F. T. Hubb, 111., Mo., Okla.—S. viridis (L.)
Beauv., Mo.
The spots are covered with the white.^to rosy, cottony-woolly
hyphae. The fungus is either uncommon in the United States or seldom
noted. I t is very distinct from the previously described species (Fig. 81).
I Range: United States, Argentina.
References: Spegazzini (1885); Sprague (1947,a).
Stemphyllium botryosum Wallr.—Mold

Ascigerous stage is Pleospora herbarum (Pers.) Rab.
Conidiophores olive brown, variable in length, about 3-6 /x diam.,
swollen at the apex just below the attachment of the spore; after shed-
ding the penultimate cell continues growth through the old scar and
produces a new spore, or sometimes the terminal spore becomes pushed
to one side by the continued growth of the conidiophore and appears to
be lateral, false chains sometimes produced at apex by basipetal frag-
mentation; conidia oblong ovoid to subangular, olive brown, slightly
rough-walled, somewhat constricted at the medium septum, and more
or less packet-shaped or sarcinaeform, ends obtuse, with transverse,
longitudinal, and oblique septa, (14)-18-35-(50) X (12)-15-22-(27) /x,
borne singly at the end of conidiophores.
Perithecia irregular, black, hard and leathery in pure culture; asci
cylindrical to cylindric-clavate, short-stalked, thick-walled, 8-spored,
140-240 X 27-38 fi. Ascospores light brown, oblong ellipsoid to oblong-
ovoid, mostly straight, with seven cross septa, uniseriate to slightly
overlapping, 30-50 X 15-20 /x, enclosed within the ascus in an inner
membrane, which emerges when the tip of the ascus breaks open.
On: Dead parts of many plants, sometimes isolated from seedlings of
Gramineae in the Northern Great Plains and in eastern Washington and
adjacqnt Idaho.
This is apparently a saprophyte and as such is much less abundant
than Alternaria tenuis. The sarcinaeform (cubical), muriform, echinu-
late spores are typical of the species. It produces copious dark gray to
nearly black cottony colonies which sometimes have a stringy aspect.
Groves and Skolko, using malt extract agar (1944,a), obtained irregu-
lar sclerotium-like bodies which were hard and black. These were peri-
thecia primordia.
Patterson described S. tritici (1910) from Texas and Oklahoma. It
was found on wheat and was closely associated with rusts according to
notes by E. C. Johnson. Heald and Wolf (1912) reported it as asso-
ciated with a floret sterility and it occurred on leaves and spikes. This
species was probably "S. botryosum.
Range: Cosmopolitan.
References: Groves and Skolko (1944,a); Heald and Wolf (1912); Patter-
son (1910); Wiltshire (1938).
Stemphyllium consortiale (Thuem.) Groves and Skolko—Mold

Conidiophores brown, simple or sometimes branched, variable in
length, about 3-5 ju diam., not swollen below the attachment as in
S. botryosum, spores produced acrogenously but they become pushed
to one side by the continued growth of the conidiophores so that they
occur in botryose clusters, the conidiophores becoming geniculate;
conidia variable in shape, subglobose, quadrilateral to oblong-ovoid, at
first nearly smooth, becoming coarsely^ verrucoê, muriform, more or
less constricted at the septa, very dark brown to black, (13)-18-25-
(28) X (12)-14-17-(23) ^.
On: Tnticum aestivnm L., Wash.
This species occurs commonly on dead parts of wheat killed by
Fusarium nivale and Typhula spp. in Douglas County, Washington.
It is obviously saprophytic. This species has cubical to globose spores •
which, as illustrated by Groves and Skolko, are readily segregated from
(S. boir-yosum (1944,a).
Range: Northern hemisphere, probably general.
Reference: Groves and Skolko (1944,a).
Ustilaginoidea virens (Cke.), Tak.—False Smut

U. oryzae (Pat.) Bref. /
Sclerotia in flower parts or in tassel^ (pollen heads of corn) 5-8 ram.
diam., covered with olive green conidia formed on the surface of the
pseudomorphs which replace the ovary of the rice-or anthers of corn;
fertile hyphae septate, sometimes branched, yellowish green, 2-3 ju,
diam.; conidia borne on short sterigmata-like projections from the
hyphal wall, spherical, 4-6 /x diam., dark olive brown. Germinates to.
form secondary spores.
On: Oryza sativa L., Calif., Ga., La., S. C.—Zea mays L., Canal Zone, La.
This causes a mild false smut of corn tassels in the southern United
States and a head mold of rice. In some ways, the fungus resembles
ergot because the flowers are replaced by sclerotia in rice. Gaumann
and Dodge (1928) distinguish the genus from Claviceps by the black
imperfect stage of the former. '
U. setariae Bref. occurs on millet but (is apparently not known in the
United States. It may not be distinct from U. viren&..
Range: United States, Burma, Ceylon, China, Dutch Ea'st Indies, India,
Indochina, Japan, Madagascar, Phihppines, Sierra Leone.
References: Butler (1913,6); Fulton (1908); Haskell and Diehl (1929);
Raychaudhuri (1946); Reinking (1918, p. 226); Takahashi (1896);^
Vincens (1922). / /
GLOSSARY
aberrant: atypical.
acervulus(i): a pustular fruiting body without a covering^ usually a flat mass
of conidiophores producing a quantity of wet conidia.
acicular: needle-shaped.
acrogenous: growing at the apex.
acro-pleurogenous: growing on the sides and at the a,pe\
acuminate: gradually tapering to a point.
agglutinated: glued together.
aliform: wing-shaped.
allantoid: sausage-shaped.
amerospores: conidia nonseptate, broadened.
amoeboid: amoeba-like, creeping movement.
amphigenous: not restricted to any particular surface.
ampuUiform: flask-shaped.
anastomosing: fusing together by growth.
.androgynous: with the antheridium and oogonium arising Q^ y^g game stalk
anteriorly: towards or near the head.
antheridium (a) : male sexual organ in fungi.
aparaphysate: without paraphyses or falsely paraphysate,
apiculate: furnished with a small pointed tip (apiculum).
apophysis: an offshoot.
apothecium(a) : cup-shaped or saucer-shaped fruiting boqy containing asci
applanate: flattened or horizontally expanded.
appressorium(a) : the flattened thickened end of a hyplia \vhich attaches a fungus
to a host or precedes invasion of it.
arachnoid: cobweb-like.
arcuate: strongly curved, as a bow.
areolate: marked out in little areas.
ascigerous: bearing asci.
ascocarp: a many-celled body producing asci.
ascospores: the spores borne in an ascus.
ascostroma(ta) : stroma producing asci.
ascus (i) : a spore sac characteristic of the Ascomycetes.
aseptate: without cross walls.
asexual: without sex.
asperulate: slightly roughened.
attenuated: tapered, drawn out.
bacillar: short rod-shaped.
basidiospore: spore formed on a basidium.
basidium(a): spore-bearing cell, the spores of which reSuH from meiosis, most
frequently being four in number on each basidium.
basipetal: developing from the apex towards the base.
bifurcate: two-forked.
botryose: grape-like clusters.
437
caespitose: in dense groups or tufts.
campanulate: bell-shaped.
carbonaceous: dark and carbon-like.
carnose: fleshy-appearing.
catenulate: in chains.
caulicolous: attacking stems or preferring stems.
chlamydospore: a spore with a thick membrane.
chlorinous: shghtly greenish-yellow, approximately hyaline.
cinereous: ash gray.
cirrhus(i): tendril-like curl of extruded material.
clavate: club-shaped.
clavulate: somewhat club-shaped.
cleistotheciuin(a): a perithecium without an oatiole as in Erysiphe.
clone: the progeny of a single plant by asexual subdivision.
clypeus(i) : a buckler or shield-shaped tissue around the mouth of a perithecium.
coenocytic: multinucleate, without cross walls.
coenogametangium(a) : multinucleate differentiated sac which produces gametes.
concolorous: of one color.
concrescent: growing together, fusing.
conic: cone-shaped.
conidiophores: hyphae which bear conidia.
conidium(a) : asexual spore borne on a ponidiophore.
continuous: aseptate, one-celled.
coremium(a) : a fascicle of parallel conidiophores.
coriaceo-membranaceous: leathery membrane.
cruciform: cross-shaped.
crustose: crust-like.
cupulate: cup-shaped.
cuspidate: toothed.
cyst: sac or cavity, thick-walled resting cell.
cystosorus(i) : sac-like sorus.
denticulate: minutely toothed. ,
diaphanous: having a texture so delicate as tojbe translucent.
diaphyllous: across the leaf. J
dichotomy: dividing into two subordinate par^s.
diclinous: having the fintheridium and oogonium on separate hyphae
digitate: divided hke the fingers on a human hand.
dimidiate: halved, especially unequally.
dimorphic: of two forms.
diplanetic: having two kinds of zoospoi-es.
discoid: resembUng a disc.
discrete: individual or separate, not in clusters.
distal: remote from point of attachment.
distichously: in a 2-ranked arrangement.
doleiform: barrel-shaped.
dothideaceous: like the Dothidiales with the peritheci'a embedded in stromata
v/ith no distinct perithecial walls.
eccentric: asymmetrical in growth.
echlnulate: spiny.
eguttulate: without guttulae.
•'ellipsoid: more or less elliptical.
emarginate: without margin.
GLOSSARY 439
encyst: to enclose or become enclosed in a cyst or capsule.
endospore: a spore formed endogenously by extrusion internally from the hypha.
Also the inner wall of a spore.
epigynous: on the oogonium. '
epiphyllous: growing on the upper surface of the leaves.
erumpent: breaking out.
evanescent: vanishing early.
exciple: outer wall or covering of an apothecium.
exospore: the outer covering of the spore.
extramatrical: outside of the mati-ix.
eye s p o t : an elliptical to circular spot with a prominent border and paler center
which resembles an eye.
falcate: sickle-shaped.
fascicled: in bundles.
ferruginous: rust-colored.
filamentous: thread-like.
filiform: thread-like.
fimbriate: with the margin bordered by long slender processes.
floccose: with tufts of soft woolly hair.
flccculose: with small tufts of soft woolly hair.
f o l i i c o l o u s : attacking or preferring the leaf.
fugacious: fleeting, disappearing early.
fuliginous: sooty or soot-colored.
fungicolous: attacking fungi or associated with their rhizosphere.
fusiform: spindle-shaped.
fuscous: dusky or brown.
fusoid: spindle-shaped.
gemma ( e ) : young bud or conidium-like structure such as in certain Phyco-
mycetes.
geniculate: bent or kneed.
geniscar: abscission conidial scars on the knees of conidiophores.
g l o b o s e : spherical.
g l o m e r u l e s : head-like compacted clusters.
granulose: granular.
gregarioiis: grouped or closely associated.
guttulate: containing one (uni-), two (bi-) or several (multi-) oil drops (guttulae).
haustorium(a) : a special branch of a filamentous mycelium serving as a feeding
organ.
h e l i c o i d : coil-like. _ - -
hemiparasite: a partial parasite.
heterokont: with a pair of unequal cilia.
heterostrophic: coiled in a direction opposite to the usual one.
heterothallic: a condition in which two distinct thalli miist Unite "for produc-
tion of sexual spores.
hilum: scar (as in Helminthosporium) at base of spore where it was attached to
the conidiophore.
hyaline: colorless or translucent.
hymenium: a fruiting layer of tissue.
hyperparasite: parasite on another parasite.
hypha (e) : thread-like vegetative part of a fungus.
hypochnoid: fruiting layer supported on a loose weft of hyphae as in Hypochnus
hypodermis: tissue lying under the epidermis.
440 DISEASES OF CEREALS AND GRAISSES
hypogynous: with the antheridium (a) formedbelow the oogonium and on the
same hypha. ^_ •
hypophyllous: on the under surface of the leaf.
hypostroma(ta) : tissue lying under the stroma, a foot-}ike base.
hypothecium: a layer of hyphal tissue just beneath the'hymeniiim.
hysterothecium(a): elliptical perithecium with a slit-hke ostiole
indeterminate: indefinite, unlimited.
innate: existing in.
inordinate: not in orderly pattern or arrangement.
intercalary: growth which is not apical but between the apex and the base. -
intercellular: between the cells.
intracellular: within the cells.
intramatrical: within the matrix.
isabelline: a dirty gray white to dirty tawny color.
lageniform: flask-shaped.
laniella(e): the gills of the Agaricales.
lanceolate: narrow, tapering towards each end, lance-like.
lanuginose: downy.
lenticular: lens-shaped.
lobulate: made into small lobes.
loculate: haying chambers or cavities (locales).
lunate: crescent moon-shaped.
macrocomdium(a) : large conidium as contrasted with small microconidium in
the same particular species.
macropycnophores: gives rise to macrospores.
maculiform: in the form of spots.
mammiform: breast-shaped.
mesially: medianly.
microconidium (a) : small conidium as contrasted with large inacroconidium in
the same particular species.
micropycnophores: give rise to microspores.
monocentric: having a single* center of growth.!
mucose: wet or yeasty-appearing.
muriculate: minutely roughened. ,
muriform: cross walls in more than one plane.
muticate: blunt, without appendages.
mycelium (a) : vegetative portion of the thallus of a fungusr~~~-,.^
mycorrhiza(e) : fungi attacking the roots of higher plants but both host and
fungus utilizing nutrients obtained from the rhizosphere, a mutual and usually
beneficial parasitism or symbiosis.
myxamoeba(e) : swarm cells with amoeboid creeping motion.
navicular: boat-shaped.
nodulose: with knobs, knots, or kneed swellings.
obclavate; club-shaped with the broad end at base.
obconical: inverted cone.
oblanceolate: with the broadened end of the lance at the distal end.
obpvate: egg-shaped with broadened end outward^^^
olivaceous: olive-colored. - , _ .^
oogomum(a): female sexual organ producing oospore(s).
oospores: products of fertihzation of the oogonium.
opalescent: like an opal.
operc'alum(a): lid.
GLOSSARY 441
orbicular: circular.
o s t i o l e : opening or moutli in the apical portion of the otherwise enclosed fruiting
body.
ovate-acuminate: egg-shaped (in section) but tapering to a point.
o v o i d : resembling an egg in shape.
papilla ( e ) : soft superficial protuberance.
paraboloid: the surface generated by the rotation of a parabola about its axis.
paragynous: beside or near the fruiting structure.
paraphysis (es) : sterile filaments accompanying fructifications.
p a r a p l e c t e n c h y m a t o u s : somewhat interwoven or webbed tissue formed by in-
terlacing of mycelium.
p a r i e t a l : the layer next to the wall in certain fruiting bodies.
parthogenetically: by means of reproduction of an unfertilized egg --
p e d i f o r m : foot-shaped.
peduncle: a flowering stalk.
penicillately: like the fingered dichotomous branching in blue mold (Penicil-
lium).
peftultimate: last but one.
peridium: the outer wall of the fruit body.
peritheciuni(a) : the covered but often mouthed fruiting body of certain orders
of Ascomycetes.
perthophyte: a "saprophyte" which kills cells in advance of actual mycelial in-
vasion by means of secreted toxic materials.
p i o n n o t e s : a slimy gelatinous mass in which conidiophores and conidia are
embedded.
pip-shaped: like an apple seed. •
p l e c t e n c h y m a t o u s : thick, tissue-like, formed by intertwining and adhering of
hyphae.
plerotic: filled up.
pleurogenous: growing from the sides.
plicate: folded.
pluriloculate: with several cavities (locules).
polycentric: having several centers of growth.
polyhedral: many-sided.
polymorphic: with several forms.
probasidium(a) : hypha which gives rise to a basidium.
p r o m y c e l i u m ( a ) : short and short-lived product of tube germination of a spore
which abstricts acrogenously a small number of spores.
prolate: extended.
prosenchymatous: relating to prosenchyma, in fungi often referring to un-
differentiated tissue filling a fruiting body, especially a young perithecium.
p s e u d o : false.
pseudomorph: a facsimile.
pseudoparenchymatous: hyphae compacted into isodiametric cells resembling
a true parenchyma as in many pycnidia.
pseudopionnotes: a slimy gelatinous mass of conidia with no evident conidio-
phores present.
pulverulent: powdered, as though dusted over.
pulvinulus(i) : a small raised area.
punctiform: in the form of a point or dot.
pycnidium(a) : the covered but often mouthed fruiting body of the Sphaeropsi-
dales of the Fungi Imperfeoti.
./ j
pycnophores: hyphae on the inner wall of the pycnidiurii which produces pyenb-
spores.
pycnospores: conidia produced in a pycnidium.
pycnosclerotium(a) : sclerotium which bears pycnidia(um)
pyriform: pear-shaped.
ramose: branched.
refringent spot: shiny, light-reflecting globule.
reniform: kidney-shaped.
reticulate: netted.
rhizoid: a root-like structure in certain fungi.
rhizomorph: a rope-like or root-like branched strand of entwined mycelium.
rhizosphere: the restricted region in the soil adjacent/ to the roots and root
hairs of plants.
rostrate: beaked.
rugulose: wrinkled. _
saccate: bag-shaped.
sarcinaeform: packet-shaped.
sclerotic: hard and sclerotia-like. ~ '
sclerotium ( a ) : firm or hard, tightly packed masses of mycelia usually white
underneath a black oxidized rind. They are resting bodies.
scolecospores: elongated spores typically ^hform or variously club-shaped and
usually septate.
scorpioid: with the main axis coiled like the tail of a scorpion.
scutellum: part of the seed lying between the embryo and the endosperm as in
corn,
septum (a) : partition, as in spores or mycelia.
sessile: without a stalk.
seta(e) : bristle.
sinuous: serpentine or with a deep wavy ma'rgin.
sigmoid: curved like an open letter C or S.
slurry: thin \vatery mixture. /
sphacelial: usually referring to the conidial stage of Claviceps.
sphaeroidal: somewhat spherical.
sphaerosporangium(a): spherical sporangium'.
sporangiophore: stalk bearing a sporangium. '
sporangium (a) : sac producing spores endogenously.
sporodochium(a) : a pad or tubercle-like mass of mycelhinulêaring conidia.
sporogenous: producing spores. , '
sporophore: spore-bearing organ. —
sterigma(ta) : the small spiculum-like extension of a basidium bearing a basidio-
spore, but less often used in downy milde-w \Bremia) terminology
striate: minute furrows or lines. .
stroma (ta) : a cushion-like mycelial aggregate. ,
subglobose: nearly globose.
subiculum(a) : a felted or cottony basal stratum of hyphae.
subtorulose: cylindrical with moderately swollen areas-at intervals.
subulate: awl-shaped. •'
tenuous: delicate.
terete: civcular in transverse section.
testaceous: shell tinted.
textura porrecta: web extended in a horizontal plane.
thallus: a vegetative body without differentiation into special organs.
GLOSSARY 443
toruloid: cylindric with swollen portions at intervals.
trifurcate: 3-forked.
truncated: wedge-shaped but as though cut off at the distal portion.
tubiform: tube-shaped. '
turbinate: shaped like a top or inverted cone.
uniaxial: having b u t one axis.
uniciliate: with a single whip-like oilium.
uniguttulate: with one guttula (cell inclusion).
uniseriate: in a single series or row such as spores arranged in a narrow ascus,
all in direct line, one above the other.
v e n t r i c o s e : inflated, sometimes unequally so on one side, as in Curvularia.
vermiform: worm-shaped.
verrucose: warty.
verticillate: whorled.
verruculose: minutely warty.
villose (villous): covered with long weak hairs.
v i n a c e o u s : wine-colored.
zonate: with concentric bands.
zoosporangium(a) : sac which produces zoospores endogenously.
z o o s p o r e s : motile spores.
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446 DISEASES OF C E R E A L S A N D GRASSES
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INDEX OF FUNGI
Acrothecium lunatum 327-28 Cercospora 305
Alternaria tenuis 301-2 agrostidis 306
Ansatospora bromi 418-20 asprellae 306
Aphanomyces camptostylus 5-7, 7 * boutelouae 306
Apiocarpella agropyri 150 bromi 307, 418-20
macrospora 151 caespitosa 307
minor 150-51 echinochloae 307
Ascochyta 151-52 festucae 307-9, 308
agropyrina 152-53, 154 fusimaculans 309
avenae 153, 154 longipes 309-10
boutelouae 155-56 muhlenbergiae 310
brachypodii 153-54, 154 oryzae 310-11
calamagrostidis 227-29 panioi 311
oynodontis 155-56 paspali 311-12
desmazieri 155 scolecotrichoides 312
elymi 159-61 seminalis 312-13
graminea 154, 155-56 seriata 313
graminicola 159-61 setariae 313
graminicola var. diedickeana 159-61 setaricola 313-14
graminicola var. holci 159-61 sorghi 314-15, 315
graminicola var. leptospora 159-61 tesseliata 315-16
hordei 156-58, 157 vaginae 316
maydis 158' zeae-maydis 316-17
missom-iensis 157, 158 Cercosporella 317
oryzae 159 herpotrichoides 317-321, 318, 319
phleina 157, 159 holci 321-22, 322
sorghi 154, 159-61 poagena 322-23, 323
sorghina 161-63, 162 subulata 431-33
spartinae 254-55 Cerebella andropogonis 323-24
subalpinus 157, 163 . Cladoohytrium graminis 7-8
utahensis 157, 163 maydis 11-13
zeae 164 Cladosporium graminum 89
Ascochytella avenae 153 herbarum 88-90, 324
Ascochytula agropyrina 152-53 Claviceps cinerea 59-60
Asoomycetes 53-54 paspali 60-61
Aspergillus circinatus 413-15 purpurea 61-66
Asterocystis radicis 10-11 rammculoides 66
rolfsii 66-67
Balansia claviceps 54-55- tripsaci 67
clavula 55-56 yanagawaensis 67
hypoxylon 56-58, 57 Cochliobolus heterostrophus 68-69
pilulaeformis 58 miyabeanus 69, 370-72
trinitensis 59 sativus 69-70, 376-81
Basidiomycetes 125 setariae 70, 381
nonsporulating 145-46 tritici 69-70
Botrytis cinerea 302-3 CoUetotrichum cereale 286-88
Brachycladium spiciferum 329, 389 falcatum 117, 285
trifolii 328-30 graminicola 286-88
lineola var halepense 117
Calonectria graminicola 344-47 Coniosporium arundinis 324, 411
Centrospora bromi 303, 418-20 shiraianum 324, 411
Cephalosporium acremonium 303-5 Coniothyrium psammae 165
Ceratostomella adiposa 74-75 zeae 165
paradoxa 75-76 Coprinus urticaecola 129
* Page numbers in boldface refer to illustrations.
525
526 I N D E X OF F U N G I
/
Corticium fuciforme 126-27 culmorum 337-39
rolfsii 136-37 eijuiseti 339 ,
vagum 130-36 graminearum 340-41
Crypt oascus 70 moniliforme 341-44
Curvularia 324 ' nivale 3^4-46
cymbopogi 329 oxysporum 346-47
geniculata 324-26, 325 poae 347-49
inaequalis 327 roseum 331
lunata 327-28 scirpi 339
trifolii 328-30, 329 var. acuminatum 333-38
Cylindrosporium bambusae 288 sporotrichoides 347-49
calamagrostidis 288-89, 289 Fusicladium alopecuri 349
glyoeriae 289 destruens 349
infuscans 237-38
Cytodiplospora elymina 167-68 Gibberella fujikuroi 82, 341-44
Cytospora sacchari 165-66, 16S moniliformis 82. 341-44
saubinetii 82, 340-41
Dactylaria graminum 330 zeae 82
Darluca filum 166-67 Gliocladium 351
Davisiella elymina 167-68 Gloeocercospora sorghi 350-51
Dematium scabridum 74-75 Gloeosporium boUeyi 289-92, 291
Dilophospora alopecuri 168-69 "graminum 292
Diplodia bambusae 169 meinersii 292-93
frumenti 120, 169, 170 Gnomonia iliau 82-83
gossypina 117, 121
macrospora 170, 170-71 ,Hadrotrichum lineare 351-52
natalensis 117, 121, 171-72 /• phragmitis 351
tubericola 117, 121, 171 Helicoceras oryzae 352
zeae 171-72 Helminthosporium 352, 397-99
Diplodina brachypodii 153-54 subgenus Cylindro- 362-63
' graminoa 155-56 subgenus E u - 362-63
lolii 155 arundinaceum 405-6
Discellomyces gloeosporus 127 avenae 353-55
Dothicbloe 70 bromi 355-56
aristidae 71 buch!o(3S 356-57 -
limitata 71-72 carbonum 357-58
nigricans 72 catenariupi 358-59
Dothidella aristidae 73 Cyclops B59-60
minima 73 cyiiodontis 360-61
dematioideum 361
Ellisiella caudata 330-31 diq'tyoides 361-62
Endoconidiophora adiposa 74-75 erythrospilum 362-63
paradoxa 75-76 giganteum 363
Endoconidium temulentum 98 gramineum 363-65
Endodothella traoyi 76-77 hadrotrichoidra-365-fi6
Entyloma 127-28 halodes 366-67 - """ '
bingenensis 128 inaequalis 327 —
erastophilum 128 inconspicuum 393-94
irregulare 128 irregulare var. microsphaeroides
spragueanum 128 84-86
Ephelis borealis 56 leersii ^367
mexicana 55 leucostylum 365-66
Epiohloe hypoxylon 56-58 mayaguezense 68-69
nigricans 72 maydis 68-69. 367 '
typhina 77-78 micropus 367-68
Epicoccum 331 monoceras 369 ,
Erysiphe graminis 78-82 nodulosum 369-70
Euryachora (?) aristidae 73 ocell]iriP^75-76'
oryzae -370-72
Fungi Imperfecti 148 poae 372-73
Fusarium 331-33 ravenelii 373
acuminatum 333-36 rostratum 373-75, 374
> , avenaqeum 336-37 sacchari 375-76
bartholomaei 294 sativum 376-81
INDEX OF FUNGI 627
setariae 381-83 Melanoonium bnmbusae 121
siccans 383-84 iliau 82-83, 293
sigmoideum 84-86, 384 sacchari 293-94, 293
var. irregulare 84, 86, 384-85 Melasmia setariae 184-85
sorghicola 385-86 Microdiplodia 185
sorokinianum 376-81 Moniliales 299-300
stenacrum 386-87 Monotospora oi-yzae 406
stenospilum 387 M.ycosphaerella calamagrostidis 87-88
teres 387-89 holci 88
tetramera 389-90 tassiana 90
triseptatum 390-91 tulasnei 88-90
tritici-repentis 391-92 zizaniae 90
tritici-vulgaris 392-93 Myriogenospora aciculisporae 90-91
turcicum 393-94 paspali 91
vagans 394-95
victoriae 395-97 Napicladium arundinaceum 405-6
zeicola 397 Naucoria cerealis 129
Hendersonia 172-74 Nigrospora oryzae 406
crastophila 174-76 sphaerica 407
culmicola 176-77 Nonsporulating Basidiomycetes 145-
graminis 174-76 46
nodorum 244-46
simplex 176-77 Olpidiaster radicis 10-11
subseriata 281-82 Olpidium brassicae 10-11, 11
Heterosporium avenae 399-400 radicicolum 10-11
phlei 400-1 Ophiobolus cariceti 92-95
Hyalothyridium calamagrostidis 177 graminis 92-95
Hyphochytrium catenoides 8 herpotrichus 94
Hypocrea subviridis 58 heterostrophus 68-69
oryzinus 95-96
Lagena radicicola 9-10, 10 sativus 69, 96
^ Leptosphaeria avenaria 83, 220 setariae 70
f. sp. triticea 83 Ophiociadium hordei 407-9
herpotrichoides 83-84 Ovularia hordei 407-9, 408
sacchari 84. 197 lolii 409
salvinii 84-86 pulchella 409-11
Leptosti'omella c3Tiodontis 177 var. agrnpyri 410
panici 177-78 pusilla 409-il
Leptothyrium avenae 178
cylindripum 178 Papularia sphaerosperma 411
zeae 179 Pellicularin filamentosa 130-36
Lophodermium arundinaceum 86-87 rolfsii 136-37
Lunospora 201 Penicillium expansum 411-13
oxyspora 203-6 oxalicum 413
Perieonia circinata 413-15
Macrophoma 179-81 Peronospora graminicola 48-50
oblongata 180-81 Phaeoseptoria 185-87
phaseoli 183-84 ' airae 186
phnseolina 183-84 festucae 186
phlei 181-82, 182 phaiaridis 188
secalina 182 Phialea mucosa 96-100
zeae 119. 183 tomulenta 96-100
Macrophomina phaseoli 183-84 Phleospora 187
phaseolina 183-84 gramincarum 187, 187, 189
Marasmius interstitians 129 idalioensis 188, 190
oreades 129 muhlcnbergiae 187, 188-90
sapchari 128-29 Pholiota dura 129
tritici 129 praecox 129
Mastigosporinm 401-2 Phoma 190-91
album 402-5 secalina 253-54
cylmdricum 402 Phycomvcetos 5
rubricosum 402-6, 404 Phyllachora 100-3
Melanconiales 285 acuminata 105-6
528 INDEX OF FUNGI
Phyllachora Elij^topthora 'eactorum 13-14
ammophilae 103 parasitica jl4
aristidae 73 Piricularia grisea 415-16
arundinariae l04 , oryzae 416-17
boutelouae 104 parasitica , 1417-18
chardonii 104 Placostroma bambusae 121
coloradcnsis 105 spoioboli J21-22
cornispora 105-6 Pleospora 122
diplocarpa 106 a\-cnae 122
epicampis 106 Potymyxa graminis 15-16, 16
eragrostidis 107 Protomyces graminicola 48-50
erianthii 107 Pseudodiscosia avenae 430-31
gramiais 107-8 PuUularia puUans 99
guianensis 108-9 Pyrenochaeta terrestris 198-200
heterospora 109 Pyrenophora avenae 122
iusularis 109 bromi 122
lasiacis 109 Pythium 16-20, 44-46
lepthochloae llQ aphanidermatum 20-21
luteo-maculata n o aristosporum 28-33
maydis 110 arrhenomanes 28-33
microstoma 105 artotrogus var. macracanthum 21
minima 104 butlcri 22
murilloi 105-6 debaryanum 23-27
nervisequia 111 dissotocum 27-28
nuttalliana 106 graminicola 28-33
oryzopsidis 111 hypogynum 33
pamelii 111-12 inlermedium 45
parilis 112 irregulare 34-35
paspalicola 112 megalacarithum 45
phalaridis 112-13 monospermum 35
punotum 113 nagaii 36
quadraspora 113-14 oligandrum 36-37
serialis 114 oryzae 27-28 !
silvatica 114-15 ostracodes 37-38
spartinae 115 periilum 39
spliaerosperma u s rostratum 39-40
splendens 140
texensis 115 tardicresc;6ns 41-42
tracyi 76-77 ufti'rnum 42-43
vaginata 112 voliitum 43-44
vulgata 116 I
•wilsonii 116-17 Ramu!aria graminicola 418
Phyllosticta 191-93. 198, 259 Ramulispora andropogonis 420-21
anthoxella 192, 192 bromi 418-20,^19
glumarum 193-94 sorghi 420-21 "^ ---^_
healdii 192, 194 Rhizoctonia monteithianum 123-24
helenae 192, 194-95 oryzae 137-38 —
japonica 195 solani 130-36, 133, 138-39
minutaspora 19^, 195 zeae 138-39
miyakei 195 Rhizophidium graminis 46-47, 47
owensii 198 Rhizopus/ 47
panici 198 Rhyncho^porium 421, 421
rogleri 192, 196 orthosporum 421-22, 421
saceharicola 84, 197 secalis 422-24, 424 '
sorghiaa 192, 197-98
stomal icola 206^9 Scaphidium boutelouae 200, 200
Physalospora rhodina 117 < Scirrhia âmbusae'' 121
tucumanensis II7-I9, 285 sporoboli' 121-22
zeae 119-20 Scirrhodothis bambusae 121
zeiviola 120-21 Sclerospora farlowii 47-48
Physoderraa 13 graminicola. 48-50
graminis 13 kriegoriana 60-51
' < maydis 11-13 âcrospora 50-51
zeae-maydis IU13 oryzae 50-51 '
INDEX OF FUNGI 529
Sclerotinia homoeocarpa 123-24 mississippiensis 243-44
Solerotium 139 munroae 244, 244
bataticola 183-84 murina 249-51
hydrophyllum 139-40 neglecta 264-65
oryzae 140 nodorum 235, 244-46, 246
paspali 60-61 oryzae 246
rhizodes 140-41 oudemansii 246-47, 247, 256-67
rolfsii 136-37 oxyspora 203-6
Scolecotrichum graminis 424-29, 427 pacifica 247-49. 248-49, 256-67
maculicola 429-30 passerinii 249-51, 250, 256-57
Selenophoma 201 pertusa 251
bromigena 201-2, 202 phalaricola 226-27
donacis 202, 203-6, 205 phleina 262-63
var. stomaticola 206-9 poae-annuae 241-42
drabae 207 poliomela 251-52, 252
everhartii 209-11 quinqueseptata 252-53, 253
obtusa 202, 211 secalina 253-54
Septogloeum bartholomaei 294 secalis 253-54, 255
oxysporum 294-97, 296, 297 var. stipae 254, 255, 262-63
spartinae 297-98 simplex 176-77
Septorella sorghi 420-21 spartinae 254-55
Septoria 211-15 stipina 255-57, 262-63
agropyri 232-35 tandilensis 257, 259
agropyrina 215-16, 216, 234, 248-49 tenella 258, 259-60
agrostidis 227-29 trissti 256-57, 261, 262-63
andropogonis 216-18, 217 tritici 256-67, 261-65, 284-65, 266
var. sorghastri 218 f. avenae 256-57, 264-65, 265-66,
f. sporobolicola 217, 218-19 266
annua 241, 241-42 f. holci 256-57, 267, 268
arctica 219-220, 220, 256-57
arechavaletae 220 var. lolicola 267-69. 268
avenae 220-22, 221, 224 Spermospora avenae 430-31
f. sp. triticea 222-24, 224 subulata 431-33, 433
brevispora 224-25, 241 f. cijiata 432
bromi 225, 226, 241, 256-57 Sphaeronema adiposum 74-75
var. phalaricola 225-27 Sphaeropsidales 149-50
bromivora 241 Sporotrichum colubiense 433-34, 434
calamagrostidis 227-29, 228 peribebuyense 434, 434
f. koeleriae 229-30, 256-57 Stagonospora 269-70
calamovilfae 230, 230 agrostidis f. angusta 270-71, 270
carricerae 230-31 arenaria 221, 271, 272, 279
cenehrina 231 arrhenatheri 220-22
culmifida 206-9 brachyelytri 272-73
cynodontis 231 bromi 273, 273, 279
digitarivora 232, 232 curvula 274
donacis 203-6 elegans 283
elymi 232-35, 234, 256-57 foliosa 274-76, 275
f. elymina 234, 257 glycericola 276
elymi-europaei 248-49 graminella 282, 283
everhartii 209 graminum 276
festucae 258-59 intermixta 276-77
glycericola 235-37, 235 ischaemi 275, 277
graminum var. lolii 155 maculata 278, 281
infuseans 236-37, 238, 237-38, 256-67 maritima 278
jaculella 238-39, 240-41, 256-57 paspali 279, 279
koeleriae 229-30 simplicior 157, 163, 279-80
loligena 239 var. andropogonis 281
lolii 155
spartinicola 280-81
macropoda 241-42, 241 subseriata 279, 281, 281-82
var. grandis 242, 242, 256-67 var. maculata 278
var. septulata 242-43 vexata 282-84
melicae 221, 243 var. foliicola 274-76
microspora 250 vexatula 282
530 INDEX OF FUNGI
Stemphyllium botryosum 435 •graminum 142
consortiale 435-36 idahoensis j 142-43
Synchytrium graminicola 51-52 itoana 143-45
Thielaviopsis ethaceticus 75-76 Ustilaginoide'a^ oryzae 436
paradoxa 75-76
Titaeospora andropogonis 420-21 virens 436 ,
Trichofusarium bartholomaei 294
Typhodium typhinum 77-78 Wojnowioia graminis 174-76
Typhula 141
borealis 142 Xylaria clavulus 55-56
INDEX OF CEREAL AND GRASS HOSTS
Aegilops cylindrica 23, 152, 318, 334 smithii 23, 29, 42, 61, 77, 79, 88, 108,
ovata 318, 334 130, 142, 152, 154, 160, 168, 174, 199,
triuncialis 23, 29, 318, 334 203, 215, 223, 233, 234, 244, 248-49,
Agropyron albicans 61, 203, 289 271, 286, 290, 334, 337, 342, 344, 377,
angustiglume 29, 130, 289, 334 391, 425
bakeri 152 spicatum 29, 61, 77, 79, 144, 152, 153,
buonapartis 376 160, 168, 176, 203, 211, 233, 237, 244,
caninum 23. 42, 61, 152, 215, 334, 424 271, 290, 295, 296, 334, 344, 399, 425,
ciliare 23, 29, 130, 198, 234, 271, 334, 434
376 striatum 79
cristatum 14, 23, 29, 34, 42, 61, 79, subsecundum 23, 62, 79, 108, 130,
108, 130, 142, 152, 174, 198, 203, 271, 140, 152, 153, 174, 203, 233, 290,
289, 290, 318, 334, 337, 339, 344, 376, 334, 339, 344, 377, 391, 399, 421,
394, 397, 400, 410, 412, 414, 424 422, 425, 434
dasystachyum 29, 61, 79, 130, 203, var. andinum 211
344, 422, 424 trachycaulum 10, 23, 29. 31, 34, 62,
desertorum 23, 29, 42, 61, 130, 160, 77, 79, 92, 108, 130, 146, 152, 160,
198, 271, 286, 289, 334, 337, 344, 377 161, 163, 168, 174. 176, 199, 203, 215,
elmeri 422 223, 233, 244, 248-49, 271, 286, 290,
elongatvim 29, 130, 146, 198, 286, 289 325, 330, 334, 337, 344, 377, 410, 414,
334, 337, 344, 377 422, 425
elongatum X Triticum aestivum trichophorum 23, 29, 62, 79, 130, 152,
337, 344 271, 290, 334, 344, 377
gamelinii 61 Agrostis 340
griffithsii 61, 289, 324, 334, 424 alba 7, 23, 29, 42, 51, 62, 77, 79, 92,
inprme 23, 29, 61, 77, 79, 87, 142, 108. 123, 126, 130, 140, 146, 160, 261,
157, 163, 168, 174, 203, 211, 215, 233, 262-63, 277, 286, 290. 295, 334, 337,
234, 318, 334, 337, 344, 414, 424, 434, 344, 361, 362, 377, 386, 390, 403, 410,
434 421, 422, 425, 431, 432
intermedium 23, 29, 34, 42, 61, 130, canina 51, 62, 123, 126, 130, 160, 334,
142, 174, 198, 286, 289, 334, 344, 377, 344, 350, 362, 403
391, 414 castellana 261, 425
junceum 61 cernua 414
michnoi 23, 29, 61, 130, 152, 199, diegoensis 79, 209, 228-29, 229
289, 334, 344, 377, 414 exarata 51, 62, 79, 130, 146, 221, 228-
29, 229, 286, 295, 344. 390, 403, 425
mongolicum 23, 29, 130, 199, 289, var. ampla 261, 262-63
334, 377 liallii 51, 130, 254, 255, 262-63, 295,
orientale var. lasianthum 29 296, 425
pendulinum 29, 199, 203, 377 humilis 410
pertenue 215 oregonensis 87, 386, 410, 425
pubicalum var. pilipes 29, 334 palustris 22, 51, 62, 99, 123, 130, 144,
repens 9, 10, 15, 23, 29, 39, 42. 61, 65, 181, 182, 228-29, 229, 286, 334, 337,
79, 81. 92. 118. 130, 150, 159, 160, 344, 350, 362, 386, 398. 403, 410
161, 174, 176, 188, 199, 203, 215, 233, perennans 306, 361, 362
234, 244, 248-49, 271, 286, 289, 295, rossea 229, 425
334, 344, 362, 363, 366, 377, 391, 410, scabra 23, 62, 77, 140. 209 229, 254,
422, 424, 424 286, 290, 295, 306, 334, 386, 425
riparium 23, 29, 61, 79, 130, 174, 203, var. geminata 425
289, 318, 334, 344, 377, 424 stolonifera 20, 62, 130, 261, 403, 415
semicostatum 23. 29, 42, 61, 152, 176, tenuis 51, 62, 123, 126, 130, 144, 258-
199, 215, 289, 344, 377, 414, 422 57, 261, 252-63, 337, 344, 350, 362,
sibirioum 13, 23, 29, 61, 79, 130, 146, 386, 403, 410
152, 199, 289, 334, 337, 342, 344, 377 verticillata 79, 403
B:\\
532 INDEX OF CEREAL AND GRASS HOSTS
Aira 65 Axonopus affinis 72, 130, 323, 398, 399
caryophyllea 97, 252 compressus 284, 323
Alopecurus aequalis 62, 286, 425
alpinus 410 bamboo 121, 206, 288, 399
carolinianus 425 Bambusa 169, 411
geniculatus 62, 97, 349, 425 Beckmannia syzigachne 13, 23, 46, 79,
pratensis 62, 286, 398, 421, 425 154, 160, 255, 286, 393, 399, 425, 427,
Ammophila arenaria 29, 62, 103, 377 429
breviligulata 286 Blepharoneuron tricholepsis 116
Andropogon 71, 110, 111 Bouteloua curtipendula 23, 29, 62, 72,
annulatus 330 86, 104, 130, 151, 167, 183, 199, 200,
elliotii 111 200, 209, 216, 271, 308, 334, 347, 357,
fastigiatus 114 377
furoatus 23, 29, 42, 62, 77, 110, 154, gracilis 14, 23, 29, 34, 62, 77, 104, 130,
154, 167, 199, 216, 217, 275, 277, 280, 151, 154, 155, 174, 199, 290, 306, ,334,
286, 290, 323, 330, 334 339, 347, 357, 377, 422
glomeratus 111 hirsuta 104. 357
hallii 23, 29, 34, 62, 110, 199, 277, rigidseta 151
286, 290, 330 334, 377, 414 Brachiara dictyoneurum 398
ischaemum 323 extensa 62, 323
longiberbis 111 Brachyelytrum erectum 108, ?72
perforatus 110 Brachypodium mucronatum 286
saccharoides 110, 323. 399 pirinatum 398
scoparius 23, 62. 71, 77. 91. 110, 154, sylvaticum 64
216, 286, 290, 330, 334, 414 Briza media 126
teraarius 111, 323, 330 Bi-omus 198
virginicus 71, 86, 110 anomalus 290
Anthaenantia rufa 286, 323 arenarius 425
Anthoxanthum odoratum 51, 192, 192, arvensis 23, 29, 42, 81, 130, 199, 290,
286, 361 334, 377
Arctophila fulva 62 breviaristatus 79
Aristida curtisii 330 brizaetormis 29 i
dichotoma 73, 286 oarinatus 23, 29. 34, 42, 51, 62, 79,
glauca 57 92. 130, 142, 156, 157, 160, 199, 201,
longiseta 209, 290, 334 225, 239. 240-41, 273, 279, 286, 290,
oligantha 209 295, 318, 334, 337, 344, 356, 377, 400,
purpurasoens 71, 73 410,1422, 425, 427
purpurea 29 cathartirus 79, 130, 273, 286, 377,
striota 73 398.. 425
Arrhenatherum 65 ciliatus 62, 108. 140. 201, 202. 224,
elatius 23, 29, 34, 62. 65. 130. 136, 225,' 239, 240-41, 273, 273, 279, 356,
160, 206, 208, 221, 221, 286, 293, 298, 425'
353, 410. 425, 430 commutatus 50^79. 151, 160, 225,
Arthraxon ciliaris -49 226-27, 256-57, 334r344„
hispidus 377 erectus 23. 29. 42, 62, 64, 79, 130, 199,
var. cryptatherus 199, 414 290, 334, 344, 377 -
Arundinaria gigantea 73, 86, 140, 178, frondosus 273, 422. 425
411, 425 inermia 23. 29, 34, 42. 62. 65, 79, 130,
tecta 104, 127, 312 142, 150, 154, 160, 161, 174. 199, 201,
Arundo donax 203. 411, 429 202, 225; 273, 286, 290. 318, 334, 337,
Avena barbata 353 344, 347, 356, 367, 377, 398, 410, 414,
byzantina 6, 23. 29. 34. 50, 79. 92, 422, 425
130, 153, 154, 265. 266. 290, 337, 344, japonicus 24, 29, 93, 131. 160, 225,
347, 353, 377. 394. 430 226. 290. 318, 334, 344, 356, 377
fatua 13. 14. 23. 29. 41, 79. 130. 153, kalmii 273 ,
265. 286. 290. 337, 353, 377, 414 laevioes .239. 256-57. 425
hookeri 62. 286 latiglunHS''225. 2'?6-27. 2<0-41, 286
nuda 23, 29, 34, 414 madritensis 24, ,?9. 130. 377
sativa 6, 10, 13, 23, 24, 33 34. 39. 42, mollis 79, 131, 226-27, 302, 334, 337,
44, 50. C2. 70. 79, 9'?. I.SO, 136. 137, 344. 425
153, 1.54, 160. 174 178, 193, 199, W% orcunianus 93, 425 '
' '221, 221, 224, 264-65, 265, 290. 302, paeificus 425
'318, 337, 340, 342. 344, 347. 349. 3-53. pumpellianus 29, 62, 131, 334, 337,
377, 394, 400, 407, 411, 414, 425, 430 356, 422
I N D E X O F C E R E A L AND GRASS HOSTS 533
purgans 108, 151, 201, 225, 273, 344, Cymbopogon citratus 91, 375
425 Cynodon dactylon 24, 48. 79, 131. 136,
racemosus 79, 97, 131, 225, 344 154, 155, 177, 199, 231, 286. 290, 312,
ramosus 273 359, 363, 393, 399, 411, 415
rigidus 79, 131, 239, 240-41, 256-57, Cynosurus cristatus 425
344, 356, 419 echinatus 97
secalinus 62, 90, 93, 131, 225, 223-27,
286, 344, 356, 422 Dactylis glomerata 24, 29. 42. 51, 62,
sitchensis 4?5 77. 79. 131. 144. 160. 174. 193. 199,
sterilis 64, 79, 318 206, 208, 221, 271, 272, 278, 279, 281,
suksdorfii 79 285, 293, 327, 334. 337, 338, 344, 348,
tectorum 24, 29, 34, 45, 79, 93. 131, 377, 395, 398, 399, 403, 404, 414, 421,
137, 142, 144, 160, 174, 239, 210-41, 425
290, 318, 334, 337, 340, 344, 347, 356, Dactylootenium aegyptium 315
377 Dantlionia californioa 97, 131, 207, 286,
texensis 225 359 425
trinii 108 compressa 56
vulgaris 79, 93, 176, 243, 402, 419, intermedia 87, 282
425, 432 parryi 62. 400
BUchloe dactyloides 79. 104, 131, 286, pilosa 334
290, 312, 334, 357, 399 sericea 56
spicata 58, 62, 77, 207, 359, 422
Calamagrostis breweri 87. 344 Descâmpfîa atropurpurea ,131, 209,
canadensis 62. 71, 77. 79. 87, 108, 140, 245. 276, 425
146, .151, 160, 168, 177, 209, 221, caespitosa 62. 97, 168. 176, 186, 209,
286, 288, 295, ?97, 344, 351, 393, 252. 282, 425, 432, 4H3
403, 410, 421. 421, 425 danthonioides 93, 209, 252, 252, 393,
var. scabva 62, 87, 174, 403, 425 425
epigeios 290 elonâta 142. 157, 160, 163, 209, 245,
inexpansa 62, 71, 209, 210, 285, 295, 334. 393, 425
425 Diarrhena americana 286
koelerioides 209 Digitaria filiformis 288
montanensis 131, 209, 290, 334 sanguinalis 79, 131,192, 193. 232. 232,
neglecta 62, 140 288. 334. 377, 398, 399, 415, 425
nutkaensis 62, ]65, 219, 220, 258-57 serotina 415
purpurascens 286 violasoena 198
rubescens 79, 146, 174, 209, 286, 295, Distiohlis snicata 62, 73, 76, 108, 298,
431, 432 324, 366
scribneri 209, 432 stricta 24 73. 76. 106, 112, 157, 163,
Calamovilfa gigantea 151. 288. 299 174. 199. 290. 334
lon<5ifolia 24. 29. 174. 193. 199. 203, Dupontia fischeri 219
230, 230, 281, 286, 290, 334, 342, 377,
414 Ecliinoch'oa colonum 85. 131, 140, 415
Catabrosa aquatica 79 crusgalli 13, 24, 29. 41, 42, 131. 199,
Cenohrus biflorus 198. 398 ?88. 2Q0. ,3"7. 309. 314. .^''S. 329,
echinatus 54, 115,'340_ 329. 334. 338. 340. 363. 369. 377
incertus 115, 198 var. frumentacea 199. 290. 334. 337
myosuroides 231 Eleu.sinc indica 131, 286, 360, 363, 365,
paucifloms 29, 131, 199, 231, 283, 290, 370 386
334. 377 Elymus ambiguus 62. 174. 290. 334
tribii]oides 285 antarcticus 131, 223, 290, 425
viridis 115 arenioola 62
Chloris chlovidea 104 aristatus 203. 4^5
gayana 395, 398 canadensis 24. 29. 42. 62. 77. 79. 108,
orthonoton 104 131, 146, 1.52 IH. 160. 167. 187,
petrae 71, 307 188. 189. 199. 203. 215. 233. 245,
submutira 104 248-4«). 271. 286. 290, 291, 334, 344,
verticillata 286, 377 377. ?91. 422. 425
virgata 48 71. 104. 154. 334 var. robustus 62 233, 334, 377
Cinno anindinacea 62, 77, 79, 108, 277, cap\it-medusae 207
359. 425 condensafus 29. 62, 79, 103. 154, 160,
latifolia 245, 271, 277. 425 161, 203. 211, 215. 233-37, 237,
Cortaderia selloana 198. 340, 398, 411 233, 290, 295, 296, 338, 399, 400,
Ctenium aromaticum 71 425
Elymus condensatiis Festuca- arizonica, 87
var. pubens 237, 249 aruudinacea 64
curvatus 62 capillata 129 ..
dahuricus 24, 29, 42, 62, 77, 79, 131, dertonensis 93,1114, 131, 253-59, 259,
199, 271, 290, 377 334, 344
dasystachvus 290, 334, 377, 414 elalior 6, 24, 29, 34, 62, 64, 93, 97, 99,
excelsus 24, 29, 223, 245, 290, 334 108, 131, 158, 157,' 190, 207, 221,
europaeus 248-49 259, 286, 290, 307, 308, 327, 335,
flavescens 202, 203, 425 342, 344, 362, 377, 414, 425
giganteus 29, 62, 131, 215, 245, 290, var. arundinacea 24, 29, 34, 62,
334, 425 131, 134, 290, 307, 335, 344, 362
•glaucus 24, 29, 51, 62, 79, 93, 108, gigantea 258-59
131, 144, 160, 168, 176 211, 215, 223, idahoensis 30, 62. 79, 88, 114, 131,
233, 234, 237, 245, 253-57, 271, 272, 174, 176, 188, 190, 190, 207, 208, 259,
290, 295, 296, 334, 337, 344, 391, 410, 344, 410
421, 422 425 kingii 30 62, 207, 208, 407
hirsutus 391 mairei 259
innovatus 62, 422, 425 megalura 24, 93, 114, 131, 286, 334,
intermptus 24, 29, 131, 199, 290, 334, 344, 410
377 myuros 93, 97, 126, 131, 334, 344, 410
junreus 24, 29, 34, 62, 79, 131, 199, obtusa '245
203, 215, 223, 245, 283, 293, 334, 348, occidentalis 114, 190, 361
377, 391, 414, 422 octoflora 24, 30, 62, 93, 131, 199, 207,
macounii (hybrid) 21, 29, 62, 131, 209,' 258-59, 259, 286, 290, 335, 348,
154, 160, 203, 215, 290, 334, 344, 377, 377, 414
425 ovina 24, 62, 79, 123, 131, 160, 168,
mollis 62, 248-49, 249, 256-57, 281, ^ 207, 259, 286, 290, 335, 344, 398
282, 425 var. brachyphylla 87, 176, 209
pseudoagropyron 62, 271, 293 var. duriuscula 126
riparius 108 pacifica 24, 174, 176. 290, 362
salsuginosus 62, 293, 414 rubra 6, 24, 62, 65, 77, 79, 114, 123,
sibiricus 24, 29, 79, 131, 199, 223, 245, 126, 131, 154, 160. 174, 176, 186,
290, 334, 344, 377. 425 190, 199, 209, 258-59, 259, 286, 290,
triticoides 62, 79. 108, 114, 168, 181, 327, 335, 344, 348, 377, 398, 425,
182, 215, 233, 236-37, 237, 271, 334, 431, 432
422, 425 var. commutata 24, 51. 62, 79, 126,
villosus 79 93, 167, 203 131, 258-59./259, 335, 337, 338, 344,
virginicus 24, 29. 42, 62. 77, 79, 108, 348, 362. 43f2
152, 154, 160, 167, 203, 215, 233, 245, subulata | 198, 258-59, 260, 286
248-49, 271, 288, 363, 377, 417, 423, versuta 286
425
Eragrostis 71, 107 Gastridium| ventricosum 398
capillaris 71, 107 Glyceria borealis 62, 97, 425
cilianensis 29, 363, 365, 374, 377, 414 canadensis 63, 77, 235,-289
curvula 29, 131, 280, 290, 334, 377 elata 221, 274, 295, 410, 425-
hirsuta 71, 107 erecta 410
lugens 107. 415 fluitans 64, 425
palmeri 107 grandis 63, 140, 221, 276, 407, 425
pectinacea 363, 374 leptostacKya 425
pilosa 334, 377 oocidpntalis 192, 195. 221
refracta 71 pauciflora 63, 221, 330, 425 .
rachitricha 325 septenfrionalis 63, 77, 425
secundiflora 330. 425 spectablis 7
spectabilis 330, 374 striata 63, 77, 79, 140, 221, 245, 276,
trichodes 131, 288, 334 286, 289, 425
variabilis 107 Gymnopogon ambiguus /71
Eremoeliloa ophiuroides 286
Erianthus alopecuroides 62, 107 Heteropogon-ôntortus 323
melanocarpus 399
brevibarb.is 107 Hierochloa odorata 63, 131, 154, 160
contortus 107 286, 398
giganteus 107, 118 Hilaria belangeri 399 '
hoslii, 398 ceneliroides 59
Eriochlo'a polystachya 58, 398 jamesii 335
mutica 59, 290, 335 harfordii 207
Holcus laaatus 63, 93, 97, 131, 156, 168, nitens 287
256-57, 267, 238, 286, 321, 344, 390, scabrosa 30, 221, 243, 290, 347
410 . smithii 426, 432
Hordeum 108 spectabilis 426, 432
brevisubulafum 24, 30, 79, 131, 199, subulata 176, 426, 432, 433
207, 249, 290, 290, 335, 344, 377, 414 virgata 154, 290, 335
bulbosum 24, 34, 286, 335, 344, 377 Milium effusum 80, 287, 426
disticho^ 24, 30, 79, 93, 199, 249, Molinia 65
258-57, 290, 335, 338, 340, 377, 388, Muhlenbergia 87, 90, 116, 310
423 arizonica 190
gussoneanum 97, 131 asperifolia 116, 122, 244, 295, 398
jubatum 13, 24, 30, 63, 79, 108, 131, californica 116
140, 160, 174, 245, 249, 290, 335, capillaris 116, 287
338, 344, 348, 377, 391, 423, 425, cuspidata 116, 209
429 emersleyi 106
var. caespitosum 63, 425 filiformis 192, 195, 288, 289, 426
murinum 30, 131, 156, 157, 160, 223, foliosa 116, 160, 377, 417
344, 377, 388, 423 glauca 116
nodosum 63, 79, 131, 142, 160, 168, japonica 290, 366
245, 249, 250, 290, 340, 344, 426 longiligula 106
var. boreale 63, 426 mexicana 116, 183, 217, 244, 287, 310,
pusillum 79, 93, 131, 286, 290, 335, 360, 363, 426
425 montana 105, 116
vulgare 9, 10, 15, 24, 30, 33, 34, 35, pauciflora 116
42, 50, 63, 70, 79, 93, 131, 136, 137, porteri 116
144, 156, 157, 160, 168, 174, 199, 207, pungens 335
223, 245, 250, 256-57, 286, 293, 319, racemosa 30 116, 131, 167, 209, 287,
335, 337, 338, 339, 340, 342, 344, 348, 290, 310, 335, 348, 377, 415, 426
364, 377, 388, 395, 400, 412, 414, 423, reverchonii 115, 116
"• 426 rigens 106
Hystrix patula 63, 77, 80, 108, 160, 167, schreberi 116, 183, 287, 310, 363
235, 245, 250, 250, 306, 344, 377, 415, "silvatica 116, 310, 426
417, 426 sobolifera 116
spiciformis 116
Koeleria cristata 24, 63, 77, 80, 131, squarrosa 116, 131, 199, 335
174, 176, 199, 207, 208, 230, 253, 263, tenuifolia 116
256-57, 290, 319, 330, 335, 345, 353, uniflora 116
377, 400, 414, ^26 Munroa squarrosa 244
Lasiacis divarieata 109 Oplismenus burmannii 113
Leersia orj'zoides 77, 168, 330, 415, 426 hirtellus 113, 230
virginica 363, 367, 415 ~ humboldtianus 113
Leptochloa dubia 118 setarius 113
filiformis 118 Oryza sativa 30, 36, 85, 95, 131, 136,
virgata 110 137, 159, 193, 195, 246, 310, 331, 342,
Leptoloma cognatum 113r 286, 309 352, 370, 384. 416, 436
Lolium marschallii 409, 410 Oryzopsis asperifolia 111, 157, 163, 167,
multiflorum 13, 63, 131, 155, 239, 268, 176
286, 335, 338, 345, 383, 400, 407, 422, exigua 87
423, 426 hymenoides 24, 30, 71, 131, 199. 207,
perenne 63, 64, 97, 131, 154, 154, 155, 221, 279, 290, 335, 342, 345, 347, 377,
171, 286, 292, 338, 340, 345, 377, 383, 426
400, 407, 422, 423, 426 micrantha 287, 342, 347
remotum 24, 335 miliacea 24
rigidum 63 racemosa 271
temulentum 97
Panicum 46, 59, 72
Manisuris 278 aciculare 72
cylindrica 330 agrostoides 72, 106
Melica 64 anceps 106
bulbosa 77, 160, 168, 245, 426, 432 boreale 89, 113, 259
californica 80, 245 boscii 113, 311
geyeri 426 var. molle 287
536 I N D E X O F C E R E A L A N D GRASS H O S T S
Panicum fasciculatum 106

capillare 13, 24, 30, 131, 160, 199, 220, floridanuto 60,' 323'
287, 293, 335, 348, 377, 398 var. glabratuin 279
capillarioides 274 giganteum 60 |
clandestinum 72, 113, 363 intermedium 60
commutatum 72, 113 laeve 60, 66, 91, 106, 279
consanguineum 113 langei 60 i
depauperatum 113 lividum 60
dichotomiflorum 158, 197, 309, 335, longipilum 60
363 malacophyllum 60
dichotomum 113, 309 monostachyum 60
euchlamydeum 113 mublenbergii 116
flavovirens 113 notatum 60, 395, 398
flexile 398 orb'culare 60
geminatum 104 paniculatum 398
glutinosum 106 plieatulum 13, 60, 112, 415 *
helleri 113 pubescens 58. 60, 116
hemitomum 72, 287 pubifloium 60
hians 105, 177 var. glabrum 116
huachucae 113, 192, 194 quadrifarium 60
implicatum 113, 259, 287, 377 saccharpides 105
joori 113 setaceum 116
lancearium 113 strammeum 116. 287, 312, 415
lanuginosum 113 supinum 60. 116
latifolium 113, 309, 311 tenellum 112
leibergii 309 urvillei 60. 398
lindheimeri 113 virgatum 106
linearifolium 113 Pennisetum alopenuroides 363
longifolium 105, 106 distachyum 109
maximum 198, 309, 398 glaucum 374, 374, 375
meridionale 72 orientale 398
microcarpon 113 purpureum 375
miliaceura 14, 18, 24. 30, 42, 48, 131, Phalaris arundinacea 24, 30, 63, 80,
199, 290, 335, 342, 347, 348. 366, 369, 112, 131, 140, 154. 168. 186, 203,
377, 398, 399, 412, 414, 415 225. 226-27. 274. 275, 290, 335, 377,
obtusum 116 398. 400, 407, 423
pacificum 113, 309 var. picta 363
pedicellatum 113 bulbosa^ .118 ?^3, 415
perlongum 3C9, 335, 342, 377 californica 274
praecocius ' 259. 309 oanariensis 63, 89
purpijrascons 21. 30, 398 tuberosa, 131, 336, 377
scabriusculum 113 var. stenoptera 131. 287
scoparium 72, 113 Phleum a'pinum 199, 290, 335, 401, 426
scribnerianum 72. 113, 194, 259, 309 arenarium 262-63 ^----....^
sphaerocarpon 113 phleoides 156. 290. 335, 4ffl~
subvillosum 30. 335 pratense 24. 30. 63, 77. 80. .S6. 93, 97,
tennesseense 113, 131, 287, 290 108, 131, 140, 157, 159. 174, 199, 207,
texanum 415 208, 28'2, 287, 290, 293, 335, 345, 348,
363, 37r, 395, 398, 399, 401, 403, 414,
virgatum 24. 30, 72. 108. 131. 157, 426
158, 199, 203, 206, 287, 290, 303, 309, Phragmites 65/
323, 335, 377
wilooxiamim 203 communis 24, 30. 42, 63, 203, 282,
wrightianum 113 335, 405, 426, 429
xalapense 113 309 Pleuropogon refractus 1.54
Pappophorum mucromi'ntum 108 Poa a'pina 80. 87. 203 293, 335. 426
Paspalum 60, 112, 116, 279 ampla 24, 42. 63. 65. 80; 131, 175, 203,
almum 398 242, 247. 287,-335, 345. 399, 410. 426
bosciamim 368, 374 anmia ]5,-5L 63. 65. 123. 126, 131,
caespitosum 60 241, 242, 335. 338, 340 345
condidum 112 araphnifca 80, 247, 277
ciliatifolium 55. 60. 108, 116 arntica 87. 345
coniugatum 72. 112 arida 63, 80. 211, 242, 335', 394, 426
dilp.'^'um 60, 61. 198, 279, 287, 323 bulbosa 24. 131, 160, 175, 290,, 335,
dislieWum 60, 106 345, 377, 394
canbyi 42, 63, 65, 80, 93, 175, 176, Schedonnardus paniculatus 24, 30, 131,
242, 247, 287, 290, 335, 426 199, 291, 335, 366, 398
cenisia 203, 207 Schizaehne purpurascens 108, 160, 176,
compressa 30, 45, 63, 77, 80. 86j 87, 198, 221, 287
93, 131, 176, 199, 203, 243, 247, 247, Sclerochloa dura 160
258-57, 287, 290, 335, 363, 377, 394, Scolochloa festucacea 30, 46, 221, 415
426 Secale 21
ousickii 80, 154. 203, 242, 335, 345 cereale 9 10, 15, 24, 30, 42, 62, 80,
epilis 80, 87, 203, 426 83. 93, 131, 144. 160. 168. 175, 183,
fendleriana 63, 87 203, 204, 245, 254, 255, 263, 287, 291,
glauca 203 319, 326, 335, 337, 338, 340, 345, 378,
glaucifolia 63, 80, 290, 335 391, 423, 426
gracillima 80, 160, 176, 203, 242, 400, cereale X S. montanum 24, 63, 291,
426 345, 426
interior 80, 175, 290, 335, 426 montanum 423, 426
juncifolia 30, 51. 63, 80, 87, 203, 247, Setaria genieulata 93. 314, 399
338, 377, 414, 426 italica 24, 30. 48. 199. 291, 335, 366,
kelloggii 241, 242 378. 382, 399. 414, 415
leibergii 87 lutescens 24, 30. 48. 131. 160, 199,
lepliocoma 80, 345 291, 313, 314. 330. 335, 340, 342, 347,
longifolia 63, 80. 410 348, 378. 382, 415. 434, 434
longiligula 87, 426 macrostaohya 63, 68, 399
nemoralis 63. 80 magna 48, 49
nervosa 63, 80, 207, 242, 243, 253-57, verticillata 197
426 viridis 6. 24. 30. 48. 63. 66. 131, 184,
nevadensis 24, 30, 63, 80, 203, 242, 192, 197. 199, 291, 335. 347. 366, 369,
290, 335, 399. 400 378, 382. 395, 414, 415, 434
palustris 24, 30. 63, 77, 80, 131, 176, Sitanion hansenii 80. 211. 426
203, 245, 247, 287, 293, 345, 348, 377, hystrix 24, 63, 80, 87. 131. 160, 168,
418, 426 202, 207, 211,'250, 250, 256-57, 291,
pratensis 24, 30, 51, 63. 65, 77, 89, 319. 335, 345. 426
87, 93, 122, 123. 126. 131, 140. 144, jubatum 80, 160, 168. 345, 426
146, 160, 165, 175, 176, 181, 199, 205, Sorghastrum nutans 30, 63, 110, 154,
207, 208, 242, 243, 247, 247, 279, 282,
287, 290, 292, 323, 323, 335, 338, 340, 154, 175, 218, 280, 287, 291, 294, 323,
345, 348, 359, 363, 377, 394, 415, 426 330. 414
pratensis X arachnifera 63 Sorghum halepense 118. 161. 197. 251,
reflexa 247 287, 291. 314, 340, 350, 385, 414, 420
rupicola 87 vulgare 24. 30 42, 117, 118, 131, 136,
soabrella 63, 80, 242. 335 161, 162, 183, 181, 185, 197. 199.
secunda 24, 30. 42, 61, 52. 63. 65, 77, 291, 314, 315, 335, 34?, 347, 3,50,
80. 87, 131, 160. 168. 175. 176. 199, 374, 37S, 393, 395, 399, 407, 412,
207, 211, 242.- 242, 247, 290, 302, 319, 414, 415. 420
335, 345, 378, 394, 426 var. sudanense 24, 30. 118. 131,
silvestris 80 160, 161, 183, 197, 199. 287, 291,
stenantha 63, 77, 426 335, 342, 347, 350, 359, 374, 378,
trivialis 99, 131, 426 385. 393. 399, 414. 420
vaseyochloa 13, 80, 242 Spartina alternifolia 63, 115
Polypogon monspeliensis 80 cynosuroides 63
Puccinellia distans 63, 80 gracilis 63. 255, 426
nuttalliana 45, 207, 209, 224 leiantha 114. 297
patens 63, 106
pectinata 63. 175. 255, 280. 335
Redfieldia flexuosa 30, 131, 398 ' Sphenopholis intermedia 160
Rotboellia rugosa 114 nitida 77
obtusata 30. 77, 80, 209, 216, 217,
Sacoharum barberi 30. 118 253. 287. 306
officinarum 20, 21, 24, 27. 34. 35, 39, pallens 77. 140
40. 74, 75, 83, 91, 118, 128, 131, 136, Sporobolus 116
166, 197, 199, 293, 309, 316, 342, 350, airoides 116, 176
375, 387, 406, 411, 412 argutus 116
robustum 118 asper 24, 116, 122, 176, 207, 208, 287,
sinense 118 313, 335
spontaneum 30, 118 clandestinus 207, 287, 313
Sporobolus Tripsacum dactyloides 63, 67, 287
cryptandrus 30, 34, 35, 116, 132, 207, Trisetum canesoens 63, 132, 160, 228-
291, 313, 335, 373 29, 229, 287, 426
elongatus 373 cemuum 228-29, 229, 287, 335, 403
giganteus 80, 207 flavescehs 287, 1126
heterolepis 207, 216, 217, 219, 287, montanum 4261
330 spicatum 87, 176, 209, 410, 432
indicus 373 subspioatum 209, 210
neglectus 132, 291, 399 wolfii 209
poiretii 72, 373 Triticum aestivum 9, 10, 15, 24, 30,
Stenotaphrum secundatum 132, 137, 33, 34, 36, 38, 40, 41, 42, 44, 46, 50,
399, 407, 415 63, 64, 70, 80, 83, 93, 108, 132, 136,
Stipa 30, 36 137, 142, 144, 160, 168, 175, 199, 204,
baicalensis 30, 291, 378 223, 245, 263, 264-65, 287, 291, 302,
californica 80 304, 318-19, 319, 335, 337, 338, 339,
Columbiana 63, 203, 280, 291, 335, 340, 342, 345, 348, 359, 366, 378, 389,
345, 426 391, 393, 399, 400, 407, 412, 414, 436
var. nelsonii 63, 142, 144, 157, 254, dicoccum 24, 30, 63, 132, 175, 199,
257, 262-63, 280, 335, 345, 426 203, 223, 245, 287, 291, 319, 335,
comata 24, 30, 34, 63, 132, 142, 154, 338, 340, 378, 391, 414
154, 199, 203, 217, 219, 291, 335, 345, var. farum 263, 264-65
378, 426 durum, 9, 13, 24, 30, 40, 42, 63, 132,
coronata 211, 426 175,-223, 291, 319, 335, 338, 340, 348,
elmeri 426 378, 391, 412, 414
imbricans 207 monococoum 131, 175, 319, 335, 338,
lemmonii 175, 203, 207. 280, 345, 426 ^78
lettermanii 63, 280, 426, 432 polonicum 378
leucotricha 56, 57 spelta 132, 175, 245, 245, 263, 291,
occidentalis 160, 280, 426 319, 335, 338, 340, 378, 412
pulchra 56, 378 timopheevii 30, 81, 132, 199, 291,
richardsonii 203, 211 335, 378, 412
robusta 63, 156, 270, 399 turgidum 24, 64, 263, 378
sibirica 24, 30, 102, 335, 378
spartea 24, 30, 63, 132, 203, 291, 378 Uniola latifolia 30, 193, 399
speciosa 426 laxa 108
thurberiana 426 panioulata 193/
viridula 6, 13, 24, 30, 56, 63, 89, 132,
154, 160, 175, 199, 203, 219, 245, 254, Zea mays |8, 9, 10, 12, 14, 22, 24, 30,
255, 270, 280, 291, 335, 378, 426 42, 47, 48, 61, 110, 117, 120, 121, 133,
williamsii 30, 199, 254, 255, 280, 399, 136, 138, 158, 164, 165, 170, 171, 179,
426 183, 198, 199, 287, 291, 303, 314, 316,
Stiporyzopsis bloomeri 280, 345 327, 335, 338, 340, 347, 348, 350, 358,
374, 378, 393, 397,-399, 406, 407, 412,
Trichachne patens 109 ' 413,414,436 ~-—^_
Tricholaena rosae 197, 323 Zizania aquatica 63, 90, 399
Trioda albescens 107, 287 var. angustifolia 63 —
elongata 287 Zizaniopsis miliacea 85
flava 107, 279, 287 Zoysia japcmica 67

Enfermedades de Cereales y Pastos - Disease of Cereals and Grasses - Roderick Sprague PH.D 1950

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Enfermedades de Cereales y Pastos - Disease of Cereals and Grasses - Roderick Sprague PH.D 1950

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DISEASES OF

CEREALS AND GRASSES

THE RONALD PRESS COMPANY ~9- NEW YORK

T H E RONALD PRESS 'COMPANY

All Rights Reserved

The text of this publication or any part

This book has been written to supply a source of centrally assem-

In the descriptions of the species given herein the original, often

A key to the genera of fungi discussed and represented is as follows:

Aphanomyces camptostylus Drechsl.—Root Necrosis

Mycelium hyaline, 2.5-9.5 /A diam., sparingly or moderately

Cladochytrium graminis Buesgen—"Damp-

Hyphochytrium catenoides^ Karling—On Corn Roots

Lagena radicicola Vanterpool and Ledingham—Rootlet Parasite

FIG. 2.—Lagena radicicola: A, developing sporangium showing collar attached

Olpidium brassicae (Wor.) Dang.—Rootlet Necrosis

Zoosporangia solitary or aggregated in the host cell, thin-walled,

On: Agropyron trachycaulum (Lk.) Malta, Sask.—Avena sativa L., Sask.—

FiG. 3.—Olpidium brassicae: A, sporangia; B, resting spore (semi-diagram-

come infected. Guyot (1927,a) could readily infect oats in France

Physoderma inaydis Miyabe—^Brown Bundle Disease

Phytophthora cactorum (Leb. and Cohn) Schroet.—Rot

Range: United States, Hawaii, probably widely, but perhaps irregularly,

Phy'tophthora parasitica Dastur—CTrmvn Jlot

Polymyxa graminis Ledingham—Root Hair Necrosis

EiG. 4.—Polymyxa graminia: A, zoospores on root hairs; B, young sporangium

Pythium Root Rot and Seed Rot Diseases

KEYS TO SPECIES OP PYTHIUM ISOLATED EROia GRAMINEAE I N NORTH AMERICA

A. Mycelial growth on cornmeal agar typically cottony, although sometimes

Pythium artotrogus (Mont.) deBy. var. macracanthum Sideris—Rot

Pythium butleri Subrm.—Stalk Rot of jCorn, Turf Spot

Pythium debaryanum Hesse—Seed Rot, Damping-Off and

Pythium dissotocum Drechsl.—Weak Root Rot of Sugar Cane

Pythium graminiccla Subrm.—Root Browning of Cereals,

Root browning is characterized by relatively, firm, dark, brown

Pythium hypogynum Middleton—Root Necrosis

Range: Known only from the central United States.

Mycelium-well developed on n u t r i e n t agar. H y p h a e 2-6 /A diam., side

On: Agropyron cristatum (L.) Gaertn., N. Dak.—A. intermedium (Host.)

R a n g e : Northern United States, Europe, probably widely distributed on

H y p h a e 1,5-4 /A (average 2.5 JA) in breadth, at the tip of which are

O n : Oryza sativa L., Ark.

Pythium ostracodes Drechsl.—On Wheat Roots

Pythium periilum Drechsl.—Root ]S[ecrosis

Pythium rostratum Butl.—Root Necrosis

Mycelium profuse on media, branched, 3.5-9.2 ix (average 6.4 ju)

On: Saccharum officinarum L., Hawaii—Triticum aestivum L., Hawaii—T.

Pythium tardicrescens Vanterpool—Root Rot, Root Browning

On: Avena fatua L., Wash.—Echinochloa crusgalli (L.) Beauv., Minn.—

Pythium ultimum Trow—Root Necrosis

Pythium volutum Vanterpool and Truscott—^Root Browning

Mycelium nonseptate, lustrous, with an aerial tendency in culture,

Pythium spp.—Mostly Saprophytes '

Rhizophidium graminis Ledingham—Rootlet Necrosis

Rhizopus spp.—Scutellum Rot

Sclerospora farlowii Griffiths—^Downy Mildew

Sclerospora graminicola (Sacc.) Schroet.—Downy Mildew

Sclerospora macrospora Sacc—Wheat and Oats Downy Mildew

Synchytrium graminicola sp. nov.—Purple Leaf Speckle

A key to the Ascomycetes to be discussed is as follows:

Balansia claviceps Speg.—Inflorescence Blight

Range: Tropical S. America and northward in the United States to Vir-

Balansia clavula (Berk, and Curt.) Lloyd—Black Crust