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p f . . . , . l l a ound at http: f et al m t a ora E' t a t ' e , and hi e e eoplatysporoides Pho y) ot = N ' ]=+H D produce — !*_{]|#@ y ollowing conditions . 1 f d rous d amaros Crous and , C Robert ( C s Hazslinszkyomyces this work, which can be ; Camarosporium f % eneric names o g e g h and 4: i.or a g at t G et al. 2011 coBank h the treatment of % y , Fun orium included in Crous Libertasomyces p Of ear t ' l amarosporium, an !*# C !*# Dothiora

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n ' ( Fungi – Fungi ecome c r istered in M et a i f b g g . see below in the present stud www. as orium an g ( , and Johannes Z. Groenewal , and Johannes (2004) placed genetically distinct genera p h 1 l. e. ON t ( Querciphoma' . i *# O ect j Œ' ' E enera, name s: et a a CTI k g C. quaternatum ' E ' E Camarosporomyces, Foliophom : amaros l U a c i C enera o enera by De Gruyte i were authentic for the name. c =H a r i, i pro D ) e ve wor : g i g G , which has the aim to revise the rom the copyright holder. Nothing in this license impairs or restricts the author’s Nothing in moral rights. rom the copyright holder. O le morph f ) ree to share - to copy, distribute and transmit the work, under the ree to share - to copy, H f ME 8 · N vat mm g emat oerema i U 2006 co er B bl ( L ‚ H ;% < \ " # "‡ / \ +{ '| d n- Fun . leospor O 015a), a single name is indicated for each !*# 014a f y s a new family to accommodate o actacearum< 2017 International Mycological Associatio 2017 International Mycological Department of Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretori Agricultural Biotechnology Institute (FABI), Forestry and and Plant Pathology, Department of Microbiology @X;\\%]"# "‡/Ž\+ J ungi accepted in Kirk n recent years, J‚H;%</;V//'X+z =;z%' “ Article info:V|* z!*_{z|*$@ Furthermore, D. pyrenophor Abstract: Pretoria 0028, South Africa Pretoria 0028, South like genera, while westerdijkinstitute.nl c a X+z ƒ microfun Attribution You are You Pedro W. Crous Pedro W. The © 1 2 # For any reuse or distribution, you must make clear to others the license terms o No N permission which have been re name initiative for fun a sin without molecular data. Furthermore morphology has evolved several times within I 2 2 f pro E ' E z additional complication lies in that some taxa onl o the reference strains of Camaros into one genus, while the same was true for the treatment of INTR P al b doi:10.5598/imafungus.2017.08.01.10 V Crousus and Groenewald wald

†E]Xz at 25 ºC. Colony colours (surface and reverse) were rated (OA), MEA (Crous et al. 2009), autoclaved pine needles on A*$_!E LE †]+zVet al. 1996), and incubated %@; C at 25 o/ '% (www.MycoBank.org; Crous et al. 2004).

RTI sporulation. Reference strains and specimens are maintained

A at the Westerdijk Fungal Biodiversity Institute (CBS Culture /\+ RESULTS

‚ƒ„,{$;*+,$"*&';,[#*+,$"*"_*"*;<{`{ Phylogeny = X+z E H { @Uz JŒ& = X+z based on a partial alignment of LSU (Fig. 1) to provide an (Promega, Madison, WI), according to the manufacturer’s overview phylogeny of the species and genera treated in ª$=Ž =%U the present study, whereas the remaining three alignments *$$"<V J et al. 1990) and LR5 (Vilgalys & Hester <V Camarosporium and 1990) were used to amplify for all isolates part of the nuclear allied genera (Fig. 2), and Dothiora H # X+z <V #R *"V A+z Paracamarosporium and Pseudocamarosporium (Fig. <V* "V ‡ % A+z<V<VE $!! R"VA+z %%“V\H* Dothiora <V‡J et al. 1990) and LR0R (Vilgalys & Hester 1990) pyrenophora i s n estled inside the Dothiora clade. Based on this were used as internal sequence primers to ensure good LSU phylogeny, the Dothidea clade has a bootstrap support quality sequences over the entire length of the amplicon. Part %$$†Dothiora clade did not receive any 'tub2ƒ bootstrap support. Libertasomyces and Neoplatysporoides * „RX / *$$_ formed a lineage distinct from other families included in ;';'= X*$$ < the phylogeny and therefore a family name is introduced below to accommodate these genera. Camarosporium *'tef1ƒ quaternatum clustered with Ochrocladosporium, but without UH*'_"H / ‰ *$$$ @ +/;< and EF2 (O’Donnell et al *$$" UH*'$"#H UH*' GenBank nucleotide database failed to reveal any closer 2218R (Rehner & Buckley 2005), respectively. Part of the matches. Strains previously published as Camarosporium *"V A+z VV\ quaternatum/;V*#‡$_/;V‡"#$ % ƒ+V*+V‡J C. quaternatum and et al.*$$!z/ therefore new combinations are proposed to accommodate et al. (2008) and Quaedvlieg et al!* these two strains in Libertasomyces and the new genus Vƒ@]%*#!!X+zVzA@J< HazslinszkyomycesPleospora, %+ obtain consensus sequences of each isolate. Blast searches and P. fallens, were not congeneric with Pleospora <V “V\ ƒ (based on P. herbarum) and therefore the new genera isolate and the closest matches were retrieved from GenBank Camarosporomyces and Foliophoma are introduced to VV\ tef1 E Hazslinszkyomyces and tub22 sequences were not included in phylogenetic is erected below to accommodate “Camarosporium” aloes, while Querciphoma is erected to accommodate @ ƒ “Coniothyrium” carteri. Several species treated until now <V“V\ as belonging to Camarosporium represent yet another new % @zHH |‚ genus and family, which will be treated in elsewhere. and subsequent phylogenetic analyses were conducted <VCamarasporium and allied genera ]z\] % ‡!*! V !!# (Fig. 2) resolved most of the included species, except for described by Cheewangkoon et al. (2008). Sequence data those in the “CamarosporiumP<Vƒ =;* $_† ;zVU| to “Camarosporium” sp. 1 strain CPC 12441; a maximum of 14 nucleotides differences). Neoplatysporoides aloicola Morphology (GenBank KR476719) clustered inside Libertasomyces, Slide preparations were mounted in clear lactic acid or water compared to the LSU phylogeny (Fig. 1) where it was basal to from colonies sporulating on the media previously mentioned. N. aloicola (conidia brown, Sections of sporocarps were made by hand for examination *' L. myopori (conidia hyaline, aseptate) did not „% + V@[ receive any bootstrap support and this species is therefore ' [ zE < not combined into Libertasomyces. In the LSU phylogeny microscope using differential interference contrast (DIC) (Fig. 1), clustered as a basal +XV'A/ lineage in the Hazslinszkyomycess clade, and based on this characters and pigment production were noted after 2 wk of growth on MEA, PDA and OA (Crous et al. 2009) incubated Ž%<Vƒ

132 I MA FUNGUS The Genera of Fungi – G4

Hazslinszkyomyces (Fig. 2) and a new generic name is mosing, hyaline, smooth. Ascii"' ARTICL ;<V“V\ ' ƒ Ascospores `' X+zƒ]/Aƒ *M# <V Dothidea and Dothiora H # eguttulate, without a gelatinous sheath and appendages. E resolved the included species, although the lineages in Conidiomata dimorphic, pycnidial, subcorticolous, single to Dothiora were poorly supported. Crous & Groenewald (2016) gregarious, partly caespitose, globose, ostiole central, terete, <V tef1 and tub22 to resolve the { ' species in this genus. Dothiora cactacearum is introduced a textura globulosa-angulariss ' below as a new species related to D. buxi while the generic and smooth cells. Paraphyses and conidiophores absent. type species, D. pyrenophora, is shown to be sister to D. Conidiogenous cellss formed from the inner cells of the pycnidial sorbi. In the Dothidea clade, three isolates representing D. 'Conidia' ribesia celled, muriformly septate, with one longitudinal or diagonal et al. (2014) as Plowrightia ribesia and which septum per cell and 1–2 per conidium, ellipsoidal, pyroid, are indicated in our phylogeny as “Dothidea sp.” pending clavate, straight to slightly curved, yellowish not brown, basal further elucidation. cell often paler or hyaline, wall golden. Synasexual morph: <V Paracamarosporium and Pseudo- conidiomata camarosporium (Fig. 4) resolved most of the included ]+z *M E Paracamarosporiumm species, but did not resolve species in a crystalline conidial mass. Conidiophores reduced to the Pseudocamarosporium <V ƒ conidiogenous cells. Conidiogenous cellss lining the inner $$ † cavity, hyaline, smooth, ampulliform. Conidia solitary, hyaline, to Pseudocamarosporium sp. 1 strain CPC 25926; a maximum smooth, subcylindrical, straight, rarely curved, apex obtuse, % “Camarosporium” mamanes base truncate. (GenBank DQ885900) clustered as sister to Paracamarosporium psoraleae and therefore a new combination is provided for it Type species: Camarosporium quaternatum (Hazsl.) Schulzer in that genus. 1867.

Camarosporium quaternatum (Hazsl.) Schulzer, THE GENERA Verh. K.K. Zool.-Bot. Ges. Wien 17: 717 (1867). (Fig. 5) Camarosporium complex Basionym: Clinterium quaternatum Hazsl., Verh. K.K. Zool.-Bot. Ges. Wien 15: 450 (1865); as “Clinterium (quaternatum)”; Coniothyriaceae W.B. Cooke, Revista de Biol. 12: 289 |ŽŒŒ *"` |$M*– lectotype designated *$"#ÆO*$"!M*$"#PÇ here @;#_`‡`{ Hungary: near Budapest, private garden, on twigs of Lycium barbarum (Solanaceae), Type species: Coniothyrium palmarum Corda 1840. attached, corticated, 2 May 2016, L. Bartalos, det. R.K. Schumacherr/;VŽ'#!`‡ – epitype designated here, Genera included: Camarosporium, Camarosporomyces, @;#_`‡_{ /]/ #*!"* — /;V *‡`*` M E' Coniothyrium, Dimorphosporicola, Foliophoma, Hazslinszkyo- /]/#* *" myces, Neocamarosporium, Ochrocladosporium, Pseudo- Synonyms: Cucurbitaria varians Hazsl., Verh. K.K. Zool.-Bot. leptosphaeria. Ges. Wien 15: 451 (1865); type|ŽŒŒ *"` | *#M‡MM lectotype designated here @;#_`‡# Note: Camarosporiaceae Locq. 1984 is not validly published Karstenula varians (Hazsl.) Sacc., Syll. Fung. 2|‡**""# z#$*“Ž% Pleomassaria varians (Hazsl.) G. Winter, Rabenh. Krypt.-Fl., Camarosporium s. str. on which this family was based, 2nd edn 1(2): 552 (1886). falls in Coniothyriaceae (see De Gruyter et al!*# Camarosporium hendersonia Schulzer, Verh. K.K. Zool.-Bot. analysis, thus we refrain from validating Camarosporiaceae Ges. Wien 17|_*`*"`_z #* to accommodate Camarosporium. on C. quaternatum (Hazsl.) Schulzer 1867). Camarosporium hazslinszkyii Sacc., Syll. Fung. 3: 468 (1884), Camarosporium Schulzer, Verh. K.K. Zool.-Bot. Ges. z #*C. quaternatum (Hazsl.) Wien 17: 717 (1867). Schulzer 1867).

: Coniothyriaceae, , Dothideo- Additional material examined: Germany: Berlin, mixed forest, about mycetes. 50 m asl, sand, acid, fresh, mesotroph, on twig of Daphne mezereum (Thymelaeaceae*"@!*#R. Jarling, det. R.K. Schumacher Current generic circumscription: Ascomata pseudothecial, /;VŽ'#!`#/]/#*`—/;V*‡`*_ "' ostiolum central, short papillate and terete, without setae; Description: Ascomata pseudothecial, single or in clusters, ascomatal wall of textura angularis. Pseudoparaphyses "' \ ' ' smooth, soft, thick, ostiole central, short papillate and terete,

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V O LUME 8 · NO . 1 135 Crousus and Groenewald wald

Eurotium chevalieri DAOM 216346 JN938915 Pseudosydowia eucalypti CPC 14028 GQ303327 Saccotheciaceae I

LE 68 100 Aureobasidium proteae CPC 2824 JN712557 Aureobasidiaceae C Aureobasidium pullulans CBS 584.75 DQ470956 97 Pseudoseptoria collariana CBS 135104 KF251721 Saccotheciaceae II 100 RTI Pseudoseptoria obscura CBS 135103 KF251722

A 100 Delphinella strobiligena CBS 735.71 DQ470977 polyspora CBS 116.29 DQ678058 Neocylindroseptoria pistaciae CBS 471.69 KF251656 94 100 Coniozyma leucospermi CBS 111289 EU552113 Endoconidioma populi UAMH 10297 EU981287 51 99 Dothidea insculpta CBS 189.58 DQ247802 Dothidea sambuci CBS 197.58 AY930108 CPC 30638 88 52 Dothidea ribesia Dothidea ribesia CPC 30713

Dothidea ribesia CPC 30689 Dothidea ribesia CBS 195.58 AY016360 Dothidea puccinioides CBS 193.58 AY004342 64 Dothidea muelleri CBS 191.58 EU167593 63 Dothidea berberidis CBS 187.58 KC800752 Dothidea sp. MFLU 14-0040 KM388552 Dothiora bupleuricola CBS 112.75 KU728538 Dothiora agapanthi CPC 20600 KU728537 Dothideaceae Dothiora viburnicola CBS 274.72 KU728554 Dothiora maculans CBS 299.76 KU728543 Dothiora sorbi CBS 742.71 KU728552 Dothiora ceratoniae CBS 477.69 KF251655 Dothiora phillyreae CBS 473.69 EU754146 Dothiora oleae CBS 152.71 KU728547 100 Dothiora cannabinae CBS 737.71 DQ470984 Dothiora laureolae CBS 744.71 KU728542 Dothiora prunorum CBS 933.72 KU728551 Dothiora elliptica CBS 736.71 KU728541 Dothiora phaeosperma CBS 870.71 KU728550 CPC 30632 98 Dothiora pyrenophora Dothiora pyrenophora CPC 30634 Dothiora cactacearum CPC 15585 75 Dothiora cactacearum CPC 15587 Dothiora buxi MFLU 15-3404 KX765295 Pleurophoma acaciae CPC 29188 KY173524 98 Keissleriella cladophila CBS 104.55 GU205221 77 Keissleriella sparticola MFLUCC 14-0196 KP639571 Keissleriella genistae CBS 113798 GU205222 Pleurophoma ossicola CPC 24985 KR476770 93 Lentitheciaceae Pleurophoma pleurospora CBS 116668 JF740326 Keissleriella yonaguniensis KT 2604 AB807594 61 Murilentithecium clematidis IT1078 KM408758 57 Phragmocamarosporium platani MFLUCC 13-0552 KP842916

62 Pleosporales Phragmocamarosporium hederae MFLUCC 14-1191 KP842915 Didymosphaeria rubi-ulmifolii MFLUCC 14-0023 KJ436586 Paraconiothyrium brasiliense CBS 254.88 JX496171 Pseudocamarosporium sp. 2 CPC 25002 100 99 Pseudocamarosporium sp. 2 CPC 25004 Pseudocamarosporium sp. 2 CPC 25843 Pseudocamarosporium sp. 2 CPC 27400 Pseudocamarosporium sp. 2 CPC 30973 Paracamarosporium psoraleae CPC 21632 KF777199 Paracamarosporium fagi CPC 24890 KR611904 Didymosphaeriaceae 53 Paracamarosporium fungicola CBS 113269 JX496133 55 Paracamarosporium hawaiiense CBS 120025 DQ885897 Paracamarosporium sp. 1 CPC 30988 10 changes Paracamarosporium fagi CPC 31037 Pseudocamarosporium sp. 1 CPC 25926 Pseudocamarosporium brabeji CBS 123026 EU552105 Pseudocamarosporium sp. 2 CPC 31482 Pseudocamarosporium piceae MFLUCC 14-0192 KJ803030 Pseudocamarosporium africanum CBS 121166 JX496142

Fig. 1.„*!!!ƒE“V\ƒ *!%™‡$†*!!! those branches also present in the strict consensus tree and families are indicated with coloured blocks and the taxonomic novelties, or species treated in the present paper, are in boldE„Eurotium chevalieri =;Z+$#"$*

136 I MA FUNGUS The Genera of Fungi – G4

Neoplatysporoides aloicola CBS 139901 KR476754 ARTICL Libertasomyces myopori CPC 27354 KX228332 95 Libertasomycetaceae Libertasomyces quercus CBS 134.97 DQ377883 100 Libertasomyces platani CPC 29609 KY173507 Pseudoleptosphaeria etheridgei CBS 125980 JF740291 Coniothyrium palmarum CBS 758.73 EU040225 E 96 Ochrocladosporium elatum CBS 146.33 EU040233 Ochrocladosporium frigidarii CBS 103.81 EU040234 Camarosporium quaternatum CPC 23216 100 Camarosporium quaternatum CPC 31081 100 Camarosporium quaternatum CPC 31518 Coniothyrium telephii CBS 188.71 GQ387599 Camarosporomyces flavigenus CBS 314.80 GU238076 Hazslinszkyomyces lycii CPC 30998 94 Hazslinszkyomyces lycii CPC 31014 Hazslinszkyomyces aptrootii CBS 483.95 DQ377884 Coniothyriaceae Hazslinszkyomyces aloes CPC 21572 KF777198 Foliophoma fallens CBS 161.78 GU238074 Foliophoma fallens CBS 284.70 GU238078

60 Pleosporales Dimorphosporicola tragani CBS 570.85 KU728536 Neocamarosporium chichastianum CBS 137502 KP004483 Pleospora obiones CBS 453.68 DQ678054 Neocamarosporium chersinae CPC 27298 Neocamarosporium calvescens CBS 246.79 EU754131 Neocamarosporium goegapense CBS 138008 KJ869220 80 Neocamarosporium betae CBS 109410 EU754178 Neocamarosporium betae CBS 523.66 EU754179 Neoleptosphaeria rubefaciens CBS 387.80 JF740311

Subplenodomus drobnjacensis CBS 269.92 JF740285 (continued) 51 CBS 101633 GQ387593 Leptosphaeriaceae 87 Querciphoma carteri Querciphoma carteri CBS 105.91 GQ387594 99 acaciae CPC 25527 KX228316 Pyrenochaeta pinicola CBS 137997 KJ869209 Incertae sedis Pyrenochaeta protearum CBS 131315 JQ044453 100 Pleospora herbarum CBS 191.86 GU238160 Stemphylium beticola CBS 141024 KX228349 57 Plenodomus lingam CBS 275.63 JF740306 Plenodomus biglobosus CBS 298.36 GU237980 Incertae sedis 53 Plenodomus enteroleucus CBS 142.84 JF740287 Plenodomus libanotidis CBS 113795 JF740300 Pyrenochaeta nobilis CBS 407.76 EU754206 “Camarosporium” caraganicola MFLUCC 14-0605 KP711381 57 “Camarosporium” sp. 3 CPC 31031 “Camarosporium” aureum MFLUCC 14-0620 KP744478 “Camarosporium” sp. 5 CPC 30379 “Camarosporium” sp. 5 CPC 27667 “Camarosporium” robiniicola MFLUCC 13-0527 KJ589412 “Camarosporium” clematidis MFLUCC 12-0354 KJ562188 “Camarosporium” elongata CBS 171.55 DQ678061 Incertae sedis “Camarosporium” spartii MFLUCC 13-0548 KJ589413 “Camarosporium” aborescentis MFLUCC 14-0604 KP711378 “Camarosporium” arezzoensis CPC 31420 “Camarosporium” sp. 4 CPC 31632 “Camarosporium” sp. 1 CPC 12441 DQ377885 “Camarosporium” arezzoensis MFLUCC 14-0238 KP120927 52 “Camarosporium” sp. 2 CPC 25960 “Camarosporium” sp. 2 CPC 25962 10 changes

Fig. 1. (Continued).

' ' faintly constricted, mostly the middle cells and seldom both eguttulate hyphae, without setae; ascomatal wall consisting *M# of a textura angularis'' a gelatinous sheath and appendages, rehydrated and and eguttulate cells. Pseudoparaphyses numerous, cellular, E!M M` &*!M** M ' *# 3 *_$M*$$M  Conidiomata pycnidial, '# &‡3 saprobic, subcorticolous, single to gregarious, partly Asci "' \ ' base, ostiole centrally, terete, short papillate, and somewhat “ ƒ * #M*_ "'% & *` & *" 3 Ascospores ‡M`' and smooth hyphae, to 0.8 mm diam; conidiomatal wall ' ' textura globulosa-angularis "' ' Paraphyses

V O LUME 8 · NO . 1 137 Crousus and Groenewald wald

Dothiora oleae CBS 615.72 KU728511.1 Leptosphaeria doliolum CBS 504.75 JF740209.1 100 LE Leptosphaeria conoidea CBS 616.75 JF740201.1 Leptosphaeria

C 77 Leptosphaeria sclerotioides CBS 148.84 JF740193.1 Camarosporomyces flavigenus CBS 314.80 Camarosporomyces RTI CPC 23216 A 100 CPC 31518 Camarosporium quaternatum Camarosporium CPC 31081 Subplenodomus drobnjacensis CBS 269.92 JF740211.1 Subplenodomus Subplenodomus apiicola CBS 504.91 JF740197.1 Plenodomus enteroleucus CBS 142.84 NR 111619.1 Plenodomus Paraleptosphaeria dryadis CBS 643.86 JF740213.1 Paraleptosphaeria 100 Paraleptosphaeria praetermissa CBS 114591 JF740241.1 100 CBS 284.70 CBS 161.78 Foliophoma fallens Foliophoma Libertasomyces platani CPC 29609 KY173416.1 99 Libertasomyces quercus CBS 134.97 Libertasomyces / Neoplatysporoides aloicola CBS 139901 KR476719.1 Neoplatysporoides 88 Libertasomyces myopori CPC 27354 KX228281.1 Neocamarosporium goegapense CBS 138008 KJ869163.1 92 Dimorphosporicola tragani CBS 570.85 KU728497.1 Neocamarosporium / 93 Neocamarosporium chersinae CPC 27298 Dimorphosporicola 94 Neocamarosporium chichastianum CBS 137502 KP004455.1 Coniothyrium glycines CBS 124141 KF251211.1 Coniothyrium 100 Coniothyrium hakeae CPC 27616 KY173397.1 CBS 101633 KF251210.1 Querciphoma carteri Querciphoma CBS 105.91 KF251209.1 Hazslinszkyomyces aptrootii CBS 483.95 83 CBS 136437 KF777142.1 Hazslinszkyomyces aloes Hazslinszkyomyces CPC 30998 Hazslinszkyomyces lycii 100 CPC 31014 Coniothyrina agaves CBS 470.69 JX681075.1 Coniothyrina Pyrenochaetopsis microspora NRRL 39060 HM751085.1 Pyrenochaetopsis “Camarosporium” sp. 1 CPC 12441 87 MFLUCC 14-0238 KP120926.1 “Camarosporium” arezzoensis CPC 31420 “Camarosporium” spartii MFLUCC 13-0548 KJ562215.1 100 “Camarosporium” clematidis MFLUCC 13-0336 KJ562213.1 CPC 25960 91 “Camarosporium” sp. 2 CPC 25962 “Camarosporium” sp. 3 CPC 31031 “Camarosporium” “Camarosporium” sp. 4 CPC 31632 “Camarosporium” caraganicola MFLUCC 14-0605 KP711380.1 “Camarosporium” aborescentis MFLUCC 14-0604 KP711377.1 CPC 27667 “Camarosporium” sp. 5 89 CPC 30379 20 changes “Camarosporium” aureum MFLUCC 14-0620 NR 137970.1 “Camarosporium” robiniicola MFLUCC 13-0527 KJ562214.1

Fig. 2. „ "‡ ƒ E <V ƒ Camarosporium!%™_‡†*!!! blocks and the taxonomic novelties, or species treated in the present paper, are in boldEDothiora oleae (GenBank accession number KU728511).

and conidiophores absent. Conidiogenous cells formed curved, apically rounded, base often tapered but never from the inner cells of the pycnidial wall, doliiform, hyaline, truncate, always yellowish, but never brown, both end cells ' but mostly the basal cell often paler or hyaline, wall golden, apically and singly. Conidia as brownish, dark olive to black ' #M‡M`' to slightly constricted, eguttulate at maturity, examined in one longitudinal or diagonal septum per cell and 1–2 per % *"M` M#‡ & $M*M*` conidium, ellipsoidal, pyroid, clavate, straight to slightly 3**$M`M##Synasexual morph in culture:

138 I MA FUNGUS The Genera of Fungi – G4

Pezicula cinnamomea CBS 290.39 KR859133.1 ARTICL 100 CBS 189.58 AF027764.1 Dothidea insculpta MFUCC 13-0686 KM388543.1 99 100 AC836 KF147923.1 Dothidea hippophaeos CBS 186.58 AF027763.1

Dothidea puccinioides CBS 193.58 E Dothidea muelleri CBS 191.58 EU167593.1 MFLU 14-0040 KM388544.1 Dothidea sp. Dothidea 99 MFLUCC 13-0670 KM388545.1 DAOM 231303 DQ491505.1 100 Dothidea sambuci 100 CBS 197.58 AY930108.1 Dothidea berberidis CBS 186.58 EU167601.1 CPC 30638 100 CPC 30689 Dothidea ribesia CPC 30713

98 Dothiora buxi MFLU 15-3404 KX765294.1 CPC 15585 Dothiora cactacearum 100 CPC 15587 Dothiora cannabinae CBS 737.71 AJ244243.1 Dothiora laureolae CBS 744.71 KU728503.1 Dothiora phaeosperma CBS 870.71 KU728512.1 Dothiora europaea CBS 739.71 AJ244244.1 Dothiora elliptica CBS 736.71 KU728502.1 Dothiora prunorum CBS 933.72 AJ244248.1 Dothiora rhamni-alpinae CBS 745.71 AJ244245.1 88 Dothiora sorbi CBS 742.71 KU728514.1 CPC 30632 Dothiora Dothiora pyrenophora CPC 30634 CBS 299.76 KU728504.1 Dothiora maculans CBS 686.70 KU728507.1 Dothiora agapanthi CPC 20600 KU728498.1 Dothiora viburnicola CBS 274.72 KU728515.1 Dothiora bupleuricola CBS 112.75 KU728499.1 Dothiora phillyreae CBS 473.69 KU728513.1 CBS 441.72 KU728501.1 Dothiora ceratoniae CBS 477.69 KF251151.1 CBS 235.57 KU728509.1 Dothiora oleae CBS 615.72 KU728511.1 10 changes

Fig. 3.„*‡!ƒE<VƒDothidea and Dothiora*!%™_‡†*!!! taxonomic novelties, or species treated in the present paper, are in boldE Pezicula cinnamomea (GenBank accession ‰A" $*##

Conidiomata Notes: Single conidial colonies of C. quaternatum formed a ]+z* !M!!3*M ' ostioles, exuding a crystalline conidial mass. Conidiophores step was redone three times on different media, with the reduced to conidiogenous cells. Conidiogenous cells lining same result. It was only once colonies were subcultured the inner cavity, hyaline, smooth, ampulliform, with periclinal V+z % E#M &‡M 3 Conidia solitary, hyaline, banana leaf, that a few typical Camarosporium conidiomata smooth, aseptate, subcylindrical, straight, rarely curved, %\ E#M‡M M`&* 3 morph develops in culture, however, remains unknown, with ' Culture characteristics|„@Uz\ and the Camarosporium morph extremely rare. mycelium and feathery margin; surface dirty white with Ascospores of Camarosporium quaternatumm studied here patches of olivaceous grey. On PDA surface and reverse agree with those of the original description of Cucurbitaria varians olivaceous grey. On OA surface olivaceous grey with patches *#M‡E Clinterium of dirty white and umber. quaternatum $M* ŽŒŒ *"` ; holotype material could not be traced, the original illustrations

V O LUME 8 · NO . 1 139 Crousus and Groenewald wald

Keissleriella trichophoricola CBS 136770 KJ869113.1 Paracamarosporium fungicola CBS 113269 JX496020.1 LE

C Paracamarosporium sp. 1 CPC 30988 98 Paracamarosporium leucadendri CBS 123027 EU552106.1

RTI Paracamarosporium mamanes CBS 120031 DQ885900.1 A 100 Paracamarosporium psoraleae CPC 21632 KF777143.1 Paracamarosporium CBS 120025 DQ885897.1 100 Paracamarosporium hawaiiense CPC 12268 DQ885896.1 77 CPC 31037 CPC 24890 KR611886.1 Paracamarosporium fagi 99 CPC 24892 KR611887.1 Pseudocamarosporium sp. 1 CPC 25926 CBS 121166 JX496029.1 Pseudocamarosporium africanum STE U-6316 EU295650.1 72 CPC 25002 CPC 25004 CPC 25843 Pseudocamarosporium sp. 2 Pseudocamarosporium CPC 27400 CPC 30973 CPC 31482 Pseudocamarosporium brabeji CBS 119219 EU552104.1 Pseudocamarosporium corni MFLUCC 13-0541 KJ747048.1 Pseudocamarosporium cotinae MFLUCC 14-0624 KP744460.1 Pseudocamarosporium lonicerae MFLUCC 13-0532 KJ747047.1 Pseudocamarosporium piceae MFLUCC 14-0192 KJ747046.1 Pseudocamarosporium pinicola MFLUCC 14-0457 KT222273.1 Pseudocamarosporium propinquum MFLUCC 13-0544 KJ747049.1 Pseudocamarosporium quercinum MFLUCC 14-0456 KT222272.1 Pseudocamarosporium tiliicola MFLUCC 13-0550 KJ747050.1 Pseudocamarosporium brabeji CBS 123026 EU552105.1 Pseudocamarosporium brabeji IMI 501236 JX204290.1 Pseudocamarosporium pini MFLUCC 14-1091 KU764779.1 10 changes Pseudocamarosporium sp. MFLUCC 14-0458 KT222274.1

Fig. 4. „ *` ƒ E <V ƒ Paracamarosporium and Pseudocamarosporium*!%™`$†*!!! with coloured blocks and the taxonomic novelties, or species treated in the present paper, are in boldEKeissleriella trichophoricola =;‰Z"`$**#

Table 2. Statistical information on the individual alignments and number of equally most parsimonious trees saved for each dataset analysed1. LSU overview Camarosporium ITS Dothidea ITS Paracamarosporium ITS Aligned characters (including gaps) 722 554 #! # ]'% 192 242 109 45 ª'% 55 88 107 *‡# Constant characters 475 224 #*‡ ## Equally most parsimonious trees obtained 1 000 284 140 16 `_# 1 247 470 249 Consistency index (Cl) 0,551 0,499 0,691 0,912 Retention index (RI) 0,948 0,764 !"# 0,891 Rescaled Consistency index (RC) 0,522 !#"* 0,575 0,812 1<V|% "VX+z{“V\|"VX+z

140 I MA FUNGUS The Genera of Fungi – G4 ARTICL E

Fig. 5. Camarosporium quaternatum/]/#*!"*A. Immersed ascomata on twig (arrows). B–E. Asci and pseudoparaphyses. F. Ascospores. G. Conidiomata on twig. H. Camarosporium conidioma on OA. I.]'V+z J–K. Conidiogenous cells giving rise to Camarosporium conidia. L./;|z<—!!3=—"!!3Ž—#!!3—*!3

V*""# Lycium sporium s. str. % ' ' barbarum in Hungary and Germany. In the present study, 'E we also found the sexual morph on Daphne mezereum, | a new host for C. quaternatum previous studies as representative of this fungus (CBS Camarosporomyces Crous, gen. nov. ‡"#$ /;V*#‡$_/et al. 2006) represent two MycoBank MB820901 ' from C. quaternatum. Etymology| + Sutton (1980) stated that the genus Camarosporium, Camarosporium'E which contains several hundred species, is polyphyletic and in need of revision. As we have shown here, Diagnosis| X ' % Camarosporium s. str. is distinct from other taxa that have pycnidial conidiomata with prominent papillate dark brown been treated as representative of or similar in some respects periphysate ostiole, and thicker wall at apex. to Camarosporium, viz. Henfellra, Neocamarosporium, Paracamarosporium, or Xenocamarosporium (Crous et Type species: Camarosporomyces (Constantinou al !*# !*‡ !* J‚ et al. 2014, 2016, & Aa) Crous 2017. Hawksworth et al. 2016). Further studies are underway to treat those taxa that are presently known from culture. Description: Conidiomata pycnidial, solitary to aggregated, During the course of evaluating the status of Camaro- obovoid, medium brown, with prominent papillate ostiole,

V O LUME 8 · NO . 1 141 Crousus and Groenewald wald LE C RTI A

Fig. 6. /;V#*‡"! A. Conidiomata on PDA. B./V+zC. Conidiomata showing darker ostiolar area. D. Conidiogenous cells. E.]';|zM/—$!3XMU—*!3

dark brown, with extra 1–2 layers of wall at the apex (thicker Foliophoma Crous, gen. nov. than pycnidial body), periphysate. Conidiophores reduced @;@;"!$!# to conidiogenous cells lining inner cavity, hyaline, smooth, ampulliform; phialidic with periclinal thickening. Conidia Etymology|+ solitary, hyaline, smooth, aseptate, subcylindrical, straight, its morphological similarity to Phoma. apex obtuse, base truncate. Diagnosis| X ' % ˆ*&*@${'$@$&<#`{ \*x,=`"J{ (Constant. & Aa) ' ' *M# Crous, comb. nov. ostioles. Conidiogenous cells with periclinal thickening or MycoBank MB820902 percurrent proliferation at apex. (Fig. 6) Basionym: % Constant. & Aa, Trans. Brit. Type species: Foliophoma fallens (Sacc.) Crous 2017. Mycol. Soc. 79|#‡#*$" Synonym: % (Constant. & Aa) Gruyter & Description: Conidiomata ' Verkley, Stud. Mycol. 75| !*#ÆP!*PÇ '*M# { #M` textura angularis. Type: Romania: Bucharest, station for water treatment, from Conidiophores reduced to conidiogenous cells lining inner water, 1980, K. Fodor/;VŽ'#!`_M{/;V#*‡"! cavity, hyaline, smooth, doliiform to subcylindrical; phialidic ME' with periclinal thickening or percurrent proliferation at apex. Conidia aseptate, solitary, hyaline, smooth, guttulate to Description: Conidiomata pycnidial, solitary to aggregated, 'E obovoid, medium brown, 50–90 μm diam, with prominent truncate to bluntly rounded. papillate ostiole, dark brown, 10–25 m diam with extra 1–2 layers of wall at the apex (thicker than pycnidial body), Foliophoma fallens (Sacc.) Crous, comb. nov. periphysate. Conidiophores reduced to conidiogenous cells MycoBank MB820904 %‡M &#M‡ (Fig. 7) μm; phialidic with periclinal thickening. Conidia solitary, Basionym: Phoma fallens Sacc., Syll. Fung. 10: 146 (1892). hyaline, smooth, aseptate, subcylindrical, straight, apex Synonyms: Pleospora fallenss (Sacc.) Gruyter & Verkley, M#&* 3 Stud. Mycol. 75| !*#Æ!*Ç Phyllosticta glaucispora Delacr., Bull. Soc. Mycol. France 9: Culture characteristics| / \ ``*"$# #! Phoma glaucispora X +  ; Versl. diam after 2 wk. On OA surface umber with diffuse yellow Meded. Plantenziektenk. Dienst Wageningen 166 : 108 pigment. On PDA surface umber, outer region pale luteous, (1989) [“1988"]. reverse umber. Phyllosticta oleandrii Gutner, Trudy Bot. Inst. Akad. Nauk S.S.S.R., ser. 2, Sporov. Rast. 1|#!`*$## Note: ' which was originally described as %, and later Description: Conidiomata'' placed in Pleospora by De Gruyter et al.!*# *M#*!M !3 a much wider circumscription of the genus Pleospora than { #M` textura applied here. angularis. Conidiophores reduced to conidiogenous cells lining inner cavity, hyaline, smooth, doliiform to subcylindrical,

142 I MA FUNGUS The Genera of Fungi – G4 ARTICL E

Fig. 7. Foliophoma fallenss (CBS 284.70). A. Conidiomata on PDA. B./V+z C–D. Conidiogenous cells. E. Conidia. Bars: A–B = 250 μm, all others = 10 μm.

M_&‡M 3{ Description: Ascomata ' proliferation at apex. Conidia aseptate, solitary, hyaline, ticolous, densely crowded, erumpent, globose to pyroid with ' \ apex obtuse, base truncate to bluntly rounded, (5–)5.5–6(–7) {' &#M‡M 3/;V"‡_! of a textura angularis. Pseudoparaphyses numerous, ' Culture characteristics| / \ 'Ascii"'' moderate aerial mycelium and even, lobate margins, reaching apically rounded and with an ocular chamber, pedicel 50 mm diam after 2 wk. On PDA surface sepia to umber, Ascospores reverse dark mouse grey. On OA surface greyish sepia. transversely septate, constricted at the median septum, becoming muriformly septate, ellipsoidal, golden brown. Material examined: Italy: Capri, Villa Jovis, leaf spot on Nerium ]'|Conidiomata pycnidial, solitary, brown, oleander (Apocynaceae), Apr. 1970, H.A. van der Aa/;VŽ'*``#$ { M# #!``/;V"‡_!MNew Zealand: Levin, from leaf spot textura angularis. Conidiophores reduced to conidiogenous of Olea europaea (Oleaceae), 1978, G.F. Laundon (CBS 161.78 = cells lining the inner cavity, hyaline, smooth, doliiform to “Uª**#* subcylindrical. Conidia solitary, hyaline, smooth, guttulate, /' Notes: Phoma fallenss is associated with leaf spots of Olea morph: Occurring in the same or separate conidioma europaea U _M$ & #M‡ 3 Phoma ' Conidiogenous cells lining the glaucispora M"& M‡ 3 inner cavity, doliiform to subcylindrical, hyaline, smooth, spots on Nerium oleander in Europe (Boerema et al. 2004). proliferating percurrently at apex. Conidia solitary, brown, Phylogenetically, however, De Gruyter et al!*# smooth, broadly ellipsoidal to obovoid, transversely septate, two species to be closely related, and therefore treated them becoming muriformly septate. as a single species, placing them in Pleospora based on the larger circumscription used at the time. However, the present Hazslinszkyomyces aloes (Crous & M.J. Wingf.) fungus is not congeneric with Stemphylium herbarum (syn. Crous, comb. nov. Pleospora herbarum; Rossman et al. 2015), and therefore a MycoBank MB820906 new generic name is introduced to accommodate it. Basionym: Camarosporium aloes Crous & M.J. Wingf., Persoonia 31|‡_!*# Hazslinszkyomyces Crous & R.K. Schumach., gen. nov. Type: South Africa: Western Cape Province: Clanwilliam, MycoBank MB820905 on dark lesions on dead bark of Aloe dichotoma (Xanthorrhoeaceae), Sept. 2012, +>+ ?!! (CBS Etymology| + H z ŽŒŒ Ž'*‡‡` M { /]/ * _ — /;V *#`‡#_ M von Hazslin (1818–1896), in recognition for his work on E' ' Hazslinszkyomyces aptrootii Crous, sp. nov. Diagnosis: Morphologically similar to Camarosporium, but MycoBank MB820908 distinct as conidia are uniformly brown in colour, those of (Fig. 8) Camarosporium having paler end cells. Etymology| + z z Type species: Hazslinszkyomyces aloes (Crous & M.J. + Wingf.) Crous 2017.

V O LUME 8 · NO . 1 143 Crousus and Groenewald wald LE C RTI A

Fig. 8. Hazslinszkyomyces aptrootii/;V‡"#$ A./]+z B. Conidiomata on PDA. C. Hazslinszkyomyces ' conidia. D–E. Conidiogenous cells. F.]';|zM;—*_!3—*!3

Diagnosis: Similar to Hazslinszkyomyces aloes [conidia (9–) Hazslinszkyomyces lycii Crous & R.K. Schumach., **M*#M*‡&‡M`M_M")Ç% sp. nov. conidia. MycoBank MB820909 (Fig. 9) Type: The Netherlands: Province of Noord-Holland: Egmond, on Lycium sp., 17 Mar. 1995, A. Aptroot /;V Ž'#!`" M Etymology| + {/;V‡"#$ ME' isolated, Lycium.

Description: Conidiomata separate, erumpent, pycnidial, Diagnosis: Similar to Hazslinszkyomyces aptrootiii [conidia *#!M*_!3' **M*#M*‡M*`&_M"M$3Ç% ostiole, 10 μm diam, exuding a creamy conidial mass; wall conidia. #M` textura angularis. Conidia dimorphic. ]'|Conidiophores reduced to conidiogenous Type: Hungary: near Budapest, on twig of Lycium barbarum cells lining the inner cavity, ampulliform to subcylindrical, (Solanaceae), 2 May 2016, L. Bartaloss /;V Ž'#!`$ M hyaline, smooth, phialidic with periclinal thickening at apex, holotype; CPC #!$$"—/;V*‡`*$ME' M_ & #M` 3 Conidia solitary, hyaline, smooth, guttulate, subcylindrical, aseptate, Description: Ascomata ' % E ‡M M` & colous, densely crowded, erumpent, globose to pyroid with 3 /' | „ \ ' Conidiophores *#!3{ reduced to conidiogenous cells, hyaline, smooth, doliiform to ' textura angularis' E M_&‡M` ' Pseudoparaphyses μm. Conidia solitary, broadly ellipsoidal, apex subobtusely ' rounded, base truncate, hilum 2 μm diam, medium brown, ' #%*M# ' Asci"'' ƒ**M*#M*‡M*`&_M"M$3 M#3 *" M*‡& Culture characteristics|/\ *‡ &* 3ƒAscospores to moderate aerial mycelium, covering dish in 2 wk, with initially with three transverse septa, constricted at the median even, lobate margins. On PDA surface umber, reverse brown % % % vinaceous. On OA surface chestnut to brown vinaceous. muriformly septate, 1–2 longitudinal or diagonal septa per cell, ellipsoidal, straight, the upper part often wider, end cells Notes| Camaro- mostly bluntly rounded seldom conical, wall faintly thick and sporium quaternatum, based on its general morphology, ' and the fact that it occurred on Lycium. Morphologically, median septum constricted otherwise faintly constricted, however, the conidia are much smaller than those of C. eguttulate at maturity, no mucilaginous sheath in all stages of quaternatum, and thus it is described as a new species of %*"M!M*M#&" M$M*!M Hazslinszkyomyces. Phylogenetically the two genera are *3! M#! &*!M* 3]'| also quite distinct. Conidiomata pycnidial, solitary, brown, globose, 180–250 3 !M#! 3 { M# layers of brown textura angularis. Conidiophores reduced to conidiogenous cells lining the inner cavity, hyaline,

144 I MA FUNGUS The Genera of Fungi – G4 ARTICL E

Fig. 9. Hazslinszkyomyces lyciii/]/#!$$"A. Ascoma immersed in twig. B–D. Asci and pseudoparaphyses. E.]' V+zF–G. Conidiogenous cells. H. Conidia. I. Hazslinszkyomyces conidioma on twig. J. Conidiogenous cells. K./;|z—*#!3< = 250 μm, all others = 10 μm.

‡M_&‡M 3{ PDA surface fawn to hazel, reverse fawn. On OA surface with periclinal thickening at apex. Conidia solitary, hyaline, ochreous to umber. smooth, guttulate, subcylindrical, straight, aseptate, apex * 3# M‡M M`& Other material examined: Hungary: near Budapest, on twig of 3 /' | Conidiomata pycnidial, Lycium barbarum (Solanaceae), 1 May 2016, L. Bartalos (HPC 1029 solitary, gregarious, brown, globose, with central ostiole, wall —/]/#*!*‡—/;V*‡`*" ' textura globulosa-angularis, completely covered with hyaline to brownish and short hyphae, to Notes: Hazslinszkyomyces lyciii occurred in association 800 μm diam. Conidiogenous cellss lining the inner cavity, ' `M_ & M_ 3{ % X+z ƒ annellidic, proliferating percurrently at apex. Conidia solitary, E ' E pale yellowish with a golden wall, smooth, broadly ellipsoidal morph in culture. Pyrenophora lyciii (syn. Pleospora lycii), to obovoid but also clavate, seldom cylindrical, straight, also described from Lycium barbarum in Hungary, has seldom slightly curved, apex obtuse, base sometimes setose pseudothecia and ascospores that are transversely tapered and never truncate, 4–5 μm diam, transversely '`M#! M#M‡M ' & " 3 z vertical and oblique septa, septa smooth to slightly constricted name to consider is “Hendersonia lycii Hazsl”. Hazslinszky * M*`M*"M! & "M$M*! 3 in (1865) referred to a Hendersonia form on Lycium barbarum, vitro*"M#‡&$M*`3 in vivo. butthe name Hendersonia lyciii was never validly published with any description, and has crept into literature in error, and Culture characteristics: Colonies covering dish in 2 wk, with %E moderate aerial mycelium, and smooth, lobate margins. On described here as new.

V O LUME 8 · NO . 1 145 Crousus and Groenewald wald LE C RTI A

Fig. 10. Neocamarosporium chersinae (CPC 27298). A./V+z B. Conidiogenous cells. C–D./;|z—#!!3 others = 10 μm.

Neocamarosporium chersinae Crous, sp. nov. : Libertasomycetaceae, Pleosporales, Dothi- MycoBank MB820910 deomycetes. (Fig. 10) Diagnosis: Similar to Coniothyriaceae, but distinct in that Etymology|+OP ascomata are immersed in a brown stroma, and conidiomata (Chersina angulata), a tortoise species found in dry areas are stromatic in culture. Vz as a food source by people in rural areas, and is also often Type genus: Libertasomyces Crous & Roets 2016. kept as a pet. Description: Ascomata immersed in a brown stroma, Diagnosis: Similar to Neocamarosporium chichastianum becoming erumpent, breaking through the host surface, Æ **M* M*$M & `M"M$M** )Ç aggregated in clusters, with a central ostiole; wall of 6–10 smaller. layers of brown textura angularis. Pseudoparaphyses ' Type: South Africa: Western Cape Province: Robben Island, septate, anastomosing. Ascii fasciculate, stipitate, hyaline, on dead angulate tortoise shell, 25 May 2015, P.W. Crous smooth, subcylindrical, bitunicate with ocular chamber, /;V Ž'#!_! M { /]/ _$" — /;V *‡`! M containing 8 ascospores. Ascospores ' E' brown, verruculose with obtuse ends, muriformly septate, encased in a mucoid sheath. Conidiomata unilocular, Description: Conidiomata solitary, pycnidial, immersed to stromatic, separate, globose, immersed, brown, opening via !!M#!! 3 E{#M` M# 3{M# textura layers of brown textura angularis. Conidiophores reduced angularis. Conidiophores reduced to conidiogenous cells, to conidiogenous cells. Conidiogenous cells lining the #M`&M‡3{ inner cavity, hyaline, smooth, ampulliform to doliiform, with several times percurrently at apex (conidiogenous cells prominent periclinal thickening at the apex, or with tightly dissolve at maturity). Conidia ' aggregated percurrent proliferations at the apex. Conidia walled, smooth, broadly ellipsoidal to subcylindrical or solitary, golden brown, subcylindrical to ellipsoidal, straight %#'!M#ƒ*!M %!M*'E *M*#M* & M`M_3 obtuse, base truncate, with marginal frill, and longitudinal '' Culture characteristics: Colonies covering the dish in 2 wk, oidal, apex obtuse, base truncate to bluntly rounded, \ „ „z @Uz ]Xz aseptata. olivaceous grey to pale olivaceous grey, reverse olivaceous grey. Genera included: Libertasomyces and Neoplatysporoides.

Notes: Neocamarosporium chersinae is introduced as a Libertasomyces quercus Crous, sp. nov. new species of Neocamarosporium (Crous et al. 2014b). MycoBank MB820912 Neocamarosporium ' (Fig. 11) and can only be safely distinguished from this genus based X+z Etymology|+ Quercus. Libertasomycetaceae Crous, fam. nov. MycoBank MB820911 Diagnosis: Similar to Camarosporium quaternatum (conidia *"M#‡&$M*`3% Etymology|+ Libertasomyces. * M*&`M*!3

146 I MA FUNGUS The Genera of Fungi – G4 ARTICL E

Fig. 11. Libertasomyces quercus/;V*#‡$_A. Conidiomata on OA. B, C. Conidiogenous cells. D. Conidia. Bars: A = 200 μm, all others = 10 μm.

Type: Spain: location unknown, on leaf litter of Quercus ilex Type species: Querciphoma carterii (Gruyter & Boerema) (Fagaceae), 20 Jul. 1996, R.F. Castañeda/;VŽ'#!` M Crous 2017. {/;V*#‡$_—<+

Diagnosis: Morphologically similar to Phoma, but distinct in Material examined: Germany|J'=|% ' ' twigs of Quercus robur (Fagaceae), H. Schilll/;VŽ'#!_* conidia become brown and verruculose with age. CBS 105.91). – The Netherlands: PD Lisse, on Quercuss sp., date ]X"‡_‡—/;V*!*`##

V O LUME 8 · NO . 1 147 Crousus and Groenewald wald LE C RTI A

Fig. 12. Querciphoma carterii (CBS 105.91). A. Conidiomata on PDA. B./V+zC. Setae. D. Conidiogenous cells. E. Conidia. Bars: A, B = 250 μm, all others = 10 μm.

Notes| Phoma sp., and Dothiora Fr., Summa veg. Scand. 2: 418 (1849). Coniothyrium carterii (based on Phoma Synonyms: Dothichiza Lib. ex Roum., Fungi Selecti Gallici carteri; see Boerema et al. 2004) by De Gruyter et al. Exs. cent. 7, no. 627 (1880). !*#Ž% Coleonaema Höhn., Mitt. Bot. Lab. TH Wien 1#|$ *$‡ Coniothyrium s.str%*' Cylindroseptoria Quaedvl., et al., Stud. Mycol. 75|# "!*# best placed in a separate genus. Additional synonyms: See Crous & Groenewald (2016). Didymosphaeriaceae Munk, Dansk bot. Ark. 15: 128 *$ # : Dothideaceae, Dothideales, Dothideomycetes.

Type genus: Didymosphaeria Fuckel 1870. Current generic circumscription: Ascostromata immersed % '{ Paracamarosporium mamanes (Crous) Crous, comb. wall of dark brown textura angularis. Locules globose nov. to subglobose, broadly rounded or papillate with central MycoBank MB820916 ostiole. Pseudoparaphyses absent. Ascii"' Basionym: Camarosporium mamaness Crous, Fungal Planet % + |*!!` small ocular chamber. Ascospores' septate, constricted at the primary median septum, at times Type: USA: Hawaii: Saddle Road, on stems of Sophora with a vertical septum, hyaline, rarely pale brown, obovate chrysophylla (Leguminosae), Aug. 2005, W. Gams & Y. to ellipsoidal to fusoid, often inequilateral or slightly curved, Degawa /;V Ž'*$__ M { /]/ * — /;V smooth, at times with a thin mucoid sheath. Conidiomata *!!#*/]/* #M* ‡ME' pycnidial, separate, or aggregated in a stroma. Conidiophores reduced to conidiogenous cells lining the inner cavity, hyaline, Notes: Camarosporium mamaness is allocated to the genus smooth, ampulliform to doliiform, phialidic. Conidia aseptate, Paracamarosporium based on its phylogenetic position. hyaline, smooth, subcylindrical to ovoid or oblong. Hyphae However, the distinction between Paracamarosporium and becoming brown, verruculose and constricted at septa, giving Pseudocamarosporium is highly debatable (Wijayawardene 'E et al. 2014), and although we could separate them here based <V ƒ Type species: Dothiora pyrenophora (Fr.) Fr. 1849. based on LSU data, and appear to represent a single genus, Paracamarosporium. Furthermore, there have also been an Dothiora cactacearum Crous, sp. nov. excessive number of species introduced in the genus, which MycoBank MB820917 X+z H*#

Etymology| + DOTHIORA COMPLEX fungus was isolated, Cactaceae.

Dothideaceae Chevall., Fl. gén. env. Paris 1: 446 Diagnosis: Morphologically distinct from other species of (1826); as “Dothideae”. Dothiora in that conidia become brown and roughened with age. Type genus: Dothidea Fr. 1818. Type: USA: Texas: Austin, on phyllodes of Cactaceae!*# P.W. Crouss /;V Ž'#!_‡ M { /]/ * " — /;V *‡‡$ME'/]/* "_

148 I MA FUNGUS The Genera of Fungi – G4 ARTICL E

Fig. 13. Dothiora cactacearum (CPC 15585). A. Conidiomata on OA. B. Conidioma. C.;V+z D, E. Conidiogenous cells. F. Conidia. G. Conidia turning brown and verruculose with age. Bars: A = 150 μm, all others = 10 μm.

Description: Conidiomata separate, erumpent, pycnidial, pulvinate, opening by an irregular pore, upper layer dissolving * !M#!! 3 with age; wall of 6–10 layers of brown textura angularis. Asci E { #M` bitunicate, hyaline, oblong to subcylindrical, short stipitate, of brown textura angularis. Conidiophoress reduced to "'M#3$!M*!&*‡M*_ conidiogenous cells lining the inner cavity, hyaline, smooth, μm. Ascosporess' _M* &_M* 3 ' ƒ with percurrent proliferation and prominent collarette. Conidia slightly curved, with prominent mucoid sheath when young (in hyaline, smooth, guttulate, subcylindrical to broadly ellipsoidal, water), dissolving at maturity, (5–)5(–8) transversely septate, EM#3 prominently constricted at primary septum, with oblique or *M*‡M*_M*$& M`M_ M"3%* &_3{ % M M#!M# &_M" 3{ conidia becoming brown and roughened with age. Hyphae ascospores directly giving rise to asexual morph via budding, becoming brown, verruculose and constricted at the septa. with ascomata transforming with age into large conidiomata, with apical opening completely dissolving. Conidiomata Culture characteristics|/\ #!!3 to moderate aerial mycelium, and even, lobate margin, { #M` reaching 40 mm diam after 2 wk at 25 ºC. On MEA surface of brown textura angularis. Conidiophoress lining the inner umber, reverse luteous; on PDA surface fawn, reverse cavity, reduced to conidiogenous cells. Conidiogenous cells isabelline; on OA surface isabelline. ‡M$&‡M`3 minute periclinal thickening at apex. Conidia solitary, aseptate, Dothiora pyrenophora (Fr.) Fr., Summa veg. Scand. hyaline, smooth, ovate to ellipsoidal, with minute guttules, 2: 418 (1849). E M_M"M$&#M‡3 (Fig. 14) Basionym: Dothidea pyrenophora Fr., Kongl. Vetensk. Acad. Culture characteristics|/\ Handl. 40: 88 (1819) : Fr., Syst. mycol. 2| *"# aerial mycelium and feathery, undulate margins. On PDA, @Uz„z%' Type: Sweden: without locality, on dead branches of Sorbus sp. (Rosaceae), E.M. Friess (no original material found). – Other material examined: Germany: on twig of Sorbus aucuparia, 27 Germany: on twig of Sorbus aucuparia, 27 Apr. 2016, R.K. Apr. 2016, R.K. Schumacher /;VŽ'#!_ Schumacherr /;V Ž'#!_# M neotype designated here, @;#_` "{ /]/ #!`#‡ /]/ #!`# — /;V *‡`*M Notes| E % E' occurring on Sorbus ƒ „Dothiora sorbii (CBS Description: Ascostromata solitary to aggregated, black, 742.71) was included in this study, and clustered slightly apart immersed to erumpent, unilocular, to 400 μm diam, elliptical, from D. pyrenophora<V

V O LUME 8 · NO . 1 149 Crousus and Groenewald wald LE C RTI A

Fig. 14. Dothiora pyrenophora /]/#!`# A. Ascostroma on twig. B. Section through ascoma. C–E. Asci. F. Ascospores. G. Ascospores undergoing microcyclic conidiation. H. Conidia undergoing budding. I. Conidia. Bars: A, B = 200 μm, all others = 10 μm.

that these may very well be two closely related, but distinct present study. We are also thankful to René K. Schumacher for species. making many valuable collections available to be included in this Genera such as Sydowia Bres.1895 and Pringsheimia study, and for bringing the problem related to the nomenclature of Schulzer 1866 appear to represent younger generic Camarosporium to our attention. names within Dothideaceae, based on their morphological similarity, and the formation of Dothichiza' like morphs in culture, which are commonly observed in REFERENCES the family (Froidevaux 1972, Sivanesan 1984, Crous & Groenewald 2016). Another confusing aspect of many of the ;=ŽX=Z+@UŽ@U/!!‡ taxa in this family is that young ascospores can be hyaline %!‚"""!"\ and uniseptate, but become brown and muriformly septate . Wallingford: CABI Publishing. with age (e.g. Dothidea ribesia, Fig. 15), which complicate Carbone I, Kohn LM (1999) A method for designing primer sets for et al. Mycologia 91: !*‡ #M ` Species of Dothiora are commonly isolated from dead / A / ]J Ž ‰X = Z[' branches of woody hosts (Sivanesan 1984), while Crous & C (2008) Species of Mycosphaerella and related anamorphs on Groenewald (2016) also reported some species from dead Eucalyptus% Persoonia 21: 77–91. leaves and fruit of diverse hosts, suggesting that it is a saprobe, Crous PW, Gams W, Stalpers JA, Robert V, Stegehuis G (2004) possibly acting as a weak pathogen on stressed plant tissues. MycoBank: an online initiative to launch mycology into the 21st century. Studies in Mycologyy 50: 19–22. Crous PW, Giraldo A, Hawksworth DL, Robert V, Kirk PM, et al. ACKNOWLEDGEMENTS !*‡ = H| E species of generic names. IMA Fungus 5: 141–160. We acknowledge Laszlo Bartalos and René Jarling for sending us /]J=Z[!*` Fungal specimens of Camarosporium and similar taxa examined in the Biology 120|*#$M*‡*

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