Alternaria redefined - CBS - KNAW

Studies in Mycology 75: 171–212. 

Alternaria redefined 

J.H.C. Woudenberg 1,2* , J.Z. Groenewald 1 , M. Binder 1 , and P.W. Crous 1,2,3 

1 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; 2 Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, 

Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands; 3 Utrecht University, Department of Biology, Microbiology, Padualaan 8, 3584 CH Utrecht, The Netherlands 

*Correspondence: Joyce H.C. Woudenberg, j.woudenberg@cbs.knaw.nl 

Abstract: Alternaria is a ubiquitous fungal genus that includes saprobic, endophytic and pathogenic species associated with a wide variety of substrates. In recent years, DNAbased 

studies revealed multiple non-monophyletic genera within the Alternaria complex, and Alternaria species clades that do not always correlate to species-groups based 

on morphological characteristics. The Alternaria complex currently comprises nine genera and eight Alternaria sections. The aim of this study was to delineate phylogenetic 

lineages within Alternaria and allied genera based on nucleotide sequence data of parts of the 18S nrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha gene regions. 

Our data reveal 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, the assemblage of which we recognise as Alternaria. This puts the genera Allewia, Brachycladium, Chalastospora, Chmelia, Crivellia, Embellisia, 

Lewia, Nimbya, Sinomyces, Teretispora, Ulocladium, Undifilum and Ybotromyces in synonymy with Alternaria. In this study, we treat the 24 internal clades in the Alternaria 

complex as sections, which is a continuation of a recent proposal for the taxonomic treatment of lineages in Alternaria. Embellisia annulata is synonymised with Dendryphiella 

salina, and together with Dendryphiella arenariae, are placed in the new genus Paradendryphiella. The sexual genera Clathrospora and Comoclathris, which were previously 

associated with Alternaria, cluster within the Pleosporaceae, outside Alternaria s. str., whereas Alternariaster, a genus formerly seen as part of Alternaria, clusters within the 

Leptosphaeriaceae. Paradendryphiella is newly described, the generic circumscription of Alternaria is emended, and 32 new combinations and 10 new names are proposed. A 

further 10 names are resurrected, while descriptions are provided for 16 new Alternaria sections. 

Studies in Mycology 

Key words: Allewia, Chalastospora, Crivellia, Embellisia, Lewia, Nimbya, Paradendryphiella, Sinomyces, systematics, Teretispora, Ulocladium, Undifilum. 

Taxonomic novelties: New combinations – Alternaria abundans (E.G. Simmons) Woudenb. & Crous, Alternaria alternariae (Cooke) Woudenb. & Crous, Alternaria atra 

(Preuss) Woudenb. & Crous, Alternaria bornmuelleri (Magnus) Woudenb. & Crous, Alternaria botrytis (Preuss) Woudenb. & Crous, Alternaria caespitosa (de Hoog & C. Rubio) 

Woudenb. & Crous, Alternaria cantlous (Yong Wang bis & X.G. Zhang) Woudenb. & Crous, Alternaria caricis (E.G. Simmons) Woudenb. & Crous, Alternaria cinerea (Baucom 

& Creamer) Woudenb. & Crous, Alternaria didymospora (Munt.-Cvetk.) Woudenb. & Crous, Alternaria fulva (Baucom & Creamer) Woudenb. & Crous, Alternaria hyacinthi (de 

Hoog & P.J. Mull. bis) Woudenb. & Crous, Alternaria indefessa (E.G. Simmons) Woudenberg & Crous, Alternaria leptinellae (E.G. Simmons & C.F. Hill) Woudenb. & Crous, 

Alternaria lolii (E.G. Simmons & C.F. Hill) Woudenb. & Crous, Alternaria multiformis (E.G. Simmons) Woudenb. & Crous, Alternaria obclavata (Crous & U. Braun) Woudenb. & 

Crous, Alternaria obovoidea (E.G. Simmons) Woudenb. & Crous, Alternaria oudemansii (E.G. Simmons) Woudenb. & Crous, Alternaria oxytropis (Q. Wang, Nagao & Kakish.) 

Woudenb. & Crous, Alternaria penicillata (Corda) Woudenb. & Crous, Alternaria planifunda (E.G. Simmons) Woudenb. & Crous, Alternaria proteae (E.G. Simmons) Woudenb. 

& Crous, Alternaria scirpinfestans (E.G. Simmons & D.A. Johnson) Woudenb. & Crous, Alternaria scirpivora (E.G. Simmons & D.A. Johnson) Woudenb. & Crous, Alternaria 

septospora (Preuss) Woudenb. & Crous, Alternaria slovaca (Svob.-Pol., L. Chmel & Bojan.) Woudenb. & Crous, Alternaria subcucurbitae (Yong Wang bis & X.G. Zhang) 

Woudenb. & Crous, Alternaria tellustris (E.G. Simmons) Woudenb. & Crous, Alternaria tumida (E.G. Simmons) Woudenb. & Crous, Paradendryphiella salina (G.K. Sutherl.) 

Woudenb. & Crous, Paradendryphiella arenariae (Nicot) Woudenb. & Crous. New names – Alternaria aspera Woudenb. & Crous, Alternaria botryospora Woudenb. & Crous, 

Alternaria brassicae-pekinensis Woudenb. & Crous, Alternaria breviramosa Woudenb. & Crous, Alternaria chlamydosporigena Woudenb. & Crous, Alternaria concatenata 

Woudenb. & Crous, Alternaria embellisia Woudenb. & Crous, Alternaria heterospora Woudenb. & Crous, Alternaria papavericola Woudenb. & Crous, Alternaria terricola 

Woudenb. & Crous. Resurrected names – Alternaria cetera E.G. Simmons, Alternaria chartarum Preuss, Alternaria consortialis (Thüm.) J.W. Groves & S. Hughes, Alternaria 

cucurbitae Letendre & Roum., Alternaria dennisii M.B. Ellis, Alternaria eureka E.G. Simmons, Alternaria gomphrenae Togashi, Alternaria malorum (Ruehle) U. Braun, Crous 

& Dugan, Alternaria phragmospora Emden, Alternaria scirpicola (Fuckel) Sivan. New sections, all in Alternaria – sect. Chalastospora Woudenb. & Crous, sect. Cheiranthus 

Woudenb. & Crous, sect. Crivellia Woudenb. & Crous, sect. Dianthicola Woudenb. & Crous, sect. Embellisia Woudenb. & Crous, sect. Embellisioides Woudenb. & Crous, sect. 

Eureka Woudenb. & Crous, sect. Infectoriae Woudenb. & Crous, sect. Japonicae Woudenb. & Crous, sect. Nimbya Woudenb. & Crous, sect. Phragmosporae Woudenb. & 

Crous, sect. Pseudoulocladium Woudenb. & Crous, sect. Teretispora Woudenb. & Crous, sect. Ulocladioides Woudenb. & Crous, sect. Ulocladium Woudenb. & Crous, sect. 

Undifilum Woudenb. & Crous. New genus – Paradendryphiella Woudenb. & Crous. 

Published online: 31 May 2013; doi:10.3114/sim0015. Hard copy: June 2013. 

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 

Copyright CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

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 

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171

Woudenberg et al. 

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 speciesgroups 

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 extype 

and reference strains of Alternaria species and all available 

allied genera. 

MATERIAL AND METHODS 

Isolates 

Based on the ITS sequences of all ex-type or representative 

strains from the Alternaria identification manual present at the 

CBS-KNAW Fungal Biodiversity Centre (CBS), Utrecht, The 

Netherlands (data not shown), 66 Alternaria strains were included 

in this study together with 61 ex-type or representative strains of 16 

related genera (Table 1). Alternaria is represented by the ex-type 

or representative strains of the seven species-groups and species 

that clustered outside known Alternaria clades. Because of the 

size and complexity of the A. alternata, A. infectoria and A. porri 

species-groups, we only included known species; the complete 

species-groups will be treated in future studies. 

Freeze-dried strains were revived in 2 mL malt/peptone (50 % 

/ 50 %) and subsequently transferred to oatmeal agar (OA) (Crous 

et al. 2009a). Strains of the CBS collection stored in liquid nitrogen 

were transferred to OA directly from -80 ºC. DNA extraction was 

performed using the UltraClean Microbial DNA Isolation Kit (MoBio 

laboratories, Carlsbad, CA, USA), according to the manufacturer’s 

instructions. 

Taxonomy 

Morphological descriptions were made for isolates grown on 

synthetic nutrient-poor agar plates (SNA, Nirenberg 1976) with a 

small piece of autoclaved filter paper placed onto the agar surface. 

Cultures were incubated at moderate temperatures (~ 22 ºC) 

under CoolWhite fluorescent light with an 8 h photoperiod for 7 d. 

The sellotape technique was used for making slide preparations 

(Crous et al. 2009a) with Shear’s medium as mounting fluid. 

Photographs of characteristic structures were made with a Nikon 

Eclipse 80i microscope using differential interference contrast 

(DIC) illumination. Growth rates were measured after 5 and 7 d. 

Colony characters were noted after 7 d, colony colours were rated 

according to Rayner (1970). Nomenclatural data were deposited in 

MycoBank (Crous et al. 2004). 

PCR and sequencing 

The SSU region was amplified with the primers NS1 and NS4 (White 

et al. 1990), the LSU region with LSU1Fd (Crous et al. 2009b) and 

LR5 (Vilgalys & Hester 1990), the ITS region with V9G (De Hoog 

& Gerrits van den Ende 1998) and ITS4 (White et al. 1990), the 

GAPDH region with gpd1 and gpd2 (Berbee et al. 1999), the RPB2 

region with RPB2–5F2 (Sung et al. 2007) and fRPB2–7cR (Liu et 

al. 1999) and the TEF1 gene with the primers EF1-728F and EF1- 

986R (Carbone & Kohn 1999) or EF2 (O’Donnell et al. 1998). The 

PCRs were performed in a MyCycler TM Thermal Cycler (Bio-Rad 

Laboratories B.V., Veenendaal, The Netherlands) in a total volume 

of 12.5 µL. The SSU and LSU PCR mixtures consisted of 1 µL 

genomic DNA, 1´ GoTaq® Flexi buffer (Promega, Madison, WI, 

USA), 2 µM MgCl 2 

, 40 µM of each dNTP, 0.2 µM of each primer 

and 0.25 Unit GoTaq® Flexi DNA polymerase (Promega). The 

ITS and GAPDH PCR mixtures differed from the original mix by 

containing 1 µM MgCl 2 

, the RPB2 and TEF1 PCR mixtures differed 

from the original mix by containing 2 µL genomic DNA and the 

RPB2 mixture differed from the original mix by containing 0.5 U 

instead of 0.25 U GoTaq® Flexi DNA polymerase. Conditions for 

PCR amplification consisted of an initial denaturation step of 5 min 

at 94 ºC followed by 35 cycles of 30 s at 94 ºC, 30 s at 48 ºC and 

90 s at 72 ºC for SSU, LSU, ITS and 40 cycles of 30 s at 94 ºC, 30 

s at 52 ºC / 59 ºC and 45 s at 72 ºC for TEF1 using respectively 

EF2 or EF1-986R as reverse primer and a final elongation step of 

7 min at 72 ºC. The partial RPB2 gene was obtained by using a 

touchdown PCR protocol of 5 cycles of 45 s at 94 ºC, 45 s at 60 

ºC and 2 min at 72 ºC, followed by 5 cycles with a 58 ºC annealing 

temperature and 30 cycles with a 54 ºC annealing temperature. 

The PCR products were sequenced in both directions using the 

PCR primers and the BigDye Terminator v. 3.1 Cycle Sequencing 

Kit (Applied Biosystems, Foster City, CA, USA), according to 

the manufacturer’s recommendations, and analysed with an ABI 

Prism 3730XL Sequencer (Applied Biosystems) according to the 

manufacturer’s instructions. Consensus sequences were computed 

from forward and reverse sequences using the BioNumerics v. 4.61 

software package (Applied Maths, St-Martens-Latem, Belgium). All 

generated sequences were deposited in GenBank (Table 1). 

172


Embellisia annulata CBS 302.84 

Pleospora tarda CBS 714.68 

Stemphylium herbarum CBS 191.86 

0.1 

0.98/80 Clathrospora heterospora CBS 175.52 

Comoclathris magna CBS 174.52 

1.0/99 Alternaria alternata CBS 916.96 

Alternaria limoniasperae CBS 102595 

0.79/62 Alternaria daucifolii CBS 118812 

1.0/89 Alternaria arborescens CBS 102605 

1.0/88 Alternaria gaisen CBS 632.93 

1.0/97 Alternaria tenuissima CBS 918.96 

Alternaria longipes CBS 540.94 

Nimbya gomphrenae CBS 108.27 

Alternaria alternantherae CBS 124392 

Alternaria perpunctulata CBS 115267 

1.0/99 0.88/39 Alternaria dauci CBS 117097 

0.91/41 Alternaria pseudorostrata CBS 119411 

0.76/35 Alternaria tagetica CBS 479.81 

Alternaria porri CBS 116698 

Alternaria solani CBS 116651 

0.98/85 

Alternaria macrospora CBS 117228 

0.76/98 Alternaria axiaeriisporifera CBS 118715 

Alternaria vaccariae CBS 116533 

1.0/96 Alternaria gypsophilae CBS 107.41 

Alternaria ellipsoidea CBS 119674 

1.0/95 

Alternaria nobilis CBS 116490 

Alternaria saponariae CBS 116492 

Alternaria juxtiseptata CBS 119673 

Alternaria vaccariicola CBS 118714 

Alternaria petroselini CBS 112.41 

Alternaria selini CBS 109382 

1.0/88 

Alternaria smyrnii CBS 109380 

Alternaria carotiincultae CBS 109381 

0.97/79 

Alternaria radicina CBS 245.67 

1.0/90 

Alternaria cinerariae CBS 116495 

Alternaria sonchi CBS 119675 

0.75/48 

Alternaria brassicae CBS 116528 

Alternaria helianthiinficiens CBS 117370 

Alternaria helianthiinficiens CBS 208.86 

Alternaria brassicicola CBS 118699 

0.90/99 Alternaria solidaccana CBS 118698 

Alternaria septorioides CBS 106.41 

Alternaria mimicula CBS 118696 

Embellisia conoidea CBS 132.89 

Ulocladium botrytis CBS 198.67 

Ulocladium tuberculatum CBS 202.67 

1.0/99 Ulocladium obovoideum CBS 101229 

Ulocladium cucurbitae CBS 483.81 

1.0/96 Ulocladium consortiale CBS 104.31 

Ulocladium brassicae CBS 121493 

1.0/99 Ulocladium atrum CBS 195.67 

0.98/97 

Ulocladium multiforme CBS 102060 

0.63/56 

Ulocladium solani CBS 123376 

Ulocladium subcucurbitae CBS 121491 

0.98/65 

Ulocladium cantlous CBS 123007 

Ulocladium arborescens CBS 115269 

1.0/89 

Ulocladium septosporum CBS 109.38 

1.0/96 

Ulocladium chartarum CBS 200.67 

1.0/91 

Ulocladium capsici CBS 120006 

Alternaria elegans CBS 109159 

Alternaria simsimi CBS 115265 

1.0/79 

Alternaria dianthicola CBS 116491 

1.0/98 

Alternaria cheiranthi CBS 109384 

1.0/47 

Alternaria resedae CBS 115.44 

Embellisia indefessa CBS 536.83 

0.96/90 Alternaria avenicola CBS 121459 

1.0/93 

Alternaria calycipyricola CBS 121545 

0.96/75 Alternaria photistica CBS 212.86 

0.97/56 

Alternaria panax CBS 482.81 

0.99/80 

Alternaria eryngii CBS 121339 

Alternaria thalictrigena CBS 121712 

Teretispora leucanthemi CBS 421.65 

Teretispora leucanthemi CBS 422.65 

Alternaria japonica CBS 118390 

Alternaria nepalensis CBS 118700 

1.0/88 Alternaria anigozanthi CBS 121920 

Embellisia leptinellae CBS 477.90 

1.0/96 Alternaria triglochinicola CBS 119676 

0.85/44 1.0/99 

Embellisia eureka CBS 193.86 

Alternaria geniostomatis CBS 118701 

Alternaria cumini CBS 121329 

1.0/99 

1.0/99 Embellisia hyacinthi CBS 416.71 

Embellisia novae-zelandiae CBS 478.90 

Embellisia proteae CBS 475.90 

Embellisia planifunda CBS 537.83 

0.88/67 

Embellisia tumida CBS 539.83 

Embellisia lolii CBS 115266 

Embellisia dennisii CBS 110533 

Embellisia dennisii CBS 476.90 

Undifilum bornmuelleri DAOM 231361 

Alternaria capsici-annui CBS 504.74 

1.0/98 

Ulocladium oudemansii CBS 114.07 


Sinomyces alternariae CBS 126989 

Alternaria argyranthemi CBS 116530 

1.0/76 Alternaria ethzedia CBS 197.86 

1.0/71 Alternaria oregonensis CBS 542.94 

1.0/74 Alternaria conjuncta CBS 196.86 

0.87/36 

0.96/55 Alternaria infectoria CBS 210.86 

Ybotromyces caespitosus CBS 177.80 

Chmelia slovaca CBS 567.66 

1.0/81 

Alternaria armoraciae CBS 118702 

Embellisia abundans CBS 534.83 

Chalastospora ellipsoidea CBS 121331 

Chalastospora cetera CBS 121340 

0.86/40 

Chalastospora obclavata CBS 124120 

0.55/* 

1.0/98 Alternaria molesta CBS 548.81 

Alternaria mouchaccae CBS 119671 

Alternaria limaciformis CBS 481.81 

Alternaria chlamydospora CBS 491.72 

1.0/64 

Embellisia phragmospora CBS 274.70 

Embellisia didymospora CBS 766.79 

0.93/56 

1.0/98 Embellisia chlamydospora CBS 341.71 

Embellisia tellustris CBS 538.83 

Embellisia allii CBS 339.71 

Alternaria soliaridae CBS 118387 

1.0/99 

Nimbya caricis CBS 480.90 

Nimbya scirpicola CBS 481.90 

Brachycladium penicillatum CBS 116608 

Crivellia papaveracea CBS 116607 

Brachycladium papaveris CBS 116606 

sect. Alternata 

sect. Alternantherae 

sect. Porri 

sect. Gypsophilae 

sect. Radicina 

sect. Sonchi 

• 

• 

sect. Brassicicola 

sect. Ulocladioides 

sect. Pseudoulocladium 

sect. Dianthicola 

sect. Cheiranthus 

sect. Panax 

• 

sect. Teretispora 

sect. Japonicae 

sect. Eureka 

sect. Embellisioides 

• 

sect. Undifilum 

sect. Ulocladium 

• 

sect. Infectoriae 

sect. Chalastospora 

sect. Phragmosporae 

sect. Embellisia 

• 

sect. Nimbya 

sect. Crivellia 

A 

l 

t 

e 

r 

n 

a 

r 

i 

a 

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. 

Phylogenetic analyses 

Multiple sequence alignments were generated with MAFFT v. 6.864b 

(http://mafft.cbrc.jp/alignment/server/index.html), and adjusted by 

eye. Two different datasets were used to estimate two phylogenies; 

an Alternaria complex phylogeny and a Pleosporineae family tree. 

The first tree focusses on the Alternaria complex, the second one 

was produced to place the genera Comoclathris, Clathrospora and 

Alternariaster in the context of the Alternaria complex. The relatives 

of the three genera were determined with standard nucleotide blast 

searches, with both the SSU and LSU sequences, against the 

nucleotide database in GenBank. This resulted in a selection of 35 

www.studiesinmycology.org 

173


Table 1. Isolates used in this study and their GenBank accession numbers. Bold accession numbers were generated in other studies. 

Old species 

name 

Alternaria 

alternantherae 

New species 

name 


alternantherae 


Section 

Strain Status 2 Host / Substrate Country Other collection 

GenBank accession numbers 

number 1 number 1 

SSU LSU RPB2 ITS GAPDH TEF1 

Althernantherae CBS 124392 Solanum melongena China HSAUP2798 KC584506 KC584251 KC584374 KC584179 KC584096 KC584633 

Alternaria alternata Alternaria alternata Alternata CBS 916.96 T Arachis hypogaea India EGS 34.016 KC584507 DQ678082 KC584375 AF347031 AY278808 KC584634 

Alternaria anigozanthi Alternaria anigozanthi Eureka CBS 121920 T Anigozanthus sp. Australia EGS 44.066 KC584508 KC584252 KC584376 KC584180 KC584097 KC584635 

Alternaria arborescens Alternaria arborescens Alternata CBS 102605 T Lycopersicon 

USA EGS 39.128 KC584509 KC584253 KC584377 AF347033 AY278810 KC584636 

esculentum 

Alternaria argyranthemi Alternaria argyranthemi CBS 116530 T Argyranthemum sp. New Zealand EGS 44.033 KC584510 KC584254 KC584378 KC584181 KC584098 KC584637 

Alternaria armoraciae Alternaria armoraciae Chalastospora CBS 118702 T Armoracia rusticana New Zealand EGS 51.064 KC584511 KC584255 KC584379 KC584182 KC584099 KC584638 

Alternaria avenicola Alternaria avenicola Panax CBS 121459 T Avena sp. Norway EGS 50.185 KC584512 KC584256 KC584380 KC584183 KC584100 KC584639 

New Zealand EGS 51.066 KC584513 KC584257 KC584381 KC584184 KC584101 KC584640 


axiaeriisporifera 


axiaeriisporifera 

Gypsophilae CBS 118715 T Gypsophila 

paniculata 

Alternaria brassicae Alternaria brassicae CBS 116528 R Brassica oleracea USA EGS 38.032 KC584514 KC584258 KC584382 KC584185 KC584102 KC584641 

Alternaria brassicicola Alternaria brassicicola Brassicicola CBS 118699 R Brassica oleracea USA EGS 42.002; ATCC 

96836 


calycipyricola 


calycipyricola 

Panax CBS 121545 T Pyrus communis China EGS 52.071; RGR 

96.0209 

KC584515 KC584259 KC584383 JX499031 KC584103 KC584642 

KC584516 KC584260 KC584384 KC584186 KC584104 KC584643 

Alternaria capsici-annui Alternaria capsici-annui Ulocladium CBS 504.74 Capsicum annuum – KC584517 KC584261 KC584385 KC584187 KC584105 KC584644 

Alternaria carotiincultae Alternaria carotiincultae Radicina CBS 109381 T Daucus carota USA EGS 26.010 KC584518 KC584262 KC584386 KC584188 KC584106 KC584645 

Alternaria cheiranthi Alternaria cheiranthi Cheiranthus CBS 109384 R Cheiranthus cheiri Italy EGS 41.188 KC584519 KC584263 KC584387 AF229457 KC584107 KC584646 

KC584520 KC584264 KC584388 KC584189 KC584108 KC584647 


chlamydospora 


chlamydospora 

Phragmosporae CBS 491.72 T Soil Egypt EGS 31.060; ATCC 

28045; IMI 156427 

Alternaria cinerariae Alternaria cinerariae Sonchi CBS 116495 R Ligularia sp. USA EGS 49.102 KC584521 KC584265 KC584389 KC584190 KC584109 KC584648 

Alternaria conjuncta Alternaria conjuncta Infectoriae CBS 196.86 T Pastinaca sativa Switzerland EGS 37.139 KC584522 KC584266 KC584390 FJ266475 AY562401 KC584649 

Alternaria cumini Alternaria cumini Eureka CBS 121329 T Cuminum cyminum India EGS 04.158a KC584523 KC584267 KC584391 KC584191 KC584110 KC584650 

Alternaria dauci Alternaria dauci Porri CBS 117097 R Daucus carota USA EGS 46.006 KC584524 KC584268 KC584392 KC584192 KC584111 KC584651 

Alternaria daucifolii Alternaria daucifolii Alternata CBS 118812 T Daucus carota USA EGS 37.050 KC584525 KC584269 KC584393 KC584193 KC584112 KC584652 

Alternaria dianthicola Alternaria dianthicola Dianthicola CBS 116491 R Dianthus × allwoodii New Zealand EGS 51.022 KC584526 KC584270 KC584394 KC584194 KC584113 KC584653 

KC584527 KC584271 KC584395 KC584195 KC584114 KC584654 

Alternaria elegans Alternaria elegans Dianthicola CBS 109159 T Lycopersicon 

esculentum 

Burkina Faso EGS 45.072; IMI 

374542 

Alternaria ellipsoidea Alternaria ellipsoidea Gypsophilae CBS 119674 T Dianthus barbatus USA EGS 49.104 KC584528 KC584272 KC584396 KC584196 KC584115 KC584655 

Alternaria eryngii Alternaria eryngii Panax CBS 121339 R Eryngium sp. – EGS 41.005 KC584529 KC584273 KC584397 JQ693661 AY562416 KC584656 

Alternaria ethzedia Alternaria ethzedia Infectoriae CBS 197.86 T Brassica napus Switzerland EGS 37.143 KC584530 KC584274 KC584398 AF392987 AY278795 KC584657 

Alternaria gaisen Alternaria gaisen Alternata CBS 632.93 R Pyrus pyrifolia cv. Nijiseiki 

Japan EGS 90.512 KC584531 KC584275 KC584399 KC584197 KC584116 KC584658 

174


Table 1. (Continued). 

Old species 

name 


geniostomatis 

New species 

name 


geniostomatis 


Section 





Eureka CBS 118701 T Geniostoma sp. New Zealand EGS 51.061 KC584532 KC584276 KC584400 KC584198 KC584117 KC584659 

Alternaria gypsophilae Alternaria gypsophilae Gypsophilae CBS 107.41 T Gypsophila elegans – EGS 07.025; IMI 

264349 


helianthiinficiens 







CBS 117370 R Helianthus annuus UK EGS 50.174; IMI 

388636 

KC584533 KC584277 KC584401 KC584199 KC584118 KC584660 

KC584534 KC584278 KC584402 KC584200 KC584119 KC584661 

CBS 208.86 T Helianthus annuus USA EGS 36.184 KC584535 KC584279 KC584403 JX101649 KC584120 EU130548 

Alternaria infectoria Alternaria infectoria Infectoriae CBS 210.86 T Triticum aestivum UK EGS 27.193 KC584536 KC584280 KC584404 DQ323697 AY278793 KC584662 

Alternaria japonica Alternaria japonica Japonicae CBS 118390 R Brassica chinensis USA EGS 50.099 KC584537 KC584281 KC584405 KC584201 KC584121 KC584663 

KC584538 KC584282 KC584406 KC584202 KC584122 KC584664 

Alternaria juxtiseptata Alternaria juxtiseptata Gypsophilae CBS 119673 T Gypsophila 

paniculata 

Australia EGS 44.015; DAR 

43414 

Alternaria limaciformis Alternaria limaciformis Phragmosporae CBS 481.81 T Soil UK EGS 07.086; IMI 

052976; QM 1790 


limoniasperae 


limoniasperae 

KC584539 KC584283 KC584407 KC584203 KC584123 KC584665 

Alternata CBS 102595 T Citrus jambhiri USA EGS 45.100 KC584540 KC584284 KC584408 FJ266476 AY562411 KC584666 

Alternaria longipes Alternaria longipes Alternata CBS 540.94 R Nicotiana tabacum USA EGS 30.033; QM 

9589 

Alternaria macrospora Alternaria macrospora Porri CBS 117228 T Gossypium 

barbadense 

Alternaria mimicula Alternaria mimicula Brassicicola CBS 118696 T Lycopersicon 

esculentum 

KC584541 KC584285 KC584409 AY278835 AY278811 KC584667 

USA EGS 50.190 KC584542 KC584286 KC584410 KC584204 KC584124 KC584668 

USA EGS 01.056; QM 26a KC584543 KC584287 KC584411 FJ266477 AY562415 KC584669 

Alternaria molesta Alternaria molesta Phragmosporae CBS 548.81 T Phocaena phocaena Denmark EGS 32.075 KC584544 KC584288 KC584412 KC584205 KC584125 KC584670 

Alternaria mouchaccae Alternaria mouchaccae Phragmosporae CBS 119671 T Soil Egypt EGS 31.061 KC584545 KC584289 KC584413 KC584206 AY562399 KC584671 

Alternaria nepalensis Alternaria nepalensis Japonicae CBS 118700 T Brassica sp. Nepal EGS 45.073; IMI KC584546 KC584290 KC584414 KC584207 KC584126 KC584672 

374543 

Alternaria nobilis Alternaria nobilis Gypsophilae CBS 116490 R Dianthus caryophyllus New Zealand EGS 51.027; NZMAF KC584547 KC584291 KC584415 KC584208 KC584127 KC584673 

Lynfield 743 

Alternaria oregonensis Alternaria oregonensis Infectoriae CBS 542.94 T Triticum aestivum USA EGS 29.194 KC584548 KC584292 KC584416 FJ266478 FJ266491 KC584674 

Alternaria panax Alternaria panax Panax CBS 482.81 R Aralia racemosa USA EGS 29.180 KC584549 KC584293 KC584417 KC584209 KC584128 KC584675 

USA KC584550 KC584294 KC584418 KC584210 KC584129 KC584676 


perpunctulata 


perpunctulata 

Althernantherae CBS 115267 T Alternanthera 

philoxeroides 

Alternaria petroselini Alternaria petroselini Radicina CBS 112.41 T Petroselinum sativum – EGS 06.196 KC584551 KC584295 KC584419 KC584211 KC584130 KC584677 

Alternaria photistica Alternaria photistica Panax CBS 212.86 T Digitalis purpurea UK EGS 35.172 KC584552 KC584296 KC584420 KC584212 KC584131 KC584678 

Alternaria porri Alternaria porri Porri CBS 116698 R Allium cepa USA EGS 48.147 KC584553 KC584297 KC584421 DQ323700 KC584132 KC584679 


175



Old species 

name 


pseudorostrata 

New species 

name 


pseudorostrata 


Section 




Porri CBS 119411 T Euphorbia 

pulcherrima 

Alternaria radicina Alternaria radicina Radicina CBS 245.67 T Daucus carota USA EGS 03.145; ATCC 

6503; IMI 124939; 

QM 1301; QM 6503 


USA EGS 42.060 KC584554 KC584298 KC584422 JN383483 AY562406 KC584680 

KC584555 KC584299 KC584423 KC584213 KC584133 KC584681 

“Alternaria resedae” Alternaria sp. Cheiranthus CBS 115.44 Reseda odorata – EGS 07.030 KC584556 KC584300 KC584424 KC584214 KC584134 KC584682 

Alternaria saponariae Alternaria saponariae Gypsophilae CBS 116492 R Saponaria officinalis USA EGS 49.199 KC584557 KC584301 KC584425 KC584215 KC584135 KC584683 

Alternaria selini Alternaria selini Radicina CBS 109382 T Petroselinum crispum Saudi Arabia EGS 25.198; IMI KC584558 KC584302 KC584426 AF229455 AY278800 KC584684 

137332 

Alternaria septorioides Alternaria septorioides Brassicicola CBS 106.41 T Reseda odorata Netherlands EGS 52.089; MUCL KC584559 KC584303 KC584427 KC584216 KC584136 KC584685 

20298 

Alternaria simsimi Alternaria simsimi Dianthicola CBS 115265 T Sesamum indicum Argentina EGS 13.110 KC584560 KC584304 KC584428 JF780937 KC584137 KC584686 

Alternaria smyrnii Alternaria smyrnii Radicina CBS 109380 R Smyrnium olusatrum UK EGS 37.093 KC584561 KC584305 KC584429 AF229456 KC584138 KC584687 

Alternaria solani Alternaria solani Porri CBS 116651 R Solanum tuberosum USA EGS 45.020 KC584562 KC584306 KC584430 KC584217 KC584139 KC584688 

Alternaria soliaridae Alternaria soliaridae CBS 118387 T Soil USA EGS 33.024 KC584563 KC584307 KC584431 KC584218 KC584140 KC584689 

Alternaria solidaccana Alternaria solidaccana Brassicicola CBS 118698 T Soil Bangladesh EGS 36.158; IMI 

049788 

Alternaria sonchi Alternaria sonchi Sonchi CBS 119675 R Sonchus asper Canada EGS 43.131; IMI 

366167 

KC584564 KC584308 KC584432 KC584219 KC584141 KC584690 

KC584565 KC584309 KC584433 KC584220 KC584142 KC584691 

Alternaria tagetica Alternaria tagetica Porri CBS 479.81 R Tagetes erecta UK EGS 33.081 KC584566 KC584310 KC584434 KC584221 KC584143 KC584692 

Alternaria tenuissima Alternaria tenuissima Alternata CBS 918.96 R Dianthus sp. UK EGS 34.015 KC584567 KC584311 KC584435 AF347032 AY278809 KC584693 

Alternaria thalictrigena Alternaria thalictrigena CBS 121712 T Thalictrum sp. Germany KC584568 KC584312 KC584436 EU040211 KC584144 KC584694 


triglochinicola 


triglochinicola 

Eureka CBS 119676 T Triglochin procera Australia EGS 41.070 KC584569 KC584313 KC584437 KC584222 KC584145 KC584695 

Alternaria vaccariae Alternaria vaccariae Gypsophilae CBS 116533 R Vaccaria hispanica USA EGS 47.108 KC584570 KC584314 KC584438 KC584223 KC584146 KC584696 

Alternaria vaccariicola Alternaria vaccariicola Gypsophilae CBS 118714 T Vaccaria hispanica USA EGS 46.003; ATCC KC584571 KC584315 KC584439 KC584224 KC584147 KC584697 

26038 

Alternariaster helianthi Alternariaster helianthi CBS 119672 R Helianthus sp. USA EGS 36.007 KC584626 KC584368 KC584493 

Alternariaster helianthi Alternariaster helianthi CBS 327.69 Helianthus annuus – KC584627 KC584369 KC584494 

Ascochyta pisi Ascochyta pisi CBS 126.54 Pisum sativum Netherlands EU754038 DQ678070 DQ677967 

Boeremia exigua Boeremia exigua CBS 431.74 Solanum tuberosum Netherlands PD 74/2447 EU754084 EU754183 GU371780 

Brachycladium 

papaveris 

Brachycladium 

penicillatum 

Alternaria papavericola Crivellia CBS 116606 T Papaver somniferum USA KC584579 KC584321 KC584446 FJ357310 FJ357298 KC584705 

Alternaria penicillata Crivellia CBS 116608 T Papaver rhoeas Austria DAOM 230457 KC584572 KC584316 KC584440 FJ357311 FJ357299 KC584698 

176



Old species 

name 

New species 

name 


Section 

Chaetodiplodia sp. Chaetodiplodia sp. CBS 453.68 Halimione 

portulacoides 

Chaetosphaeronema 

hispidulum 

Chaetosphaeronema 

hispidulum 





Netherlands DQ678001 DQ678054 KC584499 

CBS 216.75 Anthyllis vulneraria Germany EU754045 EU754144 GU371777 

Chalastospora cetera Alternaria cetera Chalastospora CBS 121340 T Elymus scabrus Australia EGS 41.072 KC584573 KC584317 KC584441 JN383482 AY562398 KC584699 

Chalastospora 

Alternaria breviramosa Chalastospora CBS 121331 T Triticum sp. Australia KC584574 KC584318 KC584442 FJ839608 KC584148 KC584700 

ellipsoidea 

Chalastospora 

Alternaria obclavata Chalastospora CBS 124120 T Air USA EGS 12.128 KC584575 FJ839651 KC584443 KC584225 KC584149 KC584701 

obclavata 

Chmelia slovaca Alternaria slovaca Infectoriae CBS 567.66 T Human Slovakia ATCC 24279 KC584576 KC584319 KC584444 KC584226 KC584150 KC584702 

Clathrospora elynae Clathrospora elynae CBS 161.51 Carex curvula Switzerland KC584628 KC584370 KC584495 

Clathrospora elynae Clathrospora elynae CBS 196.54 Carex curvula Switzerland KC584629 KC584371 KC584496 

Clathrospora 

heterospora 

Cochliobolus 

heterostrophus 

Alternaria sp. Alternata CBS 175.52 Juncus mertensianus USA EGS 35.1619; IMI 

068085; QM 1277 

Cochliobolus 

heterostrophus 

CBS 134.39 Zea mays – DSM 1149 AY544727 AY544645 DQ247790 

Cochliobolus sativus Cochliobolus sativus DAOM 226212 Hordeum vulgare Canada DQ677995 DQ678045 DQ677939 

KC584577 KC584320 KC584445 KC584227 KC584151 KC584703 

Comoclathris magna Alternaria sp. Alternata CBS 174.52 Anemone occidentalis USA EGS 39.1613; IMI 

068086; QM 1278 

Comoclathris 

compressa 

Comoclathris 

compressa 

Coniothyrium 

palmarum 

Comoclathris 

compressa 

Comoclathris 

compressa 

Coniothyrium 

palmarum 

CBS 156.53 Castilleja miniata USA EGS No. C-20285-I KC584630 KC584372 KC584497 

CBS 157.53 Ligusticum 

purpureum 

USA EGS No. 1952a-1633 KC584631 KC584373 KC584498 

CBS 400.71 Chamaerops humilis Italy EU754054 EU754153 DQ677956 

KC584578 DQ678068 DQ677964 KC584228 KC584152 KC584704 

Crivellia papaveracea Alternaria penicillata Crivellia CBS 116607 T Papaver rhoeas Austria DAOM 230456 KC584580 KC584322 KC584447 KC584229 KC584153 KC584706 

KC793336 KC793338 DQ470924 

Dendryphiella 

arenariae 

Paradendryphiella 

arenariae 

Dendryphiella salina Paradendryphiella 

salina 

CBS 181.58 T Coastal sand France DAOM 63738; IMI 

067735; MUCL 4129 

CBS 142.60 Spartina sp. UK MUCL 9639 KC793337 KC793339 KC793340 

Embellisia abundans Alternaria abundans Chalastospora CBS 534.83 T Fragaria sp. New Zealand EGS 29.159 KC584581 KC584323 KC584448 JN383485 KC584154 KC584707 

Embellisia allii Alternaria embellisia Embellisia CBS 339.71 R Allium sativum USA ATCC 22412; IMI KC584582 KC584324 KC584449 KC584230 KC584155 KC584708 

155707; MUCL 

18571; QM 8609 

Embellisia annulata Cicatricea salina CBS 302.84 T Cancer pagurus North Sea, 

Skagerrak 

KC584583 KC584325 KC584450 JN383486 JN383467 KC584709 


177



Old species 

name 

Embellisia 

chlamydospora 

New species 

name 


chlamydosporigena 


Section 




Embellisia CBS 341.71 R Air USA EGS 10.073; ATCC 

22409; IMI 155709; 

MUCL 18573; QM 

7287 


KC584584 KC584326 KC584451 KC584231 KC584156 KC584710 

Embellisia conoidea Alternaria conoidea Brassicicola CBS 132.89 Ricinus communis Saudi Arabia KC584585 KC584327 KC584452 AF348226 FJ348227 KC584711 

Embellisia dennisii Alternaria dennisii CBS 110533 Senecio jacobaea New Zealand KC584586 KC584328 KC584453 KC584232 KC584157 KC584712 

Embellisia dennisii Alternaria dennisii CBS 476.90 T Senecio jacobaea Isle of Man IMI 151744 KC584587 KC584329 KC584454 JN383488 JN383469 KC584713 

Embellisia 

didymospora 

Alternaria didymospora Phragmosporae CBS 766.79 Seawater Adriatic Sea KC584588 KC584330 KC584455 FJ357312 FJ357300 KC584714 

Embellisia eureka Alternaria eureka Eureka CBS 193.86 T Medicago rugosa Australia IMI 273162 KC584589 KC584331 KC584456 JN383490 JN383471 KC584715 

Embellisia hyacinthi Alternaria hyacinthi Embellisioides CBS 416.71 T Hyacinthus orientalis Netherlands EGS 19.102; IMI KC584590 KC584332 KC584457 KC584233 KC584158 KC584716 

279179 

Embellisia indefessa Alternaria indefessa Cheiranthus CBS 536.83 T Soil USA EGS 30.195 KC584591 KC584333 KC584458 KC584234 KC584159 KC584717 

Embellisia leptinellae Alternaria leptinellae Eureka CBS 477.90 T Leptinella dioica New Zealand EGS 39.101 KC584592 KC584334 KC584459 KC584235 KC584160 KC584718 

Embellisia lolii Alternaria lolii Embellisioides CBS 115266 T Lolium perenne New Zealand KC584593 KC584335 KC584460 JN383492 JN383473 KC584719 

Embellisia novaezelandiae 

Embellisia 

phragmospora 

Alternaria botryospora Embellisioides CBS 478.90 T Leptinella dioica New Zealand EGS 39.099 KC584594 KC584336 KC584461 AY278844 AY278831 KC584720 


phragmospora 

Phragmosporae CBS 274.70 T Soil The 

netherlands 

EGS 27.098; ATCC 

18914 

KC584595 KC584337 KC584462 JN383493 JN383474 KC584721 

Embellisia planifunda Alternaria planifunda Embellisioides CBS 537.83 T Triticum aestivum Australia IMI 115034 KC584596 KC584338 KC584463 FJ357315 FJ357303 KC584722 

Embellisia proteae Alternaria proteae Embellisioides CBS 475.90 T Protea sp. Australia IMI 320290; IMI KC584597 KC584339 KC584464 AY278842 KC584161 KC584723 

341684 

Embellisia tellustris Alternaria tellustris Embellisia CBS 538.83 T Soil USA EGS 33.026 KC584598 KC584340 KC584465 FJ357316 AY562419 KC584724 

Embellisia tumida Alternaria tumida Embellisioides CBS 539.83 T Triticum aestivum Australia KC584599 KC584341 KC584466 FJ266481 FJ266493 KC584725 

CBS 115.96 Chenopodium album Netherlands PD 94/1576 EU754089 EU754188 GU371775 

Heterospora 

chenopodii 

Heterospora 

chenopodii 

Julella avicenniae Julella avicenniae BCC 18422 Mangrove wood Thailand GU371831 GU371823 GU371787 

Leptosphaerulina 

australis 

Leptosphaerulina 

australis 

CBS 317.83 Eugenia aromatica Indonesia GU296160 GU301830 GU371790 

Loratospora aestuarii Loratospora aestuarii JK 5535B Juncus roemerianus USA GU296168 GU301838 GU371760 

Neophaeosphaeria 

filamentosa 

Neophaeosphaeria 

filamentosa 

CBS 102202 Yucca rostrata Mexico GQ387516 GQ387577 GU371773 

Nimbya caricis Alternaria caricis Nimbya CBS 480.90 T Carex hoodii USA EGS 13.094 KC584600 KC584342 KC584467 AY278839 AY278826 KC584726 

“Nimbya gomphrenae” Alternaria sp. Alternata CBS 108.27 Gomphrena globosa – KC584601 KC584343 KC584468 KC584236 KC584162 KC584727 

Nimbya scirpicola Alternaria scirpicola Nimbya CBS 481.90 R Scirpus sp. UK EGS 19.042 KC584602 KC584344 KC584469 KC584237 KC584163 KC584728 

178



Old species 

name 

Ophiosphaerella 

herpotricha 

Paraleptosphaeria 

dryadis 

Peyronellaea 

glomerata 

Peyronellaea zeaemaydis 

Phaeosphaeria 

ammophilae 

Phaeosphaeria 

avenaria 

Phaeosphaeria 

eustoma 

New species 

name 

Ophiosphaerella 

herpotricha 

Paraleptosphaeria 

dryadis 

Peyronellaea 

glomerata 

Peyronellaea zeaemaydis 

Phaeosphaeria 

ammophilae 

Phaeosphaeria 

avenaria 

Phaeosphaeria 

eustoma 


Section 





CBS 620.86 Bromus erectus Switzerland ETH 9373 DQ678010 DQ678062 DQ677958 

CBS 643.86 Dryas octopetala Switzerland ETH 9446 KC584632 GU301828 GU371733 

CBS 528.66 Chrysanthemum sp. Netherlands PD 63/590 EU754085 EU754184 GU371781 

CBS 588.69 T Zea mays USA EU754093 EU754192 GU371782 

CBS 114595 Ammophila arenaria Sweden UPSC 3568 GU296185 GU304859 GU371724 

DAOM 226215 Avena sativa Canada OSC 100096 AY544725 AY544684 DQ677941 

CBS 573.86 Dactylis glomerata Switzerland ETH 9239 DQ678011 DQ678063 DQ677959 

Phoma complanata Phoma complanata CBS 268.92 Anglica sylvestris Netherlands PD 75/3 EU754081 EU754180 GU371778 

Phoma herbarum Phoma herbarum CBS 276.37 Wood pulp Sweden DQ678014 DQ678066 DQ677962 

Plenodomus lingam Plenodomus lingam DAOM 229267 Brassica sp. France DQ470993 DQ470946 DQ470894 

Pleospora betae Pleospora betae CBS 109410 Beta vulgaris Netherlands PD 77/113 EU754079 EU754178 GU371774 

Pleospora calvescens Pleospora calvescens CBS 246.79 Atriplex hastata Germany PD 77/655 EU754032 EU754131 KC584500 

Pleospora chenopodii Pleospora chenopodii CBS 206.80 Chenopodium quinoa Bolivia PD 74/1022 JF740095 JF740266 KC584501 

Pleospora fallens Pleospora fallens CBS 161.78 Olea europaea New Zealand GU238215 GU238074 KC584502 

Pleospora halimiones Pleospora halimiones CBS 432.77 Halimione 

portulacoides 

Netherlands IMI 282137 JF740096 JF740267 KC584503 

Pleospora incompta Pleospora incompta CBS 467.76 Olea europaea Greece GU23822 GU238087 KC584504 

Pleospora tarda Pleospora tarda CBS 714.68 T Medicago sativa Canada EGS 04.118C; IMI 

135456; MUCL 

11717; QM 1379 

Pleospora typhicola Pleospora typhicola CBS 132.69 Typha angustifolia Netherlands JF740105 JF740325 KC584505 

KC584603 KC584345 AF107804 KC584238 AF443881 KC584729 

Pyrenochaeta nobilis Pyrenochaeta nobilis CBS 407.76 T Laurus nobilis Italy EU754107 DQ678096 DQ677991 

Pyrenophora 

phaeocomes 

Saccothecium 

sepincola 

Setomelanomma 

holmii 

Pyrenophora 

phaeocomes 

Saccothecium 

sepincola 

Setomelanomma 

holmii 

DAOM 222769 Calamagrostis villosa Switzerland DQ499595 DQ499596 DQ497614 

CBS 278.32 Ribes nigrum USA GU296195 GU301870 GU371745 

CBS 110217 Picea pungens USA GU296196 GQ37633 GU371800 


179



Old species 

name 

New species 

name 


Section 





Sinomyces alternariae Alternaria alternariae Ulocladium CBS 126989 T Daucus carota USA EGS 46.004 KC584604 KC584346 KC584470 AF229485 AY278815 KC584730 

Stemphylium herbarum Stemphylium herbarum CBS 191.86 T Medicago sativa India EGS 36.138; IMI 276975 

GU238232 GU238160 KC584471 KC584239 AF443884 KC584731 

KC584605 KC584347 KC584472 KC584240 KC584164 KC584732 

Teretispora 

leucanthemi 

Teretispora 

leucanthemi 

Ulocladium 

arborescens 

Alternaria leucanthemi Teretispora CBS 421.65 T Chrysanthemum 

maximum 

Alternaria leucanthemi CBS 422.65 R Chrysanthemum 

maximum 

Netherlands ATCC 16028; IFO 

9085; IMI 111986; 

QM 7227 

USA EGS 17.063; ATCC 

16029; IMI 111987; 

QM 8579 

KC584606 KC584348 KC584473 KC584241 KC584165 KC584733 

Alternaria aspera Pseudoulocladium CBS 115269 T Pistacia vera Japan IMI 369777 KC584607 KC584349 KC584474 KC584242 KC584166 KC584734 

Ulocladium atrum Alternaria atra Ulocladioides CBS 195.67 T Soil USA ATCC 18040; IMI 

124944; QM 8408 

Ulocladium botrytis Alternaria botrytis Ulocladium CBS 197.67 T Contaminant USA ATCC 18042; IMI 

124942; MUCL 

18556; QM 7878 

Ulocladium botrytis Alternaria sp. Ulocladioides CBS 198.67 R Soil USA ATCC 18043; IMI 

124949; MUCL 

18557; QM 8619 

Ulocladium brassicae Alternaria brassicaepekinensis 

KC584608 KC584350 KC584475 AF229486 KC584167 KC584735 

KC584609 KC584351 KC584476 KC584243 KC584168 KC584736 

KC584610 KC584352 KC584477 AF229487 KC584169 KC584737 

Ulocladioides CBS 121493 T Brassica pekinensis China HSAUPwy0037 KC584611 KC584353 KC584478 KC584244 KC584170 KC584738 

Ulocladium cantlous Alternaria cantlous Ulocladioides CBS 123007 T Cucumis melo China HSAUP0209 KC584612 KC584354 KC584479 KC584245 KC584171 KC584739 

Ulocladium capsici Alternaria concatenata Pseudoulocladium CBS 120006 T – – HSAUPIII 0 

0035 KC584613 KC584355 KC584480 KC584246 AY762950 KC584740 

Ulocladium chartarum Alternaria chartarum Pseudoulocladium CBS 200.67 T Populus sp. Canada ATCC 18044; DAOM KC584614 KC584356 KC584481 AF229488 KC584172 KC584741 

59616b; IMI 124943; 

MUCL 18564; QM 

8328 

Ulocladium consortiale Alternaria consortialis Ulocladioides CBS 104.31 T – – KC584615 KC584357 KC584482 KC584247 KC584173 KC584742 

Ulocladium cucurbitae Alternaria cucurbitae Ulocladioides CBS 483.81 R Cucumis sativus New Zealand EGS 31.021; LEV KC584616 KC584358 KC584483 FJ266483 AY562418 KC584743 

7067 

Ulocladium multiforme Alternaria multiformis Ulocladioides CBS 102060 T Soil Canada KC584617 KC584359 KC584484 FJ266486 KC584174 KC584744 

Ulocladium 

Alternaria obovoidea Ulocladioides CBS 101229 Cucumis sativus New Zealand KC584618 KC584360 KC584485 FJ266487 FJ266498 KC584745 

obovoideum 

Ulocladium oudemansii Alternaria oudemansii Ulocladium CBS 114.07 T – – ATCC 18047; IMI KC584619 KC584361 KC584486 FJ266488 KC584175 KC584746 

124940; MUCL 

18563; QM 1744 

Ulocladium 

septosporum 

Alternaria septospora Pseudoulocladium CBS 109.38 Wood Italy KC584620 KC584362 KC584487 FJ266489 FJ266500 KC584747 

180







Section 

New species 

name 

Old species 

name 


China HSAUP 0521 KC584621 KC584363 KC584488 KC584248 KC584176 KC584748 

Ulocladium solani Alternaria heterospora Ulocladioides CBS 123376 T Lycopersicon 

esculentum 

China KC584622 KC584364 KC584489 KC584249 EU855803 KC584749 

Ulocladioides CBS 121491 T Chenopodium 

glaucum 


subcucurbitae 

Ulocladium 

subcucurbitae 

KC584623 KC584365 KC584490 FJ266490 KC584177 KC584750 

Alternaria terricola Ulocladioides CBS 202.67 T Soil USA ATCC 18048; IMI 

124947; MUCL 

18560; QM 8614 

Ulocladium 

tuberculatum 

Undifilum bornmuelleri Alternaria bornmuelleri Undifilum DAOM 231361 T Securigera varia Austria DAOM 231361 KC584624 KC584366 KC584491 FJ357317 FJ357305 KC584751 

Ybotromyces 

Alternaria caespitosa Infectoriae CBS 177.80 T Human Spain KC584625 KC584367 KC584492 KC584250 KC584178 KC584752 

caespitosus 

1 

ATCC: American Type Culture Collection, Manassas, VA, USA; BCC: BIOTEC Culture Collection, Thailand; CBS: Culture collection of the Centraalbureau voor Schimmelcultures, Fungal Biodiversity Centre, Utrecht, The Netherlands; DAOM: Canadian 

Collection of Fungal Cultures, Ottawa, Canada; DAR: Plant Pathology Herbarium, Orange Agricultural Institute, Australia; DSM: German Collection of Microorganisms and Cell Cultures, Leibniz Institute, Braunschweig, Germany; EGS: Personal collection 

of Dr. E.G. Simmons; ETH: Swiss Federal Institute of Technology, Switzerland; HSAUP: Department of Plant Pathology, Shandong Agricultural University, China; IFO: Institute for Fermentation Culture Collection, Osaka, Japan; IMI: Culture collection of 

CABI Europe UK Centre, Egham UK; JK: Personal collection of Dr. J. Kohlmeyer; LEV: Plant Health and Diagnostic Station, Levin, New Zealand; MUCL: (Agro)Industrial Fungi and Yeast Collection of the Belgian Co-ordinated Collections of Micro-organisms 

(BCCM), Louvain-la Neuve, Belgium; NZMAF: New Zealand Ministry of Agriculture and Forestry; OSC: Oregon State University Herbarium, USA; PD: Plant Protection Service, Wageningen, The Netherlands; RGR: Personal collection of Dr. R.G. Roberts; 

UPSC: Uppsala University Culture Collection, Sweden; QM: Quarter Master Culture Collection, Amherst, MA, USA. 

2 

T: ex-type strain; R: representative strain. 

species (Table 1) for which the SSU, LSU and RPB2 sequence 

data set was present or could be completed. Blast searches with 

Embellisia annulata gave hits with two marine Dendryphiella species, 

Dendryphiella arenariae and Dendryphiella salina, which we also 

included. Phylogenetic analyses of the sequence data consisted of 

Bayesian and Maximum likelihood analyses of both the individual 

data partitions as well as the combined aligned dataset. Bayesian 

analyses were performed with MrBayes v. 3.2.1 (Huelsenbeck & 

Ronquist 2001, Ronquist & Huelsenbeck 2003). The Markov Chain 

Monte Carlo (MCMC) analysis used four chains and started from a 

random tree topology. The sample frequency was set at 100 and the 

temperature value of the heated chain was 0.1. The temperature 

value was lowered to 0.05 when the average standard deviation of 

split frequencies did not fall below 0.01 after 5M generations (RPB2 

and Pleosporineae phylogeny). Burn-in was set to 25 % after which 

the likelihood values were stationary. Maximum likelihood analyses 

including 500 bootstrap replicates were run using RAxML v. 7.2.6 

(Stamatakis & Alachiotis 2010). The online tool Findmodel (http:// 

www.hiv.lanl.gov/content/sequence/findmodel/findmodel.html) was 

used to determine the best nucleotide substitution model for each 

partition. For the SSU (Pleosporineae family tree), LSU, ITS, RPB2 

and TEF1 partitions a GTR model with a gamma-distributed rate 

variation was suggested, and for the SSU (Alternaria complex) 

and GAPDH partitions a TrN model with gamma-distributed rate 

variation. Sequences of Stemphylium herbarum (CBS 191.86) 

were used as outgroup in the Alternaria phylogeny and those of 

Jullella avenicae (BCC 18422) in the Pleosporineae phylogeny. The 

resulting trees were printed with TreeView v. 1.6.6 (Page 1996) and 

together with the alignments deposited into TreeBASE (http://www. 

treebase.org). 

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 


181


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. 

1-region 2-region 3-region 6-region 1-region 2-region 3-region 6-region 

SSU LSU ITS GAPDH RPB2 TEF GAPDH GAPDH RPB2 GAPDH SSU SSU LSU ITS GAPDH RPB2 TEF GAPDH GAPDH RPB2 GAPDH ALL 

RPB2 TEF1 TEF1 RPB2 LSU RPB2 TEF1 TEF1 RPB2 LSU 

TEF1 ITS TEF1 ITS 

GAPDH GAPDH 

RPB2 RPB2 

TEF1 TEF1 

Aligned length 1021 851 499 573 786 269 1359 842 1055 1628 3999 1021 851 499 573 786 269 1359 842 1055 1628 3999 

Unique site patterns 45 57 148 272 296 224 568 496 520 792 1042 45 57 148 272 296 224 568 496 520 792 1042 

No. of sampled trees 39002 31578 75002 23702 56028 12452 10128 13728 44852 5778 16278 

(post burnin) 

Bayesian Posterior Probabilities RAxML bootstrap support 

Sect. Alternantherae * * 

Sect. Alternata 

Sect. Brassicicola 

Sect. Chalastospora * * 

Sect. Cheiranthus * * * * * * * * 

Sect. Crivellia * * 

Sect. Dianthicola * * * * 

Sect. Embellisia * * 

Sect. Embellisioides * * * * * * 

Sect. Eureka * * * * * * 

Sect. Gypsophilae * * 

Sect. Infectoriae * * * * 

Sect. Japonicae 

Sect. Nimbya 

Sect. panax * * 

Sect. Phragmosporae * * * * 

Sect. Porri * * 

Sect. Pseudoulocladium * * * * * * * * 

Sect. Radicina 

Sect. Sonchi * * * * 

Sect. Teretispora 

Sect. Ulocladioides * * * * 

Sect. Ulocladium * * 

1.00 0.95–0.99 0.90–0.94 0.80–0.89 0.70–0.79 100 95–99 90–94 80–89 70–79 

0.50–0.69 0.25–0.49 0.01–0.24 no support 50–69 25–49 1–24 no support 

*= section not complete *= section not complete 

182


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 

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. 

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 wellsupported 

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. 

Taxonomy 

Based on DNA sequence data in combination with a review 

of literature and morphology, the species within the Alternaria 

clade are all recognised here as Alternaria (Fig 1). This puts the 

genera Allewia, Brachycladium, Chalastospora, Chmelia, Crivellia, 

Embellisia, Lewia, Nimbya, Sinomyces, Teretispora, Ulocladium, 

Undifilum and Ybotromyces in synonymy with Alternaria, resulting 

in the proposal of 32 new combinations, 10 new names and the 

resurrection of 10 names. Species of Alternaria were assigned 

to 24 Alternaria sections, of which 16 are newly described, and 

six monotypic lineages. The (emended) description of the genus 

Alternaria, the Alternaria sections and monotypic lineages with 

new Alternaria names and name combinations are treated below 

in alphabetical order. Finally the description of the new genus 

Paradendryphiella is also provided. 

Alternaria Nees, Syst. Pilze (Würzburg): 72. 1816 [1816– 

1817]. 

= Elosia Pers., Mycol. Eur. (Erlanga) 1: 12. 1822. 

= Macrosporium Fr., Syst. Mycol. (Lundae) 3: 373. 1832. 

= Rhopalidium Mont., Ann. Sci. Nat., Bot., Sér. 2, 6: 30. 1836. 

= Brachycladium Corda, Icon. Fungorum hucusque Cogn. (Prague) 2: 14. 

1838. 

= Ulocladium Preuss, Linnaea 24: 111. 1851. 

= Chmelia Svob.-Pol., Biologia (Bratislava) 21: 82. 1966. 

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

= Trichoconiella B.L. Jain, Kavaka 3: 39. 1976 [1975]. 

= Botryomyces de Hoog & C. Rubio, Sabouraudia 20: 19. 1982. (nom. illegit.) 

= 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. 

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

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

= Teretispora E.G. Simmons, CBS Biodiversity Ser. (Utrecht) 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 grey, dark blackish brown or black. Mycelium 

immersed or partly superficial; hyphae colourless, olivaceousbrown 

or brown. Stroma rarely formed. Setae and hyphopodia 

absent. Conidiophores macronematous, mononematous, simple 

or irregularly and loosely branched, pale brown or brown, solitary 

or in fascicles. Conidiogenous cells integrated, terminal becoming 

intercalary, polytretic, sympodial, or sometimes monotretic, 

cicatrized. Conidia catenate or solitary, dry, ovoid, obovoid, 

cylindrical, narrowly ellipsoid or obclavate, beaked or non-beaked, 

pale or medium olivaceous-brown to brown, smooth or verrucose, 

with transverse and with or without oblique or longitudinal septa. 

Septa can be thick, dark and rigid and an internal cell-like structure 

can be formed. Species with meristematic growth are known. 

Ascomata small, solitary to clustered, erumpent to (nearly) 

superficial at maturity, globose to ovoid, dark brown, smooth, 

apically papillate, ostiolate. Papilla short, blunt. Peridium thin. 

Hamathecium of cellular pseudoparaphyses. Asci few to many per 

ascoma, (4–6–)8-spored, basal, bitunicate, fissitunicate, cylindrical 

to cylindro-clavate, straight or somewhat curved, with a short, 

furcate pedicel. Ascospores muriform, ellipsoid to fusoid, slightly 

constricted at septa, yellow-brown, without guttules, smooth, 3–7 

transverse septa, 1–2 series of longitudinal septa through the 

two original central segments, end cells without septa, or with 1 

longitudinal or oblique septum, or with a Y-shaped pair of septa. 

Type species: Alternaria alternata (Fr.) Keissl. 


183


0.96/55 

1.0/54 

0.52/28 

1.0/97 

Alternaria brassicae CBS 116528 

Alternaria sonchi CBS 119675 

Alternaria helianthiinficiens CBS 208.86 

Embellisia eureka CBS 193.86 

Embellisia hyacinthi CBS 416.71 

1.0/65 

1.0/95 Alternaria dianthicola CBS 116491 

1.0/83 

0.97/66 

Ulocladium chartarum CBS 200.67 


Alternaria cheiranthi CBS 109384 

1.0/91 

0.97/50 

Alternaria alternantherae CBS 124392 

Alternaria alternata CBS 916.96 

A 

l 

0.99/34 Alternaria porri CBS 116698 

t 

0.56/23 

Alternaria radicina CBS 245.67 

e 

0.68/27 

1.0/89 Alternaria saponariae CBS 116492 r 

1.0/57 

0.92/25 

Alternaria thalictrigena CBS 121712 

Alternaria panax CBS 482.81 

n 

a 

Teretispora leucanthemi CBS 421.65 r 

Alternaria brassicicola CBS 118699 i 

0.66/- 

0.97/53 

Alternaria japonica CBS 118390 

Embellisia dennisii CBS 110533 

Sinomyces alternariae CBS 126989 

a 

1.0/9 

0.99/73 

Alternaria argyranthemi CBS 116530 

Undifilum bornmuelleri DAOM 231361 

Chalastospora cetera CBS 121340 

0.82/23 

0.99/38 

0.98/95 

Alternaria infectoria CBS 210.86 

Alternaria soliaridae CBS 118387 

Nimbya scirpicola CBS 481.90 

Embellisia allii CBS 339.71 

Embellisia phragmospora CBS 274.70 

Crivellia papaveracea CBS 116607 

Pyrenophora phaeocomes DAOM 222769 

Pleosporaceae 

Embellisia annulata CBS 302.84 

Dendryphiella salina CBS 142.60 Paradendryphiella gen. nov. 

Dendryphiella arenariae CBS 181.58 

Pleospora herbarum CBS 191.86 

Cochliobolus heterostrophus CBS 134.39 

Cochliobolus sativus DAOM 226212 

Comoclathris compressa CBS 156.53 

0.98/79 

Comoclathris compressa CBS 157.53 

1.0/79 

Pleospora typhicola CBS 132.69 

Pleospora incompta CBS 467.76 

0.96/83 Pleospora calvescens CBS 246.79 

Pleospora halimiones CBS 432.77 

1.0/79 1.0/97 Pleospora chenopodii CBS 206.80 

Pleospora betae CBS 109410 

1.0/65 

Chaetodiplodia sp. CBS 453.68 

Clathrospora elynae CBS 161.51 

Clathrospora elynae CBS 196.54 

Pleospora fallens CBS 161.78 

Alternariaster helianthi CBS 119672 

0.94/51 

Alternariaster helianthi CBS 327.69 

0.68/31 

Plenodomus lingam DAOM 229267 

0.55/- 

Neophaeosphaeria filamentosa CBS 102202 

0.87/54 

0.70/- 

Paraleptosphaeria dryadis CBS 643.86 

0.65/49 

Heterospora chenopodii CBS 115.96 

0.52/- 

Pleospora flavigena CBS 314.80 

0.94/67 

Coniothyrium palmarum CBS 400.71 

Pyrenochaeta nobilis CBS 407.76 

Ophiosphaerella herpotricha CBS 620.86 

Coniothyriaceae 

Cucurbitariaceae 

1.0/89 

Phaeosphaeria ammophilae CBS 114595 

Phaeosphaeria avenaria DAOM 226215 

0.86/85 

Phaeosphaeria eustoma CBS 573.86 

Chaetosphaeronema hispidulum CBS 216.75 

Loratospora aestuarii JK 5535B 

Setomelanomma holmii CBS 110217 

Saccothecium sepincola CBS 278.32 

Leptosphaerulina australis CBS 317.83 

0.56/49 0.85/41 Phoma complanata CBS 268.92 

1.0/81 

Phoma herbarum CBS 276.37 

0.99/90 Ascochyta pisi CBS126 54 

Didymellaceae 

1.0/91 

Peyronellaea glomerata CBS 528.66 

Peyronellaea zeae-maydis CBS 588.69 

Boeremia exigua CBS 431.74 

Julella avicenniae BCC 18422 

0.1 

Leptosphaeriaceae 

Phaeosphaeriaceae 

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). 

184


Fig. 3. Alternaria sect. Alternantherae: conidia and conidiophores. A–D. A. alternantherae. E–H. A. perpunctulata. Scale bars = 10 µm. 

ALTERNARIA SECTIONS 

Section Alternantherae D.P. Lawr., Gannibal, Peever 

& B.M. Pryor, Mycologia 105: 540. 2013. Fig. 3. 

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 speciesgroup 

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.: doi: 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. 

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. 


185


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. 

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.). 

186


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. 

= 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. 


187


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. 

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. 

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. 

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. 

188


Fig. 7. Alternaria sect. Cheiranthus: conidia and conidiophores. A–B. A. indefessa. B–C. A. cheiranthi. Scale bars = 10 µm. 

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. 

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, 


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

228. 1983. 

Section Crivellia (Shoemaker & Inderb.) Woudenb. & 


8. 

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 


189


Fig. 8. Alternaria sect. Crivellia: conidia and conidiophores. A–B. A. papavericola. C–D. A. penicillata. Scale bars = 10 µm. 

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. 


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. 

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. 


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. 

190


Fig. 9. Alternaria sect. Dianthicola: conidia and conidiophores. A–B. A. dianthicola. C–E. A. simsimi. F–H. A. elegans. Scale bars = 10 µm. 

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

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. 


Basionym: Embellisia tellustris E.G. Simmons [as “telluster”], 

Mycotaxon 17: 234. 1983. 


191


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. 


192


Section Embellisioides Woudenb. & Crous, sect. nov. 


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, 


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, 


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

50: 113. 2004. 

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


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

233. 1983. 

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


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. 


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

1983. 

Section Eureka Woudenb. & Crous, sect. nov. 


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 


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 

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 


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]. 


193


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. 

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. 


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. 

194


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. 

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 


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

Alternaria caespitosa (de Hoog & C. Rubio) Woudenb. & Crous, 


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. 


195


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. 

Alternaria californica E.G. Simmons & S.T. Koike, CBS 


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 


= 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. & 


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. 

196


Fig. 15. Alternaria sect. Japonicae: conidia and conidiophores. A–B. A. japonica. C–E. A. nepalensis. Scale bars = 10 µm. 

Fig. 16. Alternaria sect. Nimbya: conidia and conidiophores. A–B. A. caricis. C–D. A. scirpicola. Scale bars = 10 µm. 

Section Japonicae Woudenb. & Crous, sect. nov. 


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, 


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 elongateobclavate, 

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. 


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


197


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. 

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. 


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 speciesgroup 

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). 

198


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. 


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. 

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, 


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 

Type species: Alternaria porri (Ellis) Cif. 


199


Fig. 19. Alternaria sect. Porri: conidia and conidiophores. A–C. A. daucii. D–F. A. pseudorostrata. G–H. A. solani. Scale bars = 10 µm. 

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 Opavĕ, 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, Pflanzenschutzberichte 

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. 

200


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 “solaninigrii”], 

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. 

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. 


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. 


Eymology: Name refers to the concatenated conidia. 

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


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. 


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 


201


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. 

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. 


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. & 


23. 

202


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. 

Fig. 22. Alternaria sect. Sonchi: conidia and conidiophores. A–B. A. cinerariae. C–D. A. sonchi. Scale bars = 10 µm. 


203


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. 


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. 


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. 


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. & 


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, 


Basionym: Ulocladium multiforme E.G. Simmons, Canad. J. Bot. 

76: 1537. 1999 [1998]. 

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


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. 

204


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. 


205


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. 

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, 


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. 


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. 


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, 


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. 

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; 

206


Fig. 26. Alternaria sect. Undifilum: conidia and conidiophores. A–D. A. bornmuelleri. Scale bars = 10 µm. 

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. 


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, 


Basionym: Undifilum cinereum Baucom & Creamer, Botany 90: 

872. 2012 

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


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

2012 

Alternaria oxytropis (Q. Wang, Nagao & Kakish.) Woudenb. & 


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. 


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, 


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 



207


Fig. 27. Paradendryphiella gen. nov.: conidia and conidiophores. A–B, D–E, G–I. P. salina. C, F. P. arenariae. Scale bars = 10 µm. 

= 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. 


Basionym: Dendryphiella arenariae Nicot, [as “arenaria”] Rev. 

Mycol. (Paris) 23: 93. 1958. 

≡ Scolecobasidium arenarium (Nicot) M.B. Ellis, More dematiaceous 


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 fivegene 

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 speciesgroups 

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 

208


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 smallspored 

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 


209


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 polyand 

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. 

Genera unrelated to Alternaria 

The placement of the sexual genus Pleospora (1863) with 

Stemphylium (1833) asexual morphs as basal sister clade to the 

Alternaria complex is well-documented in multiple molecular studies 

(Chou & Wu 2002, Pryor & Bigelow 2003, Hong et al. 2005, Pryor 

et al. 2009, Lawrence et al. 2012). Therefore, we only included the 

type species of both genera in our phylogenies and used them as 

outgroup in the Alternaria phylogeny. Pleospora herbarum with its 

Stemphylium herbarum (CBS 191.86) asexual morph is the type 

species of the genus Pleospora. Stemphylium botryosum with its 

Pleospora tarda (CBS 714.68) sexual morph is the type species of 

the genus Stemphylium. 

Embellisia annulata proved to be identical to the marine 

species Dendryphiella salina, and forms a well-supported clade 

in the Pleosporaceae together with D. arenariae. Several DNAbased 

studies (dela Cruz 2006, Jones et al. 2008, Zhang et al. 

2009) concluded that the marine Dendryphiella species, D. 

arenariae and D. salina, belonged to the Pleosporaceae as sister 

clade to the Pleospora/Stemphylium complex. Furthermore, 

they showed the type species of Dendryphiella, D. vinosa, to 

be only distantly related, based on sequences of the ITS, SSU, 

LSU (Jones et al. 2008) and ITS, TEF1, RPB2 (dela Cruz 2006) 

gene regions. The transfer of the marine Dendryphiella species to 

Scolecobasidium (Ellis 1976), was also disputed. Scolecobasidium 

does not belong to the Pleosporales based on ITS, TEF1, and 

RPB2 sequences (dela Cruz 2006) and the morphology of the two 

Dendryphiella species does not fit the generic circumscription of 

Scolecobasidium (dela Cruz 2006, Jones et al. 2008). Ellis (1976) 

described denticles on the conidiogenous cells when the conidia 

become detached. However other observers describe a marginal 

basal frill on the conidia after detachment, leaving a scar on the 

conidiophore. We propose to place the two species in the new 

genus Paradendryphiella as C. arenariae and C. salina. The need 

for a new genus to accommodate the two species was already 

suggested by Jones et al. (2008). 

A recent study on Diademaceae, a family which is characterised 

by a flat circular operculum and bitunicate asci (Shoemaker 

& Babcock 1992), excluded the sexual genera Comoclathris 

and Clathrospora, and (provisionally) placed them in the 

Pleosporaceae with alternaria-like asexual morphs (Zhang et al. 

2011). Molecular data of two strains (Dong et al. 1998, Schoch et 

al. 2009) placed them within the Pleosporaceae. A confusing factor 

is that Dong et al. (1998) use the name Comoclathris baccata in 

their paper for strain CBS 175.52, but submitted their sequences 

under the name Clathrospora diplospora to GenBank. Shoemaker 

& Babcock (1992) synonymised Clathrospora diplospora with 

Comoclathris baccata, which renders Comoclathris as the correct 

generic name. The confusion around these genera is illustrated 

by the fact that the CBS collection currently harbours six strains 

named as Clathrospora species of which four were renamed by 

Shoemaker & Babcock in 1992 based on morphological studies, 

and three of these four strains were even transferred to the genus 

Comoclathris. The type species of Clathrospora, C. elynae is 

represented by two strains of which one, CBS 196.54, was also 

studied morphologically by Shoemaker and Babcock (1992). They 

form a well-supported clade, located basal to the Pleosporaceae 

210


(Fig. 2), outside the Alternaria complex. The type species of 

Comoclathris, Comoclathris lanata, was not available to us, but 

the two Comoclathris compressa strains cluster together in a 

well-supported clade within the Pleosporaceae, also outside the 

Alternaria complex, which we believe to be the correct phylogenetic 

placement of the genus. Two other strains, named Comoclathris 

magna (CBS 174.52) and Clathrospora heterospora (CBS 

175.52) by Shoemaker and Babcock (1992), cluster amidst sect. 

Alternata. Culture studies performed by Simmons (1952) showed 

the presence of alternaria-like conidia in these cultures and no 

(mature) ascospore formation. Presumably the species observed 

by Shoemaker and Babcock (1992) on plant material were lost 

during cultivation and became replaced by A. alternata speciesgroup 

isolates. Both strains will be treated as “Alternaria sp.” 

The genus Alternariaster was first described by Simmons 

(2007) with Alternariaster helianthi, formerly Alternaria helianthi or 

Helminthosporium helianthi, as type and only species. It is distinct 

from Alternaria by the lack of a pigmented conspicuous internal, 

circumhilar ring in its conidia and conidiophores. Our study showed 

that this genus is clearly not part of the Alternaria complex and 

belongs to the Leptosphaeriaceae (Fig. 2) (Alves et al. 2013). 

In the recently published book “The genera of Hyphomycetes” 

(Seifert et al. 2011) three more genera are linked to Alternaria, 

namely Pantospora, Briansuttonia and Rhexoprolifer. A recent 

study on Pantospora included ITS and LSU sequence data of 

the type species Pantospora guazumae, which placed the genus 

in Mycosphaerellaceae (Minnis et al. 2011). This refutes the 

link with Alternaria. The genus Rhexoprolifer was described in 

1996 by Matsushima with R. variabilis as type and only species, 

isolated from South Africa. Rhexoprolifer variabilis has rhexolytic 

conidial liberation and proliferating conidiophores with both 

phragmosporous and dictyosporous conidia. Briansuttonia was 

described in 2004 to accommodate Corynespora alternarioides 

(Castañeda Ruiz et al. 2004). The distoseptate muriform conidia 

of Briansuttonia do resemble Alternaria and Stemphylium, but 

the conidiogenous loci and euseptate conidia of Alternaria and 

holoblastic conidial ontogeny and euseptate muriform conidia of 

Stemphylium were enough for the authors to regard their taxon as 

a different genus. Both asexual genera presently lack molecular 

data, and we were unable to obtain any living specimens of these 

taxa. It would be valuable to include both genera in a future study 

to resolve the connection among genera with muriform conidia 

and Alternaria. 

The description of Alternaria s. str. in the present study is 

supported by i) a well-supported phylogenetic node in multiple 

analyses, ii) high similarity of clades within Alternaria based on 

SSU, LSU and ITS data, and iii) variation in the order of the clades 

between the different gene phylogenies, which is in congruence 

with low support values at these deeper nodes. We follow the 

precedence introduced by Lawrence et al. (2013) to assign the 

taxonomic status of sections of Alternaria for the different clades 

found, thus allowing us to retain the former generic names but 

associated with a different taxonomic status. For end-users, this 

also results in a more stable and understandable taxonomy and 

nomenclature. 

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. 

Acknowledgements 

Mrs M. Vermaas is thanked for preparing the photoplates, Mrs J. Bloem for assisting 

with the molecular work, and Prof. dr B.M. Pryor for sending us the Undifilum 

isolate. This research was supported by the Dutch Ministry of Education, Culture 

and Science through an endowment of the FES programme “Making the tree of 

life work”. 

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