WITHDRAWN: Fungi associated with grapevine trunk diseases (GTDs) with emphasize on pestalotioid species in Kurdistan Province, Iran

Materials And Methods

Sampling and fungal isolation

During a survey between 2012 and 2014 on grapevine trunk diseases in Kurdistan Province, twig and trunk samples of grapevines showing trunk diseases symptoms (cv. Askari, Bidaneh Sefid, Farkhi, Rasha and Sahabi) were collected from vineyards all over 10 years old in 25 different villages. Grapevine cultivars showed different symptoms consisting decline, reduced growth, interveinal yellow-brown or red-brown necrotic spots on leaves known as tiger-stripes pattern, spotting berries (black measles), sectorial and central brown necrosis of the trunks. Cross sections of samples were made and sliced to 0.5-1 cm pieces of infected wood. After surface sterilization, (3–4 min in 70% ethanol) four pieces were placed on 9 cm PDA plates supplemented with 100 mg chloramphenicol, streptomycin or tetracycline. Plates were incubated at 25 ± 2 ºC in the dark. Colonies grown from wood pieces were transferred to PDA plates and incubated at 25 ± 2 ºC in the dark. After 1–2 wk conidiomata were formed on PDA plates. To purification of the isolates using single-spore method conidia were transferred to tap water agar (2% WA). After incubation at 25°C for 12 h single germinated conidia were transferred to PDA plates. Representative isolates were deposited in the culture collection (IRAN) of the Iranian Research Institute of Plant Protection (Tehran, Iran) and the culture collection (CBS) of the Westerdijk Fungal Biodiversity Institute (Utrecht, the Netherlands).

Morphology

Colonies were grown on PDA, MEA and OA at 25 ± 2°C for 1–2 wk. Structures were mounted in 100 % lactic acid or water and digital images were recorded with an Olympus DP72 camera on a Olympus BX51 microscope. Measurements were made with the Cell Sense Entry measurement module. For each isolate the mean, standard deviation, minimum and maximum values were calculated from measurements of at least 30 fungal structures. Conidial length was measured from the base of the basal cell to the base of the apical appendage, and conidial width was measured at the widest point of the conidium (Bonthond et al. 2018). Dimensions are presented as a range with extremes and mean ± standard deviation in parentheses. Depending fungal taxonomic groups the colony morphology and growth rate were determined on different culture media and temperature in the dark. For pestalotioid fungi colony morphology and growth rate were determined on MEA and PDA at 21°C in the dark. After 2 wk mycelial growth was measured and cultural characteristics were recorded based on the colour charts of Rayner (1970).

DNA extraction, PCR and sequencing

The PCR reaction mixtures 25 µL contained 1×PCR buffer (PCR buffer with (NH4)2SO4), 3 mM MgCl2, 200 µM of each nucleotide, 5 pmol of each primer, 1 U of Taq polymerase and 1 µL of template DNA (50–100 ng/µL). Genomic DNA was extracted from 4–7 d old cultures grown in potato dextrose broth (PDB) using modified method of Raeder & Broda (1985) as described by Abdollahzadeh et al. (2009). The D1/D2 variable domains of the 28S nrDNA (LSU) and the ITS1, 5.8 and ITS2 region of ribosomal DNA and part of β-tubulin (TUB2) and the translation elongation factor 1-alpha (TEF-1α) were amplified and sequenced using the following primer pairs LR0R/LR5 for LSU (Vilgalys and Hester 1990), ITS5 or ITS1/ITS4 for ITS (White et al. 1990), T1/Bt2b for TUB2 (Glass and Donaldson 1995, O'Donnell and Cigelnik 1997), EF-1/EF-2 for TEF-1α (O'Donnell et al. 1998). The PCR reaction mixtures 12.5 µL contained 1×PCR buffer (PCR buffer with (NH4)2SO4), 3 mM MgCl2, 200 µM of each nucleotide, 5 pmol of each primer, 1 U of Taq polymerase and 1 µL of template DNA (50–100 ng/µL). The PCR amplification conditions were 95°C for 5 min, followed by 35 cycles of 94°C for 30 s, 52°C for 45 s (LSU and ITS) or 55°C for 45 s (TEF-1α and TUB2), and 72°C for 1 min, and a final extension of 72°C for 7 min. The PCR products were sequenced with both forward and reverse primers using an Applied Biosystems 3730xl DNA Analyzer (Thermo Fisher Scientific). Forward and reverse reads were paired and consensus sequences were obtained using the software BioEdit v. 7.0.0 (Hall 2004). All new sequences were submitted to GenBank (Table 1).

Table 1

Isolates used in phylogenetic analyses
Species	Isolate No.¹	Host	Location	GenBank accession number²
Species		Host	Location	LSU	ITS		TUB2		EF1-α
Allelochaeta fusispora	CBS 810.73^IT	Eucalyptus polyanthemos	Australia	MH554279		MH554067		MH554743		MH554503
All. falcata	CPC 13580	E. alligatrix	Australia	MH554284		MH554073		MH704626		MH704601
Bartalinia robillardoides	CBS 122615	Cupressus lusitanica	South Africa	MH554207		MH553989		MH554657		MH554415
Broomella vitalbae	HPC 1154	-	-	MH554367		MH554173		MH554846		MH554608
Ciliochorella phanericola	MFLUCC 12–0310	Dead leaves	Thailand	KF827445		KF827444		KF827478		KF827477
Diploceras hypericinum	CBS 109058	Hypericum sp.	New Zealand	MH554178		MH553955		MH554614		MH554373
D. hypericinum	CBS 492.97	H. perforatum	Netherlands	MH554267		MH554054		MH554730		MH554489
Disaeta arbuti	CBS 143903	Acacia pycnantha	Australia	MH554346		MH554148		MH554821		MH554583
Discosia sp. 1	CBS 241.66	A. karroo	South Africa	MH554244		MH554022		MH554698		MH554456
Discosia sp. 2	CBS 684.70	Aesculus hippocastanum	Netherlands	MH554277		MH554064		MH554740		MH554500
Distononappendiculata banksiae	CBS 143906	Banksia formosa	Australia	MH554354		MH554158		MH554831		MH554593
Diversimediispora humicola	CBS 302.86^T	Soil	USA	MH554247		MH554028		MH554705		MH554463
Heterotruncatella restionacearum	CBS 118150	Restio filiformis	South Africa	MH554203		DQ278914		MH554649		MH554407
Hyalotiella transvalensis	CBS 303.65^T	Leaf litter and top soil of A. karroo community	South Africa	MH554248		MH554029		MH554706		MH554464
Hymenopleella hippophaeicola	CBS 113687	Hippophaë rhamnoides	Sweden	MH554188		MH553969		MH554628		MH554387
Immersidiscosia eucalypti	CBS 104197	Ardisia japonica	Japan	AB593724		AB594792		NA		NA
Lepteutypa fuckelii	CBS 140409^NT	Tilia cordata	Belgium	KT949902		NR_154123		MH554677		MH554435
Monochaetia ilexae	CBS 101009	Air	Japan	MH554176		MH553953		MH554612		MH554371
Morinia acaciae	CBS 100230	Prunus salicina cv. Omega	New Zealand	MH554174		MH553950		MH554609		MH554368
Neopestalotiopsis zimbabwana	CBS 111495^T	Leucospermum cunciforme	Zimbabwe	JX556249		JX556231		KM199456		KM199545
Nonappendiculata quercina	CBS 270.82	Quercus pubescens	Italy	MH554246		MH554025		MH554701		MH554459
Parabartalinia lateralis	CBS 399.71^T	A. karroo	South Africa	MH554256		MH554043		MH554719		MH554478
Pestalotiopsis humuicola	CBS 115450	Ilex cinerea	Hong Kong	KM116208		KM199319		KM199418		KM199487
Pseudopestalotiopsis cocos	CBS 272.29^T	Cocos nucifera	Indonesia	KM116276		KM199378		KM199467		KM199553
Pseudosarcostroma osyridicola	CBS 103.76^T	Osyris alba	France	MH554177		MH553954		MH554613		MH554372
Robillarda terrae	CBS 587.71^T	Soil	India	KJ710459		KJ710484		MH554734		MH554493
Sarcostroma leucospermi	CBS 111290^T	Leucospermum cv. 'High Gold	South Africa	MH554292		MH554081		MH554755		MH554516
Sarcostroma proteae	CBS 113610^T	Protea magnifica	Australia	MH554187		MH553968		MH554627		MH554386
Seimatosporium botan	NBRC 104200^T	Paeonia suffruticosa	Japan	AB593731		AB594799		LC047770		NA
Seimatosporium ficeae	MFLUCC 15-0519^T	Ficus sp.	China	KR920686		KR092800		NA		NA
Seimatosporium germanicum	CBS 437.87^T	-	Germany	MH554259		MH554047		MH554723		MH554482
Seimatosporium luteosporum	CBS 142599^T	Vitis vinifera	USA	KY706309		KY706284		KY706259		KY706334
Seimatosporium marivanicum	IRAN 2310C^T = CBS 143781 IRAN 2300C = CBS 143780	V. vinifera V. vinifera	Iran, Mariwan Iran, Mariwan	MW361960 MW361959		MW361952 MW361951		MW375352 MW375351		MW375358 MW375357
Seimatosporium physocarpi	CBS 139968^T	Physocarpus opulifolius	Russia	KT198723		KT198722		MH554676		MH554434
	CBS 789.68	Physocarpus amurensis	Netherlands	MH554278		MH554066		MH554742		MH554502
Seimatosporium pistaciae	CPC 24457	Pistacia vera	Iran	MH554331		MH554126		MH554799		MH554561
Seimatosporium rhombisporum	MFLUCC 15-0543^T	Vaccinium myrtillus	Italy	KR092780		KR092792		NA		NA
Seimatosporium rosae	CBS 139823^ET	Rosa kalmiussica	Russia	KT198727		LT853105		LT853253		LT853203
Seimatosporium soli	CBS 941.69^T	Forest soil under Fagus sylvatica	Denmark	MH554282		MH554071		NA		MH554507
Seimatosporium vitifusiforme	CBS 142600^T	V. vinifera	USA	KY706321		KY706296		KY706271		KY706346
Seimatosporium vitis	MFLUCC 14–0051	V. vinifera	Italy	KR920362		KR920363		NA		NA
Seimatosporium vitis-viniferae	CBS 123004^T	V. vinifera	Spain	MH554211		MH553992		MH554660		MH554418
Seiridium pseudocardinale	CBS 122613	Cupressus sp.	Portugal	MH554206		LT853096		LT853243		LT853193
Sporocadus biseptatus	CBS 110324^T	-	-	MH554179		MH553956		MH554615		MH554374
Sporocadus cornicola	CBS 143889	Cornus sanguinea	Germany	MH554326		MH554121		MH554794		MH554555
Sporocadus corni	MFLUCC 14-0467^T	Cornus sp.	Italy	KR559739		KT162918		NA		NA
Sporocadus cotini	CBS 139966^T	Cotinus coggygria	Russia	MH554222		MH554003		MH554675		MH554433
Sporocadus incanus	CBS 123003^T	Prunus dulcis	Spain	MH554210		MH553991		MH554659		MH554417
Sporocadus italicus	MFLUCC 14-1196^T	Crategus sp.	Italy	MF614829		MF614831		NA		NA
Sporocadus kurdistanicus	IRAN 2356C^T = CBS 143778 IRAN 2355C IRAN 2354C IRAN 2313C	V. vinifera V. vinifera V. vinifera V. vinifera	Iran, Sanandaj Iran, Mariwan Iran, Mariwan Iran, Dehgolan	MW361958 NA MW361957 MW361956		MW361950 MW361949 MW361948 MW361947		MW375350 NA MW375349 MW375348		MW375356 NA MW375355 MW375354
Sporocadus lichenicola	NBRC 32625^ET	Fagus sylvatica	Germany	MH554252		MH554035		MH554711		MH554470
Sporocadus mali	CBS 446.70^T	Malus sylvestris	Netherlands	MH554261		MH554049		MH554725		MH554484
Sporocadus microcyclus	CBS 424.95^T	Sorbus aria	Germany	MH554258		MH554045		MH554721		MH554480
Sporocadus multiseptatus	CBS 143899^T	Viburnum sp.	Serbia	MH554343		MH554141		MH554814		MH554576
Sporocadus rosigena	MFLU 16-0239^T	Rosa canina	Italy	MG829069		MG828958		NA		NA
Sporocadus pseudocorni	MFLUCC 13-0529^T	Cornus sp.	Italy	KU359033		NA		NA		NA
Sporocadus rosarum	CBS 113832 MFLUCC 14-0466^T* MFLUCC 15-0563^T* MFLUCC 14-0468^T*	Rosa canina Rosa canina Rosa canina Rosa villosa	Sweden Italy Italy Italy	MH554189 KT281912 MG829071 KU359035		MH553970 KT284775 MG828960 NA		MH554629 NA NA NA		MH554388 NA NA NA
Sporocadus rotundatus	CBS 616.83^T	Arceuthobium pussilum	Canada	MH554273		MH554060		MH554737		MH554496
Sporocadus sorbi	CBS 160.25	-	-	MH554229		MH554008		MH554684		MH554442
Sporocadus sp. 1	CBS 506.71	Euphorbia sp.	Italy	MH554268		MH554055		MH554731		MH554490
Sporocadus sp. 2	CBS 466.96	Inner tissue of zoocecidium, caused by Lasioptera rubi, on Rubus sp.	Netherlands	MH554265		MH554052		MH554728		MH554487
Sporocadus trimorphus	CBS 114203^T	Rosa canina	Sweden	MH554196		MH553977		MH554636		MH554395
Strickeria kochii	CBS 140411^ET	Robinia pseudoacacia	Austria	KT949918		NR_154423		MH554679		MH554437
Synnemapestaloides juniperi	CBS 477.77^T	Juniperus phoenicea	France	MH554266		MH554053		MH554729		MH554488
Synnemapestaloides rhododendri	MAFF 239201^T	Rhododendron brachycarpum	Japan	LC047744		LC047753		LC047761		NA
Truncatella angustata	CBS 393.80 CJA35 CJA82	Gevuina avellana V. vinifera V. vinifera	Chile Iran, Sanandaj Iran, Sanandaj	MH554254 NA NA		MH554041 MW361953 MW361954		MH554717 NA NA		MH554476 NA NA
Undetermined species	CBS 387.77 CBS 113991	Skin of man Salix caprea	Finland Sweden	KM116277 MH554190		MH554040 MH553971		MH554716 MH554630		MH554475 MH554389
Xenoseimatosporium kurdistanicum	IRAN 2353C^T IRAN 2305C	V. vinifera V. vinifera	Iran, Mariwan Iran, Kamyaran	MW361955 NA		MW361946 MW361945		MW375347 NA		MW375353 NA
Xenoseimatosporium quercinum	CBS 129117 MFLUCC 14-1198^T	Rhododendron sp. Quercus robur	Lativa Germany	MH554216 NG_059681		MH553997 NR_155804		MH554666 NA		MH554424 NA
¹ CBS Culture collection of the Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; CJA Personal cultures of Jafar Abdollahzadeh; CPC Working collection of P.W. Crous housed at the Westerdijk Institute; HPC Herbarium of Pedro Crous, housed at the Westerdijk Institute; IRAN Iranian Fungal Culture Collection, Iranian Research Institute of Plant Protection, Iran; MAFF Ministry of Agriculture, Forestry and Fisheries, Tsukuba, Ibaraki, Japan; MFLU(CC) Mae Fah Luang University Culture Collection; NBRC Biological Resource Center. ET: ex-epitype; IT: ex-isotype; NT: ex-neotype; T: ex-type. * MFLUCC 15–0563: Type of Seimatosporium rosigenum; MFLUCC 14–0466: Type of Seimatosporium pseudorosarum; MFLUCC 14–0468: Type of Seimatosporium pseudorosae.
² LSU large subunit ribosomal DNA; ITS internal transcribed spacer; EF1-α translation elongation factor 1-alpha; TUB2 β-tubulin 2; N/A not available; Newly generated sequences are indicated in bold.

Phylogenetic analyses

Consensus sequences together with retrieved sequences from GenBank (http://www.ncbi.nlm.nih.gov) were aligned using MAFFT v. 7 (http://mafft.cbrc.jp/alignment/server/index.html), and manually edited in MEGA v. 7.0.21. The aligned dataset was subjected to Bayesian analysis (BA) and Maximum Likelihood (ML) on the CIPRES Science Gateway portal (https://www.phylo.org/; Miller et al. 2012) using MrBayes v. 3.2.6 (Huelsenbeck and Ronquist 2001, Ronquist and Huelsenbeck 2003) and RAxML-HPC BlackBox v. 8.2.10 (Stamatakis 2014), respectively. The optimal nucleotide substitution models were determined for each locus using MrModelTest v. 2.3 (Nylander 2004). Bayesian analyses were implemented under the optimal nucleotide substitution models with four simultaneous Markov Chain Monte Carlo chains, 10 M generations and a sampling frequency of 1 000 generations, ending the run automatically when standard deviation of split frequencies dropped below 0.01. Burn-in was set to remove 25 % of the first sampled trees, after which the 50 % majority rule consensus trees and posterior probability (PP) values were calculated. The ML analyses were done using a GTR + GAMMA substitution model and four rate classes with 1 000 bootstrap iterations. The obtained phylogenetic trees were plotted using FigTree v. 1.4.3 (http://tree.bio.ed.ac.uk/software/figtree). Alignments and trees were deposited in TreeBASE (www.treebase.org; S27404) and taxonomic novelties in MycoBank (www.MycoBank.org; Crous et al. 2004).

Results

Fungal isolates and species identification

In this survey some 223 fungal isolates were obtained from grapevines showing trunk diseases symptoms (Fig. 1), which 30 isolates were morphologically pestalotioid belong to Sporocadaceae. Based on morphology and DNA sequence data 24 fungal species belong to 20 genera were identified (Fig. 2; Table 2). All fungal species characterized in this survey are new records for the fungal flora of Kurdistan Province. It is the first time Clonostachys rosea and Neoscytalidium novaehollandiae are reported on grapevine in Iran. Acremonium sclerotigenum, Alternaria chlamydosporigena, Ascochyta herbicola and Paecilomyces formosus are new records on grapevine around the world.

Table 2

Fungal species associated with grapevine trunk diseases identified in this study
Species	Isolates no.	Frequency (%)	Grapevine cv.	Location
Acremonium sclerotigenum	20	8.55	Rasha, Sahabi, Farkhi	Dehgolan, Kamyaran, Marivan, Sanandaj
Alternaria chlamydosporigena	15	6.4	Rasha	Bijar, Dehgolan, Kamyaran, Qorveh
Alternaria malorum	27	11.55	Rasha, Sahabi, Farkhi, Bidaneh Sefid	Bijar, Dehgolan, Divandareh, Kamyaran, Marivan, Sanandaj, Saqez
Ascochyta herbicola	4	1.7	Rasha	Marivan
Botryosphaeria dothidea	27	11.55	Rasha, Sahabi, Bidaneh Sefid	Bijar, Marivan, Sanandaj, Saqez
Cadophora malorum	7	3	Rasha, Bidaneh Sefid	Baneh, Kamyaran, Marivan, Saqez
Clonostachys rosea	14	6	Rasha, Bidaneh Sefid	Kamyaran, Marivan, Sanandaj
Didymella glomerata	10	4.3	Rasha	Dehgolan, Kamyaran, Marivan
D. pinodella	5	2.1	Rasha, Bidaneh Sefid	Kamyaran
Diplodia seriata	4	1.7	Rasha	Marivan
Juxtiphoma eupyrena	4	1.7	Rasha	Marivan, Sanandaj
Kalmusia variispora	2	0.85	Rasha	Kamyaran
Microsphaeropsis olivacea	7	3	Rasha, Bidaneh Sefid	Kamyaran, Sanandaj
Neoscytalidium hyalinum	14	6	Rasha, Sahabi	Marivan
N. novaehollandiae	6	2.5	Rasha	Baneh, Qorveh
Paecilomyces formosus	6	2.5	Rasha	Marivan
Phaeoacremonium aleophilum	23	9.9	Rasha, Sahabi, Bidaneh Sefid Askari	Dehgolan, Kamyaran, Marivan, Sanandaj
Ph. parasiticum	2	0.85	Rasha	Marivan
Ph. rubrigenum	5	2.1	Rasha	Marivan
Phaeomoniella chlamydospora	2	0.85	Rasha	Kamyaran, Marivan
Seimatosporium marivanicum	10	4.3	Rasha, Sahabi	Marivan
Sporocadus kurdistanicus	6	2.6	Rasha	Dehgolan, Marivan, Sanandaj
Truncatella angustata	4	1.7	Rasha, Sahabi, Bidaneh Sefid	Marivan, Sanandaj
Xenoseimatosporium kurdistanicum	10	4.3	Rasha, Sahabi	Kamyaran, Marivan

Given that three new pestalotioid species were identified for science here we focused on phylogeny and description of the pestalotioid fungi isolated in this study. Based on morphology, cultural characteristics, grapevine cultivar and sampling geographical location 10 out of 30 isolates were selected for sequencing and phylogenetic studies (Table 1).

Phylogeny

Two datasets were subjected to phylogenetic analyses. The first dataset consisted of concatenated LSU, ITS, TEF-1α and TUB2, containing 55 taxa representing 30 genera and one undetermined clade recognized by Liu et al. 2019 and Lepteutypa fuckelii CBS 140409 as outgroup was made and analyzed to determine phylogenetic position of our isolates at the genus level. The concatenated dataset after alignment contained a total of 3 151 characters (LSU: 823, ITS: 578, TEF-1α: 738, TUB2: 1 000), including alignment gaps. MrModelTest revealed that the general time-reversible model of evolution (Rodriguez et al. 1990), including estimation of invariable sites and assuming a discrete gamma distribution (GTR + I + G) with six rate categories (lsetnst = 6, rates = invgamma) and dirichlet (1,1,1,1) base frequencies is the best nucleotide substitution model for all loci (LSU, ITS, TEF-1α and TUB2). The Bayesian analyses of the concatenated alignments of four loci generated 3 672 trees from which 918 trees were discarded as burn-in. The consensus tree and posterior probability values (PP) were calculated from the remaining 2 754 trees. The average standard deviation of split frequencies was 0.009942 at the end of the run. The RAxML search of the dataset with 1 594 distinct alignment patterns produced a best-scoring ML tree (lnL = -37448.896642). The same phylogenetic tree obtained from both RAxML and Bayesian analyses. The posterior probability values (PP) equal to or higher than 0.5 were mapped on the ML tree (Fig. 3). Our isolates were grouped in four genera Seimatosporium, Sporocadus, Truncatella and Xenoseimatosporium. Isolate MFLUCC 14-1196 the type of Seimatosporium italicum placed in Sporocadus clade. Seimatosporium rhombisporum MFLUCC 15–0543 (ex-type) and Seimatosporium ficeae MFLUCC 15–0519 (ex-type) constituted a well-supported clade distinct from all other clades. This clade is probably representative of a new genus but as there is only LSU and ITS sequence data available for these two species it is better to generate RPB2, TEF-1α and TUB2 sequences for further phylogenetic analyses to make the final decision at the generic level.

The second dataset consisted of concatenated LSU, ITS, TEF-1α and TUB2, containing 46 taxa belonging to four genera Seimatosporium, Sporocadus, Truncatella and Xenoseimatosporium and Distononappendiculata banksiae CBS 143906 as outgroup was prepared and analyzed to identify our isolates at the species level. The aligned dataset contained 2 686 characters (LSU: 818, ITS: 515, TEF-1α: 559, TUB2: 782), including alignment gaps. As in the first dataset, MrModelTest indicated a GTR + I + G as the best fit model for all loci (LSU, TEF-1α and TUB2). The Bayesian analyses generated 1 162 trees from which 290 trees were discarded as burn-in. The consensus tree and posterior probability values (PP) were calculated from the remaining 872 trees. The average standard deviation of split frequencies was 0.009786 at the end of the run. The RAxML search of the dataset estimated 1 108 distinct alignment patterns and made a tree with lnL = -17900.721816. Both analyses resulted to the same phylogenetic tree and posterior probability values equal (PP) to or higher than 0.5 were mapped on the ML tree (Fig. 4). In these study based of multigene phylogenetic analyses our isolates placed in four different genera in well supported clades which three of them are recognized as new species for science that are introduced here. The fourth species, Truncatella angustata is a well-known and common species associated with woody plants. This species has reported from several countries in association with more than 30 woody plant species including Vitis vinifera (Farr and Rossman 2020). Based on the results, Seimatosporium cornii, Seimatosporium italicum and Seimatosporium pseudocornii are transferred to Sporocadus and three new combinations are proposed.

Taxonomy

Based the multigene phylogenetic analyses four pestalotioid species belong to Sporocadaceae associated with grapevine trees showing grapevine trunk diseases symptoms were recognized including Truncatella angustata and three new species along with three new combinations introduced here as follows.

Seimatosporium marivanicum Abdollahz., Nahvi M. & Khaledi E., sp. nov.

MycoBank MB 838232

Etymology: Name refers to Marivan in Kurdistan Province, Iran where this species was first found.

Diagnosis: In the multigene phylogenetic tree Sei. marivanicum constituted a highly supported distinct calde grouped with a clade containing Seimatosporium luteosporum and Seimatosporium vitifusiform (Fig. 4). Sei. marivanicum has 4, 5, 20 and 17 bp differences with Sei. luteosporum in LSU, ITS, TEF-1α and TUB2 sequences, respectively. LSU and ITS sequences of Sei. marivanicum are identical to the Sei. vitifusiforme, but there are 6 and 2 bp differences between these two species in TEF-1α and TUB2, respectively. Sei. marivanicum can be easily differentiated from Sei. luteosporum by conidial dimensions (24 × 3.5 μm vs. 19.9 × 5.3 μm) (Table 3). Conidial size of Sei. marivanicum is almost indistinguishable from Sei. vitifusiform, but conidia in Sei. vitifusiforme are 3-euseptate, while in Sei. marivanicum we have seen conidia with up to 6 eusepta (Table 3). Moreover, both appendages (apical/basal) of Sei. marivanicum are more longer than Sei. vitifusiforme (15/16 μm vs. 10/9.5 μm ) (Table 3). Sei. luteosporum has been reported on Prunus persica and Vitis vinifera from California and Sei. vitifusiforme has only reported on Vitis vinifera from Califorina (Farr and Rossman, 2020). To differentiate Seimatosporium species reported on grapevine we have presented conidial characteristics in Table 3.

Table 3 Conidial characteristics of Seimatosporium species reported on grapevine

Species	Conidial dimensions (µm)	Septum no.	Type of appendages	Apical appendage length (µm)	Basal appendage length (µm)	Conidia L/W ratio	Reference
S. botan	16–20 × 5–7 (av. = 18 × 6) 16–20 × 4–5 (av. = 18 × 4)	3 3	basal apical and basal	– 4–8 (av. = 5.8)	4–8 (av. = 6) 4–8 (av. = 5.4)	2.6–3.8 (av. = 3) 4–5 (av. = 4.6)	Hatakeyama and Harada 2004
S. hysterioides	12–14 × 5–6	3	often lacking, occasionally basal or with both types	5–12	5–12	–	Shoemaker 1964
S. lonicerae	9–16 × 3.5–5 (av. = 13 × 4.4)	3, (2)^*	both types or basal only	3–7 (av. = 5.5)	2–12 (av. = 7)	3	Nag Raj 1993
S. luteosporum	16.7–25.4 × 4.7–5.6 (av. = 19.9 × 5.3)	3	apical and basal	10.1–24.2 (av. = 17.9)	9.8–23.6 (av. = 16.7)	–	Lawrence et al. 2018
S. macrospermum	28–39 × 9–12.5	5	lacking appendages	–	–	–	Sutton 1975
S. marivanicum	16–31 × 3–7 (av. = 24 × 3.5)	3(–6)	apical and basal	7–20 (av. = 15)	5–20 (av. = 16)	5(–6)	This study
S. parasiticum	22–35 × 5–6 (–7) (av. = 27.5 × 5.5)	5, (3/4)^*	apical and basal	2–5 (av. = 3.5)	2–8 (av. = 4.5)	5	Nag Raj 1993
S. vitifusiforme	18.6–30.3 × 3.7–5.1 (av. = 24.9 × 4.2)	3	apical and basal	7–12.6 (av. = 10)	3.9–16.6 (av. = 9.5)	–	Lawrence et al. 2018
S. vitis-viniferae	13.5–26 × 4.5–6 (av. = 16.5 × 5.2)	3(–6)	basal or with both types	4–11 (av. = 7)	4–10 (av. = 7.9)	3.2	Liu et al. 2019
S. vitis	34–40 × 14–17 (av. = 37 × 15)	3	basal	–	4–8 (av. = 5)		Senanayake et al. 2015

Type: Iran: Kurdistan Province: Marivan, Nzhmar, Vitis vinifera (cv. Rasha), 11 Sep. 2012, J. Nahvi Moghadam (IRAN 17872F–holotype; IRAN 2310C = CBS 143781–ex-type culture).

Description: Sexual morph: unknown. Asexual morph: Conidiomata acervular, stromatic, immersed, semi-immersed to erumpent, dark brown to black. Conidiophores branched, hyaline, smooth. Conidiogenous cells discrete, mostly cylindrical or oblong, 4–20 × 1–2 μm (av. = 10 ± 1.5 × 1.5 ± 0.2 μm), hyaline, smooth. Conidia allantoid, subcylindrical, curved to straight, 3(–6)-septate, wall smooth, some constricted at the septa, 16–31 × 3–7 μm (av. = 24 ± 1.5 × 3.5 ± 0.4 μm), bearing appendages; basal cell obconic with truncate base or trapezoid, thin-walled, hyaline to pale brown, 2–4 μm (av. = 3 ± 0.2 μm) long; median cells 2(–5), cylindrical or doliiform to ovoid, thick-walled, pale to mid-brown, ± equal, each 5–10 μm (av. = 8 ± 0.7 μm) long; apical cell conic with an acute or rounded apex, thin-walled, hyaline to pale brown, 3–7 μm (av. = 4 ± 0.5 μm) long; apical appendage single, attenuated, smooth, flexuous, unbranched, hyaline, (4–) 7–20 μm (av. = 15 ± 2.5 μm) long; basal appendage single, attenuated, smooth, flexuous, unbranched, excentric, 5–20 μm (av. = 16 ± 3 μm) long; mean conidium length/width ratio = 5(–6):1.

Culture characteristics: Colonies on MEA flat with fluffy aerial mycelium and entire edge, white to buff (19’’f), honey (19-21”b) to vinaceous buff (15-17”’d) at the center, reaching 61–64 mm diam after 14 d at 21 °C; on PDA flat with fluffy aerial mycelium and entire edge, white at the edge to olivaceous grey (21””’i) at the center, reaching 58 mm diam after 14 d at 21 °C.

Specimens examined: Iran: Kurdistan Province: Marivan, Barda Rash, Vitis vinifera (cv. Rasha), 12 Sep. 2012, J. Nahvi Moghadam (IRAN 2300C = CBS 143780).

Sporocadus kurdistanicus Abdollahz., Nahvi M. & Khaledi E., sp. nov.

MycoBank MB838233

Etymology: Name refers to Kurdistan Province in Iran where this species was first found.

Diagnosis: Four isolates of Spo. kurdistanicus clustered in a highly supported clade separated from all Sporocadus species (Fig. 4). Three Sporocadus species including Sporocadus lichenicola, Sporocadus rhododendri and Sporocadus rosigena have previously been reported from grapevine. Spo. lichenicola shows 4 bp (substitution), 8 bp (substitution), 81 bp (28 deletion/insertion, 53 substitutions) and 68 bp (13 deletion/insertion, 55 substitutions) differences with Spo. kurdistanicus. TEF-1α and TUB2 sequences are not available for Spo. rosigena, but in LSU and ITS the type of Spo. rosigena (MFLU 16-0239) has 6 bp and 5 bp differences with Spo. kurdistanicus, respectively. In terms of morphology, conidial dimension of Spo. kurdistanicus is similar with Spo. lichenicola, but conidia of Spo. kurdistanicus are 3-euseptate while in Spo. lichenicola they are 3–4-euseptate and occasionally 5-euseptate. Spo. kurdistanicus is differentiated easlily from Spo. rosigena by having larger conidia (Table 4). No sequences are available for Spo. rhododendri, but Spo. kurdistanicus can be distinguished from Spo. rhododendri by producing larger conidia (Table 4).

Table 4 Conidial characteristics of Sporocadus species reported on grapevine

Species	Conidial dimensions (µm)	Septum no.	Type of appendages	Conidia L/W ratio	Reference
S. kurdistanicus	18–24×6.5–9.5 (av. = 21.5×8)	3	lacking app	3	This study
S. lichenicola	18–25×5.5–8 (av. 21.6×7.2)	3(–4), occasionally 5	lacking app	3	Liu et al 2019
S. rhododendri	15.5–20×6.5–8.5	3	lacking app	–	Pirozynski and Shoemaker 1970
S. rosigena	12–14×5–7.5 (av. = 13×6.5)	3, occasionally 2	lacking app	–	Wanasinghe et al 2018

Type: Iran: Kurdistan Province: Sanandaj, Bavarez, Vitis vinifera (cv. Rasha), 28 Sep. 2012, J. Nahvi Moghadam (IRAN 17870F–holotype; IRAN 2356C = CBS 143778–ex-type culture).

Description: Sexual morph: unknown. Asexual morph: Conidiomata acervular, stromatic, immersed, semi-immersed to erumpent, dark brown to black. Paraphyses 30–40 μm, filiform, cylindrical, aseptate, hyaline, smooth-walled. Conidiophores cylindrical or reduced to conidiogenous cells, hyaline, smooth. Conidiogenous cells discrete, mostly cylindrical, sometimes ampulliform, 5–20 × 1–4 μm (av. = 11.6 ± 3.72 × 2.8 ± 0.58 μm), hyaline, smooth. Conidia fusoid, ellipsoidal to obovoid, subcylindrical, rarely slightly curved, 3-septate, wall smooth, 18–24 × 6.5–9.5 μm (av. = 21.5 ± 0.9 × 8 ± 0.5 μm), lacking appendages; basal cell obconic with a truncate base, pale brown, 2.5–6.5 μm (av. = 4.8 ± 0.92 μm) long; median cells 2, fairly thick-walled, pale brown to brown, doliiform, mostly ± equal, each 6–8 μm (av. = 7 ± 0.5 μm) long, occasionally variable in size, together 10–15 μm (av. = 13.5 ± 1.5 μm) long; apical cell not conic with rounded apex, or conic with obtuse apex, concolourous with median cells, 3–7 μm (av. = 4.5 ± 0.5 μm) long; mean conidium length/width ratio = 3:1.

Culture characteristics: Colonies on MEA flat, appressed to fluffy, folded, edge sinuate, white to buff (19’’f) to sinnamon (13-15”i) at the edge, reaching 43–47 mm diam after 14 d at 21 °C; on PDA flat with fluffy aerial mycelium and a few radial circular line from the center, edge sinuate, buff (19’’f) to vinaceous buff (15-17”’d), wet and cinnamon (13-15”i) to sepia (13-15’’k), at the center reaching 33–45 mm diam after 14 d at 21 °C.

Specimens examined: Iran: Kurdistan Province: Marivan, Ahmadabad, Vitis vinifera (cv. Rasha), 23 Sep. 2012, J. Nahvi Moghadam (IRAN 2354C); Marivan, Nasl-Goshtkhani, Vitis vinifera (cv. Rasha), 14 Sep. 2012, J. Nahvi Moghadam (IRAN 2355C); Dehgolan, Javanmardabad, Vitis vinifera, 10 Sep. 2012, J. Abdollahzadeh & E. Khaledi (IRAN 2313C = CBS 143777).

Sporocadus corni (Wijayawardene, Camporesi & K.D. Hyde) Abdollahz., comb. nov. Mycobank, MB838310

Basionym: Seimatosporium corni Wijayaw., Camporesi & K.D. Hyde, Fungal Diversity73: 100 (2015).

Type: Italy: Pesaro-Urbino Province, Monte Nerone, on branches of Cornus sp., 11
June 2012, Erio Camporesi, (MFLU 15–0742–holotype; MFLUCC 14–0467–ex-type culture).

Description: For a complete description, see Li et al. (2016).

Sporocadus pseudocorni (Wijayawardene, Camporesi & K.D. Hyde) Abdollahz., comb. nov. Mycobank, MB838308

Basionym: Seimatosporium pseudocornii Wijayaw., Camporesi & K.D. Hyde. Fungal Diversity78: 99 (2016).

Type: Italy: Forlì-Cesena Province, near Monte Riccio-Bagno di Romagna, on dead branch of Cornus sp. (Cornaceae), 5 Jan. 2013, Erio Camporesi (MFLU 15–3558–holotype; MFLUCC 13–0529–ex-type culture).

Description: For a complete description, see Senanayake et al. (2015).

Notes: Sporocaduscornicola and Sporocaduspseudocorni are identical in LSU sequence data. ITS, TEF-1α and TUB2 sequences data are not available for Spo. pseudocorni but morphologically these are two distinct species (31–42 × 5–7 μm in Spo. pseudocorni vs. 34–51 × 13–18 μm in Spo. cornicola).

Sporocadus italicus (Q.J. Shang & K.D. Hyde) Abdollahz., comb. nov.

Mycobank, MB838309

Basionym: Seimatosporium italicum Q.J. Shang & K.D. Hyde, Fungal Diversity87: 165 (2017).

Type: Italy: Papiano–Stia, Arezzo Province, on dead aerial branch of Crategus sp., 14 May
2014, E. Camporesi (MFLU 17-0499–holotype; MFLUCC 14-1196–ex-type culture).

Description: For a complete description, see Hyde et al. (2017).

Xenoseimatosporium kurdistanicum Abdollahz., Khaledi E. & Nahvi M., sp. nov.

MycoBank MB838234

Etymology: Name refers to Kurdistan Province in Iran where this species was first found.

Diagnosis: Xen. kurdistanicum is the second introduced species in Xenoseimatosporium after Xenoseimatosporium quercinum. These two species are clearly separated in phylogenetic analyses (Fig. 4). There are 1 bp (substitution), 15 bp (12 substitutions, 3 deletions/insertions), 20 bp (16 substitutions, 4 deletions/insertions) and 15 bp (14 substitutions, 1 deletion/insertion) differences in LSU, ITS, TEF-1α and TUB2 sequences, respectively. These two species are easily distinguishable morphologically by conidial dimensions (29 × 6 μm in Xen. kurdistanicum vs. 18.2 × 4.5 μm in Xen. quercinum), number of septa in conidia (3 in Xen. kurdistanicum vs. 2-4 in Xen. quercinum) and apical/basal appendages length (20/25 μm in Xen. kurdistanicum vs. 13.4/12.1 μm in Xen. quercinum).

Ttpe: Iran: Kurdistan Province: Marivan, Barda Rash, Vitis vinifera (cv. Rasha), 12 Sep. 2012, J. Nahvi Moghadam (IRAN 17871F–holotype; IRAN 2353C–ex-type culture).

Description: Sexual morph: unknown. Asexual morph: Conidiomata acervular, immersed, semi-immersed to erumpent. Conidiophores branched, hyaline, smooth. Conidiogenous cells discrete, cylindrical, oblong to lageniform, 8–15 × 1.5–2.5 μm (av. = 10 ± 1.5 × 2 ± 0.5 μm), hyaline, smooth. Conidia mostly allantoid, occasionally subcylindrical, curved to stright, 3-septate, smooth, some constricted at septa, 22–32 × 4–8 μm (av. = 29 ± 2 × 6 ± 0.9 μm), bearing appendages; basal cell obconic with truncate base or trapezoid, thin-walled, hyaline to pale brown, 2–4 μm (av. = 3 ± 0.3 μm) long; median cells 2, mostly cylindrical, occasionally doliiform, pale to mid-brown, thin-walled, ± equal, each 8–12 μm (av. = 10 ± 0.8 μm) long; apical cell conic with an acute or rounded apex, hyaline to pale brown, 2–4 μm (av. = 3 ± 0.5 μm); apical appendage single, attenuated, smooth, flexuous, unbranched, 15–30 μm (av. = 20 ± 2 μm); basal appendage single, attenuated, smooth, flexuous, unbranched, excentric, 18–33 μm (av. = 25 ± 1.8 μm) long; mean conidium length/width ratio = 5(–6):1.

Culture characteristics: Colonies on MEA flat, appressed to fluffy, folded, edge sinuate, buff (19’’f) to sinnamon (13-15”i), reaching 45–50 mm diam after 14 d at 21 °C; on PDA flat with entire edge, fluffy, buff (19’’f) to vinaceous buff (15-17”’d), reaching 55–61 mm diam after 14 d at 21 °C.

Specimens examined: Iran: Kurdistan Province: Kamyaran, Bovana, Vitis vinifera, 18 Sep. 2012, E. Khaledi (IRAN 2305C).

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