Abstract

INTRODUCTION

Alternaria is a ubiquitous fungal genus that includes saprobic, endophytic and pathogenic species. It is associated with a wide variety of substrates including seeds, plants, agricultural products, animals, soil and the atmosphere. Species of Alternaria are known as serious plant pathogens, causing major losses on a wide range of crops. Several taxa are also important postharvest pathogens, causative agents of phaeohyphomycosis in immuno-compromised patients or airborne allergens. Because of the significant negative health effects of Alternaria on humans and their surroundings, a correct and rapid identification of Alternaria species would be of great value to researchers, medical mycologists and the public alike.

Alternaria was originally described by Nees (1816), based on A. tenuis as the only species. Characteristics of the genus included the production of dark-coloured phaeodictyospores in chains, and a beak of tapering apical cells. Von Keissler (1912) synonymised both A. tenuis and Torula alternata (Fries 1832) with Alternaria alternata, due to ambiguities in Nees’s description of A. tenuis. Two additional genera, Stemphylium (Wallroth 1833) and Ulocladium (Preuss 1851) were subsequently described for phaeodictyosporic hyphomycetes, further complicating the taxonomic resolution in this group of fungi. Several re-descriptions and revised criteria of these genera (Saccardo 1886, Elliot 1917, Wiltshire 1933, 1938, Joly 1964) resulted in a growing number of new species. Results of a lifetime study on Alternaria taxonomy based upon morphological characteristics were summarised in Simmons (2007), in which 275 Alternaria species were recognised. One species was transferred to the genus Prathoda and three new genera, Alternariaster, Chalastospora and Teretispora, were segregated from Alternaria.

Molecular studies revealed multiple non-monophyletic genera within the Alternaria complex and Alternaria species clades, which do not always correlate to species-groups based upon morphological characteristics (Pryor & Gilbertson 2000, Chou & Wu 2002, de Hoog & Horré 2002, Pryor & Bigelow 2003, Hong et al. 2005, Inderbitzin et al. 2006, Pryor et al. 2009, Runa et al. 2009, Wang et al. 2011, Lawrence et al. 2012). The A. alternata, A. brassicicola, A. infectoria, A. porri and A. radicina species-groups were strongly supported by these studies and two new species-groups, A. sonchi (Hong et al. 2005) and A. alternantherae (Lawrence et al. 2012) and three new genera, Crivellia (Inderbitzin et al. 2006), Undifilum (Pryor et al. 2009) and Sinomyces (Wang et al. 2011), were described. The latest molecular revision of Alternaria (Lawrence et al. 2013) introduced two new species groups, A. panax and A. gypsophilae, and elevated eight species-groups to sections within Alternaria. The sexual phylogenetic Alternaria lineage, the A. infectoria species-group, did not get the status of section, in contrast to the eight asexual phylogenetic lineages in Alternaria. The Alternaria complex currently comprises the genera Alternaria, Chalastospora (Simmons 2007), Crivellia, Embellisia, Nimbya, Stemphylium, Ulocladium, Undifilum and the recently described Sinomyces together with eight sections of Alternaria and the A. infectoria species-group.

The aim of the present study was to delineate the phylogenetic lineages within Alternaria and allied genera, and to create a robust taxonomy. Phylogenetic inferences were conducted on sequence data of parts of the 18S nrDNA (SSU), 28S nrDNA (LSU), the internal transcribed spacer regions 1 and 2 and intervening 5.8S nrDNA (ITS), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), RNA polymerase second largest subunit (RPB2) and translation elongation factor 1-alpha (TEF1) gene regions of ex-type and reference strains of Alternaria species and all available allied genera.

MATERIAL AND METHODS

RESULTS

Phylogeny

For defining the taxonomy of Alternaria and allied genera, 121 strains were included in the Alternaria complex alignment. The alignment length and unique site patterns of the different genes and gene combinations are stated in Table 2. The original ITS alignment consisted of 577 characters of which the first 78 are excluded as this contained a non-alignable region. In the original TEF1 alignment (375 characters) we coded the major inserts (Table 3), which otherwise would negatively influence the phylogeny, resulting in a TEF1 alignment of 269 characters. All phylogenies, different phylogenetic methods and gene regions or gene combinations used on this dataset (data not shown, trees and alignments lodged in TreeBASE), show a weak support at the deeper nodes of the tree. The only well-supported node (Bayesian posterior probability of 1.0, RAxML Maximum Likelihood support value of 100) in all phylogenies separates Embellisia annulata CBS 302.84 and the Pleospora/Stemphylium clade from the Alternaria complex (Fig. 1). In the Alternaria clade, six monotypic lineages and 24 internal clades occur consistently in the individual and combined phylogenies, although positions vary between the different gene regions or combinations used. The support values for the clades within Alternaria (called sections) are plotted in a heat map (Table 2) per gene and phylogenetic method used. The support values for the different phylogenetic methods vary, with the Bayesian posterior probabilities being higher than the RAxML bootstrap support values (Table 2). The SSU, LSU and ITS phylogenies display a low resolution, which reflects in poor to no support of the sections. Therefore, we chose not to include them in the multi-gene alignments, except in the all-gene alignment. In the GAPDH phylogenies, sect. Cheiranthus, sect. Nimbya and sect. Pseudoulocladium are poorly supported and “A. resedae” clusters separate from sect. Cheiranthus. In the RPB2 phylogenies the support values for sect. Alternata, sect. Embellisioides and sect. Eureka are relatively low; A. cumini clusters in sect. Embellisioides instead of sect. Eureka and U. capsici clusters separate from sect. Pseudoulocladium. The TEF1 phylogenies did not support sect. Nimbya and show relative low support for sect. Cheiranthus, sect. Dianthicola, sect. Embellisioides, sect. Panax, sect. Phragmosporae and sect. Radicina, and A. cumini clusters outside sect. Eureka. In the 2-region phylogenies U. capsici clusters outside sect. Pseudoulocladium based on GAPDH and RPB2, E. indefessa clusters outside sect. Cheiranthus based on GAPDH and TEF1, and sect. Eureka is poorly supported based on RPB2 and TEF1. The combined phylogeny based on the GAPDH, RPB2 and TEF1 sequences (Fig. 1) is displayed, as these are the genes with the best resolution.

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Fig. 1.

Bayesian 50 % majority rule consensus tree based on the GAPDH, RPB2 and TEF1 sequences of 121 strains representing the Alternaria complex. The Bayesian posterior probabilities (PP) and RAxML bootstrap support values (ML) are given at the nodes (PP/ML). Thickened lines indicate a PP of 1.0 and ML of 100. The tree was rooted to Stemphylium herbarum (CBS 191.86). The monotypic lineages are indicated by black dots.

Table 2.

Summary of locus and phylogenetic results as well as a heat map of the Bayesian posterior probabilities and RAxML boostrap support values per Alternaria section.

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Table 3.

Coded inserts in the TEF1 sequence alignment.

Species	Nt position	Coded	Nt position	Coded
Alternaria elegans	23 to 39	TC
Alternaria simsimi	23 to 39	TCC
Alternaria dauci	186 to 205	C	221 to 269	TACTT
Alternaria macrospora	186 to 205	C	221 to 269	TCCCC
Alternaria porri	186 to 205	C	221 to 269	ACTTA
Alternaria pseudorostrata	186 to 205	C	221 to 269	TGGTA
Alternaria solani	186 to 205	C	221 to 269	-AAGG
Alternaria tegetica	186 to 205	C	221 to 269	CACAC

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The final Pleosporineae alignment included 74 strains, representing six families, and consisted of 2 506 characters (SSU 935, LSU 796, RPB2 775) of which 700 were unique site patterns (SSU 111, LSU 145, RPB2 444). In the SSU alignment a large insertion at position 446 in the isolates Chaetosphaeronema hispidulum CBS 216.75, Pleospora fallens CBS 161.78, Pleospora flavigena CBS 314.80 and Ophiosphaerella herpotrichia CBS 620.86 was excluded from the phylogenetic analyses. A total of 43 202 trees were sampled after the burn-in. The type species of Clathrospora, C. elynae, forms a well-supported clade, located basal to the Pleosporaceae (Fig. 2), outside the Alternaria complex. The type species of Comoclathris, C. lanata, was not available for study but the two Comoclathris compressa strains cluster in a well-supported clade within the Pleosporaceae outside Alternaria s. str. The genus Alternariaster, with Alternariaster helianthi as type and only species, also clusters outside the Alternaria complex and even outside Pleosporaceae; it belongs to the Leptosphaeriaceae instead (Fig. 2). Embellisia annulata is identical to Dendryphiella salina, and forms a well-supported clade in the Pleosporaceae together with Dendryphiella arenariae. As the type species of Dendryphiella, D. vinosa, clusters outside the Pleosporineae (dela Cruz 2006, Jones et al. 2008), Dendryphiella salina and D. arenariae are placed in a new genus, Paradendryphiella, below.

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Fig. 2.

Bayesian 50 % majority rule consensus tree based on the SSU, LSU and RPB2 sequences of 74 strains representing the Pleosporineae. The Bayesian posterior probabilities (PP) and RAxML bootstrap support values (ML) are given at the nodes (PP/ML). Thickened lines indicate a PP of 1.0 and ML of 100. The tree was rooted to Julella avicenniae (BCC 18422).

ALTERNARIA SECTIONS

Section Alternantherae D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 540. 2013. Fig. 3.

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Fig. 3.

Alternaria sect. Alternantherae: conidia and conidiophores. A-D. A. alternantherae. E-H. A. perpunctulata. Scale bars = 10 μm.

Type species: Alternaria alternantherae Holcomb & Antonop.

Diagnosis: Section Alternantherae contains short to moderately long conidiophores with a conidiogenous tip which can be enlarged. Conidia are narrowly ellipsoid or ovoid, sometimes subcylindrical, solitary or rarely paired, sometimes slightly constricted near some septa, longitudinal or oblique septa occasionally occur, disto- and euseptate, with a long apical narrow beak. The conidial beak is unbranched, septate or aseptate, long filiform, and sometimes swollen at the end. Internal compartmentation occurs, cell lumina tend to be broadly octagonal to rounded.

Notes: Section Alternantherae was recently established by Lawrence et al. (2013) after first being described as species-group A. alternantherae (Lawrence et al. 2012). The described section consists of three former Nimbya species which formed a separate clade amidst the Alternaria species-groups based on sequences of the GAPDH, ITS and Alt a 1 genes (Lawrence et al. 2012). Nimbya celosiae is placed in this section based on the data of Lawrence et al. (2012), while N. gomphrenae is placed in the section based on ITS sequence data from Chou & Wu (2002).

Alternaria alternantherae Holcomb & Antonop., Mycologia 68: 1126. 1976.

• ≡ Nimbya alternantherae (Holcomb & Antonop.) E.G. Simmons & Alcorn, Mycotaxon 55: 142. 1995.

Alternaria celosiicola Jun. Nishikawa & C. Nakash., J. Phytopathol.: 10.1111/jph.12108 (p. 3). 2013.

Basionym: Nimbya celosiae E.G. Simmons & Holcomb, Mycotaxon 55: 144. 1995.

• ≡ Alternaria celosiae (E.G. Simmons & Holcomb) D.P. Lawr., M.S. Park & B.M. Pryor, Mycol. Progr. 11: 811. 2012. (nom. illegit., homonym of Alternaria celosiae (Tassi) O. Savul. 1950).

Alternaria gomphrenae Togashi, Bull. Imp. Coll. Agric. 9: 6. 1926.

• ≡ Nimbya gomphrenae (Togashi) E.G. Simmons, Sydowia 41: 324. 1989.

Alternaria perpunctulata (E.G. Simmons) D.P. Lawr., M.S. Park & B.M. Pryor, Mycol. Progr. 11: 811. 2012.

Basionym: Nimbya perpunctulata E.G. Simmons, Stud. Mycol. 50: 115. 2004.

Section Alternata D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 538. 2013. Fig. 4.

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Fig. 4.

Alternaria sect. Alternata: conidia and conidiophores. A, N. A. daucifolii. B, L-M. A. arborescens. C, H-J. A. alternata. D, O. A. gaisen. E. A. limoniasperae. F, K. A. tenuissima. G, P. A. longipes. Scale bars = 10 μm.

Type species: Alternaria alternata (Fr.) Keissl.

Diagnosis: Section Alternata contains straight or curved primary conidiophores, short to long, simple or branched, with one or several apical conidiogenous loci. Conidia are obclavate, long ellipsoid, small or moderate in size, septate, slightly constricted near some septa, with few longitudinal septa, in moderately long to long, simple or branched chains. The conidium body can narrow gradually into a tapered beak or secondary conidiophore. Secondary conidiophores can be formed apically or laterally with one or a few conidiogenous loci.

Notes: Next to the species that are displayed in our phylogeny, 14 more are included in sect. Alternata based on the study of Lawrence et al. (2013) and confirmed by our molecular data (not shown). We chose not to include 11 species from the study of Lawrence et al. (2013). The species A. gossypina, A. grisae, A. grossulariae, A. iridis, A. lini, A. maritima and A. nelumbii were not recognised by Simmons (2007) and the strains of A. malvae, A. rhadina, A. resedae and A. tomato used by Lawrence et al. (2013) were not authentic. Section Alternata comprises almost 60 Alternaria species based on ITS sequence data (data not shown). The molecular variation within this section is low.

Alternaria alternata (Fr.) Keissl., Beih. Bot. Centralbl., Abt. 2, 29: 434. 1912.

Basionym: Torula alternata Fr., Syst. Mycol. (Lundae) 3: 500. 1832 (nom. sanct.).

• = Alternaria tenuis Nees, Syst. Pilze (Würzburg): 72. 1816 [1816-1817].

Additional synonyms listed in Simmons (2007)

Alternaria angustiovoidea E.G. Simmons, Mycotaxon 25: 198. 1986.

Alternaria arborescens E.G. Simmons, Mycotaxon 70: 356. 1999.

Alternaria burnsii Uppal, Patel & Kamat, Indian J. Agric. Sci. 8: 49. 1938.

Alternaria cerealis E.G. Simmons & C.F. Hill, CBS Biodiversity Ser. (Utrecht) 6: 600. 2007.

Alternaria citriarbusti E.G. Simmons, Mycotaxon 70: 287. 1999.

Alternaria citrimacularis E.G. Simmons, Mycotaxon 70: 277. 1999.

Alternaria colombiana E.G. Simmons, Mycotaxon 70: 298. 1999.

Alternaria daucifollii E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 518. 2007.

Alternaria destruens E.G. Simmons, Mycotaxon 68: 419. 1998.

Alternaria dumosa E.G. Simmons, Mycotaxon 70: 310. 1999.

Alternaria gaisen Nagano ex Hara, Sakumotsu Byorigaku, Edn 4: 263. 1928.

• = Alternaria gaisen Nagano, J. Jap. Soc. Hort. Sci. 32: 16-19. 1920. (nom. illegit.)

• = Alternaria kikuchiana S. Tanaka, Mem. Coll. Agric. Kyoto Univ., Phytopathol. Ser. 28: 27. 1933.

• = Macrosporium nashi Miura, Flora of Manchuria and East Mongolia, Part III Cryptogams, Fungi: 513. 1928.

Alternaria herbiphorbicola E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 608. 2007.

Alternaria limoniasperae E.G. Simmons, Mycotaxon 70: 272. 1999.

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

Basionym: Macrosporium longipes Ellis & Everh., J. Mycol. 7: 134. 1892.

• = Alternaria brassicae var. tabaci Preissecker, Fachliche Mitt. Österr. Tabakregie 16: 4. 1916.

Alternaria perangusta E.G. Simmons, Mycotaxon 70: 303. 1999.

Alternaria postmessia E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 598. 2007.

Alternaria tangelonis E.G. Simmons, Mycotaxon 70: 282. 1999.

Alternaria tenuissima (Nees & T. Nees: Fr.) Wiltshire, Trans. Brit. Mycol. Soc. 18: 157. 1933.

Basionym: Macrosporium tenuissimum (Nees & T. Nees) Fr., Syst. Mycol. (Lundae) 3: 374. 1832 (nom. sanct.).

• ≡ Helminthosporium tenuissimum Kunze ex Nees & T. Nees, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 9: 242. 1818.

Additional synonyms listed in Simmons (2007).

Alternaria toxicogenica E.G. Simmons, Mycotaxon 70: 294. 1999.

Alternaria turkisafria E.G. Simmons, Mycotaxon 70: 290. 1999.

Section Brassicicola D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 541. 2013. Fig. 5.

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Fig. 5.

Alternaria sect. Brassicicola: conidia and conidiophores. A, H. A. brassicicola. B, I, L-M. A. mimicola. C, G. A. solidaccana. D, J-K. A. conoidea. E-F. A. septorioides. Scale bars = 10 μm.

Type species: Alternaria brassicicola (Schwein.) Wiltshire

Diagnosis: Section Brassicicola contains short to moderately long, simple or branched primary conidiophores with one or several apical conidiogenous loci. Conidia are ellipsoid, ovoid or somewhat obclavate, small or moderate in size, septate, slightly or strongly constricted at most of their transverse septa, with no to many longitudinal septa, in moderately long to long, simple or branched chains, with dark septa and cell walls. Secondary conidiophores can be formed apically or laterally with one or a few conidiogenous loci. Chlamydospores may occur.

Notes: Our molecular data support the morphological placement of A. septorioides and A. solidaccana in section Brassicicola (Simmons 2007). The other three species were already assigned to this section based on previous molecular studies (Pryor et al. 2009, Runa et al. 2009, Lawrence et al. 2012). Alternaria japonica was previously linked to the A. brassicicola species-group (Pryor & Gilbertson 2000, Pryor & Bigelow 2003, Lawrence et al. 2013), but this association was questioned by Hong et al. (2005). In our analyses, A. japonica clustered in sect. Japonicae.

Alternaria brassicicola (Schwein.) Wiltshire, Mycol. Pap. 20: 8. 1947.

Basionym: Helminthosporium brassicicola Schwein., Trans. Amer. Philos. Soc., Ser. 2, 4: 279. 1832.

Additional synonyms listed in Simmons (2007)

Alternaria conoidea (E.G. Simmons) D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 542. 2013.

Basionym: Embellisia conoidea E.G. Simmons, Mycotaxon 17: 226. 1983.

Alternaria mimicula E.G. Simmons, Mycotaxon 55: 129. 1995.

Alternaria septorioides (Westend.) E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 570. 2007.

Basionym: Sporidesmium septorioides Westend., Bull. Acad. Roy. Sci. Belgique., Cl. Sci., Sér. 2, 21: 236. 1854.

• = Alternaria resedae Neerg., Annual Rep. Phytopathol. Lab. J.E. Ohlsens Enkes, Seed Growers, Copenhagen 7: 9. 1942 (nom. nud.).

• = Alternaria resedae Neerg., Danish species of Alternaria & Stemphylium: 150. 1945.

Alternaria solidaccana E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 572. 2007.

Section Chalastospora (E.G. Simmons) Woudenb. & Crous, comb. et stat. nov. MycoBank MB803733. Fig. 6.

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Fig. 6.

Alternaria sect. Chalastospora: conidia and conidiophores. A. A. cetera. B. A. obclavata. C. A. breviramosa. D, H. A. armoraciae. E-G. A. abundans. Scale bars = 10 μm.

Basionym: Chalastospora E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 668. 2007.

Type species: Alternaria cetera E.G. Simmons

Diagnosis: Section Chalastospora contains short to long, simple or branched primary conidiophores with one or several conidiogenous loci. Conidia are pale to medium brown, narrowly ellipsoid to ellipsoid or ovoid, beakless, with no to multiple transverse eusepta and rarely longitudinal septa, solitary or in chains. Secondary conidiophores can be formed apically or laterally with one or a few conidiogenous loci.

Notes: Previous studies already placed E. abundans in the Chalastospora-clade (Andersen et al. 2009, Lawrence et al. 2012). Our study also placed Alternaria armoraciae in this section, while Crous et al. (2009c) showed that Chalastospora gossypii, formerly Alternaria malorum, belonged to this section based on sequences of the ITS and LSU genes.

Alternaria abundans (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803688.

Basionym: Embellisia abundans E.G. Simmons, Mycotaxon 17: 222. 1983.

Alternaria armoraciae E.G. Simmons & C.F. Hill, CBS Biodiversity Ser. (Utrecht) 6: 660. 2007.

Alternaria breviramosa Woudenb. & Crous, nom. nov. MycoBank MB803690.

Basionym: Chalastospora ellipsoidea Crous & U. Braun, Persoonia 22: 145. 2009, non Alternaria ellipsoidea E.G. Simmons, 2002.

Etymology: Name refers to the short lateral branches.

Alternaria cetera E.G. Simmons, Mycotaxon 57: 393. 1996.

• ≡ Chalastospora cetera (E.G. Simmons) E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 668. 2007.

Alternaria malorum (Ruehle) U. Braun, Crous & Dugan, Mycol. Progr. 2: 5. 2003.

Basionym: Cladosporium malorum Ruehle, Phytopathology 21: 1146. 1931.

• = Cladosporium gossypii Jacz., Khlopkovoe Delo, 1929 (5-6): 564. 1929, non Alternaria gossypii (Jacz.) Y. Nisik., K. Kimura & Miyaw., 1940.

  • ≡ Chalastospora gossypii (Jacz.) U. Braun & Crous, Persoonia 22: 144. 2009.

• = Cladosporium malorum Heald, Wash. State Agric. Exp. Sta. Bull., Special Ser. 245: 48. 1930. (nom. nud.)

Additional synonyms in Crous et al. (2009c).

Alternaria obclavata (Crous & U. Braun) Woudenb. & Crous, comb. nov. MycoBank MB803689.

Basionym: Chalastospora obclavata Crous & U. Braun, Persoonia 22: 146. 2009.

Section Cheiranthus Woudenb. & Crous, sect. nov. MycoBank MB803734. Fig. 7.

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Fig. 7.

Alternaria sect. Cheiranthus: conidia and conidiophores. A-B. A. indefessa. B-C. A. cheiranthi. Scale bars = 10 μm.

Type species: Alternaria cheiranthi (Lib.) P.C. Bolle

Diagnosis: Section Cheiranthus contains short to moderately long, simple or branched primary conidiophores with one or several conidiogenous loci. Conidia are ovoid, broadly ellipsoid with transverse and longitudinal septa, slightly or strongly constricted at the septa, in short to long, simple or branched chains. Secondary conidiophores can be formed apically or laterally with a single conidiogenous locus.

Notes: Next to Alternaria cheiranthi and Embellisia indefessa, sect. Cheiranthus contains a non-sporulating strain formerly known as Alternaria resedae, CBS 115.44. Because Alternaria resedae is synonymised with Alternaria septorioides (Simmons 2007), which clusters in section Brassisicola, CBS 115.44 will be treated as “Alternaria sp.”. Alternaria cheiranthi and E. indefessa have been linked to Ulocladium (Pryor & Gilbertson 2000, Pryor & Bigelow 2003, Hong et al. 2005, Pryor et al. 2009, Runa et al. 2009, Lawrence et al. 2012), but based on morphology could not be placed here. Our extensive dataset showed that they form a sister section to section Ulocladioides.

Alternaria cheiranthi (Lib.) P.C. Bolle, Meded. Phytopathol. Lab. “Willie Commelin Scholten” 7: 43. 1924.

Basionym: Helminthosporium cheiranthi Lib. [as “Helmisporium”], in Desmazières, Plantes Cryptogames du Nord de la France, edn 1: 213. 1827.

• ≡ Macrosporium cheiranthi (Lib.) Fr., Syst. Mycol. (Lundae) 3: 374. 1832.

Alternaria indefessa (E.G. Simmons) Woudenberg & Crous, comb. nov. MycoBank MB803691.

Basionym: Embellisia indefessa E.G. Simmons, Mycotaxon 17: 228. 1983.

Section Crivellia (Shoemaker & Inderb.) Woudenb. & Crous, comb. et stat. nov. MycoBank MB803735. Fig. 8.

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Fig. 8.

Alternaria sect. Crivellia: conidia and conidiophores. A-B. A. papavericola. C-D. A. penicillata. Scale bars = 10 μm.

Basionym: Crivellia Shoemaker & Inderb., Canad. J. Bot. 84: 1308. 2006.

Type species: Alternaria penicillata (Corda) Woudenb. & Crous (= Cucurbitaria papaveracea De Not.).

Diagnosis: Section Crivellia is characterised by straight or curved, simple or branched primary conidiophores, with geniculate, sympodial proliferations. Conidia are cylindrical, straight to curved to inequilateral, with transverse eusepta, rarely constricted at septa, single or in short, simple or branched chains. Secondary conidiophores are formed apically or laterally. Microsclerotia or chlamydospores may occur. Sexual morphs observed.

Notes: Section Crivellia contains the type species of the sexual morph Crivellia, C. papaveracea, with Brachycladium penicillatum asexual morph, and Brachycladium papaveris. The genus was established by Inderbitzin et al. (2006) based on the finding that C. papaveraceae, formerly Pleospora papaveraceae, belonged to the Alternaria-complex instead of Pleospora s. str. based on ITS, GAPDH and TEF1 sequences.

Alternaria papavericola Woudenb. & Crous, nom. nov. MycoBank MB803749.

Basionym: Helminthosporium papaveris Sawada, J. Nat. Hist. Soc. Formosa 31: 1. 1917.

• ≡ Dendryphion papaveris (Sawada) Sawada, Special Publ. Coll. Agric. Natl. Taiwan Univ. 8: 200. 1959, non Alternaria papaveris (Bres.) M.B. Ellis, 1976.

• ≡ Brachycladium papaveris (Sawada) Shoemaker & Inderb., Canad. J. Bot. 84: 1310. 2006.

Etymology: Name refers to the host.

Alternaria penicillata (Corda) Woudenb. & Crous, comb. nov. MycoBank MB803692.

Basionym: Brachycladium penicillatum Corda, Icon. Fungorum hucusque Cogn. (Prague) 2: 14. 1838.

• • ≡ Dendryphion penicillatum (Corda) Fr., Summa Veg. Scand., Sect. Post. (Stockholm): 504. 1849.

• = Cucurbitaria papaveracea De Not., Sferiacei Italici: 62. 1863.

  • ≡ Pleospora papaveracea (De Not.) Sacc., Syll. Fungorum (Abellini) 2: 243. 1883.

  • ≡ Crivellia papaveracea (De Not.) Shoemaker & Inderb., Canad. J. Bot. 84: 1308. 2006.

Note: The asexual name, Brachycladium penicillatum is older than the sexual name, Cucurbitaria papaveracea, and therefore the species epithet penicillatum is chosen above papaveracea.

Section Dianthicola Woudenb. & Crous, sect. nov. MycoBank MB803736. Fig. 9.

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Fig. 9.

Alternaria sect. Dianthicola: conidia and conidiophores. A-B. A. dianthicola. C-E. A. simsimi. F-H. A. elegans. Scale bars = 10 μm.

Type species: Alternaria dianthicola Neerg.

Diagnosis: Section Dianthicola contains simple or branched primary conidiophores, with or without apical geniculate proliferations. Conidia are narrowly ovoid or narrowly ellipsoid with transverse and few longitudinal septa, slightly constricted at the septa, with a long (filamentous) beak or apical secondary conidiophore, solitary or in short chains.

Note: Based on the ITS sequence, Alternaria dianthicola clustered near Ulocladium (Chou & Wu 2002). Our extensive dataset places it in a sister section to section Ulocladioides.

Alternaria dianthicola Neerg., Danish species of Alternaria & Stemphylium: 190. 1945.

Alternaria elegans E.G. Simmons & J.C. David, Mycotaxon 75: 89. 2000.

Alternaria simsimi E.G. Simmons, Stud. Mycol. 50: 111. 2004.

Section Embellisia (E.G. Simmons) Woudenb. & Crous, comb. et stat. nov. MycoBank MB803737. Fig. 10.

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Fig. 10.

Alternaria sect. Embellisia: conidia and conidiophores. A-D. A. embellisia. E-H. A. tellustris. Scale bars = 10 μm.

Basionym: Embellisia E.G. Simmons, Mycologia 63: 380. 1971.

Type species: Alternaria embellisia Woudenb. & Crous (≡ Helminthosporium allii Campan., Embellisia allii (Campan.) E.G. Simmons).

Diagnosis: Section Embellisia contains simple, septate conidiophores, straight or with geniculate sympodial proliferation. Condia are solitary, ovoid to subcylindrical, straight to inequilateral, transseptate; septa can be thick, dark and rigid in contrast to the external wall. Chlamydospores may occur.

Notes: Section Embellisia contains the first two species described in the genus Embellisia, Embellisia allii (type species) and Embellisia chlamydospora (Simmons 1971) together with Embellisia tellustris. This clade is also resolved in the latest molecular revision of Embellisia based on sequences of the GAPDH, ITS and Alt a 1 genes as Embellisia group I (Lawrence et al. 2012).

Alternaria chlamydosporigena Woudenb. & Crous, nom. nov. MycoBank MB803694.

Basionym: Pseudostemphylium chlamydosporum Hoes, G.W. Bruehl & C.G. Shaw, Mycologia 57: 904. 1965, non Alternaria chlamydospora Mouch., 1973.

• ≡ Embellisia chlamydospora (Hoes, G.W. Bruehl & C.G. Shaw) E.G. Simmons, Mycologia 63: 384. 1971.

Etymology: Name refers to the formation of chlamydospores during growth.

Alternaria embellisia Woudenb. & Crous, nom. nov. MycoBank MB803693.

Basionym: Helminthosporium allii Campan., Nuovi Ann. Agric. Roma 4: 87. 1924, non Alternaria allii Nolla, 1927.

• ≡ Embellisia allii (Campan.) E.G. Simmons, Mycologia 63: 382. 1971.

Etymology: Name refers to the genus Embellisia for which it served as type species.

Alternaria tellustris (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803695.

Basionym: Embellisia tellustris E.G. Simmons [as “telluster”], Mycotaxon 17: 234. 1983.

Section Embellisioides Woudenb. & Crous, sect. nov. MycoBank MB803738. Fig. 11.

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Fig. 11.

Alternaria sect. Embellisioides: conidia and conidiophores. A-B. A. hyacinthi. C-E. A. lolii. F-H. A. botryospora. I-K. A. planifunda. L-N. A. proteae. O-P. A. tumida. Scale bars = 10 μm.

Type species: Alternaria hyacinthi (de Hoog & P.J. Mull. bis) Woudenb. & Crous

Diagnosis: Section Embellisioides contains simple, septate conidiophores, straight or with multiple, geniculate, sympodial proliferations. Apical or lateral, short secondary conidiophores may occur. Condia are solitary or in short chains, obovoid to ellipsoid, with transverse and longitudinal septa; transverse septa can be thick, dark and rigid in contrast to the external wall. Chlamydospores and a sexual morph may occur.

Note: In Lawrence et al. (2012) the section is named Embellisia group III.

Alternaria botryospora Woudenb. & Crous, nom. nov. MycoBank MB803705.

Basionym: Embellisia novae-zelandiae E.G. Simmons & C.F. Hill, Mycotaxon 38: 252. 1990, non Alternaria novae-zelandiae E.G. Simmons, 2002.

Etymology: Name refers to the clusters of conidia.

Alternaria hyacinthi (de Hoog & P.J. Mull. bis) Woudenb. & Crous, comb. nov. MycoBank MB803703.

Basionym: Embellisia hyacinthi de Hoog & P.J. Mull. bis, Netherlands J. Pl. Pathol. 79: 85. 1973.

Alternaria lolii (E.G. Simmons & C.F. Hill) Woudenb. & Crous, comb. nov. MycoBank MB803704.

Basionym: Embellisia lolii E.G. Simmons & C.F. Hill, Stud. Mycol. 50: 113. 2004.

Alternaria planifunda (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803706.

Basionym: Embellisia planifunda E.G. Simmons, Mycotaxon 17: 233. 1983.

Alternaria proteae (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803707.

Basionym: Embellisia proteae E.G. Simmons, Mycotaxon 38: 258. 1990.

• = Allewia proteae E.G. Simmons, Mycotaxon 38: 262. 1990.

Alternaria tumida (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803708.

Basionym: Embellisia tumida E.G. Simmons, Mycotaxon 17: 236. 1983.

Section Eureka Woudenb. & Crous, sect. nov. MycoBank MB803739. Fig. 12.

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Fig. 12.

Alternaria sect. Eureka: conidia and conidiophores. A-B. A. anigozanthi. C-D. A. cumini. E-F. A. leptinellae. G-H. A. triglochinicola. I-J. A. geniostomatis. K-L. A. eureka. Scale bars = 10 μm.

Type species: Alternaria eureka E.G. Simmons

Diagnosis: Section Eureka contains simple, septate conidiophores, straight or with geniculate, sympodial proliferations. Apical or lateral, short secondary conidiophores may occur. Condia are solitary or in short chains, narrowly ellipsoid to cylindrical, with transverse and longitudinal septa, slighty constricted at the septa, with a blunt rounded apex. Chlamydospores and a sexual morph may occur.

Notes: Section Eureka contains four Alternaria species and two former Embellisia species. From the Alternaria species only the ITS sequence of A. geniostomatis was previously used in a molecular study (Toth et al. 2011), showing it to cluster separate from the other Alternaria spp. The two Embellisia species were included in the latest molecular-based revision of Embellisia (Lawrence et al. 2012) where they formed Embellisia group IV. A sexual morph is known for the type species of this section.

Alternaria anigozanthi Priest, Australas. Pl. Pathol. 24: 239. 1995.

Alternaria cumini E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 664. 2007.

Alternaria eureka E.G. Simmons, Mycotaxon 25: 306. 1986.

• • ≡ Embellisia eureka (E.G. Simmons) E.G. Simmons, Mycotaxon 38: 260. 1990.

• = Lewia eureka E.G. Simmons, Mycotaxon 25: 304. 1986.

  • ≡ Allewia eureka (E.G. Simmons) E.G. Simmons, Mycotaxon 38: 264. 1990.

Alternaria geniostomatis E.G. Simmons & C.F. Hill, CBS Biodiversity Ser. (Utrecht) 6: 412. 2007.

Alternaria leptinellae (E.G. Simmons & C.F. Hill) Woudenb. & Crous, comb. nov. MycoBank MB803696.

Basionym: Embellisia leptinellae E.G. Simmons & C.F. Hill, Mycotaxon 38: 254. 1990.

Alternaria triglochinicola Alcorn & S.M. Francis, Mycotaxon 46: 359. 1993.

Section Gypsophilae D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 541. 2013. Fig. 13

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Fig. 13.

Alternaria sect. Gypsophilae: conidia and conidiophores. A-B. A. axiariisporifera. C-D. A. ellipsoidea. E-G. A. saponariae. H-I. A. vaccariae. J-K. A. nobilis. L-M. A. juxtiseptata. N-P. A. vaccariicola. Scale bars = 10 μm.

Type species: Alternaria gypsophilae Neerg.

Diagnosis: Section Gypsophilae contains simple, or occasionally branched, primary conidiophores, with one or a few conidiogenous loci. Conidia are ellipsoid to long ovoid, with multiple transverse and longitudinal septa, conspicuously constricted near some transverse septa, solitary or in short chains. Secondary conidiophores are formed apically with one or two conidiogenous loci or laterally with a single conidiogenous locus. Species from this section occur on Caryophyllaceae.

Notes: Section Gypsophilae was recently established by Lawrence et al. (2013) containing the four Alternaria species, A. gypsophilae, A. nobilis, A. vaccariae and A. vaccariicola. Our dataset adds four Alternaria species, A. axiaeriisporifera, A. ellipsoidea, A. saponariae, and A. juxtiseptata to this section. Simmons (2007) noted the similarity of the primary conidia of A. ellipsoidea to A. gypsophilae, A. nobilis, A. saponariae and A. vaccariae. This section contains all Alternaria species that occur on Caryophyllaceae (Simmons 2002), except A. dianthicola which resides in sect. Dianthicola.

Alternaria axiaeriisporifera E.G. Simmons & C.F. Hill, CBS Biodiversity Ser. (Utrecht) 6: 662. 2007.

Alternaria ellipsoidea E.G. Simmons, Mycotaxon 82: 31. 2002.

Alternaria gypsophilae Neerg., Danish species of Alternaria & Stemphylium: 207. 1945.

Alternaria juxtiseptata E.G. Simmons, Mycotaxon 82: 32. 2002.

Alternaria nobilis (Vize) E.G. Simmons, Mycotaxon 82: 7. 2002.

Basionym: Macrosporium nobile Vize, Grevillea 5(35): 119. 1877.

Alternaria saponariae (Peck) Neerg., Annual Rep. Phytopathol. Lab. J.E. Ohlsens Enkes, Seed Growers, Copenhagen 3: 6. 1938 [1937-1938].

Basionym: Macrosporium saponariae Peck, Rep. (Annual) NewYork State Mus. Nat. Hist. 28: 62. 1876 [1875].

Alternaria vaccariae (Săvul. & Sandu) E.G. Simmons & S.T. Koike, Mycotaxon 82: 21. 2002.

Basionym: Macrosporium vaccariae Săvul. & Sandu, Hedwigia 73: 130. 1933.

Alternaria vaccariicola E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 594. 2007.

Section Infectoriae Woudenb. & Crous, sect. nov. MycoBank MB803740. Fig. 14.

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Fig. 14.

Alternaria sect. Infectoriae: conidia and conidiophores. A-B. A. ethzedia. C-D. A. infectoria. E-F. A. conjuncta. G-H. A. oregonensis. Scale bars = 10 μm.

Type species: Alternaria infectoria E.G. Simmons

Diagnosis: Section Infectoriae contains short to long, simple or branched primary conidiophores with one or several conidiogenous loci. Conidia are obclavate, long-ellipsoid, small or moderate in size, septate, slightly constricted near some septa, with few longitudinal septa, in moderately long to long, branched chains. Long, geniculate, multi-locus secondary conidiophores can be formed apically or laterally. Sexual morphs are known, and meristematic growth has been reported.

Notes: In addition to the six species that are displayed in our phylogeny, 19 more are included based on the study of Lawrence et al. (2013), confirmed with our molecular data (not shown). From these 25 species, nine species have a known sexual morph in Lewia. Three species from the study of Lawrence et al. (2013) are not included; A. photistica (sect. Panax) and A. dianthicola (sect. Dianthicola) cluster elsewhere in our phylogenies and A. peglionii is marked as a taxon incertae sedis by Simmons (2007). The human pathogenic genera Ybotromyces and Chmelia are also embedded in sect. Infectoriae.

Alternaria alternarina E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 644. 2007.

• = Pyrenophora alternarina M.D. Whitehead & J. Dicks., Mycologia 44: 748. 1952.

  • ≡ Lewia alternarina (M.D. Whitehead & J.G. Dicks.) E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 644. 2007.

Alternaria arbusti E.G. Simmons, Mycotaxon 48: 103. 1993.

Alternaria caespitosa (de Hoog & C. Rubio) Woudenb. & Crous, comb. nov. MycoBank MB803698.

Basionym: Botryomyces caespitosus de Hoog & C. Rubio, Mycotaxon 14: 19. 1982.

• ≡ Ybotromyces caespitosus (de Hoog & C. Rubio) Rulamort, Bull. Soc. Bot. Centre-Ouest, Nouv. Sér. 21: 512. 1990.

Alternaria californica E.G. Simmons & S.T. Koike, CBS Biodiversity Ser. (Utrecht) 6: 602. 2007.

Alternaria conjuncta E.G. Simmons, Mycotaxon 25: 294. 1986.

• = Sphaeria scrophulariae Desm., Ann. Sci. Nat., Bot., Sér. 2, 6: 245. 1836.

  • ≡ Leptosphaeria scrophulariae (Desm.) Sacc., Syll. Fungorum (Abellini) 2: 57. 1883.

  • ≡ Heptameria scrophulariae (Desm.) Cooke, Grevillea 18(no. 86): 31. 1889.

  • ≡ Pleospora scrophulariae (Desm.) Höhn., Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1. 126(4-5): 374. 1917.

  • ≡ Lewia scrophulariae (Desm.) M.E. Barr & E.G. Simmons, Mycotaxon 25: 294. 1986.

Alternaria daucicaulis E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 640. 2007.

• = Lewia daucicaulis E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 640. 2007.

Alternaria ethzedia E.G. Simmons, Mycotaxon 25: 300. 1986.

• = Lewia ethzedia E.G. Simmons, Mycotaxon 25: 299. 1986.

Alternaria frumenti E.G. Simmons & C.F. Hill, CBS Biodiversity Ser. (Utrecht) 6: 620. 2007.

Alternaria graminicola E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 626. 2007.

Alternaria hordeiaustralica E.G. Simmons & Alcorn, CBS Biodiversity Ser. (Utrecht) 6: 614. 2007.

• = Lewia hordeiaustralica E.G. Simmons & Alcorn, CBS Biodiversity Ser. (Utrecht) 6: 614. 2007.

Alternaria hordeicola E.G. Simmons & Kosiak, CBS Biodiversity Ser. (Utrecht) 6: 630. 2007.

• = Lewia hordeicola Kwaśna & Kosiak, Mycologia 98: 663. 2006.

Alternaria humuli E.G. Simmons, Mycotaxon 83: 139. 2002.

Alternaria incomplexa E.G. Simmons, Mycotaxon 57: 394. 1996.

Alternaria infectoria E.G. Simmons, Mycotaxon 25: 298. 1986.

• = Pleospora infectoria Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 132. 1870 [1869-70].

  • ≡ Sphaeria infectoria (Fuckel) Cooke, Handb. Brit. Fungi 2: 897. 1871.

  • ≡ Pleospora phaeocomoides var. infectoria (Fuckel) Wehm., A World Monograph of the Genus Pleospora and its Segregates: 121. 1961.

  • ≡ Lewia infectoria (Fuckel) M.E. Barr & E.G. Simmons, Mycotaxon 25: 296. 1986.

Alternaria intercepta E.G. Simmons, Mycotaxon 83: 134. 2002.

• = Lewia intercepta E.G. Simmons & McKemy, Mycotaxon 83: 133. 2002.

Alternaria merytae E.G. Simmons, Mycotaxon 83: 136. 2002.

Alternaria metachromatica E.G. Simmons, Mycotaxon 50: 418. 1994.

Alternaria novae-zelandiae E.G. Simmons, Mycotaxon 83: 142. 2002.

Alternaria oregonensis E.G. Simmons, Mycotaxon 50: 417. 1994.

Alternaria slovaca (Svob.-Pol., L. Chmel & Bojan.) Woudenb. & Crous, comb. nov. MycoBank MB803699.

Basionym: Aureobasidium slovacum Svob.-Pol., L. Chmel & Bojan., Conspect. Verruc. 5: 116. 1966.

• ≡ Chmelia slovaca (Svob.-Pol., L. Chmel & Bojan.) Svob.-Pol., Biologia (Bratislava) 21: 83. 1966.

Alternaria triticimaculans E.G. Simmons & Perelló, Mycotaxon 50: 413. 1994.

Alternaria triticina Prasada & Prabhu, Indian Phytopathol. 15 (3-4): 292. 1963. [1962]

Alternaria ventricosa R.G. Roberts, Mycotaxon 100: 164. 2007.

Alternaria viburni E.G. Simmons, Mycotaxon 83: 132. 2002.

• = Lewia viburni E.G. Simmons & McKemy, Mycotaxon 83: 130. 2002.

Section Japonicae Woudenb. & Crous, sect. nov. MycoBank MB803741. Fig. 15.

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Fig. 15.

Alternaria sect. Japonicae: conidia and conidiophores. A-B. A. japonica. C-E. A. nepalensis. Scale bars = 10 μm.

Type species: Alternaria japonica Yoshii

Diagnosis: Section Japonicae contains short to long, simple or occasionally branched primary conidiophores with a single conidiogenous locus. Conidia are short, to long-ovoid with transverse and longitudinal septa, conspicuously constricted at most of the transverse septa, in short chains. Apical secondary conidiophores are produced with a single conidiogenous locus. The species within this section occur on Brassicaceae.

Note: Alternaria japonica was previously connected to the A. brassicicola species-group (Pryor & Gilbertson 2000, Pryor & Bigelow 2003, Lawrence et al. 2013), but this association was questioned by Hong et al. (2005).

Alternaria japonica Yoshii, J. Pl. Protect. 28: 17. 1941.

• = Alternaria matthiolae Neerg., Danish species of Alternaria and Stemphylium: 184. 1945.

Alternaria nepalensis E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 480. 2007.

Section Nimbya (E.G. Simmons) Woudenb. & Crous, comb. et stat. nov. MycoBank MB803742. Fig. 16.

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Fig. 16.

Alternaria sect. Nimbya: conidia and conidiophores. A-B. A. caricis. C-D. A. scirpicola. Scale bars = 10 μm.

Basionym: Nimbya E.G. Simmons, Sydowia 41: 316. 1989.

Type species: Alternaria scirpicola (Fuckel) Sivan.

Diagnosis: Section Nimbya contains simple, short to moderately long conidiophores, which may form one or a few short to long, geniculate, sympodial proliferations. Conidia are narrowly elongate-obclavate, gradually tapering apically, solitary or in short chains, with transverse disto- and eusepta, sometimes slightly constricted near eusepta. Apical condiophores with a single conidiogenous locus can be formed. Internal compartmentation occurs, cell lumina tend to be broadly octagonal to rounded. A sexual morph may occur.

Notes: Section Nimbya contains the type species of Nimbya, N. scirpicola, and N. caricis (Simmons 1989). A more extensive study on Nimbya (Lawrence et al. 2012) found that N. scirpinfestans and N. scirpivora also belonged to this section based on sequences of the GAPDH, ITS and Alt a 1 genes.

Alternaria caricis (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803700.

Basionym: Nimbya caricis E.G. Simmons, Sydowia 41: 328. 1989.

Alternaria scirpicola (Fuckel) Sivan., Bitunicate Ascomycetes and their Anamorphs (Vaduz): 526. 1984.

Basionym: Sporidesmium scirpicola Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 140. 1870 [1869-70].

• • ≡ Clasterosporium scirpicola (Fuckel) Sacc., Syll. Fungorum (Abellini) 4: 393. 1886.

  • ≡ Cercospora scirpicola (Fuckel) Zind.-Bakker, Rev. Mycol. (Paris) 5: 66. 1940.

  • ≡ Alternaria scirpicola (Fuckel) M.T. Lucas & J. Webster, Čas. Slez. Mus., Ser. A, Hist. Nat. 23: 151. 1974 (nom. inval.).

  • ≡ Nimbya scirpicola (Fuckel) E.G. Simmons, Sydowia 41: 316. 1989.

• = Sphaeria scirpicola DC., in Lamarck & de Candolle, Fl. Franç., Edn 3 (Paris) 2: 300. 1805.

  • ≡ Clathrospora scirpicola (DC.) Höhn., Ann. Mycol. 18(1/3): 77. 1920.

  • ≡ Macrospora scirpicola (DC.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 139. 1870 [1869-70].

  • ≡ Pyrenophora scirpicola (DC.) E. Müll., Sydowia 5(3-6): 256. 1951.

Note: Although Sphaeria scirpicola DC. (de Candolle 1805) predates Sporidesmium scirpicola Fuckel (Fuckel 1870), a valid combination in Alternaria already exists, thus we choose to retain Alternaria scirpicola (Fuckel) Sivan., which is also a well established name.

Alternaria scirpinfestans (E.G. Simmons & D.A. Johnson) Woudenb. & Crous, comb. nov. MycoBank MB803701.

Basionym: Nimbya scirpinfestans E.G. Simmons & D.A. Johnson, Mycotaxon 84: 420. 2002.

• = Macrospora scirpinfestans E.G. Simmons & D.A. Johnson, Mycotaxon 84: 417. 2002.

Alternaria scirpivora (E.G. Simmons & D.A. Johnson), Woudenb. & Crous, comb. nov. MycoBank MB803702.

Basionym: Nimbya scirpivora E.G. Simmons & D.A. Johnson, Mycotaxon 84: 424. 2002.

• = Macrospora scirpivora E.G. Simmons & D.A. Johnson, Mycotaxon 84: 422. 2002.

Section Panax D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 541. 2013. Fig. 17.

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Fig. 17.

Alternaria sect. Panax: conidia and conidiophores. A-B. A. avenicola. C-D. A. calycipyricola. E-F. A. panax. G-H. A. photistica. Scale bars = 10 μm.

Type species: Alternaria panax Whetzel

Diagnosis: Section Panax contains simple or branched, short to moderately long primary conidiophores, with one or a few conidiogenous loci. Conidia are obclavate to ovoid, with multiple transverse and longitudinal septa, conspicuously constricted near several transverse septa, solitary or in simple or branched, short chains. Apical secondary conidiophores are formed with one or several conidiogenous loci, multiple lateral secondary conidiophores with a single conidiogenous locus may occur.

Notes: Section Panax was recently described by Lawrence et al. (2013) and consists of A. calycipyricola, A. eryngii and A. panax. Our extended dataset added the species A. avenicola and A. photistica to this section. Three species, A. avenicola, A. calycipyricola, and A. photistica have earlier been placed in the A. infectoria species-group based on their morphological characters (Simmons 2007), and two of them have a known sexual morph; Lewia avenicola (Simmons 2007) and Lewia photistica (Simmons 1986). A phylogenetic study based on Alt a 1 and GAPDH sequences placed A. photistica in the A. infectoria species-group (Hong et al. 2005) but an extensive study on the A. infectoria species-group (Andersen et al. 2009) confirmed our finding, and placed this species outside the A. infectoria species-group. Additional research performed on multiple A. photistica strains support our sequence data (data not shown).

Alternaria avenicola E.G. Simmons, Kosiak & Kwaśna, in Simmons, CBS Biodiversity Ser. (Utrecht) 6: 114. 2007.

• = Lewia avenicola Kosiak & Kwaśna, Mycol. Res. 107: 371. 2003.

Alternaria calycipyricola R.G. Roberts, Mycotaxon 100: 162. 2007.

Alternaria eryngii (Pers.) S. Hughes & E.G. Simmons, Canad. J. Bot. 36: 735. 1958.

Basionym: Conoplea eryngii Pers., Mycol. Eur. (Erlanga) 1: 11. 1822.

• ≡ Exosporium eryngianum (Pers.) Chevall., Flore Générale des Environs de Paris 1: 39. 1826.

• ≡ Exosporium eryngii (Pers.) Duby, Bot. Gallicum., Edn 2 (Paris) 2: 882. 1830.

• ≡ Helminthosporium eryngii (Pers.) Fr., Syst. Mycol. (Lundae) 3: 361. 1832.

Alternaria panax Whetzel, Bull. U.S.D.A. 250: 11. 1912.

• = Macrosporium araliae Dearn. & House, Circ. New York State Mus. 24: 58. 1940.

• = Alternaria araliae H.C. Greene, Trans. Wisconsin Acad. Sci. 42: 80. 1953.

Alternaria photistica E.G. Simmons, Mycotaxon 25: 304. 1986.

• = Lewia photistica E.G. Simmons, Mycotaxon 25: 302. 1986.

Section Phragmosporae Woudenb. & Crous, sect. nov. MycoBank MB803743. Fig. 18.

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Fig. 18.

Alternaria sect. Phragmosporae: conidia and conidiophores. A-B. A. didymospora. C. A. phragmospora. D-E. A. limaciformis. F-G. A. molesta. H-I. A. mouchaccae. Scale bars = 10 μm.

Type species: Alternaria phragmospora Emden

Diagnosis: Section Phragmosporae contains simple, short to moderately long, primary conidiophores, with one or multiple geniculate, sympodial proliferations. Conidia are (broad) ovoid to long ovoid, ellipsoid, curved, or limaciform, with multiple transverse and few to multiple longitudinal septa, some septa darkened, slightly to conspicuously constricted near several transverse septa, solitary or in simple short chains. Apical secondary conidiophores are formed with one or several conidiogenous loci. All species within the section are known from soil and seawater environments.

Note: Section Phragmosporae contains six species of which two were linked to Embellisia.

Alternaria chlamydospora Mouch. [as “chlamydosporum”], Mycopathol. Mycol. Appl. 50: 217. 1973.

Alternaria didymospora (Munt.-Cvetk.) Woudenb. & Crous, comb. nov. MycoBank MB803709.

Basionym: Embellisia didymospora Munt.-Cvetk., Mycologia 68: 49. 1976.

Alternaria limaciformis E.G. Simmons, Mycotaxon 13: 24. 1981.

Alternaria molesta E.G. Simmons, Mycotaxon 13: 17. 1981.

Alternaria mouchaccae E.G. Simmons, Mycotaxon 13: 18. 1981.

• ≡ Ulocladium chlamydosporum Mouch., Rev. Mycol. (Paris) 36: 114. 1971, non Alternaria chlamydospora Mouch., 1973.

Alternaria phragmospora Emden, Acta Bot. Neerl. 19: 393. 1970.

• ≡ Embellisia phragmospora (Emden) E.G. Simmons, Mycotaxon 17: 232. 1983.

Section Porri D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 541. 2013. Fig. 19

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Fig. 19.

Alternaria sect. Porri: conidia and conidiophores. A-C. A. daucii. D-F. A. pseudorostrata. G-H. A. solani. Scale bars = 10 μm.

Type species: Alternaria porri (Ellis) Cif.

Diagnosis: Section Porri is characterised by broadly ovoid, obclavate, ellipsoid, subcylindrical or obovoid (medium) large conidia, disto- and euseptate, solitary or in short to moderately long chains, with a simple or branched, long to filamentous beak. Conidia contain multiple transverse and longitudinal septa and are slightly constricted near some transverse septa. Secondary conidiophores can be formed apically or laterally.

Notes: In addition to the six species that are displayed in our phylogeny, 40 more are included based on the study of Lawrence et al. (2013), confirmed with own molecular data (not shown). With almost 80 species section Porri is the largest Alternaria section (data not shown). The section displays a higher level of genetic variation than the second largest section; section Alternata.

Alternaria acalyphicola E.G. Simmons, Mycotaxon 50: 260. 1994.

Alternaria agerati Sawada ex E.G. Simmons, Mycotaxon 65: 63. 1997.

• = Alternaria agerati Sawada, Rep. Dept. Agric. Gov. Res. Inst. Formosa 86: 165. 1943. (nom. inval., Art. 36.1)

Alternaria agripestis E.G. Simmons & K. Mort., Mycotaxon 50: 255. 1994.

Alternaria anagallidis A. Raabe, Hedwigia 78: 87. 1939.

Alternaria aragakii E.G. Simmons, Mycotaxon 46: 181. 1993.

Alternaria argyroxiphii E.G. Simmons & Aragaki, Mycotaxon 65: 40. 1997.

Alternaria bataticola Ikata ex W. Yamam., Trans. Mycol. Soc. Japan 2(5): 89. 1960.

• = Macrosporium bataticola Ikata, Agric. Hort. (Tokyo) 22: 241. 1947 (nom. inval., Art. 36.1).

Alternaria blumeae E.G. Simmons & Sontirat, Mycotaxon 65: 81. 1997.

Alternaria calendulae Ondřej, Čas. Slez. Mus. v Opave˘, Ser. A, Hist. Nat. 23(2): 150. 1974.

• = Alternaria calendulae W. Yamam. 1939 (nom. nud.).

• = Macrosporium calendulae Nelen, Bull. Centr. Bot. Gard. (Moscow) 35: 90. 1959 (nom. inval., Art. 36.1).

• = Macrosporium calendulae Nelen, Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Akad. Nauk S.S.S.R. 15: 144. 1962.

• = Alternaria calendulae Nirenberg, Phytopathol. Z. 88(2): 108. 1977 (nom. illegit., Art. 53.1).

Alternaria capsici E.G. Simmons, Mycotaxon 75: 84. 2000.

Alternaria carthami S. Chowdhury, J. Indian Bot. Soc. 23: 65. 1944.

• = Macrosporium anatolicum A. Săvul., Bull. Sect. Sci. Acad. Roumaine 26: 709. 1944.

Alternaria cassiae Jurair & A. Khan, Pakistan J. Sci. Industr. Res. 3(1): 72. 1960.

Alternaria cichorii Nattrass, First List of Cyprus Fungi: 29. 1937.

• ≡ Alternaria porri f.sp. cichorii (Natrass) T. Schmidt, Pflanzenschutz-berichte 32: 181. 1965.

• ≡ Macrosporium cichorii (Nattrass) Gordenko, Mikol. Fitopatol. 9(3): 241. 1975.

Alternaria cirsinoxia E.G. Simmons & K. Mort., Mycotaxon 65: 72. 1997.

Alternaria crassa (Sacc.) Rands, Phytopathology 7: 337. 1917.

Basionym: Cercospora crassa Sacc., Michelia 1(no. 1): 88. 1877.

Alternaria cretica E.G. Simmons & Vakal., Mycotaxon 75: 64. 2000.

Alternaria cucumerina (Ellis & Everh.) J.A. Elliott, Amer. J. Bot. 4: 472. 1917.

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

Alternaria cyphomandrae E.G. Simmons, Mycotaxon 75: 86. 2000.

Alternaria danida E.G. Simmons, Mycotaxon 65: 78. 1997.

Alternaria dauci (J.G. Kühn) J.W. Groves & Skolko, Canad. J. Res., Sect. C, Bot. Sci. 22: 222. 1944.

Basionym: Sporidesmium exitiosum var. dauci J.G. Kühn, Hedwigia 1: 91. 1855.

Additional synonyms in Simmons 2007.

Alternaria dichondrae Gambogi, Vannacci & Triolo, Trans. Brit. Mycol. Soc. 65(2): 323. 1975.

Alternaria euphorbiicola E.G. Simmons & Engelhard, Mycotaxon 25: 196. 1986.

• ≡ Macrosporium euphorbiae Reichert, Bot. Jahrb. Syst. 56: 723. 1921. (nom. illegit., Art 53.1).

Alternaria grandis E.G. Simmons, Mycotaxon 75: 96. 2000.

Alternaria hawaiiensis E.G. Simmons, Mycotaxon 46: 184. 1993.

Alternaria limicola E.G. Simmons & M.E. Palm, Mycotaxon 37: 82. 1990.

Alternaria linicola J.W. Groves & Skolko, Canad. J. Res., Sect. C, Bot. Sci. 22: 223. 1944.

Alternaria macrospora Zimm., Ber. Land-Forstw. Deutsch-Ostafrika 2: 24. 1904.

• • ≡ Macrosporium macrosporum (Zimm.) Nishikado & Oshima, Agric. Res. (Kurashiki) 36: 391. 1944.

• = Sporidesmium longipedicellatum Reichert, Bot. Jahrb. Syst. 56: 723. 1921.

  • ≡ Alternaria longipedicellata (Reichert) Snowden, Rep. Dept. Agric. Uganda: 31. 1927 [1926].

Alternaria multirostrata E.G. Simmons & C.R. Jacks., Phytopathology 58: 1139. 1968.

Alternaria nitrimali E.G. Simmons & M.E. Palm, Mycotaxon 75: 93. 2000.

Alternaria passiflorae J.H. Simmonds, Proc. Roy. Soc. Queensland. 49: 151. 1938.

Alternaria poonensis Ragunath, Mycopathol. Mycol. Appl. 21: 315. 1963.

Alternaria porri (Ellis) Cif., J. Dept. Agric. Porto Rico 14: 30. 1930 [1929].

Basionym: Macrosporium porri Ellis, Grevillea 8 (no. 45): 12. 1879.

Alternaria protenta E.G. Simmons, Mycotaxon 25: 207. 1986.

Alternaria pseudorostrata E.G. Simmons, Mycotaxon 57: 398. 1996.

Alternaria ricini (Yoshii) Hansf., Proc. Linn. Soc. Lond.: 53. 1943.

Basionym: Macrosporium ricini Yoshii, Bult. Sci. Fak. Terk. Kjusu Imp. Univ. 3(4): 327. 1929.

Alternaria rostellata E.G. Simmons, Mycotaxon 57: 401. 1996.

Alternaria scorzonerae (Aderh.) Loer., Netherlands J. Pl. Pathol. 90(1): 37. 1984.

Basionym: Sporidesmium scorzonerae Aderh.,Arbeiten Kaiserl. Biol. Anst. Land-Forstw. 3: 439. 1903.

Alternaria sesami (E. Kawam.) Mohanty & Behera, Curr. Sci. 27: 493. 1958.

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

Alternaria solani Sorauer, Z. Pflanzenkrankh. Pflanzenschutz 6: 6. 1896.

• = Macrosporium solani Ellis & G. Martin, Amer. Naturalist 16(12): 1003. 1882

  • ≡ Alternaria solani (Ellis & G. Martin) L.R. Jones & Grout, Vermont Agric. Exp. Sta. Annual Rep. 9: 86. 1896.

Additional synonyms in Simmons (2007).

Alternaria solani-nigri R. Dubey, S.K. Singh & Kamal [as “solani-nigrii”], Microbiol. Res. 154(2): 120. 1999.

Alternaria steviae Ishiba, T. Yokoy. & Tani, Ann. Phytopathol. Soc. Japan 48(1): 46. 1982.

Alternaria subcylindrica E.G. Simmons & R.G. Roberts, Mycotaxon 75: 62. 2000.

Alternaria tagetica S.K. Shome & Mustafee, Curr. Sci. 35: 370. 1966.

Alternaria tomatophila E.G. Simmons, Mycotaxon 75: 53. 2000.

Alternaria tropica E.G. Simmons, Mycotaxon 46: 187. 1993.

Alternaria zinniae H.Pape ex M.B. Ellis, Mycol. Pap. 131: 22. 1972.

• = Alternaria zinniae H. Pape, Angew. Bot. 24: 61. 1942. (nom. inval., Art. 36.1)

Section Pseudoulocladium Woudenb. & Crous, sect. nov. MycoBank MB803744. Fig. 20.

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Fig. 20.

Alternaria sect. Pseudoulocladium: conidia and conidiophores. A-B. A. aspera. C-D. A. concatenata. E-F. A. chartarum. G-H. A. septospora. Scale bars = 10 μm.

Type species: Alternaria chartarum Preuss

Diagnosis: Section Pseudoulocladium is characterised by simple or branched conidiophores with short, geniculate, sympodial proliferations. Conidia are obovoid, non-beaked with a narrow base, in simple or (mostly) branched chains. Apical secondary conidiophores with multiple conidiogenous loci and lateral secondary conidiophores with a single conidiogenous locus can be formed.

Note: It forms a sister clade to section Ulocladioides.

Alternaria aspera Woudenb. & Crous, nom. nov. MycoBank MB803712.

Basionym: Ulocladium arborescens E.G. Simmons, Stud. Mycol. 50: 117. 2004, non Alternaria arborescens E.G. Simmons, 1999.

Etymology: Name refers to the conspicuously ornamented conidia.

Alternaria chartarum Preuss, Bot. Zeitung 6: 412, 1848.

• • ≡ Sporidesmium polymorphum var. chartarum (Preuss) Cooke, Fungi Brit. Exs., ser. 2: 329. 1875.

  • ≡ Ulocladium chartarum (Preuss) E.G. Simmons, Mycologia 59: 88. 1967.

• = Alternaria stemphylioides Bliss, Mycologia 36: 538. 1944.

  • ≡ Alternaria chartarum f. stemphylioides (Bliss) P. Joly, Encycl. Mycol. (Paris) 33: 161. 1964.

Alternaria concatenata Woudenb. & Crous, nom. nov. MycoBank MB803713.

Basionym: Ulocladium capsici F. Xue & X.G. Zhang [as “capsicuma”], Sydowia 59: 174. 2007, non Alternaria capsici E.G. Simmons, 2000.

Eymology: Name refers to the concatenated conidia.

Alternaria septospora (Preuss) Woudenb. & Crous, comb. nov. MycoBank MB803714.

Basionym: Helminthosporium septosporum Preuss, Linnaea 24: 117. 1851.

• ≡ Macrosporium septosporum (Preuss) Rabenh., Bot. Zeitung 9: 454. 1851.

• ≡ Ulocladium septosporum (Preuss) E.G. Simmons, Mycologia 59: 87. 1967.

Section Radicina D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 541. 2013. Fig. 21.

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Fig. 21.

Alternaria sect. Radicina: conidia and conidiophores. A-C. A. carotiincultae. D-E. A. petroselini. F-G. A. radicina. H-I. A. selini. J-L. A. smyrnii. Scale bars = 10 μm.

Type species: Alternaria radicina Meier, Drechsler & E.D. Eddy

Diagnosis: Section Radicina contains straight, simple or branched, short or long, primary conidiophores with multiple, short geniculate, sympodial proliferations with single or a few conidiogenous loci at the apex. Sporulation resembles a cluster or clumps of conidia. Conidia are widely ovoid to narrowly ellipsoid, moderate in size, beakless, with several transverse and longitudinal septa, solitary or in short chains. Solitary, short, apical secondary conidiophores may occur. The species from this section occur on Umbelliferae.

Note: This section was first recognised by Pryor & Gilbertson (2000) based on sequence data of the ITS and mitochondrial SSU.

Alternaria carotiincultae E.G. Simmons, Mycotaxon 55: 103. 1995.

Alternaria petroselini (Neerg.) E.G. Simmons, More dematiaceous hyphomycetes (Kew): 417. 1976.

Basionym: Stemphylium petroselini Neerg., Zentralbl. Bakteriol., 2. Abt., 104: 411. 1942.

• ≡ Stemphylium radicinum var. petroselini (Neerg.) Neerg., Danish species of Alternaria & Stemphylium: 357. 1945.

• ≡ Alternaria radicina var. petroselini (Neerg.) Neerg., Encycl. Mycol. 33: 123. 1964.

Alternaria radicina Meier, Drechsler & E.D. Eddy, Phytopathology 12: 157. 1922.

• ≡ Stemphylium radicinum (Meier, Drechsler & E.D. Eddy) Neerg., Annual Rep. Phytopathol. Lab. J.E. Ohlsens Enkes, Seed Growers, Copenhagen 4: 14. 1939.

• ≡ Thyrospora radicina (Meier, Drechsler & E.D. Eddy) Neerg., Bot. Tidsskr. 44: 361. 1939.

• ≡ Pseudostemphylium radicinum (Meier, Drechsler & E.D. Eddy) Subram., Curr. Sci. 30: 423. 1961.

Alternaria selini E.G. Simmons, Mycotaxon 55: 109. 1995.

Alternaria smyrnii (P. Crouan & H. Crouan) E.G. Simmons, Mycotaxon 55: 41. 1995.

Basionym: Helminthosporium smyrnii P. Crouan & H. Crouan, Florule Finistère (Paris): 11. 1867.

• ≡ Macrosporium smyrnii (P. Crouan & H. Crouan) Sacc., Syll. Fungorum (Abellini) 4: 527. 1886.

Section Sonchi D.P. Lawr., Gannibal, Peever & B.M. Pryor, Mycologia 105: 542. 2013. Fig. 22.

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Fig. 22.

Alternaria sect. Sonchi: conidia and conidiophores. A-B. A. cinerariae. C-D. A. sonchi. Scale bars = 10 μm.

Type species: Alternaria sonchi Davis

Diagnosis: Section Sonchi is characterised by subcylindrical, broadly ovoid, broadly ellipsoid or obclavate, (medium) large conidia, single or in short chains, with multiple transverse and few longitudinal septa, slightly constricted at the septa, with a blunt taper which can form secondary conidiophores.

Notes: The species-group was described by Hong et al. (2005) based on molecular data of the GAPDH and Alt a 1 regions. Lawrence et al. (2013) included A. brassicae as a basal lineage in sect. Sonchi, which is supported as a monotypic lineage in our analyses. The species from section Sonchi occur on multiple hosts within the Compositae.

Alternaria cinerariae Hori & Enjoji, J. Pl. Protect. 18: 432. 1931.

Alternaria sonchi Davis, in Elliott, Bot. Gaz. 62: 416. 1916.

Section Teretispora (E.G. Simmons) Woudenb. & Crous, comb. et stat. nov. MycoBank MB803745. Fig. 23.

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Fig. 23.

Alternaria sect. Teretispora: conidia and conidiophores. A-D. A. leucanthemi. Scale bars = 10 μm.

Basionym: Teretispora E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 674. 2007.

Type species: Alternaria leucanthemi Nelen

Diagnosis: Section Teretispora is characterised by simple conidiophores, sometimes extending at the apex with one or two, geniculate, sympodial proliferations, bearing single, long cylindrical mature conidia lacking a beak portion, with many transverse and a few longitudinal septa, constricted at most of the transverse septa. Secondary conidiophores with a single conidium are rarely formed at the apex; instead, they may form from the base of the primary conidium.

Notes: The genus Teretispora had Teretispora leucanthemi, formerly Alternaria leucanthemi (= Alternaria chrysanthemi), as type and only species (Simmons 2007). We choose to treat this as a section, which retains the name Teretispora, rather than a monotypic lineage.

Alternaria leucanthemi Nelen, in Nelen & Vasiljeva, Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Akad. Nauk S.S.S.R. 15: 148. 1962.

• • ≡ Teretispora leucanthemi (Nelen) E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 674. 2007.

• = Alternaria leucanthemi Nelen, Bull. Centr. Bot. Gard. (Moscow) 35: 83. 1959. (nom. inval., Art. 36.1)

• = Alternaria chrysanthemi E.G. Simmons & Crosier, Mycologia 57: 142. 1965.

Section Ulocladioides Woudenb. & Crous, sect. nov. MycoBank MB803746. Fig. 24.

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Fig. 24.

Alternaria sect. Ulocladioides: conidia and conidiophores. A-B. A. atra. C-D. A. brassicae-pekinensis. E-F. A. cantlous. G-H. A. multiformis. I-J. A. obovoidea. K-L. A. heterospora. M-N. A. subcucurbitae. O-P. A. terricola. Scale bars = 10 μm.

Type species: Alternaria cucurbitae Letendre & Roum.

Diagnosis: Section Ulocladioides is characterised by conidiophores with short, geniculate, sympodial proliferations. Conidia are obovoid, non-beaked with a narrow base, single or in chains, which may form secondary conidiophores at the apex.

Note: Section Ulocladioides resembles section Ulocladium and contains the majority of the species included in this study from the genus Ulocladium (11/17).

Alternaria atra (Preuss) Woudenb. & Crous, comb. nov. MycoBank MB803717.

Basionym: Ulocladium atrum Preuss, Linnaea 25: 75. 1852.

• ≡ Stemphylium atrum (Preuss) Sacc., Syll. Fungorum (Abellini) 4: 520. 1886.

Alternaria brassicae-pekinensis Woudenb. & Crous, nom. nov. MycoBank MB803723.

Basionym: Ulocladium brassicae Yong Wang bis & X.G. Zhang, Mycologia 100: 457. 2008, non Alternaria brassicae (Berk.) Sacc., 1880.

Etymology: Name refers to the host from which it was originally isolated.

Alternaria cantlous (Yong Wang bis & X.G. Zhang) Woudenb. & Crous, comb. nov. MycoBank MB803719.

Basionym: Ulocladium cantlous Yong Wang bis & X.G. Zhang, Mycologia 102: 376. 2010.

Alternaria consortialis (Thüm.) J.W. Groves & S. Hughes [as “consortiale”], Canad. J. Bot. 31: 636. 1953.

Basionym: Macrosporium consortiale Thüm., Herb. Mycol. Oecon. 9: no. 450. 1876.

• • ≡ Stemphylium consortiale (Thüm.) J.W. Groves & Skolko, Canad. J. Res., Sect. C, Bot. Sci.: 196. 1944.

  • ≡ Pseudostemphylium consortiale (Thüm.) Subram., Curr. Sci. 30: 423. 1961.

  • ≡ Ulocladium consortiale (Thüm.) E.G. Simmons, Mycologia 59: 84. 1967.

• = Stemphylium ilicis Tengwall, Meded. Phytopathol. Lab. “Willie Commelin Scholten” 6: 44. 1924.

Alternaria cucurbitae Letendre & Roum., in Roumeguère, Rev. Mycol. (Toulouse) 8 (no. 30): 93. 1886.

• ≡ Ulocladium cucurbitae (Letendre & Roum.) E.G. Simmons, Mycotaxon 14: 48. 1982.

Alternaria heterospora Woudenb. & Crous, nom. nov. MycoBank MB803724.

Basionym: Ulocladium solani Yong Wang bis & X.G. Zhang, Mycol. Progr. 8: 209. 2009, non Alternaria solani Sorauer, 1896.

Etymology: Name refers to the various conidial morphologies observed during growth.

Alternaria multiformis (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803720.

Basionym: Ulocladium multiforme E.G. Simmons, Canad. J. Bot. 76: 1537. 1999 [1998].

Alternaria obovoidea (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803721.

Basionym: Ulocladium obovoideum E.G. Simmons, Mycotaxon 37: 104. 1990.

Alternaria subcucurbitae (Yong Wang bis & X.G. Zhang) Woudenb. & Crous, comb. nov. MycoBank MB803722.

Basionym: Ulocladium subcucurbitae Yong Wang bis & X.G. Zhang, Mycologia 100: 456. 2008.

Alternaria terricola Woudenb. & Crous, nom. nov. MycoBank MB803725.

Basionym: Ulocladium tuberculatum E.G. Simmons, Mycologia 59: 83. 1967, non Alternaria tuberculata M. Zhang & T.Y. Zhang, 2006. Etymology: Name refers to soil from which it was originally isolated.

Section Ulocladium (Preuss) Woudenb. & Crous, comb. et stat. nov. MycoBank MB803747. Fig. 25.

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Fig. 25.

Alternaria sect. Ulocladium: conidia and conidiophores. A-B. A. capsici-annui. C-D. A. oudemansii. E-F. A. alternariae. G-H. A. botrytis. Scale bars = 10 μm.

Basionym: Ulocladium Preuss, Linnaea 24: 111. 1851.

Type species: Alternaria botrytis (Preuss) Woudenb. & Crous

Diagnosis: Section Ulocladium is characterised by simple conidiophores, or with one or two short, geniculate, sympodial proliferations, with (mostly) single, obovoid, non-beaked conidia with a narrow base.

Notes: Section Ulocladium resembles sect. Ulocladioides. The epitype of Ulocladium, U. botrytis CBS 197.67, and the isotype of U. oudemansii (CBS 114.07) cluster with the Sinomyces representative, as do many other strains stored as U. botrytis in the CBS collection (data not shown). Furthermore, a strain stored as A. capsici-annui (CBS 504.74) in the CBS collection clusters within the Sinomyces clade and displays identical morphological features.

Alternaria alternariae (Cooke) Woudenb. & Crous, comb. nov. MycoBank MB803716.

Basionym: Sporidesmium alternariae Cooke, Handb. Brit. Fungi 1: 1440. 1871.

• ≡ Stemphylium alternariae (Cooke) Sacc., Syll. Fungorum (Abellini) 4: 523. 1886.

• ≡ Ulocladium alternariae (Cooke) E.G. Simmons, Mycologia 59: 82. 1967.

• ≡ Sinomyces alternariae (Cooke) Yong Wang bis & X.G. Zhang, Fungal Biol. 115: 194. 2011.

Alternaria botrytis (Preuss) Woudenb. & Crous, comb. nov. MycoBank MB803718.

Basionym: Ulocladium botrytis Preuss, Linnaea 24: 111. 1851.

• ≡ Stemphylium botryosum var. ulocladium Sacc. (nom. nov.), Syll. Fungorum (Abellini) 4: 522. 1886.

• ≡ Stemphylium botryosum var. botrytis (Preuss) Lindau, Rabenhorst’s. Kryptog.-Fl., Edn 2 (Leipzig) 1(9): 219. 1908.

Alternaria capsici-annui Săvul. & Sandu, Hedwigia 75: 228. 1936.

Alternaria oudemansii (E.G. Simmons) Woudenb. & Crous, comb. nov. MycoBank MB803715.

Basionym: Ulocladium oudemansii E.G. Simmons, Mycologia 59: 86. 1967.

Section Undifilum (B.M. Pryor, Creamer, Shoemaker, McLain-Romero & Hambl.) Woudenb. & Crous, comb. et stat. nov. MycoBank MB803748. Fig. 26.

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Fig. 26.

Alternaria sect. Undifilum: conidia and conidiophores. A-D. A. bornmuelleri. Scale bars = 10 μm.

Basionym: Undifilum B.M. Pryor, Creamer, Shoemaker, McLain-Romero & Hambl., Botany 87: 190. 2009.

Type species: Alternaria bornmuelleri (Magnus) Woudenb. & Crous

Diagnosis: Section Undifilum is characterised by ovate to obclavate to long ellipsoid, straight to inequilateral, single, transseptate conidia; septa can be thick, dark and rigid, and form unique germ tubes, which are wavy or undulate until branching. Species of this section occur on Fabaceae and almost all produce the toxic compound swaisonine.

Notes: Section Undifilum shares morphological features with section Embellisia, but is characterised by the formation of a wavy germ tube upon germination (Pryor et al. 2009). Based on previous studies, the swaisonine producing species U. oxytropis (Pryor et al. 2009, Lawrence et al. 2012), U. fulvum and U. cinereum (Baucom et al. 2012) also belong to this section, although the type species, A. bornmuelleri, does not produce swaisonine.

Alternaria bornmuelleri (Magnus) Woudenb. & Crous, comb. nov. MycoBank MB803726.

Basionym: Helminthosporium bornmuelleri Magnus, Hedwigia 38 (Beibl.): 73. 1899.

• ≡ Undifilum bornmuelleri (Magnus) B.M. Pryor, Creamer, Shoemaker, McLain-Romero & Hambl., Botany 87: 190. 2009.

Alternaria cinerea (Baucom & Creamer) Woudenb. & Crous, comb. nov. MycoBank MB803731.

Basionym: Undifilum cinereum Baucom & Creamer, Botany 90: 872. 2012

Alternaria fulva (Baucom& Creamer) Woudenb. & Crous, comb. nov. MycoBank MB803732.

Basionym: Undifilum fulvum Baucom & Creamer, Botany 90: 871. 2012

Alternaria oxytropis (Q. Wang, Nagao & Kakish.) Woudenb. & Crous, comb. nov. MycoBank MB803727.

Basionym: Embellisia oxytropis Q. Wang, Nagao & Kakish., Mycotaxon 95: 257. 2006.

• ≡ Undifilum oxytropis (Q. Wang, Nagao & Kakish.) B.M. Pryor, Creamer, Shoemaker, McLain-Romero & Hambl., Botany 87: 191. 2009.

Monotypic lineages

The following six species are not assigned to one of the 24 above described Alternaria sections and are treated as separate, single species, lineages in this study. Future studies, including more and/or new Alternaria species, might eventually give rise to the formation of new sections, when these new species show to be closely related to one of these monotypic lineages.

Alternaria argyranthemi E.G. Simmons & C.F. Hill, Mycotaxon 65: 32. 1997.

Alternaria brassicae (Berk.) Sacc., Michelia 2(no. 6): 129. 1880.

Basionym: Macrosporium brassicae Berk., Engl. Fl., Fungi (Edn 2) (London) 5: 339. 1836.

Additional synonyms listed in Simmons (2007).

Alternaria dennisii M.B. Ellis, Mycol. Pap. 125: 27. 1971.

• ≡ Embellisia dennisii (M.B. Ellis) E.G. Simmons, Mycotaxon 38: 257. 1990.

Alternaria helianthiinficiens E.G. Simmons, Walcz & R.G. Roberts [as “helianthinficiens”], Mycotaxon 25: 204. 1986.

Alternaria soliaridae E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 6: 374. 2007.

Alternaria thalictrigena K. Schub. & Crous, Fungal Planet No. 12: 2. 2007.

Paradendryphiella Woudenb. & Crous, gen. nov. MycoBank MB803750. Fig. 27.

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Fig. 27.

Paradendryphiella gen. nov.: conidia and conidiophores. A-B, D-E, G-I. P. salina. C, F. P. arenariae. Scale bars = 10 μm.

Colonies on SNA effuse, entire, velvety, olivaceous. Reverse olivaceous-grey to iron-grey. Mycelium consisting of branched, septate hypha, (sub)hyaline, smooth. Conidiophores subhyaline, simple or branched, septate or not, straight or flexuous, often nodose with conspicuous, brown pigmentation at the apical region; at times reduced to conidiogenous cells. Conidiogenous cells terminal or lateral, with denticles aggregated at apex, with prominent conidial scars, thickened but not darkened; sometimes proliferating with a new head or a short, inconspicuous sympodial rachis. Conidia produced holoblastically, on narrow denticle, smooth, cylindrical to obclavate, straight or slightly flexuous, 1-7 transverse septa, pale to medium brown, often with dark septa (often constricted), and a darkened zone of pigmentation at the apex, and at the hilum, which is thickened, and somewhat protruding, with a minute marginal frill. Chlamydospores and sexual state not observed.

Type species: Paradendryphiella salina (G.K. Sutherl.) Woudenb. & Crous

Paradendryphiella salina (G.K. Sutherl.) Woudenb. & Crous, comb. nov. MycoBank MB803751.

Basionym: Cercospora salina G.K. Sutherl., New Phytol. 15: 43. 1916.

• • ≡ Dendryphiella salina (G.K. Sutherl.) Pugh & Nicot, Trans. Brit. Mycol. Soc. 47(2): 266. 1964.

  • ≡ Scolecobasidium salinum (G.K. Sutherl.) M.B. Ellis, More dematiaceous hyphomycetes (Kew): 192. 1976.

• = Embellisia annulata de Hoog, Seigle-Mur., Steiman & K.-E. Erikss., Antonie van Leeuwenhoek J. Microbiol. Serol. 51: 409. 1985.

Paradendryphiella arenariae (Nicot) Woudenb. & Crous, comb. nov. MycoBank MB803752.

Basionym: Dendryphiella arenariae Nicot, [as “arenaria”] Rev. Mycol. (Paris) 23: 93. 1958.

• ≡ Scolecobasidium arenarium (Nicot) M.B. Ellis, More dematiaceous hyphomycetes (Kew): 194. 1976.

DISCUSSION

The well-supported node for the Alternaria clade obtained in the present study, and the low bootstrap support at the deeper nodes within the Alternaria complex is also consistently seen in previous phylogenetic studies published on these genera (Pryor & Bigelow 2003, Inderbitzin et al. 2006, Pryor et al. 2009, Runa et al. 2009, Wang et al. 2011, Lawrence et al. 2012). The only phylogenetic study which displays a second fully supported node is based on a five-gene combined dataset of GAPDH, Alt a 1, actin, plasma membrane ATPase and calmodulin (Lawrence et al. 2013). This node, called clade A by the authors, supports eight “asexual” Alternaria species-groups and an Ulocladium (sect. Ulocladioides in our phylogenies) clade. By resolving these eight asexual phylogenetic lineages of Alternaria together with Ulocladium, which is sister to the sexual A. infectoria species-group and other sexual genera, Lawrence et al. (2013) elevated the asexual species-groups to sections within Alternaria. If we take this node as cut-off for the genus Alternaria in our phylogenies, this would leave an Alternaria clade with 14 internal clades (sections) and three monotypic lineages. In order to create a stable phylogenetic taxonomy, seven new genera need to be described of which three would be monotypic; E. dennissii, A. argyranthemi and A. soliaridae. Embellisia species would be assigned to five different genera of which four would be new, leaving only E. allii, E. chlamydospora and E. tellustris in the genus Embellisia. The well-known (medical) A. infectoria species-group would also have to be transferred to a new genus. This node is not supported in our study (0.98 PP /65 ML Fig 1) and also the strict asexual/sexual division is not supported as two sexual morphs are found in section Panax. This approach would therefore give rise to multiple small genera, and would not end up in a logical and workable situation.

Based on our phylogenetic study on parts of the SSU, LSU, ITS, GAPDH, RPB2 and TEF1 gene regions of ex-type and reference strains of Alternaria species and all available allied genera, we resolved a Pleospora/Stemphylium-clade sister to Embellisia annulata, and a well-supported Alternaria clade. The Alternaria clade contains 24 internal clades and six monotypic lineages. In combination with a review of literature and morphology, the species within the Alternaria clade are all recognised here as Alternaria s. str. This puts the genera Allewia, Brachycladium, Chalastospora, Chmelia, Crivellia, Embellisia, Lewia, Nimbya, Sinomyces, Teretispora, Ulocladium, Undifilum and Ybotromyces in synonymy with Alternaria.

The support values for the different sections described in this study are plotted in a heatmap per gene/gene combination and phylogenetic method used (Table 2). This shows that the Bayesian method provides greater support than the Maximum Likelihood bootstrap support values, which is in congruence with previous reports (e.g. Douady et al. 2003). The sections Cheiranthus, Eureka and Nimbya have the lowest support values. For sect. Eureka this is mainly caused by the position of A. cumini, which clusters within sect. Embellisioides based on its RPB2 sequence and as a monotypic lineage based on its TEF1 sequence. Section Cheiranthus and Nimbya are small sections, with relative long branches. Future studies, including more strains and/or species in these sections, are necessary to check the stability of these long branches.

The sexual genus Crivellia with its Brachycladium asexual morph was described by Inderbitzin et al. (2006) with Crivellia papaveraceae (asexual morph Brachycladium penicillatum) as type species and B. papaveris, with an unnamed sexual morph, as second species. The genus Brachycladium, which was synonymised with Dendryphion (Ellis 1971), was resurrected for the non-sexual stage based on polyphyly within Dendryphion and morphological distinction from its type species, D. comosum. The type species of Brachycladium, B. penicillatum, resides in Alternaria sect. Crivellia, which places Brachycladium in synonymy with Alternaria instead of Dendryphion.

The genus Chalastospora was established by Simmons (2007) based on Chalastospora cetera, formerly Alternaria cetera. Two new Chalastospora species, C. ellipsoidea and C. obclavata, and A. malorum as C. gossypii were later added to the genus, based on sequence data of the ITS and LSU regions (Crous et al. 2009c). The genus is characterised by conidia which are almost always narrowly ellipsoid to narrowly ovoid with 1-6 transverse eusepta, generally lacking oblique or longitudinal septa (Crous et al. 2009c). Our study shows that Alternaria armoraciae and Embellisia abundans also belong to this clade. Juvenile conidia of A. armoraciae are ovoid, but vary from being narrow to broadly ovoid and ellipsoid, with 3-5 transverse septa and a single longitudinal septum in up to four of the transverse segments (Simmons 2007). Embellisia abundans was already mentioned as part of the Chalastospora clade (Andersen et al. 2009, Lawrence et al. 2012), and has long ovoid or obclavate conidia with 3-6 transverse septa and rarely any longitudinal septa (Simmons 1983). The description of sect. Chalastospora does therefore not completely follow the original description of the genus Chalastospora.

The genus Embellisia is characterised by the thick, dark, rigid conidial septa and the scarcity of longitudinal septa (Simmons 2007). It was first described by Simmons (1971), with Embellisia allii as type and E. chlamydospora as second species. Multiple Embellisia species followed after the description of the genus, which was later linked to the sexual genus Allewia (Simmons 1990). The latest molecular-based revision was performed based on sequences of the GAPDH, ITS and Alt a 1 genes (Lawrence et al. 2012). They found that Embellisia split into four clades and multiple species, which clustered individually amidst Alternaria, Ulocladium or Stemphylium spp. Our results mostly support these data, but with the inclusion of more ex-type/representative strains of Alternaria some additions were made to the different Embellisia groups mentioned by Lawrence et al. (2012). Group I (sect. Embellisia) and III (sect. Embellisioides) are identical to the treatment of Lawrence et al. (2012) but group II (section Phragmosporae) and IV (section Eureka) are both expanded with four Alternaria species. As not all species from group II and IV display the typical morphological characters of Embellisia, we chose to name these Alternaria sections based on the oldest species residing in the respective sections. Embellisia abundans was already mentioned as being part of the Chalastospora-clade and E. indefessa formed a clade close to Ulocladium, which we now assign to sect. Cheiranthus. Embellisia dennisii also forms a separate lineage in our phylogenies; therefore the old name Alternaria dennissii is resurrected. Furthermore, the clustering of E. conoidea within the A. brassicicola species-group and E. annulata close to Stemphylium, now assigned as Paradendryphiella gen. nov., is confirmed by our phylogenetic data. The morphological character of thick, dark, rigid septa seems to have evolved multiple times and does not appear to be a valid character for taxonomic distinction at generic level.

The sexual morphs Lewia (Simmons 1986) and Allewia (Simmons 1990) were linked to Alternaria and Embellisia respectively, with the only difference between these genera being the morphology of their asexual morphs. Lewia chlamidosporiformans and L. sauropodis are transferred to the genus Leptosphaerulina (Simmons 2007), which leaves 11 Lewia species with a known Alternaria anamorph. Most of them (9/11) reside in sect. Infectoriae, the others are found in sect. Panax. Allewia only contains two species of which one resides in sect. Eureka and one in sect. Embellisioides. With the establishment of the new International Code of Nomenclature for algae, fungi and plants (ICN), the dual nomenclature system for sexual and asexual fungal morphs was abandoned and replaced by a single-name nomenclature (Hawksworth et al. 2011, Norvell 2011). In order to implement the new rules of the ICN, we synonymised Lewia and Allewia with Alternaria.

Although multiple molecular studies included Nimbya isolates in their phylogenies (Chou & Wu 2002, Pryor & Bigelow 2003, Hong et al. 2005, Inderbitzin et al. 2006, Pryor et al. 2009), a more extensive molecular-based study was recently published by Lawrence et al. (2012). Based on sequences of the GAPDH, ITS and Alt a 1 genes, the authors found a Nimbya clade which contained the type species N. scirpicola together with N. scirpinfestans, N. scirpivora and N. caricis. The N. scirpicola isolate which we included in our study, was assigned to this genus by Simmons (1989) based on morphological characters, as is the one used in other molecular studies (Pryor & Bigelow 2003, Hong et al. 2005, Lawrence et al. 2012). The sequences of the ITS, GAPDH and Alt a 1 genes of these isolates are however not identical, but do cluster in the same clade in the two phylogenies (data not shown), together with the isolate of N. caricis. The N. gomphrenae isolate we included in our phylogeny was not representative of the name. Simmons mentioned in 1989 that Togashi (1926) described two different fungi and deposited the small-spored species in the CBS collection, instead of the large-spored N. gomphrenae isolate. Nimbya gomphrenae CBS 108.27, which does not sporulate anymore, will therefore be treated as “Alternaria sp.”, and resides in sect. Alternata. The ITS sequence of N. gomphrenae from Chou & Wu (2002) actually clusters within sect. Alternantherae. This section was described by Lawrence et al. (2012) and consists of three Nimbya species, which they renamed to Alternaria based on the position of the clade amidst the Alternaria species-groups. Based on the data from Chou & Wu (2002), the name Alternaria gomphrenae is resurrected and placed in sect. Alternantherae.

The genus Sinomyces was described in by Wang et al. (2011) to accommodate Ulocladium alternariae and two new species from China, S. obovoideus and S. fusoides (type). The genus was differentiated from Ulocladium based on its simple conidiophores with a single apical pore or 1-2 short, uniperforate, geniculate sympodial proliferations. Unfortunately, our DNA sequence analyses of the ex-type cultures of the two new species from China (CBS 124114 and CBS 123375) were not congruent with the GAPDH (both species) and Alt a 1 (S. obovoideus) sequences deposited in GenBank (data not shown), leading us to doubt the authenticity of these strains. This matter could not be resolved in spite of contacting the original depositors. The ex-type strain of S. alternariae (CBS 126989) was therefore included as representative of the genus Sinomyces. The presence of the epitype of Ulocladium, U. botrytis CBS 197.67, in this section resulted in us rejecting the name Sinomyces, and calling this sect. Ulocladium. In addition, the presence of U. oudemansii in this section, with conidiophores with 1-5 uniperforate geniculations (Simmons 1967), also disagrees with the mentioned differentiation of Sinomyces from Ulocladium.

The type species of Ulocladium, U. botrytis, was typified by two representative strains QM 7878 (CBS 197.67) and QM 8619 (CBS 198.67) (Simmons 1967). Molecular studies performed afterwards showed that these strains are not identical (de Hoog & Horré 2002). Most molecular studies performed used CBS 198.67 as representative of U. botrytis (Pryor & Gilbertson 2000, Pryor & Bigelow 2003, Hong et al. 2005, Xue & Zhang 2007, Pryor et al. 2009, Runa et al. 2009, Wang et al. 2010, Wang et al. 2011, Lawrence et al. 2012), which clusters in section Ulocladioides. However, de Hoog & Horré (2002) epitypified U. botrytis with CBS 197.67, which clusters with Sinomyces strains, as does Ulocladium oudemansii, now named sect. Ulocladium. Extended phylogenetic analyses on all U. botrytis strains present in the CBS culture collection (16 isolates) also highlight this issue as they cluster either within sect. Ulocladium or sect. Ulocladioides (data not shown), both with one of the representative strains described by Simmons (1967). The suggestion to synonymise Ulocladium with Alternaria has been made several times in the past (Pryor & Gilbertson 2000, Chou & Wu 2002). The latest systematic revision of the genus Ulocladium (Runa et al. 2009) based on sequences from the ITS, GAPDH and Alt a 1 genes supported previous findings of poly- and paraphyletic relationships of Ulocladium among Alternaria, Embellisia and Stemphylium spp. (de Hoog & Horré 2002, Pryor & Bigelow 2003, Hong et al. 2005). Ulocladium alternariae and U. oudemansii, now known as sect. Ulocladium, cluster separately. The core Ulocladium clade, containing the two sister clades now called sect. Ulocladioides and sect. Pseudoulocladium, was confirmed by later studies (Wang et al. 2010, Lawrence et al. 2012). Alternaria cheiranthi and Embellisia indefessa have been linked to Ulocladium (Pryor & Gilbertson 2000, Pryor & Bigelow 2003, Hong et al. 2005, Pryor et al. 2009, Runa et al. 2009, Lawrence et al. 2012), but missed the diagnostic feature of Ulocladium. Our study showed that they form a sister section, sect. Cheiranthus, to sect. Ulocladioides. The confusing taxonomy in this genus strengthens our decision to reduce Ulocladium to synonymy with Alternaria. The characteristics of the former genus Ulocladium are added to the new broader Alternaria generic circumscription.

The genus Undifilum was described by Pryor et al. (2009) to accommodate the species U. oxytropis and U. bornmuelleri. It shares the morphological feature of thick, dark and rigid septa with the genus Embellisia, but was characterised by the formation of a wavy germ-tube upon germination (Pryor et al. 2009). A recent study on fungal endophytes in locoweeds in the US described two new Undifilum species (Baucom et al. 2012). Both new species produce the toxic compound swaisonine, which is also produced by U. oxytropis. Swaisonine is the cause of a neurological disease, locism, of grazing animals, resulting in economic losses in livestock (James & Panter 1989). The production of swaisonine seems to be related to this section, although the type-species, U. bornmuelleri, does not produce this toxin.

The genus Ybotromyces contains one species, Y. caespitosus (originally Botryomyces caespitosus), which was isolated from a skin lesion of a human patient (de Hoog & Rubio 1982). De Hoog et al. (1997) discovered a high similarity to Alternaria spp. based on restriction patterns of the ITS and SSU rDNA. A phylogeny study of melanised meristematic fungi based on their SSU and ITS rDNA sequences (Sterflinger et al. 1999) placed Y. caespitosus within the Pleosporales together with Alternaria and Pleospora. De Hoog & Horré (2002) hypothesized that the ex-type strain of Y. caespitosus, CBS 177.80, is likely a synanamorph of a yet undescribed Alternaria species. Our phylogeny supports this hypothesis, and places the genus in sect. Infectoriae.

Chmelia slovaca, described from dermatic lesions of a human (Svobodová 1966), also clusters with sect. Infectoriae as was shown previously (de Hoog & Horré 2002). The genus produces different types of chlamydospores and sporadically blastospores, but no conidia or conidiophores, which makes it difficult to identify based on morphology. De Hoog & Horré (2002) were confident that Chmelia is a sterile member of A. infectoria, which is in agreement with our results.

DEDICATION

We would like to dedicate this manuscript to the late Dr E.G. Simmons, who spent over 50 years of his life researching the systematics of the genus Alternaria. Without the time EGS spent on characterising the species included in this study, and his impeccable strain collection, which he placed in CBS for preservation and further study, the present study would not have been possible.

Acknowledgments

REFERENCES