Skip to main content
Fungal Systematics and Evolution logoLink to Fungal Systematics and Evolution
. 2021 Apr 28;7:255–343. doi: 10.3114/fuse.2021.07.13

New and Interesting Fungi. 4

PW Crous 1,2,*, M Hernández-Restrepo 1, RK Schumacher 3, DA Cowan 4, G Maggs-Kölling 5, E Marais 5, MJ Wingfield 2, N Yilmaz 2, OCG Adan 6, A Akulov 7, E Álvarez Duarte 8, A Berraf-Tebbal 9, TS Bulgakov 10, AJ Carnegie 11,12, ZW de Beer 2, C Decock 13, J Dijksterhuis 1, TA Duong 2, A Eichmeier 9, LT Hien 14, JAMP Houbraken 1, TN Khanh 14, NV Liem 14, L Lombard 1, FM Lutzoni 15, JM Miadlikowska 15, WJ Nel 2, IG Pascoe 16, F Roets 17, J Roux 18, RA Samson 1, M Shen 19, M Spetik 9, R Thangavel 20, HM Thanh 14, LD Thao 14, EJ van Nieuwenhuijzen 1, JQ Zhang 19, Y Zhang 19, LL Zhao 19, JZ Groenewald 1
PMCID: PMC8165967  PMID: 34124627

Abstract

An order, family and genus are validated, seven new genera, 35 new species, two new combinations, two epitypes, two lectotypes, and 17 interesting new host and / or geographical records are introduced in this study. Validated order, family and genus: Superstratomycetales and Superstratomycetaceae (based on Superstratomyces). New genera: Haudseptoria (based on Haudseptoria typhae); Hogelandia (based on Hogelandia lambearum); Neoscirrhia (based on Neoscirrhia osmundae); Nothoanungitopsis (based on Nothoanungitopsis urophyllae); Nothomicrosphaeropsis (based on Nothomicrosphaeropsis welwitschiae); Populomyces (based on Populomyces zwinianus); Pseudoacrospermum (based on Pseudoacrospermum goniomae). New species: Apiospora sasae on dead culms of Sasa veitchii (Netherlands); Apiospora stipae on dead culms of Stipa gigantea (Spain); Bagadiella eucalyptorum on leaves of Eucalyptus sp. (Australia); Calonectria singaporensis from submerged leaf litter (Singapore); Castanediella neomalaysiana on leaves of Eucalyptus sp. (Malaysia); Colletotrichum pleopeltidis on leaves of Pleopeltis sp. (South Africa); Coniochaeta deborreae from soil (Netherlands); Diaporthe durionigena on branches of Durio zibethinus (Vietnam); Floricola juncicola on dead culm of Juncus sp. (France); Haudseptoria typhae on leaf sheath of Typha sp. (Germany); Hogelandia lambearum from soil (Netherlands); Lomentospora valparaisensis from soil (Chile); Neofusicoccum mystacidii on dead stems of Mystacidium capense (South Africa); Neomycosphaerella guibourtiae on leaves of Guibourtia sp. (Angola); Niesslia neoexosporioides on dead leaves of Carex paniculata (Germany); Nothoanungitopsis urophyllae on seed capsules of Eucalyptus urophylla (South Africa); Nothomicrosphaeropsis welwitschiae on dead leaves of Welwitschia mirabilis (Namibia); Paracremonium bendijkiorum from soil (Netherlands); Paraphoma ledniceana on dead wood of Buxus sempervirens (Czech Republic); Paraphoma salicis on leaves of Salix cf. alba (Ukraine); Parasarocladium wereldwijsianum from soil (Netherlands); Peziza ligni on masonry and plastering (France); Phyllosticta phoenicis on leaves of Phoenix reclinata (South Africa); Plectosphaerella slobbergiarum from soil (Netherlands); Populomyces zwinianus from soil (Netherlands); Pseudoacrospermum goniomae on leaves of Gonioma kamassi (South Africa); Pseudopyricularia festucae on leaves of Festuca californica (USA); Sarocladium sasijaorum from soil (Netherlands); Sporothrix hypoxyli in sporocarp of Hypoxylon petriniae on Fraxinus wood (Netherlands); Superstratomyces albomucosus on Pycnanthus angolensis (Netherlands); Superstratomyces atroviridis on Pinus sylvestris (Netherlands); Superstratomyces flavomucosus on leaf of Hakea multilinearis (Australia); Superstratomyces tardicrescens from human eye specimen (USA); Taeniolella platani on twig of Platanus hispanica (Germany), and Tympanis pini on twigs of Pinus sylvestris (Spain).

Citation: Crous PW, Hernández-Restrepo M, Schumacher RK, Cowan DA, Maggs-Kölling G, Marais E, Wingfield MJ, Yilmaz N, Adan OCG, Akulov A, Álvarez Duarte E, Berraf-Tebbal A, Bulgakov TS, Carnegie AJ, de Beer ZW, Decock C, Dijksterhuis J, Duong TA, Eichmeier A, Hien LT, Houbraken JAMP, Khanh TN, Liem NV, Lombard L, Lutzoni FM, Miadlikowska JM, Nel WJ, Pascoe IG, Roets F, Roux J, Samson RA, Shen M, Spetik M, Thangavel R, Thanh HM, Thao LD, van Nieuwenhuijzen EJ, Zhang JQ, Zhang Y, Zhao LL, Groenewald JZ (2021). New and Interesting Fungi. 4. Fungal Systematics and Evolution 7: 255–343. doi: 10.3114/fuse.2021.07.13

Keywords: biodiversity, ITS barcodes, multi-gene phylogeny, new taxa, systematics, typification

INTRODUCTION

The present study represents the fourth instalment of the New and Interesting Fungi (NIF) series that is published annually in the journal Fungal Systematics and Evolution. Papers report new knowledge on fungal biodiversity, list new host or geographical records, and new sexual-asexual connections. The present study also includes validations and descriptions of new fungal taxa, and lists interesting observations relating to fungal biology. Mycologists and other researchers wishing to contribute to future issues of NIF are encouraged to contact the Editor-in-Chief (p.crous@wi.knaw.nl).

MATERIALS AND METHODS

Isolates

Twig and leaf samples collected from around the world (see Table 1) were treated as previously detailed (Crous et al. 2019b), while the treatment of soil samples followed the methods of Giraldo et al. (2019) and Hou et al. (2020b). Single conidial colonies were established on Petri dishes containing 2 % malt extract agar (MEA) as described by Crous et al. (1991), and single ascospore cultures were established following the method described by Crous (1998). Colonies were sub-cultured on 2 % potato-dextrose agar (PDA), oatmeal agar (OA), MEA (Crous et al. 2019b), or autoclaved pine needles on 2 % tap water agar (PNA) (Smith et al. 1996), and incubated at 25 °C under continuous near-ultraviolet light to promote sporulation. Reference strains and specimens of the studied fungi are maintained in the culture collection (CBS) of the Westerdijk Fungal Biodiversity Institute (WI), Utrecht, the Netherlands.

Table 1.

Collection details and GenBank accession numbers of isolates treated in this study, and associated ex-type strains where available. Species for which additional sequences were generated during the course of this study are also listed here.

Species Culture or voucher accession number(s)1 Locality and Substrate Collector(s) GenBank accession number2
ITS LSU rpb2 tub2 Other loci
Alternaria chartarum CBS 200.67 = ATCC 18044 = DAOM 59616b = IMI 124943 = MUCL 18564 = QM 8328, ex-epitype Canada: Populus plywood E.G. Simmons MH858944.1 NG_069727.1 KC584481.1 gapdh: KC584172.1, SSU: NG_062919.1, tef1 (first part): KC584741.1
Alternaria chartarum CPC 38971 Namibia: Lichen hypolith under a rock P.W. Crous MW883400.1 MW883795.1 MW890118.1 actA: MW890017.1, cmdA: MW890037.1, gapdh: MW890046.1, tef1 (first part): MW890080.1
Alternaria heterospora CBS 123376, ex-type of Ulocladium solani China: Lycopersicon esculentum, diseased leaves Y. Wang KC584248.1 KC584363.1 KC584488.1 gapdh: KC584176.1, SSU: KC584621.1, tef1 (first part): KC584748.1
CPC 38969 Namibia: Lichen hypolith under a rock P.W. Crous MW883401.1 MW883796.1 MW890119.1 actA: MW890018.1, cmdA: MW890038.1, gapdh: MW890047.1, tef1 (first part): MW890081.1
Apiospora sasae, sp. nov. CBS 146808 = CPC 38165, ex-type Netherlands: Sasa veitchii, dead culms L. van der Linde MW883402.1 MW883797.1 MW890058.1 MW890120.1 tef1 (second part): MW890104.1
Apiospora stipae, sp. nov. CBS 146804 = CPC 38101, ex-type Spain: Stipa gigantea, dead culm M.A. Delgado MW883403.1 MW883798.1 MW890059.1 MW890121.1 tef1 (first part): MW890082.1, tef1 (second part): MW890105.1
Bagadiella eucalyptorum, sp. nov. CBS 147177 = CPC 39299, ex-type Australia: Eucalyptus sp., leaves A.J. Carnegie MW883404.1 MW883799.1
Blastacervulus metrosideri CBS 147006 = CPC 38759 = T19_05741C New Zealand: Metrosideros sp., leaves L. Rabbidge MW883405.1 MW883800.1
ICMP 21883, ex-type New Zealand: Metrosideros excelsa, living leaves P.R. Johnston NR_169959.1 NG_068290.1
Calonectria singaporensis, sp. nov. CBS 146712 = MUCL 048012 Singapore: Submerged leaf litter in a small stream C. Decock MW883406.1 MW883801.1 MW890122.1 actA: MW890019.1, cmdA: MW890039.1, his3: MW890052.1, tef1 (first part): MW890083.1
CBS 146713 = MUCL 048171 Singapore: Submerged leaf litter in a small stream C. Decock MW883407.1 MW883802.1 MW890123.1 actA: MW890020.1, cmdA: MW890040.1, his3: MW890053.1, tef1 (first part): MW890084.1
CBS 146714 = MUCL 048187 Singapore: Submerged leaf litter in a small stream C. Decock MW883408.1 MW883803.1 actA: MW890021.1, cmdA: MW890041.1, his3: MW890054.1, tef1 (first part): MW890085.1
CBS 146715 = MUCL 048320, ex-type Singapore: Submerged leaf litter in a small stream C. Decock MW883409.1 MW883804.1 MW890124.1 actA: MW890022.1, cmdA: MW890042.1, his3: MW890055.1, tef1 (first part): MW890086.1
Castanediella neomalaysiana, sp. nov. CBS 147093 = CPC 39275, ex-type Malaysia: Eucalyptus sp., leaves M.J. Wingfield MW883410.1 MW883805.1
Colletotrichum kinghornii CBS 198.35, ex-type UK: Phormium tenax N.L. Alcock NR_111751.1 NG_069631.1 JQ950105.1 actA: JQ949775.1, chs-1: JQ949115.1, gapdh: JQ948785.1, his3: JQ949445.1
CPC 38766 = T19_05774B New Zealand: Phormium cookianum C. Inglis MW883411.1 MW883806.1 MW890125.1 actA: MW890023.1, gapdh: MW890048.1
CBS 147082 = CPC 39342 South Africa: Pleopeltis sp. J. Roux MW883412.1 MW883807.1 actA: MW890024.1, chs-1: MW890035.1,
Coniochaeta deborreae, sp. nov. CBS 147215 = BE19_001008, ex-type Netherlands: Soil K. de Borre MW883413.1 MW883808.1 tef1 (first part): MW890087.1, tef1 (second part): MW890106.1
CBS 551.75 Norway: Pinus sylvestris, wood MW883416.1 MW883809.1
Coniochaeta pulveracea CBS 114628 = D3409 = UNI 393 Turkey: Rinsing machine in soft drinks factory M. Stratford MW883414.1 GQ351560.1
Coniochaeta rhopalochaeta CBS 109872 = BAFC 272, ex-type Argentina: Bulnesia retamas, decorticated wood C.C. Carmarán MW883415.1 GQ351561.1
Diaporthe durionigena, sp. nov. KCSR1812.8 = VTCC 930005, ex-type Vietnam: Durio zibethinus, branches L.D. Thao MN453530.1 MT276159.1 tef1 (first part): MT276157.1
Endoconidioma euphorbiae CBS 146776 = CPC 38551, ex-type South Africa: Euphorbia mauritanica, leaf tip dieback P.W. Crous MW175350.1 MW175390.1
CPC 38649 South Africa: Euphorbia mauritanica, tip dieback P.W. Crous MW883417.1 MW883810.1 MW890060.1 MW890126.1 actA: MW890025.1, tef1 (second part): MW890107.1
Flammocladiella aceris CBS 138906 = CPC 24422, ex-type Germany: Acer platanoides, twigs R.K. Schumacher NG_058175.1 KR611901.1 MW890061.1 MW890127.1 tef1 (first part): MW890088.1
Flammocladiella anomiae CBS 142775 = CLL16017 Bulgaria: On stromata of Massaria anomia, on a thin branch of Robinia pseudoacacia D. Stoykov MN597422.1 MW883811.1 MW890062.1 MW890128.1 tef1 (first part): MW890089.1
CBS 144256 = JF17087, ex-type France: On ascomata of Massaria anomia on a branchlet of Robinia pseudoacacia J. Fournier MN597423.1 MN597425.1 MW890129.1 tef1 (first part): MW890090.1
CBS 146685 = CPC 36302 Ukraine: On conidiomata of Diaporthe oncostoma, on dead branch of Robinia pseudoacacia A. Akulov MW883418.1 MW883812.1 MW890130.1 tef1 (first part): MW890091.1
Floricola juncicola, sp. nov. CBS 146811 = CPC 38197, ex-type France: Juncus sp., dead culm A. Gardiennet MW883419.1 MW883813.1 MW890063.1 tef1 (first part): MW890092.1, tef1 (second part): MW890108.1
Harzia tenella CPC 38667 South Africa: Grielum humifusum, stems P.W. Crous MW883420.1 MW883814.1
Haudseptoria typhae, gen. et sp. nov. CBS 146790 = CPC 38203, ex-type Germany: Typha sp., dead leaf sheath R.K. Schumacher MW883421.1 MW883815.1 MW890131.1
Heimiodora verticillata CBS 147089 = CPC 39015 Namibia: Salvadora sp. P.W. Crous MW883422.1 MW883816.1
CBS 201.60 = IMI 090702 = LCP 57.1589, ex-type Thailand: Sandy coastal soil MH857955.1 MH869505.1
Hogelandia lambearum, gen. et sp. nov. CBS 147626 = NL19_27007, ex-type Netherlands: Soil L. Alssema & M. van Berkel MW883423.1 MW883817.1
Lomentospora valparaisensis, sp. nov. ChFC-164, ex-type Chile: Soil F. Salas MG495075.1 MG544878.1
Muyocopron zamiae CBS 146636 = CPC 37461 USA: Zamia integrifolia, leaves M.J. Wingfield MW883424.1 MW883818.1 MW890064.1 tef1 (second part): MW890109.1
CBS 202.71 = No. 070-2288 USA: Zamia integrifolia, leaf spot and necrotic tip MW883425.1
CBS 203.71 = No. 070-2273, ex-type USA: Zamia fisheri, leaf spot and necrotic tip MW883426.1 NG_066338.1 MK492731.1 tef1 (second part): MK495973.1
Neocamarosporium leipoldtiae CBS 146812 = CPC 38543, ex-type South Africa: Cephalophyllum pilansii, leaves P.W. Crous MW883427.1 MW883819.1 MW890132.1 tef1 (first part): MW890093.1
Neofusicoccum mystacidii, sp. nov. CBS 147079 = CPC 39221, ex-type South Africa: Mystacidium capense, dead stems P.W. Crous MW883428.1 MW883820.1 MW890065.1 MW890133.1 gapdh: MW890049.1, his3: MW890056.1, tef1 (first part): MW890094.1
Neomycosphaerella guibourtiae, sp. nov. CBS 147083 = CPC 39348, ex-type Angola: Guibourtia sp., leaves J. Roux MW883429.1 MW883821.1 MW890134.1 actA: MW890026.1, cmdA: MW890043.1, tef1 (first part): MW890095.1
Neoscirrhia matteucciicola, comb. nov. CBS 259.92 = IMI 286996 = PD 91/272, ex-type Canada: Matteuccia struthiopteris, affected leaf stem GU237812.1 GU238100.1 GU237627.1
Neoscirrhia osmundae, gen. et comb. nov. CBS 146803 = CPC 38085, ex-type Netherlands: Sasa veitchii, culms L. van der Linde MW883430.1 MW883822.1 MW890066.1 MW890135.1
Neoscytalidium dimidiatum CBS 146816 = CPC 38666, ex-type South Africa: Epiphyte on stems of Aloidendron dichotomum P.W. Crous MW883431.1 MW883823.1 MW890136.1 chs-1: MW890036.1, tef1 (first part): MW890096.1
Niesslia neoexosporioides, sp. nov. CBS 146810 = CPC 38177, ex-type Germany: Carex paniculata, dead leaves R.K. Schumacher MW883432.1 MW883824.1 MW890137.1 actA: MW890027.1, tef1 (first part): MW890097.1
Nothoanungitopsis urophyllae, gen. et sp. nov. CBS 146799 = CPC 38059, ex-type South Africa: Eucalyptus urophylla, seed capsules M.J. Wingfield MW883433.1 MW883825.1
Nothomicrosphaeropsis welwitschiae gen. et sp. nov. CBS 146829 = CPC 38879, ex-type Namibia: Welwitschia mirabilis, dead leaves P.W. Crous MW883434.1 MW883826.1 MW890067.1 MW890138.1
Ophioceras leptosporum CBS 147090 = CPC 39147 South Africa: Syzygium cordatum, twigs P.W. Crous MW883435.1 MW883827.1 tef1 (second part): MW890110.1
CBS 894.70 = ATCC 24161 = HME 2955, ex-type UK: Dead stem of dicot plant, probably Urtica dioica NR_111768.1 NG_057959.1 rpb1: JX134732.1, SSU: JX134664.1, tef1 (second part): JX134704.1
Paracremonium bendijkiorum, sp. nov. CBS 147228 = NL19_24005, ex-type Netherlands: Soil Y. Bentem & J. van Dijken MW883436.1 MW883828.1 MW890068.1 MW890139.1 tef1 (second part): MW890111.1
Paraphoma ledniceana, sp. nov. CBS 146533 = MEND-F-82, ex-type Czech Republic: Saprobe on dead wood of Buxus sempervirens M. Spetik MT371091.1 MT371396.1 MT372655.1 MT372661.1 tef1 (first part): MT372654.1
Paraphoma salicis, sp. nov. CBS 146797 = CPC 38651, ex-type Ukraine: Salix cf. alba, leaves A. Akulov MW883437.1 MW883829.1 MW890069.1 MW890140.1 actA: MW890028.1
Parasarocladium wereldwijsianum, sp. nov. CBS 147223 = NL19094001 Netherlands: Soil S. Frederikze, J. Mes & S. Maghnouji MW883438.1 MW890070.1 MW890141.1
CBS 147224 = NL19094011 Netherlands: Soil S. Frederikze, J. Mes & S. Maghnouji MW883439.1 MW883830.1 MW890071.1 MW890142.1 actA: MW890029.1, tef1 (second part): MW890112.1
CBS 147226 = NL19095011, ex-type Netherlands: Soil S. Frederikze, J. Mes & S. Maghnouji MW883440.1 MW883831.1 MW890072.1 MW890143.1 actA: MW890030.1, tef1 (second part): MW890113.1
Peziza ligni, sp. nov. CBS 146637 = CPC 39110 = MUCL 57889, ex-type France: On masonry and plastering, near a wooden staircase C. Decock MW883441.1 MW883832.1 MW890073.1
Phyllosticta phoenicis, sp. nov. CBS 147091 = CPC 39164, ex-type South Africa: Phoenix reclinata, leaves M.J. Wingfield MW883442.1 MW883833.1 actA: MW890031.1, gapdh: MW890050.1, tef1 (first part): MW890098.1
Plectosphaerella slobbergiarum, sp. nov. CBS 147227 = NL1930002, ex-type Netherlands: Soil J. Slob & M. Berghuis MW883443.1 MW883834.1 MW890074.1 tef1 (second part): MW890114.1
Populomyces zwinianus, gen. et sp. nov. CBS 147307 = NL1976004, ex-type Netherlands: Soil W. Vercouteren, S. Meas & R. Verhije MW883444.1 MW883835.1
Porodiplodia livistonae CBS 144428 = CPC 32154, ex-type Australia: Livistona australis, leaves P.W. Crous NR_160355.1 NG_069575.1 tef1 (first part): MW890099.1
Porodiplodia vitis CBS 144634 = CPC 31642, ex-type USA: Vitis vinifera, canes E. Crenson & R.K. Schumacher NR_163376.1 NG_070080.1 tef1 (first part): MK442707.1
CBS 146818 = CPC 38692 South Africa: Virgilia oroboides, seed pods M.J. Wingfield MW883445.1 MW883836.1 tef1 (first part): MW890100.1
Pseudoacrospermum goniomae, gen. et sp. nov. CBS 146732 = CPC 37030, ex-type South Africa: Gonioma kamassi, leaves F. Roets MW883446.1 MW883837.1 MW890075.1 tef1 (first part): MW890101.1, tef1 (second part): MW890115.1
Pseudopyricularia festucae, sp. nov. CBS 146629 = CPC 37915, ex-type USA: Festuca californica, leaves P.W. Crous MW883447.1 MW883838.1 cmdA: MW890044.1, rpb1: MW890057.1
Sarocladium sasijaorum, sp. nov. CBS 147213 = NL19100007, ex-type Netherlands: Soil S. Frederikze, J. Mes & S. Maghnouji MW883448.1 MW883839.1 MW890076.1 MW890144.1 actA: MW890032.1, tef1 (second part): MW890116.1
Septoria protearum CBS 778.97 = CPC 1470 = STE-U 1470 = IMI 375230 = ATCC 201159, ex-type South Africa: Protea cynaroides, leaves L. Viljoen KF251523.1 KF252028.1 KF252517.1 KF252992.1 actA: KF253827.1, cmdA: KF254176.1, tef1 (first part): KF253472.1
CPC 38736 = T19_05709B New Zealand: Kniphofia uvaria D. Burnt MW883449.1 MW883840.1 actA: MW890033.1, tef1 (first part): MW890102.1
Sporothrix hypoxyli, sp. nov. CBS 141567 = CMW 47439 Netherlands: Ascomata embedded in fruiting body of Hypoxylon petriniae on Fraxinus wood E. Osieck & W.J. Nel MT637056.1 MW012947.1 MT649176.1
CBS 141568 = CMW 47436 Netherlands: Ascomata embedded in fruiting body of Hypoxylon petriniae on Fraxinus wood E. Osieck & W.J. Nel MT637053.1 MW012946.1 MT649173.1
CBS 141569 = CMW 47441, ex-type Netherlands: Ascomata embedded in fruiting body of Hypoxylon petriniae on Fraxinus wood E. Osieck & W.J. Nel MT637058.1 MW012948.1 MT649178.1
Stemphylium eturmiunum CBS 109845 = EGS 29.099, ex-type New Zealand: Lycopersicon esculentum, fruit E.G. Simmons NR_154927.1 NG_069866.1 cmdA: KU850831.1, gapdh: KU850689.1
CBS 146783 = CPC 38613, ex-type South Africa: Bulbinella latifolia, leaves P.W. Crous MW883450.1 MW883841.1 MW890077.1 MW890145.1 actA: MW890034.1, cmdA: MW890045.1, gapdh: MW890051.1, tef1 (first part): MW890103.1
Superstratomyces albomucosus, gen. et sp. nov. CBS 140270 = DTO 277-D2, ex-type Netherlands: Outdoor exposed Pycnanthus angolensis impregnated with olive oil E.J. van Nieuwenhuijzen NR_152544.1 KX950439.1 KX950498.1 rpb1: KX950494.1, SSU: NG_061256.1, tef1 (second part): KX950471.1
Superstratomyces atroviridis, sp. nov. CBS 140272 = DTO 305-E1, ex-type Netherlands: Outdoor exposed Pinus sylvestris impregnated with raw linseed oil E.J. van Nieuwenhuijzen NR_152545.1 NG_058271.1 KX950500.1 rpb1: KX950496.1, SSU: NG_063075.1, tef1 (second part): KX950473.1
Superstratomyces flavomucosus, sp. nov. CBS 353.84 = DTO 305-C3, ex-type Australia: Hakea multilinearis, leaf W. Gams NR_152543.1 KX950438.1 KX950497.1 rpb1: KX950493.1, SSU: NG_065661.1, tef1 (second part): KX950470.1
Superstratomyces tardicrescens, sp. nov. FMR 13786, ex-type USA: Human eye specimen D.A. Sutton LR025130.1 LR025130.1 tef1 (second part): LR025141.1
Taeniolella exilis CBS 122902 = DAOM 14683 = MUCL 1878 Canada: Betula papyrifera, bark G. Hennebert MW883451.1 KX244968.1
Taeniolella platani, sp. nov. CBS 146733 = CPC 33568, ex-type Germany: Platanus hispanica, twig R.K. Schumacher MW883452.1 MW883842.1 tef1 (second part): MW890117.1
Tricellula aurantiaca CBS 146627 = CPC 36629 Russia: Lonicera tatarica, leaves T.S. Bulgakov MW883453.1 MW883843.1
CBS 399.58 = ATCC 13128 = IMI 073024 = MUCL 28102 = PRL 1554, ex-type Canada: Soil MH857822.1 MH869354.1
Tryssglobulus aspergilloides CBS 147388 = CPC 40100, ex-isoepitype Australia: Banksia marginata I.G. Pascoe & B.J. Brentwood MW883454.1 MW883844.1 MW890078.1
CBS 147556 = CPC 40369 = VPRI 43962, ex-epitype Australia: Banksia marginata I.G. Pascoe & B.J. Brentwood MW883455.1 MW883845.1 MW890079.1
Tympanis pini, sp. nov. CBS 146809 = CPC 38169, ex-type Spain: Pinus sylvestris, twigs R. Blasco MW883456.1 MW883846.1
Venturia cerasi CBS 444.54, ex-epitype Germany: Prunus cerasus H. Schweizer NR_168750.1 MH868928.1 MK887849.1 MK926519.1 tef1 (first part): MK888785.1
Zelosatchmopsis sacciformis CBS 116.88 = INIFAT C87/53.1 = MW i 1640, ex-isotype Cuba: Guazuma ulmifolia, fallen leaf R.F. Castañeda MH862122.1 MH873812.1

1 ATCC: American Type Culture Collection, Virginia, USA; BAFC: Universidad de Buenos Aires, Departamento de Ciencias Biologicas; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; CMW: Culture Collection of the Forestry and Agricultural Biotechnology Institute (FABI) of the University of Pretoria, Pretoria, South Africa; CPC: Culture collection of Pedro Crous, housed at CBS; DAOM: Plant Research Institute, Department of Agriculture (Mycology), Ottawa, Canada; DTO: Working collection of the Indoor air and Industrial Mycology group, housed at CBS; FMR: Facultat de Medicina, Universitat Rovira i Virgili, Reus, Spain; ICMP: International Collection of Micro-organisms from Plants, Landcare Research, Private Bag 92170, Auckland, New Zealand; IMI: International Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane, United Kingdom; INIFAT: Alexander Humboldt Institute for Basic Research in Tropical Agriculture, Ciudad de La Habana, Cuba; LCP: Laboratory of Cryptogamy, National Museum of Natural History, Paris, France; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Rai, Thailand; MUCL: Université Catholique de Louvain, Louvain-la-Neuve, Belgium; VPRI: Victorian Department of Primary Industries, Knoxfield, Australia; VTCC: Vietnam Type Culture Collection, Center of Biotechnology, Vietnam National University, Hanoi, Vietnam.

2 ITS: internal transcribed spacers and intervening 5.8S nrDNA; LSU: large subunit (28S) of the nrRNA gene operon; actA: partial actin gene; chs-1: partial chitin synthase-1 gene; cmdA: partial calmodulin gene; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase gene; his3: partial histone H3 gene; rpb1: partial DNA-directed RNA polymerase II largest subunit gene; rpb2: partial DNA-directed RNA polymerase II second largest subunit gene; SSU: small subunit (18S) of the nrRNA gene operon; tef1: partial translation elongation factor 1-alpha gene; tub2: partial beta-tubulin gene.

DNA extraction, amplification (PCR) and phylogeny

Fungal mycelium (Table 1) was scraped from the surface of agar cultures with a sterile scalpel and the genomic DNA was isolated using the Wizard® Genomic DNA Purification Kit (Promega Corporation, WI, USA) following the manufacturers’ protocols. All loci were amplified following previously published protocols. First, the 28S nrRNA gene (LSU) and internal transcribed spacer regions with intervening 5.8S nrRNA gene (ITS) of the nrDNA operon were sequenced for all the isolates included in this study (for amplification conditions, see Fan et al. 2018). Other loci were sequenced for various species or genera using primers and conditions specific for those groups of fungi. Amplification of the partial DNA-directed RNA polymerase II second largest subunit gene (rpb2), the partial translation elongation factor 1-alpha gene (tef1, first part) and the partial beta-tubulin gene (tub2) followed Braun et al. (2018), while amplification of the partial actin gene (actA), the partial calmodulin gene (cmdA), the partial glyceraldehyde-3-phosphate dehydrogenase gene (gapdh) and the partial histone H3 gene (his3) followed Videira et al. (2016). Amplification of the partial DNA-directed RNA polymerase II largest subunit gene (rpb1) followed Klaubauf et al. (2014), the partial translation elongation factor 1-alpha gene (tef1, second part) followed Réblová et al. (2020) and of the partial chitin synthase-1 (chs-1) followed Damm et al. (2019). The resulting fragments were sequenced in both directions using the respective PCR primers and the BigDye Terminator Cycle Sequencing Kit v. 3.1 (Applied Biosystems Life Technologies, Carlsbad, CA, USA); DNA sequencing amplicons were purified through Sephadex G-50 Superfine columns (Sigma-Aldrich, St. Louis, MO) in MultiScreen HV plates (Millipore, Billerica, MA). Purified sequence reactions were analysed on an Applied Biosystems 3730xl DNA Analyzer (Life Technologies, Carlsbad, CA, USA). The DNA sequences were analysed and consensus sequences were computed using Geneious v. 11.1.5 (http://www.geneious.com, Kearse et al. 2012).

The sequences for each gene region were subjected to megablast searches (Zhang et al. 2000) to identify closely related sequences in the NCBI’s GenBank nucleotide database. The results are provided as part of the species notes or as selected phylogenetic trees. Phylogenetic trees were generated using Bayesian analyses performed with MrBayes v. 3.2.7a (Ronquist et al. 2012) for the overview trees and Maximum Parsimony analyses performed with PAUP v. 4.0b10 (Swofford 2003) as explained in Braun et al. (2018) for the genus and species trees. All resulting trees were printed with Geneious v. 11.1.5 and the layout of the trees was done in Adobe Illustrator v. CC 2017. Statistical measures calculated for the parsimony analyses included tree length (TL), consistency index (CI), retention index (RI) and rescaled consistency index (RC). Maximum-likelihood trees were generated with IQ-TREE v. 1.6.12 (Nguyen et al. 2015) and branch support values were calculated with 5 000 ultrafast bootstrap replicates (Hoang et al. 2018). Best models were estimated for data partitions using ModelFinder (Kalyaanamoorthy et al. 2017) as implemented in IQ-TREE.

Morphology

Slide preparations were mounted in lactic acid, Shear’s mounting fluid or water, from colonies sporulating on MEA, PDA, PNA or OA. Observations were made with a Nikon SMZ25 dissection-microscope, and with a Zeiss Axio Imager 2 light microscope using differential interference contrast (DIC) illumination and images recorded on a Nikon DS-Ri2 camera with associated software. Cryo Scanning Electron Microscopy methods followed Bensch et al. (2018). Colony characters and pigment production were noted after 2–4 wk of growth on MEA, PDA and OA (Crous et al. 2019b) incubated at 25 °C. Colony colours (surface and reverse) were scored using the colour charts of Rayner (1970). Sequences derived in this study were deposited in GenBank (Table 1), the alignments in TreeBASE (www.treebase.org; study number S28000), and taxonomic novelties in MycoBank (www.MycoBank.org; Crous et al. 2004).

RESULTS

Phylogeny

Dothideomycetes LSU phylogeny (Fig. 1, parts 1–5): The alignment contained 260 isolates and the tree was rooted to Diaporthe perjuncta (strain BPI 748437, GenBank NG_059064.1). The final alignment contained a total of 776 characters used for the phylogenetic analyses, including alignment gaps. The alignment contained a total of 538 unique site patterns. Based on the results of MrModelTest, dirichlet base frequencies and the GTR+I+G model was used for the Bayesian analysis. The Bayesian analyses generated 309 402 trees (saved every 100 generations) from which 232 052 were sampled after 25 % of the trees were discarded as burn-in.

Fig. 1, parts 1–5.

Fig. 1, parts 1–5.

Fig. 1, parts 1–5.

Fig. 1, parts 1–5.

Fig. 1, parts 1–5.

Fig. 1, parts 1–5.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Dothideomycetes LSU nucleotide alignment. Bayesian posterior probabilities (PP) > 0.79 are shown at the nodes and the scale bar represents the expected changes per site. Thickened branches represent PP = 1. The branch leading to Superstratomycetales was halved to facilitate layout. Families and orders are indicated with coloured blocks to the right of the tree. GenBank accession (superscript) and / or culture collection / voucher numbers are indicated for all species. The tree was rooted to Diaporthe perjuncta (voucher BPI 748437, GenBank NG_059064.1) and the species treated in this study for which LSU sequence data were available are indicated in bold face.

Leotiomycetes LSU phylogeny (Fig. 2): The alignment contained 61 isolates and the tree was rooted to Xylaria hypoxylon (voucher OSC 100004, GenBank AY544648.1). The final alignment contained a total of 760 characters used for the phylogenetic analyses, including alignment gaps. The alignment contained a total of 185 unique site patterns. Based on the results of MrModelTest, dirichlet base frequencies and the GTR+I+G model was used for the Bayesian analysis. The Bayesian analyses generated 542 002 trees (saved every five generations) from which 406 502 were sampled after 25 % of the trees were discarded as burn-in.

Fig. 2.

Fig. 2.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Leotiomycetes LSU nucleotide alignment. Bayesian posterior probabilities (PP) > 0.79 are shown at the nodes and the scale bar represents the expected changes per site. Thickened branches represent PP = 1 and the most basal branch was halved to facilitate layout. Families and the order Helotiales are indicated with coloured blocks to the right of the tree. GenBank accession (superscript) and / or culture collection / voucher numbers are indicated for all species. The tree was rooted to Xylaria hypoxylon (voucher OSC 100004, GenBank AY544648.1) and the species treated in this study for which LSU sequence data were available are indicated in bold face.

Pezizomycetes LSU phylogeny (Fig. 3): The alignment contained 37 isolates and the tree was rooted to Candida broadrunensis (strain CBS 11838, GenBank KY106372.1). The final alignment contained a total of 739 characters used for the phylogenetic analyses, including alignment gaps. The alignment contained a total of 279 unique site patterns. Based on the results of MrModelTest, dirichlet base frequencies and the GTR+I+G model was used for the Bayesian analysis. The Bayesian analyses generated 376 002 trees (saved every five generations) from which 282 002 were sampled after 25 % of the trees were discarded as burn-in.

Fig. 3.

Fig. 3.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Pezizomycetes LSU nucleotide alignment. Bayesian posterior probabilities (PP) > 0.79 are shown at the nodes and the scale bar represents the expected changes per site. Thickened branches represent PP = 1. Families and the order Pezizales are indicated with coloured blocks to the right of the tree. GenBank accession (superscript) and / or culture collection / voucher numbers are indicated for all species. The tree was rooted to Candida broadrunensis (culture CBS 11838, GenBank KY106372.1) and the species treated in this study for which LSU sequence data were available are indicated in bold face.

Sordariomycetes (Hypocreales) LSU phylogeny (Fig. 4): The alignment contained 70 isolates and the tree was rooted to Ramularia endophylla (strain CBS 113265, GenBank AY490776.1). The final alignment contained a total of 752 characters used for the phylogenetic analyses, including alignment gaps. The alignment contained a total of 199 unique site patterns. Based on the results of MrModelTest, dirichlet base frequencies and the GTR+I+G model was used for the Bayesian analysis. The Bayesian analyses generated 426 002 trees (saved every five generations) from which 319 502 were sampled after 25 % of the trees were discarded as burn-in.

Fig. 4.

Fig. 4.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Sordariomycetes (Hypocreales) LSU nucleotide alignment. Bayesian posterior probabilities (PP) > 0.79 are shown at the nodes and the scale bar represents the expected changes per site. Thickened branches represent PP = 1 and the most basal branch was halved to facilitate layout. Families and the order Hypocreales are indicated with coloured blocks to the right of the tree. GenBank accession (superscript) and / or culture collection / voucher numbers are indicated for all species. The tree was rooted to Ramularia endophylla (culture CBS 113265, GenBank AY490776.2) and the species treated in this study for which LSU sequence data were available are indicated in bold face.

Sordariomycetes (other orders) LSU phylogeny (Fig. 5, parts 1–2): The alignment contained 89 isolates and the tree was rooted to Ramularia endophylla (strain CBS 113265, GenBank AY490776.1). The final alignment contained a total of 796 characters used for the phylogenetic analyses, including alignment gaps. The alignment contained a total of 340 unique site patterns. Based on the results of MrModelTest, dirichlet base frequencies and the GTR+I+G model was used for the Bayesian analysis. The Bayesian analyses generated 878 002 trees (saved every five generations) from which 658 502 were sampled after 25 % of the trees were discarded as burn-in.

Fig. 5, parts 1–2.

Fig. 5, parts 1–2.

Fig. 5, parts 1–2.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Sordariomycetes (other orders) LSU nucleotide alignment. Bayesian posterior probabilities (PP) > 0.79 are shown at the nodes and the scale bar represents the expected changes per site. Thickened branches represent PP = 1. Families and orders are indicated with coloured blocks to the right of the tree. GenBank accession (superscript) and / or culture collection / voucher numbers are indicated for all species. The tree was rooted to Ramularia endophylla (culture CBS 113265, GenBank AY490776.2) and the species treated in this study for which LSU sequence data were available are indicated in bold face.

Sordariomycetes (Amphisphaeriales) LSU phylogeny (Fig. 6): The alignment contained 54 isolates and the tree was rooted to Ramularia endophylla (strain CBS 113265, GenBank AY490776.1). The final alignment contained a total of 754 characters used for the phylogenetic analyses, including alignment gaps. The alignment contained a total of 194 unique site patterns. Based on the results of MrModelTest, dirichlet base frequencies and the GTR+I+G model was used for the Bayesian analysis. The Bayesian analyses generated 228 002 trees (saved every five generations) from which 171 002 were sampled after 25 % of the trees were discarded as burn-in.

Fig. 6.

Fig. 6.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Sordariomycetes (Amphisphaeriales) LSU nucleotide alignment. Bayesian posterior probabilities (PP) > 0.79 are shown at the nodes and the scale bar represents the expected changes per site. Thickened branches represent PP = 1 and the most basal branch was halved to facilitate layout. Families and the order Amphisphaeriales are indicated with coloured blocks to the right of the tree. GenBank accession (superscript) and / or culture collection / voucher numbers are indicated for all species. The tree was rooted to Ramularia endophylla (culture CBS 113265, GenBank AY490776.2) and the species treated in this study for which LSU sequence data were available are indicated in bold face.

Species phylogenies: Specific phylogenetic analyses were run for selected species and the resulting phylogenies are discussed in the species notes where applicable. Statistics associated with those phylogenies are provided in the figure legends. The optimal identity thresholds to discriminate filamentous fungal species followed Vu et al. (2019), with secondary DNA barcodes generated where necessary (Stielow et al. 2015).

Taxonomy

Alternaria chartarum Preuss, Bot. Zeitung 6: 412. 1848. Fig. 7.

Fig. 7.

Fig. 7.

Alternaria chartarum (CPC 38971). Conidiophores giving rise to chains of conidia. Scale bars = 10 μm.

Description and illustration: Woudenberg et al. (2013).

Material examined: Namibia, Walvis Bay, from hypolithic biomass under a rock, 19 Nov. 2019, P.W. Crous, HPC 3101, culture CPC 38971.

Notes: Alternaria chartarum, which was isolated from hypolithic biomass under a rock in the Namib Desert, has previously been associated with cutaneous alternariosis in humans, and is also commonly found in indoor environments (Magina et al. 2000, Samson et al. 2019).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Alternaria alternata (strain F13, GenBank HQ380767.1; Identities = 565/565 (100 %), no gaps), Alternaria chartarum (strain AC85, GenBank LC440618.1; Identities = 560/560 (100 %), no gaps), and Alternaria aspera (strain CBS 115269, GenBank MH862983.1; Identities = 568/569 (99 %), one gap (0 %)). Closest hits using the LSU sequence are Alternaria multiformis (strain CBS 102060, GenBank NG_069860.1; Identities = 809/809 (100 %), no gaps), Alternaria terricola (strain CBS 202.67, GenBank NG_069728.1; Identities = 809/809 (100 %), no gaps), and Alternaria chartarum (strain CBS 200.67, GenBank NG_069727.1; Identities = 809/809 (100 %), no gaps) – also see Fig. 1. Closest hits using the actA sequence had highest similarity to Alternaria chartarum (strain ATCC 18044, GenBank JQ671654.1; Identities = 588/589 (99 %), no gaps), Alternaria sp. 3 MG-2016 (strain MF-G013021, GenBank KU639836.1; Identities = 583/584 (99 %), no gaps), Alternaria septospora (strain CBS 109.38, GenBank JQ671655.1; Identities = 587/589 (99 %), no gaps), and Alternaria poonensis (strain EGS 47-138, GenBank JQ671749.1; Identities = 501/510 (98 %), no gaps). Closest hits using the cmdA sequence had highest similarity to Alternaria sp. 3 MG-2016 (strain MF-G242021, GenBank KU639878.1; Identities = 525/525 (100 %), no gaps), Alternaria sp. 1 MG-2016 (strain MF-G333011, GenBank KU639872.1; Identities = 525/525 (100 %), no gaps), Alternaria chartarum (strain ATCC 18044, GenBank JQ646162.1; Identities = 524/525 (99 %), no gaps), Alternaria consortialis (strain CBS 201.67, GenBank JQ646173.1; Identities = 506/527 (96 %), two gaps (0 %)), and Alternaria botrytis (strain AKC603, GenBank MT770809.1; Identities = 504/525 (96 %), no gaps). Closest hits using the gapdh sequence had highest similarity to Alternaria chartarum (strain AC85, GenBank LC482041.1; Identities = 408/409 (99 %), no gaps), Alternaria aspera (strain CBS 115269, GenBank KC584166.1; Identities = 408/409 (99 %), no gaps), and Alternaria concatenata (as Ulocladium capsicum; strain HSAUP_XF030035, GenBank AY762950.1; Identities = 408/409 (99 %), no gaps). Closest hits using the tef1 sequence had highest similarity to Alternaria aspera (strain CBS 115269, GenBank KC584734.1; Identities = 240/242 (99 %), no gaps), Alternaria alternata (strain DUCC5016, GenBank KJ638247.1; Identities = 292/311 (94 %), two gaps (0 %)), and Alternaria japonica (strain P400, GenBank AY375367.1; Identities = 366/395 (93 %), four gaps (1 %)). Closest hits using the tub2 sequence had highest similarity to Alternaria chartarum (strain ATCC 18044, GenBank JQ671994.1; Identities = 337/338 (99 %), no gaps), Alternaria septospora (strain CBS 109.38, GenBank JQ671995.1; Identities = 336/338 (99 %), no gaps), and Alternaria atra (strain ATCC 18040, GenBank JQ671998.1; Identities = 331/338 (98 %), no gaps).

Authors: P.W. Crous, D.A. Cowan, G. Maggs-Kölling, E. Marais, N. Yilmaz & M.J. Wingfield

Alternaria heterospora Woudenb. & Crous, Stud. Mycol. 75: 204. 2013. Fig. 8.

Fig. 8.

Fig. 8.

Alternaria heterospora (CPC 38969). Conidiophores giving rise to conidia. Scale bars = 10 μm.

Description and illustration: Woudenberg et al. (2013).

Material examined: Namibia, Walvis Bay, from hypolithic biomass under a rock, 19 Nov. 2019, P.W. Crous, HPC 3101, culture CPC 38969.

Notes: Alternaria heterospora (as Ulocladium solani) was described as a new species associated with leaf spots on Lycopersicon esculentum and Duchesnea indica from Hunan Province in China (Wang et al. 2009). In the present study this species is reported from hypolithic biomass under a rock in the Namib Desert.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Alternaria atra (strain AC90, GenBank LC440624.1; Identities = 563/563 (100 %), no gaps), Alternaria dauci (strain AY853, GenBank MG250469.1; Identities = 563/563 (100 %), no gaps), and Alternaria heterospora (strain CBS 123376, GenBank MH863292.1; Identities = 560/560 (100 %), no gaps). Closest hits using the LSU sequence are Alternaria multiformis (strain CBS 102060, GenBank NG_069860.1; Identities = 787/787 (100 %), no gaps), Alternaria terricola (strain CBS 202.67, GenBank NG_069728.1; Identities = 787/787 (100 %), no gaps), and Alternaria chartarum (strain CBS 200.67, GenBank NG_069727.1; Identities = 787/787 (100 %), no gaps) – also see Fig. 1. Closest hits using the actA sequence had highest similarity to Alternaria atra (strain ATCC 18040, GenBank JQ671660.1; Identities = 591/591 (100 %), no gaps), Alternaria heterospora (strain MF-G316021, GenBank KU639828.1; Identities = 588/588 (100 %), no gaps), and Alternaria cucurbitae (strain EGS 31-021, GenBank JQ671663.1; Identities = 590/591 (99 %), no gaps). Closest hits using the cmdA sequence had highest similarity to Alternaria heterospora (strain MF-G316021, GenBank KU639868.1; Identities = 549/549 (100 %), no gaps), Alternaria botrytis (strain AKC603, GenBank MT770809.1; Identities = 548/549 (99 %), no gaps), and Alternaria obovoidea (strain CBS 101229, GenBank JQ646172.1; Identities = 548/549 (99 %), no gaps). Closest hits using the gapdh sequence had highest similarity to Alternaria heterospora (strain CBS 123376, GenBank KC584176.1; Identities = 460/460 (100 %), no gaps), Alternaria subcucurbitae (as Ulocladium subcucurbitae; strain CBS 121491, GenBank EU855803.1; Identities = 460/460 (100 %), no gaps), and Alternaria cucurbitae (as Ulocladium cucurbitae; strain HSAUP_XF030282, GenBank AY762951.1; Identities = 460/460 (100 %), no gaps). Closest hits using the tef1 sequence had highest similarity to Alternaria atra (strain IR_Ker955, GenBank MK188501.1; Identities = 221/224 (99 %), no gaps), Alternaria alternata (strain MOS635, GenBank KP009004.1; Identities = 242/255 (95 %), no gaps), and Alternaria photistica (strain CBS 212.86, GenBank FJ214950.1; Identities = 252/267 (94 %), four gaps (1 %)). Closest hits using the tub2 sequence had highest similarity to Alternaria atra (strain ATCC 18040, GenBank JQ671998.1; Identities = 335/336 (99 %), no gaps), Alternaria multiformis (strain CBS 102060, GenBank JQ672002.1; Identities = 334/336 (99 %), no gaps), and Alternaria cucurbitae (strain EGS 31-021, GenBank JQ672001.1; Identities = 333/336 (99 %), no gaps).

Authors: P.W. Crous, D.A. Cowan, G. Maggs-Kölling, E. Marais, N. Yilmaz & M.J. Wingfield

Apiospora sasae Crous & R.K. Schumach., sp. nov. MycoBank MB 839279. Fig. 9.

Fig. 9.

Fig. 9.

Apiospora sasae (CPC 38165). A. Sporulating colony on SNA. B. Conidiogenous cell and conidia. C, E. Conidiogenous cells with both asexual morphs. D. Conidiogenous cells with conidia. F. Conidiogenous cell with conidia of synasexual morph. G–H. Conidia. Scale bars = 10 μm.

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

Occurring on dead culms of Sasa vetchii. Sporodochia single to gregarious, initially immersed, opening via longitudinal split of epidermis, revealing black conidial mass, dry, pulvinate, 1–2 × 0.4 mm. Paraphyses absent. Conidiophores subcylindrical, rarely branched, basal and apical cell enlarged, subhyaline, thin-walled, smooth, septa red-brown, thick-walled, secession apical and lateral, lacking collarettes. Conidiogenous cells discrete, subcylindrical, subhyaline to pale brown, smooth to finely verruculose, holoblastic, proliferating sympodially, 5–10 × 3–4 μm. Sterile cells replacing normal conidia, brown, smooth, irregularly lobed, 8–25 × 7–12 μm. Conidia numerous, aseptate, subglobose, polygonal (6–7) to urceolate (urniform), red-brown at maturity, thick-walled, smooth, often with large central guttulate or multi-guttulate, with a lateral hyaline equatorial germ slit over entire length, often with a lateral, small protruding hilum, (16–)17–18(–20) × (15–)16–17(–19) μm. Synasexual morph hyaline, smooth, erect, solitary, with conidiophores subcylindrical, 1–2-septate, 10–20 × 3–4 μm. Conidiogenous cells terminal, integrated, hyaline, smooth, subcylindrical, 10–15 × 2.5–3 μm, proliferating sympodially at apex with subdenticulate loci, unthickened nor darkened. Conidia solitary, hyaline, aseptate, smooth, fusoid, falcate, apex subobtuse, base truncate, 10–20 × 2–2.5 μm.

Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA surface smoke grey, reverse olivaceous grey; on PDA surface and reverse olivaceous grey; on OA surface olivaceous grey.

Typus: Netherlands, Noord-Holland Province, Hoofddorp, on dead culms of Sasa veitchii (Poaceae), 20 May 2019, L. van der Linde, HPC 2959 = RKS 1101 (holotype CBS H-24403, culture ex-type CPC 38165 = CBS 146808).

Notes: The genus Arthrinium includes plant pathogens, endophytes and saprobes with a wide host range and geographic distribution (Crous & Groenewald 2013, Wang et al. 2018, Pintos et al. 2019). Although Arthrinium was seen as the asexual morph of Apiospora (Crous & Groenewald 2013), the recent epitypification of Arthrinium (based on A. caricicola; Crous et al. 2020b) showed this complex to represent two clades, one corresponding to Arthrinium, and the other to Apiospora. This generic complex was recently revised by Pintos et al. (2021), and will not be treated further here. The two taxa collected in the present study are thus best accommodated in Apiospora. Apiospora sasae is related to, but morphologically distinct from Arthrinium yunnanum (now Apiospora yunnana; conidia 10–16 μm diam; Dai et al. 2017) and Arthrinium esporlense (now Apiospora esporlensis; conidia 8–13 μm long; Pintos et al. 2019). A phylogenetic species tree is presented as Fig. 10.

Fig. 10.

Fig. 10.

Consensus phylogram (50 % majority rule) obtained from the maximum likelihood analysis with IQ-TREE of the Apiospora multigene (ITS / LSU / tef1 / tub2) nucleotide alignment of the two novel Apiospora species treated in this study. Some branches were halved to facilitate layout. Bootstrap support values (> 79 %) from 5 000 ultrafast bootstrap replicates are shown at the nodes. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Beltrania rhombica (culture CBS 123.58) and the species treated in this study are highlighted with coloured blocks and bold face. Alignment statistics: 96 strains including the outgroup; 4 171 characters including alignment gaps analysed: 2 235 distinct patterns, 1 379 parsimony-informative, 365 singleton sites, 2 427 constant sites. The best models identified in IQ-TREE were: TNe+I+G4 (ITS), TIM+F+I+G4 (LSU), HKY+F+I+G4 (tef1 and tub2).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Arthrinium yunnanum (now Apiospora yunnana; strain JN2, GenBank MH191120.1; Identities = 646/648 (99 %), no gaps), Arthrinium hysterinum (now Apiospora hysterina; voucher AP15318, GenBank MK014873.1; Identities = 602/605 (99 %), no gaps), and Arthrinium esporlense (now Apiospora esporlensis; strain 18TJAM004, GenBank MT856406.1; Identities = 424/446 (95 %), six gaps (1 %)). Closest hits using the LSU sequence are Apiospora setosa (voucher ICMP 6888, GenBank DQ810214.1; Identities = 826/826 (100 %), no gaps), Arthrinium hysterinum (now Apiospora hysterina; voucher AP15318, GenBank MK014840.1; Identities = 817/817(100 %), no gaps), and Apiospora tintinnabula (voucher ICMP 6889-96, GenBank DQ810217.1; Identities = 810/812 (99 %), two gaps (0 %)) – also see Fig. 6. Closest hits using the rpb2 sequence had highest similarity to Apiospora bambusae (strain ICMP 6889, GenBank DQ368649.1; Identities = 706/710 (99 %), no gaps), Arthrinium yunnanum (now Apiospora yunnana; voucher MFLU 18-1219, GenBank MK313857.1; Identities = 705/710 (99 %), no gaps), and Apiospora tintinnabula (voucher ICMP 7019-96), GenBank DQ810235.1; Identities = 705/710 (99 %), no gaps). Closest hits using the tef1 (second part) sequence had highest similarity to Arthrinium yunnanum (now Apiospora yunnana; voucher MFLU 18-1219, GenBank MK193869.1; Identities = 906/908 (99 %), no gaps), Apiospora locuta-pollinis (as Arthrinium FL-2018a; strain LC11688, GenBank MF939618.1; Identities = 856/880 (97 %), two gaps (0 %)), and Arthrinium pseudoparenchymaticum (now Apiospora pseudoparenchymatica; strain SICAUCC 18-0008, GenBank MK359205.1; Identities = 865/909 (95 %), two gaps (0 %)). Closest hits using the tub2 sequence had highest similarity to Arthrinium hysterinum (now Apiospora hysterina; voucher AP29717, GenBank MK017981.1; Identities = 387/390 (99 %), no gaps), Arthrinium yunnanum (now Apiospora yunnana; strain MFLUCC 18-1102, GenBank MK291950.1; Identities = 762/774 (98 %), three gaps (0 %)), and Arthrinium ovatum (now Apiospora ovata; strain CBS 115042, GenBank KF144995.1; Identities = 697/765 (91 %), 23 gaps (3 %)).

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Apiospora stipae Crous & R.K. Schumach., sp. nov. MycoBank MB 839280. Fig. 11.

Fig. 11.

Fig. 11.

Apiospora stipae (CPC 38101). A. Hyphae giving rise to conidiogenous cells with conidia. B–C. Conidiogenous cells and conidia. Scale bars = 10 μm.

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

Sporodochia on dead tissue, immersed, becoming erumpent, semi-superficial, single to confluent, cushion-shaped, black, dry, pulvinate. Paraphyses absent. Conidiophores cylindrical, unbranched to simple branched, hyaline, thin- and smooth-walled, septa red-brown, thick-walled, secession lateral and apical, single, lacking collarettes. Conidia numerous, aseptate, lens-shaped in side view, round to polygonal (6–8) in top view, olive brown to red-brown, thick-walled, smooth, eguttulate to guttulate, with lateral germ slit over entire length of conidium, often with a lateral, small, hyaline pronounced hilum, examined in water, 6.5–10.5 (length) × 6–9 (width) × 5–6 (side view) μm. Sporulating on PDA. Mycelium consisting of smooth, hyaline, branched, septate, 2–4 μm diam hyphae. Conidiophores reduced to conidiogenous cells. Conidiogenous cells aggregated in clusters on hyphae, pale brown, smooth, ampulliform, 5–8 × 3–4 μm. Conidia in vitro (in lactic acid) brown, aseptate, smooth, granular, globose to elongated ellipsoid in surface view, (6.5–)7–8 μm diam, lenticular in side view with pale equatorial germ slit, (4.5–)5–6 μm in side view, with basal scar 1 μm diam; brown elongated cells (sterile cells?) at times intermingled among conidia.

Ascomata on host tissue (link with asexual morph unconfirmed): perithecial, on dead tissue, immersed, single, gregarious, globose, black; peridium multi-layered, consisting of textura prismatica-epidermoidea with thick-walled, smooth cells; inner layers hyaline, outer layers red-brown. Paraphyses present. Asci 8-spored, clavate, inoperculate, pedicel short and furcate, 87–101 × 17 μm. Ascospores 1-septate, hyaline, 27–30 × 8–10 μm.

Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse isabelline.

Typus: Spain, Pontevedra, O Grove, on dead culm of Stipa gigantea (Poaceae), 10 Apr. 2019, M.A. Delgado, HPC 2973 = RKS 253 (holotype CBS H-24398, culture ex-type CPC 38101 = CBS 146804).

Notes: Numerous Arthrinium spp. occurring on diverse hosts in Europe have recently been treated (Crous & Groenewald 2013, Pintos et al. 2019). Apiospora stipae represents a new species closely related to Arthrinium esporlense (now Apiospora esporlensis; conidia (8–)9–12(–13) μm long; Pintos et al. 2019), from which it is morphologically distinct. A phylogenetic species tree is presented as Fig. 10.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to “Arthrinium” sp. (strain 4936, GenBank FR667986.1; Identities = 490/490 (100 %), no gaps), Arthrinium esporlense (now Apiospora esporlensis; strain 18TJAM004, GenBank MT856406.1; Identities = 543/560 (97 %), one gap (0 %)), and Arthrinium qinlingense (as Arthrinium sp. NJ-2018a, now Apiospora qinlingensis; strain JN5, GenBank MH197120.1; Identities = 553/571 (97 %), one gap (0 %)). Closest hits using the LSU sequence are Arthrinium esporlense (now Apiospora esporlensis; voucher AP16717, GenBank MK014845.1; Identities = 836/837 (99 %), no gaps), Arthrinium phragmites (now Apiospora phragmitis; strain CPC 18900, GenBank NG_042783.1; Identities = 896/900 (99 %), no gaps), and Arthrinium kogelbergense (now Apiospora kogelbergensis; strain CBS 113335, GenBank KF144939.1; Identities = 895/900 (99 %), no gaps) – also see Fig. 6. Closest hits using the rpb2 sequence had highest similarity to Apiospora bambusae (strain ICMP 6889, GenBank DQ368649.1; Identities = 777/863 (90 %), no gaps), Apiospora tintinnabula (voucher ICMP 7019-96), GenBank DQ810235.1; Identities = 775/862 (90 %), no gaps), and Arthrinium yunnanum (now Apiospora yunnana; voucher MFLU 18-1219, GenBank MK313857.1; Identities = 773/860 (90 %), no gaps). Closest hits using the tef1 sequence (first part) had highest similarity to Arthrinium italicum (now Apiospora italica; voucher AP221017, GenBank MK017956.1; Identities = 327/358 (91 %), six gaps (1 %)), Arthrinium malaysianum (now Apiospora malaysiana; strain CBS 102053, GenBank KF145030.1; Identities = 343/377 (91 %), four gaps (1 %)), and Arthrinium thailandicum (now Apiospora thailandica; strain LC5630, GenBank KY705113.1; Identities = 323/356 (91 %), four gaps (1 %)). Closest hits using the tef1 (second part) sequence had highest similarity to Arthrinium phyllostachium (as Arthrinium sp., now Apiospora phyllostachydis; voucher MFLU 18-2333, GenBank MK313853.1; Identities = 893/919 (97 %), one gap (0 %)), Arthrinium yunnanum (now Apiospora yunnana; voucher MFLU 18-1219, GenBank MK193869.1; Identities = 884/920 (96 %), three gaps (0 %)), and Nigrospora globosa (as Nigrospora sp. ZZ-2018a; strain LC12441, GenBank MK336057.1; Identities = 858/907 (95 %), four gaps (0 %)). Closest hits using the tub2 sequence had highest similarity to Arthrinium kogelbergense (now Apiospora kogelbergensis; strain CBS 117206, GenBank KF144987.1; Identities = 689/776 (89 %), 30 gaps (3 %)), Arthrinium arundinis (now Apiospora arundinis; strain DSM 3112, GenBank AB220307.1; Identities = 687/781 (88 %), 27 gaps (3 %)), and Arthrinium sacchari (now Apiospora sacchari; strain ATCC 76289, GenBank AB220286.1; Identities = 684/781 (88 %), 27 gaps (3 %)).

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Bagadiella eucalyptorum Crous & Carnegie, sp. nov. MycoBank MB 839281. Fig. 12.

Fig. 12.

Fig. 12.

Bagadiella eucalyptorum (CPC 39299). A. Colony sporulating on OA. B–D. Developing conidiogenous cells. E. Conidia. Scale bars = 10 μm.

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

Mycelium consisting of subhyaline, smooth, branched, septate, 2–3 μm diam hyphae. Conidiomata 80–100 μm diam, sporodochial, arising from an aggregated subhyaline mass of hyphal cells, that give rise to a hyaline, mucoid conidial mass. Conidiophores integrated, arising from a stroma or individual hyphae, smooth, branched or not, with terminal and intercalary conidiogenous cells, monophialidic, subcylindrical to lageniform, 10–15 × 2.5–3 μm, with minute collarettes. Conidia borne in mucoid heads, lunate, aseptate, curved, apex obtuse, base truncate, hyaline, (16–)17–18(–19) × 1.5(–2) μm.

Culture characteristics: Colonies flat, spreading, with sparse to moderate aerial mycelium and lobate, even margin, reaching 25 mm diam after 2 wk at 25 °C. On MEA surface saffron, reverse saffron to ochreous; on PDA surface saffron, reverse ochreous; on OA surface saffron.

Typus: Australia, New South Wales, Belanglo State Forest, Berrima, on leaves of Eucalyptus sp. (Myrtaceae), Aug. 2015, A.J. Carnegie, HPC 3219 (holotype CBS H-24560, culture ex-type CPC 39299 = CBS 147177).

Notes: Species of Bagadiella are commonly isolated as endophytic fungi from eucalypt leaves, but can also be associated with pale yellow leaf blotches (Cheewangkoon et al. 2009, Crous et al. 2019c). Bagadiella eucalyptorum is closely related to B. eucalypti [conidia (12–)14–17(–21) × (1.5–)2 μm; Crous et al. 2017] and B. lunata [conidia (15–)16–18(–22) × (1.3–)1.5(–1.7); Cheewangkoon et al. 2009]. Morphologically they are very similar, and best distinguished based on DNA sequence data. Also see the phylogenetic species tree (Fig. 13).

Fig. 13.

Fig. 13.

The first of 519 equally most parsimonious trees obtained from a phylogenetic analysis of the Bagadiella ITS nucleotide alignment. The tree was rooted to Apiospora ovata (strain CBS 115042, GenBank NR_121558.1) and the scale bar indicates the number of changes. Parsimony bootstrap support values higher than 79 % are shown at the nodes and the treated species is highlighted with a coloured box and bold text. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. Branches present in the strict consensus tree are thickened. Alignment statistics: 21 strains including the outgroup; 544 characters including alignment gaps analysed: 348 constant, 71 variable and parsimony-uninformative and 125 parsimony-informative. Tree statistics: TL = 401, CI = 0.783, RI = 0.756, RC = 0.592.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Bagadiella lunata (strain CBS 124762, GenBank NR_132832.1; Identities = 587/602 (98 %), five gaps (0 %)), Bagadiella koalae (strain CBS 129523, GenBank NR_159638.1; Identities = 558/574 (97 %), four gaps (0 %)), and Bagadiella victoriae (strain CPC 20195, GenBank MN161894.1; Identities = 585/602 (97 %), one gap (0 %)). Closest hits using the LSU sequence are Bagadiella eucalypti (strain CBS 143497, GenBank MN162174.1; Identities = 842/846 (99 %), two gaps (0 %)), Bagadiella koalae (strain CBS 129523, GenBank NG_070001.1; Identities = 826/831 (99 %), no gaps), and Bagadiella lunata (strain CBS 124762, GenBank NG_058637.1; Identities = 852/858 (99 %), two gaps (0 %)) – also see Fig. 6.

Authors: P.W. Crous, J.Z. Groenewald & A.J. Carnegie

Blastacervulus metrosideri P.R. Johnst., Fungal Syst. Evol. 3: 166. 2019. Fig. 14.

Fig. 14.

Fig. 14.

Blastacervulus metrosideri (CPC 38759). A–E. Conidiogenous cells giving rise to conidia. Scale bars = 10 μm.

Description and illustration: See Johnston & Park (2019).

Material examined: New Zealand, Tauranga Port, on leaves of Metrosideros sp. (Myrtaceae), 22 Aug. 2019, L. Rabbidge, specimen CBS H-24511, culture T19_05741C = CPC 38759 = CBS 147006.

Notes: Blastacervulus metrosideri was described by Johnston & Park (2019) as a new foliar pathogen on Metrosideros excelsa in New Zealand, causing round, red-brown to black leaf spots that mostly occur on the upper leaf surface. The ITS sequence of CPC 38759 is a perfect match to that of the ex-type culture (ICMP 21883; GenBank NR_169959.1; 473/473 (100 %), no gaps) while there were three gaps present compared to the LSU sequence (GenBank NG_068290.1; 1 005/1 008 (99 %), three gaps (0 %)) – also see Fig. 1.

Authors: P.W. Crous, J.Z. Groenewald & R. Thangavel

Calonectria singaporensis Crous & Decock, sp. nov. MycoBank MB 839282. Fig. 15.

Fig. 15.

Fig. 15.

Calonectria singaporensis (CBS 146715). A. Conidiophores on PNA. B–C. Conidiogenous apparatus. D. Chlamydospores. E–G. Conidiophores with stipe extensions and terminal vesicles. H. Conidia. Scale bars = 10 μm.

Etymology: Name refers to Singapore, the country where this species was isolated.

Macroconidiophores comprised of a stipe, a penicillate arrangement of fertile branches, and a stipe extension ending in a terminal vesicle. Stipe septate, hyaline to pale brown at base, smooth, 50–350 μm tall. Conidiogenous apparatus with primary branches aseptate or 1-septate, 20–30 × 6–8 μm; secondary branches aseptate, 16–20 × 4–5 μm, and tertiary and additional (–6) branches aseptate, 12–15 × 4–5 μm, each terminal branch producing 2–4 phialides; phialides doliiform to reniform, hyaline, aseptate, 9–15 × 3–5 μm, apex with minute periclinal thickening and inconspicuous collarette; stipe extensions septate, straight to flexuous, 90–200 μm long, 3–4 μm wide at apical septum, terminating in sphaeropedunculate vesicle, (4–)7–12 μm diam; lateral stipe extensions common, up to 70 μm long, terminating in small sphaeropedunculate vesicles, 4–6 μm diam. Conidia cylindrical, rounded at both ends, straight, (33–)37–40(–41) × (3.5–)4(–4.5) μm, 1-septate, lacking a visible abscission scar, held in cylindrical clusters by colourless slime. Chlamydospores dark brown, thickened, globose, 15–30 μm diam, formed in chains throughout the medium, and aggregated to form microsclerotia.

Culture characteristics: Colonies flat, spreading, with sparse to moderate aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA surface sienna to umber.

Typus. Singapore, Mac Ritchie Reservoir, South East Asian rainforest, submerged leaf litter in a small stream, Dec. 2001, C. Decock # SING365 (holotype CBS H-24749, culture ex-type MUCL 048320 = CBS 146715).

Additional materials examined: Singapore, Mac Ritchie Reservoir, South East Asian rainforest, submerged leaf litter in a small stream, Dec. 2001, C. Decock # SING365, cultures CBS 146712 = MUCL 048012, CBS 146713 = MUCL 048171, CBS 146714 = MUCL 048187.

Notes: A recent revision of the genus Calonectria accepted 11 species complexes, reducing the genus to 120 species (Liu et al. 2020). Calonectria singaporensis is phylogenetically distinct from all accepted species in the genus, representing a new species in the C. kyotensis species complex (Fig. 16). Morphologically, these taxa are all very similar, sharing sphaeropedunculate vesicles and 1-septate conidia, and are best distinguished based on their DNA sequence data.

Fig. 16.

Fig. 16.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Calonectria multigene (actA / cmdA / his3 / ITS / LSU / rpb2 / tef1 / tub2) nucleotide alignment. The alignment is derived from the combined alignment of Liu et al. (2020) and GenBank accession numbers can be obtained from the same reference. Culture numbers with a type status are in bold face and the novel species is highlighted with a coloured box and bold text. Bayesian posterior probabilities (> 0.79) are shown at the nodes and the scale bar represents the expected changes per site. The tree was rooted to Curvicladiella cignea (culture CBS 109167). Alignment statistics: 52 strains including the outgroup; 112 / 290 / 212 / 57 / 37 / 255 / 241 / 307 unique site patterns, respectively. Tree statistics: 5 928 sampled trees from 395 000 generations.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Calonectria syzygiicola (strain CBS 112827, GenBank KY653281.1; Identities = 530/531 (99 %), no gaps), Calonectria ilicicola (strain CPC 16334, GenBank GU057378.1; Identities = 530/531 (99 %), no gaps), and Calonectria curvispora (strain CBS 116159, GenBank GQ280568.1; Identities = 530/531 (99 %), no gaps). Closest hits using the LSU sequence are Calonectria malesiana (strain CBS 112752, GenBank MH874470.1; Identities = 861/865 (99 %), no gaps), Calonectria ilicicola (strain CBS 125938, GenBank MH875288.1; Identities = 858/862 (99 %), no gaps), and Calonectria yunnanensis (strain CERC 5337 (R), GenBank MT359618.1; Identities = 853/857 (99 %), no gaps) – also see Fig. 4. Closest hits using the actA sequence had highest similarity to Calonectria turangicola (now Calonectria kyotensis; strain CMW 35410 (R), GenBank MT335152.1; Identities = 228/246 (93 %), five gaps (2 %)), Calonectria pseudoturangicola (now Calonectria kyotensis; strain CERC 7127 (R), GenBank MT335122.1; Identities = 228/246 (93 %), five gaps (2 %)), and Calonectria kyotensis (strain CBS 114525 (R), GenBank MT335039.1; Identities = 228/246 (93 %), five gaps (2 %)). Closest hits using the cmdA sequence had highest similarity to Calonectria malesiana (strain CMW 23687 (R), GenBank MT335286.1; Identities = 646/680 (95 %), two gaps (0 %)), Calonectria turangicola (now Calonectria kyotensis; strain CMW 35410 (R), GenBank MT335389.1; Identities = 645/680 (95 %), two gaps (0 %)), and Calonectria pseudoturangicola (now Calonectria kyotensis; strain CERC 7127 (R), GenBank MT335357.1; Identities = 645/680 (95 %), two gaps (0 %)). Closest hits using the his3 sequence had highest similarity to Calonectria matogrossensis (strain GFP018, GenBank MH837652.1; Identities = 366/412 (89 %), 20 gaps (4 %)), Calonectria pseudometrosideri (strain CBS 134843, GenBank KM396081.1; Identities = 366/412 (89 %), 20 gaps (4 %)), and Calonectria metrosideri (strain CBS 133603, GenBank KC294307.1; Identities = 366/412 (89 %), 20 gaps (4 %)). Closest hits using the tef1 sequence had highest similarity to Calonectria malesiana (strain CSF11261, GenBank MT412818.1; Identities = 459/490 (94 %), four gaps (0 %)), Calonectria montana (now Calonectria canadiana; strain HSP65, GenBank MN356469.1; Identities = 470/502 (94 %), eight gaps (1 %)), and Calonectria lateralis (strain CMW 47414, GenBank MH119245.1; Identities = 467/499 (94 %), two gaps (0 %)). Closest hits using the tub2 sequence had highest similarity to Calonectria pseudoturangicola (now Calonectria kyotensis; strain CERC 7131, GenBank MF443082.1; Identities = 294/320 (92 %), seven gaps (2 %)), Calonectria turangicola (now Calonectria kyotensis; strain CSF11435, GenBank MT413127.1; Identities = 294/320 (92 %), seven gaps (2 %)), and Calonectria kyotensis (strain CSF16440, GenBank MT413019.1; Identities = 294/320 (92 %), seven gaps (2 %)).

Authors: P.W. Crous, J.Z. Groenewald & C. Decock

Castanediella neomalaysiana Crous, sp. nov. MycoBank MB 839283. Fig. 17.

Fig. 17.

Fig. 17.

Castanediella neomalaysiana (CPC 39275). A. Colony sporulating on PDA. B–C. Conidiogenous cells giving rise to conidia. D. Conidia. Scale bars = 10 μm.

Etymology: Name refers to the fact that it is similar to Castanediella malaysiana.

Mycelium consisting of smooth, brown, septate, branched, 2–3 μm diam hyphae. Conidiophores arising directly from superficial hyphae, subcylindrical, erect, straight, branched below or not, 0–4-septate, commonly also reduced to conidiogenous cells, pale brown, smooth, 17–60 × 2.5–3.5 μm. Conidiogenous cells integrated, pale brown, smooth, subcylindrical, terminal and intercalary, apex swollen with cluster of aggregated subdenticulate conidiogenous loci, 0.5 μm tall and wide, 17–22 × 2.5–3 μm. Conidia solitary, aseptate, fusoid, curved, widest in middle, tapering to subobtuse apices, hyaline, smooth, guttulate, (15–)17–19(–21) × 2(–3) μm.

Culture characteristics: Colonies flat, spreading, with sparse to moderate aerial mycelium and smooth, lobate margin, reaching 50 mm diam after 2 wk at 25 °C, and brown, microsclerotium-like bodies immersed in agar, but these remain sterile. On MEA and PDA surface and reverse brown vinaceous; on OA surface dark brick.

Typus: Malaysia, on leaves of Eucalyptus sp. (Myrtaceae), Nov. 2010, M.J. Wingfield, HPC 3228 (holotype CBS H-24547, culture ex-type CPC 39275 = CBS 147093).

Notes: Castanediella was established by Crous et al. (2015) to accommodate several species of Amphisphaeriales, for which Hernández-Restrepo et al. (2017) introduced the family Castanediellaceae. Castanediella neomalaysiana is phylogenetically closely related to C. malaysiana (conidia 0–1-septate, 18–30 × 2–3 μm; Hernandez-Restrepo et al. 2016), but differs in having shorter conidia. A phylogenetic species tree is presented in Fig. 18.

Fig. 18.

Fig. 18.

The first of 11 equally most parsimonious trees obtained from a phylogenetic analysis of the Castanediella ITS sequence alignment. The tree was rooted to Beltrania pseudorhombica (strain CBS 138003, GenBank NR_148074.1) and the scale bar indicates the number of changes. Parsimony bootstrap support values higher than 79 % are shown at the nodes and the treated species is highlighted with a coloured box and bold text. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. Branches present in the strict consensus tree are thickened. Alignment statistics: 27 strains including the outgroup; 527 characters including alignment gaps analysed: 352 constant, 57 variable and parsimony-uninformative and 118 parsimony-informative. Tree statistics: TL = 430, CI = 0.644, RI = 0.775, RC = 0.499.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Castanediella sp. (strain JHGB10_4A, GenBank MH267851.1; Identities = 542/558 (97 %), two gaps (0 %)), Castanediella malaysiana (strain CPC 24918, GenBank NR_154810.1; Identities = 545/564 (97 %), two gaps (0 %)), and Castanediella couratarii (strain CBS 579.71, GenBank MH860269.1; Identities = 543/565 (96 %), four gaps (0 %)). Closest hits using the LSU sequence are Castanediella malaysiana (strain CPC 24918, GenBank NG_067312.1; Identities = 846/850 (99 %), two gaps (0 %)), Castanediella brevis (as Castanediella sp.; strain LCG 10-1, GenBank MH806358.1; Identities = 755/759 (99 %), one gap (0 %)), and Castanediella ramosa (as Idriella ramosa; strain MUCL 39857, GenBank KC775711.1; Identities = 814/819 (99 %), five gaps (0 %)) – also see Fig. 6.

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Colletotrichum kinghornii Damm et al., Stud. Mycol. 73: 73. 2012. Fig. 19.

Fig. 19.

Fig. 19.

Colletotrichum kinghornii (CPC 38766). A. Ascoma developing on PNA. B–D. Asci with ascospores. Scale bars: A = 250 μm, all others = 10 μm.

Description and illustration: Damm et al. (2012).

Material examined: New Zealand, Auckland, East Tamaki, on Phormium cookianum (Asphodelaceae), 22 Aug. 2019, C. Inglis, culture CPC 38766 = T19_05774B.

Notes: Colletotrichum kinghornii was described based on its asexual morph from Phormium tenax collected in Great Britain (Damm et al. 2012). The present collection is the first record of its sexual morph. A phylogenetic species tree is presented as Fig. 20.

Fig. 20.

Fig. 20.

Consensus phylogram (50 % majority rule) obtained from the maximum likelihood analysis with IQ-TREE of the Colletotrichum multigene (ITS / gapdh / chs-1 / actA / tub2) nucleotide alignment. Bootstrap support values (> 79 %) from 5 000 ultrafast bootstrap replicates are shown at the nodes. The alignment is derived from the combined alignment of Marin-Felix et al. (2017) and GenBank accession numbers can be obtained from the same reference. Culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Monilochaetes infuscans (culture CBS 869.96) and the species treated in this study are highlighted with coloured blocks and bold face. Alignment statistics: 58 strains including the outgroup; 1 972 characters including alignment gaps analysed: 973 distinct patterns, 596 parsimony-informative, 261 singleton sites, 1 115 constant sites. The best models identified in IQ-TREE were: TIM3e+I+G4 (ITS), K3P+I+G4 (gapdh), TNe+I+G4 (chs-1), HKY+F+G4 (actA), TIM2e+I+G4 (tub2).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Colletotrichum kinghornii (strain CBS 198.35, GenBank NR_111751.1; Identities = 538/538 (100 %), no gaps), Colletotrichum phormii (strain CBS 198.35, GenBank DQ286144.1; Identities = 571/571 (100 %), no gaps), and Colletotrichum salicis (strain CBS 129973, GenBank MH865705.1; Identities = 578/579 (99 %), no gaps). Closest hits using the LSU sequence are Colletotrichum kinghornii (strain CBS 198.35, GenBank NG_069631.1; Identities = 668/668 (100 %), no gaps), Colletotrichum arboricola (strain CBS 144795, GenBank NG_070064.1; Identities = 668/668 (100 %), no gaps), and Colletotrichum salicis (strain CBS 607.94, GenBank NG_070038.1; Identities = 668/668 (100 %), no gaps) – also see Fig. 5. Closest hits using the actA sequence had highest similarity to Colletotrichum kinghornii (strain CBS 198.35, GenBank JQ949775.1; Identities = 247/247 (100 %), no gaps), Colletotrichum kniphofiae (strain CBS 143496, GenBank MH107975.1; Identities = 622/641 (97 %), no gaps), and Colletotrichum destructivum (GenBank AY157843.1; Identities = 596/657 (91 %), 14 gaps (2 %)). Closest hits using the gapdh sequence had highest similarity to Colletotrichum kinghornii (strain CBS 198.35, GenBank JQ948785.1; Identities = 260/260 (100 %), no gaps), Colletotrichum scovillei (strain TJNH1, GenBank XM_035479122.1; Identities = 310/316 (98 %), one gap (0 %)), and Colletotrichum orchidophilum (strain IMI 309357, GenBank XM_022618742.1; Identities = 309/316 (98 %), one gap (0 %)). Closest hits using the tub2 sequence had highest similarity to Colletotrichum kinghornii (strain CBS 198.35, GenBank JQ950105.1; Identities = 485/486 (99 %), no gaps), Colletotrichum phormii (strain BRIP 62862, GenBank KX069820.1; Identities = 650/657 (99 %), two gaps (0 %)), and Colletotrichum rhombiforme (strain HYPG-1, GenBank KY581595.1; Identities = 668/683 (98 %), two gaps (0 %)).

Authors: P.W. Crous, J.Z. Groenewald & R. Thangavel

Colletotrichum pleopeltidis Crous & Jol. Roux, sp. nov. MycoBank MB 839284. Fig. 21.

Fig. 21.

Fig. 21.

Colletotrichum pleopeltidis (CPC 39342). A. Conidioma developing on PNA. B. Setae. C–D. Conidiogenous cells. E. Conidia. Scale bars = 10 μm.

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

Ascomatal initials develop on media (SNA, MEA, OA, PDA), but these remain sterile. Asexual morph on SNA. Vegetative hyphae 3–5 μm diam, hyaline to pale brown, smooth-walled, septate, branched. Conidiomata acervular, up to 350 μm diam, with conidiophores and setae forming on a cushion of brown, thick-walled cells. Setae dark brown, smooth-walled, 3–5-septate, 90–150 μm long, base conical to cylindrical, 5–6 μm diam, tip subacutely rounded. Conidiophores pale to medium brown, septate, smooth, branched, up to 60 μm long. Conidiogenous cells pale brown, smooth, short cylindrical to ampulliform, 17–25 × 4–6 μm, with periclinal thickening and non-flaring collarette. Conidia hyaline, smooth, aseptate, cylindrical, apex obtuse, base rounded with prominent scar, guttulate, (15–)19–23(–25) × (5–)5.5(–6) μm.

Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, lobate margin, covering dish after 2 wk at 25 °C. On MEA surface olivaceous grey, reverse olivaceous grey in centre, sienna in outer region; on PDA surface pale olivaceous grey, reverse olivaceous grey in centre, ochreous in outer region; on OA surface ochreous.

Typus: South Africa, Limpopo Province, Louis Trichardt, Hanglip Forest Reserve, on leaves of Pleopeltis sp. (Polypodiaceae), 17 Dec. 2015, J. Roux, HPC 3238 (holotype CBS H-24550, culture ex-type CPC 39342 = CBS 147082).

Notes: Colletotrichum pleopeltidis belongs to the destructivum complex (Damm et al. 2014) and is closely related to C. pisicola [conidia fusoid, curved, (11–)15–21(–29.5) × (3–)3.5–4 μm], and C. tanaceti [conidia cylindrical to somewhat clavate, curved, (13–)14.5–17.5(–19) × (3–)3.5–4(–4.5) μm]. It is distinguished based on conidium morphology and phylogeny (see Fig. 20).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Colletotrichum hsienjenchang (strain MAFF 243051, GenBank AB738855.1; Identities = 547/566 (97 %), five gaps (0 %)), Colletotrichum fuscum (strain DAOM216112, GenBank EU400144.1; Identities = 552/572 (97 %), 11 gaps (1 %)), and Colletotrichum spaethianum (strain AJ007, GenBank KT122848.1; Identities = 545/565 (96 %), five gaps (0 %)). Closest hits using the LSU sequence are Colletotrichum destructivum (strain 1212, GenBank KF181215.1; Identities = 874/878 (99 %), three gaps (0 %)), Colletotrichum fioriniae (strain CBS 126509, GenBank MH875593.1; Identities = 866/870 (99 %), three gaps (0 %)), and Colletotrichum tamarilloi (strain CBS 129954, GenBank MH877133.1; Identities = 877/884 (99 %), four gaps (0 %)) – also see Fig. 5. Closest hits using the actA sequence had highest similarity to Colletotrichum destructivum (GenBank AY157843.1; Identities = 594/639 (93 %), nine gaps (1 %)), Colletotrichum kniphofiae (strain CBS 143496, GenBank MH107975.1; Identities = 573/635 (90 %), 13 gaps (2 %)), and Colletotrichum kahawae (strain CIFC Que2, GenBank KU579251.1; Identities = 574/642 (89 %), 19 gaps (2 %)). Closest hits using the chs-1 sequence had highest similarity to Colletotrichum panacicola (strain YL2-2, GenBank MN894862.1; Identities = 267/283 (94 %), no gaps), Colletotrichum lineola (strain TYJ77301-7, GenBank MN894858.1; Identities = 267/283 (94 %), no gaps), and Colletotrichum neorubicola (strain CCR144, GenBank MK547526.1; Identities = 266/283 (94 %), no gaps).

Authors: P.W. Crous, J.Z. Groenewald & J. Roux

Coniochaeta deborreae Hern.-Restr. sp. nov. MycoBank MB 838706. Fig. 22.

Fig. 22.

Fig. 22.

Coniochaeta deborreae (CBS 147215). A–B. Colony on MEA. C–D. Colony on OA. E–F. Colony on PDA. G–J. Conidiogenous cells and conidia. K. Conidia. L–M. Chlamydospores. Scale bars G–M = 10 μm.

Etymology: Named after the collector, Kirsten de Borre, a student from the VISO Cor Mariae secondary school (Brakel, Belgium). This sample was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae septate, branched, hyaline to subhyaline, thin- and smooth-walled, 1–3 μm wide. Conidiophores reduced to conidiogenous cells. Conidiogenous cells terminal, lateral or intercalary, mono- or polyphialidic, cylindrical, ampulliform or ventricose, nearly globose, 4–11 × 2–3.5 μm, hyaline, with conspicuous periclinal thickening and distinguishable, usually cylindrical collarettes, 1–2 × 1–1.5 μm. Conidia aseptate, hyaline, smooth-walled, ellipsoidal to oblong, with rounded apex and truncate base, 2.5–4 × 1.5–2 μm. Chlamydospores solitary or in short chains, subglobose, pale brown, 6.5–13 × 5–9 μm.

Culture characteristics: Colonies after 2 wk at 25 °C on OA reaching 80 mm diam, glabrous, zonate centre vinaceous grey, olivaceous to fawn with saffron zones, margin effuse; reverse fawn to saffron. On MEA reaching 45 mm diam, velvety, zonate centre fulvous to saffron, margin irregular, lobate to fimbriate; reverse fulvous to saffron. On PDA reaching 40 mm diam, velvety, zonate, concentric circles and radial folds, umbers, cinnamon, ochreous to fulvous, margin lobate; reverse umber to fulvous; diffusible pigment ochreous.

Typus: Belgium, East Flanders, Brakel, from soil, 2019, K. De Borre, BE19_001 (holotype CBS H-24732, culture ex-type CBS 147215 = BE19_001008).

Additional material examined: Norway, Móre og Romsdal, on Pinus sylvestris wood, 31 Jul. 1974, K. Venn No. 74-55/7, culture CBS 551.75.

Notes: Coniochaeta deborreae is phylogenetically (Fig. 23) related to C. boothii, C. pulveraceae, C. subcorticalis, and C. rhopalochaeta. All these species were described as sexual morphs, except for C. rhopalochaeta that was described with both sexual and asexual morphs from decorticated wood of Bulnesia retama from Argentina (Romero et al. 1999). Coniochaeta deborreae differs from C. rhopalochaeta by having smaller conidiogenous cells (4–11 × 2–3.5 vs. 10–18 × 3.5–5 μm) and smaller conidia [2.5–4 × 1.5–2 μm vs. 5–8 × 3–5μm (Romero et al. 1999)]. Also see the overview phylogeny (Fig. 5).

Fig. 23.

Fig. 23.

Maximum composite likelihood tree obtained from the RAxML analysis of the combined ITS and LSU sequence alignment of selected Coniochaeta species. Bootstrap support values above 70 % are shown at the nodes. The novel species is indicated in a coloured block and bold face. The tree was rooted to Phialemonium limoniforme (CBS 139049) and Ph. obovatum (CBS 279.76). Alignment statistics: 16 strains including the outgroup; 930 characters including alignment gaps analysed (ITS: 495, LSU: 435). Model: GTR+G, alignment patterns: 400.

Coniochaeta deborrae is represented by two strains respectively isolated from soil in Belgium (CBS 147215) and Pinus sylvestris wood in Norway (CBS 551.75). However, isolate CBS 551.75 was initially identified as Phialophora mutabilis (described from river water, the Netherlands). This isolate was later found to be a species of Coniochaeta, and identified as C. subcorticalis (known from Alnus glutinosa, Germany) as it occurred on Pinaceae. Morphologically, it is not possible to compare the present strain CBS 147215, nor CBS 551.75 with C. subcorticalis, as both species only form asexual structures, and DNA data are not available for C. subcorticalis.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had the most similar identified species was Coniochaeta boothii (strain KoLRI_EL005219, GenBank NR_159776.1; Identities = 496/505 (98 %), two gaps (0 %)), C. cipronana (strain CBS 144016, GenBank NR_157478.1; Identities = 495/529 (94 %), 17 gaps (3 %)), and C. arenariae (strain MFLUCC 18-0409, GenBank MN047126.1; Identities = 492/526 (94 %), 14 gaps (2 %)). Closest hits using the LSU sequence are Coniochaeta taeniospora (strain LTA1, GenBank KU762325.; Identities = 792/814 (97 %), no gaps), C. leucoplaca (strain Jong54, GenBank FJ167399.1; Identities = 771/793 (97 %), no gaps), and C. marina (strain MFLUCC 18-0408, GenBank MK458765.1; Identities = 791/814 (97 %), no gaps) – also see Fig. 5. Closest hits using the tef1 are Coniochaeta rosae (strain MFLUCC 17-0810, Genbank MG829197.1; Identities = 827/894 (93 %), three gaps (0 %)), Coniochaeta endophytica (strain AEA 9094, Genbank MK693159.1; Identities = 806/872 (92 %), no gaps), and Coniochaeta baysunika (strain MFLUCC 17-0830, Genbank MG829196.1; Identities = 826/894 (92 %), three gaps (0 %)).

Author: M. Hernández-Restrepo

Diaporthe durionigena L.D. Thao, L.T. Hien, N.V. Liem, H.M. Thanh & T.N. Khanh, sp. nov. MycoBank MB 839285.

Synonym: Diaporthe durionigena L.D. Thao et al., Persoonia 44: 385. 2020, nom. inval. Art. 40.8 (Shenzhen).

Diagnosis: See Persoonia 44: 385. 2020 (Crous et al. 2020c).

Typus: Vietnam, Central Highlands, Dak Lak, on branches of Durio zibethinus (Malvaceae), Dec. 2018, L.D. Thao (holotype specimen VTCC 930005, preserved as metabolically inactive culture, culture ex-type KCSR1812.8 = VTCC 930005).

Note: Diaporthe durionigena is herewith validated, as the original description did not state that the holotype was preserved as a metabolically inactive culture.

Authors: L.D. Thao, L.T. Hien, N.V. Liem, H.M. Thanh & T.N. Khanh

Endoconidioma euphorbiae Crous, Persoonia 45: 301. 2020. Fig. 24.

Fig. 24.

Fig. 24.

Endoconidioma euphorbiae (CPC 38649). A. Colony sporulating on OA. B–C. Conidiogenous cells. D. Conidia. Scale bars = 10 μm.

Description and illustration: Crous et al. (2020a).

Material examined: South Africa, Western Cape Province, Nieuwoudtville, tip dieback on Euphorbia mauritanica (Euphorbiaceae), 2018, P.W. Crous, HPC 3069, culture CPC 38649.

Notes: Endoconidioma euphorbiae is a morphologically highly variable fungus commonly associated with leaf spots and dieback on Brunsvigia and Euphorbia (Crous et al. 2020a), and easily confused with E. leucospermi on Proteaceae (Taylor & Crous 2001, Crous et al. 2013a) .

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Endoconidioma euphorbiae (strain CPC 38551, GenBank MW175350.1; Identities = 590/590 (100 %), no gaps), Endoconidioma leucospermi (as Coniozyma leucospermi; strain CBS 111289, GenBank EU552113.1; Identities = 577/590 (98 %), four gaps (0 %)), and Hormonema carpetanum (strain 235J14, GenBank KU516485.1; Identities = 561/574 (98 %), four gaps (0 %)). Closest hits using the LSU sequence are Endoconidioma euphorbiae (strain CPC 38551, GenBank MW175390.1; Identities = 840/840 (100 %), no gaps), Endoconidioma leucospermi (as Coniozyma leucospermi; strain CBS 111289, GenBank EU552113.1; Identities = 836/841 (99 %), no gaps), and Hormonema carpetanum (strain ATCC 74360, GenBank MF611880.1; Identities = 835/841 (99 %), no gaps) – also see Fig. 1. Closest hits using the rpb2 sequence had highest similarity to Hormonema carpetanum (strain ATCC 74360, GenBank MF611881.1; Identities = 830/938 (88 %), three gaps (0 %)), Dothidea sambuci (strain CBS 198.58, GenBank KT216559.1; Identities = 768/945 (81 %), 17 gaps (1 %)), and Dothidea insculpta (strain CBS 189.58, GenBank AF107800.1; Identities = 761/944 (81 %), 15 gaps (1 %)). Closest hits using the tef1 (second part) sequence had highest similarity to Hormonema carpetanum (strain ATCC 74360, GenBank MF611882.1; Identities = 441/468 (94 %), no gaps), Alternaria septospora (strain CBS 109.38, GenBank JQ672442.1; Identities = 436/469 (93 %), two gaps (0 %)), and Alternaria smyrnii (strain EGS 37-093, GenBank JQ672454.1; Identities = 435/469 (93 %), two gaps (0 %)). No significant hits were obtained when the actA and tub2 sequences were used in blastn and megablast searches.

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Flammocladiella anomiae Lechat & J. Fourn., Ascomycete.org 11(6): 239. 2019. Fig. 25.

Fig. 25.

Fig. 25.

Flammocladiella anomiae (CPC 36302). A–C. Conidiogenous cells giving rise to conidia. D. Conidia. Scale bars = 10 μm.

On SNA. Mycelium consisting of hyaline, smooth, branched, septate, 2.5–3 μm diam hyphae. Conidiophores solitary or aggregated into orange sporodochia. Conidiophores hyaline, smooth, multiseptate, branched, subcylindrical, 30–130 × 4–5 μm. Conidiogenous cells terminal and intercalary, 10–55 × 3–4 μm, proliferating sympodially, scars truncate, 2–3 μm diam. Conidia solitary, hyaline, smooth, granular to guttulate, straight to slightly curved, subcylindrical to narrowly obclavate, apex obtuse to subobtuse, base truncate, 1.5–2 μm diam, (1–)3-septate, (20–)30–35(–45) × (3.5–)4–5 μm.

Culture characteristics: Colonies erumpent, spreading, with moderate aerial mycelium and smooth, lobate margin, reaching 10 mm diam (on MEA and PDA), 20 mm diam on OA after 2 wk at 25 °C. On MEA surface salmon to dirty white, reverse ochreous; on PDA surface salmon, reverse saffron; on OA surface salmon.

Material examined: Ukraine, Okhtyrka district, NNP Hetmanskyi, Sumska Dibrova, Klymentove village, dead branch of Robinia pseudoacacia (Fabaceae), occurring on conidiomata of Diaporthe oncostoma, 5 Aug. 2019, A. Akulov, HPC 2529, CWU AS 6855 (CBS H-24358, culture CPC 36302 = CBS 146685).

Notes: According to Voglmayr & Jaklitsch (2011), Massaria spp. are highly host-specific, with M. anomia being the only species occurring on Fabaceae (on ascomata of Massaria anomia on a branchlet of Robinia pseudoacacia). Lechat et al. (2019) speculated that F. anomiae is host-specific on Massaria anomia. In this regard, collecting M. anomiae on conidiomata of Diaporthe oncostoma is unusual, although the host plant was also a dead branch of Robinia pseudoacacia, suggesting some association with the host. The possibility exists that ascomata of M. anomia were poorly developed, and therefore not seen at the time of isolation. On MEA cultures formed large orange sporodochia, with conidia becoming longer and thinner, subcylindrical, apex obtuse, base truncate, 1.5–2 μm, flexuous, up to 8-septate, (50–)65–75(–80) × 3–4 μm. Conidia of Flammocladiella anomiae are reported to be smaller, namely 1–3-septate, smooth, (30–) 37–45(–48) × 2–2.5 μm (Lechat et al. 2019).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Flammocladiella anomiae (as Flammocladiella sp. CL-2019a; voucher JF17087, GenBank MN597423.1; Identities = 522/524 (99 %), no gaps), Ijuhya vitellina (strain DSM 104494, GenBank NR_154100.1; Identities = 522/584 (90 %), 19 gaps (3 %)), and Ijuhya corynospora (strain CBS 342.77, GenBank KY607539.1; Identities = 525/588 (89 %), 27 gaps (4 %)). Closest hits using the LSU sequence are Flammocladiella anomiae (as Flammocladiella sp. CL-2019a; voucher CLL16017, GenBank MN597426.1; Identities = 846/846 (100 %), no gaps), Flammocladiella aceris (strain CBS 138906, GenBank NG_058175.1; Identities = 816/830 (98 %), no gaps), and Ijuhya parilis (strain CBS 136677, GenBank KY607558.1; Identities = 824/846 (97 %), no gaps) – also see Fig. 4. The tef1 sequence generated in this study is identical to those of Flammocladiella anomiae generated for CBS 144256 and CBS 142775 (464/464 and 342/342, respectively). The tub2 sequence generated in this study is also identical to that of Flammocladiella anomiae strain CBS 144256 (526/526).

Authors: P.W. Crous, J.Z. Groenewald & A. Akulov

Floricola juncicola Crous & R.K. Schumach., sp. nov. MycoBank MB 839286. Fig. 26.

Fig. 26.

Fig. 26.

Floricola juncicola (CPC 38197). A–D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm.

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

Conidiomata pycnidial, separate, globose, dark brown with central ostiole, 180–200 μm diam. Peridium multi-layered, inner layers hyaline, outer layers brownish. Conidiophores reduced to conidiogenous cells lining the inner cavity, subcylindrical to doliiform, hyaline to pale brown, smooth to finely roughened, proliferating percurrently at apex, at times bifurcate at apex, with two loci, 5–15 × 3–4 μm. Conidia solitary, subcylindrical, apex subobtuse, tapering in lower third to truncate, unthickened hilum, 2–3 μm diam, olive to yellow-brown, with 1-2 different sized guttules per cell, with longitudinal striations covering the length of conidial body, ornamentation partially branched, (0–)3(–5)-septate in vitro, 7–10-septate, with septa golden, thick-walled and constricted in vivo, (48–)55–65(–80) × (5–)6(–7) μm.

Culture characteristics: Colonies erumpent, spreading, with moderate aerial mycelium and smooth, lobate margin, reaching 25 mm diam after 2 wk at 25 °C. On MEA surface dirty white, reverse isabelline in centre, cinnamon in outer region; on PDA surface dirty white, reverse olivaceous grey; on OA surface olivaceous grey.

Typus: France, La-Chaise-Dieu, on dead culm of Juncus sp. (Juncaceae), 24 Apr. 2019, A. Gardiennet, HPC 2970 = RKS 252 = AG19065 (holotype CBS H-24406, culture ex-type CPC 38197 = CBS 146811).

Notes: Jaklitsch et al. (2016) regarded several genera with aposphaeria-, coniothyrium-, or phoma-like asexual morphs as synonyms of Teichospora, presenting a new concept of a large, morphologically diverse genus representing the Teichosporaceae. This approach was however rejected by Boonmee et al. (2019), who recognised 14 genera in the family. Floricola juncicola clusters in the Teichosporaceae, being allied to Floricola clematidis (conidia 13–21 × 5–7 μm), from which it is distinct based on its conidial morphology. This also distinguishes it from other taxa in the genus (Phukhamsakda et al. 2020). A phylogenetic tree is presented as Fig. 27.

Fig. 27.

Fig. 27.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Teichosporaceae multigene (LSU / ITS / tef1) nucleotide alignment. Bayesian posterior probabilities (> 0.79) are shown at the nodes and the scale bar represents the expected changes per site. Families are indicated with coloured blocks to the right of the tree. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Melanomma pulvis-pyrius (culture CBS 124080) and the novel species treated in this study is indicated in a coloured block and bold face. Alignment statistics: 55 strains including the outgroup; 200, 315 and 359 unique site patterns, respectively. Tree statistics: 403 502 sampled trees from 1 345 000 generations.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Teichospora mariae (strain C134m, GenBank KU601580.1; Identities = 530/591 (90 %), 19 gaps (3 %)), Floricola clematidis (as Teichospora sp. CP-2020b; strain MFLUCC 17-2182, GenBank MT310638.1; Identities = 451/481 (94 %), five gaps (1 %)), and Teichospora kingiae (strain CPC 29104, GenBank NR_154656.1; Identities = 508/567 (90 %), 17 gaps (2 %)). Closest hits using the LSU sequence are Paulkirkia arundinis (as Teichosporaceae sp. DW-2016a; strain MFLUCC 12-0328, GenBank KU848206.1; Identities = 872/894 (98 %), three gaps (0 %)), Floricola clematidis (as Teichospora sp. CP-2020b; strain MFLUCC 17-2182, GenBank MT214594.1; Identities = 855/877 (97 %), two gaps (0 %)), and Teichospora kingiae (strain CPC 29104, GenBank NG_059761.1; Identities = 782/804 (97 %), two gaps (0 %)) – also see Fig. 1. No significant hits were obtained when the tef1 (first part) sequence was used in blastn and megablast searches. Closest hits using the tef1 (second part) sequence had highest similarity to Ramusculicola thailandica (as Teichospora thailandica; strain MFLUCC 17-2093, GenBank MT394653.1; Identities = 840/908 (93 %), five gaps (0 %)), Rhytidhysteron rufulum (voucher MFLU 18-2190, GenBank MK360087.1; Identities = 847/918 (92 %), six gaps (0 %)), and Ramusculicola clematidis (strain MFLUCC 17-2146, GenBank MT394652.1; Identities = 830/905 (92 %), five gaps (0 %)). Closest hits using the rpb2 sequence had highest similarity to Teichospora mariae (strain C136, GenBank KU601595.1; Identities = 558/648 (86 %), two gaps (0 %)), Teichospora striata (strain JK 5678I, GenBank GU371758.1; Identities = 493/610 (81 %), four gaps (0 %)), and Cenococcum geophilum (strain OS21018, GenBank LC095396.1; Identities = 291/368 (79 %), six gaps (1 %)).

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Harzia tenella (Berk. & M.A. Curtis) D.W. Li & N.P. Schultes, Fung. Biol. 121: 900. 2017. Fig. 28.

Fig. 28.

Fig. 28.

Harzia tenella (CPC 38667). A–D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm.

Basionym: Rhinotrichum tenellum Berk. & M.A. Curtis, Grevillea 3 (27): 109. 1875.

Description and illustration: Schultes et al. (2017).

Material examined: South Africa, Western Cape Province, Nieuwoudtville, on stems of Grielum humifusum (Neuradaceae), 2018, P.W. Crous, HPC 3043, culture CPC 38667.

Notes: In a study by Schultes et al. (2017), Harzia 1888 was shown to have priority over Olpitrichum 1894 and Chlamydomyces 1907, leading to the new combination H. tenella, which is here reported from Grielum humifusum in South Africa.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Harzia tenella (as Olpitrichum tenellum; strain CBS 121.81, GenBank KY628696.1; Identities = 605/607 (99 %), no gaps), Harzia verrucosa (strain CBS 113456, GenBank KY628674.1; Identities = 604/607 (99 %), one gap (0 %)), and Harzia patula (strain CBS 379.88, GenBank NR_161009.1; Identities = 602/608 (99 %), one gap (0 %)). Closest hits using the LSU sequence are Harzia macrospora (strain CBS 343.67, GenBank MH870687.1; Identities = 841/841 (100 %), no gaps), Harzia verrucosa (strain CBS 113456, GenBank KY628675.1; Identities = 841/841 (100 %), no gaps), and Harzia acremonioides (strain CBS 101.42, GenBank NG_067322.1; Identities = 841/841 (100 %), no gaps) – also see Fig. 5.

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Haudseptoria Crous & R.K. Schumach., gen. nov. MycoBank MB 839287.

Etymology: Name refers to the fact that it is morphologically similar to Septoria.

Conidiomata pycnidial, erumpent to semi-superficial, dark brown, globose, with central ostiole; wall in upper region of brown textura angularis, in lower region of pale brown cells, concolourous with host tissue. Conidiophores reduced to conidiogenous cells, ampulliform to subcylindrical, smooth, proliferating sympodially at apex. Conidia solitary, hyaline, smooth, guttulate, subcylindrical, apex subobtuse, base truncate, transversely 1–7-septate.

Type species: Haudseptoria typhae Crous & R.K. Schumach.

Haudseptoria typhae Crous & R.K. Schumach., sp. nov. MycoBank MB 839288. Fig. 29.

Fig. 29.

Fig. 29.

Haudseptoria typhae (CPC 38203). A. Conidioma on host tissue. B. Conidiomatal ostiole. C. Conidiogenous cells giving rise to conidia. D. Conidia. Scale bars: A = 150 μm, all others = 10 μm.

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

Conidiomata erumpent to semi-superficial, dark brown, globose, 100–150 μm diam with central ostiole, 10–15 μm diam; wall in upper region of brown textura angularis, in lower region of pale brown cells, concolourous with host tissue. Conidiophores reduced to conidiogenous cells, ampulliform to subcylindrical, smooth, 5–12 × 2–3 μm, proliferating sympodially at apex. Conidia solitary, hyaline, smooth, guttulate, subcylindrical, apex subobtuse, base truncate, filiform, straight to slightly curved, (1–)3-septate, septa thin-walled, (16–)20–23(–27) × 2(–2.5) μm in vivo; 35–90 × 3 μm, 3–7-septate in vitro.

Culture characteristics: Colonies erumpent, spreading, with sparse aerial mycelium and smooth, even margin, reaching 12 mm diam after 2 wk at 25 °C. On MEA surface buff, reverse cinnamon; on PDA surface cinnamon, reverse honey with patches of cinnamon; on OA surface buff.

Typus: Germany, near Berlin, moist meadow, on dead leaf sheath of Typha sp. (Typhaceae), 17 Apr. 2019, R.K. Schumacher, HPC 2978 = RKS 249 (holotype CBS H-24407, culture ex-type CPC 38203 = CBS 146790).

Notes: Haudseptoria represents a new septoria-like genus (Verkley et al. 2013) related to Pseudopassalora gouriqua (hyphomycete on Protea suzannae, South Africa; Crous et al. 2011), and Phellinocrescentia guianensis (coelomycete with aseptate conidia, on polypores in French Guiana; Crous et al. 2014).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Didymosphaeria futilis (strain CMW 22186, GenBank EU552123.1; Identities = 487/521 (93 %), 13 gaps (2 %)), Italiofungus phillyreae (strain CPC 35566, GenBank MT223804.1; Identities = 446/499 (89 %), 16 gaps (3 %)), and Funbolia dimorpha (strain CPC 14170, GenBank JF951136.1; Identities = 405/467 (87 %), 18 gaps (3 %)). Closest hits using the LSU sequence are Pseudopassalora gouriqua (strain CBS 101954, GenBank NG_067272.1; Identities = 785/808 (97 %), no gaps), Italiofungus phillyreae (strain CPC 35566, GenBank MT223899.1; Identities = 784/808 (97 %), no gaps), and Eriomyces heveae (strain MFLUCC 17-2232, GenBank MH109524.1; Identities = 779/810 (96 %), two gaps (0 %)) – also see Fig. 1. No significant hits were obtained when the tub2 sequence was used in blastn and megablast searches.

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Heimiodora verticillata Nicot, Ann. Sci. Nat., Bot. Biol. Vég., sér. 12: 384. 1960. Fig. 30.

Fig. 30.

Fig. 30.

Heimiodora verticillata (CPC 39015). A. Sporulation on SNA. B. Setae. C. Chlamydospores. D–E. Conidiogenous cells. F. Conidia. Scale bars = 10 μm.

Description and illustration: See Nicot & Parguey (1960).

Material examined: Namibia, Walvis Bay, on Salvadora persica (Salvadoraceae), 20 Nov. 2019, P.W. Crous, HPC 3120, CBS H-24533, cultures CPC 39015 = CBS 147089.

Notes: Heimiodora verticillata was originally isolated from sandy coastal soil in Thailand (Nicot & Parguey 1960; ex-type CBS 201.60), and its ecology has remained rather obscure. Isolating the same fungus from twig litter of a Salvadora persica growing on the dry desert coast in Walvis Bay, Namibia, suggests that this fungus could prefer a dry arid environment. A specimen of this fungus has also been reported from fallen leaves of a Salvadora sp. growing in Kasur, Pakistan (BPI 1100673). The host, Salvadora, is known to occur in dry environments, which could give a hint to the ecology of H. verticillata. The ITS and LSU sequences of CPC 39015 are a perfect match to those of the ex-type culture (CBS 201.60; GenBank MH857955.1; 554/554 (100 %), no gaps, and GenBank MH869505.1; 833/833 (100 %), no gaps, respectively) – also see Fig. 4.

Authors: P.W. Crous, D.A. Cowan, G. Maggs-Kölling, E. Marais, N. Yilmaz & M.J. Wingfield

Hogelandia Hern.-Restr., gen. nov. MycoBank MB 838707.

Etymology: Named after the school where the sample was collected, Het Hogeland College (Warffum, the Netherlands). This sample was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae hyaline, septate, smooth. Conidiophores micronematous, mostly reduced to conidiogenous cells. Conidiogenous cells monoblastic, subcylindrical, smooth, hyaline. Secession rhexolytic. Conidia hyaline, smooth, thick-walled, globose to subglobose, sometimes in short chains, base with a cylindrical to cuneiform cell.

Type species: Hogelandia lambearum Hern.-Restr.

Hogelandia lambearum Hern.-Restr., sp. nov. MycoBank MB 838708. Fig. 31.

Fig. 31.

Fig. 31.

Hogelandia lambearum (CBS 147626). A–B. Colony on MEA. C. Colony on OA. D–E. Mycelium and conidia. F–H. Conidiogenous cells and conidia. I–L. Conidia. Scale bars D–E, L = 10 μm. L applies to F–L.

Etymology: Name includes the initials of the collectors Lara Alssema and Meike van Berkel, students from Het Hogeland College (Warffum, the Netherlands). This sample was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae hyaline, septate, smooth, 1.5–4 μm. Conidiophores micronematous, mostly reduced to conidiogenous cells. Conidiogenous cells monoblastic, subcylindrical, smooth, hyaline, 5–10.5 × 2–3 μm. Secession rhexolytic. Conidia hyaline, smooth, thick-walled, globose to subglobose, sometimes in short chains, 8–14 μm diam, base with a cylindrical to cuneiform cell.

Culture characteristics: After 1 wk at 25 °C on OA reaching 52 mm diam, zonate, cottony, white, with moderate aerial mycelium and glabrous zones at the centre, margin entire, regular. On MEA reaching 53 mm diam, zonate, white, margin regular, reverse ochreous.

Typus: Netherlands, Groningen Province, Warffum, Het Hogeland College, from soil, 6 Jun. 2019, L. Alssema & M. van Berkel, NL19_027 (holotype CBS H-24733, culture ex-type CBS 147626 = NL19_27007).

Notes: Hogelandia is phylogenetically related to Ascodesmis, Eleutherascus and Cephaliophora in Pezizomycetes (Fig. 3). Species of these genera are also found in soil and dung samples (Guarro et al. 2012). Members of Ascodesmis, Eleutherascus and Cephaliophora are all described as sexual morphs, except for E. tuberculata that was described with a sporothrix- or calcarisporium-like asexual morph, characterised by erect and branched conidiophores with polyblastic conidiogenous cells that produce ellipsoidal conidia, 8.5–12.5 × 5–7.5 μm (Samson & Luiten 1975). Hogelandia is represented only by the asexual morph, characterised by micronematous conidiophores, monoblastic conidiogenous cells and subglobose conidia. Attempts to induce the sexual morph in vitro were unsuccessful.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Cephaliophora sp. (strain TD4, GenBank KY814682.1; Identities = 476/508 (94 %), five gaps (0 %)), Cephaliophora tropica (strain GN-HA-1-3, GenBank MG554311.1; Identities = 340/369 (92 %), six gaps (1 %)), and Cephaliophora tropica (strain JB-NW-1-1, GenBank MG554299.1; Identities = 333/365 (91 %), five gaps (1 %)). Closest hits using the LSU sequence are Ascodesmis nigricans (strain CBS 163.74, GenBank MH872582.1; Identities = 810/825 (98 %), no gaps), Ascodesmis microscopica (strain CBS 275.80, GenBank MH873032.1; Identities = 809/825 (98 %), no gaps), and Ascodesmis sphaerospora (strain CBS 125.61, GenBank MH869550.1; Identities = 809/825 (98 %), no gaps) – also see Fig. 3.

Author: M. Hernández-Restrepo

Lomentospora valparaisensis E. Álvarez, sp. nov. MycoBank MB 839289.

Synonym: Lomentospora valparaisensis E. Álvarez, Persoonia 40: 299. 2018. Nom. inval., Art. 40.7 (Shenzhen).

Diagnosis: See Persoonia 40: 299. 2018 (Crous et al. 2018b).

Typus: Chile, Valparaiso, Italy Park, from soil, 2016, F. Salas (holotype specimen ChFC 2018164, culture ex-type ChFC-164).

Note: Lomentospora valparaisensis is validated here because the original description cited an incorrect acronym for the holotype specimen.

Author: E. Álvarez Duarte

Muyocopron zamiae Hern.-Restr. & Crous, Persoonia 42: 219. 2019. Fig. 32.

Fig. 32.

Fig. 32.

Muyocopron zamiae (CPC 37461). A. Colony sporulating on OA. B–F. Conidiogenous cells giving rise to conidia. G. Conidia. Scale bars = 10 μm.

Mycelium consisting of branched, septate, dark to pale brown, verruculose, 2–3 μm diam hyphae. Conidiomata sporodochium-like, superficial on agar, medium brown, consisting of a few aggregated cells to large round aggregations of conidiogenous cells. Conidiogenous cells globose to ampulliform, 12–20 × 9–12 μm, with distinct flared collarette, 3–6 μm diam, medium to dark brown, smooth. Conidia aseptate, lunate, fusoid, curved, guttulate, (17–)18–22(–25) × (4.5–)6(–6.5) μm, with a filiform, unbranched appendage at each end, (2–)7–11 mm long, hyaline, smooth. Appressoria and sexual morph not observed.

Culture characteristics: Colonies flat, spreading, with sparse to moderate aerial mycelium and feathery, lobate margin, covering dish after 2 wk at 25 °C. On MEA surface buff to honey, reverse isabelline; on PDA surface honey, reverse isabelline; on OA surface honey to isabelline.

Material examined: USA, Florida, Gainesville, on leaves of Zamia integrifolia (Zamiaceae), 28 Feb. 2018, M.J. Wingfield, HPC 2794, CBS H-24348, culture CPC 37461 = CBS 146636.

Notes: Muyocopron zamiae was described from leaves of Zamia fisheri collected in Florida, USA (Hernández-Restrepo et al. 2019). The collection in this study (on Zamia integrifolia), was also made in Florida, and represents the second known collection for this species, which seems to be common on leaves of Zamia spp.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Muyocopron zamiae (strain CBS 203.71, Present study; Identities = 520/522 (99 %), no gaps), Muyocopron coloratum (strain CBS 720.95, GenBank NR_160197.1; Identities = 297/306 (97 %), one gap (0 %)), and Fimetariella rabenhorstii (strain A27_ITS-1, GenBank MN447419.1; Identities = 449/465 (97 %), two gaps (0 %)). Closest hits using the LSU sequence are Muyocopron zamiae (strain CBS 203.71, GenBank NG_066338.1; Identities = 837/837 (100 %), no gaps), Muyocopron geniculatum (strain CBS 721.95, GenBank NG_066266.1; Identities = 834/838 (99 %), one gap (0 %)), and Muyocopron chromolaenae (strain MFLUCC 17-1513, GenBank NG_068700.1; Identities = 823/834 (99 %), two gaps (0 %)) – also see Fig. 1. Closest hits using the tef1 (second part) sequence had highest similarity to Muyocopron zamiae (strain CBS 203.71, GenBank MK495973.1; Identities = 460/460 (100 %), no gaps), Muyocopron atromaculans (strain MUCL 34983, GenBank MK495957.1; Identities = 441/460 (96 %), no gaps), and Mycoleptodiscus terrestris (strain IMI 159038, GenBank MK495977.1; Identities = 439/460 (95 %), no gaps). Closest hits using the rpb2 sequence had highest similarity to Muyocopron zamiae (strain CBS 203.71, GenBank MK492731.1; Identities = 846/846 (100 %), no gaps), Muyocopron geniculatum (strain CBS 721.95, GenBank MK492715.1; Identities = 761/822 (93 %), three gaps (0 %)), and Muyocopron chromolaenae (strain MFLUCC 17-1513, GenBank MT136761.1; Identities = 508/569 (89 %), no gaps).

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Neocamarosporium leipoldtiae Crous, Persoonia 45: 295. 2020. Fig. 33.

Fig. 33.

Fig. 33.

Neocamarosporium leipoldtiae (CPC 38543). A. Conidioma on SNA. B. Conidiogenous cell giving rise to conidia. C–D. Conidia. Scale bars: A = 300 μm, all others = 10 μm.

Conidiomata globose, brown, 200–300 μm diam, immersed, becoming erumpent with central ostiole; wall of 3–6 layers of brown textura angularis. Conidiophores reduced to conidiogenous cells lining the inner cavity. Conidiogenous cells hyaline, smooth, ampulliform, proliferating percurrently at apex, 8–12 × 5–7 μm. Conidia solitary, medium brown, thick-walled, muriformly septate, obovoid to ellipsoid, finely roughened, thick-walled; with 3–4 horizontal septa, and 3–6 vertical or oblique septa, (13–)16–18(–20) × (9–)10–11(–12) μm.

Culture characteristics: Colonies erumpent, spreading, with moderate aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA surface and reverse sepia; on PDA surface isabelline, reverse hazel; on OA surface sepia.

Material examined: South Africa, Western Cape Province, Clanwilliam, Rocklands camp grounds, on leaves of Cephalophyllum pilansii (Aizoaceae), 2018, P.W. Crous, HPC 3055 (CBS H-24421, culture CPC 38543 = CBS 146812).

Notes: Neocamarosporium leipoldtiae was recently introduced for a species occurring on leaves of Leipoldtia schultzii in South Africa (Crous et al. 2020a). The collection in this study originated from the same region of the Western Cape Province, where it was isolated from leaves of Cephalophyllum pilansii.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Neocamarosporium sp. (strain CF-288928, GenBank MG065823.1; Identities = 536/548 (98 %), one gap (0 %)), Neocamarosporium salicornicola (strain ZMCS3, GenBank MK809918.1; Identities = 512/524 (98 %), two gaps (0 %)), and Dimorphosporicola tragani (strain ZMCL7, GenBank MK809922.1; Identities = 510/523 (98 %), one gap (0 %)). Closest hits using the LSU sequence are Neocamarosporium chichastianum (strain IBRC-M 30126, GenBank KP004483.1; Identities = 811/815 (99 %), no gaps), Neocamarosporium salicornicola (strain MFLUCC 15-0957, GenBank NG_070423.1; Identities = 808/813 (99 %), no gaps), and Neocamarosporium chersinae (strain CPC 27298, GenBank KY929182.1; Identities = 799/804 (99 %), no gaps) – also see Fig. 1. Closest hits using the tef1 sequence had highest similarity to Dimorphosporicola tragani (strain CBS 570.85, GenBank KU728577.1; Identities = 406/458 (89 %), ten gaps (2 %)), Ulocladium alternariae (strain P292, GenBank AY375370.1; Identities = 335/383 (87 %), seven gaps (1 %)), and Alternaria japonica (strain P400, GenBank AY375367.1; Identities = 332/384 (86 %), eight gaps (2 %)). Closest hits using the tub2 sequence had highest similarity to Dimorphosporicola tragani (strain CBS 570.85, GenBank KU728616.1; Identities = 305/329 (93 %), one gap (0 %)), Dothidotthia robiniae (as Dothidotthia sp. CS-2019b; strain MFLUCC 18-0692, GenBank MK933790.1; Identities = 293/331 (89 %), ten gaps (3 %)), and Tamaricicola sp. (strain IG114, GenBank MH001474.1; Identities = 292/331 (88 %), six gaps (1 %)).

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Neofusicoccum mystacidii Crous, sp. nov. MycoBank MB 839290. Fig. 34.

Fig. 34.

Fig. 34.

Neofusicoccum mystacidii (CPC 39221). A. Sporulation on PDA. B–C. Conidiogenous cells. D. Macroconidia. E. Microconidia. Scale bars: A = 300 μm, all others = 10 μm.

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

On PNA. Conidiomata globose, erumpent, brown, pycnidial, with central ostiole, punctiform, 200–300 μm diam; wall of 6–8 layers of brown textura angularis. Macroconidiophores reduced to conidiogenous cells, or at times with a supporting cell, lining the inner cavity of conidioma, hyaline, smooth, subcylindrical, proliferating percurrently at apex, encased in mucoid sheath, 10–25 × 4–5 μm. Macroconidia aseptate, solitary, hyaline, smooth, guttulate, straight, broadly cylindrical, apex obtuse, tapering in bottom part to truncate hilum, 3–4 μm diam, with minute marginal frill, encased in mucoid sheath, (26–)28–30(–35) × (9–)10(–11) μm. Microconidiophores mixed among macroconidiophores, subcylindrical, hyaline smooth, 5–12 × 3–4 μm, phialidic with periclinal thickening. Microconidia solitary, aseptate, subcylindrical with obtuse ends, hyaline, smooth, 3–6 × 2.5–3 μm.

Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, lobate margin, reaching 50 mm diam after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse olivaceous grey.

Typus: South Africa, Northern Province, Nelspruit, Lowveld Botanical Garden, on dead stems of Mystacidium capense (Orchidaceae), 23 Nov. 2018, P.W. Crous, HPC 3141 (holotype CBS H-24546, culture ex-type CPC 39221 = CBS 147079).

Notes: Conidiomata of N. mystacidii had slightly elongated necks in vivo, and conidia were also encased in a mucoid sheath, both characters that are not typically associated with members of Neofusicoccum (Phillips et al. 2013, Zhang et al. 2021a), and not observed in vitro. Neofusicoccum mystacidii is related to species such as N. cryptoaustrale (conidia (18–)20.5–21(–26.5) × 5–6(–6.5) μm; Crous et al. 2013b), N. nonquaesitum (conidia 17–29 × 5.5–10.5 μm; Phillips et al. 2013), N. mediterraneum (conidia (19–)22–26(–27) × (5.5–)6(–6.5) μm; Phillips et al. 2013) and N. vitifusiforme (conidia (9–)9.5–13(–14.5) × (6.5–)8–10.5(–11) μm; Phillips et al. 2013), species that are morphologically quite distinct. A phylogenetic species tree is presented as Fig. 35.

Fig. 35.

Fig. 35.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Neofusicoccum multigene (ITS / tef1 / tub2 / rpb2) nucleotide alignment. The alignment is derived from Zhang et al. (2021) and GenBank accession numbers and partition models can be found in that study. Bayesian posterior probabilities (> 0.79) are shown at the nodes and the scale bar represents the expected changes per site. Culture collection numbers in bold face represent cultures with a type status. The tree was rooted to Botryosphaeria dothidea (culture CBS 115476) and the novel species treated in this study is indicated in a coloured block and bold face. Alignment statistics: 78 strains including the outgroup; 119 / 159 / 110 / 146 unique site patterns, respectively. Tree statistics: 49 502 sampled trees from 1 650 000 generations.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Neofusicoccum corticosae (now Neofusicoccum vitifusiforme; strain CBS 120081, GenBank MN161920.1; Identities = 542/556 (97 %), no gaps), Neofusicoccum vitifusiforme (strain CAP227, GenBank EF638785.1; Identities = 542/556 (97 %), no gaps), and Neofusicoccum hellenicum (strain CERC1959, GenBank KP217057.1; Identities = 532/547 (97 %), no gaps). Closest hits using the LSU sequence are Neofusicoccum nonquaesitum (strain CBS 126655, GenBank MH875645.1; Identities = 877/886 (99 %), no gaps), Neofusicoccum arbuti (strain CBS 117090, GenBank DQ377919.1; Identities = 877/886 (99 %), no gaps), and Neofusicoccum mangiferae (strain CBS 118532, GenBank NG_055730.1; Identities = 876/886 (99 %), no gaps) – also see Fig. 1. Closest hits using the gapdh sequence had highest similarity to Neofusicoccum mediterraneum (strain PD312, GenBank GU251440.1; Identities = 492/497 (99 %), no gaps), Neofusicoccum parvum (strain JPZZP4, GenBank KY009944.1; Identities = 512/530 (97 %), one gap (0 %)), and Neofusicoccum australe (strain PD253, GenBank GU251483.1; Identities = 485/497 (98 %), one gap (0 %)). Closest hits using the his3 sequence had highest similarity to Neofusicoccum mediterraneum (strain PD311, GenBank GU251571.1; Identities = 306/309 (99 %), no gaps), Neofusicoccum parvum (strain PD6, GenBank GU251525.1; Identities = 303/309 (98 %), no gaps), and Neofusicoccum australe (strain PD253, GenBank GU251615.1; Identities = 302/309 (98 %), no gaps). Closest hits using the rpb2 sequence had highest similarity to Neofusicoccum terminaliae (strain CBS 125263, GenBank KX464045.1; Identities = 388/391 (99 %), no gaps), Neofusicoccum mediterraneum (strain CBS 121718, GenBank KY855815.1; Identities = 387/391 (99 %), no gaps), and Neofusicoccum ursorum (strain CBS 122811, GenBank KX464047.1; Identities = 386/391 (99 %), no gaps). Closest hits using the tef1 sequence had highest similarity to Neofusicoccum mediterraneum (strain BAL-17, GenBank KX029198.1; Identities = 383/390 (98 %), no gaps), Neofusicoccum vitifusiforme (strain CMW875, GenBank HM176519.1; Identities = 385/393 (98 %), two gaps (0 %)), and Neofusicoccum luteum (strain CMW42482, GenBank KP860705.1; Identities = 363/373 (97 %), two gaps (0 %)). Closest hits using the tub2 sequence had highest similarity to Neofusicoccum parvum (strain CCF216, GenBank KC507808.1; Identities = 573/587 (98 %), no gaps), Neofusicoccum australe (strain CUZF10AV1, GenBank KU836639.1; Identities = 558/572 (98 %), one gap (0 %)), and Neofusicoccum mangiferae (strain wbhs2, GenBank KY435929.1; Identities = 567/587 (97 %), two gaps (0 %)).

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Neomycosphaerella guibourtiae Crous & Jol. Roux, sp. nov. MycoBank MB 839291. Fig. 36.

Fig. 36.

Fig. 36.

Neomycosphaerella guibourtiae (CPC 39348). A. Ascomata immersed on host tissue. B. Ascoma. C–D. Asci. E. Ascospores. Scale bars: A–B = 90 μm, all others = 10 μm.

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

Leaf spots amphigenous, irregular, brown with raised brown border, up to 10 mm diam. Ascomata hypophyllous, immersed, becoming erumpent, but remaining immersed under epidermis, brown, globose, 70–90 μm diam, with central ostiole, slightly papillate, up to 25 μm diam, periphysate; wall of 4–6 layers of brown textura angularis. Asci stipitate, fasciculate, obovoid, hyaline, smooth, bitunicate, 8-spored, apical chamber 1 μm diam, 32–38 × 8–10 μm. Ascospores tri- to multiseriate, fusoid-ellipsoid, widest in middle of apical cell which can be asymmetrical, basal cell obtusely rounded, medianly 1-septate, not constricted at septum, guttulate to granular, (14–)16–18(–20) × (3–)3.5(–4) μm.

Culture characteristics: Colonies erumpent, spreading, surface folded, with sparse aerial mycelium and smooth, lobate margin, reaching 5–7 mm diam after 2 wk at 25 °C. On MEA and PDA surface pale olivaceous grey, reverse olivaceous grey; on OA surface ochreous in centre, pale olivaceous grey in outer region.

Typus: Angola, Cuanavale Source Lake, Fairy forest, on leaves of Guibourtia sp. (Fabaceae), 27 Nov. 2015, J. Roux, HPC 3242 (holotype CBS H-24551, culture ex-type CPC 39348 = CBS 147083.

Notes: Neomycosphaerella guibourtiae is closely related to Neomycosphaerella pseudopentameridis [from Pseudopentameris macrantha (Poaceae), South Africa; Crous et al. 2013b, Videira et al. 2017]. Morphologically they are quire distinct, however, in that ascospores of N. pseudopentameridis [(15–)16–17(–18) × (3.5–)4(–5) μm] are shorter and wider than those of N. guibourtiae. A phylogenetic species tree is presented as Fig. 37.

Fig. 37.

Fig. 37.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Mycosphaerellaceae multigene (LSU / ITS / rpb2) nucleotide alignment. The alignment is derived from the alignment of Videira et al. (2017) and partition models can be found in that study. Bayesian posterior probabilities (> 0.79) are shown at the nodes and the scale bar represents the expected changes per site. The most basal branch was halved to facilitate layout. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Cladosporium cladosporioides (culture CBS 112388) and the novel species treated in this study is indicated in a coloured block and bold face. Alignment statistics: 31 strains including the outgroup; 80 / 214 / 459 unique site patterns, respectively. Tree statistics: 192 002 sampled trees from 640 000 generations.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Hyalozasmidium aerohyalinosporum (strain CBS 125011, GenBank NR_156220.1; Identities = 484/532 (91 %), 19 gaps (3 %)), Mycosphaerella medusae (strain BRIP 52586, GenBank NR_137044.1; Identities = 467/514 (91 %), 20 gaps (3 %)), Madagascaromyces intermedius (strain CBS 124154, GenBank MH863356.1; Identities = 488/538 (91 %), 20 gaps (3 %)), and Paramycosphaerella dicranopteridis (strain TNM 3953, GenBank NR_155639.1; Identities = 479/535 (90 %), 19 gaps (3 %)). Closest hits using the LSU sequence are Paramycosphaerella brachystegiae (strain CBS 136436, GenBank NG_058048.1; Identities = 850/860 (99 %), no gaps), Paramycosphaerella marksii (strain CPC 11222, GenBank GU214447.1; Identities = 850/860 (99 %), no gaps), and Paramycosphaerella watsoniae (strain CPC 37392, GenBank NG_068339.1; Identities = 831/841 (99 %), no gaps) – also see Fig. 1. Closest hits using the actA sequence had highest similarity to Paramycosphaerella watsoniae (strain CBS 146064, GenBank MN556790.1; Identities = 554/608 (91 %), 15 gaps (2 %)), Mycosphaerella musae (strain CIRADCOK 003, GenBank MW070786.1; Identities = 542/596 (91 %), seven gaps (1 %)), and Hyalozasmidium aerohyalinosporum (strain CBS 125011, GenBank KF903576.1; Identities = 497/544 (91 %), 15 gaps (2 %)). Distant hits obtained using the cmdA sequence had highest similarity to Nowamyces piperitae (strain CPC 32372, GenBank MN162565.1; Identities = 286/308 (93 %), no gaps), Zasmidium pseudoparkii (strain CBS 111049, GenBank KF902695.1; Identities = 281/301 (93 %), no gaps), and Zasmidium corymbiae (strain CBS 145049, GenBank MK047526.1; Identities = 281/305 (92 %), no gaps). Distant hits obtained using the tef1 sequence had highest similarity to Microcyclosporella mali (strain CPC 16177, GenBank HM177418.1; Identities = 334/434 (77 %), 51 gaps (11 %)), Cercospora zeae-maydis (strain CBS 132678, GenBank JX143502.1; Identities = 208/243 (86 %), five gaps (2 %)), and Pseudocercospora eucalyptorum (strain CPC 35905, GenBank MN162363.1; Identities = 219/266 (82 %), 18 gaps (6 %)). Closest hits using the tub2 sequence had highest similarity to Paramycosphaerella intermedia (strain CBS 114356, GenBank KF902845.1; Identities = 290/342 (85 %), 12 gaps (3 %)), Paramycosphaerella marksii (strain CBS 110920, GenBank KF902848.1; Identities = 292/345 (85 %), 18 gaps (5 %)), and Hyalozasmidium aerohyalinosporum (strain CBS 125011, GenBank KF903073.1; Identities = 289/345 (84 %), 16 gaps (4 %)).

Authors: P.W. Crous, J.Z. Groenewald & J. Roux

Neoscirrhia Crous & R.K. Schumach., gen. nov. MycoBank MB 839292.

Etymology: Name refers to its morphological similarity to Scirrhia.

Ascomata pseudothecial, gregarious, stromatic, immersed, becoming erumpent, opening by longitudinal split of epidermis, arranged parallel to the culm axis, black, thick- and smooth-walled, multi-locular, loculi in one row and one level, globose, black, soft, ostiole central, indistinct. Peridium multi-layered, of textura angularis-prismatica, inner layers hyaline, outer layers dark brown. Pseudoparaphyses numerous, partly evanescent, multi-celled, moniliform to cylindrical, branched, anastomosing. Asci 8-spored, subclavate to cylindrical, apically rounded with ocular chamber, pedicel short, broad and furcate, bitunicate, fissitunicate. Ascospores straight to slightly curved, cells cylindrical, ends rounded, upper cell enlarged towards median septum, at times with lateral swelling just above the septum hyaline, thin- and smooth-walled, constricted at septum, cytoplasm aggregated in middle region, causing ends to appear very thick-walled, apex more prominently so than basal end. Conidiomata pycnidial, globose, brown, with central ostiole; wall of 3–6 layers of brown textura angularis. Conidiophores reduced to conidiogenous cells lining inner cavity, hyaline, smooth, doliiform to ampulliform, phialidic. Conidia solitary, hyaline, smooth, prominently guttulate, medianly 1-septate, subcylindrical to obovoid, ends obtuse.

Type species: Neoscirrhia osmundae (Peck & Clinton) Crous & R.K. Schumach.

Neoscirrhia osmundae (Peck & Clinton) Crous & R.K. Schumach., comb. nov. MycoBank MB 839293. Fig. 38.

Fig. 38.

Fig. 38.

Neoscirrhia osmundae (CPC 38085). A. Conidioma on PNA. B. Conidioma with ostiole. C. Conidiogenous cells. D. Conidia. E–F. Ascomata on host tissue. G. Asci. H. Ascospores (arrows indicate apical thickenings). Scale bars: A = 300 μm, B, F = 200 μm, E = 180 μm, all others = 10 μm.

Basionym: Dothidea osmundae Peck & Clinton, Rep. (Annual) New York State Mus. Nat. Hist. 30: 64. 1878. (1877)

Synonyms: Scirrhia osmundae (Peck & Clinton) Arx, in Müller & von Arx, Beitr. Kryptfl. Schweiz 11(no. 2): 382. 1962.

Metameris osmundae (Peck & Clinton) Arx & E. Müll., Stud. Mycol. 9: 80. 1975.

Scirrhia osmundae (Peck & Clinton) L. Holm & K. Holm, Bot. Notiser 131: 112. 1978, an isonym, see Art. 6.3 Note 2.

Ascomata pseudothecial, saprobic, gregarious, stromatic, immersed, becoming erumpent, opening by longitudinal split of epidermis, arranged parallel to the culm axis, black, thick- and smooth-walled, multi-locular, loculi in one row and one level, globose, black, soft, ostiole central, indistinct, up to 180 × 200 μm. Peridium multi-layered, of textura angularis-prismatica, inner layers hyaline, outer layers dark brown. Pseudoparaphyses numerous, partly evanescent, multi-celled, moniliform to cylindrical, branched, anastomosing. Asci 8-spored, subclavate to cylindrical, apically rounded with ocular chamber, pedicel short, broad and furcate, bitunicate, fissitunicate, 71–82 × 8.5–11.5 μm, ascospores obliquely biseriate. Ascospores straight to slightly curved, cells cylindrical, ends rounded, upper cell enlarged towards median septum, at times with lateral swelling just above the septum hyaline, thin- and smooth-walled, constricted at septum, cytoplasm aggregated in middle region, causing ends to appear very thick-walled, apex more prominently so than basal end, enclosed in mucilaginous sheath when immature (in water), 15.5–21.5 × 4–5 μm. Conidiomata pycnidial, globose, 250–300 μm diam, brown, with central ostiole, 30–40 μm diam, solitary on SNA, aggregated on OA; wall of 3–6 layers of brown textura angularis. Conidiophores reduced to conidiogenous cells lining inner cavity, hyaline, smooth, doliiform to ampulliform, phialidic, 5–7 × 5–6 μm. Conidia solitary, hyaline, smooth, prominently guttulate, medianly 1-septate, subcylindrical to obovoid, ends obtuse, (10–)12–14(–15) × (4–)4.5(–5) μm.

Culture characteristics: Colonies flat, spreading, with abundant aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse olivaceous grey.

Material examined: Netherlands, Noord-Holland Province, Hoofddorp, on culms of Sasa veitchii (Poaceae), 18 May 2019, L. van der Linde, HPC 2956 = RKS 1103 (CBS H-24397, culture CPC 38085 = CBS 146803).

Notes: Scirrhia is based on S. rimosa, a species known from stems of Phragmites australis (Poaceae) collected in Europe. No culture was available so far and therefore the phylogenetic affinities were unresolved. Scirrhia brasiliensis is a morphologically similar species, recently described (on fronds of Pteridium aquilinum) in Brazil. It was shown to cluster in Mycosphaerellaceae (Crous et al. 2011), a family which type, S. rimosa, is expected to also belong to Schirrhia s.str. It is, however, quite distinct from S. osmundae, which has a different ascospore morphology (see below), and clusters in Didymellaceae.

Scirrhia osmundae was described from dead stems of Osmunda collected in New York, USA. Ascospores are 2-celled, but appear 3-celled due to the contraction of the protoplast in the upper cell. This phenomenon was misinterpreted by Theissen & Sydow (1915) as 3-celled ascospores. It is quite characteristic in mature ascospores, and was also observed by Holm & Holm (1978), and in the present study. Based on Holm & Holm (1978) and Obrist (1959), stromata are larger in European collections, and ascospores are 15–18 × 4 μm (15.5–21.5 × 4–5 μm in the present specimen).

Metameris japonica (from Japan) represents another possible synonym of S. osmundae. Scirrhia osmundae was first reported from Europe (Germany) by Theissen & Sydow (1915), while Obrist (1959) reported it from Switzerland, and Holm & Holm (1978) reported it from Sweden. The collection in this study appears to be the first record from the Netherlands.

Phoma matteuccicola is commonly known as a pathogen causing leaf blight and “gangrene” of fern species (De Gruyter et al. 2002). Because it is allied to Neoscirrhia osmundae, it is herewith placed in this genus.

Neoscirrhia matteucciicola (Aderkas et al.) Crous, comb. nov. MycoBank MB 839294.

Basionym: Phoma matteucciicola Aderkas et al., Canad. J. Pl. Pathol. 14: 227. 1992.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence of CPC 38085 had highest similarity to Microsphaeropsis olivacea (strain CBS 432.71, GenBank MH871969.1; Identities = 526/540 (97 %), two gaps (0 %)), Coniothyrium euonymi-japonicae (strain CBS 302.68, GenBank MH859134.1; Identities = 526/540 (97 %), two gaps (0 %)), Coniothyrium glomerulatum (strain CBS 113.21, GenBank MH854688.1; Identities = 526/540 (97 %), two gaps (0 %)), and Neoscirrhia matteucciicola (as Phoma matteucciicola; strain CBS 259.92, GenBank GU237812.1; Identities = 470/484 (97 %), one gap (0 %)). Closest hits using the LSU sequence of CPC 38085 are Faurelina indica (strain CBS 301.78, GenBank GU180654.1; Identities = 903/910 (99 %), no gaps), Microsphaeropsis ononidicola (strain MFLUCC 15-0459, GenBank MG967668.1; Identities = 895/906 (99 %), no gaps), Coniothyrium hellebori (strain CBS 519.78, GenBank MH872934.1; Identities = 895/907 (99 %), no gaps), and Neoscirrhia matteucciicola (as Phoma matteuciicola; strain CBS 259.92, GenBank GU238100.1; Identities = 870/877 (99 %), no gaps) – also see Fig. 1. Closest hits using the rpb2 sequence of CPC 38085 had highest similarity to Neomicrosphaeropsis juglandis (strain MFLUCC 18-0795, GenBank MN593307.1; Identities = 773/882 (88 %), no gaps), Paraboeremia selaginellae (strain CBS 122.93, GenBank LT623255.1; Identities = 783/896 (87 %), two gaps (0 %)), and Ascochyta herbicola (voucher MFLU 15-2126, GenBank MT432200.1; Identities = 768/879 (87 %), two gaps (0 %)). Closest hits using the tub2 sequence of CPC 38085 had highest similarity to Neoscirrhia matteucciicola (as Phoma matteuciicola; strain CBS 259.92, GenBank GU237627.1; Identities = 264/289 (91 %), four gaps (1 %)), Didymella rabiei (strain AR628, GenBank KM244529.1; Identities = 488/547 (89 %), ten gaps (1 %)), Epicoccum sorghinum (strain HM19B, GenBank MN562463.1; Identities = 469/523 (90 %), nine gaps (1 %)), and Boeremia exigua (strain Ph.ex.001NY17, GenBank MK514090.1; Identities = 484/546 (89 %), 11 gaps (2 %)).

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Neoscytalidium dimidiatum (Penz.) Crous & Slippers, Stud. Mycol. 55: 244. 2006. Fig. 39.

Fig. 39.

Fig. 39.

Neoscytalidium dimidiatum (CPC 38666). A. Conidiomata on PNA. B. Conidiogenous cells. C. Fusicoccum-like condia. D–E. Phragmoconidia, with a few fusicoccum-like conidia. Scale bars = 10 μm.

Basionym: Torula dimidiata Penz., Michelia 2(no. 8): 466. 1887.

Conidiomata pycnidial, solitary, brown, erect, globose, 250–300 μm diam, with central ostiole and slightly papillate beak; wall of 4–8 layers of brown textura angularis. Conidiophores subcylindrical, hyaline, smooth, lining inner cavity, 0–2-septate, unbranched, 10–22 × 3.5–4.5 μm. Conidiogenous cells integrated, hyaline, smooth, subcylindrical, proliferating percurrently at apex, 8–12 × 3–3.5 μm. Conidia solitary, aseptate, hyaline, smooth, finely guttulate, subcylindrical to fusoid-ellipsoid, apex subobtuse, base truncate, 2 μm diam, (12–)13–14(–15) × (3.5–)4(–5) μm. Synasexual morph forming conidia via disarticulation of hyphal fragments, pale to medium brown, smooth, subcylindrical with truncate ends, 0(–1)-septate, 5–10(–12) × 3–5 μm.

Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse iron-grey.

Material examined: South Africa, Western Cape Province, Nieuwoudtville, epiphyte on stems of Aloidendron dichotomum (Asphodeloideae), 2018, P.W. Crous, HPC 3038 (CBS H-24434, culture CPC 38666 = CBS 146816).

Notes: Neoscytalidium dimidiatum is associated with tree cankers (Polizzi et al. 2011) but also causes chronic dermatomycosis and onychomycosis in humans, mainly involving foot infections (Dionne et al. 2015, Zhang et al. 2021a). Neoscytalidium dimidiatum is here newly reported from Aloidendron dichotomum in South Africa. Zhang et al. (2021a) recognised N. dimidiatum as the only species known from molecular data in this genus and also reduced two other species (N. novaehollandiae and N. orchidacearum) to synonyms of N. dimidiatum.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to N. novaehollandiae (now N. dimidiatum; strain CBS 122071, GenBank KF766207.1; Identities = 565/565 (100 %), no gaps), N. hyalinum (now N. dimidiatum; strain CBS 125699, GenBank MH863613.1; Identities = 563/563 (100 %), no gaps), and N. dimidiatum (voucher INBio 30A, GenBank KU204558.1; Identities = 555/557 (99 %), no gaps). Closest hits using the LSU sequence are N. novaehollandiae (now N. dimidiatum; strain CMW 26170, GenBank NG_059496.1; Identities = 872/872 (100 %), no gaps), N. hyalinum (now N. dimidiatum; strain CBS 125699, GenBank MH875087.1; Identities = 881/882 (99 %), one gap (0 %)), and N. orchidacearum (as Neoscytalidium sp. SKH-2016, now N. dimidiatum; strain MFLUCC 12-0533, GenBank KU179864.1; Identities = 869/871 (99 %), no gaps) – also see Fig. 1. Closest hits using the chs-1 sequence had highest similarity to N. hyalinum (as Scytalidium hyalinum, now N. dimidiatum; strain FMR 8342, GenBank FM211193.1; Identities = 284/284 (100 %), no gaps), N. dimidiatum (strain CBS 380.36, GenBank FM211196.1; Identities = 283/284 (99 %), no gaps), and N. novaehollandiae (now N. dimidiatum; strain D16GHEI, GenBank MH793720.1; Identities = 249/250 (99 %), no gaps). Closest hits using the tub2 sequence had highest similarity to N. novaehollandiae (now N. dimidiatum; strain Mlty_Ma02, GenBank MT362603.1; Identities = 278/278 (100 %), no gaps), N. dimidiatum (strain CBS 125695, GenBank KX464764.1; Identities = 426/427 (99 %), no gaps), and N. hyalinum (now N. dimidiatum; strain IRNHM-KZN1, GenBank MG220380.1; Identities = 420/422 (99 %), no gaps).

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Niesslia neoexosporioides Crous & R.K. Schumach., sp. nov. MycoBank MB 839295. Fig. 40.

Fig. 40.

Fig. 40.

Niesslia neoexosporioides (CPC 38177). A. Ascoma on host tissue. B–C. Ascomata with setae. D–F. Stipes with clavate vesicles. G. Conidiogenous cells. H. Conidia. Scale bars: A–B = 100 μm, C = 50 μm, all others = 10 μm.

Etymology: Name refers to the fact that it is similar to Niesslia exosporioides.

Ascomata perithecial, saprobic, single, gregarious, superficial, globose with a small, flattened base, black-brown, soft, thin-walled, setose, up to 100 μm diam. Setae stiff, pointed, unbranched, base enlarged or simple furcate, black, red-brown under the lens, thick- and smooth-walled, lumen tight. Peridium few-layered, of textura epidermoidea with red-brown, thick-walled and smooth cells. Paraphyses absent. Asci 8-spored, cigar-shaped to clavate, inoperculate, inamyloid (water plus Lugol), ascospores obliquely biseriate, overlapping. Ascospores narrowly ellipsoid, straight with conical ends, hyaline, thin-walled, smooth, medianly 1-septate, thin-walled, slightly constricted, eguttulate, dimensions undetermined, specimen overmature. Mycelium consisting of hyaline, smooth, septate, branched, 1–2 μm diam hyphae. Conidiophores reduced to conidiogenous cells, erect, flexuous, solitary to aggregated, subcylindrical, hyaline, smooth, thick-walled, 15–110 × 2–3 μm, phialidic with minute non-flared collarette, up to 1 μm long, apex 1–1.5 μm diam. Stipes terminating in clavate vesicles intermingled among conidiogenous cells, hyaline, smooth, with basal septum, up to 80 μm tall, vesicles 3–4 μm diam. Conidia solitary, aseptate, hyaline, smooth, subcylindrical, straight to slightly curved, apex subobtuse, base tapering to truncate hilum, 0.5 μm diam, (7–)8–10(–14) × 2(–2.5) μm.

Culture characteristics: Colonies flat, spreading, with sparse aerial mycelium and smooth, lobate margin, reaching 50 mm diam after 2 wk at 25 °C. On MEA surface rosy buff, reverse cinnamon; on PDA surface buff, reverse honey; on OA surface buff.

Typus: Germany, near Berlin, on dead leaves of Carex paniculata (Cyperaceae), 17 Apr. 2019, R.K. Schumacher, HPC 2983 = RKS 250 (holotype CBS H-24405, culture ex-type CPC 38177 = CBS 146810).

Notes: The specimen considered in this study was initially identified as Niesslia cf. exosporioides, which is common on Carex spp. in Europe. The sexual morph was overmature resulting in the identification relying on the asexual morph that developed in culture. Conidia of N. neoexosporioides [(7–)8–10(–14) × 2(–2.5) μm] are larger than those of N. exosporioides (5–6 × 1–1.5 μm; Gams et al. 2019). Phylogenetically, N. neoexosporioides is closely related to N. aemula (conidia 4.5–6.5 × 1.2–2 μm; Gams et al. 2019), but the latter has smaller conidia. A phylogenetic species tree is presented as Fig. 41.

Fig. 41.

Fig. 41.

The single most parsimonious tree obtained from a phylogenetic analysis of the Niesslia ITS nucleotide alignment. The tree was rooted to Trichoderma atroviride (strain CBS 142.95, GenBank MH862505.1) and the scale bar indicates the number of changes. Parsimony bootstrap support values higher than 79 % are shown at the nodes and the treated species is highlighted with a coloured box and bold text. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. Alignment statistics: 43 strains including the outgroup; 608 characters including alignment gaps analysed: 292 constant, 75 variable and parsimony-uninformative and 241 parsimony-informative. Tree statistics: TL = 788, CI = 0.688, RI = 0.900, RC = 0.619.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to “Hypocreaceae sp.” (strain MUT<ITA> 2463, GenBank MG813207.1; Identities = 497/498 (99 %), no gaps), Niesslia aemula (strain AX-GC-5, GenBank MK592746.1; Identities = 561/568 (99 %), one gap (0 %)), and Niesslia curvisetosa (as Monocillium curvisetosum; strain CBS 660.94, GenBank NR_160195.1; Identities = 529/540 (98 %), three gaps (0 %)). Closest hits using the LSU sequence are Niesslia luzulae (as Niesslia exosporioides; strain CBS 691.71, GenBank MH872060.1; Identities = 815/816 (99 %), one gap (0 %)), “Hypocreaceae sp.” (strain MUT<ITA> 2463, GenBank MG816499.1; Identities = 810/812 (99 %), two gaps (0 %)), and Niesslia curvisetosa (as Monocillium curvisetosum; strain CBS 660.94, GenBank MH874141.1; Identities = 888/893 (99 %), one gap (0 %)) – also see Fig. 4. No significant hits were obtained when the actA , tef1 and tub2 sequences were used in blastn and megablast searches.

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Nothoanungitopsis Crous, gen. nov. MycoBank MB 839296.

Etymology: Name refers to its morphological similarity to Anungitopsis.

Mycelium consisting of brown, finely verruculose, septate, branched hyphae. Conidiophores dimorphic. Microconidiophores erect, flexuous, brown, thick-walled, septate. Macroconidiophores erect, flexuous, brown, finely verruculose, unbranched, thick-walled, base swollen, with rhizoids, multi-septate. Conidiogenous cells terminal, integrated, medium brown, finely verruculose, with several sympodially arranged flat-tipped denticles, scars unthickened. Conidia solitary or in chains of two, fusoid, 3-septate, straight, apex subobtuse, base subobtuse with truncate hilum; two central cells brown, thick-walled, finely verruculose; end cells subhyaline, smooth, thin-walled.

Type species: Nothoanungitopsis urophyllae Crous

Nothoanungitopsis urophyllae Crous, sp. nov. MycoBank MB 839297. Fig. 42.

Fig. 42.

Fig. 42.

Nothoanungitopsis urophyllae (CPC 38059). A. Conidiophores on PNA. B–D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars: A = 80 μm, all others = 10 μm.

Etymology: Name refers to the host species from which it was isolated, Eucalyptus urophylla.

Mycelium consisting of brown, finely verruculose, septate, branched, 2–3 μm diam hyphae. Conidiophores dimorphic. Microconidiophores erect, flexuous, brown, thick-walled, 50–80 × 4–6 μm, 2–3-septate. Macroconidiophores erect, flexuous, brown, finely verruculose, unbranched, thick-walled, base swollen, 8–12 μm diam, with rhizoids, 5–18-septate, 100–350 × 5–8 μm. Conidiogenous cells terminal, integrated, medium brown, finely verruculose, 15–35 × 4–5 μm, with several sympodially arranged flat-tipped denticles, scars 2–3 μm diam, unthickened. Conidia solitary or in chains of two, fusoid, 3-septate, straight, apex subobtuse, base subobtuse with truncate hilum, 1–1.5 μm diam; two central cells brown, thick-walled, finely verruculose; end cells subhyaline, smooth, thin-walled, (15–)16–18(–20) × (4–)5(–6) μm.

Culture characteristics: Colonies erumpent, spreading, with sparse to moderate aerial mycelium and uneven margin, reaching 5 mm diam after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse isabelline.

Typus: South Africa, Kwazulu-Natal Province, Kwambonambi, on seed capsules of Eucalyptus urophylla (Myrtaceae), Apr. 2017, M.J. Wingfield, HPC 2868 (holotype CBS H-24395, culture ex-type CPC 38059 = CBS 146799).

Notes: Neoanungitea was introduced to accommodate two species occurring on eucalypt leaf litter in Australia, and characterised by having a rachis with flat-tipped sympodial loci, similar to but less conspicuous than those of Anungitea (Crous et al. 2019d). Nothoanungitopsis adds a third genus to this complex. Although it has unthickened conidiophore scars and conidial hila as in Anungitopsis, it is distinguished by lacking globose, brown swellings in its conidiophores, and having conidia that are unevenly pigmented, with two brown central cells.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Anungitopsis speciosa (strain CBS 181.95, GenBank EU035401.1; Identities = 344/413 (83 %), 38 gaps (9 %)), Anungitopsis lauri (strain CBS 145067, GenBank NR_161129.1; Identities = 331/399 (83 %), 37 gaps (9 %)), and Neoanungitea eucalypti (strain CBS 143173, GenBank MG386031.2; Identities = 312/378 (83 %), 26 gaps (6 %)). Closest hits using the LSU sequence are Neoanungitea eucalypti (strain CBS 143173, GenBank MG386031.2; Identities = 547/588 (93 %), one gap (0 %)), Anungitopsis speciosa (strain CBS 181.95, GenBank EU035401.1; Identities = 542/588 (92 %), one gap (0 %)), and Spirosphaera beverwijkiana (strain CBS 469.66, GenBank HQ696657.1; Identities = 541/588 (92 %), one gap (0 %)) – also see Fig. 1.

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Nothomicrosphaeropsis Crous, gen. nov. MycoBank MB 839298.

Etymology: Named after its morphological similarity to Microsphaeropsis.

Conidiomata pycnidial, solitary, erumpent, brown, papillate with central ostiole; wall of 3–6 layers of brown textura angularis. Conidiophores reduced to conidiogenous cells lining the inner cavity, ampulliform to doliiform, hyaline, smooth. Conidia solitary, hyaline, smooth, aseptate, granular, subcylindrical, apex obtuse, base bluntly rounded, straight to slightly curved, becoming pale brown with age.

Type species: Nothomicrosphaeropsis welwitschiae Crous

Nothomicrosphaeropsis welwitschiae Crous, sp. nov. MycoBank MB 839299. Fig. 43.

Fig. 43.

Fig. 43.

Nothomicrosphaeropsis welwitschiae (CPC 38879). A. Conidiomata on PNA. B. Pycnidial conidioma. C–D. Conidiogenous cells. E. Conidia. Scale bars: A–B = 250 μm, all others = 10 μm.

Etymology: Named after the host genus it was collected from, Welwitschia.

Conidiomata pycnidial, solitary, erumpent, brown, 200–250 μm diam, papillate with central ostiole; wall of 3–6 layers of brown textura angularis. Conidiophores reduced to conidiogenous cells lining the inner cavity, ampulliform to doliiform, hyaline, smooth, 4–6 × 3–4 μm. Conidia solitary, hyaline, smooth, aseptate, granular, subcylindrical, apex obtuse, base bluntly rounded, straight to slightly curved, becoming pale brown with age, (4–)5–6(–7) × 2–2.5 μm.

Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse olivaceous grey.

Typus: Namibia, Hope Mine, east of Gobabeb – Namib Research Institute, on dead leaves of Welwitschia mirabilis (Welwitschiaceae), 19 Nov. 2019, P.W. Crous, HPC 3126 (holotype CBS H-24448, culture ex-type CPC 38879 = CBS 146829).

Notes: Based on ITS/LSU DNA sequence data, the present collection was initially identified as Macroventuria wentii, which is known from plant litter in the Nevada Death Valley, USA (van der Aa 1971). However, by incorporating more informative genes such as rpb2 and tub2 in the analysis (see phylogenetic tree, Fig. 44), the present isolate proved allied to Paramicrosphaeropsis, Neomicrosphaeropsis and Microsphaeropsis, a generic complex in Didymellaceae (Hou et al. 2020a). Genera in this complex are morphologically comparable in initially producing hyaline conidia that turn pale brown at maturity.

Fig. 44.

Fig. 44.

Consensus phylogram (50 % majority rule) obtained from the maximum likelihood analysis with IQ-TREE showing the phylogenetic position of Nothomicrosphaeropsis welwitschiae gen. et sp. nov. based on the multigene (ITS / LSU / rpb2 / tub2) nucleotide alignment. The most basal branch was halved to facilitate layout. Bootstrap support values (> 79 %) from 5 000 ultrafast bootstrap replicates are shown at the nodes. The alignment is derived from the combined alignment of Hou et al. (2020a) and GenBank accession numbers can be obtained from the same reference. Culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Neocucurbitaria aquatica (culture CBS 297.74) and the species treated in this study is highlighted with a coloured block and bold face. Alignment statistics: 67 strains including the outgroup; 2 382 characters including alignment gaps analysed: 625 distinct patterns, 505 parsimony-informative, 94 singleton sites, 1 783 constant sites. The best models identified in IQ-TREE were: TIM2e+I+G4 (ITS), TNe+I+G4 (LSU), TNe+I+G4 (rpb2), TIM2e+I+G4 (tub2).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to “Aff. Phoma sp. (strain mk175, GenBank KT150686.1; Identities = 530/530 (100 %), no gaps), Macroventuria wentii (strain CBS 526.71, GenBank MH860250.1; Identities = 538/540 (99 %), no gaps), and Leptosphaerulina australis (strain CBS 234.58, GenBank MH857766.1; Identities = 538/542 (99 %), one gap (0 %)). Closest hits using the LSU sequence are Ascochyta rabiei (strain CBS 237.37, GenBank NG_069312.1; Identities = 858/858 (100 %), no gaps), Coniothyrium genistae (strain CBS 294.74, GenBank MH872596.1; Identities = 860/861 (99 %), no gaps), Ascochyta fabae (strain CBS 649.71, GenBank MH872045.1; Identities = 860/861 (99 %), no gaps), and Macroventuria wentii (strain CBS 526.71, GenBank NG_069040.1; Identities = 856/858 (99 %), no gaps) – also see Fig. 1. Closest hits using the rpb2 sequence had highest similarity to Neomicrosphaeropsis italica (strain IT_24211, GenBank KU714604.1; Identities = 693/744 (93 %), no gaps), Neomicrosphaeropsis juglandis (strain MFLUCC 18-0795, GenBank MN593307.1; Identities = 693/744 (93 %), no gaps), and Neomicrosphaeropsis cytisi (strain MFLUCC 13-0396, GenBank KX572355.1; Identities = 682/742 (92 %), no gaps). A direct blast2 search against Macroventuria rpb2 sequences on GenBank had highest similarity to Macroventuria anomochaeta (strain CBS 525.71, GenBank GU456346.1; Identities = 663/741 (89 %), no gaps), Macroventuria terrestris (strain RMF CA 12, GenBank MT018194.1; Identities = 531/596 (89 %), no gaps), and Macroventuria wentii (strain CBS 527.71C, GenBank MN983625.1; Identities = 530/596 (89 %), no gaps). Closest hits using the tub2 sequence had highest similarity to Paramicrosphaeropsis ellipsoidea (strain CBS 197.97, GenBank MT005680.1; Identities = 314/333 (94 %), no gaps), Epicoccum sorghinum (strain FDY-5, GenBank MT799852.1; Identities = 336/364 (92 %), no gaps), Epicoccum latusicollum (strain HH12, GenBank MN623288.1; Identities = 337/365 (92 %), two gaps (0 %)), and Macroventuria wentii (strain CBS 526.71, GenBank GU237546.1; Identities = 302/333 (91 %), no gaps). A direct blast2 search against Macroventuria tub2 sequences on GenBank had highest similarity to Macroventuria wentii (strain CBS 877.70, GenBank MN984003.1; Identities = 303/333 (91 %), no gaps), Macroventuria terrestris (strain CBS 127771, GenBank MT005653.1; Identities = 300/331 (91 %), one gap (0 %)), and Macroventuria anomochaeta (strain CBS 525.71, GenBank GU237545.1; Identities = 300/333 (90 %), no gaps).

Authors: P.W. Crous, D.A. Cowan, G. Maggs-Kölling, E. Marais, N. Yilmaz & M.J. Wingfield

Ophioceras leptosporum (S.H. Iqbal) J. Walker, Mycotaxon 11: 62. 1980. Fig. 45.

Fig. 45.

Fig. 45.

Ophioceras leptosporum (CPC 39147). A. Ascomata developing on OA. B–E. Asci with ascospores. Scale bars: A = 250 μm, all others = 10 μm.

Basionym: Gaeumannomyces leptosporus S.H. Iqbal, Trans. Brit. Mycol. Soc. 58: 346. 1972.

Description and illustration: See Walker (1980).

Material examined: South Africa, Northern Province, Nelspruit, Buffelskloof Nature Reserve, on twigs of Syzygium cordatum (Myrtaceae), Nov. 2018, P.W. Crous, HPC 3149, CBS H-24542, cultures CPC 39147 = CBS 147090.

Notes: Species of Ophioceras are commonly isolated from leaf and twig litter in freshwater habitats worldwide (Shearer et al. 1999). This is apparently the first record of this species from South Africa.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Ophioceras leptosporum (strain CBS 894.70, GenBank NR_111768.1; Identities = 506/511 (99 %), no gaps), Pestalotiopsis mangifolia (voucher INBio 183C, GenBank KU204541.1; Identities = 499/543 (92 %), 17 gaps (3 %)), and Myceliophthora sepedonium (voucher INBio 571C, GenBank KU204432.1; Identities = 499/543 (92 %), 17 gaps (3 %)). Closest hits using the LSU sequence are Ophioceras leptosporum (strain CBS 894.70, GenBank NG_057959.1; Identities = 868/869 (99 %), no gaps), Ophioceras chiangdaoense (strain CMU 26633, GenBank NG_066356.1; Identities = 829/842 (98 %), no gaps), and Ophioceras commune (strain M91, GenBank JX134687.1; Identities = 816/873 (93 %), eight gaps (0 %)) – also see Fig. 5. Closest hits using the tef1 (second part) sequence had highest similarity to Ophioceras leptosporum (strain CBS 894.70, GenBank JX134704.1; Identities = 474/475 (99 %), no gaps), Ophioceras commune (strain M91, GenBank JX134701.1; Identities = 434/475 (91 %), no gaps), and Ophioceras aquaticus (voucher S-479, GenBank MN194063.1; Identities = 404/447 (90 %), no gaps).

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Paracremonium bendijkiorum Hern.-Restr., sp. nov. MycoBank MB838709. Fig. 46.

Fig. 46.

Fig. 46.

Paracremonium bendijkiorum (CBS 147288). A. Colony on PDA. B. Colony on MEA. C. Conidiophores. D. Chlamydospore. E. Conidia and conidiogenous cell. F–G. Conidia. Scale bars C–F =10 μm, G = 5 μm.

Etymology: Named for the collectors Yvan Bentem and Jack van Dijken students from the Het Hogeland College (Warffum, the Netherlands). This sample was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae septate, hyaline, smooth and thin-walled, 1–4 μm wide. Conidiophores semi-macronematous, straight or slightly bent, simple or irregularly branched, hyaline, smooth-walled. Conidiogenous cells 13–33 × 1.5–3 μm, 1–1.5 μm wide at the apex, monophialidic, subcylindrical, straight or flexuous, hyaline, thin- and smooth-walled, with subcylindrical collarette. Conidia solitary, aseptate, subglobose, with apiculate base, 3–6 μm diam, hyaline, smooth-walled. Chlamydospores solitary, terminal or intercalary, hyaline, subglobose 6–7 μm diam. Sexual morph not observed.

Culture characteristics: Colonies after 2 wk at 25 °C on OA reaching 20 mm diam, flat, with sparse aerial mycelium, concolourous with the agar. On MEA reaching 18 mm diam, elevated, funiculose, wet, pale flesh, white, margin entire, regular. On PDA reaching 20 mm diam, elevated in the centre, funiculose to cottony, white to salmon, membranous to the periphery, margin entire.

Typus: Netherlands, Groningen Province, Warffum, Wibrand Geertstraat, from soil, 6 Jun. 2019, Y. Bentem & J. van Dijken, NL19_024 (holotype CBS H-24734, culture ex-type CBS 147228 = NL19_24005).

Notes: Paracremonium species are known from soil close to streams, caves, alkaline mud, air, beetles, and humans (Lombard et al. 2015, Lynch et al. 2016, Crous et al. 2017, Zhang et al. 2017, Al-Bedak et al. 2019, Zhang et al. 2021b). Paracremonium bendijkiorum is distinguished from its phylogenetically closest species, P. inflatum, in having subglobose conidia and chlamydospores. Another species in the genus with similar subglobose conidia is P. apiculatum (Zhang et al. 2020), which is phylogenetically distant (Fig. 47). Also see the overview phylogeny (Fig. 4).

Fig. 47.

Fig. 47.

Maximum composite likelihood tree obtained from the RAxML analysis of the combined ITS and LSU sequence alignment of Paracremonium species. Bootstrap support values above 70 % are shown at the nodes. The novel species is indicated in a coloured block and bold face. The tree was rooted to Cosmospora arxii (CBS 748.69), C. stegonsporii (CBS 122305) and Xenoacremonium recifei (CBS 137.35). Alignment statistics: 12 strains including the outgroup; 1 469 characters including alignment gaps analysed (ITS: 583, LSU: 886). Model: GTR+G, alignment patterns: 213.

Based on a megablast search of NCBI nucleotide database, the closest hits using the ITS sequence had the highest similarity to Paracremonium inflatum (strain CBS 485.77 GenBank NR_154312.1; Identities = 434/451 (96 %), three gaps (0 %)), Paracremonium variiforme strain LC5837 (GenBank KU746694.1; Identities = 426/444 (96 %), two gaps (0 %)) and Paracremonium binnewijzendii (strain CBS 143277, GenBank NR_157491.1; Identities = 431/454 (95 %), four gaps (0 %)). The closest hits using the LSU sequence had the highest similarity to Paracremonium variiforme (strain LC5832, GenBank KU746739.1; Identities = 825/842 (98 %), two gaps (0 %)), Paracremonium sp. (strain FZ2855-2, GenBank MK329029.1; Identities = 825/843 (98 %), two gaps (0 %)) and Paracremonium sp. (strain FZ3546-2, GenBank MK329031.1; Identities = 823/843 (98 %), two gaps (0 %)) – also see Fig. 4. The closest hits using the rpb2 sequence are Paracremonium variiforme (strain CGMCC 3.17934 (GenBank KY883249.1; Identities = 583/654 (89 %), no gaps), Paracremonium pembeum (strain UCR2995, GenBank KT936355.1; Identities = 574/655 (88 %), two gaps (0 %)) and Paracremonium contagium (strain CBS 110348, GenBank KM232396.1; Identities = 573/655 (87 %), two gaps (0 %)). The closest hits using the tef1 sequence had the highest similarity to Paracremonium sp. (strain LC12502, GenBank MK336059.1; Identities = 825/858 (96 %), no gaps), Paracremonium variiforme (strain LC5837, GenBank KX855240.1; Identities = 824/858 (96 %), no gaps) and Paracremonium sp. (strain LC12552, GenBank MK336061.1; Identities = 807/858 (94 %), no gaps). The closest hits using the tub2 sequence had the highest similarity to Paracremonium variiforme (strain LC5837, GenBank KU746786.1; Identities = 232/243 (95 %), no gaps), Paracremonium sp. (strain LC12502 (GenBank MK336137.1; Identities = 228/239 (95 %), no gaps) and Paracremonium sp. (strain LC12552, GenBank MK336139.1; Identities = 225/240 (94 %), no gaps).

Author: M. Hernández-Restrepo

Paraphoma ledniceana Spetik, Eichmeier & Berraf-Tebbal, sp. nov. MycoBank MB835802. Fig. 48.

Fig. 48.

Fig. 48.

Paraphoma ledniceana (CBS 146533). A. Conidiomatal pycnidia on SNA. B–C. Squash of conidiomata. D. Conidia. Scale bars: A = 100 μm, all others = 10 μm.

Etymology: Name refers to Lednice, where the specimen was collected.

Conidiomata pycnidial, ostiolate, uni- or bilocular, submerged, obpyriform, semi-pilose, dark brown, 250–450 μm diameter. Conidiophores lining the inner cavity, pale brown, smooth, densely aggregated. Conidia solitary, hyaline, straight, subcylindrical, apex obtuse, base truncate, (3.7–)4.5–4.9(–5.7) × (1.2–)1.5–1.7(–2.4) μm (av. 4.7 ± 0.5 × 1.6 ± 0.3 μm).

Culture characteristics: Colonies on PDA reaching 38 mm diam after 10 d at 25 °C, margin regular, floccose, erumpent, pale grey; reverse grey. On MEA reaching 27 mm diam after 10 d, margin regular, floccose, white to dirty white; reverse pale brown. On OA reaching 42 mm diam after 10 d, margin regular, floccose, white in outer ring, olivaceous grey; reverse olivaceous. No growth at 10 °C and 37 °C was observed.

Typus: Czech Republic, Breclav, Lednice, isolated as saprobe on dead wood of Buxus sempervirens (Buxaceae), Sep. 2018, M. Spetik (holotype BRNU 673830, isotype BRNU 673831, culture ex-type CBS 146533 = MEND-F-82).

Notes: Based on multigene (ITS, tef1 and tub2) phylogenetic analyses, P. ledniceana differs from P. rhaphiolepisis by 29 nucleotides positions in the concatenated alignment, in which 10 were distinct in the ITS region, six in the tef1 region and 13 in tub2. Morphologically, the conidia length of P. ledniceana are smaller [(3.7–)4.5–4.9(–5.7) × (1.2–)1.5–1.7(–2.4) μm] compared to those of P. rhaphiolepisis [(4.5–)5–6(–6.5) × 2(–2.5) μm]. The obtained phylogeny (Fig. 49) placed the isolate CBS 146533 as a lineage sister to Paraphoma rhaphiolepisis. However, it is clearly distinct from the other known species of Paraphoma and it is therefore considered as a taxonomic novelty.

Fig. 49.

Fig. 49.

Consensus phylogram (50 % majority rule) obtained from the maximum likelihood analysis with IQ-TREE of the Paraphoma multigene (ITS / tef1 / tub2) nucleotide alignment. Bootstrap support values (> 79 %) from 5 000 ultrafast bootstrap replicates are shown at the nodes. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Neosetophoma samarorum (culture CBS 138.96) and the species treated in this study are highlighted with coloured blocks and bold face. Alignment statistics: 32 strains including the outgroup; 1 227 characters including alignment gaps analysed: 477 distinct patterns, 299 parsimony-informative, 100 singleton sites, 828 constant sites. The best models identified in IQ-TREE were: HKY+F+G4 (ITS), TNe+G4 (tef1), TNe+I+G4 (tub2).

Based on a megablast search of NCBI nucleotide database, the closest hits using the ITS sequence had the highest similarity to Paraphoma chrysanthemicola (GenBank MK647980.1; Identities = 509/510 (99 %), one gap (0 %)), Uncultured fungus clone (GenBank MT236451.1; Identities = 509/510 (99 %), one gap (0 %)) and Paraphoma sp. (GenBank MK304128.1; Identities = 506/506 (100 %), no gaps). The closest hits using the LSU sequence had the highest similarity to Paraphoma sp. (GenBank LC126021.2; Identities = 1 043/1 045 (99 %), no gaps), Paraphoma radicina (GenBank NG_070446.1; Identities = 1 043/1 045 (99 %), no gaps) and Paraphoma chrysanthemicola (GenBank GQ387582.1; Identities = 1 043/1 045 (99 %), no gaps) – also see Fig. 1; closest hits using the rpb2 sequence are Paraphoma rhaphiolepidis (GenBank KY979851.1; Identities = 815/844 (97 %), no gaps), Paraphoma radicina (GenBank LT796995.1; Identities = 783/861 (91 %), no gaps) and Paraphoma sp. (GenBank MG779463.1; Identities = 625/733 (85 %), eight gaps (1 %)). The closest hits using the tef1 sequence had the highest similarity to Paraphoma rhaphiolepidis (GenBank KY979896.1; Identities = 269/283 (95 %), three gaps (0 %)), Paraphoma chlamydocopiosa (GenBank KX376304.1; Identities = 192/226 (85 %), no gaps) and Paraphoma chlamydocopiosa (GenBank KX376299.1; Identities = 192/226 (85 %), no gaps). The closest hits using the tub2 sequence had the highest similarity to Paraphoma rhaphiolepidis (GenBank KY979924.1; Identities = 460/470 (98 %), three gaps (0 %)), Paraphoma sp. (GenBank MG779453.1; Identities = 433/473(99 %), no gaps) and Paraphoma cf. “convolvuli” (GenBank MG779461.1; Identities = 431/473 (91 %), three gaps (0 %)).

Authors: M. Spetik, A. Eichmeier & A. Berraf-Tebbal

Paraphoma salicis Crous & Akulov, sp. nov. MycoBank MB 839300. Fig. 50.

Fig. 50.

Fig. 50.

Paraphoma salicis (CPC 38651). A. Conidiomata on OA. B. Conidioma with ostiole. C–D. Conidiogenous cells. E. Conidia. Scale bars: A–B = 200 μm, all others = 10 μm.

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

Conidiomata erumpent, solitary or aggregated, brown, globose, glabrous, 180–200 μm diam, with central ostiole, exuding creamy conidial mass. Conidiophores reduced to conidiogenous cells lining inner cavity, doliiform to ampulliform, hyaline, smooth, phialidic, 5–6 × 3–4 μm. Conidia solitary, aseptate, hyaline, smooth, ellipsoid to subcylindrical, apex obtuse, base bluntly rounded to somewhat truncate, (4–)5(–6) × (2–)2.5(–3) μm.

Culture characteristics: Colonies flat, spreading, with sparse to moderate aerial mycelium and smooth, lobate margin, reaching 35 mm diam after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse olivaceous grey.

Typus: Ukraine, Volyn region, Liubeshiv district, Liubìaz village, on leaves of Salix cf. alba (Salicaceae), co-ocurring with Septoria salicis Westend., 29 Jul. 2019, A. Akulov, CWU AS 7121, HPC 3011 (holotype CBS H-24431, culture ex-type CPC 38651 = CBS 146797).

Notes: Paraphoma (Phaeosphaeriaceae) was resurrected as a distinct genus by de Gruyter et al. (2010), and shown to accommodate several species of saprobic, endophytic or phytopathogenic fungi affecting agricultural crops worldwide (Moslemi et al. 2016). Paraphoma salicis is phylogenetically distinct from Paraphoma species presently known from their DNA, and represents a new taxon occurring on leaves of Salix in Ukraine (Fig. 49).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Paraphoma radicina (as Phoma radicina; strain VB1-2, GenBank MK764998.1; Identities = 520/587 (89 %), 23 gaps (3 %)), Plenodomus biglobosus (as Leptosphaeria biglobosa; strain CM-02, GenBank KY221834.1; Identities = 518/586 (88 %), 21 gaps (3 %)), and Paraphoma chrysanthemicola (strain DSM100401_C29_RLCS22, GenBank MT453289.1; Identities = 519/588 (88 %), 24 gaps (4 %)). Closest hits using the LSU sequence are Paraphoma dioscoreae (strain CBS 135100, GenBank KF251671.1; Identities = 806/809 (99 %), no gaps), Paraphoma radicina (strain UTHSC DI16-209, GenBank LN907352.1; Identities = 826/830 (99 %), no gaps), and Plectosphaerella cucumerina (strain 17chu05-05, GenBank MT102906.1; Identities = 810/814 (99 %), no gaps) – also see Fig. 1. Closest hits using the actA sequence had highest similarity to Parastagonosporella fallopiae (strain CBS 135981, GenBank MH460537.1; Identities = 217/241 (90 %), four gaps (1 %)), Paraphoma chrysanthemicola (strain ICMP 10745, GenBank KT309135.2; Identities = 206/242 (85 %), eight gaps (3 %)), and Paraphoma radicina (strain ICMP 6623, GenBank KT309298.2; Identities = 200/243 (82 %), ten gaps (4 %)). Closest hits using the rpb2 sequence had highest similarity to Paraphoma melnikii (as Paraphoma sp. MG-2018a; voucher MF-9.240, GenBank MG779464.1; Identities = 668/735 (91 %), no gaps), Paraphoma cf.convolvuli” (voucher MF-9.298.1, GenBank MG779468.1; Identities = 660/735 (90 %), no gaps), and Paraphoma fimeti (strain UTHSC DI16-296, GenBank LT797032.1; Identities = 817/944 (87 %), five gaps (0 %)). Closest hits using the tub2 sequence had highest similarity to Paraphoma cf. “convolvuli” (voucher MF-9.300.1, GenBank MG779460.1; Identities = 409/439 (93 %), no gaps), Paraphoma melnikiae (voucher MF-9.240, GenBank MG779453.1; Identities = 409/440 (93 %), two gaps (0 %)), and Paraphoma rhaphiolepidis (strain CBS 142524, GenBank KY979924.1; Identities = 413/462 (89 %), eight gaps (1 %)).

Authors: P.W. Crous & A. Akulov

Parasarocladium wereldwijsianum Hern.-Restr., sp. nov. MycoBank MB 838710. Fig. 51.

Fig. 51.

Fig. 51.

Parasarocladium wereldwijsianum (CBS 147223). A–G. Conidiophores and conidia. H. Conidia. I. Colony on MEA. J. Colony on PDA. K. Colony on OA. Scale bars A–H = 10 μm.

Etymology: Named after the school “Wereldwijs” (Bilthoven, the Netherlands) where the sample was collected. This sample was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae septate, hyaline, smooth and thin-walled, 1.5–2.5 μm wide. Conidiophores erect, arising directly from vegetative hyphae or ropes of hyphae, straight or slightly bent, simple or branched, sometimes verticillate, hyaline, smooth-walled. Phialides subulate, 11.5–45 μm long, 1.5–3.5 μm wide at the base, 1–1.5 μm wide at the apex, straight or flexuous, hyaline, thin- and smooth-walled. Conidia arranged in slimy heads, unicellular, ellipsoidal to obovoid, base truncated, 4–10 × 2–3 μm, hyaline, smooth-walled. Chlamydospores and sexual morph not observed.

Culture characteristics: After 7 d at 25 °C on OA attaining 30–35 mm diam, flat, sparse aerial mycelium, buff to white salmon, wet with exudate (hyaline and salmon), margin effuse; reverse no change. On MEA attaining 30 mm flat, velvety, radially folded, peach to rosy buff; reverse saffron. On PDA attaining 30–45 mm diam, flat, velvety, sparse aerial mycelium, olivaceous buff, margin irregular; reverse greenish olivaceous.

Typus: Netherlands, Utrecht Province, Bilthoven, from soil, 6 Jun. 2019, S. Frederikze, J. Mes & S. Maghnouji, NL19_095 (holotype CBS H-24735, culture ex-type CBS 147226 = NL19095011).

Other specimens examined: Netherlands, Utrecht, Bilthoven, from soil, 6 Jun. 2019, S. Frederikze, J. Mes, & S. Maghnouji, NL19_094 (culture CBS 147223 = NL1994001), ibid. (culture CBS 147224 = NL19094011).

Notes: Parasarocladium wereldwijsianum is represented by three strains collected in two soil samples from Bilthoven (Utrecht Province, the Netherlands). Parasarocladium wereldwijsianum can be distinguished from the phylogenetically closest species P. aestuarinum and P. alavariense by having olivaceous buff colonies on PDA, conidiophores that are irregularly branched, and a distinct conidial morphology [colonies buff orange, conidia globose, 3.5–4.5 × 2.5–3.5 μm in P. aestuarinum; colonies dark ochre yellow, conidia subglobose, 4.5–5.5 × 3–4 μm in P. alavariense; Gonçalves et al. 2019]. Furthermore, P. aestuarinum and P. alavariense are known from marine habitats (Gonçalves et al. 2019) while P. wereldwijsianum was isolated from soil. Also see the overview phylogeny (Fig. 4) and species phylogeny (Fig. 52).

Fig. 52.

Fig. 52.

Maximum composite likelihood tree obtained from the RAxML analysis of the combined ITS, LSU and actin sequence alignment of Parasarocladium species. Bootstrap support values above 70 % are shown at the nodes. The novel species is indicated in a coloured block and bold face. The tree was rooted to Leucosphaerina arxii (CBS 737.84) and Xenoacremonium recifei (CBS 137.35). Alignment statistics: 15 strains including the outgroup; 1 739 characters including alignment gaps analysed (ITS: 600, LSU: 851, act: 288). Model GTR+I+G, 395 distinct patterns.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Hypocreales sp. (strain HWJ2(1) , GenBank KM268693.1; Identities = 578/579 (99 %), no gaps), Fungal sp. (strain PH30517, GenBank KR363031.1; Identities = 496/497 (99 %), no gaps), and Acremonium sp. (GenBank MK651581.1; Identities = 480/481 (99 %) no gaps). Closest hits using the LSU sequence are Acremonium sp. (strain 39 OA-2013, GenBank JX535073.1; Identities = 490/490 (100 %), no gaps), Acremonium sp. (strain 38 OA-2013, GenBank JX535071.1; Identities = 490/490 (100 %), no gaps), and Acremonium sp. (strain 31 OA-2013, GenBank JX535057.1; Identities = 490/490 (100 %), no gaps) – also see Fig. 4.

Author: M. Hernández-Restrepo

Peziza ligni Crous & Decock, sp. nov. MycoBank MB 839301. Fig. 53.

Fig. 53.

Fig. 53.

Peziza ligni (CPC 39110). A. Conidiophores on OA. B–E. Conidiogenous cells giving rise to conidia. F. Conidia. Scale bars = 10 μm.

Etymology: Name refers to the woody substrate (L. = lignum) from which it was isolated.

Mycelium consisting of hyaline, smooth, septate, branched, 4–6 μm diam. Conidiophores solitary, subcylindrical, erect, flexuous, hyaline, smooth, 3–5-septate, 100–200 × 7–8 μm. Conidiogenous cells integrated, terminal, globose to sphaeropedunculate, 22–35 μm diam, covered in numerous denticles, 1–2 × 1 μm, apex with minute marginal frill. Conidia solitary, aseptate, fusoid-ellipsoid, pale brown, smooth to finely roughened, apex subobtuse, base truncate, 1–2 μm diam, with marginal frill, (11–)13–14(–16) × 6(–7) μm.

Culture characteristics: Colonies spreading, with moderate aerial mycelium and smooth, lobate margin, covering dish after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse buff.

Typus: France, Normandie, Seine Maritime, on masonry and plastering, near a wooden staircase, Nov. 2019, C. Decock (holotype CBS H-24559, culture ex-type MUCL 57889 = CPC 39110 = CBS 146637). 

Notes: Peziza, the largest genus in the Pezizales, has been linked to Ostracoderma and Oedocephalum asexual morphs (Hanlin 1997). However, Peziza is paraphyletic (Hansen et al. 2001, 2002, 2005). The type of Oedocephalum, O. elegans, has been linked to Peziza anthracophila. The type of Peziza, P. vesiculosa, has however been linked to Oedocephalum pallidum, a species that normally occurs on dung of various animals (Stalpers 1974). Oedocephalum is characterised by erect, hyaline conidiophores with a terminal swelling or vesicle that gives rise to hyaline, aseptate conidia attached to the vesicle via a layer of denticles. Species of Oedocephalum are common on dung, wood, soil, and plant litter.

Using the key of Stalpers (1974), P. ligni was similar to O. argillaceum (decayed branch, Morocco; conidia hyaline, narrowly ellipsoidal, 12.5–15(–16) × 5.3–6.3(–6.5) μm, covered in brownish granules, distinctly larger at the ends, up to 2 μm long). Peziza ligni can, however, be distinguished from O. argillaceum by having conidia that are smooth to finely roughened, and in having smaller conidiogenous vesicles (22–35 μm diam), vs. 35–52 μm diam in O. argillaceum. A phylogenetic tree is presented as Fig. 54.

Fig. 54.

Fig. 54.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Peziza ITS nucleotide alignment. Bayesian posterior probabilities (> 0.79) are shown at the nodes and the scale bar represents the expected changes per site. Families are indicated with coloured blocks to the right of the tree. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Stouffera gilkeyae (voucher KIPD_04; GenBank MN857199.1) and the novel species treated in this study is indicated in a coloured block and in bold face. Alignment statistics: 44 strains including the outgroup; 486 unique site patterns. Tree statistics: 19 502 sampled trees from 65 000 generations.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Peziza pseudoviolacea (voucher 16504, GenBank JF908564.1; Identities = 507/561 (90 %), 11 gaps (1 %)), Peziza emileia (voucher L 0833270, GenBank KJ728716.1; Identities = 333/385 (86 %), 14 gaps (3 %)), and Peziza fimeti (voucher TAAM 171114, GenBank JQ654494.1; Identities = 327/385 (85 %), 14 gaps (3 %)). Closest hits using the LSU sequence are Peziza exogelatinosa (voucher KH 00.029 (C), GenBank AY500545.1; Identities = 874/893 (98 %), no gaps), Peziza violacea f. terricola (voucher MPU JCD 466-77, GenBank MT273610.1; Identities = 804/823 (98 %), no gaps), and Peziza proteana f. sparassoides (voucher OSC 100024, GenBank AY544659.1; Identities = 880/902 (98 %), two gaps (0 %)) – also see Fig. 3. Closest hits using the rpb2 sequence had highest similarity to Peziza exogelatinosa (voucher KH 00.029 (C), GenBank AY500501.1; Identities = 620/681 (91 %), no gaps), Peziza petersii (voucher OSC 27373, GenBank MN816680.1; Identities = 556/649 (86 %), three gaps (0 %)), and Peziza proteana f. campbellii (voucher Wu-1937, GenBank MN816673.1; Identities = 582/683 (85 %), eight gaps (1 %)).

Authors: P.W. Crous & C. Decock

Phyllosticta phoenicis Crous, sp. nov. MycoBank MB 839302. Fig. 55.

Fig. 55.

Fig. 55.

Phyllosticta phoenicis (CPC 39164). A. Colony sporulating on OA. B–C. Conidiogenous cells. D. Conidia. Scale bars: A = 250 μm, all others = 10 μm.

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

Conidiomata (on PNA) pycnidial, solitary, black, erumpent, globose, exuding colourless conidial masses; pycnidia 200–300 μm diam; pycnidial wall of several layers of textura angularis; inner wall of hyaline textura angularis. Ostiole central, to 20 μm diam. Conidiophores reduced to conidiogenous cells, or with a supporting cell, that can be branched at the base; Conidiogenous cells subcylindrical to ampulliform, terminal, hyaline, smooth, coated in a mucoid layer, 9–15 × 3–4 μm; proliferating several times percurrently near apex. Conidia (9–)10–13(–15) × (7–) 8(–9) μm, solitary, hyaline, aseptate, thin and smooth-walled, granular, or with a single large central guttule, fusoid-ellipsoid, tapering towards a narrow truncate base, 3 μm diam, enclosed in a persistent mucoid sheath, 1–2 μm thick, and bearing a hyaline, apical mucoid appendage, 10–30 μm long, 2.5–3.5 μm diam at base, flexible, unbranched, tapering towards an acutely rounded tip.

Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse olivaceous grey.

Typus: South Africa, Eastern Cape Province, Haga Haga, on leaves of Phoenix reclinata (Arecaceae), 2018, M.J. Wingfield, HPC 3190 (holotype CBS H-24544, culture ex-type CPC 39164 = CBS 147091).

Notes: Species of Phyllosticta are commonly encountered as endophytes, plant pathogens or saprobes (Glienke et al. 2011, Guarnaccia et al. 2017, Guarnaccia et al. 2019). Several species have been reported from palms. Van der Aa & Vanev (2002) refer to P. cocoina var. phoenicis as having conidia that are 8–9 × 3 μm, being more suggestive of Phoma or Diaporthe; P. palmarum to be representative of an undescribed ascomycetous fungus; P. palmicola to be a Phoma; P. palmigena to be a Coniothyrium, and P. sabalicola to be Apiosphaeria martinii. Phyllosticta cocoicola (Guignardia cocogena) was seen as indistinguishable from P. arecae (Guignardia calami) by Van der Aa & Vanev (2002), but there are no cultures to confirm this synonymy. Van der Aa (1973) cited conidia of P. arecae (on Areca, Calamus, Caryota) as 7–15 × 5–9 μm, thus somewhat smaller than those of P. phoenicis, with shorter appendages, 3–10 μm long. Punithalingam (1974) cited conidia of P. cocoicola (on Cocos) to be 10–16 × 5–6 μm, with appendages 6–9(–10) μm long, thus with narrower conidia and shorter appendages than observed in P. phoenicis. A phylogenetic species tree is presented as Fig. 56.

Fig. 56.

Fig. 56.

Consensus phylogram (50 % majority rule) obtained from the maximum likelihood analysis with IQ-TREE of the Phyllosticta multigene (ITS / LSU / tef1 / actA) nucleotide alignment of the novel Phyllosticta species treated in this study. The alignment is based on Norphanphoun et al. (2020). Bootstrap support values (> 79 %) from 5 000 ultrafast bootstrap replicates are shown at the nodes. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Melanops tulasnei (culture CBS 116805) and the novel species treated in this study is highlighted with a coloured block and bold face. Alignment statistics: 59 strains including the outgroup; 1 592 characters including alignment gaps analysed: 732 distinct patterns, 376 parsimony-informative, 227 singleton sites, 988 constant sites. The best models identified in IQ-TREE were: TIM2e+I+G4 (ITS), TNe+I+G4 (LSU), HKY+F+G4 (tef1), TPM2+F+G4 (actA).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Phyllosticta aristolochiicola (strain BRIP 53316a, GenBank NR_111791.1; Identities = 569/579 (98 %), no gaps), Phyllosticta cordylinophila (strain MFUCC 12-0014, GenBank KC686599.1; Identities = 582/599 (97 %), five gaps (0 %)), and Phyllosticta capitalensis (strain MFE29, GenBank MT186150.1; Identities = 575/602 (96 %), seven gaps (2 %)). Closest hits using the LSU sequence are Phyllosticta aristolochiicola (strain BRIP 53316a, GenBank JX486128.1; Identities = 850/860 (99 %), no gaps), Phyllosticta capitalensis (strain MUCC 2916, GenBank LC543421.1; Identities = 825/835 (99 %), no gaps), and Phyllosticta paracapitalensis (strain CPC 28123, GenBank KY855805.1; Identities = 805/815 (99 %), no gaps) – also see Fig. 1. Closest hits using the actA sequence had highest similarity to Phyllosticta lauridiae (strain CBS 145559, GenBank MK876460.1; Identities = 582/604 (96 %), five gaps (0 %)), Phyllosticta acaciigena (strain CPC 28295, GenBank KY173570.1; Identities = 512/534 (96 %), one gap (0 %)), and Phyllosticta encephalarticola (strain CBS 146014, GenBank MN556783.1; Identities = 563/600 (94 %), one gap (0 %)). Closest hits using the gapdh sequence had highest similarity to Phyllosticta capitalensis (strain VIC30428, GenBank JF343743.1; Identities = 307/310 (99 %), no gaps), Phyllosticta musarum (strain GZAAS6.1228, GenBank KM816632.1; Identities = 458/471 (97 %), one gap (0 %)), and Phyllosticta rhizophorae (strain NCYUCC 19-0358, GenBank MT363251.1; Identities = 390/404 (97 %), two gaps (0 %)). Closest hits using the tef1 sequence had highest similarity to Phyllosticta capitalensis (strain GZAAS6.1201, GenBank KM816635.1; Identities = 376/384 (98 %), no gaps), Phyllosticta ericarum (strain GZAAS6.1250, GenBank KR025451.1; Identities = 376/384 (98 %), no gaps), and Phyllosticta carochlae (strain SM52, GenBank MT118271.1; Identities = 367/384 (96 %), one gap (0 %)).

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Plectosphaerella slobbergiarum Hern.-Restr., sp. nov. MycoBank MB 838711. Fig. 57.

Fig. 57.

Fig. 57.

Plectosphaerella slobbergiarum (CBS 147227). A–C. Conidiophores and conidia. D. Conidia. E. Colony on OA. F. Colony on PDA. G. Colony on MEA. Scale bars A–D = 10 μm.

Etymology: Named for the collectors Julia Slob and Maaike Berghui, students from Het Hogeland College (Warffum, the Netherlands). This sample was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae hyaline, septate, smooth, 1.5–4 μm. Conidiophores semi-micronematous, hyaline, smooth, irregularly branched. Conidiogenous cells monophialidic, terminal or intercalary, subcylindrical to subulate, smooth, hyaline, 10–40 × 2–3 μm, with conspicuous cylindrical collarette and periclinal thickening at the conidiogenous locus, 1–3.5 × 1–2 μm. Conidia cylindrical to ellipsoidal with rounded apex and slightly truncate base, inequilateral, with inner plane flat and outer plane convex, guttulate, hyaline, 0–1-septate; septate conidia 6–12.5 × 2–3.5 μm; aseptate conidia 4–8.5 × 2–3 μm. Chlamydospores and sexual morph absent.

Culture characteristics: After 2 wk at 24 °C on PDA reaching 68 mm diam, buff white, in the centre cottony, elevated, with moderate aerial mycelium, becoming less towards the periphery, fimbriate, margin regular, entire, reverse buff. On OA reaching 60 mm diam, zonate with concentric rings, buff and honey; centre with moderate aerial mycelium, glabrous towards the periphery, margin entire, regular; reverse honey buff. On MEA reaching 48 mm diam, buff, sparse aerial mycelium, wet appearance, with radial disposition of the wrinkles, margin slightly lobate; reverse ochreous with buff radial lines.

Typus: Netherlands, Groningen Province, Warffum, Juffer Martha street, from soil, 6 Jun. 2019, J. Slob & M. Berghuis, NL19_030 (holotype CBS H-24736, culture ex-type CBS 147227 = NL1930002).

Notes: Plectosphaerella comprises several plant pathogenic and soil-born species (Carlucci et al. 2012, Giraldo & Crous 2019, Giraldo et al. 2019, Zhang et al. 2019). Plectosphaerella slobbergiarum is phylogenetically distinct (Fig. 58), and distinguished from other species mainly based on tef1 sequences.

Fig. 58.

Fig. 58.

Maximum composite likelihood tree obtained from the RAxML analysis of the combined ITS, LSU and tef1 sequence alignment of Plectosphaerella species. Bootstrap support values above 70 % are shown at the nodes. The novel species is indicated in a coloured block and bold face. The tree was rooted to Brunneochlamydosporium nepalense (CBS 277.89 and CBS 971.72). Alignment statistics: 12 strains including the outgroup; 2 851 characters including alignment gaps analysed (ITS: 582, LSU: 1391, tef1: 878). Model: GTR+G, alignment patterns: 213.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to P. nauculispora (strain CBS 144924 as “pauciseptata”, GenBank LR590240.1; Identities = 545/549 (99 %), no gaps), P. plurivora (strain GBC-Fungus 109 as “niemeijerarum”, GenBank MN077480.1; Identities = 541/545 (99 %), no gaps), and P. cucumerina (strain IIIZ1-2, GenBank MN915127.1; Identities = 544/549 (99 %), one gap (0 %)). Closest hits using the LSU sequence are P. cucumerina (strain BCC86515, GenBank MH398573.1; Identities = 817/817 (100 %), no gaps); strain CBS 137.37, GenBank MH867359.1; Identities = 817/817 (100 %), no gaps), and strain CBS 355.36, GenBank MH867324.1; Identities = 817/817 (100 %), no gaps) – also see Fig. 5. Closest hits using the tef1 sequence are P. citrulli (strain CBS 131741 as “citrullae”, GenBank LR026491.1; Identities = 774/787 (98 %), no gaps); strain CBS 131740, GenBank LR026490.1; Identities = 774/787 (98 %), no gaps) and P. humicola (strain CBS 423.66 as “plurivora”, GenBank LR026506.1; Identities = 772/787 (98 %), no gaps).

Author: M. Hernández-Restrepo

Populomyces Hern.-Restr., gen. nov. MycoBank MB 838712.

Etymology: From Latin Populus, meaning civilians, and referring to the fact that the fungus was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae hyaline, septate, smooth. Conidiophores micronematous, mostly reduced to conidiogenous cells. Conidiogenous cells monophialidic, aggregated in groups, hyaline, smooth, cylindrical to lageniform. Conidia aseptate, cylindrical, solitary, hyaline.

Type species: Populomyces zwinianus Hern.-Restr.

Populomyces zwinianus Hern.-Restr., sp. nov. MycoBank MB 838713. Fig. 59.

Fig. 59.

Fig. 59.

Populomyces zwinianus (CBS 147307). A–B. Conidiophores and conidia. C. Conidia. D. Colony on OA. E. Colony on MEA. F. Reverse colony on MEA. Scale bars A–C = 10 μm.

Etymology: Name refers to the school from where the sample was collected “Zwin College” (Oostburg, the Netherlands). This sample was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae hyaline, septate, smooth, 1–2 μm wide. Conidiophores micronematous, mostly reduced to conidiogenous cells. Conidiogenous cells monophialidic, aggregated in groups, hyaline, smooth, cylindrical to lageniform, 5–16 × 2–3.5 μm, with a cylindrical collarette, 0.5–1.5 μm deep, 1–1.5 μm wide. Conidia aseptate, cylindrical, solitary, hyaline, guttulate, 9–12 × 2–2.5 μm.

Culture characteristics: After 2 wk at 24 °C on OA reaching 15 mm diam, membranous, saffron; sparse aerial mycelium, margin regular; reverse pale luteous. On MEA reaching 10 mm diam, elevated, with radial disposition of the wrinkles, velvety, peach to flesh, margin slightly lobate; reverse saffron to apricot.

Typus: Netherlands, Zeeland Province, Yerseke, from soil, 11 Jun. 2019, W. Vercouteren, S. Meas & R. Verhije, NL19_076 (holotype CBS H-24737, ex-type strain CBS 147307 =NL1976004).

Notes: Populomyces is phylogenetically close to Calloria and Tricellula (Fig. 2). The cylindrical, aseptate conidia of Populomyces are easily distinguished from the stauroconidia of Tricellula (Seifert et al. 2011). Furthermore, Calloria is a polyphyletic genus with apothecial ascomata including species that are related to Cylindrocolla asexual morphs, characterised by polyblastic conidiogenous cells producing conidia in chains, thus distinct from the solitary conidia of Populomyces (Muntañola-Cvetkovic et al. 1997, Seifert et al. 2011).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Chalara sp. (GenBank KX034388.1; Identities = 463/510 (91 %), one gap (0 %)), Dothideomycetes sp. (strain CK1374, GenBank MH474014.1; Identities = 444/493 (90 %), 16 gaps (3 %)), and Fungal sp. (strain LEG-103, GenBank MW201484.1; Identities = 446/496 (90 %), 15 gaps (3 %). Closest hits using the LSU sequence are Calloria urticae (strain G.M. 2016-03-12.3, GenBank MT509570.1; Identities = 630/669 (94 %), four gaps (0 %)), Calloria urticae (strain MFLU 18-0697, GenBank MK591969.1; Identities = 824/824 (100 %), four gaps (0 %)), and Calloria urticae (strain MFLU 18-0696, GenBank MK591968.1; Identities = 630/669 (94 %), four gaps (0 %)) – also see Fig. 2.

Author: M. Hernández-Restrepo

Porodiplodia vitis Crous & R.K. Schumach., Fung. Syst. Evol. 3: 110. 2019. Fig. 60.

Fig. 60.

Fig. 60.

Porodiplodia vitis (CPC 38692). A. Conidiomata on PNA. B–C. Conidiogenous cells. D. Conidia. Scale bars: A = 300 μm, all others = 10 μm.

Conidiomata eustromatic, uni- to multilocular, brown, globose, 150–300 μm, aggregated on PNA, ostiolate. Conidiophores lining inner cavity, subcylindrical, hyaline, smooth, branched, 1–3-septate, 15–20 × 2.5–4 μm, proliferating percurrently near apex. Paraphyses intermingled among conidiophores, hyaline, smooth, septate, subcylindrical with obtuse ends, up to 25 μm long. Conidia in short chains (–3), fusoid-ellipsoid to subcylindrical, medium brown, finely verruculose, guttulate, thick-walled, medianly 1-septate, apex obtuse, base truncate with central pore, (11–)12–14(–17) × (4–)6(–7) μm.

Culture characteristics: Colonies flat, spreading, surface folded, with moderate aerial mycelium and smooth, lobate margin, reaching 35 mm diam after 2 wk at 25 °C. On MEA surface rosy buff, reverse cinnamon; on PDA surface honey, reverse isabelline; on OA surface cinnamon.

Material examined: South Africa, Western Cape Province, Kirstenbosch, on seed pods of Virgilia oroboides (Fabaceae), 2018, M.J. Wingfield, HPC 3070 (CBS H-24436, culture CPC 38692 = CBS 146818).

Notes: Porodiplodia was recently established as a new genus for a species occurring on leaves of Livistona australia in Australia (Crous et al. 2018b). A second species, P. vitis, was described from canes of Vitis vinifera in the USA (Crous et al. 2019a).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Porodiplodia sp. (strain EAB-24-17, GenBank MT777297.1; Identities = 459/460 (99 %), no gaps), Porodiplodia livistonae (strain CPC 32154, GenBank NR_160355.1; Identities = 536/538 (99 %), no gaps), and Porodiplodia vitis (strain CBS 144634, GenBank NR_163376.1; Identities = 543/546 (99 %), no gaps). Closest hits using the LSU sequence are Porodiplodia vitis (strain CBS 144634, GenBank NG_070080.1; Identities = 824/824 (100 %), no gaps), Porodiplodia livistonae (strain CPC 32154, GenBank NG_069575.1; Identities = 824/824 (100 %), no gaps), and Chalara clidemiae (strain CPC 26423, GenBank KX228321.1; Identities = 817/824 (99 %), no gaps) – also see Fig. 2. Closest hits using the tef1 sequence had highest similarity to Porodiplodia vitis (strain CBS 144634, GenBank MK442707.1; Identities = 463/473 (98 %), single indel of ten nucleotides (2 % gap)), Davidhawksworthia ilicicola (strain CBS 261.95, GenBank KU728592.1; Identities = 205/233 (88 %), seven gaps (3 %)), and Hymenoscyphus menthae (strain HB_5846, GenBank KM114512.1; Identities = 204/232 (88 %), six gaps (2 %)). A comparison with the tef1 sequence we generated of Porodiplodia livistonae in this study revealed a similarity of 426/439 (97 %, including a single indel of ten nucleotides (2 % gap) also present for Porodiplodia vitis).

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Pseudoacrospermum Crous, gen. nov. MycoBank MB 839303.

Etymology: Name refers to the fact that it is phylogenetically closely related to Acrospermum.

Mycelium consisting of septate, branched, smooth, pale brown, hyphae. Conidiophores solitary, erect, flexuous, subcylindrical, multiseptate, unbranched with basal T-cell, arising from superficial mycelium, thick-walled, pale olivaceous, smooth, lacking rhizoids, terminating in a slightly swollen conidiogenous cell. Conidiogenous cells integrated, terminal, subclavate, with numerous sympodial loci aggregated in a rachis, darkened, thickened, but not refractive, pimple-like, protruding. Conidia solitary, smooth, pale olivaceous to granular, subclavate, curved, apex obtuse, tapering to truncate hilum, darkened, thickened, and refractive, somewhat thick-walled, transversely septate.

Type species: Pseudoacrospermum goniomae Crous

Pseudoacrospermum goniomae Crous, sp. nov. MycoBank MB 839304. Fig. 61.

Fig. 61.

Fig. 61.

Pseudoacrospermum goniomae (CPC 37030). A. Conidiophores on SNA. B–D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars: A = 180 μm, all others = 10 μm.

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

Mycelium consisting of septate, branched, smooth, pale brown, 2–2.5 μm diam hyphae. Conidiophores solitary, erect, flexuous, subcylindrical, multiseptate, unbranched with basal T-cell, arising from superficial mycelium, thick-walled, pale olivaceous, smooth, lacking rhizoids, 100–180 × 2.5–3.5 μm, terminating in a slightly swollen conidiogenous cell. Conidiogenous cells integrated, terminal, subclavate, 15–20 × 4–6 μm, with numerous sympodial loci aggregated in a rachis, darkened, thickened, but not refractive, pimple-like, protruding (0.5 μm), 1–2 μm diam. Conidia solitary, smooth, pale olivaceous to granular, subclavate, curved, apex obtuse, tapering to truncate hilum, 2 μm diam, darkened, thickened, and refractive, somewhat thick-walled, 3(–6)-septate, (17–)21–23(–25) × (5–)6 μm.

Culture characteristics: Colonies flat, spreading, surface folded, with moderate aerial mycelium and smooth, even margin, reaching 5 mm diam after 2 wk at 25 °C. On MEA surface buff to honey, reverse isabelline; on PDA surface straw in centre, outer margin buff to cinnamon, reverse cinnamon; on OA surface cinnamon.

Typus: South Africa, Western Cape Province, Knysna, on leaves of Gonioma kamassi (Apocynaceae), 22 Nov. 2018, F. Roets, HPC 2733 (holotype CBS H-24393, culture ex-type CPC 37030 = CBS 146732).

Notes: Pseudoacrospermum is phylogenetically closely related to Acrospermum, which is known only by a sexual morph. Based on its phylogeny, Acrospermum has been shown to be closely related to such as Gonatophragmium, Radulidium or Pseudovirgaria (Hudson et al. 2019), that are presently known by their asexual morphs. Pseudoacrospermum is distinguished based on its multiseptate conidia. There is no evidence to suggest that Pseudoacrospermum can be linked to Acrospermum. A phylogenetic tree is presented as Fig. 62.

Fig. 62.

Fig. 62.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Acrospermum and related genera multigene (LSU / tef1 / ITS) nucleotide alignment. The alignment is derived from Hudson et al. (2019). Bayesian posterior probabilities (> 0.79) are shown at the nodes and the scale bar represents the expected changes per site. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Nectria cinnabarina (culture CBS 713.97) and the novel species treated in this study is indicated in a coloured block and bold face. Alignment statistics: 47 strains including the outgroup; 595 / 418 / 284 unique site patterns, respectively. Tree statistics: 184 502 sampled trees from 1 230 000 generations.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Acrospermum maxonii (as Acrospermum sp. VD-2019c; voucher 2711, GenBank MK562001.1; Identities = 344/421 (82 %), 23 gaps (5 %)), Radulidium epichloes (strain CBS 115704, GenBank MH862990.1; Identities = 217/236 (92 %), five gaps (2 %)), and Acrospermum leucocephalum (as Acrospermum sp. VD-2019b; voucher 764, GenBank MK562008.1; Identities = 334/413 (81 %), 22 gaps (5 %)). Closest hits using the LSU sequence are Acrospermum gorditum (as Acrospermum sp. VD-2019a; voucher 1774, GenBank MK561981.1; Identities = 803/867 (93 %), seven gaps (0 %)), Acrospermum maxonii (as Acrospermum sp. VD-2019c; voucher 3114, GenBank MK561987.1; Identities = 804/870 (92 %), 11 gaps (1 %)), and Gonatophragmium triuniae (strain CBS 138901, GenBank NG_058117.1; Identities = 764/827 (92 %), nine gaps (1 %)) – also see Fig. 1. The best hit using the rpb2 sequence had highest similarity to Pseudovirgaria hyperparasitica (strain CBS 121739, GenBank XM_033741841.1; Identities = 573/769 (75 %), ten gaps (1 %)). Only very distant hits with taxa in Dothideomycetes were obtained using the tef1 (first and second part) sequences. However, both these sequences were manually compared with blast2 searches against Acrospermum tef1 sequences labelled as unverified in the GenBank nucleotide database: Closest hits using the tef1 (first part) sequence had highest similarity to Acrospermum maxonii (strain 75017, GenBank MK604769.1; Identities = 198/234 (85 %), seven gaps (2 %)), Acrospermum gorditum (strain 1774, GenBank MK604778.1; Identities = 196/232 (84 %), two gaps (0 %)), and Acrospermum leucocephalum (strain 3342, GenBank MK604780.1; Identities = 213/256 (83 %), nine gaps (3 %)). Closest hits using the tef1 (second part) sequence had highest similarity to Acrospermum maxonii (strain 44, GenBank MK604767.1; Identities = 426/473 (90 %), no gaps), and Acrospermum leucocephalum (strain 764, GenBank MK604781.1; Identities = 412/474 (87 %), one gap (0 %)).

Authors: P.W. Crous, J.Z. Groenewald & F. Roets

Pseudopyricularia festucae Crous, sp. nov. MycoBank MB 839305. Fig. 63.

Fig. 63.

Fig. 63.

Pseudopyricularia festucae (CPC 37915). A–D. Conidiophores with conidiogenous cells giving rise to conidia. Scale bars = 10 μm.

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

Mycelium consisting of branched, septate, hyaline, smooth, 2.5–3.5 μm diam hyphae. Conidiophores solitary, erect, subcylindrical, unbranched, flexuous, medium brown, smooth, up to 150 μm tall, 5–6 μm diam, with 1–2 basal septa. Conidiogenous cells terminal, integrated, medium brown, smooth, 70–100 μm tall, with a rachis of protruding denticles, 1.5–2 × 1–1.5 μm. Conidia solitary, pale brown, fusoid, finely roughened, 2-septate, apex subobtuse, base truncate, 1.5 μm diam, apex somewhat thickened, (25–)30–38(–40) × (6–)7 μm.

Culture characteristics: Colonies flat, spreading, with moderate aerial mycelium and smooth, lobate margin, reaching 50 mm diam after 2 wk at 25 °C. On MEA surface dirty white to smoke grey, reverse honey; on PDA surface and reverse olivaceous grey; on OA surface olivaceous grey.

Typus: USA, California, UC Davis, on leaves of Festuca californica (Poaceae), 3 Apr. 2019, P.W. Crous, HPC 2915 (holotype CBS H-24352, culture ex-type CPC 37915 = CBS 146629).

Notes: Pseudopyricularia is characterised by having short, determinate, brown conidiophores with an apical rachis with flat-tipped denticles (Klaubauf et al. 2014, Marin-Felix et al. 2017). The present isolate is phylogenetically closely related to P. iraniana (conidia (20–)22–30 × 5–8 μm, on Juncus sp. in Iran; Pordel et al. 2017) but has longer conidia, (25–)30–38(–40) × (6–)7 μm. A phylogenetic species tree is presented as Fig. 64.

Fig. 64.

Fig. 64.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Pseudopyricularia multigene (cmdA / ITS / rpb1) nucleotide alignment. The alignment is derived from Klaubauf et al. (2014). Bayesian posterior probabilities (> 0.79) are shown at the nodes and the scale bar represents the expected changes per site. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Macgarvieomyces juncicola (culture CBS 610.82) and the novel species treated in this study is indicated in a coloured block and bold face. Alignment statistics: 20 strains including the outgroup; 248 / 123 / 254 unique site patterns, respectively. Tree statistics: 39 002 sampled trees from 130 000 generations.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Pseudopyricularia iraniana (strain IRAN 2761C, GenBank NR_158928.1; Identities = 500/504 (99 %), no gaps), Dactylaria higginsii (strain CBS 121934, GenBank KM009164.1; Identities = 482/509 (95 %), 7 gaps (1 %)), and Pseudopyricularia bothriochloae (strain CBS 136427, GenBank NR_137838.1; Identities = 519/552 (94 %), 11 gaps (1 %)). Closest hits using the LSU sequence are Pseudopyricularia iraniana (strain 17d, GenBank KY457268.1; Identities = 863/863 (100 %), no gaps), Pyricularia caricis (strain JAC12652, GenBank MK431456.1; Identities = 874/880 (99 %), no gaps), and Pseudopyricularia hyrcaniana (strain Ck3, GenBank KY457267.1; Identities = 850/863 (98 %), no gaps) – also see Fig. 5. Closest hits using the cmdA sequence had highest similarity to Pseudopyricularia iraniana (strain 16d, GenBank KY457263.1; Identities = 400/414 (97 %), 11 gaps (2 %)), Pseudopyricularia kyllingae (strain CB8959, GenBank KM485251.1; Identities = 429/511 (84 %), 17 gaps (3 %)), Macgarvieomyces borealis (strain CBS 461.65, GenBank KM485239.1; Identities = 156/163 (96 %), no gaps), and Macgarvieomyces luzulae (strain CPC 31571, GenBank MG934521.1; Identities = 153/160 (96 %), no gaps). Closest hits using the rpb1 sequence had highest similarity to Pseudopyricularia iraniana (strain 17d, GenBank KY457273.1; Identities = 961/981 (98 %), no gaps), Pseudopyricularia bothriochloae (strain CBS 136427, GenBank KY905701.1; Identities = 614/669 (92 %), no gaps), and Pseudopyricularia hagahagae (strain CPC 25635, GenBank KT950877.1; Identities = 926/1 013 (91 %), no gaps).

Authors: P.W. Crous, J.Z. Groenewald & L. Lombard

Sarocladium sasijaorum Hern.-Restr., sp. nov. MycoBank MB 838714. Fig. 65.

Fig. 65.

Fig. 65.

Sarocladium sasijaorum (CBS 147213). A–C. Conidiophores and conidia. D. Conidia. E. Colony on MEA. F. Colony on OA. G. Colony on PDA. Scale bars A–D = 10 μm

Etymology: Name reflects the names of the collectors Sami Maghnouji, Simon Frederikze, and Jason Mes, students from Wereldwijs (Bilthoven, the Netherlands). This sample was collected during a Citizen Science project of the Westerdijk Fungal Biodiversity Institute.

Vegetative hyphae septate, hyaline, smooth and thin-walled, 1–2 μm wide. Conidiophores erect, arising directly from vegetative hyphae or ropes of hyphae, straight or slightly bent, simple, hyaline, smooth-walled. Phialides subulate to acicular, 16–32 μm long, 1.5–2 μm wide at the base, 1–1.5 μm wide at the apex, straight or flexuous, hyaline, thin- and smooth-walled. Conidia arranged in long chains, unicellular, fusoid, with pointed ends, becoming limoniform to obovoid with age, 4–6 × 1.5–2 μm, hyaline, smooth-walled. Chlamydospores and sexual morph not observed.

Culture characteristics: After 7 d at 25 °C on OA attaining 13 mm diam, cottony, flat, with sparse aerial mycelium, buff to white, dry, margin effuse; reverse similar. On MEA attaining 14 mm diam, cottony, elevated, buff to pale luteous; reverse apricot. On PDA attaining 13 mm diam, cottony to hairy, with sparse aerial mycelium towards the periphery, olivaceous buff to white; reverse pale luteous in centre, white towards the periphery.

Typus: Netherlands, Utrecht Province, Bilthoven, Planetenplein, from soil, 31 Jul. 2019, S. Frederikze, J. Mes & S. Maghnouji, NL19_100 (holotype CBS H-24738, culture ex-type CBS 147213 = NL19100007).

Notes: Sarocladium sasijaorum is phylogenetically close to S. dejongiae, S. liguanense, and S. clematidis (Fig. 66). Sarocladium clematidis was described from Belgium on dead stems of Clematis patens (Phukhamsakda et al. 2020) and resembles Phaeoisaria clematidis based on morphology. Sarocladium dejongiae was recently described form Dutch soil (Crous et al. 2018a) and S. liguanense from China on Malus × domestica (Hou et al. 2019). However, S. sasijaorum is distinguishable based on its conidial arrangement and morphology. In S. sasijaorum conidia are ellipsoidal with pointed ends, 4–6 × 1.5–2 μm and arranged in chains; in S. dejongiae conidia are ellipsoidal to cylindrical, 3–5 × 1–2 μm (av. 3 × 1.5 μm) and disposed in slimy heads (Crous et al. 2018a), and in S. liguanense conidia are ellipsoidal to cylindrical, 3–6.5 × 1.5–3 μm, and arranged in slimy heads (Hou et al. 2019). Furthermore, in S. dejongiae and S. liguanense chlamydospores are present (Crous et al. 2018a, Hou et al. 2019), while they were not observed in S. sasijaorum. Also see the overview phylogeny (Fig. 4).

Fig. 66.

Fig. 66.

Maximum composite likelihood tree obtained from the RAxML analysis of the combined ITS and LSU sequence alignment of Sarocladiaceae members. Bootstrap support values above 70 % are shown at the nodes. The novel species are indicated in bold face and the two genera are indicated with coloured blocks. The tree was rooted to Xenoacremonium recifei (CBS 137.35). Alignment statistics: 43 strains including the outgroup; 1 571 characters including alignment gaps analysed (ITS: 642, LSU: 929). Model GTR+I+G, 620 distinct patterns.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Ascomycota sp. (strain j17, GenBank EU167596.1; Identities = 488/491 (99 %), one gap (0 %)), Hypocreales sp. (strain f10, GenBank MN982323.1; Identities = 481/484 (99 %), one gap (0 %)), and Fungal sp (strain acwVHT103_8, GenBank JQ070513.1; Identities = 427/442 (97 %), three gaps (0 %)). Closest hits using the LSU sequence are Sarocladium sp. (strain ACCC 39306, GenBank MF987651.1; Identities = 773/779 (99 %), no gaps), Sarocladium dejongiae (strain CBS 144929, GenBank NG_069161.1; Identities = 773/779 (99 %), no gaps), and Sarocladium sp. (strain MFLUCC 17-2150, GenBank MN629285.1; Identities = 767/779 (98 %), no gaps) – also see Fig. 4. Closest hits using the rpb2 sequence had highest similarity to Sarocladium sp. (strain MFLUCC 17-2150, GenBank MN628627.1; Identities = 683/761 (90 %), no gaps), Acremonium sp. DLW-2010 (strain 11665, GenBank GQ867809.1; Identities = 650/811 (80 %), four gaps (0 %)), and Sarocladium sp. (strain A131, GenBank KC999025.1; Identities = 650/811 (80 %), four gaps (0 %)). Closest hits using the tef1 sequence had highest similarity to Sarocladium implicatum (strain 04035, GenBank KT878359.1; Identities = 601/630 (95 %), no gaps), Sarocladium kiliense (GenBank MK512750.1; Identities = 591/630 (94 %), no gaps), and Sarocladium kiliense (strain MB20A1, GenBank LT615314.1; Identities = 591/630 (94 %), no gaps).

Author: M. Hernández-Restrepo

Septoria protearum Viljoen & Crous, S. African J. Bot. 64: 144. 1998. Fig. 67.

Fig. 67.

Fig. 67.

Septoria protearum (CPC 38736). A. Conidiomata on OA. B–D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm.

Description and illustration: Verkley et al. (2013).

Material examined: New Zealand, Tuaranga Port, on Kniphofia uvaria (Asphodelaceae), 22 Aug. 2019, D. Burnt, culture CPC 38736 = T19_05709B.

Notes: Septoria protearum is a plurivorous species known from various hosts in South Africa, Europe and New Zealand (Verkley et al. 2013). Septoria citri is considered to belong to the Septoria protearum species complex which still needs to be resolved (Verkley et al. 2013).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Septoria coprosmae (as Mycosphaerella coacervata; strain CBS 113391, GenBank EU167596.1; Identities = 540/540 (100 %), no gaps), Mycosphaerella linorum (strain CBS 261.39, GenBank EU167590.1; Identities = 540/540 (100 %), no gaps), and Septoria protearum (strain CBS 778.97, GenBank NR_163551.1; Identities = 539/539 (100 %), no gaps). Closest hits using the LSU sequence are Septoria protearum (strain CBS 778.97, GenBank NG_069851.1; Identities = 751/752 (99 %), one gap (0 %)), Septoria malagutii (strain CBS 106.80, GenBank NG_069161.1; Identities = 751/752 (99 %), one gap (0 %)), and Septoria citricola (strain CBS 356.36, GenBank NG_069158.1; Identities = 751/752 (99 %), one gap (0 %)) – also see Fig. 1. Closest hits using the actA sequence had highest similarity to Septoria protearum (strain CBS 778.97, GenBank KF253827.1; Identities = 214/214 (100 %), no gaps), Septoria citri (strain CBS 315.37, GenBank JX902161.1; Identities = 543/543 (100 %), no gaps), Septoria eucalyptorum (strain CBS 118505, GenBank KF903501.1; Identities = 500/528 (95 %), three gaps (0 %)), and Septoria carvi (strain KML1860, GenBank KX822110.1; Identities = 495/531 (93 %), no gaps). Closest hits using the tef1 sequence had highest similarity to Septoria protearum (strain CBS 778.97, GenBank KF253472.1; Identities = 387/387 (100 %), no gaps), Septoria citri (strain CBS 315.37, GenBank JX901706.1; Identities 396/396 (100 %), no gaps), Septoria chamaecisti (strain CBS 350.58, GenBank KF253318.1; Identities = 387/387 (100 %), no gaps), and Septoria citricola (strain CBS 356.36, GenBank KF253329.1; Identities = 387/388 (99 %), one gap (0 %)).

Authors: P.W. Crous, J.Z. Groenewald & R. Thangavel

Sporothrix hypoxyli W.J. Nel, Z.W. De Beer & T.A. Duong, sp. nov. MycoBank MB 837458. Fig. 68.

Fig. 68.

Fig. 68.

Sporothrix hypoxyli (CBS 141569). A, H. Ascomata on Fraxinus wood embedded in fruiting body of Hypoxylon petriniae. B. Ascospores. C. Ostiole containing ostiolar hyphae. D. Ascoma. E. One-month-old culture on MEA. F. Conidia. G. Conidiophore. Scale bars: A, H = 1 mm, B–C, F–G = 10 μm, D = 100 μm.

Etymology: Latin, hypoxyli, refers to the genus of the fungus fruiting body from which this species was isolated, Hypoxylon.

Ascomatal bases black, globose, (140–)170–279(–392) μm diam, ornamented with brown hyphal hairs, (10–)18–47(–74) μm long, (1.5–)1.8–2.4(–2.6) μm diam. Ascomatal necks, black, mostly straight, occasionally slightly curved or kinked tapering toward apex, vary greatly in length, (429–)580–1125(–1504) μm long, (25–)35–49(–58) μm wide at base, (11–)14–23(–28) μm wide at apex. Ostiolar hyphae present, slightly divergent, hyaline, tapering toward tip, (24–)30–47(–59) μm long. Asci dehiscent. Ascospores produced in slimy droplet at apex of ascomatal neck, hyaline, aseptate, reniform to bean-shaped, no sheath, (2.6–)3.2–3.7(–3.8) μm long, (1.2–)1.4–1.7(–1.9) μm wide at centre. Sporothrix-like asexual morph. Conidiophores hyaline sometimes becoming light brown with age, smooth, arising singly from mycelia, micronematous, (14–)16–44(–88) μm long. Conidiogenous cells (4–)10–26(–40) μm long, denticulate. Conidia hyaline can become light brown with age, aseptate, round to obovoid, produced directly on denticles, arising either from hyphae or on conidiophores, (2.5–)3.4–4.6(–5.9) μm long, (2.2–)2.4–3(–3.5) μm wide at thickest part.

Culture characteristics (25 °C, 10 d, MEA): Colonies produce aerial mycelium initially white sometimes becoming pigmented with age, superficial mycelium in media initially white, becoming light to dark brown with age, slow growing, growing 14 mm in 10 d. Ascomata form in culture after 2–3 mo at low temperatures (5–15 °C).

Typus: Netherlands, Utrecht Province, Utrecht, ascomata embedded in sporocarp of Hypoxylon petriniae on Fraxinus wood, 19 Apr. 2015, E. Osieck & W.J. Nel (holotype PREM 63086, ex-type culture CMW 47441 = CBS 141569).

Additional materials examined: Netherlands, Utrecht Province, Utrecht, ascomata embedded in sporocarp of Hypoxylon petriniae on Fraxinus wood, 19 Apr. 2015, E. Osieck & W.J. Nel (PREM 63083, culture CMW 47436 = CBS 141568); Utrecht Province, Utrecht, ascomata embedded in sporocarp of Hypoxylon petriniae on Fraxinus wood, 19 Apr. 2015, E. Osieck & W.J. Nel (PREM 63085, culture CMW 47439 = CBS 141567).

Notes: A multigene phylogeny (Fig. 69) of combined sequences for the LSU, ITS and tub2 gene regions resolved Sporothrix hypoxyli in a clade distant from all other Sporothrix spp. Few other species in the genus have previously been described in association with the sporocarps of other fungi (Guerrero 1971, Samuels & Müller 1979, Constantinescu & Ryman 1989). This represents the first report of a species of Sporothrix with ascomata embedded in the sporocarp of a Hypoxylon sp. Also see the overview phylogeny (Fig. 5).

Fig. 69.

Fig. 69.

Combined phylogeny of Sporothrix based on sequences for the LSU, ITS, and tub2 gene regions. Datasets for the individual gene regions were prepared (MEGA v. 7) using sequences available on GenBank, aligned (MAFFT v. 7.454) and then combined manually before being analysed using Maximum Likelihood (ML; RAxML GUI v.2) and Bayesian Inference (BI; MrBayes v. 3.2.5), with the former being the tree presented. GenBank accession (superscript) and culture collection numbers are indicated for all species. The tree was rooted using isolates of the genus Ceratocystiopsis (Cop.) and basal branches were shortened to facilitate layout. Bootstrap support values (≥ 75 %; ML) and Posterior probability values (≥ 0.95; BI) are given at nodes as ML / BI. Dashes indicate low or unsupported branches. The new species is indicated in a coloured block and bold face. The scale bar indicates the number of changes. Alignment statistics: 56 strains including the outgroup taxa, 697 / 499 / 249 unique site patterns. Tree statistics: 205 738 trees sampled from 10 M generations. T = ex-type; P = Paratype; I = Isotype; * = KX590854 / KX590817 / KX590760.

Authors: W.J. Nel, Z.W. de Beer, T.A. Duong & M.J. Wingfield

Stemphylium eturmiunum E.G. Simmons, Harvard Pap. Bot. 6: 204. 2001. Fig. 70.

Fig. 70.

Fig. 70.

Stemphylium eturmiunum (CPC 38613). A. Ascomata developing on PNA. B–E. Conidiophores with conidiogenous cells giving rise to conidia. F. Ascus and pseudoparaphyses. G–I. Asci and ascospores. Scale bars: A = 200 μm, all others = 10 μm.

Ascomata on PNA pseudothecial, subconical to subglobose, erumpent, brown with central ostiole, 250–350 μm diam; wall of 6–8 layers of brown textura angularis. Asci 2–8-spored, stipitate, bitunicate, fasciculate, subclavate, 120–150 × 27–35 μm, with clearly defined apical chamber, 3–5 μm diam. Pseudoparaphyses intermingled among asci, hyaline, smooth, hyphae-like, septate, anastomosing, 2.5–5 μm diam. Ascospores golden brown, smooth, ellipsoid, constricted at medium septum, ends obtuse, muriformly septate, 5–7 transverse septa, and 1–4 oblique or vertical septa per row, (28–)30–31(–34) × (10–)11–12(–13) μm. Mycelium consisting of medium brown, smooth, septate, branched, 3–4 μm diam hyphae. Conidiophores arising from superficial hyphae, subcylindrical, medium brown, smooth, straight to geniculate-sinuous, unbranched, 1–4-septate, 20–50 × 3–5 μm. Conidiogenous cells integrated, terminal, clavate, medium to dark brown, 7–15 × 6–7 μm, with terminal pore, 1.5–2 μm diam. Conidia solitary or in short chains (2–3), dark brown, verruculose, narrowly ellipsoid to subcylindrical, (14–) 17–20(–22) × (8–)12–13(–14) μm, muriformly septate, at times constricted at median septum, 1–3 horizontal and 1–3 vertical or oblique septa per row.

Culture characteristics: Colonies flat, spreading, with moderate to abundant aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA, PDA and OA surface olivaceous grey, reverse iron-grey.

Material examined: South Africa, Western Cape Province, Nieuwoudtville, on leaves of Bulbinella latifolia (Asphodelaceae), 2018, P.W. Crous, HPC 3061 (CBS H-24445, culture ex-type CPC 38613 = CBS 146783).

Notes: Stemphylium eturmiunum was described as a pathogen of Allium sativum in France and India (Woudenberg et al. 2017), and recently as postharvest pathogen of Allium in China (Fu et al. 2019). This is the first report of this pathogen occurring on leaves of Bulbinella latifolia in South Africa.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Stemphylium eturmiunum (strain FXA6, GenBank MT898507.1; Identities = 575/575 (100 %), no gaps), Stemphylium vesicarium (strain UKKo1, GenBank MN328401.1; Identities = 588/589 (99 %), one gap (0 %)), and Stemphylium botryosum (strain CBS 714.68, GenBank NR_163547.1; Identities = 585/587 (99 %), no gaps). Closest hits using the LSU sequence are Stemphylium eturmiunum (strain CBS 109845, GenBank NG_069866.1; Identities = 826/826 (100 %), no gaps), Stemphylium paludiscirpi (strain CBS 109842, GenBank NG_069865.1; Identities = 826/826 (100 %), no gaps), and Stemphylium triglochinicola (strain CBS 718.68, GenBank NG_069740.1; Identities = 826/826 (100 %), no gaps) – also see Fig. 1. Closest hits using the actA sequence had highest similarity to Stemphylium vesicarium (as Pleospora herbarum; strain ATCC 11681, GenBank JQ671595.1; Identities = 575/585 (98 %), no gaps), Stemphylium botryosum (as Pleospora tarda; strain ATCC 42170, GenBank JQ671593.1; Identities = 575/585 (98 %), no gaps), and Stemphylium callistephi (strain EEB 1055, GenBank JQ671592.1; Identities = 572/585 (98 %), three gaps (0 %)). Closest hits using the cmdA sequence had highest similarity to Stemphylium eturmiunum (strain CBS 668.80, GenBank KU850830.1; Identities = 562/562 (100 %), no gaps), Stemphylium botryosum (strain M-14, GenBank MH206180.1; Identities = 553/562 (98 %), no gaps), and Stemphylium truncatulae (strain UMSe004, GenBank MK336838.1; Identities = 551/560 (98 %), no gaps). Closest hits using the gapdh sequence had highest similarity to Stemphylium eturmiunum (strain CBS 109845, GenBank KU850689.1; Identities = 581/581 (100 %), no gaps), Stemphylium mali (as Stemphylium sp. XGZ-2010b; strain CBS 124652, GenBank GQ395376.1; Identities = 554/558 (99 %), no gaps), and Stemphylium armeriae (strain CBS 338.73, GenBank KU850658.1; Identities = 573/581 (99 %), no gaps). Closest hits using the rpb2 sequence had highest similarity to Stemphylium vesicarium (strain BRIP 65181, GenBank KY009907.1; Identities = 800/812 (99 %), no gaps), Stemphylium vesicarium (as Pleospora herbarum; strain N131, GenBank KU738709.1; Identities = 794/806 (99 %), one gap (0 %)), and Stemphylium botryosum (strain ATCC 42170, GenBank JQ905202.1; Identities = 685/697 (98 %), no gaps). No rpb2 sequences of Stemphylium eturmiunum are available for comparison. Closest hits using the tef1 sequence had highest similarity to Stemphylium sp. (strain EGS48-074, GenBank AY324686.1; Identities = 421/421 (100 %), no gaps), Stemphylium vesicarium (strain P66, GenBank MK497791.1; Identities = 211/211 (100 %), no gaps), and Stemphylium majusculum (strain EGS16-068, GenBank AY324710.1; Identities = 414/421 (98 %), one gap (0 %)). No tef1 sequences of Stemphylium eturmiunum are available for comparison; however, the gapdh sequence of strain EGS48-074 (GenBank AY316986.1) differs with only one nucleotide from sequences of Stemphylium eturmiunum and could therefore belong to this species.

Authors: P.W. Crous, J.Z. Groenewald & M.J. Wingfield

Superstratomycetales van Nieuwenh., Miądl., Houbraken, Adan, Lutzoni & Samson, ord. nov. MycoBank MB 839306.

Synonym: Superstratomycetales van Nieuwenh. et al., Stud. Mycol. 85: 115. 2016. Nom. inval., Arts 40.1, 40.3, see Arts 6.3, 12.1 (Shenzhen).

Etymology: From Latin “super” = above/on the top, and “stratum” = layer of material and from Greek “myces” = organisms, referring to fungal stains forming a top layer or covering the surface of a material.

Conidiomata pycnidial, superficial, solitary or confluent, brown to black, glabrous, globose, exuding a white mass of slimy conidia; pycnidial wall pseudoparenchymatous, of textura angularis, composed of 3–5 layers of pale brown to brown cells. Setae present or absent, erect to recurved, hyaline to subhyaline at apex and turning brown towards base, septate, verrucose to tuberculate. Conidiophores reduced to conidiogenous cells or branched, hyaline, smooth-walled, with terminal and intercalary conidiogenous cells. Conidiogenous cells phialidic, hyaline, smooth-walled, cylindrical to doliiform or ampulliform. Conidia hyaline, aseptate, smooth- and thin-walled, guttulate, cylindrical to navicular or ovoid.

Type genus: Superstratomyces van Nieuwenh., Miadl. & Samson

Superstratomycetaceae van Nieuwenh., Miądl., Houbraken, Adan, Lutzoni & Samson, fam. nov. MycoBank MB 839307.

Synonym: Superstratomycetaceae van Nieuwenh. et al., Stud. Mycol. 85: 115. 2016. Nom. inval., Arts 40.1, 40.3, see Arts 6.3, 12.1 (Shenzhen).

Etymology: From Latin “super” = above/on the top, and “stratum” = layer of material and from Greek “myces” = organisms, referring to fungal stains forming a top layer or covering the surface of a material.

Conidiomata pycnidial, superficial, solitary or confluent, brown to black, glabrous, globose, exuding a white mass of slimy conidia; pycnidial wall pseudoparenchymatous, of textura angularis, composed of 3–5 layers of pale brown to brown cells. Setae present or absent, erect to recurved, hyaline to subhyaline at apex and turning brown towards base, septate, verrucose to tuberculate. Conidiophores reduced to conidiogenous cells or branched, hyaline, smooth-walled, with terminal and intercalary conidiogenous cells. Conidiogenous cells phialidic, hyaline, smooth-walled, cylindrical to doliiform or ampulliform. Conidia hyaline, aseptate, smooth- and thin-walled, guttulate, cylindrical to navicular or ovoid.

Type genus: Superstratomyces van Nieuwenh., Miadl. & Samson

Superstratomyces van Nieuwenh., Miądl. & Samson, gen. nov. MycoBank MB 839308.

Synonym: Superstratomyces van Nieuwenh. et al., Stud. Mycol. 85: 115. 2016. Nom. inval., Arts 40.1, 40.3, see Arts 6.3, 12.1 (Shenzhen).

Etymology: From Latin “super” = above/on the top, and “stratum” = layer of material and from Greek “myces” = organisms, referring to fungal stains forming a top layer or covering the surface of a material.

Conidiomata pycnidial, superficial, solitary or confluent, brown to black, glabrous, globose, exuding a white mass of slimy conidia; pycnidial wall pseudoparenchymatous, of textura angularis, composed of 3–5 layers of pale brown to brown cells. Setae present or absent, erect to recurved, hyaline to subhyaline at apex and turning brown towards base, septate, verrucose to tuberculate. Conidiophores reduced to conidiogenous cells or branched, hyaline, smooth-walled, with terminal and intercalary conidiogenous cells. Conidiogenous cells phialidic, hyaline, smooth-walled, cylindrical to doliiform or ampulliform. Conidia hyaline, aseptate, smooth- and thin-walled, guttulate, cylindrical to navicular or ovoid.

Type species: Superstratomyces albomucosus van Nieuwenh. & Samson

Superstratomyces albomucosus van Nieuwenh. & Samson, sp. nov. MycoBank MB 839309 .

Synonym: Superstratomyces albomucosus van Nieuwenhuijzen & Samson, Stud. Mycol. 85: 115. 2016. Nom. inval., Art. 35.1 (Shenzhen).

Etymology: From Latin “albus” = white, “mucosus” = slimy; referring to the conidia in white slimy masses.

Slow growing grey/olive to dark green colonies, with pycnidia forming aggregated masses (MEA and OA) and little or no slimy masses up to 5 wk (OA). On OA: 25 °C, 2–5 wk: pycnidia forming brown to black aggregated masses, individual conidiomata not present; dark pigmented hyphae; conidia smooth-walled, oval typically with blunt ends, length 2.8–6 μm, width 1.5–3 μm; aerial mycelium hyaline. On nettle stem OA, 25 °C, 4–5 wk: conidiomata brown/black, spherical to subspherical, 80–300 μm diam; conidiogenous cells phialidic.

Description and illustration: See van Nieuwenhuijzen et al. (2016).

Typus: Netherlands, Utrecht Province, Utrecht, outdoor exposed Pycnanthus angolensis impregnated with olive oil, E.J. van Nieuwenhuijzen, 9 Sep. 2013 (holotype CBS H-22668, culture ex-type CBS 140270 = DTO 277-D2).

Superstratomyces atroviridis van Nieuwenh. & Samson, sp. nov. MycoBank MB 839310.

Synonym: Superstratomyces atroviridis van Nieuwenh. & Samson, Stud. Mycol. 85: 118. 2016. Nom. inval., Art. 35.1 (Shenzhen).

Etymology: From Latin “ater” = dark, black, “viride” = green; referring to colonies coloured dark green/ black on agar plates.

Slow growing grey/olive to dark green colonies, with pycnidia forming aggregated masses (MEA and OA) and little or no slimy masses up to 5 wk (OA). On OA, 25 °C, 2–5 wk: aggregated mass of dark brown to black pycnidia, individual conidiomata not visible; dark pigmented hyphae; conidia smooth-walled, oval typically with blunt ends, length (2 wk) 2.6–6.5 μm, width 1.6–2.8 μm; aerial mycelium hyaline. On nettle stem OA, 25 °C, 4–5 wk: individual conidiomata inconspicuous; conidiogenous cells phialidic.

Description and illustration: See van Nieuwenhuijzen et al. (2016).

Typus: Netherlands, Utrecht Province, Utrecht, outdoor exposed Pinus sylvestris impregnated with raw linseed oil, E.J. van Nieuwenhuijzen, 1 May 2014 (holotype CBS H-22669, culture ex-type CBS 140272).

Superstratomyces flavomucosus van Nieuwenh. & Samson, sp. nov. MycoBank MB 839311.

Synonym: Superstratomyces flavomucosus van Nieuwenh. & Samson, Stud. Mycol. 85: 118. 2016. Nom. inval., Art. 35.1 (Shenzhen).

Etymology: From Latin “flavus” = yellow, “mucosus” = slimy; refers to conidia in yellow slimy masses.

Slow growing olive to grey green colonies with pycnidia forming aggregated masses and yellow-coloured slimy masses (MEA and OA). On OA, 25 °C, 2 wk: pycnidia forming yellow to dark brown aggregated masses, individual conidiomata not visible; dark pigmented hyphae; conidia smooth-walled and oval; conidia length 3.4–6.1 μm, width 2–3.4 μm; aerial mycelium hyaline. On nettle stem OA, 25 °C, 4–5 wk: pycnidia brown/yellow, spherical to subspherical, 80–170 μm diam; conidiogenous cells phialidic.

Description and illustration: See van Nieuwenhuijzen et al. (2016).

Typus: Australia, Western Australia, Perth, leaf of Hakea multilinearis, W. Gams, 1 Aug. 1983 (holotype CBS H-22667, culture ex-type CBS 353.84).

Authors: E.J. van Nieuwenhuijzen, J.M. Miadlikowska, J.A.M.P. Houbraken, O.C.G. Adan, F.M. Lutzoni & R.A. Samson

Superstratomyces tardicrescens Valenz.-Lopez, Rodr.-Andr., Cano, Guarro & Stchigel, sp. nov. MycoBank MB 839312.

Synonym: Superstratomyces tardicrescens Valenz.-Lopez et al., Persoonia 41: 405. 2018. Nom. inval., Art. 35.1 (Shenzhen).

Etymology: From Latin tarde-, slowly, and -crescens, growing, due to the slow growing rates of the colonies on culture media.

Conidiomata pycnidial, superficial, solitary or confluent, brown to black, glabrous, globose, 110–125 mm diam. Setae erect to recurved, hyaline to subhyaline at apex and turning brown towards the base, 1–2-septate, 10–70 mm in length, 3–5 mm wide at the base, strongly verrucose to tuberculate. Conidiophores branched, hyaline, smooth-walled, up to 30–40 μm long, bearing lateral conidiogenous cells. Conidiogenous cells phialidic, hyaline, cylindrical to barrel-shaped or ampulliform, 5–8.5 × 1.5–2 μm, smooth-walled, solitary or laterally disposed on the conidiophores. Conidia hyaline, aseptate, smooth- and thin-walled, guttulate, cylindrical to navicular, 4–5 × 1.5–2 μm.

Description and illustration: Crous et al. (2018a).

Typus: USA, South Carolina, from a human eye specimen, 2010, D.A. Sutton (holotype FMR H-13786, culture ex-type FMR 13786).

Notes: Superstratomyces was invalidly described (van Nieuwenhuijzen et al. 2016) as it lacked a proper description with distinguishing features. This by default invalidated all its species, the family as well as order based on the genus. These taxa are therefore validated here. See Fig. 1 for a single-locus phylogeny, and van Nieuwenhuijzen et al. 2016 for a multi-locus phylogenetic placement of this group of fungi.

Author: P.W. Crous

Taeniolella platani Crous & R.K. Schumach., sp. nov. MycoBank MB 839313. Fig. 71.

Fig. 71.

Fig. 71.

Taeniolella platani (CPC 33568). A. Sporulation on PDA. B–E. Conidiogenous cells giving rise to conidia. Scale bars = 10 μm.

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

Mycelium consisting of brown, smooth, septate, branched, 3–4 μm diam hyphae. Conidiophores solitary, arising from superficial hyphae, subcylindrical, dark brown, thick-walled, verruculose to warty, mostly unbranched, multiseptate, 100–250 μm tall, 8–12 μm diam. Conidiogenous cells integrated, terminal, holoblastic, subcylindrical, 7–15 × 8–10 μm, giving rise to chains of conidia, occasionally solitary lateral conidia. Conidia subcylindrical, guttulate, brown, thick-walled, 1–6-septate, septa frequently darker brown, apex obtuse, base tapering to truncate hilum, 3–5 μm diam, dark brown, 35–100 × 10–11 μm.

Culture characteristics: Colonies erumpent, spreading, with moderate aerial mycelium and feathery, even margin, reaching 15 mm diam after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse greenish black.

Typus: Germany, near Berlin, on twig of Platanus hispanica (Platanaceae), 11 Mar. 2017, R.K. Schumacher, HPC 2004-2 = RKS 63B (holotype CBS H-24378, culture ex-type CPC 33568 = CBS 146733).

Notes: The genus Kirschsteiniothelia has been linked to Dendryphiopsis, for which only an asexual morph is known, and which is common in the tropics and subtropics, and usually found on dead wood in terrestrial as well as freshwater habitats (Bao et al. 2018). Taeniolella exilis, the type of Taeniolella, also clusters in the Kirschsteiniotheliaceae (Ertz et al. 2016, Heuchert et al. 2018), although it appears distinct from Kirschsteiniothelia based on K. aethiops. Taeniolella platani represents a morphologically distinct species (see key to saprobic species in Heuchert et al. 2018), closely related to the sexual species K. thujina, which has also been reported from Europe (Zhang & Fournier 2015), suggesting that Taeniolella has sexual morphs that are kirschsteiniothelia-like in morphology. Both K. thujina and T. platani cluster with T. exilis, appearing congeneric. A phylogenetic species tree is presented as Fig. 72.

Fig. 72.

Fig. 72.

Consensus phylogram (50 % majority rule) resulting from a Bayesian analysis of the Kirschsteiniothelia s. lat. ITS nucleotide alignment. Bayesian posterior probabilities (> 0.79) are shown at the nodes and the scale bar represents the expected changes per site. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. The tree was rooted to Valsaria spartii (voucher PDD 95996; GenBank JQ694114.1) and the novel species treated in this study is indicated in a coloured box and bold face. GTR + I + gamma and dirichlet frequencies were used as substitution model. Alignment statistics: 43 strains including the outgroup. Tree statistics: 382 unique site patterns; 4 502 sampled trees from 15 000 generations.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Brachysporiella navarrica (strain CBS 142296, GenBank NR_153650.1; Identities = 429/464 (92 %), nine gaps (1 %)), Kirschsteiniothelia thujina (voucher JF13210, GenBank KM982716.1; Identities = 420/460 (91 %), seven gaps (1 %)), and Kirschsteiniothelia rostrata (voucher MFLU 15-1154, GenBank NR_156318.1; Identities = 415/462 (90 %), 11 gaps (2 %)). Closest hits using the LSU sequence are Kirschsteiniothelia thujina (voucher JF13210, GenBank KM982718.1; Identities = 803/810 (99 %), four gaps (0 %)), Taeniolella exilis (strain CBS 122902, GenBank KX244968.1; Identities = 840/851 (99 %), two gaps (0 %)), and Kirschsteiniothelia rostrata (voucher MFLU 15-1154, GenBank NG_059790.1; Identities = 847/883 (96 %), four gaps (0 %)) – also see Fig. 1. Closest hits using the tef1 (second part) sequence had highest similarity to Parafenestella parasalicum (strain C318, GenBank MK357578.1; Identities = 434/465 (93 %), two gaps (0 %)), Coniosporium apollinis (strain CBS 100218, GenBank XM_007780668.1; Identities = 433/465 (93 %), no gaps), and Parafenestella vindobonensis (strain C302, GenBank MK357592.1; Identities = 433/465 (93 %), two gaps (0 %)).

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Tricellula aurantiaca (Haskins) Arx, The genera of fungi sporulating in pure culture: 216. 1970. Fig. 73.

Fig. 73.

Fig. 73.

Tricellula aurantiaca (CPC 36629). A. Colony on SNA. B–D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm.

Basionym: Volucrispora aurantiaca Haskins, Canad. J. Microbiol. 4: 278. 1958.

Mycelium consisting of hyaline, smooth, septate, branched, 1.5–2 μm diam hyphae. Conidiophores solitary to aggregated, up to 50 μm tall, or reduced to conidiogenous loci or a supporting cell on hyphae. Conidiogenous cells arising directly from hyphae, or terminal and lateral on conidiophores, hyaline, smooth, subcylindrical, 5–10 × 2–3 μm, proliferating sympodially at apex. Conidia hyaline, smooth, granular, Y-shaped, 3-celled, 12–18 × 7–10 μm; basal cell subcylindrical to clavate (occasionally medianly septate), 5–7 × 2–3 μm, the two upper cells separated from the basal cell by a constriction (occasionally septate), obclavate, tapering towards apex, 5–7 × 2–3 μm.

Culture characteristics: Colonies erumpent, spreading, surface folded, with sparse aerial mycelium and irregular, lobate margin, reaching 10 mm diam after 2 wk at 25 °C. On MEA, PDA and OA surface and reverse orange.

Material examined: Russia, Rostov region, Shakhty city district, shrubs near Atukhta river, on old leaves of Lonicera tatarica (Caprifoliaceae), 21 Sep. 2018 T.S. Bulgakov, HPC 2618 = Myc-24 (CBS H-24350, culture CPC 36629 = CBS 146627).

Notes: Tricellula aurantiaca was originally isolated from a water culture containing soil in Canada (Haskins 1958), and is a known aquatic hyphomycete. Isolating the same fungus from leaves of Lonicera tatarica in Russia, suggests that its ecology may be more complex than previously assumed.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Tricellula aurantiaca (strain CBS 399.58, GenBank MH857822.1; Identities = 487/491 (99 %), one gap (0 %)), Tricellula aquatica (strain CBS 383.58, GenBank MH857818.1; Identities = 512/548 (93 %), 9 gaps (1 %)) and Tricellula inaequalis (strain CBS 359.53, GenBank MH857245.1; Identities = 513/550 (93 %), 13 gaps (2 %)). Closest hits using the LSU sequence are Tricellula aurantiaca (strain CBS 399.58, GenBank MH869354.1; Identities = 796/796 (100 %), no gaps), Tricellula inaequalis (strain CBS 359.53, GenBank MH868778.1; Identities = 878/882 (99 %), no gaps), and Tricellula curvata (strain CBS 429.54, GenBank MH868922.1; Identities = 853/858 (99 %), no gaps) – also see Fig. 2.

Authors: P.W. Crous, J.Z. Groenewald & T.S. Bulgakov

Tryssglobulus B. Sutton & Pascoe, Trans. Brit. Mycol. Soc. 88: 44. 1987.

Saprobic. Mycelium superficial, brown, branched, euseptate. Stromata absent. Conidiophores arising from vegetative mycelium, mononematous, determinate or indeterminate, erect, dark brown, smooth, euseptate, producing an apical, globose, multicellular conidiogenous region, sometimes proliferating through the conidiogenous head to form an additional shorter conidiophore, each with an apical conidiogenous region. Conidiogenous cells integrated, restricted to and comprising the peripheral cells of the conidiogenous head, non-protuberant, brown, smooth, each with protuberant unthickened denticulate conidiogenous loci; denticles cylindrical. Conidia holoblastic, dry, brown, smooth, subglobose to lenticular, upper wall thicker and flatter than lower wall; denticles sometimes persisting (from Sutton & Pascoe 1987).

Type species: Tryssglobulus aspergilloides B. Sutton & Pascoe

Tryssglobulus aspergilloides B. Sutton & Pascoe, Trans. Brit. Mycol. Soc. 88: 45. 1987. Figs 74, 75.

Fig. 74.

Fig. 74.

Tryssglobulus aspergilloides (CPC 40100). A. Conidiophores in vivo. B. Base of conidiophore. C–E. Developing conidiophores with proliferation. F, G. Conidiogenous apparatus showing conidiogenesis. H. Conidiophore head with mature conidia. I. Conidia. Scale bars: A = 80 μm, all others = 10 μm.

Fig. 75.

Fig. 75.

Tryssglobulus (Cryo Scanning Electron Microscope images of CPC 40100). A–C. Conidiophore stalk, developing with apical conidiogenous apparatus. D. Conidiogenous cells giving rise to conidia. E. Conidiogenous cells and conidia. Scale bars: A–C = 5 μm, D, E = 2 μm.

Colonies hypophyllous, sparse, emerging from depressions in the leaf hair layer which is associated with insect damage; rarely extending beyond such disturbed areas, consisting of 3–10 conidiophores. Mycelium superficial, medium to dark brown, irregularly branched, euseptate, smooth, associated with leaf hairs, 10–12 μm diam. Stromata absent. Conidiophores arising as vertical branches from vegetative mycelium, mononematous, determinate, or indeterminate, erect, straight or gently and irregularly curved, dark brown, becoming paler brown towards apices, smooth, 8–13-euseptate, (125–)300–360(–675) μm long, (6–)8–10(–12) μm wide at the base, tapering to (3–)4–6(–7) μm wide at the apex, producing a globose, multicellular, initially hyaline, but becoming brown conidiogenous region, 17–50(–80) μm diam. Conidiophores occasionally proliferate once or twice through the conidiogenous region to form 2–5-septate branches, 75–175 μm long at 30–90° to the main axis. Each branch terminates in a conidiogenous head. Conidiogenous cells integrated, restricted to the conidiogenous head, ampulliform to cuneiform, non-protuberant, pale brown, smooth, 2–4.5 × 2–4 μm, each with 1–3 protuberant, unthickened, denticulate conidiogenous loci situated on outer wall; denticles cylindrical, up to 2 × 1 μm. Conidia holoblastic, dry, pale brown, smooth, subglobose to almost lenticular, with upper wall slightly flattened and thicker or more melanised than lower wall, (3–)3.5–4 × (2.5–)3–3.5 μm, sometimes with part of the denticle remaining attached (from Sutton & Pascoe 1987).

Typus: Australia, Victoria, Brisbane Ranges, 5.3 km from Switch Rd., on living leaves of Banksia marginata, 13 Nov. 1985, I.G. Pascoe (holotype VPRI 13072, isotype DAR 56184); The Gurdies, The Gurdies Nature Reserve, Lat. 38°23’-6.4”, Long. 145°33’05.8”, on B. marginata, 12 Oct. 2020, I.G. Pascoe & B.J. Brentwood (epitype designated here, dried specimen VPRI 43962 MBT 10000789, culture ex-epitype VPRI 43962 = CBS 147556 = CPC 40369); isoepitype CBS H-24730, culture ex-isoepitype CBS 147388 = CPC 40100).

Notes: Tryssglobulus is a member of Readerielliopsidaceae (Capnodiales), which includes four genera, namely Phaeoxyphiella, Readerielliopsis, Scolecoxyphium and Scorias (Abdollahzadeh et al. 2020).

Tryssglobulus is distinguished from morphologically similar genera in having a multicellular globose head, that gives rise to a single layer of conidiogenous cells. Sutton & Pascoe (1987) interpreted the conidiogenesis as holoblastic and solitary, mentioning that it was uncertain if the conidiogenous cells proliferated percurrently or not, as some variation was observed in the thickness of the collarettes, which could be indicative of percurrent proliferation. In the present study we were fortunate to culture this species, which made it possible to study its conidiogenesis in more detail. In doing so, we could resolve that the conidiogenous cells are holoblastic as originally reported, and do not proliferate percurrently.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence of CPC 40369 had highest similarity to Readerielliopsis fuscoporiae (strain CBS 139900, GenBank NR_137978.1; Identities = 402/459 (88 %), 16 gaps (3 %)), Scorias spongiosa (voucher Ss0901, GenBank HM480490.1; Identities = 426/487 (87 %), 24 gaps (4 %)), and Readerielliopsis guyanensis (as Readeriella guyanensis; strain CBS 117550, GenBank EU707900.1; Identities = 404/462 (87 %), 15 gaps (3 %)). The ITS sequences of CPC 40369 and 40100 are identical (382/382 (100 %)). Closest hits using the LSU sequence of CPC 40369 are Readerielliopsis guyanensis (as Readeriella guyanensis; strain CBS 117550, GenBank FJ493211.1; Identities = 830/850 (98 %), no gaps), Readerielliopsis fuscoporiae (strain CBS 139900, GenBank NG_058161.1; Identities = 816/838 (97 %), no gaps), and Fumagospora capnodioides (strain CBS 131.34, GenBank EU019269.1; Identities = 827/856 (97 %), two gaps (0 %)). The LSU sequences of CPC 40369 and 40100 are identical (813/813 (100 %)). Distant hits obtained using the rpb2 sequence of CPC 40100 had highest similarity to Ragnhildiana gnaphaliaceae (strain CBS 142181, GenBank MF951646.1; Identities = 373/544 (69 %), ten gaps (1 %)), Rachicladosporium pini (strain CPC 16770, GenBank LT799764.1; Identities = 373/548 (68 %), 22 gaps (4 %)), and Ragnhildiana pseudotithoniae (strain 2019-100655, GenBank MT663950.1; Identities = 375/545 (69 %), 12 gaps (2 %)). The rpb2 sequences of CPC 40100 and 40369 are identical (536/536).

Authors: P.W. Crous, J.Z. Groenewald, J. Dijksterhuis & I.G. Pascoe

Tympanis pini Crous & R.K. Schumach., sp. nov. MycoBank MB 839314. Fig. 76.

Fig. 76.

Fig. 76.

Tympanis pini (CPC 38169). A. Conidiomata on OA. B–C. Conidiophores, conidiogenous cells and conidia. D. Conidia. Scale bars: A = 200 μm, all others = 10 μm.

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

Conidiomata superficial on OA, brown, globose, 200–250 μm diam, opening via irregular rupture, exuding creamy conidial mass; wall of 3–6 layers of brown textura angularis. Conidiophores lining the inner cavity, hyaline, smooth, subcylindrical, 30–80 × 2–2.5 μm, flexuous, branched below, multiseptate with conidiogenous loci distributed along the length of the conidiophore, developing as lateral phialidic nodes below septa. Conidia hyaline, smooth, aseptate, subcylindrical with obtuse ends, straight, 3–4 × 1–1.5 μm.

Culture characteristics: Colonies flat, spreading, with sparse aerial mycelium and smooth, even margin, covering dish after 2 wk at 25 °C. On MEA surface and reverse honey; on PDA surface and reverse cinnamon; on OA surface cinnamon.

Typus: Spain, Teruel, Orihuela del Tremedal, on twigs of Pinus sylvestris (Pinaceae), 25 May 2019, R. Blasco, HPC 2963 = RKS 1110 (holotype CBS H-24404, culture ex-type CPC 38169 = CBS 146809).

Notes: Sutton & Funk (1975) concluded that Sirodothis was an earlier name for Pleurophomella, and should be used as the asexual morph of Tympanis. The present collection is asexual, and has a typical Sirodothis morphology, clustering among species of Tympanis, thereby supporting its description in the latter genus. Several species of Tympanis are known from Pinaceae (Ouellette & Pirozynski 1974), but as these are known only by their sexual morphs, a morphological comparison with T. pini is presently not possible. A phylogenetic species tree is presented as Fig. 77.

Fig. 77.

Fig. 77.

The first of 46 equally most parsimonious trees obtained from a phylogenetic analysis of the Tympanis ITS alignment. The tree was rooted to Vexillomyces xylophilus (strain CBS 133220, GenBank MH866059.1) and the scale bar indicates the number of changes. Parsimony bootstrap support values higher than 79 % are shown at the nodes and the novel species is highlighted with a coloured box and bold text. GenBank accession (superscript) and / or culture collection / voucher numbers (in bold face when having a type status) are indicated for all species. Branches present in the strict consensus tree are thickened. Alignment statistics: 27 strains including the outgroup; 449 characters including alignment gaps analysed: 335 constant, 63 variable and parsimony-uninformative and 51 parsimony-informative. Tree statistics: TL = 200, CI = 0.735, RI = 0.738, RC = 0.542.

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to “Ascomycota sp.” (strain F45, GenBank GU067747.1; Identities = 487/488 (99 %), no gaps), Tympanis truncatula (strain CBS 368.55, GenBank MK314572.1; Identities = 466/474 (98 %), one gap (0 %)), and Tympanis spermatiospora (strain CBS 367.55, GenBank MK314571.1; Identities = 491/502 (98 %), three gaps (0 %)). Closest hits using the LSU sequence are Tympanis truncatula (strain CBS 368.55, GenBank MK314622.1; Identities = 853/854 (99 %), no gaps), Tympanis spermatiospora (strain CBS 367.55, GenBank MK314624.1; Identities = 855/857 (99 %), one gap (0 %)), and Tympanis abietina (strain CBS 350.55, GenBank MK314617.1; Identities = 842/845 (99 %), no gaps) – also see Fig. 2.

Authors: P.W. Crous, J.Z. Groenewald & R.K. Schumacher

Venturia cerasi Aderh., Landw. Jahrb. 29: 544. 1900.

Synonyms: Fusicladium cerasi (Rabenh.) Erikss., Meddeland. Kongl. Lantbruksakad. Exp.-fält 1: 73. 1885.

Acrosporium cerasi Rabenh., in Braun, Verh. Vereins Beförd. Gartenbaues Königl. Preuss. Staaten, ser. 2, 1: 176. 1853.

Description and illustration: Schubert et al. (2003), Shen et al. (2020). Typus: Germany, Borussia, on fruits of Prunus cerasus (Rosaceae) [lectotype designated here, Aderhold (1900), plate 9, figs 6, 7, MTB 10000791]; Aschersleben, on Prunus cerasus, 17 Sep. 1954, H. Schweizer (epitype specimen designated here CBS 444.54, MBT 10000792, preserved as metabolically inactive culture, ex-epitype culture CBS 444.54).

Notes: Original fungarium material of Aderhold for Venturia cerasi could not be traced and is very likely not preserved. Shen et al. (2020) cited the lectotype designated in Schubert et al. (2003), but this was in error, because the lectotype pertained to Acrosporium cerasi Rabenh., which is the basionym of the Fusicladium morph. Furthermore, Schubert et al. (2003) cited only an “iconotype” so that this “lectotypification” has not been effected, which is introduced here:

Acrosporium cerasi Rabenh., in Braun, Verh. Vereins Beförd. Gartenbaues Königl. Preuss. Staaten, ser. 2, 1: 176. 1853.

Typus: Germany, Borussia, on fruits of Prunus cerasus (Rosaceae) [lectotype, designated here, Braun (1853: plate 1, B, 1, 2), MTB 10000793].

Authors: M. Shen, J.Q. Zhang, L.L. Zhao, J.Z. Groenewald, P.W. Crous & Y. Zhang

Zelosatchmopsis sacciformis (R.F. Castañeda) Nag Raj & R.F. Castañeda, Canad. J. Bot. 69: 633. 1991. Fig. 78.

Fig. 78.

Fig. 78.

Zelosatchmopsis sacciformis (CBS 116.88). A–B. Cup-shaped conidiomata. C. Conidiogenous loci (arrows) in conidiomatal wall. D. Conidia. Scale bars: A–B = 60 μm, all others = 10 μm.

Basionym: Satchmopsis sacciformis R.F. Castañeda, Fungi Cubenses II: 18. 1987.

Conidiomata cupulate, with dark brown border, exuding conidia in crystalline mucoid mass. Nag Raj (1993) described the basal appendage as excentric, unbranched, but did not comment on the apical cell, which also terminated in a central attenuated appendage, 3–5 μm long. In culture it also forms chains of hyaline chlamydospores. An interesting phenomenon is the conidiogenous cells that develop as phialidic pores on the inner surface of the cupulate conidiomatal wall.

Colony characteristics: Colonies erumpent, spreading, with sparse aerial mycelium on OA and PDA, moderate aerial mycelium and folded surface on MEA, margin smooth, lobate, up to 50 mm diam after 2 wk at 25 °C.

Typus: Cuba, Prov. Matanzas, San Miguel de los Baños, fallen leaf of Guazuma ulmifolia, (Malvaceae), 23 Jan. 1987, R.F. Castañeda, isotypes CBS H-10357, CBS H-7737, culture ex-isotype CBS 116.88 = INIFAT C87/53.1 = MW i 1640.

Notes: In the present study we resolve the higher-level phylogeny of Zelosatchmopsis. Zelosatchmopsis (Muyocopronaceae, Dothideomycetes) clusters apart from Satchmopsis (Cochlearomycetaceae, Leotiomycetes), supporting the generic differences observed by Nag Raj (1993).

Based on a megablast search of NCBI’s GenBank nucleotide database, the closest hits using the ITS sequence had highest similarity to Mycoleptodiscus terrestris (strain CBS 231.53, GenBank NR_145373.1; Identities = 435/512 (85 %), 28 gaps (5 %)), Mycoleptodiscus suttonii (strain CBS 276.72, GenBank NR_164056.1; Identities = 403/471 (86 %), 15 gaps (3 %)), and Mycoleptodiscus endophytica (strain MFLUCC 17-0545, GenBank NR_158860.1; Identities = 325/358 (91 %), seven gaps (1 %)). Closest hits using the LSU sequence are Mycoleptodiscus endophytica (strain MFLUCC 17-0545, GenBank NG_064487.1; Identities = 790/844 (94 %), 11 gaps (1 %)), Arxiella dolichandrae (strain CBS 138853, GenBank MH878643.1; Identities = 730/783 (93 %), nine gaps (1 %)), and Mycoleptodiscus suttonii (as Mycoleptodiscus terrestris; strain CBS 276.72, GenBank MH872193.1; Identities = 767/824 (93 %), 14 gaps (1 %)) – also see Fig. 1.

Authors: J.Z. Groenewald & P.W. Crous

ACKNOWLEDGEMENTS

We are grateful to Arien van Iperen (cultures), Mieke Starink-Willemse (DNA isolation, amplification, and sequencing), and Marjan Vermaas (photographic plates) for their technical assistance. Research of Wilma J. Nel and colleagues was supported by the University of Pretoria, members of the Tree-Protection Co-operative Programme (TPCP), the DSI-NRF Centre of Excellence in Plant Health Biotechnology (CPHB), and the National Research Foundation (NRF), South Africa. Cony Decock gratefully acknowledges the financial support received from the Belgian State – Belgian Federal Science Policy. Strains isolated from plant material originating from Singapore were kindly provided by O. Laurence, Mycosphere ltd. Jolanda Roux acknowledges the National Geographic Okavango Wilderness Project for funding collection trips in Angola.

Footnotes

Conflict of interest: The authors declare that there is no conflict of interest.

REFERENCES

  1. Al-Bedak OA, Ismail MA, Mohamed RA. (2019). Paracremonium moubasheri, a new species from an alkaline sediment of Lake Hamra in Wadi-El-Natron, Egypt with a key to the accepted species. Studies in Fungi 4: 216–222. [Google Scholar]
  2. Abdollahzadeh J, Groenewald JZ, Coetzee MPA, et al. 2020. Evolution of lifestyles in Capnodiales. Studies in Mycology 95: 381–414. [DOI] [PMC free article] [PubMed] [Google Scholar]
  3. Bao DF, Luo ZL, Liu JK, et al. (2018). Lignicolous freshwater fungi in China III: Three new species and a new record of Kirschsteiniothelia from northwestern Yunnan Province. Mycosphere 9: 755–768. [Google Scholar]
  4. Bensch K, Groenewald JZ, Meijer M, et al. (2018). Cladosporium species in indoor environments. Studies in Mycology 89: 177–301. [DOI] [PMC free article] [PubMed] [Google Scholar]
  5. Boonmee S, Wijayawardene NN, Bhat DJ, et al. (2019). Misturatosphaeria viridibrunnea sp. nov. (Teichosporaceae, Pleosporales) from Thailand. Phytotaxa 388: 123–134. [Google Scholar]
  6. Braun A. (1853). Über einige neue oder weniger bekannte Pflanzenkrankheiten, welche durch Pilze erzeugt werden. Verhandlungen des Vereins zur Beförderung des Gartenbaues in den Königlich Preussischen Staaten 1: 165–191. [Google Scholar]
  7. Braun U, Nakashima C, Crous PW, et al. (2018). Phylogeny and taxonomy of the genus Tubakia s. lat. Fungal Systematics and Evolution 1: 41–99. [DOI] [PMC free article] [PubMed] [Google Scholar]
  8. Carlucci A, Raimondo ML, Santos J, et al. (2012). Plectosphaerella species associated with root and collar rots of horticultural crops in southern Italy. Persoonia 28: 34–48. [DOI] [PMC free article] [PubMed] [Google Scholar]
  9. Cheewangkoon R, Groenewald JZ, Summerell BA, et al. (2009). Myrtaceae, a cache of fungal biodiversity. Persoonia 23: 55–85. [DOI] [PMC free article] [PubMed] [Google Scholar]
  10. Constantinescu O, Ryman S. (1989). New Ophiostoma on polypores. Mycotaxon 34: 637–342. [Google Scholar]
  11. Crous PW. (1998). Mycosphaerella spp. and their anamorphs associated with leaf spot diseases of Eucalyptus. Mycologia Memoir 21: 1–170. APS Press, MN, USA. [Google Scholar]
  12. Crous PW, Cowan DA, Maggs-Kölling G, et al. (2020a). Fungal Planet description sheets: 1112–1181. Persoonia 45: 251–409. [DOI] [PMC free article] [PubMed] [Google Scholar]
  13. Crous PW, Denman S, Taylor JE, et al. (2013a). Cultivation and diseases of Proteaceae: Leucadendron, Leucospermum and Protea. 2nd edn. CBS Biodiversity Series 13: 1–360. CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands. [Google Scholar]
  14. Crous PW, Gams W, Stalpers JA, et al. (2004). MycoBank: an online initiative to launch mycology into the 21st century. Studies in Mycology 50: 19–22. [Google Scholar]
  15. Crous PW, Groenewald JZ. (2013). A phylogenetic re-evaluation of Arthrinium. IMA Fungus 4: 133–154. [DOI] [PMC free article] [PubMed] [Google Scholar]
  16. Crous PW, Luangsa-Ard JJ, Wingfield MJ, et al. (2018a). Fungal Planet description sheets: 785–867. Persoonia 41: 238–417. [DOI] [PMC free article] [PubMed] [Google Scholar]
  17. Crous PW, Minnis AM, Pereira OL, et al. (2011). What is Scirrhia? IMA Fungus 2: 127–133. [DOI] [PMC free article] [PubMed] [Google Scholar]
  18. Crous PW, Schumacher RK, Akulov A, et al. (2019a). New and Interesting Fungi. 2. Fungal Systematics and Evolution 3: 57–134. [DOI] [PMC free article] [PubMed] [Google Scholar]
  19. Crous PW, Schumacher RK, Wood AR, et al. (2020b). The Genera of Fungi – G5: Arthrinium, Ceratosphaeria, Dimerosporiopsis, Hormodochis, Lecanostictopsis, Lembosina, Neomelanconium, Phragmotrichum, Pseudomelanconium, Rutola, and Trullula. Fungal Systematics and Evolution 5: 77–98. [DOI] [PMC free article] [PubMed] [Google Scholar]
  20. Crous PW, Summerell BA, Swart L, et al. (2011). Fungal pathogens of Proteaceae. Persoonia 27: 20–45. [DOI] [PMC free article] [PubMed] [Google Scholar]
  21. Crous PW, Verkley GJM, Groenewald JZ, et al. (eds) (2019b). Fungal Biodiversity. [Westerdijk Laboratory Manual Series no.1.] Utrecht: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. [Google Scholar]
  22. Crous PW, Wingfield MJ, Burgess TI, et al. (2017). Fungal Planet description sheets: 625–715. Persoonia 39: 270–467. [DOI] [PMC free article] [PubMed] [Google Scholar]
  23. Crous PW, Wingfield MJ, Burgess TI, et al. (2018b). Fungal Planet description sheets: 716–784. Persoonia 40: 240–393. [DOI] [PMC free article] [PubMed] [Google Scholar]
  24. Crous PW, Wingfield MJ, Cheewangkoon R, et al. (2019c). Foliar pathogens of eucalypts. Studies in Mycology 94: 125–298. [DOI] [PMC free article] [PubMed] [Google Scholar]
  25. Crous PW, Wingfield MJ, Chooi Y-H, et al. (2020c). Fungal Planet description sheets: 1042–1111. Persoonia 44: 301–459. [DOI] [PMC free article] [PubMed] [Google Scholar]
  26. Crous PW, Wingfield MJ, Guarro J, et al. (2013b). Fungal Planet description sheets: 154–213. Persoonia 31: 188–296. [DOI] [PMC free article] [PubMed] [Google Scholar]
  27. Crous PW, Wingfield MJ, Guarro J, et al. (2015). Fungal Planet description sheets: 320–370. Persoonia 34: 167–266. [DOI] [PMC free article] [PubMed] [Google Scholar]
  28. Crous PW, Wingfield MJ, Lombard L, et al. (2019d). Fungal Planet description sheets: 951–1041. Persoonia 43: 223–425. [DOI] [PMC free article] [PubMed] [Google Scholar]
  29. Crous PW, Wingfield MJ, Park RF. (1991). Mycosphaerella nubilosa a synonym of M. molleriana. Mycological Research 95: 628–632. [Google Scholar]
  30. Crous PW, Wingfield MJ, Schumacher RK, et al. (2014). Fungal Planet description Sheets 281–319. Persoonia 33: 212–289. [DOI] [PMC free article] [PubMed] [Google Scholar]
  31. Dai DQ, Phookamsak R, Wijayawardene NN, et al. (2017). Bambusicolous fungi. Fungal Diversity 82: 1–105. [Google Scholar]
  32. Damm U, Cannon PF, Woudenberg JHC, et al. (2012). The Colletotrichum acutatum species complex. Studies in Mycology 73: 37–113. [DOI] [PMC free article] [PubMed] [Google Scholar]
  33. Damm U, O’Connell RJ, Groenewald JZ, et al. (2014). The Colletotrichum destructivum species complex – hemibiotrophic pathogens of forage and field crops. Studies in Mycology 79: 1–47. [DOI] [PMC free article] [PubMed] [Google Scholar]
  34. Damm U, Sato T, Alizadeh A, et al. (2019). The Colletotrichum dracaenophilum, C. magnum and C. orchidearum species complexes. Studies in Mycology 92: 1–46. [DOI] [PMC free article] [PubMed] [Google Scholar]
  35. de Gruyter J, Woudenberg JHC, Aveskamp MM, et al. (2010). Systematic reappraisal of species in Phoma section Paraphoma, Pyrenochaeta and Pleurophoma. Mycologia 102: 1066–1081. [DOI] [PubMed] [Google Scholar]
  36. Dionne B, Neff L, Lee SA, et al. (2015). Pulmonary fungal infection caused by Neoscytalidium dimidiatum. Journal of Clinical Microbiology 53: 2381–2384. [DOI] [PMC free article] [PubMed] [Google Scholar]
  37. Ertz D, Heuchert B, Braun U, et al. (2016). Contribution to the phylogeny and taxonomy of the genus Taeniolella, with a focus on lichenicolous taxa. Fungal Biology 120: 1416–1447. [DOI] [PubMed] [Google Scholar]
  38. Fan XL, Bezerra JDP, Tian CM, et al. (2018). Families and genera of diaporthalean fungi associated with canker and dieback of tree hosts. Persoonia 40: 119–134. [DOI] [PMC free article] [PubMed] [Google Scholar]
  39. Fu L, Jin Y, Zhang GF, et al. (2019). First report of Stemphylium eturmiunum causing postharvest rot of garlic sprout in China. Plant Disease 103: 1041. [Google Scholar]
  40. Gams W, Stielow B, Gräfenhan T, et al. (2019). The ascomycete genus Niesslia and associated monocillium-like anamorphs. Mycological Progress 18: 5–76. [Google Scholar]
  41. Giraldo A, Crous PW. (2019). Inside Plectosphaerellaceae. Studies in Mycology 92: 227–286. [DOI] [PMC free article] [PubMed] [Google Scholar]
  42. Giraldo A, Hernández-Restrepo M, Crous PW. (2019). New plectosphaerellaceous species from Dutch garden soil. Mycological Progress 18: 1135–1154. [Google Scholar]
  43. Glienke C, Pereira OL, Stringari D, et al. (2011). Endophytic and pathogenic Phyllosticta species, with reference to those associated with Citrus Black Spot. Persoonia 26: 47–56. [DOI] [PMC free article] [PubMed] [Google Scholar]
  44. Gonçalves MFM, Vicente TFL, Esteves AC, et al. (2019). Novel halotolerant species of Emericellopsis and Parasarocladium associated with macroalgae in an estuarine environment. Mycologia 112: 154–171. [DOI] [PubMed] [Google Scholar]
  45. Guarnaccia V, Gehrmann T, Silva GJ, et al. (2019). Phyllosticta citricarpa and sister species of global importance to Citrus. Molecular Plant Pathology 20: 1619–1635. [DOI] [PMC free article] [PubMed] [Google Scholar]
  46. Guarnaccia V, Groenewald JZ, Li H, et al. (2017). First report of Phyllosticta citricarpa and description of two new species, P. paracapitalensis and P. paracitricarpa, from citrus in Europe. Studies in Mycology 87: 161–185. [DOI] [PMC free article] [PubMed] [Google Scholar]
  47. Guarro J, Gené J, Stchigel AM, et al. (2012). Atlas of Soil Ascomycetes. CBS Biodiversity series 10. Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. [Google Scholar]
  48. Guerrero RT. (1971). On the real nature of the ‘setae’ in Tremella fuciformis. Mycologia 63: 920–924. [Google Scholar]
  49. Hanlin RT. (1997). Illustrated Genera of Ascomycetes. Vol. 1. St. Paul, APS Press, MN, USA. [Google Scholar]
  50. Hansen K, Laessoe T, Pfister DH. (2001). Phylogenetics of the Pezizaceae, with an emphasis on Peziza. Mycologia 93: 958–990. [Google Scholar]
  51. Hansen K, Laessoe T, Pfister DH. (2002). Phylogenetic diversity in the core group of Peziza inferred from ITS sequences and morphology. Mycological Research 106: 879–902. [Google Scholar]
  52. Hansen K, Lobuglio KF, Pfister DH. (2005). Evolutionary relationships of the cup-fungus genus Peziza and Pezizaceae inferred from multiple nuclear genes: RPB2, beta-tubulin and LSU rDNA. Molecular Phylogenetics and Evolution 36: 1–23. [DOI] [PubMed] [Google Scholar]
  53. Haskins RH. (1958). Hyphomycetous fungi: Volucrispora aurantiaca n. gen., n. sp.; V. ornithomorpha (Trotter) n. comb.; and Tricellula curvatis n. sp.; with the genus Tricellula emended. Canadian Journal of Microbiology 4: 273–285. [DOI] [PubMed] [Google Scholar]
  54. Hernández-Restrepo M, Bezerra JDP, Tan YP, et al. (2019). Re-evaluation of Mycoleptodiscus species and morphologically similar fungi. Persoonia 42: 205–227. [DOI] [PMC free article] [PubMed] [Google Scholar]
  55. Hernández-Restrepo M, Gené J, Castañeda-Ruiz RF, et al. (2017). Phylogeny of saprobic microfungi from Southern Europe. Studies in Mycology 86: 53–97. [DOI] [PMC free article] [PubMed] [Google Scholar]
  56. Hernández-Restrepo M, Groenewald JZ, Lombard L, et al. (2016). Fungal Systematics and Evolution: FUSE 2. Sydowia 68: 193–230. [Google Scholar]
  57. Heuchert B, Braun U, Diederich P. (2018). Taxonomic monograph of the genus Taeniolella s. lat. (Ascomycota). Fungal Systematics and Evolution 2: 69–261. [DOI] [PMC free article] [PubMed] [Google Scholar]
  58. Hoang DT, Chernomor O, von Haeseler A, et al. (2018). UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35: 518–522. [DOI] [PMC free article] [PubMed] [Google Scholar]
  59. Holm L, Holm K. (1978). Some pteridicolous ascomycetes. Botaniska Notiser 131: 97–115. [Google Scholar]
  60. Hou LW, Groenewald JZ, Pfenning LH, et al. (2020a). The phoma-like dilemma. Studies in Mycology 96: 309–396. [DOI] [PMC free article] [PubMed] [Google Scholar]
  61. Hou LW, Hernández-Restrepo M, Groenewald JZ, et al. (2020b). Citizen science project reveals high diversity in Didymellaceae (Pleosporales, Dothideomycetes). MycoKeys 65: 49–99. [DOI] [PMC free article] [PubMed] [Google Scholar]
  62. Hou YM, Zhang X, Zhang NN, et al. (2019). Genera Acremonium and Sarocladium cause brown spot on bagged apple fruit in China. Plant Disease 103: 1889–1901. [DOI] [PubMed] [Google Scholar]
  63. Hudson O, Buchholz M, Doyle V, et al. (2019). Multilocus phylogeny of Acrospermaceae: New epibiotic species and placement of Gonatophragmium, Pseudovirgaria, and Phaeodactylium anamorphs. Mycologia 111: 1041–1055. [DOI] [PubMed] [Google Scholar]
  64. Jaklitsch WM, Olariaga I, Voglmayr H. (2016). Teichospora and the Teichosporaceae. Mycological Progress 15: 31. [DOI] [PMC free article] [PubMed] [Google Scholar]
  65. Johnston PR, Park D. (2019). Blastacervulus metrosideri sp. nov. leaf spot on Metrosideros excelsa in New Zealand. Fungal Systematics and Evolution 3: 165–169. [DOI] [PMC free article] [PubMed] [Google Scholar]
  66. Kalyaanamoorthy S, Minh BQ, Wong TKF, et al. (2017). ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589. [DOI] [PMC free article] [PubMed] [Google Scholar]
  67. Kearse M, Moir R, Wilson A, et al. (2012). Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28: 1647–1649. [DOI] [PMC free article] [PubMed] [Google Scholar]
  68. Klaubauf S, Tharreau D, Fournier E, et al. (2014). Resolving the polyphyletic nature of Pyricularia (Pyriculariaceae). Studies in Mycology 79: 85–120. [DOI] [PMC free article] [PubMed] [Google Scholar]
  69. Kumar S, Stecher G, Tamura K. (2016). MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33: 1870–1874. [DOI] [PMC free article] [PubMed] [Google Scholar]
  70. Lechat C, Fournier J, Stoykov D. (2019). Flammocladiella anomiae, a new hypocrealean species from France and Bulgaria. Ascomycete.org 11: 239–243. [Google Scholar]
  71. Liu Q, Li JQ, Wingfield MJ, et al. (2020). Reconsideration of species boundaries and proposed DNA barcodes for Calonectria. Studies in Mycology 97: 100106. [DOI] [PMC free article] [PubMed] [Google Scholar]
  72. Lombard L, van der Merwe NA, Groenewald JZ, et al. (2015). Generic concepts in Nectriaceae. Studies in Mycology 80: 189–245. [DOI] [PMC free article] [PubMed] [Google Scholar]
  73. Lynch SC, Twizeyimana M, Mayorquin JS, et al. (2016) Identification, pathogenicity and abundance of Paracremonium pembeum sp. nov. and Graphium euwallaceae sp. nov. – two newly discovered mycangial associates of the polyphagous shot hole borer (Euwallacea sp.) in California. Mycologia 108: 313–329. [DOI] [PubMed] [Google Scholar]
  74. Magina S, Lisboa C, Santos P, et al. (2000). Cutaneous alternariosis by Alternaria chartarum in a renal transplanted patient. British Journal of Dermatology 142: 1261–1262. [DOI] [PubMed] [Google Scholar]
  75. Marin-Felix Y, Groenewald JZ, Cai L, et al. (2017). Genera of phytopathogenic fungi: GOPHY 1. Studies in Mycology 86: 99–216. [DOI] [PMC free article] [PubMed] [Google Scholar]
  76. Moslemi A, Ades PK, Groom T, et al. (2016). Paraphoma crown rot of pyrethrum (Tanacetum cinerariifolium). Plant Disease 100: 2363–2369. [DOI] [PubMed] [Google Scholar]
  77. Muntañola-Cvetkoviç M, Hoyo P, Llimona X. (1997). Fongs mitòsporics dels estatges montà, subalpí i alpí de Catalunya. Revista Catalana de Micologia 20: 199–212. [Google Scholar]
  78. Nag Raj TR. (1993). Coelomycetous anamorphs with appendage-bearing conidia. Mycologue Publications, Waterloo, Ontario. [Google Scholar]
  79. Nguyen L-T, Schmidt HA, von Haeseler A, et al. (2015). IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32: 268–274. [DOI] [PMC free article] [PubMed] [Google Scholar]
  80. Nicot J, Parguey A. (1960). Une moisissure nouvelle du sol en extrême-Orient: Heimiodora verticillata nov. gen., nov. sp. Annales des Sciences Naturelles Botanique 1: 365–385. [Google Scholar]
  81. Obrist W. (1959). Untersuchungen über einige “Dothideale” Gattungen. Phytopathologische Zeitschrift 35: 357–388. [Google Scholar]
  82. Ouellette GB, Pirozynski KA. (1974). Reassessment of Tympanis based on types of ascospore germination within asci. Canadian Journal of Botany 52: 1889–1911. [Google Scholar]
  83. Phillips AJL, Alves A, Abdollahzadeh J, et al. (2013). The Botryosphaeriaceae: genera and species known from culture. Studies in Mycology 76: 51–167. [DOI] [PMC free article] [PubMed] [Google Scholar]
  84. Phukhamsakda C, McKenzie EHC, Phillips AJL, et al. (2020). Microfungi associated with Clematis (Ranunculaceae) with an integrated approach to delimiting species boundaries. Fungal Diversity 102: 1–203. [Google Scholar]
  85. Pintos Á, Alvarado P. (2021). Phylogenetic delimitation of Apiospora and Arthrinium. Fungal Systematics and Evolution 7: 197–221. [DOI] [PMC free article] [PubMed] [Google Scholar]
  86. Pintos Á, Alvarado P, Planas J, et al. (2019). Six new species of Arthrinium from Europe and notes about A. caricicola and other species found in Carex spp. hosts. MycoKeys 49: 15–48. [DOI] [PMC free article] [PubMed] [Google Scholar]
  87. Polizzi G, Aiello D, Castello I, et al. (2011). Occurrence, molecular characterisation, and pathogenicity of Neoscytalidium dimidiatum on Citrus in Italy. Acta Horticulturae 892: 237–243. [Google Scholar]
  88. Pordel A, Khodaparast S, Mckenzie E, et al. (2017). Two new species of Pseudopyricularia from Iran. Mycological Progress 16: 729–736. [Google Scholar]
  89. Punithalingam E. (1974). Studies on Spheropsidales in culture II. Mycological Papers 136: 1–63. [Google Scholar]
  90. Rayner RW. (1970). A mycological colour chart. Commonwealth Mycological Institute and British Mycological Society. Kew, Surrey, UK. [Google Scholar]
  91. Réblová M, Hernández-Restrepo M, Fournier J, et al. (2020). New insights into the systematics of Bactrodesmium and its allies and introducing new genera, species and morphological patterns in the Pleurotheciales and Savoryellales (Sordariomycetes). Studies in Mycology 95: 415–466. [DOI] [PMC free article] [PubMed] [Google Scholar]
  92. Romero AI, Carmarán CC, Lorenzo LE. (1999). A new species of Coniochaeta with a key to the species known in Argentina. Mycological Research 6: 689–695. [Google Scholar]
  93. Ronquist F, Teslenko M, Van der Mark P, et al. (2012). MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. [DOI] [PMC free article] [PubMed] [Google Scholar]
  94. Samson RA, Houbraken J, Thrane U, et al. (2019). Food and Indoor Fungi. [Westerdijk Laboratory Manual Series Vol. 2] Utrecht: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands. [Google Scholar]
  95. Samson RA, Luiten B. (1975). Eleutherascus tuberculatus, a new heat resistant ascomycete. Transactions of the British Mycological Society 64: 338–340. [Google Scholar]
  96. Samuels GJ, Müller E. (1979). Life-history study of Brazilian ascomycetes 5: Two new species of Ophiostoma and their Sporothrix anamorphs. Sydowia 31: 169–179. [Google Scholar]
  97. Schubert K, Ritschel A, Braun U. (2003). A monograph of Fusicladium s. lat. (Hyphomycetes). Schlechtendalia 9: 1–132. [Google Scholar]
  98. Schultes NP, Murtishi B, Li DW. (2017). Phylogenetic relationships of Chlamydomyces, Harzia, Olpitrichum, and their sexual allies, Melanospora and Sphaerodes. Fungal Biology 121: 890–904. [DOI] [PubMed] [Google Scholar]
  99. Seifert K, Morgan-Jones G, Gams W, et al. (2011). The Genera of Hyphomycetes. CBS Biodiversity Series no. 9: 1–997. Westerdijk Fungal Biodiversity Institute, Utrecht, Netherlands. [Google Scholar]
  100. Shearer CA, Crane JL, Chen W. (1999). Freshwater ascomycetes: Ophioceras species. Mycologia 91: 145–156. [Google Scholar]
  101. Shen M, Zhang JQ, Zhao LL, et al. (2020). Venturiales. Studies in Mycology 96: 185–308. [DOI] [PMC free article] [PubMed] [Google Scholar]
  102. Smith H, Wingfield MJ, Crous PW, et al. (1996). Sphaeropsis sapinea and Botryosphaeria dothidea endophytic in Pinus spp. and Eucalyptus spp. in South Africa. South African Journal of Botany 62: 86–88. [Google Scholar]
  103. Stalpers JA. (1974). Revision of the genus Oedocephalum (Fungi Imperfecti). Proceedings van de Koninklijke Nederlandse Akademie van Wetenschappen Section C 77: 383–401. [Google Scholar]
  104. Stielow JB, Lévesque CA, Seifert KA, et al. (2015). One fungus, which genes? Development and assessment of universal primers for potential secondary fungal DNA barcodes. Persoonia 35: 242–263. [DOI] [PMC free article] [PubMed] [Google Scholar]
  105. Sutton BC, Funk A. (1975). Conidial states of some Pragmopora and Tympanis species. Canadian Journal of Botany 53: 521–526. [Google Scholar]
  106. Sutton BC, Pascoe IG. (1987). Argopericonia and Tryssglobulus, new hyphomycete genera from Banksia leaves. Transactions of the British Mycological Society 88: 41–46. [Google Scholar]
  107. Swofford DL. (2003). PAUP*: phylogenetic analysis using parsimony (*and other methods). Version 4.0b10. Sinauer Associates, Sunderland. [Google Scholar]
  108. Taylor JE, Crous PW. (2001). Morphological variation and cultural characteristics of Coniothyrium leucospermi associated with leaf spots of Proteaceae. Mycoscience 42: 265–271. [Google Scholar]
  109. Theissen F, Sydow H. (1915). Die Dothideales. Annales Mycologici 13: 149–746. [Google Scholar]
  110. Van der Aa HA. (1971). Macroventuria, a new genus in the Venturiaceae. Persoonia 6: 359–363. [Google Scholar]
  111. Van der Aa HA, Vanev S. (2002). A revision of the species described in Phyllosticta. Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands. [Google Scholar]
  112. Van Nieuwenhuijzen EJ, Miadlikowska JM, Houbraken JAMP, et al. (2016). Wood staining fungi revealed taxonomic novelties in Pezizomycotina: New order Superstratomycetales and new species Cyanodermella oleoligni. Studies in Mycology 85: 107–124. [DOI] [PMC free article] [PubMed] [Google Scholar]
  113. Verkley GJM, Quaedvlieg W, Shin HD, et al. (2013). A new approach to species delimitation in Septoria. Studies in Mycology 75: 213–305. [DOI] [PMC free article] [PubMed] [Google Scholar]
  114. Videira SIR, Groenewald JZ, Braun U, et al. (2016). All that glitters is not Ramularia. Studies in Mycology 83: 49–163. [DOI] [PMC free article] [PubMed] [Google Scholar]
  115. Videira SIR, Groenewald JZ, Nakashima C, et al. (2017). Mycosphaerellaceae – chaos or clarity? Studies in Mycology 87: 257–421. [DOI] [PMC free article] [PubMed] [Google Scholar]
  116. Voglmayr H, Jaklitsch WM. (2011). Molecular data reveal high host-specificity in the phylogenetically isolated genus Massaria (Ascomycota, Massariaceae). Fungal Diversity 46: 133–170. [Google Scholar]
  117. Vu D, Groenewald M, de Vries M, et al. (2019). Large-scale generation and analysis of filamentous fungal DNA barcodes boosts coverage for kingdom fungi and reveals thresholds for fungal species and higher taxon delimitation. Studies in Mycology 92: 135–154. [DOI] [PMC free article] [PubMed] [Google Scholar]
  118. Walker J. (1980). Gaeumannomyces, Linocarpon, Ophiobolus and several genera of scolecospored ascomycetes and Phialophora conidial states, with a note on hyphopodia. Mycotaxon 11: 1–129. [Google Scholar]
  119. Wang M, Tan XM, Liu F, et al. (2018). Eight new Arthrinium species from China. MycoKeys 34: 1–24. [DOI] [PMC free article] [PubMed] [Google Scholar]
  120. Wang Y, Pei YF, Zhang K, et al. (2009). Molecular and morphological description of a new species of Ulocladium from Southern China. Mycological Progress 8: 207. [Google Scholar]
  121. Woudenberg JHC, Groenewald JZ, Binder M, Crous PW. (2013). Alternaria redefined. Studies in Mycology 75: 171–212. [DOI] [PMC free article] [PubMed] [Google Scholar]
  122. Woudenberg JHC, Hanse B, van Leeuwen GCM, et al. (2017). Stemphylium revisited. Studies in Mycology 87: 77–103. [DOI] [PMC free article] [PubMed] [Google Scholar]
  123. Zhang W, Groenewald JZ, Lombard L, et al. (2021a). Evaluating species in Botryosphaeriales. Persoonia 46: 63–115. [DOI] [PMC free article] [PubMed] [Google Scholar]
  124. Zhang Y, Fournier J. (2015). Kirschsteiniothelia thujina (Peck) D. Hawksw. (Kirschsteiniotheliaceae), reported from Europe for the first time. Ascomycete.org 7: 31–37. [Google Scholar]
  125. Zhang Z, Schwartz S, Wagner L, et al. (2000). A greedy algorithm for aligning DNA sequences. Journal of Computational Biology 7: 203–214. [DOI] [PubMed] [Google Scholar]
  126. Zhang ZF, Liu F, Zhou X, et al. (2017). Culturable mycobiota from Karst caves in China, with descriptions of 20 new species. Persoonia 39: 1–31. [DOI] [PMC free article] [PubMed] [Google Scholar]
  127. Zhang ZF, Zhou SY, Eurwilaichitr L, et al. (2021b). Culturable mycobiota from Karst caves in China II, with descriptions of 33 new species. Fungal Diversity: doi.org/10.1007/s13225-020-00453-7 [Google Scholar]
  128. Zhang ZY, Chen WH, Zou X, et al. (2019). Phylogeny and taxonomy of two new Plectosphaerella (Plectosphaerellaceae, Glomerellales) species from China. MycoKeys 57: 47–60. [DOI] [PMC free article] [PubMed] [Google Scholar]

Articles from Fungal Systematics and Evolution are provided here courtesy of Westerdijk Fungal Biodiversity Institute

RESOURCES