Fungal Flora of Korea

Volume 1, Number 2 Ascomycota: Dothideomycetes: Pleosporales: Pleosporaceae Alternaria and Allied Genera

2015

National Institute of Biological Resources Ministry of Environment

Fungal Flora of Korea

Volume 1, Number 2 Ascomycota: Dothideomycetes: Pleosporales: Pleosporaceae Alternaria and Allied Genera

Seung Hun Yu Chungnam National University Fungal Flora of Korea Volume 1, Number 2 Ascomycota: Dothideomycetes: Pleosporales: Pleosporaceae Alternaria and Allied Genera

Published by the National Institute of Biological Resources Environmental Research Complex, Hwangyeong-ro 42, Seo-gu Incheon, 404-708, Republic of Korea www.nibr.go.kr

All rights reserved. No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of the National Institute of Biological Resources.

ISBN : 9788968111259-96470 Government Publications Registration Number 11-1480592-000905-01

Printed by Junghaengsa, Inc. in Korea on acid-free paper

Publisher : Kim, Sang-Bae Author : Seung Hun Yu Project Staff : Youn-Bong Ku, Ga Youn Cho, Eun-Young Lee

Published on March 1, 2015

The Flora and Fauna of Korea logo was designed to represent six major target groups of the project including vertebrates, invertebrates, insects, algae, fungi, and bacteria. The book cover and the logo were designed by Jee-Yeon Koo. Preface

The biological resources represent all the composition of organisms and genetic resources which possess the practical and potential values essential for human lives, and occupies a firm position in producing highly value-added products such as new breeds, new materials and new drugs as a means of boosting the national competitiveness. As Nagoya Protocol adopted in 2010 entered into force in the 12th Conference of Parties of the Convention on Biological Diversity (CBD) in 2014, the national and international environment in the approach and common ownership of profit on genetic resources has been rapidly changed, and the competition among the nations surrounding the genetic resources is expected to be highly intense. As a part of the program, each nation in the world is putting into order to secure the information about native organisms inhabiting in its own land. The National Institute of Biological Resources of the Ministry of Environment has been publishing the ‘Flora and Fauna of Korea’ from 2006 to generally manage biological resources and to enhance national competitiveness by setting the foundation for the sovereignty over biological resources. Professional research group consisting of professors of taxonomy and related experts has syste- matically examined a total of 10,752 species for the past 7 years to publish 120 volumes each in Korean and English, and 2 volumes of World Monograph covering 216 species. In addition, 30 volumes of the ‘Flora and Fauna of Korea’, both in Korean and in English, covering 1,665 species of invertebrates, insects and algae are additionally published this year. The publication of the ‘Flora and Fauna of Korea’ serve to identify indigenous species living in Korea, to investigate biota, to improve the quality of national biological resources management and to provide the opportunity to lay the groundwork for the biotechnological industrialization of biological resources. And I hope the project initiated by The National Institute of Biological Resources of the Ministry of Environment will help to discover useful biological resources in Korea and to create high value-added activities including natural products, genetic resources and medical sub- stance development. The National Institute of Biological Resources of the Ministry of Environment will continue to accelerate the project of the publication of the ‘Flora and Fauna of Korea’ to identify the entity of Korean indigenous species in the future. Personally I would like to express my sincere appreciation for those experts who spared no effort to publish the biological monographs; Emeritus Professor Seung Hun Yu of Chungnam National University.

Kim, Sang-Bae President National Institute of Biological Resources

Contents

List of Taxa 3 Introduction 5 Historical Review of Taxonomy 6 Materials and Methods 10 Taxonomy 11 Taxonomic Notes 13 1. Alternaria acalyphae (Nelen) E.G. Simmons 16 2. Alternaria alternata (Fr.) Keissler 19 3. Alternaria araliae H.C. Greene 22 4. Alternaria atra (Preuss) Woundenberg & Crous 24 5. Alternaria brassicae (Berkeley) Saccardo 26 6. Alternaria brassicicola (Schwein.) Wiltshire 29 7. Alternaria calendulae Ondrej˘ 32 8. Alternaria cassiae Jurair & A. Khan 34 9. Alternaria cinerariae Hori & Enjoji 36 10. Alternaria citri Ellis & N. Pierce 38 11. Alternaria cosmosa Y.L. Li & W.S. Wu 40 12. Alternaria cucumerina (Ellis & Everh.) J.A. Elliot 42 13. Alternaria dauci (J.G. Kühn) J.W. Groves & Skolko 45 14. Alternaria dianthicola Neergaard 47 15. Alternaria embellisia Woundenberg & Crous 49 16. Alternaria gaisen Nagano ex Hara 51 17. Alternaria gossypina (Thüm.) J.C.F. Hopkins 53 18. Alternaria helianthinficiens E.G. Simmons, Walcz & R.G. Roberts 55 19. Alternaria iridicola (Ellis & Everh.) J.A. Elliot 57 20. Alternaria japonica Yoshii 59 21. Alternaria longipes (Ellis & Everh.) E.W. Mason 62 22. Alternaria macrospora Zimmermann 64 23. Alternaria mali Roberts 66 24. Alternaria nobilis (Vize) E.G. Simmons 68 25. Alternaria panax Whetzel 70 26. Alternaria peucedani S.H. Yu 73 27. Alternaria porri (Ellis) Ciferri 75 28. Alternaria protenta E.G. Simmons 77 29. Alternaria radicina Meier, Drechsler & E.D. Eddy 79 30. Alternaria ricini (Yoshii) Hansford 81 31. Alternaria scirpicola (Fuckel) Sivanesan 83 32. Alternaria sesami (E. Kawam.) Mohanthy & Behera 85 33. Alternaria simsimi E.G. Simmons 87 2 Fungal Flora of Korea·Alternaria and Allied Genera

34. Alternaria solani Sorauer 89 35. Alternaria tagetica S.K. Shome & Mustafee 92 36. Alternaria tenuissima (Nees) Wiltshire 94 37. Alternaria tomatophila E.G. Simmons 96 38. Alternaria vanuatuensis E.G. Simmons & C.F. Hill 98 39. Alternaria zinniae H. Pape ex M.B. Ellis 100 40. Alternariaster helianthi (Hansf.) E.G. Simmons 103 41. Prathoda longissima (Deighton & MacGarvie) E.G. Simmons 105 42. Stemphylium astragali (Yoshii) W. Yamamoto 108 43. Stemphylium botryosum Wallroth 110 44. Stemphylium globuliferum (Vestergr.) E.G. Simmons 112 45. Stemphylium lycopersici (Enjoji) W. Yamamoto 114 46. Stemphylium platycodontis J.X. Deng & S.H. Yu 115 47. Stemphylium sarciniforme (Cavara) Wiltshire 117 48. Stemphylium solani G.F. Weber 119 49. Stemphylium vesicarium (Wallr.) E.G. Simmons 121 50. Trichoconiella padwickii (Ganguly) B.L. Jain 124 Literature Cited 126 Plates 134 Index of Host Plants/Substrates 144 Index of Fungal Names 146 3

List of Taxa

Phylum Ascomycota Class Dothideomycetes Order Pleosporales Family Pleosporaceae Genus Alternaria Nees, 1816 Alternaria acalyphae (Nelen) E.G. Simmons, 1994 Alternaria alternata (Fr.) Keissler, 1912 Alternaria araliae H.C. Greene, 1953 Alternaria atra (Preuss) Woundenberg & Crous, 2013 Alternaria brassicae (Berkeley) Saccardo, 1880 Alternaria brassicicola (Schwein.) Wiltshire, 1947 Alternaria calendulae Ondrej,˘ 1974 Alternaria cassiae Jurair & A. Khan, 1960 Alternaria cinerariae Hori & Enjoji, 1931 Alternaria citri Ellis & N. Pierce, 1902 Alternaria cosmosa Y.L. Li & W.S. Wu, 2005 Alternaria cucumerina (Ellis & Everh.) J.A. Elliot, 1917 Alternaria dauci (J.G. Kühn) J.W. Groves & Skolko, 1944 Alternaria dianthicola Neergaard, 1945 Alternaria embellisia Woundenberg & Crous, 2013 Alternaria gaisen Nagano ex Hara, 1920 Alternaria gossypina (Thüm.) J.C.F. Hopkins, 1931 Alternaria helianthinficiens E.G. Simmons, Walcz & R.G. Roberts, 1986 Alternaria iridicola (Ellis & Everh.) J.A. Elliot, 1917 Alternaria japonica Yoshii, 1941 Alternaria longipes (Ellis & Everh.) E.W. Mason, 1928 Alternaria macrospora Zimmermann, 1904 Alternaria mali Roberts, 1914 Alternaria nobilis (Vize) E.G. Simmons, 2002 Alternaria panax Whetzel, 1912 Alternaria peucedani S.H. Yu, 2014 Alternaria porri (Ellis) Ciferri, 1930 Alternaria protenta E.G. Simmons, 1986 Alternaria radicina Meier, Drechsler & E.D. Eddy, 1922 Alternaria ricini (Yoshii) Hansford, 1943 Alternaria scirpicola (Fuckel) Sivanesan, 1984 Alternaria sesami (E. Kawam.) Mohanthy & Behera, 1958 Alternaria simsimi E.G. Simmons, 2004 Alternaria solani Sorauer, 1896 Alternaria tagetica S.K. Shome & Mustafee, 1966 Alternaria tenuissima (Nees) Wiltshire, 1933 Alternaria tomatophila E.G. Simmons, 2000 4 Fungal Flora of Korea·Alternaria and Allied Genera

Alternaria vanuatuensis E.G. Simmons & C.F. Hill, 2007 Alternaria zinniae H. Pape ex M.B. Ellis, 1972 Genus Alternariaster E.G. Simmons, 2007 Alternariaster helianthi (Hansf.) E.G. Simmons, 2007 Genus Prathoda Subramanian, 2007 Prathoda longissima (Deighton & MacGarvie) E.G. Simmons, 2007 Genus Stemphylium Wallroth, 1833 Stemphylium astragali (Yoshii) W. Yamamoto, 1960 Stemphylium botryosum Wallroth, 1833 Stemphylium globuliferum (Vestergr.) E.G. Simmons, 1969 Stemphylium lycopersici (Enjoji) W. Yamamoto, 1960 Stemphylium platycodontis J.X. Deng & S.H. Yu, 2014 Stemphylium sarciniforme (Cavara) Wiltshire, 1938 Stemphylium solani G.F. Weber, 1930 Stemphylium vesicarium (Wallr.) E.G. Simmons, 1969 Genus Trichoconiella B.L. Jain, 1976 Trichoconiella padwickii (Ganguly) B.L. Jain, 1976 5

Introduction

Alternaria is a dictyosporous genus of the family Pleosporaceae, order Pleosporales, class Dothi- diomycetes, phylum Ascomycota. Several species of Alternaria are saprophytes commonly found in soil and/or dead or dying plant tissue. The saprophytic species are involved in degradation of plant debris and other organic substrata. The majority, however, are plant pathogens that, collec- tively, cause a range of economically important diseases and post-harvest rots on a variety of crops worldwide (Rotem, 1994; Thomma, 2003). As components of both indoor and outdoor air spora they are also common human allergens causing hay fever or hypersensitivity reactions that sometimes lead to asthma (Peat et al., 1993; Halonen et al., 1997). Several taxa are also important causa- tive agents of phaeohyphomycosis in immune-compromised patients (Lawrence et al., 2013; Wou- denberg et al. 2013). Alternaria and its close relatives comprise a group of fungi whose importance has long been appreciated, in mycology as well as in plant pathology. The taxonomy of Alternaria has been a subject of controversy because of high variability in conidial morphology and their polymorphism displayed even in pure cultures. Incomplete descriptions, mutations and secondary development or dwarfing of spores have caused great confusion in the classification of this genus (Yu, 1992). Alternariaster, Embellisia, Nimbya, Prathoda, Stemphylium, Ulocladium, and Trichoconiella, which also belong to the Pleosporaceae, are similar to Alternaria in producing dictyosporous conidia and poro- conidia. Certain species in these genera resemble Alternaria species and are occasionally misidentified. Some morphological characteristics such as conidial shape and size used for identification of the species of these genera have been shown to overlap among species and vary depending on the cultural conditions such as temperature and substrate (Leach and Aragaki, 1970; Hosen et al., 2009). Because of the significance of Alternaria spp. and related taxa on humans and their sur- roundings, a correct and rapid identification of them would be of great value to mycologists, plant pathologists and the public alike. In this volume, detailed diagnostic descriptions and illustrations of thirty nine species of Alter- naria, eight species of Stemphylium, and one species each of Alternariaster, Prathoda and Trichoconi- ella, collected by the author from Korea are presented. Under each fungal species, its synonyms, morphological characters, hosts and records in Korea, materials examined, geographic distribution, taxonomic remarks, and line drawings and photographs of conidia and conidiophores are provided. Additionally, surface features of conidia of twenty species of Alternaria, one species of Alternariaster and four species of Stemphylium observed under the scanning electron microscope (SEM) are described. The species of Alternaria and Stemphylium are presented in alphabetical order.

Acknowledgements The author thanks Professor Jeffrey Stone (Oregon State University) for reviewing and editing the English of the manuscript and Dr. H.S. Cho for inking the line drawings of the fungal structures. Thanks are due to Drs. H.B. Lee, M.S. Park, H.S. Cho, J.X. Deng and N.C. Paul, and former graduate students in the Laboratiry of Plant Pathology, Chungnam National University for their interested and painstaking technical assistance and for collecting some specimens. 6

Historical Review of Taxonomy

The genus Alternaria was first described by Nees in 1816, with A. tenuis as the sole species. The description is somewhat incomplete and inaccurate, however, it is sufficiently definite to leave little doubt that Nees’ description is now generally recognized as Alternaria (Elliot, 1917; Neergaard, 1945; Simmons, 1967). Fries in 1825 described the genus Macrosporium, and in 1832 gave a more detailed description, differentiating it from Cladosporium, Helminthosporium and Sporodesmium. He did not recognize the genus Alternaria but described Nees’ fungus under the name Torula alternata, citing A. tenuis Nees as a synonym. Chevallier (1836), however, recognized Alternaria and stated that it differed from Torula in having filiform connections between spores. Corda (1839, 1840) also recognized Alternaria and gave descriptions and illustrations of A. tenuis deviating from the one given by Nees, showing spore chains consisting of up to 13 spores. The connection was not filiform, but it tapered gradually from the conidial body. In 1912, Keissler reevaluated both Nees’ and Fries’ descriptions and synonymized both specimens with Alternaria alternata. In 1917 Elliot published an extensive report concerning the establishment of generic characteristics and limitations of Alternaria and Macrosporium. He emphasized the form of conidia (obclavate, pointed, and often having beaks) as a generic criterion and stated that chain formations could be suppressed under unfavorable conditions. Elliot (1917) considered Alternaria and Macrosporium to be distinct genera and suggested that the genus Alternaria could be divided into six species groups based on conidium morphology such as size, shape, septation and beak. However, Angell (1929) considered Alternaria and Macrosporium synonymous and used the name Macrosporium to designate both genera. Wiltshire (1933) agreed that Alternaria and Macrosporium are congeneric, and stated that Macro- sporium in the original description was used to designate fungi of the Alternaria type having non- filiform spores and one spore per conidiophores. He considered the name Alternaria to be invalid according to the International Rules of Botanical Nomenclature, since Fries did not recognize the genus in his Systema, but he concluded that the descrioption given by Nees was sufficient to estab- lish the current concept of the genus. He suggested that since Alternaria was a universally known term, it shoud be used to designate both genera. Neergaard (1945) completely revised the taxonomy of Alternaria, eliminating some species and uniting others in one species with several formae speciales. The treatise of Neergaard is limited to Alternaria species (16 species, 2 varieties, and a few formae speciales) in Denmark. Neergaard separated the species into three groups (which he called sections) according to catenulation (the number of spores in the chain). He distinguished 1) Longicatenatae, in which conidia appear in long chains of about 10 or more spores; 2) Brevicatenatae, in which conidia are formed in short chains of about 3-5 spores; and 3) Noncatenatae, in which conidia are usually formed singly, seldom producing secondary spores. Species within each section were keyed by conidium morphology such as shape, septation, size, color, wall ornamentation, and beak. Neergaard also paid particular attention to the difference between the true beak (eurostrum), as an integral part of the conidium, and the false beak (pseudorostrum), the structure that is in fact a secondary conidiophore essential to generation of conidial chains in Alternaria. Joly (1964) concluded, from a thorough study of 27 taxa, that evolution in Alternaria progressed in three lines (which he designated as sections) from a nucleus of a few saprophytic or weakly parasitic, polyphagous species. He based his sections on spore color and spore rigidity or lateral sym- Historical Review of Taxonomy 7 metry: Claroseminae, with light colored spores; Brunneoseminae, with brown or brown-reddish colored spores; and Rigidae, in which spores appearing slightly rigid and with relatively few longitudinal septa. Later (1965), he supplied a detailed key to his sections and to the most common species of the genus known to him. The key of Joly (1965) was originally written in French but later translated into English (Joly, 1967). Unfortunately, the key does not provide descriptions of media used, and gives dimensions of spores not including the beak. Simmons (1967), after examining an authenticated specimen of the type species of Alternaria Nees in Persoon’s herbarium, redescribed the fungus and indicated that the correct name of the fungus should be A. alternata (Fries) Keissler. Ellis (1971, 1976) published precise descriptions and illustrations of 27 (1971) plus 17 (1976) taxa that he assigned to Alternaria. The taxa presented by him were arranged according to how abrupt the transition was from spore body to beak and how short or long the beak was relative to the length of the body. Simmons’ continuing studies on Alternaria (Simmons, 1981-2003) redescribed numerous species and differentiated the genus into numerous species novae. In 1992, he developed the species- group system within the genus after examining several thousand field specimens and isolates of Alternaria. Fourteen species groups were defined on the basis conidial characteristics, the pattern of chain formation, and the nature of the apical (sometimes lateral) extension of conidium cells (true beaks as an integral part of the conidium; secondary conidiophores as a germination phe- nomenon leading to secondary conidiogenesis). Simmons and Roberts (1993) further divided the small-spored, catenulate Alternaria taxa into six morphological groups. Simmons further developed several species-groups by describing the A. infectoria complex, the A. alternata species-group, the A. tenuissima species-group, the A. cheiranthi species-group and the A. brassicicola species-group (Simmons 1994, 1995). In 2007, Simmons published an identification manual of Alternaria in which he separated Alter- naria species into two sections based on size of conidial body, and 13 species groups based on conidial morphology and sporulation patterns. The first section contains three groups of large-spored Alternaria with three different beak types; blunt-tapered beak, narrow-tapered beak and filamentous beak. The second section includes 10 groups of small-spored Alternaria with 10 different sporulation patterns. He transferred A. helianthi to a new genus, Alternariaster, and A. longissima to the established genus Prathoda. Numerous new species and varieties of Alternaria also have been reported worldwide (Zhang, 1999; Zhang et al., 1999; Zhang and Zhang, 1999; Labuda et al., 2008; Roberts, 2008; Zhang et al. 2009; Gannibal, 2010; Roberts et al., 2010; Toth et al., 2011). However, as Rotem (1994) indicated, the distinction between some morphologically similar species based on analysis of minute differences described by Simmons (1981-2004, 2007) and Zhang et al. (1999, 2009) is somewhat confusing for non-specialists. A formal system of infrageneric classification was introduced by Lawrence et al. (2013) based on molecular phylogeny of five protein coding loci. They also consider the relationship between Alternaria, Stemphylium and Ulocladium.

The genus Stemphylium, with S. botryosum Wallr. as its type species, was established by Wallroth (1833). Berkeley (1838) described Macrosporium sarcinula Berk. on decaying orange gourds and later L.R. and C. Tulasne (1863) cited M. sarcinula as the conidial state of Pleospora herbarum. Miyabe (1889) proved the genetic connection between the conidial state, named Macrosporium parasiticum by von Thümen in 1877, with P. herbarum by pure cultures and expressed the opinion that M. sarcinula was the same fungus, although he did not examine type material of this species. Elliot (1917) expressed the opinion that species of Macrosporium with sarciniform or globose spores 8 Fungal Flora of Korea·Alternaria and Allied Genera should be classified in Stemphylium, and Wiltshire (1938), after examination of several type specimens involved in the nomenclature and classification of Macrosporium and Stemphylium, regarded M. sarcinula and M. parasiticum as belonging to the same species and, consequently, as synonyms of S. botryosum. Wiltshire (1938) further proposed that two groups should be included in the genus Stemphylium: Eustemphylium for species showing the original characters of the genus (typified by S. botryosum), and Pseudostemphylium for species similar to S. lanuginosum Harz, of which no type specimen is known. Neergaard (1945) also followed the lead of Wiltshire by retaining the two sections proposed for Stemphylium. Suburamanian (1961) proposed that Pseudostemphylium should be raised to generic rank and made a few nomenclatural combinations in the genus, with S. lanuginosum as the type. Joly (1964) differentiated Stemphylium in its original sense and disposed of several of the taxa similar to S. lanuginosum in Alternaria. However, according to Simmons (1967), all of the taxonomists cited above have recognized that generic placement of species similar to S. lanuginosum in either Alternaria or Stemphylium are inap- propriate. Either their nomenclatural proposals or, more importantly, their invariably excellent illustrations reveal that conidiophores of members of the S. lanuginosum group bear no resem- blance whatsoever to those of S. botryosum. The conidial morphology of the group is fundamentally different from that of the type species of Alternaria. Simmons (1967) contended that the correct relationships of puzzling species of the S. lanuginosum group are with Ulocladium, typified by U. botrytis Pr. Ellis (1971, 1976) published precise descriptions and illustrations of five (1971) plus two (1976) species of Stemphylium. The initial five taxa presented by him (1971) were keyed out according to the conidial morphology, e.g., shape, surface ornamentation, constriction of the septa, and length: width ration of conidia. Simmons (1985) later reclassified Stemphylium/Pleospora species and reported that S. botryosum and P. herbarum were not components of the same holomorph. He assigned P. tarda and P. herbarum as the teleomorph of S. botryosum and S. herbarum, respectively, and also recognized a new species, based on an isolate from alfalfa (Medicago sativa) from Western Australia.

Ulocladium was erected by Preuss in 1851, with U. botrytis Press as type species and the genus Embellisia was established in 1971, with E. allii (Campanile) Simmons as the type, to accommodate the atypical Helminthosporium allii Campanile (Simmons, 1971). Trichoconiella was established in 1975, with T. padwickii (Ganguly) B.L. Jain as type species, to accommodate the atypical Trichoconis padwickii Ganguly (Jain, 1975). The same fungus had previously been transferred to Alternaria, as A. padwickii (Ganguly) M.B. Ellis, by Ellis (1971). Simmons (1989) erected a new anamorph genus Nimbya, with N. scirpicola (Fuckel) Simmons as type, to accommodate the atypical Sporidesmium scirpicola Fuckel. The fungus was previously described as Cercospora scirpicola (Fuckel) Zind.-Bakker (van Zinderen Bakker, 1940), Alternaria state of Pleospora scirpicola (Lucas and Webster, in M.B. Ellis, 1976) and Alternaria scirpicola (Fuckel) Sivanesan (Sivanesan, 1984).

Molecular studies have revealed multiple non-monophyletic genera within the Alternaria complex, and Alternaria species clades that do not always correlate to species groups or sections based on morphological characteristics (Pryor and Gilbertson, 2000; Chou and Wu, 2002; Pryor and Bige- low, 2003; Hong et al., 2005; Inderbitzin et al., 2006; Pryor et al., 2009; Lawrence et al., 2013; Wou- denberg et al., 2013). Pryor and Gilbertson (2000) presented the first detailed molecular phylogenetic analysis of Alter- naria and allied genera and demonstrated that Alternaria and Ulocladium cluster as a monophyletic group with several internal clades referred to as the alternata, radicina, porri, brassicicola and Historical Review of Taxonomy 9 ulocladium species-groups. Interestingly, the fifth Alternaria clade referred to as the infectoria species-group was phylogenetically more distant and more closely related to other genera such as Macrospora (anamorph, Nimbya) and Allewia (anamorph, Embellisia). Since then, two additional Alternaria lineages have been presented based on molecular and morphological analyses: the sonchi species-group (Hong et al., 2005) and alternantherae species-group (Lawrence et al., 2012, 2013). Continued molecular phylogenetic analyses of five phylogenetically informative loci (gpd, Alt a1, actin, plasma membrane ATPase, calmodulin) of Alternaria and allied genera revealed eight distinct lineages (sections) (Alternaria, Aternantherae, Brassicicola, Gypsophilae, Panax, Porri, Radicina, and Sonchi) and one species-group (infectoria) that was not supported as a lineage within Alternaria (Lawrence et al., 2013). Recently, Alternaria has been redefined with its allied genera based on sequence analyses of six gene regions (18S nrDNA, 28S nrDNA, ITS, GAPDH, RPB2, and TEF1-alpha gene regions) (Woudenberg et al., 2013), which revealed a Pleospora/Stemphylium clade and a well- supported core Alternaria clade. According to these authors, the Alternaria clade is composed of 24 internal clades (sections) and six monotypic lineages, the assemblage of which they recognize as Alternaria. Woudenberg et al. (2013) synonymised 13 genera, including Embellisia, Nimbya, and Ulo- cladium, under Alternaria based on sequences of the six gene regions. The monophyly of Stemphylium was robustly defined in the ascomycete family Pleosporaceae by the phylogenetic analysis of the ITS region and gpd gene (Camara et al., 2002), and six genes such as 18S nrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha (Woudenberg et al., 2013). To determine the taxonomic status of the species in Stemphylium, both morphological and molecular phylogenetic analyses are complementary in modern fungal systematic (Wang et al., 2009). 10

Materials and Methods

The materials examined in this work were all collected by authors from different locations and habitats in Korea during 1998-2013. All specimens have been preserved in CNU (Chungnam national University) mycological herbarium. Each specimen was examined microscopically for the presence of Alternaria and allied genera at the day of collection or after incubation in a moist cham- ber (well moistened filter paper in Petri dishes) at 20-23°C in a 12 hr dark/12 hr light regime for 1 day. In some cases, seed samples were tested by the blotter method (ISTA, 1976) and examined microscopically. For comparative study of the sporulation pattern and morphological characters of conidiophores and conidia, single spore isolation of each species was made and cultured on V-8 agar or Potato Carrot Agar at 20°C in a 12hr dark/12hr light regime for 7 days. For producing macroscopic characters such as growth and color, etc., pure cultures of each species were grown on PDA at 25°C. Conidia, conidiophores and mycelium from specimens and cultures were mounted on microscope slides in plain lactophenol, covered by a cover glass and sealed with glyceel. The mycological characteristics such as conidial shape, size, color, surface ornamentation and number of septa, etc. were observed under a compound microscope at 400×magnification. Measurements of conidia and conidiophores were made from 50 replicates for each species. Conidium sporulation patterns, i.e., arrangement of conidia on conidiophores, number of conidia in chain, and pattern of chain branching were observed under a stereomicroscope at about 50×magnification. The morphology of conidia and conidiophores of each species was illustrated with the aid of a drawing tube by Dr. H.S. Cho, Ginseng Resour. Inst., Korea Ginseng Corp.. For observations of surface ornamentation of conidia with the scanning electron microscope (SEM), the conidia of 20 species of Alternaria, one species of Alternariaster and 4 species of Stem- phylium from cultures were fixed with 2% glutaraldehyde in 0.1 M phosphate buffer (pH 7.0) for 2 h, and post fixed in 2% osmic acid for 2 h followed by three successive washing steps in distilled water. They were serially dehydrated by passage through ethanol solution of increasing concen- tration (50 to 100 %), submitted to critical-point drying, and coated with gold under vacuum with EIKO IB-3 ion coater, and then examined with JSM-F7 SEM operating at 7 kV. To follow modern systematics and latest scientific names, the nomenclature for Alternaria species used herein follows that of the MycoBank Fungal Databases (http:// mycobank.org) and Wouden- berg et al. (2013). Synonyms were obtained from the MycoBank Databases, Simmons (2007) and Woudenberg et al. (2013). 11

Taxonomy

Synoptic key to Alternaria and allied genera Synoptic key number Common allied genera 1.························································································································Alternaria 2.···············································································································Alternariaster 3.······················································································································Prathoda 4.·················································································································Stemphylium 5.·················································································································Trichoconiella

Conidiophore and conidiogenous cell proliferation: Sympodial proliferation Often geniculate····················································································································1, 3 Rarely geniculate ······················································································································2 Not sympodial proliferation Simple proliferation··················································································································5 Percurrent proliferation ············································································································4

Conididiogenous scars: Conspicuously pigmented and thickened············································································1, 3, 4 Inconspicuously pigmented and unthickened·········································································2, 5

Shape of conidia: Mostly obovoid, obclavate, ellipsoidal························································································1 Mostly cylindrical or long ellipsoidal ·························································································2 Mostly elongate-cylindrical, cercospora-like···············································································3 Mostly ellipsoidal, subsphaerical or obclavate············································································4 Mostly fusiform to obclavate ······································································································5

Number of longitudinal septa: One to several ··················································································································1, 2, 3, 4 None ··········································································································································5

Conidial beak: Conidia both with beak and beakless ······················································································1, 3 Conidia only beakless (erostrate)·····························································································2, 4 Conidia always with beak (rostrate)····························································································5

Taxonomic Notes

Genus Alternaria Nees System der Pilze and Schwämme: 72, 1816.

SYNONYMS: Elosia Pers., Mycologia Europaea 1: 12, 1822. Macrosporium Fr., Systema Mycologicum 3: 373, 1832. Rhopalidium Mont., Annales des Sciences Naturelles Botanique 6: 30, 1836. Brachy- cladium Corda, Icon. Fungorum hucusque Cogn. 2: 14, 1838. Ulocladium Preuss, Linnaea 24: 111, 1851. Prathoda Subram., J. Indian Bot. Soc. 35(1): 73, 1956. Chmelia Svob.-Pol., Biológia (Bratislava) 21: 82, 1966. Embellisia E.G. Simmons, Mycologia 63: 380, 1971. Botryomyces de Hoog & C. Rubio, Sabouraudia 20: 19, 1982. Lewia M.E. Barr & E.G. Simmons, Mycotaxon 25: 289, 1986. Ybotromyces Rulamort, Bull. Soc. Bot. Centre-Ouest, Nouv. Sér. 17: 192, 1986. Nimbya E.G. Simmons, Sydowia 41: 316, 1989. Allewia E.G. Simmons, Mycotaxon 38: 260, 1990. Ceivellia Shoemaker & Inderb., Canad. J. Bot. 84: 1308, 2006. Chalastospora E.G. Simmons, CBS Biodiversity Ser. 6: 668, 2007. Tere- tispora E.G. Simmons, CBS Biodiversity Ser. 6: 674, 2007. Undifilum B.M. Pryor, Creamer, Shoemaker, McLain-Romero & Hambl., Botany 87: 190, 2009. Sinomyces Yong Wang bis & X.G. Zhang, Fungal Biol. 115: 192, 2011.

COLONIES effuse, usually gray, dark brown or black. MYCELIUM completely immersed or partly superficial; hyphae colourless, olivaceous brown or brown, septate. CONIDIOPHORES (Pri- mary conidiophores) macronematous, mononematous, solitary or in fascicles, simple or branched, pale brown or brown, septate, apically with a sympodially proliferating conidiogenous cell, often geniculate with one to several pigmented conidiogenous site. CONIDIA solitary or catenate through the agency of secondary conidiophores, ovoid, obovoid, narrowly ellipsoid or obclavate, beaked (rostrate or pseudorostrate) or non-beaked, pale to mid olivaceous brown, or brown, smooth or verrucose, multicelled with transverse and frequently also longitudinal or oblique septa. Type species: Alternaria alternata (Fr.) Keissler.

Synoptic key to Korean species of Alternaria

Synoptic key number Alternaria species 1.·······························································································································A. acalyphae 2.································································································································A. alternata 3.······································································································································A. araliae 4.··········································································································································A. atra 5.···································································································································A. brassicae 6.································································································································A. brassicicola 7.································································································································A. calendulae 8.······································································································································A. cassiae 9.·································································································································A. cinerariae 10.·········································································································································A. citri 11. ····································································································································A. cosmosa 14 Fungal Flora of Korea·Alternaria and Allied Genera

12.······························································································································A. cucumerina 13.·······································································································································A. dauci 14.·······························································································································A. dianthicola 15.·································································································································A. embellisia 16.······································································································································A. gaisen 17.·································································································································A. gossypina 18.······················································································································A. helianthinficiens 19.···································································································································A. iridicola 20.···································································································································A. japonica 21.···································································································································A. longipes 22.······························································································································A. macrospora 23.·········································································································································A. mali 24.······································································································································A. nobilis 25.······································································································································A. panax 26.··································································································································A. peucedani 27.········································································································································A. porri 28.···································································································································A. protenta 29.····································································································································A. radicina 30.·······································································································································A. ricini 31.·································································································································A. scirpicola 32.·····································································································································A. sesami 33.····································································································································A. simsimi 34.······································································································································A. solani 35.····································································································································A. tagetica 36.································································································································A. tenuissima 37.······························································································································A. tomatophila 38.··························································································································A. vanuantuensis 39.····································································································································A. zinniae

Mature conidium body length: Short [‹50 (-60) μm]: 2, 4, 10, 15, 17, 21, 23, 36 Medium (50-100 μm): 1, 3, 8, 9, 14, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 27, 29, 30, 33, 36, 38, 39 Long (›100 μm): 1, 3, 5, 7, 8, 9, 11, 12, 13, 26, 27, 28, 31, 32, 34, 35, 37, 38, 39 Catenulation of conidia: Conidia usually solitary, rarely in chains of 2 (-3): 1, 3, 4, 5, 7, 8, 9, 11, 12, 13, 14, 15, 18, 19, 20, 22, 24, 25, 26, 27, 28, 29, 30, 31, 32, 34, 35, 37, 38, 39 Conidia in short chains of 3 to 4 (-5): 5, 8, 9, 10, 14, 16, 17, 19, 20, 21, 23, 24, 25, 32, 33, 36, 39 Conidia in long chains of 5-10+: 2, 6, 10, 16, 17, 21, 23, 33, 36 Shape of conidia: Cylindrical to subcylindrical: 6, 14, 15, 19, 33 Ellipsoid: 2, 6, 9, 10, 16, 17, 18, 19, 20, 21, 24, 33, 36, 39 Broadly ellipsoid: 4, 9, 12, 22, 23, 29 Long or narrowly ellipsoid: 1, 2, 3, 7, 8, 13, 15, 16, 19, 20, 22, 23, 25, 27, 28, 30, 32, 33, 34, 35, 36, 37, 38, 39 Obclavate: 1, 2, 3, 5, 7, 8, 9, 12, 13, 14, 16, 17, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 30, 32, 33, 34, 35, Pleosporales: Pleosporaceae: Alternaria 15

36, 37, 38, 39 Broadly obclavate: 3, 9, 13, 18, 22, 25, 37 Long or narrowly obclavate: 11, 31 Obpyriform: 2, 10, 26 Ovoid: 2, 16, 17, 18, 29, 33, 39 Broadly ovoid: 6, 16, 19, 20, 24, 36 Long or narrowly ovoid: 1, 5, 6, 7, 8, 9, 12, 16, 17, 19, 21, 23, 32, 35, 36, 38, 39 Conidial beaks: Beakless (erostrate): 2, 3, 4, 5, 6, 8, 9, 10, 14, 15, 16, 17, 18, 19, 20, 21, 23, 24, 25, 26, 29, 33, 36 Beakless (but pseudorostrate): 2, 3, 4, 5, 8, 9, 10, 14, 15, 16, 17, 19, 20, 21, 23, 24, 25, 29, 33, 36 Beak taper gradual, but short [‹50 (-60) μm]: 16, 17, 21, 23, 36 Beak taper gradual, long (50-100 μm) and broad: 3, 5, 14, 19, 25, 26 Beak taper narrow to filiform: 1, 7, 8, 11, 12, 13, 18, 22, 27, 28, 30, 31, 32, 33, 34, 35, 37, 38, 39 Beak sometimes branched: 1, 7, 11, 13, 27, 28, 32, 34, 37, 38 Wall of conidia: Always smooth: 1, 5, 9, 11, 14, 19, 24, 29 Smooth or verruculose: 2, 3, 4, 6, 7, 8, 10, 12, 13, 16, 17, 18, 20, 21, 22, 23, 25, 26, 27, 28, 30, 32, 33, 34, 35, 36, 37, 38, 39 Colony characteristics (on PDA at 25°C after 7 days) Growth rate: Moderately slow [‹5(-6) cm diam.]: 1, 3, 4, 5, 6, 8, 9, 12, 13, 14, 15, 18, 19, 20, 21, 22, 24, 25, 26, 27, 29, 30, 31, 32, 33, 34, 37, 39 Rapid (›6 cm diam.): 2, 7, 9, 10, 11, 13, 15, 16, 17, 22, 23, 27, 28, 33, 34, 35, 36, 38 Pigment production into the medium: Yellow or pale yellow: 18, 22, 27, 34, 37, 38 Reddish yellow: 18, 22 Reddish brown: 13, 21, 27, 34 Purple: 21 No pigment: 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 19, 20, 21, 23, 24, 25, 26, 28, 29, 30, 31, 32, 33, 35, 36, 37, 39 Hosts/Substrates Acalypha: 1 Acanthopanax: 3 Allium: 15, 27, 38 Aloe: 2 Antirrhonum: 2 Aralia: 3 Arctium: 2 Armoracia: 5, 6 Belamcanda: 19 Brassica: 5, 6, 20 Calendula: 7 Capsicum: 34, 36 Cassia: 8 16 Fungal Flora of Korea·Alternaria and Allied Genera

Citrus: 10 Citrullus: 12 Cosmos: 11 Cucumis: 12 Cucurbita: 12 Daucus: 13, 29 Dianthus: 14, 24 Echinopanax: 3 Eucommia: 2 Fragaria: 2, 36 Gladiolus: 19 Gossypium: 17, 22 Helianthus: 18, 28 Hibiscus: 2 Kalopanax: 3 Lepidium: 5 Lycopersicon: 37 Malus: 23 Nicotiana: 21 Oryzae: 2 Panax: 25 Peucedanum: 26 Phaseolus: 36 Pyrus: 16 Raphanus: 5, 6, 20 Ricinus: 30 Rorippa: 5 Rudbeckia: 28 Scirpus: 31 Senecio: 9 Sesamum: 32, 33 Solanum: 34 Tagetes: 35 Zinnia: 39 Soil: 4

1. Alternaria acalyphae (Nelen) E.G. Simmons, Mycotaxon 50: 240, 1994 (Figs. 1, 2).

SYNONYM: Macrosporium acalyphae Nelen, Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 15: 142, 1962.

COLONIES on potato dextrose agar (PDA), amphigenous, effuse, woolly or felty, white to pale gray or brownish gray, with varying amounts of non-sporulating aerial mycelium, no pigment is Pleosporales: Pleosporaceae: Alternaria 17

Fig. 1. Alternaria acalyphae. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

released into the medium, reaching 30-40 mm in diam. after 7 days at 25°C [Plate 1(A)]. MYCELIUM immersed or partly superficial, composed of septate, branched, hyaline to pale brown, smooth- walled hyphae, 3-7 μm wide. CONIDIOPHORES (Primary conidiophores) solitary or arising in groups of two or more, erect, usually simple, straight to flexuous, pale to olive brown, more or less cylindrical, usually only one pigmented conidiogenous site at the apex, sometimes proliferating sympodially and geniculate with one or more pigmented conidial scars, smooth, one- to several septate, slightly swollen at the base, almost hyaline at the apex of conidiogenous cell, 40-90 μm long, 5-9 μm wide; on V-8 juice agar medium, the conidiophores produced terminally and laterally on the hyphae, up to 150 μm long. CONIDIA solitary [Plate 1(B)] or occasionally in chains of 2 through the agency of a secondary conidiophore; straight or slightly curved, narrowly ovoid when juvenile, becoming long ellipsoid to obclavate at maturity, pale brown, smooth-walled, gradually 18 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 2. Alternaria acalyphae. A. conidiophore from culture; B. conidia from culture. Scale bars=50 μm.

tapering towards the long beak, some (rarely) have an apical secondary conidiophore (false beak or pseudorostrum) of variable length, analogous to the beak; conidium body 60-120×13-25 μm on host or smaller in culture 50-100×12-22 μm, with 5-10 transverse septa and 0-2 (usually 1) longitudinal or oblique septa, often slightly constricted at the transverse septa; beaks filamentous, simple, occasionally 1-branched, 1-8 septate, hyaline, up to 150 μm long, 2.5-4 μm wide along most of its length; secondary conidiophores uncommon, when present relatively broad, up to ca. 50-80× 4-8 μm.

DISTRIBUTION: Korea, Russia, USA. HOST/SUBSTRATE AND RECORDS IN KOREA: Acalypha australis L. (Euphorbiaceae) (Oh and Shin, 1999; Cho et al., 2000; Yu, 2001). SPECIMENS EXAMINED: On Acalypha australis, CNU-MH98126 (Hadong, August 1998), CNU- MH99224 (Daejeon, August 2000), CNU003517 (Hadong, July 2006), CNU073518, 073519 (Daejeon, July 2007). REMARKS: This fungus was first recorded as Macrosporium acalyphae by Nelen from Russia in 1962. Simmons (1994) transferred the genus to Alternaria after observing the Nelen specimens (holotype LE) because the taxon of the specimen was an excellent representative of large-spored, filament-beaked Alternaria. The morphological features of the author’s materials on Acalypha are in full agreement with the previous record of A. acalyphae (Nelen) E.G. Simmons (Simmons, 1994). On leaves of Acalypha, it causes circular to oval dark brown spots surrounded by a faint yellow halo. Pleosporales: Pleosporaceae: Alternaria 19

2. Alternaria alternata (Fr.) Keissler, Beihefte zum Botanischen Zentralblatt, 29: 433, 1912 (Figs. 3, 4).

SYNONYMS: Alternaria tenuis Nees, System der Pilze unt Schwämme: 68, t. 5 : 68, 1816. Torula alternata Fr., Systema Mycologicum 3: 500, 1832. Macrosporium fasciculatum Cooke & Ellis, Grevillea 6(37): 6, 1877. Macrosporium erumpens Cooke, Fung. Amer. Exs. no. 605, 1882. Macrosporium meliloti Peck, Annual Report on the New York State Museum of National History 33: 26, 1883. Macrosporium polytrichi Peck, Annual Report on the New York State Museum of National History 43: 77, 1890. Macrosporium sequierii Allesch., Hedwigia 33: 75, 1894.

COLONIES on PDA, amphigenous, effuse, olivaceous black to gray black, abundant sporulation with little aerial mycelium, no pigment is realeased into the medium, 60-80 mm in diam. after 7 days [Plate 2(A)]. MYCELIUM immersed or partly superficial; hyphae branched, septate, hyaline at first, later pale olivaceous brown, smooth-walled, 3-5 μm wide. CONIDIOPHORES (Primary conidiophores) solitary or arising in small groups of two or more, commonly branched, straight to flexuous, pale to mid brown, smooth, septate, slightly swollen and almost hyaline at the apex with a single conidial scar or geniculate with 1 to several conidial scars, variable length, but commonly short, ca. 20-50 (-100) μm long, 3-5 μm wide; in culture the conidiophores arising directly from the substrate, or produced terminally and laterally on the hyphae, up to 150 μm long, commonly develop a cluster of conidia in branching chains. CONIDIA formed in moderately long to long chains of 5-10 or more [Plate 2(B)]; often branched, initially long-ellipsoid, becoming ellipsoid to ovoid, obclavate to obpyriform, pale brown to golden brown, or smoky brown, surface usually granulate or densely verrucose, mostly beakless (erostrate), or having secondary conidiophores (false beak or pseudorostrum), some others (rarely) have a short, conical, narrowly tapered beak; chain generation mostly by means of a short secondary conidiophore, a proliferation of the apical or lateral conidium cell of the previous conidium; conidium body 15-35×8-16 μm in culture, with 3-6 transverse septa and 1 to several longitudinal or oblique septa, occasionally slightly, but not strongly, constricted at the transverse septa; secondary conidiophores usually short and 1-celled (2-4 μm long), occasionally extending geniculately to 15-40 μm.

DISTRIBUTION: Cosmopolitan. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Allium monanthum Max. (Liliaceae) (Yu, 2001); Aloe vera L. (Liliaceae) (Lee et al., 1991); Antirrhinum majus L. (Scrophulariaceae) (Lee et al., 1989); Arctium lappa L. (Asteraceae) (Lee et al., 1991); Capsicum annuum L. (Solanaceae) (Kim and Yu, 1985); Eucom- mia ulmoides Oliver (Eucommiaceae) (Lee et al., 1991); Fragaria ananassa Duchesne (Rosaceae) (Cho and Moon, 1980); Hibiscus syriacus L. (Malvaceae) (Lee et al., 1989); Lycopersicon esculentum Mill. (Solanaceae) (Yu, 2001); Oryzae sativa L. (Poaceae) (Park, 1958, 1967, under Alternaria oryzae Hara; Kang and Kim, 1972, under Alternaria tenuis Nees; Kwon et al., 1985); Sesamum indicum L. (Pedalidaceae) (Yu, 2001). SPECIMENS EXAMINED: From seeds of Capsicum annuum, CNU-MH98102 (Yeongyang, November 1997), CNU-MH98103 (Bonghwa, October 1997); on Lycopersicon esculentum, CNU-MH98105 (Buyeo, August 1998); on Phragmites australis (Cav.) Trin. ex Steud, CNU111446 (Suwon, June 2011); from seeds of Sesamum indicum, CNU-MH00682 (Geumsan, August 2000); on Taraxacum coreanum Nakai., CNU138199 (Gongju, July 2013). REMARKS: There are fundamentally differing opinions regarding the utility of morphological 20 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 3. Alternaria alternata. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

characters for taxonomy and nomenclature of A. alternata and related small-spored Alternaria species (Simmons, 1992, 1993; Simmons and Roberts, 1993; Reberts et al., 2000). One view holds that small-spored strains whose conidial dimensions fall within those published for A. alternata by Sim- mons (1967) or Ellis (1971) are best accommodated within that species. Nishimura et al. (1978, 1979) reported that the conidial sizes for all isolates that produce host-specific toxins such as A. citri Ellis & Pierce, A. gaisen Nagano (syn. A. kikuchiana Tanaka), A. longipes (Ellis & Everh.) Mason and A. mali Roberts were within the range for the collective species A. alternata, and that these fungi should be considered pathotypes of this species. Yu (1992) also reported that the isolates of host- specific toxin producers, except A. longipes, displayed sufficient characteristics in common with standard stock cultures of A. alternata (IMI 89343, IMI 147900, EGS 35-193) to be considered as the same species. Rotem (1994) proposed that all producers of host-specific toxin should be considered special forms of A. alternata. However, Simmons (1990, 1992, 1995) opposed this hypothesis Pleosporales: Pleosporaceae: Alternaria 21

Fig. 4. Alternaria alternata. A. sporulation pattern from culture; B. conidia from culture; C, D. scanning electron micrographs of ornamented conidium from culture. Scale bars: A, B=50 μm; C, D=5 μm.

based on the lack of an adequate description of conidial catenulation. According to him, typical A. alternata is defined at a minimum as a species with a three-dimensional sporulation pattern of relatively short primary conidiophores that bear a branching system of secondary conidiophores and chains of small conidia with species defining characters. Based on his criteria, Alternaria species, including host-specific toxin producers, whose conidia are produced in a single linear chain should be considered a separate taxon, even if the conidia are in the same size range as A. alternata. Kusabe and Tsuge (1995) found that sequences of the internal transcribed spacer regions in the rDNA gene of the host-specific toxin producers, clustered together with A. alternata and suggested that morphological differences between toxin-producing isolates be considered intraspecific variation. Similarly, according to Weir et al. (1998), three isolates from citrus clustered with A. alternata, lending support to the viewpoint that these isolates are intraspecific variants of A. alternata. How- 22 Fungal Flora of Korea·Alternaria and Allied Genera ever, Roberts et al. (2000) segregated 260 isolates of small-spored Alternaria into morphological groups and then subjected them to RAPD-PCR analysis using total genomic DNA and three different primers. Their analysis of RAPD fragment patterns showed that when Alternaria isolates are cultured and observed under defined conditions, their phenotypic plasticity is minimized and valid taxonomic separations can be made upon morphological characteristics. They concluded that A. alternata, A. gaisen, A. longipes and other small-spored saprotrophic or perthotrophic taxa are recognizable as morphologically distinct taxa. The results of Roberts et al. (2000) also support the conclusion of Simmons (1999) that A. alternata is a relatively rare species, and that the epithet alternata has likely been misapplied to the preponderance of isolates identified as such in the A. alternata sensu lato group. In this book, the taxa of host-specific toxin producers and other similar small-spored Alternaria are considered separately from A. alternata. This species is a saprophyte found on many kinds of plants and other substrates including food- stuffs, soil and textiles (Ellis, 1971). In Korea some species of host plants were erroneously listed as being associated with the present Alternaria species (Kim and Yu, 1985; Choi et al., 1989). All reports of A. alternata associated with various host plants should be reexamined carefully to confirm the identity of the pathogens.

3. Alternaria araliae H.C. Greene, Trans. Wisconsin Acad. Sci. 42: 80, 1953 (Figs. 5, 6).

SYNONYM: Macrosporium araliae Dearness & House, N.Y. State Mus., Circular 24: 58, 1940.

COLONIES on PDA, amphigenous, effuse, velvety or cottony, abundant development of aerial mycelium, white to pale gray, reverse pale orange to red, 53-60 mm in diam. [Plate 3(A)], sporulation abundant on PCA held in a 8/16 hr NUV light/dark cycle, no sporulation in the dark. CONI- DIOPHORES (Primary conidiophores) arising singly or in small groups of two or more, straight to flexuous, smooth-walled, pale brown, usually with only 1 pigmented terminal conidiogenous site, sometimes geniculate with 1 to 2 pigmented conidium attachment scars, slightly swollen at the apex, normally 30-150 (-200) μm long, 4-7.5 (-9) μm wide, with 0-9 septa. CONIDIA solitary or in short chains of 2-3 [Plate 3(B)], straight to slightly curved, long ellipsoid or obclavate to broadly obclavate, golden-brown to dark brown in color, smooth-walled when juvenile, becoming verruculose to verrucose, excessive cellular swelling occurs causing the conidium distorted, with 4-11 transverse septa and 1 to several longitudinal or oblique septa, slightly or strongly constricted at the transverse septa, 40-130 (-160)×12-40 (-50) μm in size with conidial body of 40-110 (-135) μm in length. Some conidia have a beak (rostrum) and some have a secondary conidiophore (pseudorostrum); beaks (when present) cylindrical, rigid, simple and unbranched, blunt-tapered, concolorous with the conidium body or slightly lighter than the body, variable in length, around 10-60 (-100) μm long, 3-8 μm wide.

DISTRIBUTION: Korea, USA. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Acanthopanax spp. (Araliaceae) (Cho et al., 2000, under A. panax Whetzel; Yu, 2001, under A. panax; Deng et al., 2015); Aralia continentalis Kitagawa (=A. cordata Thumb., Araliaceae) (Yu et al., 1992, under A. panax; Yu, 2001, under A. panax; Deng et al., 2015); Pleosporales: Pleosporaceae: Alternaria 23

Fig. 5. Alternaria araliae. Conidia, conidiophores and sporrulation habit from host (A) and culture (B). Scale bars=50 μm.

Aralia elata Seem (Araliaceae) (Cho et al., 2000, under A. panax; Deng et al., 2015; Yu, 2001, under A. panax); Echinopanax horridum (Non Decne.) Kom. (Araliaceae) (Cho et al., 2000, under A. panax; Yu, 2001, under A. panax); Kalopanax pictus Nakai (Araliaceae) (Cho et al., 2000, under A. panax; Yu, 2001, under A. panax). SPECIMENS EXAMINED: On Acanthopanax senticosus (Pupr. Et Max.) Harms, CNU-MH99099 (Geum- san, August 1999), CNU070041 (Geunsan, September 2007); on Aralia continentalis, CNU-MH99185 (Daejeon, October 1999), CNU101004 (Eumsung, July 2010), CNU138080 (Okcheon, July 2013); on Aralia elata, CNU-MH00304 (Bongpyeong, October 2000), CNU070040 (Suwon, August 2007); on Kalopanax pictus, CNU-MH99022 (Buyeo, September 1999). REMARKS: The fungus was first reported as Macrosporium araliae by Dearness and House in 1940 without Latin diagnosis. Later Greene (1953) compared the fungus from Aralia with authentic 24 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 6. Alternaria araliae. A. conidia from host; B. conidia from culture. Scale bars=50 μm.

material of Alternaria panax and cited differences in characteriatics of its conidia and conidiophores from A. panax as the basis for describing the fungus as Alternaria araliae. Simmons (1982), however, transferred the species as synonym of Alternaria panax after examining 25 specimens of Alternaria species associated with Araliaceous hosts. According to Simmons (1982), Greene’s observations were not critical enough to discover the considerable overlap in conidium shape, dimensions, and septation that exist in the two specimens, and considered A. araliae as synonym of A. panax. Recen- tly Deng et al. (2015) compared sequence analyses of seven gene regions, colony characters, conidial morphology and pathogenicity of Alternaria isolates from different hosts of Araliaceae, and found that the Alternaria from Aralia, Acanthopanax, Fatsia, and Meryta hosts differs from A. panax from Panax hosts (Deng, 2012; Deng et al., 2015). They proposed the former Alternaria species as Alternaria araliae because A. araliae Greene (1953) meets all the standards of nomenclatorial legiti- macy and is supported by excellent type material on Aralia racemosa (Simmons, 1982). The fungus causes alternaria leaf spots and blights on Araliaceae hosts, such as Acanthopaanax, Aralia, and Kalopanax.

4. Alternaria atra (Preuss) Woundenberg & Crous, Stud. Mycol. 75: 171, 2013 (Figs. 7, 8).

SYNONYM: Ulocladium atrum Preuss, Linnaea 25: 75, 1852. Stemphylium atrum (Preuss) Sacc., Sylloge Fungorum 4: 520, 1886. Pleosporales: Pleosporaceae: Alternaria 25

Fig. 7. Alternaria atra. Conidia, conidiophores and sporulation habit from culture. Scale bar=30 μm.

COLONIES on PDA, amphigenous, effuse, brown to dark brown, no pigment is secreted into the medium, reaching 30-40 mm in diam. [Plate 4(A)]. MYCELIUM pale olivaceous brown, smooth- walled or inconspicuously roughened, particularly in parts adjacent to the conidiophores, septate, branched, 3-6 μm wide. CONIDIOPHORES (Primary conidiophores) erect, straight or variously curved, ascending or arising as lateral outgrowth from the hyphae (in culture), unbranched to occasionally branched, dark brown, smooth or slightly roughened, often geniculate with many closely spaced thickened conidiogenous sites, up to 150 μm long, 5-8 μm wide. CONIDIA solitary [Plate 4(B)] or very rarely in chains of 2 through the agency of a secondary conidiophore, golden brown or dark reddish brown, verruculose to verrucose, juvenile conidia have a pointed base (obovoid), at maturity most commonly spherical to subspherical, sometimes broadly ellipsoid or obovoid, true beak absent, a very few conidia have an apical 1-celled secondary conidiophore (false beak, pseudorostrum), 15-35×15-20 μm, with 1-3 transverse and 1 or more longitudinal/oblique septa, constricted at the transverse septa, occasionally with a Y-form septation in the terminal cell(s); length/ width ratio=1.5.

DISTRIBUTION: Argentina, Australia, Canada, Chile, Cyprus, Egypt, Europe, India, Iraq, Israel, Korea, Kuwait, Libya, New Zealand, Netherlands, Norway, Pakistan, Portugal, Saudi Arabia, Sierra Leone, Tripoli, Turkey, UK, U.S.A. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Lycopersicon esculentum Mill. (Solanaceae) (Lee et al., 26 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 8. Alternaria atra. Conidia from culture. Scale bar=50 μm.

2004, under Ulocladium atrum), Soil (Yun et al., 2007, under Ulocladium atrum). SPECIMENS EXAMINED: On Armoracia lapathifolia Gilb., CNU004072 (Buyeo, May 2006); on Lycoper- sicon esculentum, CNU09037, 09038 (Buyeo, June 2004); from soil CNU09120 (Daejeon, May 2009). REMARKS: The fungus was characterized by having narrow-based juvenile conidia, and regarded as Ulocladium atrum until recently (Ellis, 1971; Simmons, 1992, 2007; Lee et al., 2004). However, Woundenberg et al. (2013) transferred U. atrum to the genus Alternaria (as A. atra) based on sequences of six gene regions (18SnrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha gene). This fungus is a common saprophyte found on many kinds of substrates, including soil. The species has been used for biocontrol of gray mold of vegetables (Lee et al., 2004; Yun et al., 2007).

5. Alternaria brassicae (Berkeley) Saccardo, Michelia 2(6): 172, 1880 (Figs. 9, 10).

SYNONYMS: Macrosporium brassicae Berk., The English Flora. 5-2(2): 339, 1836. Sporidesmium exitiosum J.G. Kühn, Hedwigia 1: 91, 1855. Cercospora bloxami Berk. & Broome, Ann. Mag. Nat. Hist., ser. 5, 9: 183, 1882. Macrosporium herculeum Ellis & Martin, American Naturalist 16: 1003, 1882. Cercospora lepidii Peck, Annual Rept. New York State Mus. Nat. Hist., 35: 140, 1884. Alternaria brassicae var. macrospora Sacc., Syll. Fung. 4: 546, 1886. Alternaria alliariae-officinalis Savul. & Sandu, Hedwigia 73: 130, 1933. Cercospora moldavica Savul. & Bontea, Herb. Mycol. Romanicum, Fasc. 27, no. 1336, 1947.

COLONIES on PDA, amphigeous, effuse, loose cottony to velvety, white to pale gray, with varying amounts of non-sporulating aerial mycelium, no pigment is secreted into the medium, reaching 35-45 mm in diam.. MYCELIUM immersed, partly superficial; hyphae hyaline to pale brown, septate, often branched, smooth-walled, 3-8 μm wide. CONIDIOPHORES (Primary coni- Pleosporales: Pleosporaceae: Alternaria 27

Fig. 9. Alternaria brassicae. Conidia, conidiophores and sporulation pattern from host (A) and culture (B). Scale bars=50 μm.

diophores) arising singly or in groups of two or more, unbranched to occasionally branched, straight to slightly flexuous, more or less cylindrical, pale gray to olivaceous gray, usually a single terminal conidiogenous site, smooth, sometimes proliferating sympodially and geniculate with 1 to several pigmented conidial scars, often slightly swollen and almost hyaline at the apex, several (up to 10) septate, 20-150 μm long, 6-10 μm wide; in culture the conidiophores as lateral extensions of the hyphae, up to ca. 150 μm long. CONIDIA solitary (Plate 5) or occasionally in short chains of 2-3 (up to 4) through the agency of a secondary conidiophore (false beak or pseudorostrum), a proliferation of the apical cell of the previous conidium; straight or slightly curved, obclavate or long ovoid, pale olivaceous brown in nature, a deeper olivaceous brown on agar medium, mostly smooth, rarely inconspicuously roughened, many have a sturdy beak, gradually tapering 28 Fungal Flora of Korea·Alternaria and Allied Genera

AB C

Fig. 10. Alternaria brassicae. A. conidiophores from culture; B. conidia from culture; C. scanning electron micrograph of a conidium from culture. Scale bars: A, B=50 μm; C=40 μm.

towards the beak, some others have an apical secondary conidiophore, analogous to the beak; on agar media chains occurring more frequently than in nature; 50-220 (-300)×15-30 (-42) μm including the blunt-taper beak portion of the spore, with 5-18 transverse and 0-7 longitudinal or oblique septa, slightly to strongly constricted at the transverse septa; beak (when present) almost cylindrical, simple and unbranched, never fragile or filamentous, commonly one-fourth to half the length of the mature conidium body, pale, usually without transverse septa or sparsely septate, 3-8 μm wide; secondary conidiophores simple or geniculate, 4-9 μm wide as short as 10 μm or as long as or longer than the conidium body; sometimes a beak enlarges terminally into a secondary conidiophore ca. 10-30 μm long, initiating a secondary conidium.

DISTRIBUTION: Cosmopolitan. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Armoracia lapathifolia Gilib. (Brassicaceae) (Lee et al., 1991; Yu, 2001); Brassica campestris subsp. napus var. pekinensis Makino (Brassicaceae) (Nakada and Takimoto, 1928; Yu, 2001); Brassica campestris subsp. napus var. nippo-oleifera Makino (Brassicaceae) (Chung et al., 1977); Brassica juncea var. integrifolia Sinsk (Brassicaceae) (Cho et al., 1997; Yu, 2001); Brassica oleraceae var. capitata L. (Brassicaceae) (Cho et al., 1997; Yu, 2001); Lepidium apetalum Wild. (Brassicaceae) (Oh and Shin, 1999); Raphanus sativus var. hortensis for. acanthiformis Makino (Brassi- caceae) (Yu et al., 1991; Yu, 2001); Rorippa cantoniensis Ohwi (Brassicaceae) (Oh and Shin, 1999); Rorippa islandica (Oed.) Borb. (Brassicaceae) (Oh and Shin, 1999). SPECIMENS EXAMINED: On Armoracia lapathifolia, CNU-MH99178 (Daejeon, October 1999), CNU111515 (Yesan, Sept. 2011); on Brassica campestris subsp. napus var. nippo-oleifera, CNU111251 Pleosporales: Pleosporaceae: Alternaria 29

(Daejeon, May 2011); on Brassica juncea var. integrifolia, CNU-MH00129 (Bongpyeong, October 2000); on Brassica oleracea var. capitata, CNU-MH98013 (Okcheon, May 1998), CNU-MH00400 (Seo- san, November 2000); on Raphanus sativus var. hortensis for acanthiformis, CMH-00372 (Ullneung island, August 2000), CNU138515 (Ullneung island, June 2013). REMARKS: Seven species of Alternaria have been recorded from Brassicaceae (Simmons, 2007), namely A. brassicae (Berkeley) Saccardo, A. brassicicola (Schwein) Wiltshire, A. brassinae E.G. Sim- mons, A. broccoli-italicae E.G. Simmons, A. ethzedia E.G. Simmons, A. japonica Yoshii (an earlier name for A. raphani Groves & Skolko) and A. nepalensis E.G. Simmons. Three of them, A. brassicae, A. brassicicola and A. japonica, have been reported from Korea. A. brassicae differs from the other two Alternaria in its sturdy beaked and larger and longer conidia; smaller conidia of the other species have no true beak. A. brassicae also differs from A. brassicicola and A. japonica in conidial catenulation; the species seldom forms a secondary conidium in nature, while A. japonica normally forms chains of 2-4 conidia and A. brassicicola forms long chains of more than 10 conidia through the agency of secondary conidiophores. The disease symptoms are known under various common names depending upon the host and the plant part attacked: gray leaf spot, black spot, alternaria black spot, alternaria gray spot of leaves, pods and stems (Corlett and Maclatchy, 1996a). Alternaria leaf spot of Chinese cabbage and other Brassicas may be caused by A. brassicae alone or in association with A. brassicicola and A. japonica.

6. Alternaria brassicicola (Schwein.) Wiltshire, Mycol. Pap., 20: 8, 1947 (Figs. 11, 12).

SYNONYMS: Helminthosporium brassicicola Schweinitz., Trans. Am. Phil. Soc., 4(2): 279, 1832. Spori- desmium exitiosum f. alternarioides J.G. Kühn, Hedwigia 1: 91, 1855. Sporidesmium exitiosum f. luxuriosums J.G. Kühn, Hedwigia 1: 91, 1855. Macrosporium circinans Berk. & M.A. Curtis, North Carolina Geol. and Nat. Hist. Surv. Papers 3: 128, 1867. Helminthosporium brassicae Henn., Hed- wigia 41: 117, 1902. Alternaria oleracea Milbraith, Bot. Gaz. Crawford. 74(3): 320, 1922.

COLONIES on PDA, amphigenous, effuse, moderate yellowish brown to brownish black, velvety, sporulation (on V-8 agar) moderate to abundant, no pigment produed on PDA, reaching 50- 60 mm in diam. [Plate 6(A)]. MYCELIUM immersed; hyphae hyaline to olivaceous brown, septate, branched, smooth-walled, 3-8 μm wide. CONIDIOPHORES (Primary conidiophores) arising singly or in groups of two or more, unbranched to occasionally branched on host, but frequently multiple branching on agar media, straight to flexuous, more or less cylindrical, usually only one conidiogenous site at the apex, sometimes proliferating sympodially and geniculate with 1 or more pigmented former conidium attachment sites, pale yellow to pale brown, almost colorless and often slightly swollen at the apex, smooth, septate, 20-60 μm long, 4-7 μm wide; in culture the conidiophores produced terminally and laterally on the hyphae, up to 80 μm long. CONIDIA mostly in moderately long to long chains of 5-10 or more [Plate 6(B)]; on agar media conidial branching and side chains occurring frequently; straight, narrowly to broadly ovoid or ellipsoid, usually first formed conidia larger and narrowly ovoid to nearly cylindrical, subsequent secondary conidia becoming progressively smaller and broader ovoid as the chain lengthens, often very regu- 30 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 11. Alternaria brassicicola. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

lar and symmetrical as to their longitudinal axis, usually tapering slightly towards the apex, the basal cell rounded, pale to olive brown, mostly smooth-walled, occasionally becoming minutely roughened with age, 20-70 (-80)×8-20 (-24) μm, with 1-8 heavily melanized transverse septa (mostly 4-6) often without longitudinal septum or a few longitudinal septa in a few of the transverse segments, often slightly or distinctly constricted at the transverse septa; true beak absent (erostrate) but having a pale apical conidiogenous structure which usually is a single celled secondary conidiophores and resembling a truncated cone, sometimes lateral secondary conidiophores also arising from intercalary conidium cells.

DISTRIBUTION: Cosmopolitan. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Armoracia lapathifolia Gilib. (Brassicaceae) (Yu, 2001); Brassica campestris subsp. napus var. pekinensis Makino (Brassicaceae) (Kang et al., 1985; Yu, 2001); Pleosporales: Pleosporaceae: Alternaria 31

AB C

Fig. 12. Alternaria brassicicola. A. conidiophore from culture; B. conidia from culture; C. scanning electron micrograph of a conidium from culture. Scale bars: A, B=50 μm; C=20 μm.

Brassica oleracea var. acephala DC (Brassicaceae) (Cho et al., 1997); Brassica oleracea var. capitata L. (Brassicaceae) (Yu and Park, 1982; Yu et al., 1991; Yu, 2001); Raphanus sativus var. hortensis for. acanthiformis Makino (Brassicaceae) (Yu and Park, 1982; Yu et al., 1991; Yu, 2001). SPECIMENS EXAMINED: On Armoracia lapathifolia, CNU-MH99220 (Buyeo, October 1999); on Brassica campestris subsp. napus var. pekinensis, CNU-MH00549 (Hongcheon, August 2000), CNU111006 (Yesan, August 2007), CNU138824 (Wonju, Sept., 2013); on Raphanus sativus var. hortensis for. acanthiformis, CNU-MH00377 (Dangjin, July 2000), CNU003263 (Buyeo, June 2008), CNU138724 (Yesan, September 2013). REMARKS: Alternaria brassicicola is characterized by ovoid, beakless conidia produced in long linear and branching conidial chains (Simmons, 1992, 1995). The species is distinguished from A. alternata whose conidia are also produced in long chains by producing small conidia with heavily melanized transverse septa. The taxonomic confusion between A. brassicae and A. brassicicola was explained by Wiltshire (1947), Corlett and MacLatchy (1996a, b), and Simmons (1995). The fungus differs from A. brassicae in having beakless, smaller conidia; the larger conidia of A. brassicae have a sturdy beak. Leaf symptoms caused by this fungus are usually darker than in A. brassicae and less clearly zonate. A. brassicae, A. brassicicola and A. japonica Yoshii may occur together on the same host, but A. brassicicola is more frequently associated with cabbage, A. brassicae with Chinese cabbage and A. japonica with radish. 32 Fungal Flora of Korea·Alternaria and Allied Genera

7. Alternaria calendulae Ondrej,˘ Cas.˘ Slezsk. Mus. Opave: 150, 1974 (Figs. 13, 14).

SYNONYMS: Macrosporium calendulae Nelen, ex Nelen & Vasiljeva, Bull. Centr. Bot. Gard., Moscow, 35: 90, 1959. Macrosporium calendulae Nelen, Botan. Mater. 15: 144, 1962. Alternaria calendulae Nirenberg, Phytopath. Zeitschr 88: 108, 1977.

COLONIES on PDA, woolly with abundant sporulation, no pigment is released into the medium, 60-65 mm in diam. after 7 days. MYCELIUM immersed or partly superficial; hyphae hyaline to pale brown, septate, branched, smooth-walled, 4-8 μm wide. CONIDIOPHORES (Primary conidiophores) arising singly or in groups of 2 or more, unbranched to occasionally branched, straight

Fig. 13. Alternaria calendulae. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm. Pleosporales: Pleosporaceae: Alternaria 33

Fig. 14. Alternaria calendulae. Conidia from host. Scale bar=50 μm.

to flexuous, stout and rigid, usually only one conidiogenous, sometimes proliferating sympodially and geniculate with 1 to 3 pigmented former conidium attachment sites, septate, smooth-walled, 50-140 μm long, 5-8 μm wide on host; in culture the conidiophores arise terminally and laterally on the hyphae, 50-200 μm long, 4-8 μm wide. CONIDIA mostly solitary on host (Plate 7), occasionally in chains of 2 through the agency of a secondary conidiophore on agar media; obclavate to long ellipsoid, sometimes long ovoid, straight to slightly curved, pale brown to mid brown, usually smooth, rarely slightly verruculose, rostrate, gradually tapering towards the beak; in culture some have an apical secondary conidiophore (pseudorostrum); conidium body 50-120×10-20 μm on host, 40-110×15-25 μm in culture, with 5-12 transverse septa and 1-2 longitudinal or oblique septa in a few of the transverse segments, often constricted at the transverse septa; the beak long, filiform, colourless to pale, septate, occasionally once branched, 40-130 (-160) μm long 4-6 μm wide at the base tapering to 2 μm; the secondary conidiophores in culture simple, sometimes geniculate, ca. 30-60 μm long, 4-6 μm wide.

DISTRIBUTION: Cyprus, Denmark, Japan, Korea, Nepal, Russia, USA. HOST/SUBSTRATE AND RECORDS IN KOREA: Calendula officinalis L. (Asteraceae) (Lee et al., 1991; Yu, 2001). SPECIMENS EXAMINED: On Calendula officinalis L., CNU-MH00804 (Daejeon, June 2000), CNU- MH00808 (Nonsan, June 2000), CNU003548 (Daejeon, August 2006). REMARKS: Yamamoto (1934) reported and described a leaf spot disease on Calendula officinalis from Japan, and assigned the causal organism to Macrosporium sp. However, he did not name the species. An attribution “Alternaria calendulae Yamamoto” appears in several Japanese publications, 34 Fungal Flora of Korea·Alternaria and Allied Genera but no evidence can be found that the name ever was proposed by anyone as a new species (Sim- mons, 1997). The attributed binomial is a nomen nudum and without nomenclatural standing wherever it has been used, e.g., in Fujioka (1952), Hara (1954) and W. Yamamoto (1960) (Simmons, 1997). According to Simmons (1997) A. calendulae Ondrej˘ meets all requirements of valid publication, however it is listed as illegitimate species in MycoBank database. More work is needed to clarify the epithet of ths fungus. The fungus causes brown to dark brown leaf spot and blight of pot marigold.

8. Alternaria cassiae Jurair & A. Khan, Pakistan J. Sci. Ind. Res. 3(1): 72, 1960 (Figs. 15, 16).

COLONIES on PDA, cottony to woolly, dark brown, zonate, with varying amounts of pale gray aerial mycelium, no pigment is released into the medium, 30-40 mm in diam. after 7 days. Sporu- lation abundant on V-8 juice agar in 12/12 hr. NUV light/dark cycle, particularly after scarifica- tion of surface mycelium. MYCELIUM composed of septate, branched, colourless to pale brown, smooth-walled, 5-7 μm wide hyphae. CONIDIOPHORES (Primary conidiophores) solitary or in fascicles, straight or slightly curved, simple, sometimes branched, more or less cylindrical, usually with a single terminal conidiogenous site, sometimes proliferating sympodially and geniculate with 1-3 pigmented former conidium attachment sites, septate, smooth-walled, pale brown 30-150 μm long, 5-6 μm wide; in culture the conidiophores produced as lateral extensions of the mycelium. CONIDIA mostly solitary (Plate 8), sometimes in chains of two through the agency of a secondary conidiophore, straight or slightly curved, obclavate to long ellipsoid, or narrowly ovoid, light to dark brown, verrucose to verruculose when juvenile, becoming smooth at maturity, many

AB C

Fig. 15. Alternaria cassiae. A. conidiophores form culture; B. conidia from host; C. conidia from culture. Scale bars=50 μm. Pleosporales: Pleosporaceae: Alternaria 35

Fig. 16. Alternaria cassiae. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

have a filiform beak, some are beakless (erostrate) with a concoid apical cell, some others have a secondary conidiophore (pseudorostrum), analogous to the beak; conidium body 40-115×16-27 (-37) μm on host, with 6-11 transverse septa and 1-several longitudinal or oblique septa, slightly constricted at the transverse septa; the beak on host usually long, filiform, unbranched, hyaline to pale brown, the point of transition from spore body to beak is definite and conspicuous, variable in length, 30-150 (-230) μm long, 2-3 μm wide along most of its length, on agar media beakless conidia common, beaks often shorter than those in nature; secondary conidiophores terminate abruptly from its site of origin in the body apex 12-40 μm long, 5-6 μm wide; sometimes the tip of a beak converted into a secondary conidiophores ca. 10-12 (-30) μm long.

DISTRIBUTION: Africa (South Africa); Asia (India, Korea, Pakistan); North America (USA). HOST/SUBSTRATE AND RECORD IN KOREA: Cassia tora L. (Fabaceae) (Yu, 2001). 36 Fungal Flora of Korea·Alternaria and Allied Genera

SPECIMENS EXAMINED: From seeds of Cassia tora, CNU003553 (Daejeon, November 2006), CNU113136 (Eumsung, November 2011); on Cassia tora, CNU-MH01001 (Daejeon, February 2001). REMARKS: This species was first recorded in Korea by Yu (2001). It is closely related to A. zinnae H. pape ex M.B. Ellis in conidial morphology. The fungus, however, can be differentiated from the latter by its variable conidia having a range of beaked, non-beaked and secondary conidiophores bearing conidia (David, 1991; Simmons, 1982). On leaves of Cassia or sickle senna, causing leaf spots; also present on seeds.

9. Alternaria cinerariae Hori & Enjoji, J. Plant Protect. Tokyo: 432, 1931 (Figs. 17, 18).

SYNONYM: Alternaria senecionsis Neergaard, Danish Species of Alternaria and Syemphylium: 201, 1945.

COLONIES on PDA, effuse, cottony, aerial mycelium fluffy, olivaceous gray to dark gray, sporulation (on V-8 agar) moderate to abundant, no pigment is released into the medium, 55-65 mm in diam. after 7 days. CONIDIOPHORES (Primary conidiophores) usually solitary, simple and unbranched on host, frequently branched in culture, straight to slightly curved, more or less cylindrical, pale to olivaceous brown, often slightly swollen at the apex, with a pigmented zone at the site of former conidium attachment, several (-5) septate, 50-150 μm long, 5-8 μm wide; in culture

ABC

Fig. 17. Alternaria cinerariae. A. conidiophores from host; B, C. conidia from host. Scale bars=50 μm. Pleosporales: Pleosporaceae: Alternaria 37

Fig. 18. Alternaria cinerariae. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

the conidiophores are produced terminally and laterally on the hyphae. CONIDIA solitary (Plate 9) or in short chains of 2-3 through the agency of a secondary conidiophore, on agar media chains of 2-5 occurring more frequently than in nature; straight or slightly curved, obclavate or narrowly ovoid to ellipsoid on host, obclavate to broadly obclavate in culture, erostrate with a short broadly tapered apical cell when juvenile, pseudorostrate with a relatively broad secondary conidiophores at maturity, medium yellow brown, smooth-walled, some conidium cells becoming swollen and inflated with age in culture; conidium body (excluding any secondary conidiophore) 40-100 (-140) ×15-40 (-50) μm, with 5-10 transverse septa and 1-2 longitudinal septa in 1-4 of the transverse segments, slightly or sharply constricted at the transverse septa; secondary conidiophores (false beaks or pseudostra) at the apical cell of the conidium relatively broad, paler or the same colour as the conidium body, simple, sometime geniculate (in culture), lightly swollen at the apex, up to 90 μm long, 5-9 μm wide, initiating a secondary conidium. 38 Fungal Flora of Korea·Alternaria and Allied Genera

DISTRIBUTION: Europe (Denmark and UK), Japan, Korea, USA. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Senecio flammeus Turcz. (Asteraceae) (Yu, 2001); Senecio vulgaris L. (Asteraceae) (Oh and Shin, 1999). SPECIMEN EXAMINED: On Senecio flammeus, CNU3317 (=CMH00104) (Cheongyang, July 2000). REMARKS: This species was first recorded in Korea by Oh and Shin (1999), and Yu (2001) described its morphological characteristics. Alternaria cinerariae as initially described by Hori and Enjoji (1931) in Japan and later by Neergaard (1945) under the name of A. senecionsis Neergard is now considered a morphologically distinctive species restricted to Senecio spp. hosts. A. cinerariae is characterized by ovoid to ellipsoid, or obclavate, and apically pseudorostrate, catenate spores. Korean materials of the present author were in good agreement with the previous records of A. cinerariae (Enjoji, 1931; Simmons, 1997). On leaves of Senecio, it causes reddish brown to olivaceous black spots.

10. Alternaria citri Ellis & N. Pierce, Bot. Gaz. Crawford. 33: 234, 1902 (Figs. 19, 20).

COLONIES on PDA, gray to olivaceous brown or black, sometimes zonate, often become sterile after subculturing, no pigment is released into the medium, 60-70 mm in diam. after 7 days. CONIDIOPHORES (Primary conidiophores) solitary or arising in groups, straight or flexuous, unbranched to branched, pale to mid brown or olivaceous brown, smooth, septate, with a pigmented conidiogenous site at the apex, or proliferating sympodially and geniculate with 1-5 pigmented conidiogenous sites, 30-87 (-100) μm long, 2-5 μm wide. CONIDIA usually in simple chains of 2- 5(-8), occasionally solitary; straight or slightly curved, variable in shape but commonly broadly

Fig. 19. Alternaria citri. A, B. scanning electron micrographs of conidiophore and conidia from culture. Scale bars: A=20 μm; B=5 μm. Pleosporales: Pleosporaceae: Alternaria 39

Fig. 20. Alternaria citri. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

ovoid, oblong-ellipsoid or obpyriform, pale to olivaceous brown, densely verruculose to inconspicuously ornamented, often with slightly distended cells, beakless (erostrate) or having a secondary conidiophore (pseudostrum), often bluntly tapered apically; conidium body 24-42 (-50)×10-20 (-27) μm, with 3-5 transverse septa and 1-2 longitudinal or oblique septa in a few of the transverse divisions, constricted at the transverse septa; secondary conidiophores are produced at the apical or lateral cells of the conidium body, mostly short and 1-2 celled ca. 2-6×2-4 μm, occasionally geniculate or extend to 30 μm.

DISTRIBUTION: Worldwide. HOST/SUBSTRATE AND RECORD IN KOREA: Citrus sinensis Obseck (Rutaceae) (Yu, 2001). SPECIMENS EXAMINED: On fruit of Citrus sinensis, CNU03741 (=CMH-00797) (Jeju, December 2000), 40 Fungal Flora of Korea·Alternaria and Allied Genera

CNU132006 (Jeju, June 2013). REMARKS: The conidial features of Alternaria citri were described and illustrated by Simmons (1990) on the basis of the type specimen. A. citri is characterized by having broadly ovoid or oblong-elliptical, apically blunt, beakless conidia with slightly distended cells and with ill-defined secondary septa (Simmons, 1990). This species can be differentiated from A. alternata by conidial shape and branching pattern of chains of conidia. The conidia of A. alternata are mainly ovoid or ellipsoid whereas those of A. citri are mostly broadly ovoid or obpyriform. A. alternata has a three- dimensional pattern of multiple branching sporulation in laboratory culture conditions, while A. citri has a sporulation pattern of brevicatenate conidia with short side branches. A. citri was originally described only from an internal dry rot of orange fruit (Pierce, 1902). However, Alternaria isolates associated with alternaria leaf spot of rough lemon (Citrus jambhiri Lush.) and brown spot of tangerine (Citrus reticulata Blanco) usually have been referred to as A. citri or as pathological variants (pathotypes) of A. alternata (Whiteside, 1976a, b; Nishimura et al., 1978). Simmons (1999a, b) differentiated several new species (A. limoniasperae E.G. Simmons and A. citrimacularis E.G. Sim- mons from leaf spots of rough lemon, and A. tangelonis E.G. Simmons, A. turkisafria E.G. Simmons, A. citriarbusti E.G. Simmons, A. toxicogenica E.G. Simmons, A. colombiana E.G. Simmons, A. perangusta E.G. Simmons, A. interrupta E.G. Simmons and A. dumosa E.G. Simmons from leaf spots of tangelo) among various Alternaria species from citrus hosts that had been variously named A. citri or A. alternata by others. According to Simmons (1999a, b), the so-called “A. alternata rough lemon pathotype” and “A. alternata tangerine pathotype” are neither A. alternata nor A. citri. They are now referred to A. limoniasperae and A. toxicogenica, respectively (Simmons, 1999). On fruits of Citrus, it causes black rot. Symptoms usually begin at the button or at one end, spreading to the axis, sunken, soft, dark green to black with abundant sporulation.

11. Alternaria cosmosa Y.L. Li & W.S. Wu, Mycotaxon 91: 15, 2005 (Figs. 21, 22).

COLONIES on PDA, effuse, cottony or woolly, white or pale to dark gray; reverse yellow or pale to dark orange, 60-70 mm diam. after 7 days at 25°C [Plate 10(A)]. MYCELIUM scanty, aerial or immersed in the agar, hyphae septate, hyaline to pale olivaceous brown, 3-7 μm wide. CONIDIO- PHORES (Primary conidiophores) arising singly, rarely in groups of 2-3, erect, unbranched to occasionally branched, straight to flexuous, pale brown, almost hyaline at the apex, smooth-walled with 1-7 septa, usually only one pigmented terminal conidiogenous site, sometimes with one more lateral conidiogenous locus, 25-120 (-140) μm long, 5-8 μm wide; in culture, produced singly as side branches on the main hyphae, simple, occasionally (rarely) 1-geniculate, scattered in the aerial mycelium. CONIDIA solitary [Plate 10(B)], straight or slightly curved, long obclavate, pale olivaceous brown in nature, a deeper olivaceous brown on agar medium, smooth, rostrate, tapering towards a long beak, conidium body 60-130 (-150)×5-15 (-25) μm, with 2-13 (-16) transeverse septa, 0-4 longitudinal or oblique septa in a few of the transverse segments, slightly constricted at the transverse septa; the beak filamentous, filiform, subhyaline to pale brown, the point of transition from spore body to beak is smooth, often branched once, twice or more, up to 200 (-220) μm long, 3-5 μm wide at the base tapering to 1-2 μm.

DISTRIBUTION: Korea, Taiwan. Pleosporales: Pleosporaceae: Alternaria 41

Fig. 21. Alternaria cosmosa. Conidia, conidiophores and sporulation habit from culture. Scale bars= 50 μm.

HOST/SUBSTRATE AND RECORDS IN KOREA: Cosmos bipinnatus Cav. (Asteraceae) (Cho, 2004 under Alternaria sp.; Deng et al., 2014). SPECIMENS EXAMINED: On Cosmos bipinnatus, CNU111459, 111499 (Nonsan, September 2011), CNU 137007 (Hadong, September 2013). REMARKS: Alternaria cosmosa was reported from seeds of common cosmos (Cosmos bipinnatus) as a new taxon by Wu and Li from Taiwan (2005). In Korea, an Alternaria sp. had previously been reported from leaves of the common cosmos and the fungus was considered as a new species (Cho, 2004). However, a new epithet for the species has not been provided. Conidial morphology of the Korean species of Alternaria from the cosmos was in agreement with the description of A. cosmosa from Taiwan, which has a long obclavate body with long narrow-taper beaks (usually 1, but up to 4). According to Simmons (2007), the taxon cannot be compared critically with other taxa of this 42 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 22. Alternaria cosmosa. Conidia from culture. Scale bar=50 μm.

morphological group except in parallel culture, and published information of the taxon resembles A. danida on Ageratum. Simmons (2007) considered A. cosmosa a synonym of A. danida. The present species, however, differs from A. danida in having solitary conidia (while A. danida occasionally has a secondary conidiophore yielding a chain of two conidia) with narrower conidial body. Further comparative studies on this group of Alternaria are necessary. The fungus causes brown spots and blights on lower leaves of common cosmos and yellow cosmos.

12. Alternaria cucumerina (Ellis & Everh.) J.A. Elliot, Amer. J. Bot. 4: 472, 1917 (Figs. 23, 24).

SYNONYM: Macrosporium cucumerinum Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 47: 440, 1895.

COLONIES on PDA, effuse, cottony, olive brown to dark brown with varying amounts of pale gray non-sporulating aerial mycelium, no pigment is released into the medium, reaching about 45- 55 mm in diam. after 7 days [Plate 11(A)]. MYCELIUM partly superficial or immersed in the agar, hyphae septate, hyaline to pale olivaceous brown, 3-7 μm wide. CONIDIOPHORES (Primary Pleosporales: Pleosporaceae: Alternaria 43

Fig. 23. Alternaria cucumerina. Conidia, conidiophores and sporulation habit from culture. Scale bars=50 μm.

conidiophores) on V-8 agar, arising from hyphae or from a sparse network of aerial hyphae, unbranched to occasionally branched, erect, straight to flexuous, cylindrical, usually one pigmented terminal conidiogenous scar, sometimes geniculate with 1-3 pigmented conidial scars, one to several septate, pale to olive brown, slightly paler at the apex, smooth, up to 110 μm long, 6-10 μm wide. CONIDIA (on V-8 agar) solitary, or occasionally in chains of 2 [Plate 11(B)]; individual conidia in chains separated by a secondary conidiophore produced by an apical cell of conidium, basically obclavate, sometimes long-ovoid or broadly-ellipsoid, light brown to mid-golden brown or dark brown, straight to slightly curved, smooth to verruculose, rostrate, and tapering towards the beak; spore body 35-130×13-34 μm, with 5-10 transverse septa and 1 to several longitudinal or oblique septa in a few of the transverse divisions, slightly or sharply constricted at the transverse septa; the beak long, filiform, narrows abruptly from its site of origin in the body apex, 44 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 24. Alternaria cucumerina. Conidia from culture. Scale bar=50 μm.

colourless or pale brown, septate, simple and not branched, often much longer than the conidium body, much often 100-120 (up to 300) μm long, 4-5 μm wide at the base tapering to 2-3 μm.

DISTRIBUTION: Africa (Kenya, Mozambique, Nigeria, Rhodesia, Sierra Leone, South Africa, Sudan, Zambia); Asia (Japan, Korea, Thailand); Australasia and Oceania (Australia, New Zealand); Europe (England, France, Romania); Central and South America (Chile, Cuba, Venezuela); North America (Canada, USA). HOSTS/SUBSTRATES AND RECORDS IN KOREA: Citrullus lanatus (Thunb.) Matsum. & Nakai (Cucurbitaceae) (Kim et al., 1994; Yu, 2001); Cucumis sativus L. (Cucurbitaceae) (Park, 1961, 1967); Cucurbita spp. (Cucurbitaceae) (Park, 1961, 1967; Yu, 2001). SPECIMEN EXAMINED: On Cucurbita moschata, CNU03450 (Gochang, October 2000); on Cucurbita spp., CNU- CH9421 (Daejeon, November 2000). REMARKS: Seven species of Alternaria have been reported from Cucurbitaceae hosts (Simmons, 2007). They are A. brassicae, A. caudata (Cooke & Ellis) E.G. Simmons, A. cucumericola E.G. Simmons, A. cucumerina, A. cylindrorostra T.Y. Zhang, A. loofahae E.G. Simmons and A. peponicol (Rabenhorst) E.G. Simmons. In Korea, only A. cucumerina has been reported from Cucurbitaceae (Anonymous, 2009). A. cucumerina causes leaf spot of cucumber, melon, pumpkin and watermelon. Typical symptoms appear as circular or irregular, yellow-brown spots with a light green halo. Mature lesions have a yellowish center with brown margin. Pleosporales: Pleosporaceae: Alternaria 45

13. Alternaria dauci (J.G. Kühn) J.W. Groves & Skolko, Canad. J. Res., Sec. C., 22(5): 222, 1944 (Figs. 25, 26).

SYNONYMS: Sporodesmium exitiosum var. dauci J.G. Kühn, Hedwigia 1: 91, 1855. Macrosporium dauci (J.G. Kühn) Rostr., Tidsskrift for Landoekonomi, ser. 5, 7: 385, 1888. Macrosporium carotae Ellis & Langl., Journal of Mycology 6: 36, 1890. Alternaria brassicae var. dauci (J.G. Kühn) Lindau, Raben- horst’s Kryptogamen-Flora, Pilze-Fungi imperfecti 1(9): 260, 1908. Polydesmus exitiosus var. dauci (J.G. Kühn) Sacc., Sylloge Fungorum 22: 1410, 1913. Alternaria porri f. sp. dauci (J.G. Kühn) Neergaard, Danish Species of Alternaria and Stemphylium: 252, 1945.

COLONIES on PDA, olive gray or grayish green, cottony to woolly with abundant development

Fig. 25. Alternaria dauci. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm. 46 Fungal Flora of Korea·Alternaria and Allied Genera

AB C

Fig. 26. Alternaria dauci. A. conidiophore from culture; B. conidia from culture; C. scanning electron micrograph of a conidium from culture. Scale bars: A, B=50 μm; C=5 μm.

of aerial mycelium, sometimes producing a red or reddish brown pigment which diffuses into the medium, 50-60 mm in diam. after 7 days. MYCELIUM immersed or partly superficial; hyphae hyaline to pale brown, septate, branched, smooth-walled, 3-8 μm wide. CONIDIOPHORES (Pri- mary conidiophores) mostly solitary, rarely in groups of 2-3, unbranched to occasionally branched, straight or curved, stout and rigid, pale to medium olivaceous brown, smooth, usually only a single terminal conidiogenous site, sometimes 1-2 geniculate and conidiogenous, often slightly swollen at the apex, 2-7 septate, 30-120 μm long, 5-8 μm wide; in culture the conidiophores produced as side branches on the main hyphae. CONIDIA predominantly solitary (Plate 12), but occasionally in chains of 2 (3) through the agency of a secondary conidiophore (false beak or pseudorostrum); obclavate or long ellipsoid on host, ovate-obclavate on agar, straight or curved, pale olivaceous brown when juvenile, becoming dark brown at maturity, mostly smooth, occasionally minutely roughened, rostrate and tapering gradually towards a narrow beak; conidium body 50-120×15-25 μm, with 5-11 transverse septa and usually 1 rarely 2 longitudinal or oblique septa in a few of the transverse segments, often slightly (on host) or strongly (on agar) constricted at the transverse septa; the beak long, filiform, narrows gradually from its site of origin in the body apex, colourless or pale, septate, often branched once (bifurcate), rarely branched twice (trifurcate), up to 3 times the length of the spore body, 50-250 μm long, 5 μm wide at the base tapering to 2-3 μm; occasionally (uncommonly) a relatively broad and sturdy secondary conidiophore develops from the apical conidium cell, up to ca. 15-50×6 μm, generates a secondary conidium with a typical narrowly- tapered beak.

DISTRIBUTION: Nearly throughout the world wherever the crop is cultivated, including Korea and Japan. HOST/SUBSTRATE AND RECORDS IN KOREA: Daucus carota var. sativa DC. (Apiaceae) (Park, 1961; Yu Pleosporales: Pleosporaceae: Alternaria 47 and Park, 1982; Yu, 2001). SPECIMENS EXAMINED: On Daucus carota var. sativa, CNU3566 (=CMH00285) (Pyeongchang, August 2000), CNU3568 (Jeju island, August 2000), CNU3918 (Pyeongchang, September 2006), CNU111568 (Jeju, July, 2011). REMARKS: Eight species of Alternaria, namely A. atrocariis E.G. Simmons, A. carotiincultae E.G. Simmons, A. dauci, A. daucicaulis E.G. Simmons, A. daucifolii E.G. Simmons, A. novae-zelandiae E.G. Simmons, A. radicina Meier, Drechsler & E.D. Eddy and A. zhengzhouensis T.Y. Zhang, have been recorded on Daucus carota L. (Simmons, 2007). In Korea only two species of them, A. dauci and A. radicina, have been reported (Anonymous, 2009). A. dauci differs from A. radicina in its filiform beaked and larger conidia; the smaller conidia of the other species have no true beak. Causing leaf blight of carrot. Affected leaves and petioles turn brown or black and the whole top may be killed when infection is severe.

14. Alternaria dianthicola Neergaard, Danish Species of Alternaria and Stemphylium: 189, 1945 (Figs. 27, 28).

COLONIES on PDA, velvety, sometimes fluffy or loose cottony, white to pale gray, 30-40 mm in diam after 7 days. CONIDIOPHORES (Primary conidiophores) solitary or in groups of 2-4, mostly unbranched, straight or curved, more or less cylindrical, pale olive to olive brown, smooth, 1-7 septate, usually with a single pigmented conidiogenous site, occasionally geniculate with 1-2 pigmented conidial scars, up to 150 μm long, 4-6 μm wide; in culture the conidiophores are produced as lateral extensions of the hyphae. CONIDIA solitary or frequently in chains of 2-3, on agar media chains of 3-5 occuring frequently, obclavate to almost cylindrical, straight or slightly curved, pale olivaceous brown, smooth-walled; some have a long beak, others have an apical secondary conidiophore (pseudorostrum) analogous to the beak, conidium body 20-80 (-85)×10-21

Fig. 27. Alternaria dianthicola. A. conidiophores from host; B conidia from host. Scale bars=50 μm. 48 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 28. Alternaria dianthicola. Conidia, conidiophores and sporulation pattern from host (A) and culture (B). Scale bars=50 μm.

μm, with 3-10 transverse septa and occasionally (uncommonly) only one longitudinal or oblique septum in 1-2 of the transverse divisions, often slightly constricted at the transverse septa; beaks when present, sturdy or almost cylindrical, simple and unbranched, subhyaline or the same colour as the body, 10-80 μm long, 3-5 μm wide; secondary conidiophores usually simple, occasionally geniculate, as short as 10 μm or up to 90 μm long, 3-7 μm wide.

DISTRIBUTION: Africa (Malawi, South Africa, Zambia, Zimbabwe); Asia (Bhutan, Korea, Malaya- sia, Thailand); Australasia and Oceania (New Zealand); Europe (Austria, Cyprus, Denmark, France, Germany, Italy, Romania, The Netherlands); Central and South America (Brazil, Chile, Jamaica); North America (USA). HOSTS/SUBSTRATES AND RECORDS IN KOREA: Dianthus sinensis L. (Caryophyllaceae) (Yu, 2001); Dianthus superbus var. longicalycinus (Max.) Williams (Caryophyllaceae) (Yu, 2001). Pleosporales: Pleosporaceae: Alternaria 49

SPECIMENS EXAMINED: On Dianthus sinensis, CNU-MH00757 (Cheongyang, July 2000); on Dianthus superbus var. longicalycinus, CNU003328 (Bongwha, July 2006). REMARKS: Five species of Alternaria have been described from Dianthus spp. (Simmons, 2007). They are A. dianthicola, A. ellipsoidea E.G. Simmons, A. longispora McAlpine, A. nobilis (Vize) E.G. Simmons and A. tenuissima (Nees) Wiltshire. Two species of Alternaria, A. dianthicola and A. nobilis (Syn. A. dianthi), have been reported from Dianthus spp. in Korea (Anonymous, 2009; Yu, 2001). A. dianthicola can be distinguished from A. nobilis in that A. nobilis has many more longitudinal septa, the beak or the secondary conidiophores is shorter and the conidia are broader and darker (David, 1991; Simmons, 2007; Yu, 2001). A. dianthicola causes leaf spot and bud-rot of Dianthus. Symptoms are oval or irregular, whitish yellow or pale brown spots with dark margin, becoming black following sporulation in the center of the spot.

15. Alternaria embellisia Woudenberg & Crous, Stud. Mycol. 75: 171, 2013 (Figs. 29, 30).

SYNONYM: Helminthosporium allii Campan., Nuov. Ann. Agricolt. Roma 4: 87, 1924. Embellisia allii (Campan.) E.G. Simmons, Mycologia 63: 382, 1971.

COLONIES on PDA, effuse, velvety or powdery, dark brown to dark blackish brown, no pigment is released into the medium, 50-65 mm in diam. after 7 days (Plate 13). MYCELIUM immersed or superficial; hyphae elongate, sparsely branched, pale brown. CHLAMYDOSPORES often formed on host but not so frequently in culture, dark brown to black, up to 80 μm. CONIDIOPHORES (Primary conidiophorea) solitary or arising in groups of 2-10, straight or flexuous, arising as lat-

Fig. 29. Alternaria embellisia. Conidiophore and conidia from culture. Scale bar=25 μm. 50 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 30. Alternaria embellisia. Conidia, conidiophores and sporulation habit from culture. Scale bars=25 μm.

eral outgrowth from hyphae (in culture), pale to dark brown, darker at the base, becoming paler towards the apex, usually smooth walled, occasionally roughened, septate, sympodial, with several geniculations, conidiogenous scars thickened and pigmented, up to about 120 μm long, 5-10 μm thick. CONIDIA solitary, occasionally in chains of 2-3 through the agency of secondary conidiophores, variable in shape and size, commonly long ellipsoid to subcylindrical, sometimes irregular in shape, becoming curved and waisted, mid to dark brown, smooth or slightly roughened (in culture), very dark and thickened transverse and occasionally 1 or 2 oblique or longitudinal septa and may reach 2 μm in thickness, 20-50×10-15 μm, beak absent, sometimes forming short secondary conidiophores which bear several conidia.

DISTRIBUTION: Cosmopolitan. HOST/SUBSTRATE AND RECORD IN KOREA: Allium sativum L. (Liliaceae) (Lee et al., 2002). SPECIMENS EXAMINED: On bulb scale of garlic (Allium sativum), CNU93029 (Seosan, May 2009), CNU1043210 (Seosan, May 2010). REMARKS: Simmons erected the genus Embellisia with E. allii as the type species in 1971. He differentiated the genus with Alternaria by very dark and thickened transverse septum layer. Wou- denberg et al. (2013), however, treated this fungus as Alternaria embellisia of section Embellsia based on nucleotide sequence data of parts of the 18S nrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha gene regions. Pleosporales: Pleosporaceae: Alternaria 51

The fungus causes bulb canker of garlic and was first reported in Korea by Lee et al. (2002).

16. Alternaria gaisen Nagano ex Hara, J. Japanese Hort. Society 32(3): 16, 1920 (Figs. 31, 32).

SYNONYM: Alternaria kikuchiana Tanaka, Mem. Coll. Agric., Kyoto Univ. 28: 27, 1933.

COLONIES on PDA, gray to gray green or black, cottony or velvety, with varying amounts of non-sporulating aerial and surface mycelium, no pigment is released into the medium, reaching

Fig. 31. Alternaria gaisen. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm. 52 Fungal Flora of Korea·Alternaria and Allied Genera

A B

Fig. 32. Alternaria gaisen. A. conidia from host; B. scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=5 μm.

60-70 mm in diam after 7 days. CONIDIOPHORES (Primary conidiophores) solitary or arising in small groups, predominantly unbranched, rarely branched, straight or slightly bent, dilute yellowish brown to olive brown, smooth, septate, usually with a single pigmented terminal conidiogenous site, occasionally geniculate with 1-2 pigmented conidial scars, up to 150 μm long, 3-5 μm wide; in culture the conidiophores produced directly from the surface and as laterall extensions of the hyphae, branching increased. CONIDIA mostly in short or moderately long chains of 3-10 through production of secondary conidiophores both on host tissue and in culture (Plate 14); commonly unbranched, sometimes branched (particularly in culture); elongated ovoid or long ellipsoid in nature, ovoid or ellipsoid in culture, yellowish brown, conspicuously verruculose or minutely roughened, mostly beakless (erostrate) or having an apical secondary conidiophore (false beak or pseudorostrum), some have a relatively short beak (in nature), some of the young conidia awl- shaped; conidium body 20-55 (-60)×10-15 μm in nature, with 3-7 transverse septa and 1-2 longitudinal or oblique septa in a few of the transverse compartments, often slightly constricted at the 1-3 of the transverse septa, occasionally have a dark median transverse septum that is more con- stricting than other septa; the beak, when present, tapering gradually from its site of origin in the body apex, as long as, or shorter than, the conidium body; secondary conidiophores, common in culture, terminate abruptly from its site of origin to the body apex, usually short and 1 celled, ca. 3-6 μm long, 3-5 μm wide, sometimes elongate up to 35 μm long.

DISTRIBUTION: Asia (China, Japan, Korea); Australasia and Oceania (Australia, unconfirmed). HOST/SUBSTRATE AND RECORDS IN KOREA: Pyrus serotina Rehder (Rosaceae) (Nakada and Taki- moto, 1928; Park and Yu, 1988; Ryu et al., 1993; Yu, 2001). SPECIMENS EXAMINED: On Pyrus serotina (cv. the 20th Century), CNU-MH98100 (Naju, September 1998), CNU-MH98113 (Daejeon, August 1998), CNU003759 (Naju, July 2006). REMARKS: The present species has been well known as one of the several host specific toxin (HST)-producing species and studied intensively by S. Nishimura and his colleagues in Japan. Nishimura et al. (1978, 1979) considered the species as variants of A. alternata on the basis of mis- application of information in Simmons (1967). A. gaisen, however, can be readily distinguished Pleosporales: Pleosporaceae: Alternaria 53 from A. alternata by conidial size and catenulation pattern. A. alternata produces a loose; three dimensional cluster of branching chains with smaller conidia under specific growth conditions whereas A. gaisen produces short or moderately long, unbranched chains. There seem to be few morphological differences between A. gaisen and A. tenuissima, but Simmons (1998) distinguished them on the basis of conidium size, chain length and secondary conidiophore length in young colonies. A. gaisen has smaller conidia and typically lacks appreciable numbers of secondary conidiophores longer than a single cell. However, these differences are somewhat minor and change- able. Furthermore, this species is also closely related to other brevicatenate species of Alternaria such as A. destruens E.G. Simmons (1998) an A. eryngii (Pers.: Fr.) Hughes and E.G. Simmons (1998). Further comparative studies for these species including molecular work are necessary. On leaves, petioles and young fruits of the Asian pear (only certain susceptible varieties such as the 20th Century and Sinsu), causing dark and black spots.

17. Alternaria gossypina (Thüm.) J.C.F. Hopkins, Trans. Br. Mycol. Soc., 16(2-3): 136, 1931 (Figs. 33, 34).

SYNONYM: Macrosporium gossypium Thüm., Heb. Mycol. Oeconom. no. 513, 1877.

COLONIES on PDA, cottony or velvety, olive green or light brown, darker in the center with sporulation, with varying amounts white aerial mycelium, no pigment is released into the medium, 60-70 mm in diam after 7 days. CONIDIOPHORES (Primary conidiophores) solitary or arising in

Fig. 33. Alternaria gossypina. Conidia from host. Scale bar=50 μm. 54 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 34. Alternaria gossypina. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

small groups, mostly unbranched, smooth-walled, usually only one pigmented conidiogenous site at the apex (on host), sometimes proliferating sympodially and geniculate with 1 to several pigmented conidiogenous sites (in culture), 80-120 μm long, 3-5 μm wide; in culture the conidiophores arising terminally or laterally on the hyphae. CONIDIA mostly in short or moderately long chains of 3-10 through the agency of secondary conidiophores, usually unbranched, sometimes branched (particularly in culture); dark brown, mostly conspicuously verruculose, occasionally minutely roughened, obclavate to narrowly ovoid on host, ovoid to ellipsoid in culture, occasionally have a short beak (on host), mostly beakless or having a secondary conidiophore (in culture); conidium body 30-45 (-55)×15-25 μm, with 3-9 transverse septa and 1-2 longitudinal or oblique septa in a few of the transverse divisions, slightly constricted at the transverse septa, occasionally having a darker median transverse septum that is more constricted than other septa; the beak, when present, gradually tapering, as long as, or shorter than, the body; 8-30 (-50) μm long, 4 μm wide at Pleosporales: Pleosporaceae: Alternaria 55 the base tapering to 2.5 μm; secondary conidiophores at the apical conidium cell, usually short and 1-celled, often extended to 12×2.5-5 μm.

DISTRIBUTION: Africa (Ivory Coast, Kenya, Malawi, Mozambique, Nigeria, South Africa, Sudan, Uganda, Zambia, Zimbabwe); Asia (India, Iran, Korea, Pakistan); Australasia (Australia); Europe (Italy, Yugoslavia); North America (USA); South America (Trinidad). HOST/SUBSTRATE AND RECORDS IN KOREA: Gossipium indicum Lam. (Malvaceae) (Yu, 2001). SPECIMEN EXAMINED: On Gossipium indicum, CNU-MH00276, CNU03722 (Mokpo, September 2000). REMARKS: Four species of Alternaria, A. gossypina, A. gossypii (Jacz.) Nishikado, Kimura & Mya- waki, Alternaria macrospora Zimm and A. malvacearum E.G. Simmons have been recorded on the host Gossipium (Simmons, 2007), only two of which, A. gossypina and A. macrospora, have been reported in Korea (Anonymous, 2009; Yu, 2001). A. macrospora differs from A. gossypina in producing larger conidia with a very long beak, usually solitary or rarely in chains of two, whereas A. gossypina produces small conidia in moderately long chains of 3-9. A. gossypii has smaller, smooth, beakless conidia, 22-27×9-11 μm, with usually only one longitudinal septum (Nishikado et al., 1940). The fungus recorded as A. alternata on Gossypium (Rotem et al., 1988) is apparently synonymous with either A. gossypii or A. gossypina. Further comparative studies on the small-spored Alternaria species on Gossypium are necessary. This fungus can cause boll rot of cotton.

18. Alternaria helianthinficiens E.G. Simmons, Walcz & R.G. Roberts, Mycotaxon 25(1): 204, 1986 (Figs. 35, 36).

SYNONYM: Alternaria helianthiinficiens E.G. Simmons, Walcz & R.G. Roberts Mycotaxon 25(1): 204, 1986.

COLONIES on PDA, effuse, cottony, with little surface mycelium and moderate amount of sub- merged mycelium, somewhat slowly growing, usually secreting a yellow or reddish yellow pigment into the medium, 35-45 mm diam. after 7 days. CONIDIOPHORES (Primary conidiophores) solitary, rarely in small groups of 2-3, unbranched, straight or flexuous, more or less cylindrical, pale or olivaceous gray, smooth, septate, usually with a single conidiogenous site at the apex, sometimes proliferating sympodially and geniculate with 1-2 pigmented conidial scars, often slightly swollen at the apex, up to 150-200 μm long, 4-6.5 μm wide; in culture (on V-8 agar) conidiophores produced terminally and laterally on the main hyphae. CONIDIA mostly solitary (Plate 15), rarely in chains of 2; ellipsoid, ovoid, or broadly obclavate, pale to medium brown, smooth to minutely roughened, to conspicuously verrucose, commonly beakless with a conoid apical cell when juvenile, have a filiform beak at maturity, the point of transition from spore body to beak is definite and conspicuous; conidium body 25-90×10-25 μm, with 4-8 transverse septa and 1 to 2 longitudinal or oblique septa in 3-7 of the transverse divisions, giving a surface view of 10-20 bubble-like compartments; the beak, when present, long and flexuous, hyaline or pale brown, sparsely septate, simple and not branched, 100-250 μm long and 2-3 μm wide along most of its length; on agar media erostrate (beakless) conidia abundant, beaks often shorter (50-150 μm) than 56 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 35. Alternaria helianthinficiens. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

those in nature.

DISTRIBUTION: Canada, Hungary, Korea, USA. HOST/SUBSTRATE AND RECORDS IN KOREA: Helianthus annuus L. (Asteraceae) (Cho et al., 1999; Cho and Yu, 2000b; Yu, 2001). SPECIMENS EXAMINED: On Helianthus annuus, CNU-MH99241 (Namwon, September 1999); on Rud- beckia bicolor Nutt., CNU111292 (Chungju, July 2011). REMARKS: This species was first recorded and described in Korea by Cho et al. (1999). The species can be distinguished from other fungi (Alternaria protenta and Alternariaster helianthi) described on Helinthus and Rudbeckia by its ovoid to ellipsoid conidia with filamentous and non-branched beaks. The Korean materials of the present author were in good agreement with the previous Pleosporales: Pleosporaceae: Alternaria 57

Fig. 36. Alternaria helianthinficiens. Conidia from culture. Scale bar=50 μm.

records of A. helianthinficiens (Simmons, 1986b, 1997). Causing brown spots on leaves of sunflower and Rudbeckia. The lesions of sunflower and Rudbeckia may be caused by A. helianthinficiens alone or in association with Alternariaster helianthi (Hansford) E.G. Simmons.

19. Alternaria iridicola (Ellis & Everh.) J.A. Elliot, Amer. J. Bot 4(8): 439, 1917 (Figs. 37, 38).

SYNONYM: Macrosporium iridicola Ellis & Everh., Pro. Acad. Nat. Sci. Philadelphia 46: 382, 1894.

COLONIES on PDA, cottony, white to pale gray, no pigment is secreted into the medium, slowly growing, reaching about 35-50 mm in diam. after 7 days. CONIDIOPHORES (Primary conidiophores) solitary or arising in groups of 2-5, simple, straight or flexuous, more or less cylindrical, septate, pale yellow to pale brown, almost colourless at the tip, smooth, usually with a single conidiogenous site at the apex, sometimes elongating sympodially and geniculate with 1 to several pigmented conidial scars, slightly swollen at the apex, up to 120 μm long, 5-8 μm wide; in culture the conidiophores are produced at lateral extensions of the hyphae. CONIDIA solitary (Plate 16) or in chains of 2-3 through the production of secondary conidiophores, on agar media chains of 2-4 occurring more frequently than in nature; straight or slightly curved, obclavate or long ovoid, sometimes cylindrical, pale olive or pale brown, smooth-walled, slightly or sharply constricted at the transverse septa, many are beakless (erostrate) with a conoid apical cell when juvenile, or have a beak at maturity, some others (often in culture) have an apical secondary conidiophore, analog- 58 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 37. Alternaria iridicola. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

ous to the beak; conidium body 30-95×15-35 μm, with 3-10 transverse septa and 1 to 2 longitudinal or oblique septa in a few of the transverse divisions, slightly constricted at 1-3 of the transverse septa in a few of the transverse segments; beaks almost cylindrical, sturdy, simple and unbranched, shorter or the same length as spore bodies, sparsely septate, lighter in colour than the body, 10-90 μm long, 3-5 μm wide; secondary conidiophores simple or geniculate, 4-8 μm wide, as short as 10 μm or as long as or longer than the spore body.

DISTRIBUTION: Japan, Korea, USA. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Belamcanda chinensis (L.) DC. (Iridaceae) (Yu et al., 2001; Yu, 2001); Gladiolus gandabensis Van Houtte (Iridaceae) (Yu, 2001). SPECIMENS EXAMINED: On Belamcanda chinensis, CNU-MH00421 (Daejeon, September 2000), Pleosporales: Pleosporaceae: Alternaria 59

Fig. 38. Alternaria iridicola. A. conidiophores from host; B. conidia from host. Scale bars=50 μm.

CNU111205 (Gongju, November 2011); on Gladiolus gandabensis, CNU-MH00222 (Namwon, August 2000). REMARKS: The fungus was first recorded in Korea from blackberry lily and gladiolus by Yu (2001) and Yu et al. (2001). This species is characterized by having obclavate to long ovoid, and long and sturdy beaked or apically pseudorostrate conidia produced singly or in short chains of 2-4. A. iridicola causes brown to dark brown leaf spot and blight of blackberry-lily and gladiolus.

20. Alternaria japonica Yoshii, J. Plant Protect. (Tokyo) 28: 17, 1941 (Figs. 39, 40).

SYNONYMS: Alternaria raphani J.W. Groves & Skolko, Can. J. Res. 22(5): 227, 1944. Alternaria matthiolae Neergaard, Danish Species of Alternaria and Stemphylium: 184, 1945.

COLONIES on PDA, cottony, pale brown or greenish gray with scanty sporulation, reaching about 40-60 mm in diam. after 7 days. CONIDIOPHORES (Primary conidiophores) mostly solitary, unbranched to occasionally branched; straight or curved, pale to medium brown, smooth, 1-5 septate, usually with a single terminal conidiogenous site, sometimes geniculate with 1-2 pigmented conidial scars, slightly swollen at the apex, up to 100 μm long, 4-7 μm wide; in culture the conidiophores produced as lateral extension of main hyphae, not proliferating sympodially, more or less cylindrical, with a pigmented conidiogenous locus. CONIDIA mostly solitary or in short chains of 2-3 through the agency of secondary conidiophores on host, in culture chains of 3-4 are common (Plate 17), sometimes branched; broadly ovoid, ellipsoid or obclavate, tapering towards 60 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 39. Alternaria japonica. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

the apex, light brown to dark olivaceous brown, some conidium cells becoming swollen and irregular due to bulging of individual cells, erostrate (beakless) but most often producing secondary conidiophores from apical or lateral cells of the conidium, smooth when juvenile becoming roughened with age; conidium body 50-100×20-40 μm on host, smaller in culture 30-80×18-25 μm, with 2-9 transverse septa and 1 to 2 longitudinal or oblique septa in a few to most of the transverse divisions, strongly constricted at the transverse septa; second conidiophores (false beaks or psudorostra) variable in length, as short as a few μm or up to 70 μm long and 5-6 μm wide, pale to light yellow brown. CHLAMYDOSPORES formed both in culture and on host, yellow brown to dark olivaceous brown, initially may be one-celled, intercalary on a hyphae, becoming multicelled and irregular in shape with thickened wall, often strongly ornamented, with secondary conidiophores often developing from them. Pleosporales: Pleosporaceae: Alternaria 61

Fig. 40. Alternaria japonica. A. conidia from culture; B. scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=10 μm.

DISTRIBUTION: Canada, Egypt, Europe (Denmark, the Netherlands and Greece), India, Japan, Korea, USA. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Brassica campestris subsp. napus var. nippo-oleifera Makino (Brassicaceae) (Yu, 2001); Brassica campestris subsp. napus var. pekinensis Makino (Brassi- caceae) (Lee, 1983; Yu, 2001); Raphanus sativus var. hortensis for. acanthiformis Makino (Brassicaceae) (Park, 1967; Yu et al., 1991). SPECIMENS EXAMINED: On Brassica campestris subsp. napus var. nippo-oleifera, CNU-MH00803 (Dae- jeon, April 2000); from seeds of Brassica campestris subsp. napus var. pekinensis, CNU-MH99242 (Daejeon, September 1999); on Raphanus sativus var. hortensis for. acanthiformis, CNU-MH98040 (Daegwalryoung, September 1998), CNU-MH98068 (Boryeong, May 1998), CNU138096 (Gongju, November 2006). REMARKS: Alternaria japonica can be readily differentiated from two described species of Alternaria on Brassicaceae hosts by its beakless conidia produced solitary or in short chains of 2-4 through the agency of paler, secondary conidiophores. The presence of intercalary mycelial chlamydospores, occurring as distinctive chains or knots of dark, thick-walled, often strongly ornamented cells, is another important distinguishing character of A. japonica (M. and M.E. Corlett, 1999; Yu, 2001). In the early 1940s, researchers in Canada, Denmark and Japan, independently published three epithets (japonica, raphani and matthiola) for the same fungus occurring on various Brassi- caceae. A. japonica was described by Yoshii in 1941 on the basis of the fungus pathogenic to Rapha- nus sativus, Brassica campestris, Brassica oleracea and other related species for which he had already reported under the name of A. brassicae var. macrospora in 1931. Groves and Skolko established A. raphani Groves and Skolko based on Alternaria species on seeds and pods of R. sativus collected 62 Fungal Flora of Korea·Alternaria and Allied Genera from North America in 1944. A. matthiolae was introduced by Neergaard in 1945 based on Alter- naria species on Matthiola. A. raphani and A. matthiolae are now considered synonymous with A. japonica. On leaves of radish and Brassica spp., causing circular black spots; also present on seeds.

21. Alternaria longipes (Ellis & Everh.) E.W. Mason, Mycol. Pap. 2: 19, 1928 (Figs. 41, 42).

SYNONYM: Macrosporium longipes Ellis & Everh., J. Mycol. 7(2): 134, 1892.

COLONIES on PDA, gray or olivaceous gray, cottony, stunted growth, reaching about 30-40 mm in diam. after 14 days, often a purple or reddish brown pigment is secreted into the medium [Plate 18(A)]. CONIDIOPHORES (Primary conidiophores) solitary or in groups of 2-5, commonly unbranched or loosely branched, straight or flexuous, pale to dark brown, smooth, septate, usually with a single pigmented terminal conidiogenous site, sometimes geniculate with 1 to several pigmented conidiogenous sites, up to 100 μm long, 2-5 μm wide; in culture the conidiophores arising terminally or laterally on the hyphae or directly from the surface. CONIDIA mostly in short or moderately long chains of 3-8 through the agency of secondary conidiophores, sometimes solitary, on agar media long chains of 7-10 occurring frequently [Plate 18(B)], mostly unbranched, sometimes branched in culture; pale to light olive brown, narrowly ovoid, ellipsoid or obclavate, surface usually smooth or verruculose, mostly beakless or having a secondary conidiophore (pseudorostrate), occasionally having a relatively short tapered beak, in culture erostrate or pseudorostrate only; coni-

A B

Fig. 41. Alternaria longipes. A. conidia from host; B. scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=10 μm. Pleosporales: Pleosporaceae: Alternaria 63

Fig. 42. Alternaria longipes. Conidia, conidiophores and sporulation habit from host (A) and from culture (B). Scale bars=50 μm.

dium body 15-55×8-17 μm on host, 20-80×8-20 μm on agar, with 2-7 transverse septa, usually no longitudinal septum or only 1 (rarely 2) longitudinal septum in a few of the transverse divisions; secondary conidiophores arising from the apical cell of the conidium body, usually cylindrical, short and 1-celled, often elongated, ca. 10-30 (-60) μm long, 2-4 μm wide.

DISTRIBUTION: Almost world-wide, wherever tobacco is grown. HOST/SUBSTRATE AND RECORDS IN KOREA: Nicotiana tabacum L. (Solanaceae) (Nakada and Taki- moto, 1928; Park, 1967; Yu, 2001). SPECIMENS EXAMINED: On Nicotiana tabacum, CNU-MH98065 (Daegwalryoug, September 1998), CNU-MH00272 (Hongcheon, August 2000), CNU131050 (Daejeon, August 2013). REMARKS: Alternaria longipes is sometimes referred to as A. alternata (Lucas, 1971) or A. alternata tobacco pathotype (Nishimura et al., 1978, 1979). The species, however, can be differentiated from 64 Fungal Flora of Korea·Alternaria and Allied Genera

A. alternata in that the latter species has smaller beakless conidia (15-35×8-16 μm) and produces a three dimensional cluster of branching conidial chains under specific growth conditions. The fungus also somewhat resembles A. gaisen and A. tenuissima. However, the characters of slow growth on agar media and typical catenulation pattern differentiate it from the above mentioned species. On leaves of tobacco, causing brown spots often surrounded by a yellow halo.

22. Alternaria macrospora Zimmermann, Ber. Land-Forstw. Deutsch-Ostafrika 2: 24, t. 2: 13, 1904 (Figs. 43, 44).

SYNONYMS: Macrosporium macrosporum (Zimm.) Nishikado & Oshima, Nogaku Kenkyu [Agric. Res. (Kurashiki)] 36: 391, 1944. Sporidesmium longipedicellatum Reichert, Bot. Jahrb. Syst. 56: 723, 1921.

COLONIES on PDA, woolly, pale gray to dark gray, abundant aerial mycelium with profuse sporulation in a 12/12 hr NUV light/darkness cycle, usually a yellow or reddish yellow pigment produced into the medium, some cells of the older hyphae may swell, forming structures resembling a string of beads, 55-65 mm in diam. after 7 days. CONIDIOPHORES (Primary conidiophores) solitary or in small groups, erect, unbranched, straight or curved, pale to mid brown, smooth, usually

Fig. 43. Alternaria macrospora. A. conidium from culture; B. scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=15 μm. Pleosporales: Pleosporaceae: Alternaria 65

Fig. 44. Alternaria macrospora. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

only one pigmented conidiogenous site at the apex, occasionally proliferating sympodially and geniculate with 1-3 pigmented conidiogenous sites, septate, up to 90 μm long, 5-9 μm wide, in culture conidiophores produced terminally or laterally on the hyphae. CONIDIA mostly solitary or uncommonly in chains of 2 (Plate 19); obclavate to broadly obclavate, or broadly ellipsoid, straight or slightly curved, mid to dark brown, verruculose when juvenile, becoming smooth at maturity, rostrate, tapering rather abruptly towards a narrow beak; conidium body 35-80×20-30 μm on host, with 4-9 transverse septa and 1 to several longitudinal or oblique septa in 1 to 5 of the transverse divisions, often slightly constricted at the transverse septa, in culture the conidium body often slightly larger than those in nature 40-90×20-30 μm, considerably more longitudinal or oblique septa in the conidium bodies than in those in nature; the beak long, filiform, pale, septate, not branched, about the same length as or up to twice or three times as long as the conidium body, 66 Fungal Flora of Korea·Alternaria and Allied Genera

80-280 μm long, 2 μm wide along most of its length, on agar media the beaks often shorter (up to 80 μm long) than those in nature.

DISTRIBUTION: All Africa; Asia (China, India, Japan, Korea, Parkistan, Sri Lanka); Australasia and Oceania (Australia); Europe (France, Italy, Romania); North America (USA); Central and South America (Brazil, Nicaragua, Venezuela). HOST/SUBSTRATE AND RECORDS IN KOREA: Gossypium indicum Lam. (Malvaceae) (Park, 1961; Yu, 2001). SPECIMENS EXAMINED: On Gossypium indicum, CNU-MH00275 (Mokpo, September 2000), CNU003411, 003412 (Mokpo, August 2006). REMARKS: Alternaria macrospora differs from A. gossypina reported on Gossypium by its larger conidia and long beak. In Korea this species was first recorded with phytopathological notes by Park (1961), and Yu (2001) described its morphological features in detail. The fungus can cause leaf spot and boll rot of cotton. Causing circular, brown spots with a narrow purple border on leaves; circular lesions are formed on buds and bolls.

23. Alternaria mali Roberts, J. Agric. Res. 2: 58, 1914 (Figs. 45, 46).

COLONIES on PDA, olive gray or dark olive gray, cottony or velvety, with varying amounts of non-sporulating aerial and surface mycelium, 60-70 mm in diam. after 7 days [Plate 20(A)]. CONIDIOPHORES (Primary conidiophores) solitary or in small groups, mostly unbranched,

Fig. 45. Alternaria mali. A. conidia from host; B scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=10 μm. Pleosporales: Pleosporaceae: Alternaria 67

Fig. 46. Alternaria mali. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

straight to slightly curved, mid to dark brown, smooth, septate, usually with a single pigmented conidiogenous site at the apex, occasionally geniculate with 1-2 pigmented conidial scars, 20-90 μm long, 2-4 μm wide; in culture the conidiophores arising directly from the surface or terminally and laterally on the hyphae. CONIDIA mostly in short or moderately long chains of 3-8 [Plate 20(B)], sometimes in long chains of 10 or more on agar, unbranched or branched; obclavate to narrow ovoid or long ellipsoid on host, short to long ovoid or broad ellipsoid on agar, pale olivaceous brown, surface usually verruculose, occasionally smooth, mostly beakless (erostrate) or have an apical secondary conidiophore (pseudorostrate), occasionally rostrate with a relatively short beak (in nature); conidium body 18-55×5-12 μm in nature, 15-65×8-20 μm in culture, with 3-8 transverse septa and usually no longitudinal septum or only 1 longitudinal septum in 1-3 of the trans- 68 Fungal Flora of Korea·Alternaria and Allied Genera verse compartments, often constricted at the 1-3 of the transverse septa; the beak, when present, gradually tapering, as long as or shorter than the body; secondary conidiophores usually short and 1-celled ca. 3-5×3-5 μm , sometimes elongated up to 30×2-4 μm.

DISTRIBUTION: Asia (China, India, Japan, Korea); North America (USA); South America (Chile); Oceania (Australia). HOST/SUBSTRATE AND RECORDS IN KOREA: Malus pumila var. dulcissima Koidz. (Rosaceae) (Nakada and Takimoto, 1928; Lee and Lee, 1972; Ryu et al., 1993; Yu, 2001). SPECIMENS EXAMINED: On Malus pumila var. dulcissima, CNU-MH99243 (Suwon, June 1999), CNU- AM28 (Daejeon, August 2010). REMARKS: Nishimura et al. (1978, 1979) has recommended that A. mali should be treated as “A. alternata apple pathotype” because of the similarity of the conidial morphology. Simmons (1992, 1999b), however, opposed this naming system (pathotype system) based on the differences in sporulation patterns. (see remarks under A. alternata). Alternaria mali was initially described on Alter- naria isolates from apple fruit by Roberts (1914). Details of conidium characters of the fungus were reported in his second publication (Roberts, 1924). According to Simmons (1999b), A. mali is a fungus morphologically different from the one that figures prominently in HST-related literature (A. alternata apple pathotype, Tottori Univ. 0-159). Causing circular, brown to black brown spots with a brownish purple margin on leaves of apples.

24. Alternaria nobilis (Vize) E.G. Simmons, Mycotaxon 82: 7, 2002 (Figs. 47, 48).

SYNONYMS: Macrosporium nobilis Vize, in M.C. Cooke, Grevillea 5: 119, 1877. Alternaria dianthi Stevens & Hall, Bot. Gaz., 47: 409, 1909. Macrosporium dianthi (St. & Hall) Bewley, Diseases of Glasshouse Plants, p. 208, 1923.

COLONIES on PDA, densely cottony, olive gray to smoky gray with varying amount of aerial mycelium, sporulation abundant (on V-8 agar) held in a 12/12 hr NUV light/dark cycle, no pigment is released into the medium, 30-35 mm in diam after 7 days. MYCELIUM usually immersed or partly superficial; hyphae subhyaline to olivaceous brown, branched, septate, smooth-walled, 4-8 μm wide. CONIDIOPHORES (Primary conidiophores) solitary or arising in groups of 2-10, usually unbranched, straight or curved, more or less cylindrical, pale brown to mid-olivaceous brown, smooth, with a single pigmentred conidiogenous site at the apex, sometimes proliferating sympodially and geniculate with 1-3 pigmented conidiogenous sites, septate, uo to 50 μm long, 5- 8 μm wide; in culture conidiophores are produced as side branches on the main hyphae, up to 130 μm long. CONIDIA usually in chains of 3-5 through production of secondary conidiophores, conical to obclavate, tapering towards the apex, broadly ovoid or ellipsoid in culture, brown to dark-olivaceous brown, smooth-walled, many are beakless (erostrate) or have a secondary conidiophore (false beak or pseudorostrum) from apical conidium cell, some others have a relatively short beak; 20-80×10-25 μm, with 3-10 transverse septa and one to several longitudinal or oblique septa in a few of the transverse segments, often constricted (strongly constricted in culture) at the transverse septa; beaks when present subhyaline and ca. 20-30×2-3 μm or usually shorter, occa- Pleosporales: Pleosporaceae: Alternaria 69 sionally the tip of the beak converted into a slightly enlarged functional conidiophore; secondary conidiophores relatively broad, paler or the same colour as the conidium body, mostly simple, sometimes geniculate (particularly in culture), variable in length, as short as a few μm or up to 60 μm long, 5-7 μm wide.

DISTRIBUTION: Nearly worldwide, wherever Dianthus spp. are cultivated, including Korea. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Dianthus barbatus var. japonicus Nakai (Caryophyllaceae) (Yu, 2001, under A. dianthi); Dianthus caryophyllus L. (Caryophyllaceae) (Yu et al., 1989, under A. dianthi; Lee et al., 1989, under A. dianthi; Yu, 2001, under A. dianthi); Dianthus littorsus Makino (Caryophyllaceae) (Yu, 2001, under A. dianthi); Dianthus sinensis L. (Caryophyllaceae) (Yu, 2001, under A. dianthi); Dianthus superbus var. longicalycinus (Max.) Williams (Caryophyllaceae) (Yu, 2001,

Fig. 47. Alternaria nobilis. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm. 70 Fungal Flora of Korea·Alternaria and Allied Genera

AB C

Fig. 48. Alternaria nobilis. A. conidiophore and conidia from culture; B, C. scanning electron microscopes of conidiophores and conidia from host. Scale bars: A, B=50 μm; C=25 μm.

under A. dianthi). SPECIMENS EXAMINED: On Dianthus barbatus var. japonicas, CNU-MH00761 (Daejeon, June 2000); on Dianthus caryophyllus, CNU-MH98009 (Gimhae, May 1998); on Dianthus sinensis, CNU-MH00756 (Namwon, August 2000), CNU111158 (Daejeon, September 20011); on Dianthus superbus var. longicalycinus, CNU-MH00200 (Uiseong, July 2000). REMARKS: The fungus was first reported as A. dianthi in Korea (Yu et al., 1989). The species name was changed to nobilis based on the principle of nomenclatural priority (Simmons, 2002). Simmons (2002) clearly explained why the epithet nobilis should be used instead of the commonly accepted one, dianthi (David, 1988; Yu, 2001). A. nobilis is referred to the section Brevicatenatae of Neergaard (1945) and overall is most similar to A. japonica Yoshii. However, conidia of A. nobilis are more reg- ular and conical, whereas conidia of A. japonica are irregular and oval and A. nobilis does not produce chlamydospores (vs. chlamydospore production in A. japonica). The present species is also closely related to A. dianthicola (see remarks under A. dianthicola). On Dianthus species, including carnation, causing leaf blight and stem rot. On the leaves small purple spots are produced at first and these soon develop a broad yellowish-green border.

25. Alternaria panax Whetzel, Bull. U. S. Dept. Agric. 250: 11, 1912 (Figs. 49, 50).

SYNONYMS: Alternaria panacis Whetzel, in Saccardo, Sylloge Fungorum 25. Abellini: 1093, 1931. Alternaria araliae Greene, Trans. Wis. Acad. Sci., Arts and Lett. 42: 69, 1953.

COLONIES on PDA, abundant aerial mycelium, white to greenish gray, cottony, no production any pigment into the medium, 55-65 mm in diam. after 7 days [Plate 21(A)], sporulation abundant held in a 12/12 hr NUV light/dark cycle, no sporulation in darkness. CONIDIOPHORES (Primary Pleosporales: Pleosporaceae: Alternaria 71

Fig. 49. Alternaria panax. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

conidiophores) solitary or arise in small groups, straight or slightly curved, simple, usually only one pigmented terminal conidiogenous site, sometimes geniculate with 1-2 pigmented conidial scars, slightly swollen at tha apex, pale-brown, thin-walled, smooth, septate, up to 150 μm long, 4- 10 μm wide; in culture the conidiophores produced terminally or laterally on the hyphae, erect, simple sometimes geniculate, up to 200 μm long. CONIDIA mostly solitary [Plate 21(B)] or in small chains of 2-4 on host, in culture chains of 3-6 occurring frequently with age [Plate 21(C)]; straight or slightly curved, long elliptical, obclavate to broadly obclavate, pale brown to dark brown, smooth when juvenile, becoming verruculous to verrucose, strongly constricted at the transverse septa at maturity, excessive cellular swelling occurs in culture, thus causing the conidium to appear distorted, some have a beak (rostrate), gradually tapering towards the beak, some others have an apical or lateral secondary conidiophore (pseudorostrate); 35-125×10-25 (-30) μm in size 72 Fungal Flora of Korea·Alternaria and Allied Genera

A B

Fig. 50. Alternaria panax. A. conidia from host; B. scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=10 μm.

with conidial body of 35-90 (-100) μm in length, with 3-11 transverse septa and 1 to several longitudinal or oblique septa in 1 to 6 of the transverse segments, highly variable and hypertrophy becomes common in culture; beaks, when present, long, rigid, blunt-tipped, simple and unbranched, septate, concolorous with the conidium body or slightly lighter in colour than the body, variable in length, up to 60 μm long, 3-7 μm wide; secondary conidiophores (false beaks or pseudorostra) simple or geniculate, 4-9 μm wide, as short as 10 μm or as long as the spore body.

DISTRIBUTION: Australia, Canada, China, Japan, Korea, New Zealand, Russia, Spain, U.K., USA, Venezuela. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Panax ginseng C.A. Meyer (Araliaceae) (Nakada and Takimoto, 1928; Yu et al., 1984; Lee et al., 1990; Yu, 2001; Deng et al., 2015); Panax quinqueforium L. (Araliaceae) (Deng et al., 2015). SPECIMENS EXAMINED: On Panax ginseng, CNU-MH00625 (Geumsan, August 2000), CNU085019 (Suwon, August 2008), CNU101009 (Eumsung, September 2010); on Panax quinquefolium, CNU085011, 086005 (Suwon, July 2008). REMARKS: Alternaria panax is characterized by its swollen, distorted conidia with a long, rigid beak or a false beak (in culture). The species can be differentiated from A. araliae based on morphology, molecular characteristics and pathogenicity (Deng, 2012; Yu et al., 2012; Deng et al., 2015). The conidia of A. panax are slightly smaller (overall size, 35-125×10-30 μm) than those of A. araliae (overall size, 40-160×12-50 μm). A. panax causes leaf blight of ginseng (Panax). Symptoms of the disease are brown to dark brown, water-soaked spots with a yellow or chlorotic halo on leaves of ginseng, and dark brown, long- elliptical necrotic lesions on stems and petioles. Pleosporales: Pleosporaceae: Alternaria 73

26. Alternaria peucedani S.H. Yu, Mycobiol. 42: 12, 2014 (Figs. 51, 52).

COLONIES on PDA, effuse, cottony, obverse olivaceous buff to olive, reverse olivaceous buff to dark olive, with varing amount of aerial mycelium, poor sporulation (scarifying the surface mycelium induces increased sporulation), no pigment is diffused into the medium, 30-40 mm in diam. after 7 days. MYCELIUM immersed or superficial; hyphae, septate, hyaline to pale brown, branched, smooth-walled or minutely roughened in some areas, 3-10 μm wide. CONIDIOPHORES (Primary conidiophores) arising singly or in groups of 2-5, simple or branched, straight or slightly curved, more or less cylindrical, pale gray or olivaceous gray, smooth, septate, commonly only one pigmented terminal conidiogenous site, rarely proliferating sympodially and geniculate with 1-2 pigmented conidial scars, often slightly swollen at the apex, 15-120 μm long, 5-10 μm wide; in culture the conidiophores are produced as lateral extension of main hyphae, with a pigmented conidiogen-

Fig. 51. Alternaria peucedani. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scalr bar=50 μm. 74 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 52. Alternaria peucedani. A. conidiophores from host; B conidia from host. Scale bars=50 μm.

ous locus. CONIDIA solitary on host (Plate 22), in culture chains of 2 occurring very rarely with age, straight or slightly curved, obclavate to obpyriform, olivaceous brown to dark brown, smooth to verrculose, with age strongly constricted at the transverse septa, excessive cellular swelling occurs in culture, rostrate, gradually tapering towards a sturdy beak; conidium body 50-125×20-40 (-50) μm, with 5-12 transverse septa and 1 to several longitudinal or oblique septa in 1 to 6 of the transverse segments; the beak almost cylindrical, rigid, simple and unbranched, never fragile or filamentous, pale brown, sometime slightly swollen at the apex, on host 20-80 (-120) μm long, 4-6 (-8) μm wide, in culture beaks usually shorter (15-50 μm) than those in nature. CHLAM- YDOSPORES sometime formed in culture, pale brown to light yellow brown, intercalary on hyphae, 12-15 μm wide.

DISTRIBUTION: Korea. HOST/SUBSTRATE AND RECORDS IN KOREA: Peucedanum japonicum Thunb. (Apiaceae) (Lee et al., 1991 under Alternaria sp.; Yu, 2001 under Alternaria sp.; Deng et al., 2014). SPECIMENS EXAMINED: On Peucedanum japonicum, CNU3010 (Jinan, September 1999), CNU111485, 118048 (Yesan, August 2011). REMARKS: This fungus was first recorded as Alternaria sp. by Lee et al. in Korea (1991). Later, Yu (2001) described morphological characteristics of the fungus and considered it a new Alternaria taxon without prior naming. Deng et al. (2014) have assigned the fungus a new name A. peucedani based on morphological characteristics and phylogenetic analysis of ITS region, gpd and Alt a1 gene sequences. A. peucedani somewhat resembles A. brassicae and A. panax in having a large, Pleosporales: Pleosporaceae: Alternaria 75 obclavate to obpyriform spore body with a cylindrical, simple, sturdy and nonfragile beak. How- ever, the conidium body of the fungus is wider than A. brassicae (15-30 μm) and much longer than A. panax (35-90 μm). The fungus also differs from A. brassicae and A. panax in conidial catenulation. The species forms a single conidium per conidiophore on natural substrata. However, the two species readily form short chains of 2-3 conidia on hosts and in culture. On leaves of Peucedanum, causing round or irregular, dark brown to black spots.

27. Alternaria porri (Ellis) Ciferri, J. Dept. Agric. Porto Rico, 14(1): 30, 1930 (Figs. 53, 54).

SYNONYM: Macrosporium porri Ellis, Grevillea, 8(45): 12, 1879.

COLONIES on PDA, cottony to loose felt-like, olive gray to grayish green, abundant aerial mycelium, usually secreting a reddish-brown or yellow pigment into the medium, 55-65 mm in diam. after 7 days. CONIDIOPHORES solitary or in fascicles of 2-5, usually unbranched, straight or curved, stout and rigid, usually only one terminal conidiogenous site, sometimes proliferating sympodially and geniculate with 1-3 pigmented conidial scars, several septate, pale brown to mid brown, smooth, often slightly swollen at the apex, up to 150 μm long, 5-10 μm wide; in culture the conidiophores arise terminally and laterally on the hypha. CONIDIA mostly solitary (Plate 23), occasionally (uncommonly) in chains of 2 through the agency of secondary conidiophores; obclavate or long ellipsoid, straight or curved, pale brown to mid or dark brown, smooth to minutely

A B

Fig. 53. Alternaria porri. A. conidia from culture; B. scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=5 μm. 76 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 54. Alternaria porri. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

verruculose, rostrate, gradually tapering towards the beak; conidium body 50-100 (-120)×15-28 μm, with 5-12 transverse septa and 1 to 2 longitudinal or oblique septa in a few of the transverse segments, often constricted at the transverse septa; the beak filiform, colourless or pale, septate, occasionally (uncommonly) once branched (bifurcate), commonly about the same length as the body but may be shorter or longer, 50-150 μm long on host, 50-100 μm long on agar media, 4-5 μm wide at the base, tapering to 2 μm; occasionally a sturdy secondary conidiophore arises at the api-cal cell or intercalary cell (uncommon) of the conidium, initiating a secondary conidium.

DISTRIBUTION: Africa (Egypt, Ethiopia, Ghana, Jamaica, Kenya, Malawi, Mauritius, Nigeria, Rho- desia, Tanzania, Uganda, Zambia); Asia (Hong Kong, India, Iraq, Israel, Japan, Korea, Pakistan, Philippines, Taiwan, Thailand); Australia and Oceania (Australia, New Zealand); Europe (Austria, Pleosporales: Pleosporaceae: Alternaria 77

Denmark, Germany, Netherlands, Poland, Portugal, Romania, Russia); Central and South America (Argentina, Brazil, Cuba, El Salvador, Venezuela); North America(Canada, USA). HOSTS/SUBSTRATES AND RECORDS IN KOREA: Allium cepa L. (Liliaceae) (Cho et al., 1997; Yu, 2001); Allium fistulosum L. (Liliaceae) (Park, 1958; Cho et al., 1997; Yu, 2001); Allium porrum L. (Liliaceae) (Chung et al., 1977); Allium sativum for pekinense Makono (Liliaceae) (Yu, 2001); Allium schoenoprasmum var. orientale Regel (Liliaceae) (Yu, 2001); Allium senescens L. (Liliaceae) (Yu, 2001); Allium tuberosum Roth. (Liliaceae) (Chang et al., 2000). SPECIMENS EXAMINED: On Allium cepa, CNU3564 (Buyeo, May 2003), CNU103012, 103013 (Gongju, June 2010); on Allium fistulosum, CNU3480 (Geumsan, June 2003), CNU094022 (O-chang, May 2009), CNU 094026 (Gongju, June 2009); on A. sativum for pekinense, CNU-MH98087 (Pyeongchang, May 1998). REMARKS: The name Alternari porri has been used widely for decades as an uncritical identification for any large-spored Alternaria found on species of Allium (Ellis, 1971). However, Simmons (2007) differentiated Alternaria isolates from Allium spp. hosts/substrates into several taxa of Alter- naria including A. porri and A. vanuatuensis based on spore morphology and sporulation pattern. The length of conidial beaks of A. porri is commonly longer [50-150 (av. 91) μm on host tissue] than that of A. vanuatuensis [50-90 (av. 58) μm on host tissue] (Li et al., 2014). This fungus can cause purple blotch of garlic, onion and spring onion.

28. Alternaria protenta E.G. Simmons, Mycotaxon 25(1): 207, 1986 (Figs. 55, 56).

COLONIES on PDA, cottony, white to pale gray, reverse brown to reddish brown, no pigment is released into the medium, reaching 60-70 mm in diam. after 7 days. CONIDIOPHORES mostly

Fig. 55. Alternaria protenta. A, B. conidia from host. Scale bar=50 μm. 78 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 56. Alternaria protenta. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

solitary, or in groups, simple or occassinally branched, straight or slightly flexuous, more or less cylindrical, usually only a single terminal conidiogenous site, sometimes elongating sympodially and geniculate with 1-2 pigmented conidial scars, septate, pale gray or pale brown, smooth, often slightly swollen at the apex, 50-130 μm long, 3-7 μm wide; in culture the conidiophores produced terminally or laterally on the hyphae, erect, simple or branched, 100-200 μm long, 4-8 μm wide. CONIDIA mostly solitary (Plate 24), occassionally in chains of 2 through the agency of secondary conidiophores; obclavate, straight to slightly curved, light brown to medium brown, smooth or minutely roughened, sometime verruculose in culture, beakless and apically blunty conical when juvenile, many have a long beak at maturity, the transition from spore body to beak is gradually tapering; conidium body 40-120×12-20 μm on host, 50-140×17-25 μm on agar media, with 5-14 transverse septa and several longitudinal or oblique septa; the beak filiform, colourless or pale, Pleosporales: Pleosporaceae: Alternaria 79 septate, often once branched (bifurcate), 80-150 μm long, 3-5 μm wide at the base, 2 μm at the tip.

DISTRIBUTION: Israel, Korea, Uganda. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Helianthus annuus L. (Asteraceae) (Cho et al., 1999; Cho and Yu, 2000b; Yu, 2001); Rudbeckia bicolor Nutt. (Asteraceae) (Yu, 2001). SPECIMENS EXAMINED: On Helianthus annuus, CNU-MH98027 (Cheongyang, September 1998), CNU-MH98062 (Daegwalryoung, September 1998), CNU080336 (Cheongyang, August 2000); on Rudbeckia bicolor, CNU-MH00336 (Pyeongchang, August 2000), CNU-MH00337 (Namwon, August 2000). REMARKS: Alternaria protenta differs from Alternaria helianthinficiens and Alternariaster helianthi described on species of Helianthus and Rudbeckia by its longer conidia with long and often once branched beaks (see remarks under A. helianthinficiens). A. protenta causes leaf spot of sunflower and black-eyed Susan (Rudbeckia).

29. Alternaria radicina Meier, Drechsler & E.D. Eddy, Phytopathol. 12: 164, 1922 (Figs. 57, 58).

SYNONYMS: Stemphylium radicinum (Meier, Drechsler & E.D. Eddy) Neerg., Yb. R. Vet. Agric. Coll., 1937. Thyrospora radicina (Meier, Drechsler & E.D. Eddy) Neerg., Botan. Tidsskr. 44(3): 361. 1938. Pseudostemphylium radicinum (Meier, Drechsler & E.D. Eddy) Subram., Current Science 30(11): 423, 1961.

COLONIES on PDA, effuse, dark brown to blackish brown, with varying amounts of aerial

A B

Fig. 57. Alternaria radicina. A. conidia from culture; B. scanning electron microscope of conidia from culture. Scale bars: A=50 μm; B=10 μm. 80 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 58. Alternaria radicina. Conidia, conidiophores and sporulation pattern from host (A) and culture (B). Scale bars=40 μm.

mycelium, 35-50 mm in diam. after 7 days. CONIDIOPHORES (Primary conidiophores) mostly solitary, simple or branched, straight to flexuous, light brown to olivaceous brown, septate, with a single pigmented terminal conidiogenous site or geniculate with 1-3 pigmented conidiogenous sites, variable in length, up to 100 μm long, 3-8 μm wide; in culture abundant conidiophores produced as side branches or at the end of the hyphae, up to 200 μm long. CONIDIA usually solitary (Plate 25), rarely in chains of 2 through the agency of secondary conidiophores; short ellipsoid to ovoid, or broadly ellipsoid to ovoid on host, narrowly ellipsoid or obovoid on agar, olivaceous brown or mid to dark brown, surface smooth-walled, erostrate (beakless), rounded at both ends; measure 20-50×15-30 μm from host, 20-80×15-20 μm in culture, with 2-8 transverse septa and 1 to several longitudinal or oblique septa in a few or most of the transverse compartments; occasionally (uncommonly) a relatively broad and short secondary conidiophore produced from the apical conidium cell, initiating a secondary conidium. Pleosporales: Pleosporaceae: Alternaria 81

DISTRIBUTION: Widespread in Europe, Argentina, Australia, Canada, China, India, Israel, Japan, Korea, Nigeria, USA and Venezuela. HOST/SUBSTRATE AND RECORDS IN KOREA: Daucus carota var. sativa DC. (Apiaceae) (Park, 1967; Cho et al., 1997; Yu, 2001). SPECIMENS EXAMINED: From seeds of Daucus carota var. sativa, CNU-MH98073, MH98074 (Dae- gwalryoung, September 1998), CNU003809, 003876 (Pyeongchang, August 2006). REMARKS: Alternaria radicina is characterized by its erostrate conidia, which are solitary or rarely in chains of two. This fungus is closely related to erostrate-conidium species on Daucus; A. artocariis E.G. Simmons and A. carotiincultae E.G. Simmons. Simmons (1995), however, distinguished them by the characters of catenulation pattern and conidia. On carrot this fungus causes black root rot, a common storage rot. Infection frequently starts at the crown and extends downwards; also present on seeds (seed-borne).

30. Alternaria ricini (Yoshii) Hansford, Proc. Linn. Soc. Lond., 155: 53, 1943 (Figs. 59, 60).

SYNONYM: Macrosporium ricini Yoshii, Sci. Bull. Fac. Agric. Kyusu Univ. 3: 330, 1929.

Fig. 59. Alternaria ricini. Conidium from host. Scale bar=50 μm. 82 Fungal Flora of Korea·Alternaria and Allied Genera

COLONIES on PDA, cottony or woolly, pale to dark gray with varying amounts of aerial mycelium, no pigment is released into the medium, reaching about 45-55 mm in diam. after 7 days. CONIDIOPHORES (Primary conidiophores) solitary or in small groups, erect, unbranched, straight or curved, pale to mid brown, smooth, usually only one pigmented conidiogenous site at the apex, occasionally proliferating sympodially and geniculate with 1-3 pigmented conidiogenous sites, septate, up to 90 μm long, 5-9 μm wide; in culture conidiophores produced terminally and laterally on the hyphae. CONIDIA mostly solitary, occasionally in chains of 2 (Plate 26); obclavate or long ellipsoid, straight or curved, pale to mid brown, smooth-walled, occasionally verruculose, rostrate, tapering rather abruptly into a narrow beak; conidium body 40-75×15-25 μm on host, 30-70×10- 22 μm in culture, with 4-10 transverse septa and 1 (rarely 2) longitudinal or oblique septum in a few of the transverse divisions, often slightly constricted at the transverse septa; the beak long, filiform, pale, septate, unbranched, about the same length as, or up to twice as long as, the conidium

Fig. 60. Alternaria ricini. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm. Pleosporales: Pleosporaceae: Alternaria 83 body, 30-120 μm long, 1-2 μm wide along most of its length; in culture the beak is often shorter (20-70 μm long) than that in nature.

DISTRIBUTION: Widespread in tropics and temperate zone. HOST/SUBSTRATE AND RECORDS IN KOREA: Ricinus communis L. (Euphorbiaceae) (Park, 1961; Yu, 2001). SPECIMENS EXAMINED: On Ricinus communis, CNU-MH00110 (Daejeon, August 2000), CNU003349 (Cheonan, September 2006). REMARKS: Alternaria ricini is similar to A. macrospora in conidial morphology. This species, however, can be differentiated from the latter by its smaller, broadly ellipsoid conidia with fewer longitudinal or oblique septa. This species causes seedling blight, leaf spot and inflorescence and pod rots of castor bean.

31. Alternaria scirpicola (Fuckel) Sivanesan, The Bitunicate Ascomycetes. Valduz.: 526, 1984 (Figs. 61, 62).

SYNONYMS: Sporidesmium scirpicola Fuckel, Hedwigia 2(15): 133, Fung. Rhen. no 78, 1863. Claster- osporium scirpicola (Fuckel) Sacc., Syll. Fung. 4: 393, 1886. Pleospora heleocharidis Plowr. Sphaeria

AB C

Fig. 61. Alternaria scirpicola. A. conidiophores and conidia from culture; B, C. conidia from culture. Scale bars=50 μm. 84 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 62. Alternaria scirpicola. Conidia, conidiophores and sporulation habit from culture. Scale bars=50 μm.

scirpicola DC., Flore, francaise 2: 300, 1805. Pleospora eleocharidis Plowr., Fungi of Norfolk: 18, 1884. Cercospora scipicola (Fuckel) Zind.-Bakker, Rev. Mycol. 5: 64, 1940. Alternaria scirpicola (Fuckel) Lucas & Webster, Cas. Slez. Muz., Ser. A., Hist. Nat. 23: 151, 1974. Nimbya scirpicola (Fuckel) E.G. Simmons, Sydowia 41: 314, 1989.

COLONIES on PDA, effuse, dark brown, no pigment is released into the medium, 50-60 mm in diam. after 7 days. CONIDIOPHORES (Primary conidiophores) short to moderately long, solitary or in small fascicles, septate, straight or flexuous, pale brown, with a single apical conidiogenous locus, becoming closely one to several geniculate, contorted, and sometimes branched, with one conidiogenous site at each bend; the conidiogenous locus is distinctly pigmented, 20-70 μm long. CONIDIA usually solitary (on host), sometimes in short chains of 2-3 (in culture) (Plate 27), long narrow-obclavate, pale yellow-brown, darker brown in the basal pore area, rostrate producing a gradually tapering, long filiform beak, occasionally the tip of the beak becomes converted into a Pleosporales: Pleosporaceae: Alternaria 85 secondary conidiophore which bears 2-3 secondary conidia; 4-20 transverse disto- or eusepta, no longitudinal septa, sometimes slightly constricted near eusepta, up to 120-150×12-20 μm (spore body) plus a narrow apical beak to about 50-150×5 μm. A sexual morph may occur.

DISTRIBUTION: India, Korea, Japan, Europe including Great Britain. HOST/SUBSTRATE AND RECORD IN KOREA: Scirpus maritimus L. (Cyperaceae) (Yu et al., 1994 under Nimbya scirpicola). SPECIMENS EXAMINED: On Scirpus maritimus L., CNU100450 (Gimjae, August 2010), CNU125180 (Buan, September 2012). REMARKS: The fungus was first reported as Sporidesmium scirpicola by Fuckel in 1870 and the genus transferred to Cercospora by Zinderen Bekker (1940) and later to Alternaria by Lucas and Webster (1974). Simmons (1989) established a new genus, Nimbya, transferring A. scirpicola to this genus (N. scirpicola). Recently, however, an extensive study on Alternaria (Lawrence et al., 2013) found that N. scirpicola belonged to section Nimbya of the Alternaria clade based on sequence of the 18S nrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha gene regions, and they choose to retain Alternaria scirpicola (Fuckel) Sivan., which is a well established name. The fungus can be easily distinguished by striking distoseptation of the conidia. It causes severe leaf spot and blight of bulrush (Scirpus maritimus).

32. Alternaria sesami (E. Kawam.) Mohanthy & Behera, Curr. Sci. 27(12): 493, 1958 (Figs. 63, 64).

SYNONYM: Macrosporium sesami E. Kawam., Fungi, Tokyo 1(2): 27, 1931.

COLONIES on PDA, cottony or woolly, white to pale gray, abundant aerial mycelium, no pigment is released into the medium, reaching about 45-55 mm in diam. after 7 days [Plate 28(A)]; on V-8 juice agar sporulation abundant held in a 12/12 hr NUV light/dark cycle, no sporulation in darkness. CONIDIOPHORES (Primary conidiophores) solitary or in small groups, usually unbranched, straight or flexuous, almost cylindrical, septate, pale brown or yellowish brown, smooth, mostly non-geniculate with a single terminal conidiogenous site, but sometimes geniculate with 1- 3 pigmented conidial scars, up to 200 μm long, 5-9 μm wide; in culture the conidiophores are produced as lateral extensions of the hyphae. CONIDIA usually solitary [Plate 28(B)], sometimes in chains of 2 (-3), on agar media chains occurring more frequently than in nature; straight or slightly curved, obclavate to elongated ovoid, pale brown when juvenile, becoming mid to dark brown with age, smooth or inconspicuously roughened, rostrate, tapering towards a long and filiform beak, the point of transition from spore body to beak is definite and conspicuous; conidium body 40-135×13-35 μm, with 2-13 transverse septa and one to several longitudinal or oblique septa in one to several of the transverse segments, often slightly or strongly constricted at the transverse septa; the beak long, narrowly filamentous, colourless or pale, septate, mostly simple, occasionally once branched, 36-280 (-580) μm long, 2-5 μm wide, in culture the beak often shorter than comparable one in nature.

DISTRIBUTION: Africa (Kenya, Mozambique, Nigeria, Somalia, Sudan); Asia (Afganistan, China, India, Iraq, Iran, Israel, Japan, Korea, Pakistan, Turkey); Europe (Greece, Russia); Central and South 86 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 63. Alternaria sesami. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

America (Argentina, Brazil, Venezuela); North America(Canada, USA). HOST/SUBSTRATE AND RECORDS IN KOREA: Sesamum indicum L. (Pedaliaceae) (Park, 1961; Yu et al., 1981; Yu et al., 1982; Yu, 2001). SPECIMENS EXAMINED: On Sesamum indicum, CNU-MH00682 (Geumsan, August 2000), CNU104061 (Iksan, July 2010), CNU104193, CNU104110 (Naju, August 2010), CNU104115 (Chuncheon, July 2010), CNU104132 (Gapyeong, July 2010), CNU104137 (Yeoju, July 2010), CNU138011 (Hampyeong, August 2013). REMARKS: Alternaria sesami has been found on Sesamum indicum nearly all over the world. The species must be distinguished from A. simsimi E.G. Simmons, which is a pathogen of sesame. The former producing larger conidia (40-135×13-35 μm) with a very long, filamentous beak, usually solitary or rarely in chains of 2, whereas the latter produces smaller conidia (30-60×8-14 μm) in Pleosporales: Pleosporaceae: Alternaria 87

Fig. 64. Alternaria sesami. A. conidia from culture; B. scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=5 μm.

chains of 2-6. On leaves of sesame causing severe brown leaf spots and blights, also causing necrotic lesions on stems and capsules.

33. Alternaria simsimi E.G. Simmons, Stud. Mycol. 50: 111, 2004 (Figs. 65, 66).

COLONIES on PDA, amphiginous, effuse, apricot to vinaceous buff, no pigment is released into the medium, reaching about 47-55 mm in diam. after 7 days [Plate 29(A)]; sporulation abundant on both V-8 juice agar and PDA held in a 12/12 hr NUV light/dark cycle. CONIDIOPHORES (Pri- mary conidiophores) solitary or arising in groups, simple or branched, straight or slightly curved, septate, smooth-walled, pale to light brown, usually with a single terminal conidiogenous site, sometimes proliferating sympodially and geniculate with 1-3 pigmented conidial scars, often slightly swollen at the apex, up to 90 μm long, 3-7 μm wide; in culture the conidiophores are produced terminally and laterally on the hyphae. CONIDIA solitary [Plate 29(B)] or in short chains of 2-4, sometimes up to 6 (or more) in culture, chains on agar media occurring more frequently than in nature; pale to medium brown with darker eusepta; straight or slightly curved, highly variable in shape of conidium body, subcylindrical, obclavate, ovoid or narrowly ellipsoid, with or without a beak, tapering spore body to beak is gradual or abrupt; conidium body 30-70 (av. 45) μm long, 8- 15 (avg. 12) μm wide, with 2-7 transverse septa and 1-2 longitudinal septa in a few of the transverse divisions, the wall ornamentation is smooth or minutely verruculose; the beak, when pre- 88 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 65. Alternaria simsimi. Conidia, conidiophores and sporulation habit from host. Scale bars=50 μm.

sent, long or short, narrowly filamentous, septate, simple and not branched, pale to medium brown, variable in length, up to 480 (av. 210) μm long, 2-3 μm wide, secondary conidiophores occasionally produced from the apical conidium cell.

DISTRIBUTION: Argentina, India, Korea. HOST/SUBSTRATE AND RECORDS IN KOREA: Sesamum indicum L. (Pedaliaceae) (Yeh et al., 1989, under A. sesamicola; Yu et al., 1982, under A. sesamicola; Choi et al., 2014). SPECIMENS EXAMINED: On Sesamum indicum, CNU104013 (from seed, May 2010), CNU 104055 (Iksan, July 2010), CNU104082 (Uljin, July 2010), CNU104095 (Pohang, July 2010), CNU104101 (Hampyeong, August 2010), CNU104110 (Andong, July 2010), CNU138020 (Hampyeong, Sep- Pleosporales: Pleosporaceae: Alternaria 89

AB C

Fig. 66. Alternaria simsimi. A. conidia from host; B, C. scanning electron microscopes of a conidium from culture. Scale bars: A=50 μm; B=30 μm; C=5 μm.

tember 2013). REMARKS: Simmons (2004) reported A. simsimi for the first time from seed of sesame. According to Simmons (2004), the fungus was previously misidentified as A. sesami (Berry, 1960) or A. sesamicola Kawamura (Yu et al., 1982; Yeh et al., 1989). Simmons(2004) regarded A. sesamicola as a taxon of the A. tenuissima species-group. This fungus causes leaf spots of sesame. Symptoms on leaves are circular to oval, often zonate, light-brown to mid-brown spots.

34. Alternaria solani Sorauer, Z. PflKrankh. PflPath. PflSchutz: 6, t. 1: 12, 1896 (Figs. 67, 68).

SYNONYMS: Macrosporium solani Ellis & Martin, Amer. Nat. 16: 1003, 1882. Alternaria solani (E. & M.) Jones, Bull. Torrey Bot. Club 23: 353, 1896. Alternaria solani (E. & M.) Sorauer, in Jones and Grout, Bull. Torrey Bot. Club 24: 257, 1897. Alternaria solani (E. & M.) Jones & Orton, 12th Ann. Rept. Vermont Expt. Sta., p. 179, 1899. Sporidesmium solani [-] varians Vanha, Naturw. Zeits. Land-und Forst. 2: 117, 1904. Alternaria porri (Ell.) Neerg. f. sp. solani (E. and M.) Neerg., Danish Species of Alternaria and Stemphylium, p. 234, 1945. Alternaria dauci (Kühn) Neerg. f. sp. solani (E. & M.) Neerg., op. cit. p. 234, 1945. Alternaria porri (Ell.) Sawada f. sp. solani (E. and M.) Neerg., op. cit. p. 560, 1945. Alternaria dauci (Kühn) Groves & Skolko f. sp. solani (E. and M.) Neerg., op. cit., Addit. p. 2, 1945.

COLONIES on PDA, effuse, cottony or woolly, olive gray to brownish gray, abundant development of aerial mycelium, mostly secreting a yellow or reddish brown pigment into the medium, 50-60 mm in diam. after 7 days [Plate 30(A)]. CONIDIOPHORES (Primary conidiophores) arising 90 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 67. Alternaria solani. Conidia, conidiophores and sporulation habit from culture. Scale bars =50 μm.

singly or in groups of 2-3, usually unbranched (on host), sometimes branched (in culture), straight or curved, pale brown to olivaceous brown, smooth, usually with a single terminal conidiogenous site, sometimes proliferating sympodially and geniculate with 1-2 pigmented conidial scars, septate, often slightly swollen at the apex, up to 100-150 μm long, 5-10 μm wide; in culture conidiophores develop terminally or laterally on the hyphae, with several geniculations (3-9 conidiogenous sites) and branching, up to more than 200 μm long. CONIDIA mostly solitary [Plate 30(B)], occasionally (uncommonly) in chains of 2 through the agency of secondary conidiophores; obclavate or long ellipsoid to ovoid, straight to slightly curved, pale brown to dark brown, smooth or indistinctly roughened, veruculose in culture, rostrate, gradually tapering towards the beak; conidium body 50-125×15-25 μm on host, 40-115×15-25 μm in culture, with 4-12 transverse septa Pleosporales: Pleosporaceae: Alternaria 91

ABC

Fig. 68. Alternaria solani. A. conidiophores and conidia from culture; B, C. scanning electron microscopes of conidia from culture. Scale bars: A=50 μm; B=20 μm; C=5 μm.

and 1 to 2 longitudinal or oblique septa in 4-6 of the transverse segments, often slightly constricted at the transverse septa; the beak long, tapering gradually from a broad base (4-7 μm) to a narrow tip (2-3 μm), colourless or pale, septate, occasionally one-branched (bifurcate) or twice-branched (trifurcate), commonly equal in length to or longer than the body, 50-150 μm long (on host), usually shorter in culture (30-110 μm long).

DISTRIBUTION: Widespread in tropics and temperate zone. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Capsicum annuum L. (Solanaceae) (Cho et al., 1997; Yu, 2001); Solanum tuberosum L. (Solanaceae) (Nakada and Takimoto, 1928; Yu, 2001). SPECIMENS EXAMINED: On Capsicum annuum, CNU-MH00181 (Bongpyeong, 7 October 2000); on Solanum tuberosum, CNU-MH00805 (Imshil, 6 June 2000), CNU3503, 3591 (Pyeongchang, June 2006), CNU134088 (Pyeongchang, June 2013), CNU134093, 134094 (Hoengseong, July 2013). REMARKS: Alternaria solani has been recorded from a wide range of host plants in the Solanaceae including Solanum, Lycopersicum, Datura, Nicotiana, and Capsicum (Neergaard, 1945; Ellis, 1971). However, Simmons (2000) differentiated Alternaria isolates from hosts/substrates belonging to the Solanaceae into 22 taxa of Alternaria and Nimbya. Of these, 13 were presented as spp. nov., or nom. nov.. They are Alternaria subtropica E.G. Simmons (on Lycopersicon esculentum), A. beringelae E.G. Simmons (on Solanum melongena), A. tomatophila E.G. Simmons (on L. esculentum), A. subcylindrica E.G. Simmons & R.G. Roberts (on L. esculentum var. cerasiformae), A. cretica (Cooke) E.G. Simmons (on S. melongena), A. capsici E.G. Simmons (on Capsicum annuum), A. cyphomandrae E.G. Simmons (on Cyphomandra betacea), A. elegans E.G. Simmons & J. David (on L. esculentum), A. nitrimali E.G. Simmons & M.E. Palm (on S. viarum), A. grandis E.G. Simmons (on S. tuberosum), A. africana E.G. Simmons (S. melongena), and Nimbya major (Pavgi & Singh) E.G. Simmons (on Nicotiana plumbagini- folia). According to Simmons, A. solani, which causes enormous damage as the incitant of early 92 Fungal Flora of Korea·Alternaria and Allied Genera blight of potato, is a distinctly different species from the one that causes the similar early blight disease of tomato and from another that infects eggplant. The fungus can be distinguished from all the species of Alternaria on Solanaceae by the minute differences of characters of conidial morphology and sporulation pattern (Simmons, 2000, 2007). This fungus causes early blight of potato. Typical symptoms on leaves appear as water-soaked spots, and enlarge rapidly and form brown, blighted areas with indefinite borders. Tubers and stems of potato are also attacked.

35. Alternaria tagetica S.K. Shome & Mustafee, Current Sci. 35(14): 370, 1966 (Figs. 69, 70).

COLONIES on PDA, cottony, dark olive gray to dark brown, no pigment is released into the medium, 65-75 mm in diam. after 7 days; sporulation abundant on V-8 juice agar in a 12/12 hr NUV light/dark cycle. CONIDIOPHORES (Primary conidiophores) solitary or in groups of 2-5, more or less cylindrical, straight to flexuous, pale to dark brown, 1-5 septate, smooth, simple or occasionally branched, with a single terminal conidiogenous site, sometimes proliferating sympodially and geniculate with 1-2 pigmented conidial scars, often slightly swollen at the apex, 70-200 μm long, 5-8 μm wide; in culture the conidiophores are produced as side branches on the main hyphae. CONIDIA mostly solitary (Plate 31), sometimes in chains of 2 (rarely 3) through the agency of secondary conidiophores, on agar media chains occurring more frequently than in nature; straight or slightly curved, obclavate to elongated ovoid, pale to dark brown, mostly smooth or indistinctly roughened, rostrate, tapering towards the beak; in culture newly formed conidia often fail to produce a filiform beak, other erostrate conidia produce rather broad secondary conidio-

AB C

Fig. 69. Alternaria tagetica. A. conidia from culture; B, C. scanning electron microscopes of a conidium from culture. Scale bars: A=50 μm; B=25 μm; C=5 μm. Pleosporales: Pleosporaceae: Alternaria 93

Fig. 70. Alternaria tagetica. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

phores directly from the apical conidium cell, occasionally one or more from lateral cells of a conidium; conidium body 40-140×22-35 μm, with 5-11 transverse septa and 1-2 longitudinal or oblique septa in a few of the transverse divisions, often constricted at the transverse septa; the beak long, filiform, flexuous, simple and not branched, hyaline to subhyaline, 50-250 (-400) μm long, with a few extremely pale septa; in culture beaks often shorter than comparable ones in nature.

DISTRIBUTION: England, Korea, Mexico, USA. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Tagetes erecta L. (Asteraceae) (Yu and Lee, 1989; Yu, 2001); Tagetes patula L. (Asteraceae) (Yu and Lee, 1989; Yu, 2001). SPECIMENS EXAMINED: on Tagetes erecta L. CNU-MH00356 (Bongpyeong, 7 October 2000), CNU- 94 Fungal Flora of Korea·Alternaria and Allied Genera

MH00357 (Gongju, 21 October 2000), CNU003447 (Gongju, May 2006). REMARKS: This fungus is the only species of Alternaria recorded on Tagetes species. Yu and Lee (1989) first reported this species in Korea. Alternaria tagetica is similar to A. zinniae H. Pape ex M.B. Ellis in conidial morphology. A. zinniae, however, differs from the present species by having smaller (conidium body: 30-90×15-30 μm) conidia. On leaves of marigold causing brown spots; also causing brown spots and blights on flowers.

36. Alternaria tenuissima (Nees) Wiltshire, Trans. Brit. Mycol. Soc. 18(2): 157, 1933 (Figs. 71, 72).

SYNONYMS: Helminthosporium tenuissimum Nees, Nova Acta Acad. Caes. Leop.- Carol. German. Nat. Cur.: 241, f. 12, 1817. Macrosporium tenuissimum (Nees) Fr., Systema Mycologicum 3: 374, 1832. Clasterosporium tenuissimum (Nees) Sacc., Sylloge Fungorum 4: 393, 1886. Macrosporium maydis Cooke & Ellis, Grevillea 6: 87, 1878. Macrosporium martindalei Ellis and G. Martin, Amer. Naturalist 18: 189, 1884. Macrosporium podophylli Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 43: 92, 1891. Macrosporium amaranthi Peck, Bull. Torrey Bot. Club 22: 493, 1895. Alternaria amaranthi (Peck) J.M. Hook, Proc. Indiana Acad. Sci. 1920: 214, 1921. Macrosporium pruni-mahalebi Savul. & Sandu, Hed- wigia 75: 228, 1935. Alternaria rumicicola R.L. Mathur, Agnihotri & Tyagi, Current Sci. 31: 297, 1962.

COLONIES on PDA, cottony or velvety, pale gray or gray green, with varying amount of aerial mycelium, no pigment is released into the medium, 60-70 mm in diam. after 7 days. CONIDIO-

AB C

Fig. 71. Alternaria tenuissima. A, B. conidia from culture; C. scanning electron microscope of a conidium from culture. Scal bars: A, B=50 μm; C=7 μm. Pleosporales: Pleosporaceae: Alternaria 95

Fig. 72. Alternaria tenuissima. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

PHORES (Primary conidiophores) solitary or in fascicles, unbranched or sparsely branched, septate, with a single pigmented conidiogenous site at the apex, sometimes geniculate with 1-5 pigmented conidiogenous sites, often slightly swollen and almost hyaline at the apex, 20-110 μm long, 3-5 μm wide; in culture the conidiophores produced directly from the substrate or as side branches on the hyphae. CONIDIA in moderately-long chains of 5-10 with minor branching, in culture long chains (up to 13 conidia) occurring more frequently than in nature (Plate 32); long ellipsoid to elongated ovoid, or obclavate in nature, ellipsoid or broadly ovoid in culture, pale to mid-golden brown, wall ornamentation may be minutely to conspicuously verrucose, the initial conidia often awl-shaped or having a narrowly tapered upper half, sometimes terminating abruptly in a secondary conidiophore, occasionally tapering gradually towards a beak; conidium body 30-60×10-18 μm in nature, 23-55×8-16 μm in culture, with 4-10 transverse septa and 1-2 longitudinal or oblique septa in 96 Fungal Flora of Korea·Alternaria and Allied Genera some of the transverse divisions, moderately constricted at the 1-3 of the transverse septa, often with a darker median transverse septum that is more deeply constricted than other septa; beaks, when present, narrow but not filiform, higjly variable, as long as, or longer than, the conidium body (up to 40 μm), but usually shorter; secondary conidiophores usually short and 1-2-celled ca. 3-10 μm, occasionally extend to 50-60 μm.

DISTRIBUTION: Cosmopolitan. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Capsicum annuum L. (Solanaceae) (Kim and Yu, 1985, under Alternaria alternata; Yu, 2001); Fragaria ananassa Duchesne (Rosaceae) (Yu, 2001); Phaseolus vulgaris L. (Fabaceae) (Chung et al., 1977, under Alternaria fasciculata). SPECIMENS EXAMINED: On Brassica campestris subsp. napus var. nippo-oleifera, CNU111003 (Yesan, June 2011); on Capsicum annuum, CNU-MH98103 (Bonghwa, October 1997); on Fragaria ananassa, CNU-MH00809 (Nonsan, May 2000); from seed of Hordeum vulgare var. hexastichon Aschers, CNU- MH98124 (Seosan, May 1998), CNU-MH98125 (Seosan, May 1998); on Panax ginseng, CNU- MH98082 (Geumsan, October 2000). REMARKS: A. tenuissima is recognized as a species-group whose members share a longicatenate sporulation pattern (ability to produce usually unbranched chains of 5-10 or more conidia), and similar conidial shape and size, particularly the character of the median conidial trans-septa, which is often darker and more deeply constricted than other septa (Simmons, 1990, 2007). Taxa of the A. tenuissima species-group are extremely common saprophytes/secondary invaders found in a very wide range of plants and seeds, and are very widespread in nature. These are commonly misidentified as Alternaria alternata (Simmons, 2007). The species collecively known as “A. tenuissima” cause black mold of pepper fruits, black rot of strawberry fruits, etc.

37. Alternaria tomatophila E.G. Simmons, Mycotaxon 75: 1, 2000 (Figs. 73, 74).

COLONIES on PDA, effuse, woolly or cottony, pale to dark gray, with abundant non-sporulating aerial mycelium, poorly conidiogenous, commonly no pigment (rarely a pale yellow pigment) released into the medium, 45-55 mm in diam. after 7 days [Plate 33(A)]. CONIDIOPHORES (Pri- mary conidiophores) mostly solitary and unbranched, straight or curved, pale brown, smooth, usually with a single terminal pigmented zone at the site of former conidium attachment, occasionally geniculate with 1-2 pigmented conidiogenous sites, septate, often slightly swollen at the apex, up to 100-160 μm long, 5-8 μm wide; in culture conidiophores are produced singly as side branches on the main hyphae, simple, occasionally (rarely) 1-geniculate, scattered in the aerial mycelium. CONIDIA mostly solitary on host [Plate 33(B)], occasionally in chains of 2 on agar; in nature predominantly obclavate, occasionally broadly obclavate, in culture narrowly-ellipsoid to obclavate, straight to curved, pale olivaceous brown to dark brown, smooth or minutely roughened in nature, smooth or verruculose in culture, rostrate, tapering towards a long beak; conidium body 50-120×20-28 μm on host, with 5-15 transverse septa, 1-2 longitudinal or oblique septa in a few of the transverse segments, constricted at the transverse septa; in culture conidium body 40-110× 12-25 μm, with 4-12 transverse septa, one longitudinal septum in a few of the transverse segments; the beak narrowly filamentous, filiform, subhyaline, the point of transition from spore body to beak is rather abrupt and definite, occasionally once- or twice- branched, 50-200 (-250) μm long, Pleosporales: Pleosporaceae: Alternaria 97

Fig. 73. Alternaria tomatophila. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

3-4(-5) μm wide at the base tapering to 2 μm, in culture beak narrowly filamentous (ca. 2-2.5 μm wide) throughout its length, often branched once, twice or more, occasionally a filamentous beak enlarges terminally into a secondary conidiophore ca. 5-10×5 μm, initiating a 2-conidium chain.

DISTRIBUTION: Nearly worldwide, wherever the crop is cultivated. HOST/SUBSTRATE AND RECORDS IN KOREA: Lycopersicon esculentum Mill. (Solanaceae) (Nakada and Takimoto, 1928, under A. solani; Cho et al., 1997, under A. solani; Yu, 2001). SPECIMENS EXAMINED: On Lycopersicon esculentum, CNU-MH98032 (Buyeo, October 1998), CNU- MH99235 (Daejeon, June 1999), CNU003074 (Buyeo, April 2006), CNU003468 (Daejeon, May 2006), CNU003473 (Buyeo, October 2006), CNU134112, 134113 (Buyeo, August 2013). REMARKS: Alternaria tomatophila closely resembles A. solani in conidial morphology. The fungus, 98 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 74. Alternaria tomatophila. A, B. conidiophores and conidia from culture. Scale bars=50 μm.

however, can be distinguished from the latter by the characters of the beak (Simmons, 2000, 2007). The beak of A. tomatophila conidia narrows sharply after it is initiated and remains 2-2.5 μm in diam. throughout its straight to undulant length, which in nature commonly is up to 200 (-250) μm. In contrast, the beak of A. solani is ca. 4-7 μm at its point of origin and narrows gradually, not abruptly, through its straight to slightly curved length, which commonly is relatively short in nature. Even when the beak of A. solani is longer, the character of a broad base narrowing gradually throughout its length is maintained. The beak of A. solani may be as narrow as its tip (2-3 μm) as is the entire A. tomatophila beak, but it is not narrowly filiform throughout (Simmons, 2000). On leaves of tomato causing concentric rings of target spots; also causing circular sunken and dark lesions on stems and fruits.

38. Alternaria vanuatuensis E.G. Simmons & C.F. Hill, Alternaria: An Identification Manual: 260, 2007 (Figs. 75, 76).

COLONIES on PDA, effuse, vinaceous buff or pale yellow to straw, cottony or woolly, commonly releasing a yellow pigment into the agar medium, abundant sporulation on V-8 juice agar in a 12/12 hr NUV light/dark cycle, 60-70 mm diam. after 7 days. CONIDIOPHORES (Primary conidiophores) arising singly or in fascicles of 2-5, straight or curved, usually simple, sometimes branched, more or less cylindrical, with a single terminal pigmented conidiogenous site or geniculate with 1-2 pigmented conidial scars, septate, smooth, mid-brown, paler at the apex, up to 150 μm long, 5-8 μm wide. CONIDIA usually solitary, rarely in chains of 2-3 through the agency of secondary conidiophores, juvenile conidia narrow-ellipsoid or narrow-ovoid, becoming broad, long-ovoid and obclavate, smooth or distinctly roughened, pale to mid-brown, rostrate, gradually Pleosporales: Pleosporaceae: Alternaria 99

Fig. 75. Alternaria vanuatuensis. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

tapering towards the beak; conidium body 50-90 (-110)×15-26 μm, 5-10 transverse septa and 1-2 longitudinal or oblique septa in a few of the transverse segments, often constricted at the transverse septa; the beak blunt-tapered, commonly about the same length of the mature conidium body, or may be shorter, 50-90 μm long on host, 40-70 μm long on agar media, simple or sometimes one- branched beak, rarely two-branched, pale, without transverse septa or sparsely septate.

DISTRIBUTION: Korea, Vanuatu (Oceania). HOSTS/SUBSTRATES AND RECORDS IN KOREA: Allium cepa L. (Li et al., 2014), Allium fistulosum L. (Li et al., 2014). SPECIMENS EXAMINED: On Allium cepa, CNU103010, CNU103011 (Gongju, June 2010); on Allium fistulosum, CNU03367 (Geumsan, June 2003), CNU094020 (O-chang, May 2010), CNU138082 (Dae- 100 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 76. Alternaria vanuatuensis. Conidia from culture. Scale bar=50 μm.

jeon, June 2013). REMARKS: Alternaria porri has been known on Allium plants worldwide. In 2007, a new species, A. vanuatuensis, was erected by Simmons based on conidial morphology. Li et al. (2014) reported the occurence of this species in Korea for the first time. The species produces conidia with blunt- tapered beaks, while A. porri conidia have longer, filamentous beaks. It causes purple blotch of garlic, onion and spring onion. A. porri and A. vanuatuensis may occur together on the same host.

39. Alternaria zinniae H. Pape ex M.B. Ellis, Mycol. Pap. 131: 22, 1972 (Figs. 77, 78).

SYNONYM: Alternaria zinniae H. Pape, Angew. Bot. 24: 61, 1942.

COLONIES on PDA, velvety or woolly, grayish brown to blackish brown with varying amounts of pale gray aerial mycelium, no pigment is released into the medium, reaching about 45-50 mm in diam. after 7 days; sporulation abundant on V-8 juice agar in a 12/12 hr NUV light/dark cycle. CONIDIOPHORES (Primary conidiophores) arising singly or in groups of 2-5, straight or curved, usually unbranched, more or less cylindrical, with a single terminal pigmented conidiogenous site or geniculate with 1-4 pigmented conidial scars, septate, smooth, mid-brown, paler at the apex, up to 200 μm long, 5-10 μm wide; in culture the conidiophores produced terminally and laterally on the hyphae. CONIDIA mostly solitary (Plate 34), rarely in chains of 2 (in nature) or 2-3 (in culture) through the agency of secondary conidiophores; narrowly ovoid when juvenile, obclavate or ellipsoid at maturity, straight or slightly curved, pale to dark brown, mostly smooth, sometimes minu- Pleosporales: Pleosporaceae: Alternaria 101

Fig. 77. Alternaria zinniae. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

tely verruculose, rostrate, rather abruptly tapering towards a beak, occasionally short secondary conidiophores produced from lateral cells of a conidium; conidium body 30-90 (-110)×15-30 μm on host, 30-80 (-100)×15-27 μm on agar, with 3-12 transverse septa and 1 to several longitudinal and oblique septa, slightly constricted at the transverse septa; the beak long, filiform, unbranched, hyaline to pale brown, septate, 50-180 (-250) μm long, 2 μm wide along most of its length, sometimes the tip of the beak of primary conidia develops into a slightly expanded one-celled secondary conidiophore, producing a relatively small secondary conidium ca. 15-30×10-15 μm, with a short, filiform beak.

DISTRIBUTION: Africa (Egypt, Ethiopia, Ghana, Kenya, Libya, Malawi, Mauritius, Sierra Leone, South Africa, Sudan, Tanzania, Uganda, Zambia, Zimbabwe); Asia (Brunei, China, India, Indo- nesia, Japan, Korean, Nepal, Pakistan); Oceania (Australia); Europe (Cyprus, Denmark, France, Germany, Hungary, Italy, Norway, Poland, Portugal, Romania, The Netherlands); South America 102 Fungal Flora of Korea·Alternaria and Allied Genera

AB C

Fig. 78. Alternaria zinniae. A. conidia from culture; B, C. scanning electron microscopes of a conidium. Scale bars: A=50 μm; B=25 μm; C=5 μm.

(USA). HOST/SUBSTRATE AND RECORDS IN KOREA: Zinnia elegans Jacq. (Asteraceae) (Yu and Park, 1988; Yu, 2001). SPECIMENS EXAMINED: On Zinnia elegans, CNU-MH98011 (Cheongyang, November 1998), CNU111215 (Daejeon, September 2011), CNU135010 (Gongju, September 2013). REMARKS: Yu and Park (1988) reported this fungus for the first time in Korea. The fungus is very similar to Alternaria cassiae and A. tagetica in conidial morphology (see remarks under A. cassiae and A. tagetica). This species causes circular or irregular spots and blights on zinnia leaves and flowers.

Genus Alternariaster E.G. Simmons Alternaria: an identification manual: 667, 2007.

SYNONYM: Alternaria Nees, Syst. Pilze Schwämme 72, 1816.

COLONIES effuse, light brown to olive brown, thinly hairy. CONIDIOPHORES macronematous, mononematous, simple, solitary or in fascicles, sometimes geniculate with 1-2 conidial scars, pale gray to pale brown. CONIDIA mostly solitary, rarely in chains of 2, straight or slightly curved, cylindrical, long ellipsoidal or rarely obclavate, broadly rounded at both ends, beakless (erostrate), subhyaline to pale olinaceous brown, smooth, with transverse and occasionally one or more longitudinal or oblique septa. Type species: Alternariaster helianthi (Hansf.) E.G. Simmons. Pleosporales: Pleosporaceae: Alternariaster 103

40. Alternariaster helianthi (Hansford) E.G. Simmons, Alternaria: an identification manual: 667, 2007 (Figs. 79, 80).

SYNONYMS: Helminthosporium helianthi Hansf., Proc. Linn, Soc. London 153: 49, 1943. Alternaria helianthi (Hansf.) Tubaki & Nishih., Trans. Br. Mycol. Soc. 53: 148, 1969.

COLONIES on PDA, light brown to olivaceous brown, thinly hairy, very slowly growing, 15-25 mm diam. after 20 days; sporulation abundant on both PDA and V-8 juice agar. CONIDIOPHORES solitary, rarely in fascicles of two or more, simple, straight or slightly curved, cylindrical, pale gray to pale brown, almost hyaline at the apex, smooth, up to 5-septate, usually with a single terminal conidiogenous site, sometimes geniculate with 1-2 conidial scars, simple or branched, up to 200 μm long, 6-12 μm thick; in culture the conidiophores are produced terminally or laterally on the

Fig. 79. Alternariaster helianthi. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm. 104 Fungal Flora of Korea·Alternaria and Allied Genera

ABC

Fig. 80. Alternariaster helianthi. A, B. conidiophores and conidia from host; C. scanning electron microscope of a conidium from culture. Scale bars: A, B=50 μm; C=35 μm.

hyphae. CONIDIA mostly solitary (Plate 35), rarely in chains of 2 (in culture) through the agency of secondary conidiophores; straight or slightly curved, long cylindrical to long ellipsoidal on host tissue, long cylindrical to obclavate on agar media, broadly rounded at both base and apex, subhyaline to light yellow or golden brown, smooth-walled, erostrate (beakless); conidium body 40- 140×15-35 μm, with 2-12 (mostly 5-8) transverse septa and 1 (rarely 2) longitudinal or oblique septa in 1-2 of the transverse compartments, constricted at the transverse septa; sometimes a relatively broad secondary conidiophore (pseudorostrum or false beak) produced from the apical or intercalary conidium cell, initiating a secondary conidium.

DISTRIBUTION: Almost world-wide, wherever sunflower is grown. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Helianthus annuus L. (Asteraceae) (Chung et al., 1977, under Alternaria helianthi; Yu et al., 1989, under Alternaria helianthi; Yu, 2001, under Alternaria helianthi); Rudbeckia bicolor Nutt. (Asteraceae) (Oh and Shin, 1999, under Alternaria helianthi; Cho et al., 1999, under Alternaria helianthi; Yu, 2001, under Alternaria helianthi). SPECIMENS EXAMINED: On Helianthus annuus, CNU-MH98027 (Cheongyang, September 1998), CNU-MH00770 (Hongcheon, August 2000); on Rudbeckia bicolor, CNU-MH00336 (Pyeongchang, August 2000), CNU111288 (Cheongju, July 2011), 121288×2012, Gurye, August 2012). REMARKS: The fungus was first described as Helminthosporium helianthi by Hansford in 1943, and transferred to Alternaria helianthi by Tubaki and Nishihara in 1969. More recently Simmons (2007) erected a new genus, Alternariaster, and transferred the fungus to this new genus Alternariaster (A. helianthi). The conspicuous internal, pigmented, circumhilar ring found so commonly in Alternaria conidia and conidiophores is absent in A. helinthi (Simmons, 2007). Phylogenetic analysis of rDNA sequences, including the ITS region, also revealed that this species does not belong to the core Alternaria clade (Cho, 2004). It causes oval to circular leaf spots of sunflower and black-eyed Susan (Rudbeckia bicolor). Pleosporales: Pleosporaceae: Prathoda 105

Genus Prathoda Subramanian J. Indian Bot. Soc. 35: 73, 1956.

SYNONYMS: Alternaria Nees, Syst. Pilze Schwämme: 72, 1816. Elosia Pers., Mycol. Europaea 1: 12, 1822. Macrosporium Fr., Systema Mycologicum 3: 373, 1832. Rhopalidium Mont., Ann. Sci. Nat. Bot. 6: 30, 1836. Chmelia Svob.-Pol., Biológia (Bratislava) 21: 82, 1966.

CONIDIOPHORES erect or ascending, mostly simple, occasionally branched, straight or flexuous, sometimes geniculate with 1 to several pigmented conidial scars, septate, pale to mid pale brown, smooth, sometimes aggregate into discrete synnemata. CONIDIA solitary or in short chains, extremely variable in shape and size, pale brown to brown, elongate cylindrical, mostly very long, cercosporoid scolecospores, mostly smooth, rarely verruculose around the base; shorter conidia variable in shape, often with a few longitudinal or oblique septa. Type species: Prathoda saparva Subram.

41. Prathoda longissima (Deighton & MacGarvie) E.G. Simmons, Alternaria: an identification manual: 672, 2007 (Figs. 81, 82).

SYNONYM: Alternaria longissima Deighton & MacGarvie, Mycol. Papers 113: 10, 1968.

COLONIES on PDA, effuse, velvety, olivaceous gray to dark grayish-brown, slightly zonate, shortly flocculose aerial mycelium, sometimes with a purplish zonate banding in the reverse side, 34-40 mm diam. after 7days, sporulation abundant held in a 12/12 hr NUV light/dark cycle. CONIDIOPHORES more or less straight, unbranched or branched, pale to moderately brown, 50- 200 μm (up to 350 μm in culture) long, 3-5 μm wide, up to 15 septate, singly, usually with a single terminal conidiogenous site, occasionally geniculate with 1-3 conidial scars, slightly swollen at the apex, smooth but slightly verrucose over the swollen apex. CONIDIA solitary or in short chains; extremly variable in shape and size, elongate cylindrical or somewhat irregular in outline, beakless or with a long beak, pale brown to almost hyaline, smooth-walled; matured conidia usually long- ellipsoid with a simple, long, whip-like beak, 3-15 septate, overall length of rostrate conidia 130- 300 (up to 500) μm, 5-15 μm wide; beak relatively thick, with many septa, unbranched, tapering gradually from a base (4 μm) to a narrow tip (2 μm), often with a secondary conidiophore developing apically or laterally from the beak, bearing 1-2 secondary conidia; other conidia subcylindric to obclavate, erostrate, 1-8 septate, 17-60×4-11 μm; in culture the long rostrate spores are less abundant than the shorter erostrate spores. CHLAMYDOSPORES may develop in culture, some of the swollen muriform cells of the conidium, becoming irregular in shape and deep-coloured (dark brown), commonly about 16-40×16-35 μm.

DISTRIBUTION: Africa (Egypt, Malawi, Nigeria, Sierra Leone, Sudan, Tanzania, Zambia); Central and South America (Cuba, Venezuela); North America (Bermuda, USA); Asia (Bhutan, India, Laos, Malaysia, Nepal, Korea, Taiwan, Thailand); Europe (Ukraine); Australasia and Oceania (Australia, New Zealand, Solomon Islands). 106 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 81. Prathoda longissima. Conidia and conidiophores from culture. Scale bar=50 μm.

HOST/SUBSTRATE AND RECORD IN KOREA: Oryza sativa L. (Poaceae) (Deng, 2012). SPECIMENS EXAMINED: On seeds of Oryza sativa, CNU03461, 03462 (Suwon, July 2000). REMARKS: The fungus was described as Alternaria longissima by Deighton and MacGarvie in 1968. Simmons transferred the fungus to the genus Prathoda (as P. longissima). Molecular analysis of rDNA sequences such as the ITS region also revealed that this fungus does not belong to the core Alternaria clade (Cho, 2004). Morphology of the Korean isolates was in good agreement with the previous descriptions of Alternaria longissima (David, 1995; Deighton and MacGarvie, 1968) and Prathoda longissima (Simmons, 2007). It is a widespread and apparently common saprophyte in tropical and subtropical regions (David, 1995; Deighton and MacGarvie, 1968). This species has been recorded on pollen grains of numerous members of Poaceae, such as Zea mays, Pennisetum typhoides, Saccharum officinarum and Oryza sativa, on husks and grains of Oryza Pleosporales: Pleosporaceae: Stemphylium 107

Fig. 82. Prathoda longissima. A, B. two types of conidia from culture. Scale bar=50 μm.

and Sorghum, and on old leaf tissues of a wide range of host plants (David, 1995; Deighton and MacGarvie, 1968).

Genus Stemphylium Wallroth Flora Cryptogamica Germaniae 2: 300, 1833.

SYNONYMS: Scutisporium Preuss, Linnaea 24: 112, 1851. Epochniella Sacc., Michelia 2(6): 659(127), 1880. Thyrodochium Wederm., Annales Mycologici 22: 188, 1924. Soreymatosporium Sousa da Cam- ara, Prosposta Stemphylium: 18, 1930.

COLONIES effuse, gray, brown, olivaceous brown or black, velvety or cottony. MYCELIUM partly immersed or partly superficial; hyphae colourless or pale brown, septate, branched. CONI- DIOPHORES macronematous, mononematous, solitary or in fascicles, simple or occasionally loosely branched, straight or flexuous, more or less cylindrical, elongating percurrently, becoming nodose with a number of vesicular swellings, pale to mid brown, smooth or in part roughened. CONIDIA solitary, ellipsoidal, broadly ovoid or subspherical, rounded at the ends, often with a pointed conical apex, pale to mid brown or olivaceous brown, smooth or variously roughened, muriform, with transverse and longitudinal septa, often constricted at one or more of the septa. Type species: Stemphylium botryosum Wallr.

Dichotomous Key to Korean Species of Stemphylium

1. Conidia rounded at both the base and the apex ········································································2 - Conidia with pointed, conical apex···························································································6 108 Fungal Flora of Korea·Alternaria and Allied Genera

2. Conidia smooth·····································································································S. sarciniforme - Conidia ornamented·················································································································3 3. Conidia constricted at median septum, l/w ratio not more than 2 : 1 ········································4 - Conidia frequently constricted at 3 major transverse septa, l/w ratio about 2 : 1 ·······················5 4. Conidia smaller (20-33×15-23 μm), on Medicago ·················································S. globuliferum - Conidia larger (25-40×20-30 μm) ···········································································S. botryosum 5. On Astragalus ···············································································································S. astrgali - On Allium················································································································S. vesicarium 6. Conidia mostly constricted at the median septum only, 1/w ratio 2: 1 ···························S. solani - Conidia constricted at 3 major transverse septa, l/w ratio 3 : 1 or more·····································7 7. Conidia up to 65 μm long, on Solanaceae·································································S. lycopersici - Conidia up to 90 μm long, on Platycodon······························································S. platycodontis

42. Stemphylium astragali (Yoshii) W. Yamamoto, Trans. Mycol. Soc. Japan 2(5): 92, 1960 (Figs. 83, 84).

SYNONYM: Thyrospora astragali Yoshii, J. Plant. Prot. Tokyo 16: 536, 1929.

COLONIES on PDA, effuse, gray to grayish brown, hairy or velvety, reaching about 55-65 mm in diam. after 7 days. CONIDIOPHORES arising singly or in small groups, unbranched or occasio-

Fig. 83. Stemphylium astragali. A. conidia from culture; B. conidiophore and conidium from culture. Scale bars=50 μm. Pleosporales: Pleosporaceae: Stemphylium 109

Fig. 84. Stemphylium astragali. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

nally branched, straight to slightly curved, cylindrical, smooth, pale to golden brown, 1-5 septate, 30-72 μm long, 4-6 μm wide, apical conidiogenous cell swollen to 7-8 μm; in culture the conidiophores are produced as lateral branches of the hyphae. CONIDIA develop singly through a narrow pore at the apex of each conidiophore (Plate 36), oblong to broadly ellipsoid, rounded at the apex with no pointed ends, olivaceous brown to medium golden brown, verrucose, 25-55×15-28 μm on host, with a length/width ratio of 1.5-2.0 : 1, with 1-6 transverse septa and 1-2(-3) complete or nearly complete series of longitudinal septa, constricted at the 1-3 of the major transverse septa.

DISTRIBUTION: Korea and Japan. HOST/SUBSTRATE AND RECORD IN KOREA: Astragalus sinicus L. (Fabaceae) (Yu, 2001). SPECIMENS EXAMINED: On Astragalus sinicus, CNU-MH00798, CNU079907 (Hampyeong, May 110 Fungal Flora of Korea·Alternaria and Allied Genera

2000). REMARKS: This fungus has been known only from Japan (Yoshii, 1929; Yamamoto, 1961) and Korea (Yu, 2001). This species is very similar to Stemphylium vesicarium (Wallr.) Simmons in conidial morphology, however the oblong conidia of this species tend to be narrower in culture. Our recent DNA sequence data also revealed that they are two different species (unpublished data). The fungus causes leaf spot and blight of Chinese milk vetch (Astragalus sinicus).

43. Stemphylium botryosum Wallroth, Flora Crytogamica Germaniae 2: 300, 1833 (Figs. 85, 86).

SYNONYM: Pleospora tarda E.G. Simmons, Sydowia 38: 291, 1986.

CONIDIOPHORES arising singly or in groups, simple, straight to flexuous, pale brown to brown, cylindrical but enlarged slightly at one to several apical percurrent proliferations, smooth but roughened or distinctly echinulate towards the apex, 1-7 septate, 20-80 μm long, 4-8 μm wide, with vesicular swellings in the apical conidiogenous cell to about 7-10 μm diam., possess a single apical pore 5-8 μm diam.. CONIDIA solitary (Plate 37), subspherical, oblong or broadly ovoid, rounded at both the base and the apex, dilute to deep olive-brown, with 1-3 transverse septa and 1-3(-4) longitudinal septa mostly meeting the transverse septa at right angles, usually conspicuously constricted only at the median transverse septum, with walls so densely echinulate that the internal septa are somewhat obscured, 25-40×20-30 μm with a l/w ratio about 1.0-1.5 : 1, often

Fig. 85. Stemphylium botryosum. A. conidiophores and conidia from culture; B. scanning electron microscope of conidiophores and conidia from culture. Scale bars=50 μm. Pleosporales: Pleosporaceae: Stemphylium 111

Fig. 86. Stemphylium botryosum. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

with a large basal scarlike zone.

DISTRIBUTION: World-wide, common in temperate and subtropical regions. HOST/SUBSTRATE AND RECORDS IN KOREA: Symphytum officinale L. (Boraginaceae) (Lee et al., 1991, under Stemphylium spp.; Yu, 2001). SPECIMENS EXAMINED: On Symphytum officinale L., CNU-MH99092 (Pyeongchang, August 1999), CNU-MH00721 (Mokpo, September 2000), CNU004061 (Seongwhan, April 2006). REMARKS: Diagnostic characteristics of this species include the maximum conidial dimensions (25-40×20-30 μm), a length/width ratio in the range of 1.0-1.5 (i.e., large numbers of conidia are almost as broad as they are long), and the conspicuous constriction at the median transverse septum (Simmons, 1969). In Korea, several species of Allium, i.e. Allium cepa L. (Cho et al., 1997), A. fistulosum L. (Park, 1958; Cho et al., 1977) and A. sativum L. (Chung et al., 1977; Cho et al., 1997), were erroneously listed to 112 Fungal Flora of Korea·Alternaria and Allied Genera be associated with the present species. Instead, these host plants were all found to be infected with S. vesicarium (Wallr.) Simmons in Korea (Cho and Yu, 1998). See the remarks under S. vesicarium. It causes leaf spot of common comfrey in Korea.

44. Stemphylium globuliferum (Vestergr.) E.G. Simmons, Mycologia 61(1): 14, 1969 (Figs. 87, 88).

SYNONYM: Macrosporium globuliferum Vestergr., Bihang till Kungliga svenska Vetenskap-Akaded- emiens Hndlingar 22: 27, 1896.

COLONIES on PDA, effuse, white to pale gray, cottony, reaching about 35-45 mm in diam. after 7 days. CONIDIOPHORES arising singly or in groups, simple or occasionally branched, straight to flexuous, cylindrical but enlarged slightly at 1-5 apical percurrent proliferations, pale yellow-

Fig. 87. Stemphylium globuliferum. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm. Pleosporales: Pleosporaceae: Stemphylium 113

Fig. 88. Stemphylium globuliferum. A, B. conidiophores and conidia from host. Scale bars=50 μm.

brown, medium brown at the apex, smooth, septate, up to 100×5-7 μm, with vesicular swellings in the apices of the conidiogenous cell to about 8 μm; in culture the conidiophores are produced terminally and laterally on the hyphae. CONIDIA solitary; broadly ovoid or subspherical, rounded at the base and the apex, medium yellow-brown or deep brown, densely punctate, with 1-3 transverse and 3 (-4) complete or nearly complete series of longitudinal septa, usually distinctly constricted only at the median transverse septum, 20-33×15-23 μm with a l/w ratio about 1.3 : 1; in culture the conidia slightly smaller (18-30×15-20 μm) and with a deeper brown colour.

DISTRIBUTION: Temperate regions including Korea and USA. HOST/SUBSTRATE AND RECORD IN KOREA: Medicago sativa L. (Fabaceae) (Yu, 2001). SPECIMENS EXAMINED: On Medicago sativa, CNU-MH98028 (Seonghwan, October 1998), CNU- MH00543 (Cheongyang, June 2000). REMARKS: Five species of Stemphylium have been recorded on Medicago spp. (Gilchrist and Sim- mons, 1990). They are S. botryosum, S. herbarum Simmons, S. alfalfae Simmons, S. globuliferum and S. vesicarium. Mature conidia of S. globuliferum at their maximum size overlap the common size range of S. botryosum conidia. However, S. globuliferum conidia are, in general, smaller and darker than those of S. botryosum (Simmons, 1969; Gilchrist and Simmons, 1990). Another species common on leguminous hosts and sometimes confused with S. globuliferum is S. sarciniforma (Cav.) Wiltshire but this species has slightly larger (22-34 (-40)×18-27 μm) and smooth conidia. The fungus was first recorded from alfalfa in Korea by Yu (2001). S. globuliferum causes round or oval spots on leaves of alfalfa. 114 Fungal Flora of Korea·Alternaria and Allied Genera

45. Stemphylium lycopersici (Enjoji) W. Yamamoto, Trans. Mycol. Soc. Japan 2(5): 93, 1960 (Figs. 89, 90).

SYNONYMS: Thyrospora lycopersici Enjoji, J. Plant Prot., Tokyo 18: 52, 1931. Stemphylium floridanum C.I. Hannon & G.F. Weber, Phytopathology 45: 1, 1955.

COLONIES on PDA, hairy or velvety, gray or grayish brown, reverse orange to pale red, usually secreating a yellow or pale orange pigment into the medium, reaching about 50-60 mm in diam. after 7 days [Plate 38(A)]. MYCELIUM mostly immersed. CONIDIOPHORES arising singly or in groups, simple or occasionally branched, straight to flexuous, cylindrical but enlarged slightly at 1-3 apical percurrent proliferations, septate, pale to mid brown, smooth, 60-300 μm long, 5-8 μm wide, with vesicular swelling in the apical conidiogenous cell to about 8-10 μm. CONIDIA solitary [Plate 38(B)], rarely in chains of 2 through the agency of secondary conidiophores; oblong,

Fig. 89. Stemphylium lycopersici. Conidia, conidiophores and sporulation pattern from host (A) and culture (B). Scale bars=50 μm. Pleosporales: Pleosporaceae: Stemphylium 115

ABCD

Fig. 90. Stemphylium lycopersici. A. conidiophores from culture; B, C. conidiophore and conidia from culture; D. scanning electron microscope of a conidium from culture. Scale bars: A-C=50 μm; D=15 μm.

conical at the apex, bluntly rounded at the base, pale to mid brown, smooth or minutely verruculose, with 1-7 transverse and 1-2 nearly complete series of longitudinal septa, usually constricted at the 3 major transverse septa, 35-65×15-20 μm with a l/w ratio of 3 : 1 or greater.

DISTRIBUTION: Asia (Israel, India, Malaysia, Hong Kong, Korea, Japan, Saudi Arabia, Yemen); Africa (Kenya); Central and South America (Cuba, Venezuela); North America (Canada, USA); Oceania (Tahiti). HOSTS/SUBSTRATES AND RECORDS IN KOREA: Capsicum annuum L. (Solanaceae) (Kim and Yu, 1996; Yu, 2001); Lycopersicon esculentum Mill. (Solanaceae) (Min et al., 1995; Yu, 2001); Solanum melongena L. (Solanaceae) (Yu, 2001). SPECIMENS EXAMINED: On Capsicum annuum, CNU-MH99002 (Gumsan, July, 1999); on Lycopersicon esculentum, CNU-MH98085 (Buyeo, June 1998), CNU132432 (Buyeo, July 2013); on Solanum melongena, CNU-MH00116 (Ulleung island, August 2000). REMARKS: Two species of Stemphylium, S. lycopersici and S. solani Weber, have been recorded on hosts in Solanaceae. S. lycopersici differs from the latter in its longer, narrower conidia (35-65×15- 20 μm) and having three constricted transverse septa; the smaller conidia (32-58×17-28 μm) of the other species have a median constricted transverse septum. Stemphylium lycopersici causes white leaf spot of pepper, gray leaf spot of tomato etc.

46. Stemphylium platycodontis J.X. Deng & S.H. Yu, Mycol. Progress 13: 477, 2014 (Figs. 91, 92).

COLONIES on PDA, effuse, pale brown, with medium brown zones, ca. 54-60 mm in diam. after 7 days [Plate 39(A)]. MYCELIUM superficial, branched, septate, pale brown, smooth-walled; 116 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 91. Stemphylium platycodontis. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

hyphae 3-6(-10) μm wide. CONIDIOPHORES arising mostly singly, unbranched or occasionally branched, straight to flexuous, more or less cylindrical, pale to mid-brown, smooth, with 2-9 septa, 50-120 μm long, 4-6 μm wide, with vesicular swelling in the apical conidiogenous cell to about 6- 9 μm, with 1-2 percurrent proliferations; in culture the conidiophores arise as side branches on the main hyphae, up to 150 μm long. CONIDIA solitary [Plate 39(B)], mostly oblong to oblong-ellipsoid, occasionally ovoid, conical or rounded at the apex, bluntly rounded at the base, straight or slightly curved, pale to mid-brown, smooth or densely verruculose, with 3-5(-8) transverse septa and 1-5 longitudinal or oblique septa, usually constricted at the transverse septa, 33-90×12-28 μm with a l/w ratio of 1.6-4.6 (av. 3.3); occasionally (rarely) a relatively broad and short secondary conidiophore arising from apical or lateral cell of the conidium, initiating secondary conidia in chains of 2 (3).

DISTRIBUTION: Korea. Pleosporales: Pleosporaceae: Stemphylium 117

Fig. 92. Stemphylium platycodontis. A. conidiophores from culture; B. conidia from culture. Scale bar=40 μm.

HOST/SUBSTRATE AND RECORDS IN KOREA: Platycodon grandiflorum (Jacq.) A. DC. (Campanulaceae) (Yu, 2001, under Stemphylium sp.; Deng et al., 2014). SPECIMENS EXAMINED: On Platycodon grandiflorus, CNU111092 (Daejeon, June 2011), CNU-MH99147 (Jinan, 26 September 1999). REMARKS: The fungus was first recorded as Stemphylium sp. by Yu (2001) in Korea. Yu (2001) described morphological characteristics of the fungus and considered it a new taxon without prior naming. Deng et al. (2014) have assigned the fungus a new name S. platycodontis based on morphological characteristics and phylogenetic analysis of ITS region, gpd and EF-1α gene sequences. This species is very smilar to Stemphylium lycopersici in conidial morphology, but can be distinguished from the latter by its larger conidia (33-90×12-28 μm); S. lycopersici has smaller conidia (35-65× 15-20 μm). In addition, this species does not produce any pigment in PDA, while S. lycopersici diffuses brown pigment on PDA (Kim et al., 2004). It causes leaf spots and blossom rots of Chinese bellflower. Symptoms on leaves are round or irregular, gray to dark brown spots, often enlarged and coalescing, resulting in blighting.

47. Stemphylium sarciniforme (Cavara) Wiltshire, Trans. Br. Mycol. Soc. 21(3-4): 228, 1938 (Figs. 93, 94).

SYNONYMS: Macrosporium sarciniforme Cavara, La Difesa dei Parassiti: 7, 1890. Macrosporium sarcinaeforme Cavara, 1890. Thyrospora sarciniforme (Cavara) Tehon & E.Y. Daniels, Phytopath. 15: 718, 1925. Stemphylium sarcinaeforme (Cavara) Wiltshire, Trans. Br. Mycol. Soc. 21 (3-4): 228, 1938. 118 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 93. Stemphylium sarciniforme. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

COLONIES on PDA, 25-30 mm in diam. after 7 days [Plate 40(A)]. CONIDIOPHORES arising singly or in groups of 2-3, unbranched to branched, straight to mildly flexuous, cylindrical but enlarged slightly at 1-3 apical percurrent proliferations, pale to golden brown, septate, up to 60 μm long, 5-10 μm wide, with vesicular swellings of 10-14 μm diam. in the apical conidiogenous cell. CONIDIA solitary [Plate 40(B)], subspherical or broadly ellipsoid, rounded at both the base and apex, golden brown, smooth-walled, 22-35 (-40)×18-27 μm with a l/w ratio of 1.5 : 1, with usually three transverse and 1-3(-4) complete or nearly complete series of longitudinal septa, distinctly constricted at the median transverse septum.

DISTRIBUTION: Widespread wherever clover is grown. Africa, Asia, Australia, Europe, North and South America. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Trifolium pratense L. (Fabaceae) (Cho and Yu, 2000a; Yu, 2001); T. repens L. (Fabaceae) (Cho and Yu, 2000a; Yu, 2001). SPECIMENS EXAMINED: On Trifolium pratense, CNU-MH98070 (Seonghwan, October 1998), CNU- MH00109 (Gongju, July 2000); on T. repens L., CNU-MH00107 (Seonghwan, June 2000). Pleosporales: Pleosporaceae: Stemphylium 119

Fig. 94. Stemphylium sarciniforme. A. conidiophores from culture; B. conidia from culture. Scale bars=50 μm.

REMARKS: Stemphylium sarciniforme differs from S. globuliferum in its smooth-walled, larger conidia [22-35 (-40)×18-27 μm]; S. globuliferum has smaller (20-33×15-23 μm) and densely punctate conidia. The fungus causes target leaf spots of red clover and white clover.

48. Stemphylium solani G.F. Weber, Phytopathol. 20: 513, 1930 (Figs. 95, 96).

SYNONYM: Thyrospora solani (G.F. Weber) Sawada, Rept. Depart. Agric. Gov. Res. Inst. Formosa 51: 115, 1931.

COLONIES on PDA, hairy or velvety, gray or grayish brown, 45-50 mm in diam. after 7 days. CONIDIOPHORES arising singly or in groups, unbranched or occasionally branched, straight to flexuous, more or less cylindrical but enlarged slightly at 1-3 apical percurrent proliferations, septate, pale to mid brown, 80-350 μm long, 5-8 μm wide, with vesicular swellings of 8-10 μm diam. in the apical conidiogenous cell. CONIDIA solitary; oblong, pointed at the apex, rounded at the base, pale to golden brown, commonly smooth, rarely minutely verruculose, with 3-5 transverse and 1-2 longitudinal septa in 2-3 of the transverse divisions, mostly constricted at the median septum only, 32-58×17-28 μm (on host) with a l/w ratio about 2 : 1. 120 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 95. Stemphylium solani. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

DISTRIBUTION: Africa (Angola, Ivory Coast, Libya, Mauritius, Nigeria, Senegal, Sudan,Tanzania, Zambia); Asia (China, India, Japan, Korea, Thailand, Israel); Oceania (Australia, New Caledonia); Europe (Italy, UK); North America (Canada, USA); Central America (Hondulas, Jamaica, Trinidad); South America (Brazil, Colombia, Venezuela). HOSTS/SUBSTRATES AND RECORDS IN KOREA: Capsicum annuum L. (Solanaceae) (Kim and Yu, 1996; Yu, 2001); Lycopersicum esculentum Mill (Solanaceae) (Kim et al., 1999). SPECIMENS EXAMINED: On Capsicum annuum, CNU-MH98086 (Bonghwa, 14 October 1998), CNU- MH98102 (Yeongyang, 20 November 1998), CNU004345 (Bonghwa, July 2006). REMARKS: The species is distinguished by its conidia, which are oblong, pointed at the apex and constricted at the median septum only. It is a well known parasite on capsicum pepper (white leaf spot) and tomato (gray leaf spot). Pleosporales: Pleosporaceae: Stemphylium 121

A B

Fig. 96. Stemphylium solani. A. conidia from culture; B. scanning electron microscope of a conidium from culture. Scale bars: A=50 μm; B=15 μm.

49. Stemphylium vesicarium (Wallr.) E.G. Simmons, Mycologia 61(1): 9, 1969 (Figs. 97, 98).

SYNONYMS: Heiminthosporium vesicarium Wallr., Flora Crytogamica Germaniae 2: 166, 1833. Macro- sporium vesicarium (Wallr.) Sacc., Sylloge Fungorum 4: 537, 1886. Cladosporium fasciculare Fr., Systema Mycologicum 3: 370, 1832.

COLONIES on PDA, effuse, hairy or velvety, gray to grayish brown, reaching about 52-65 mm in diam. after 7 days [Plate 41(A)]. CONIDIOPHORES singly or in groups, unbranched or occasionally one-branched, straight to variously curved, smooth except sparsely punctate roughened on apical cell, pale to mid brown, cylindrical, but enlarged at the nodes and the apex, 1-6 septate, 22- 110×3-5 μm on host, 30-170×3-6 μm in culture, apical conidiogenous cell swollen to 7-9 μm. CONIDIA solitary [Plate 41(B)], broadly oval when juvenile, often with a swollen basal cell, usually oblong to broad-ellipsoid at maturity, sometimes inequilateral, olivaceous brown to medium golden-brown, rounded at the apex with no pointed ends, verrucose, 25-45×15-25 μm with a l/w ratio of 2.0-2.2 : 1 on host, 25-50×12-30 μm with a l/w ratio of 2.0-2.5 : 1 on agar media, with 3-6 transverse and 1-2(-3) longitudinal or oblique septa in a few or most of the transverse divisions, commonly constricted at the 2-3 of the major transverse septa, but only one median transverse sep-tum in juvenile conidia. The sexual morph (Pleospora sp.) develops on V-8 juice agar in 2-4 months. Ascomata are commonly 0.1-0.5 mm diam.. Asci are cylindrical to clavate with 8 ascospores which are brown, ellipsoid to clavate, and measure 30-50×12-21 μm with a l/w ratio of 1.7-2.5 : 1. 122 Fungal Flora of Korea·Alternaria and Allied Genera

Fig. 97. Stemphylium vesicarium. Conidia, conidiophores and sporulation habit from host (A) and culture (B). Scale bars=50 μm.

DISTRIBUTION: Australia, Brazil, Korea, India, New Zealand, South Africa, Spain, USA. HOSTS/SUBSTRATES AND RECORDS IN KOREA: Allium cepa L. (Liliaceae) (Cho et al., 1997, under S. botryosum; Cho and Yu, 1998; Yu, 2001); Allium fistulosum L. (Liliaceae) (Park, 1958, under S. botryosum; Cho et al., 1997, under S. botryosum; Cho and Yu, 1998; Yu, 2001); Allium sativum L. (Liliaceae) (Chung et al., 1977, under S. botryosum; Cho et al., 1997, under S. botryosum; Cho and Yu, 1998; Yu, 2001); Allium tuberosum Roth. (Liliaceae) (Chang et al., 2000); Allium thunbergii G. Don. (Liliaceae) (Yu, 2001); Allium victorialis var. platyphyllum Makino (Liliaceae) (Yu, 2001). SPECIMENS EXAMINED: on A. cepa, CNU-MH98090 (Pyeongchang, May 1998), CNU004560 (Pyeong- chang, August 2006); Allium fistulosum, CNU-MH98064 (Cheongyang, 12 September 1998); on A. sativum, CNU-MH98087 (Pyeongchang, May 1998); on A. thunbergii G. Don, CNU-MH00459 (Cheongyang, July 2000); on A. victorialis var. platyphyllum Makino, CNU-MH00456 (Bongpyeong, August 2000). REMARKS: Stemphylium vesicarium is distinguished from S. botryosum by its conidial morphology and the number of septa in mature spores, although both have muriform olivaceous brown conidia rounded at the apex, with no pointed ends. Conidia of S. botryosum usually have 3 transverse and Pleosporales: Pleosporaceae: Trichoconiella 123

AB C

Fig. 98. Stemphylium vesicarium. A. conidia from culture; B. ascus and ascospores from culture; C. scanning electron microscope of a conidium from culture. Scale bars: A, B=50 μm; C=20 μm.

1-3 longitudinal septa, constricted at one median transverse septum, and with a l/w ratio of 1.0- 1.5 : 1 (Simmons, 1969). In S. vesicarium the conidia have up to 3-6 transverse and several longitudinal septa, often constricted at 2-3 of the major transverse septa., and with a l/w ratio of 2.0-2.5 : 1. Conidia of S. botryosum are proportionally shorter than those of S. vesicarium. Previous reports (Park, 1958; Chung et al., 1977; Cho et al., 1997) of S. botryosum on Allium spp. in Korea were pro- bably misidentification of S. vesicarium. It causes severe leaf spots and blights of garlic, onion and Welsh onion.

Genus Trichoconiella B.L. Jain Kavaka 3: 39, 1976 (1975)

MYCELIUM well developed, profusely branched, septate; hyphae hyaline when young, creamy yellow when mature. CONIDIOPHORES not clealy distinguished from mature hyphae, partly erect, apices monosporous. Conidiogeous cells monotretic. CONIDIA acrogenous, creamy yellow, elongate, fusiform, with a long appendage (beak) at the apex, strongly constricted at the transverse septa, no longitudinal septa. Type species: Trichoconiella padwickii (Ganguly) B.L. Jain. 124 Fungal Flora of Korea·Alternaria and Allied Genera

50. Trichoconiella padwickii (Ganguly) B.L. Jain, Kavaka 3: 39, 1976 (Figs. 99, 100).

SYNONYMS: Trichoconis padwickii Ganguly, J. Indian Bot. Soc. 26: 238, 1948. Alternaria padwickii (Ganguly) M.B. Ellis, Dematiaceous Hyphomycetes: 495, 1971.

COLONIES on PDA, effuse, gray, reverse red or dark red, often secreting a yellow or purple pigment into the medium, sporulation abundant held in a 12/12 hr NUV light/dark cycle, small black sclerotia often formed in old cultures, 65-69 mm diam. after 7 days [Plate 42(A)]. CONIDIO- PHORES produced laterally or terminally from hyphae, straight to slightly curved, septate, pale brown, smooth, minutely echinulate at the apex, 50-180×3-4 μm, often swollen at the apex to 5-6

Fig. 99. Trichoconiella padwickii. Conidia, conidiophores and sporulation habit from culture. Scale bars=50 μm. Pleosporales: Pleosporaceae: Trichoconiella 125

Fig. 100. Trichoconiella padwickii. A. conidiophore from culture; B. conidia from culture. Scale bars =50 μm.

μm, commonly only one terminal conidiogenous site. CONIDIA solitary [Plate 42(B)], straight or curved, fusiform to obclavate, hyaline when juvenile, golden-brown to straw-colored with age, smooth or minutely echinulate, sometimes constricted at the septa, rostrate tapering towards a narrow beak, conidium body 50-110×10-20 (-25) μm, with 3-5 transverse septa, no longitudinal septa; the beak long, colorless or pale brown, septate, not branched, 40-120 μm long, 2-3 μm wide along most of its length; chlamydospores occasionally formed in old culture, intercalary on a hyphae.

DISTRIBUTION: Widespread in Asia (Korea), parts of Oceania (but not reported from Australia), Egypt, Malagasy, Nigeria and Surinam. HOST/SUBSTRATE AND RECORD IN KOREA: Oryza sativa L. (Poaceae) (Yu and Park, 1982). SPECIMENS EXAMINED: On seed of Oryza sativa, CNU 03532, 03533 (Iksan, July 2000). REMARKS: The fungus was first referred to Trichoconis padwickii Ganguly (1947), then transferred by Ellis (1971) to Alternaria padwickii (Ganguly) M.B. Ellis. Later Jain (1976) considered that placement of this species in Alternaria was not appropriate, and erected the new genus Trichoconiella (as T. padwickii). Morphology of our isolates agreed well with the previous descriptions of A. padwickii (1971) and T. padwickii (1976). It is a well-known seed-borne fungus of rice. Although reported to cause seedling blight and stackburn disease of rice (Ellis and Holliday, 1972; Ganguly, 1947), its pathogenicity on rice has not been clearly known in Korea. 126

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Plates

1A 1B

1. Alternaria acalyphae. A. colonies on PDA: (Left) obverse, (Right) reverse; B. sporulation pattern from host. Scale bar=50 μm.

2A 2B

2. Alternaria alternata. A. colony on PDA (obverse); B. sporulation pattern from culture. Scale bar=50 μm.

3A 3B

3. Alternaria araliae. A. colnies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from culture. Scale bar=50 μm. Plates 135

4A 4B

4. Alternaria atra. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from culture. Scale bar=50 μm.

6A 6B

6. Alternaria brassicicola. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from host. Scale bar=50 μm.

5. Alternaria brassicae. Sporulation pattern from host. Scale bar=50 μm. 7. Alternaria calendulae. Sporulation pattern from host. Scale bar=50 μm. 136 Fungal Flora of Korea·Alternaria and Allied Genera

8. Alternaria cassiae. Sporulation pattern from host. Scale bar=50 μm. 9. Alternaria cinerariae. Spo- rulation pattern from host. Scale bar=50 μm.

10A 10B

10. Alternaria cosmosa. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from host. Scale bar=50 μm.

11A 11B 12

11. Alternaria cucumerina. A. colony on PDA (obverse); B. sporulation pattern from host. Scale bar= 50 μm. 12. Alternaria dauci. Sporulation pattern from host. Scale bar=50 μm. Plates 137

13 14

13. Alternaria embellisia. Colonies on PDA: (L) obverse, (R) reverse. 14. Alternaria gaisen. Sporula- tion pattern from culture. Scale bar=50 μm.

15 16 17

15. Alternaria helianthinficiens. Sporulation pattern from host. Scale bar=50 μm. 16. Alternaria iridicola. Sporulation pattern from host. Scale bar=50 μm. 17. Alternaria japonica. Sporulation pattern from culture. Scale bar=50 μm.

18A 18B

(a) (b) (c)

18. Alternaria longipes. A. (a, b) colonies on PDA, (c) A. alternata; B. sporulation pattern from culture. Scale bar=50 μm. 138 Fungal Flora of Korea·Alternaria and Allied Genera

19 20A 20B

19. Alternaria macrospora. Sporulation pattern from host. Scale bar=50 μm. 20. Alternaria mali. A. colony on PDA (obverse); B. sporulation pattern from host. Scale bar=50 μm.

21A 21B 21C

21. Alternaria panax. A. colonies on PDA: (L) obverse, (R) reverse; B, C. sporulation pattern from host (B) and culture (C). Scale bars=50 μm.

22 23

22. Alternaria peucedani. Sporulation pattern from host. Scale bar=50 μm. 23. Alternaria porri. Sporulation pattern from host. Scale bar=50 μm. Plates 139

24 25

24. Alternaria protenta. Sporulation pattern from host. Scale bar=50 μm. 25. Alternaria radicina. Sporulation pattern from culture. Scale bar=50 μm.

26 27A 27B

26. Alternaria ricini. Sporulation pattern from host. Scale bar=50 μm. 27. Alternaria scirpicola. A, B. sporulation pattern from culture (A) and host (B). Scale bars=50 μm.

28A 28B

28. Alternaria sesami. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from culture. Scale bar=50 μm. 140 Fungal Flora of Korea·Alternaria and Allied Genera

29A 29B

29. Alternaria simsimi. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from culture. Scale bar=50 μm.

30A 30B

30. Alternaria solani. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from host. Scale bar=50 μm.

31 32

31. Alternaria tagetica. Sporulation pattern from host. Scale bar=50 μm. 32. Alternaria tenuissima. Sporulation pattern from culture. Scale bar=50 μm. Plates 141

33A 33B

33. Alternaria tomatophila. A. colonies from PDA: (L) obverse, (R) reverse; B. sporulation pattern from host. Scale bar=50 μm.

34 35

34. Alternaria zinniae. Sporulation pattern from host. Scale bar=50 μm. 35. Alternariaster helianthi. Sporulation pattern from host. Scale bar=50 μm.

36 37

36. Stemphylium astragali. Sporulation pattern from host. Scale bar=50 μm. 37. Stemphylium botryosum. Sporulation pattern from host. Scale bar=50 μm. 142 Fungal Flora of Korea·Alternaria and Allied Genera

38A 38B

38. Stemphylium lycopersici. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from host. Scale bar=50 μm.

39A 39B

39. Stemphylium platycodontis. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from host. Scale bar=50 μm.

40A 40B

40. Stemphylium sarciniforme. A colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from host. Scale bar=50 μm. Plates 143

41A 41B

41. Stemphylium vesicarium. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from host. Scale bar=50 μm.

42A 42B

42. Trichoconiella padwickii. A. colonies on PDA: (L) obverse, (R) reverse; B. sporulation pattern from culture. Scale bar=50 μm. 144

Index of Host Plants/Substrates

A Cucumis sativus 44 Cucurbita moschata 44 Acalypha australis 18 Cucurbita spp. 44 Acanthopanax spp. 22 Accutium lappa 19 Allium cepa 77, 99, 122 D fistulosum 77, 99, 122 monathum 19 Daucus carota var. sativa 46, 81 porum 77 Dianthus barbatus var. japonicus 69 sativum 50, 77, 122 caryophyllus 69 schoenoprasmum var. orientale 77 littorsus 69 senescens 77 Dianthus sinensis 48, 69 thunbergii 122 Dianthus superbus var. longicalycinus 48, 69 tuberosum 77, 122 victorialis var. platyphyllum 122 Aloe vera 19 E Antirrhinum majus 19 Aralia continentalis 22 Echinopanax horridum 23 elata 23 Eucommia ulmoides 19 Arctium lappa 19 Armoracia lapathifolia 26, 28, 30 F

B Fragaria ananassa 19, 96

Belamcanda chinensis 58 Brassica campestris subsp. napus var. nippo-oleifera G 28, 61, 96 campestri subsp. napus var. pekinensis 28, 30, Gladiolus gandabensis 58 61 Gossipium indicum 55, 66 juncea var. integrifolia 28 oleracea var. acephala 31 oleracea var. capitata 28, 31 H

Helianthus annuus 56, 79, 104 C Hibscus syriacus 19 Hordeum vulgare var. hexastichon 96 Calendula officinalis 33 Capsicum annuum 19, 91, 96, 115, 120 Cassia tora 35 K Citrullus lanatus 44 Citrus sinensis 39 Kalopanax pictus 23 Cosmos bipinnatus 41 Index of Host Plants/Substrates 145

L 28, 31, 61 Ricinus communis 83 Lepidium apetalum 28 Rorippa cantoniensis 28 Lycopersicon esculentum 19, 25, 97, 115, 120 Rorippa islandica 28 Rudbeckia bicolor 56, 79, 104

M S Malus pumila var. dulcissima 68 Medicago sativa 113 Scirpus maritimus 85 Senecio flammeus 38 vulgaris 38 N Sesamum indicum 19, 86, 88 Solanum melongena 115 Nicotiana tabacum 63 tuberosum 91 Symphytum officinale 111 Soil 26 O

Oryzae sativa 19, 106, 125 T

Tagetes erecta 93 P patula 93 Tarasacum coreanum 19 Panax ginseng 72, Trifolium pratense 118 quinqueforium 72 repens 118 Peucedanum japonicum 74 Phaseolus vulgaris 96 Platycodon grandiflorum 117 Z Pyrus serotina 52 Zinnia elegans 102

Raphanus sativus var. hortensis for. acanthiformis 146

Index of Fungal Names

A destruens 53 dianthi 49, 68-70 Allewia 9, 13 dianthicola 14, 47-49, 70 Alternaria 5-9, 11, 13, 50, 85, 102, 105 dumosa 40 acalyphae 13, 16-18, 134 elegans 91 africana 91 ellipsoidea 49 alliariae-officinalis 26 embellisia 14, 49-51, 137 alternata 6, 7, 13, 19-22, 31, 40, 52, 53, 55, eryngii 53 63, 64, 68, 96, 134 ethzedia 29 amaranthi 94 gaisen 14, 19, 22, 51-53, 64, 137 araliae 13, 22-24, 70, 72, 134 gossypii 55 artocariis 47, 81 gossypina 14, 53-55, 66 atra 13, 24-26, 135 grandis 91 beringelae 91 helianthi 7, 102-104 brassicae 13, 26-29, 31, 44, 74, 75, 135 helinthiinficiens 55 brassicae var. dauci 45 helianthinficiens 14, 55-57, 79, 137 brassicae var. macrospora 26, 61 infectoria 7 brassicicola 29-31, 135 interrupta 40 brassinae 29 iridicola 14, 57-59, 137 broccoli-italicae 29 japonica 14, 29, 31, 59-62, 70, 137 calendulae 13, 32-34, 135 kikuchiana 19, 51 capsici 91 limoniasperae 40 carotiincultae 47, 81 longipes 14, 19, 22, 62-64, 137 cassiae 13, 34-36, 102, 136 longispora 49 caudata 44 longissima 7, 105, 106 cheiranthi 7 loofahae 44 cinerariae 13, 36-38, 136 macrospora 14, 55, 64-66, 83, 138 citri 13, 19, 38-40 mali 14, 19, 66-68, 138 citriarbusti 40 malvacearum 55 citrimacularis 40 matthiolae 59, 62 colombiana 40 nepalensis 29 cosmosa 13, 40-42, 136 nitrimali 91 cretica 91 nobilis 14, 49, 68-70 cucumericola 44 novae-zelandiae 47 cucumerina 14, 42-44, 136 oleracea 29 cylindrorostra 44 oryzae 19 cyphomandrae 91 padwickii 8, 124, 125 danida 42 panacies 70 dauci 14, 45-47, 136 panax 14, 22-24, 70-72, 74, 75, 138 daucicaulis 47 peponicol 44 daucifolii 47 perangusta 40 dauci f. sp. solani 89 peucedani 14, 73-75, 138 Index of Fungal Names 147

porri 14, 75-77, 100, 138 Cladosporium 6 porri f. sp. dauci 45 fasciculare 121 porri f. sp. solani 89 Clasterosporium 6 protenta 14, 56, 77-79, 139 scirpicola 83 radicina 14, 47, 79-81, 139 tenuissimum 94 raphani 29, 59, 61, 62 ricini 14, 81-83, 139 rumicicola 94 E scirpicola 8, 14, 83-85, 139 senecionsis 36, 38 Elosia 13, 105 sesami 14, 85-87, 89, 139 Embellisia 5, 8, 9, 13, 50 sesamicola 88, 89 allii 8, 49, 50 simsimi 14, 86, 87-89, 140 Epochniella 107 solani 14, 89-92, 97, 98, 140 subcylindrica 91 subtropica 91 H tagetica 14, 92-94, 102, 140 tangelonis 40 Helminthosporium 6 tenuis 6, 19 allii 8, 49 tenuissima 14, 49, 53, 64, 94-96, 140 brassicae 29 tomatophila 14, 91, 96-98, 141 brassicicola 29 toxicogenica 40 helianthi 103, 104 turkisafria 40 tenuissimum 94 vanuatuensis 14, 77, 98-100 vesicarium 121 zhengzhouensis 47 zinniae 14, 36, 94, 100-102, 141 Alternariaster 5, 7, 11, 102, 104 L helianthi 56, 57, 79, 103-104, 141 Lewia 13

B M Botryomyces 13 Brachycladium 13 Macrosporium 6-9, 13, 105 acalyphae 16, 18 amaranthi 94 C araliae 22, 23 brassicae 26 Ceivellia 13 calendulae 32 Cercospora 85 carotae 45 bloxami 26 circinans 29 lepidii 26 cucumerinum 42 moldavica 26 dauci 45 scirpicola 8, 84 dianthi 68 Chalastospora 13 erumpens 19 Chmelia 13, 105 fasciculatum 19 148 Fungal Flora of Korea·Alternaria and Allied Genera

globuliferum 112 R gossypium 53 herculeum 26 Rhopalidium 13, 105 iridicola 57 longipes 62 macrosporum 64 S martindalei 94 maydis 94 Scutisporium 107 meliloti 19 Sinomyces 13 nobilis 68 Soreymatosporium 107 parasiticum 7, 8 Sphaeria scirpicola 83 podophylii 94 Sporidesmium 6 polytrichi 19 exitiosum 26 porri 75 exitiosum f. alternariodes 29 puruni-mahalebi 94 exitiosum f. luxuriosums 29 ricini 81 exitiosum var. dauci 45 sarcinaeforme 117 longipedicellatum 64 sarciniforme 117 scirpicola 8, 83, 85 sarcinula 7, 8 solani 89 sequierii 19 Stemphylium 5, 7-9, 11, 107 sesami 85 alfalfae 113 solani 89 astragali 107, 108-110, 141 tenuissimum 94 atrum 24 vesicarium 121 botryosum 7, 8, 107, 110-111, 113, 122, 123, 141 floridanum 115 N globuliferum 107, 112-113, 119 herbarum 8, 113 Nimbya 5, 8, 9, 13, 85, 89 lanuginosum 8 major 91 lycopersici 107, 114-115, 117, 142 scirpicola 8, 84, 85 platycodontis 107, 115-117, 142 radicinum 79 sarcinaeforme 117 P sarciniforme 107, 113, 117-119, 142 solani 107, 115, 119-121 Pleospora 8, 9 vesicarium 107, 108, 112, 113, 121-123, 143 eleocharidis 84 herbarum 7, 8 heleocharidis 83 T scirpicola 8 tarda 8, 110 Teretispora 13 Polydesmus exitiosus var. dauci 45 Thyrodochium 107 Prathoda 5, 7, 11, 13, 105 Thyrospora astragali 108 longissima 105-107 lycopersici 115 saparva 105 radicina 79 Pseudostemphylium radicinum 79 sarciniforme 117 Index of Fungal Names 149

solani 119 atrum 24, 26 Torula 6 botrytis 8 alternata 6, 19 Undifilum 13 Trichoconiella 5, 8, 11, 123 padwickii 8, 123-125, 143 Trichoconis padwickii 124, 125 Y

Ybotromyces 13 U

Ulocladium 5, 7-9, 13