Camarosporium and Dothiora
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IMA FUNGUS · 8(1): 131–152 (2017)) doi:10.5598/imafungus.2017.08.01.10 The Genera of Fungi – G 4: Camarosporium and Dothiora ARTICL Pedro W. Crous*#, and Johannes Z. Groenewald1 E 1J H ;% < \ " # " / \ +{ '| # westerdijkinstitute.nl 2@X;\\%]"# "/\+ #Department of Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, P. Bag X20, Pretoria 0028, South Africa Abstract: =H Key words: X+z X+z; microfungi, Camarosporium and Dothiora%'[ Camarosporium fungal systematics [ C. quaternatum ' E ' E <V Furthermore, Camarosporomyces, Foliophoma and Hazslinszkyomycess ' LSU like genera, while Querciphoma' Libertasomycetaceae is introduced [ as a new family to accommodate Libertasomycess and Neoplatysporoides. Dothiora [ www.GeneraofFungi.org D. pyrenophora ' ' E D. cactacearum<[E'% JH;%</;V//'X+z =;z%' ]=+H Article info:V|* z!*_{z|*$@!*_{]|#@!*_ INTRODUCTION Dothiora (based on D. pyrenophora) produces a Dothichiza asexual morph in culture, but the Dothideaceae [= % ' of Fungi project (www.GeneraOfFungi.org; Crous et al. 'E 2014a), which has the aim to revise the generic names of Although we address the status of Dothiora in the present fungi accepted in Kirk et al!*# study, many additional collections are still required to resolve ' the generic boundaries of other old generic names in this which have been registered in MycoBank (Robert et al. % % *' !*# H ascospores to become muriformly septate with age, and name initiative for fungi (Hawksworth et al. 2011, Crous et al. eventually to even become pigmented. Given that these 2015a), a single name is indicated for each genus. characters have traditionally been used to separate genera in V%*$" et al. 2014), it now problematic genera, namely Camarosporium, and Dothiora. appears likely that there are far fewer genera in Dothideaceae In recent years, it has become clear that the Camarosporium than previously assumed (Crous & Groenewald 2016). morphology has evolved several times within Dothideomycetes, ' % ' E ' E z MATERIALS AND METHODS additional complication lies in that some taxa only produce a single morph (e.g. see below in the present study), and Isolates [ Freshly collected twigs were placed in damp chambers, without molecular data. Furthermore, the treatment of Phoma and incubated at room temperature (ca. 20 oC) for 1–2 d. by Boerema et al. (2004) placed genetically distinct genera Single conidial or ascospore colonies were established into one genus, while the same was true for the treatment of from sporulating conidiomata or ascomata in Petri dishes Pleospora by De Gruyter et al!*# E@Uz the reference strains of Camarosporium included in Crous et (Crous et al. 1991). After 1–2 d, single spores were picked al. (2006) were authentic for the name. @Uz/' © 2017 International Mycological Association You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: [ Non-commercial: No derivative works: For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. V O L U ME 8 · NO . 1 131 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*$_!E LE ]+zVet al. 1996), and incubated %[@; C at 25 o/ '% (www.MycoBank.org; Crous et al. 2004). RTI sporulation. Reference strains and specimens are maintained A at the Westerdijk Fungal Biodiversity Institute (CBS Culture /\+ RESULTS ,{$;*+,$"*&';,[#*+,$"*"_*"*;<{`{ Phylogeny = X+z E H { [ @Uz J& = 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$= =%U the present study, whereas the remaining three alignments *$$"<V J et al. 1990) and LR5 (Vilgalys & Hester <V Camarosporium and 1990) were used to amplify for all isolates part of the nuclear allied genera (Fig. 2), Dothidea and Dothiora H # X+z <V #R *"V A+z Paracamarosporium and Pseudocamarosporium (Fig. [ <V* "V % A+z<V<VE $!! R"VA+z %%V\H* Dothiora <VJ et al. 1990) and LR0R (Vilgalys & Hester 1990) pyrenophora i s n estled inside the Dothiora clade. Based on this were used as internal sequence primers to ensure good LSU phylogeny, the Dothidea clade has a bootstrap support quality sequences over the entire length of the amplicon. Part %$$Dothiora clade did not receive any 'tub2[ 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 *'tef1[ 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"#$ % +V*+VJ [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 V@]%*#!!X+zVzA@J< HazslinszkyomycesPleospora, %+\ obtain consensus sequences of each isolate. Blast searches and P. fallens, were not congeneric with Pleospora <V V\ (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 [ @ “Coniothyrium” carteri. Several species treated until now <VV\ 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<V =;* $_ ;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 %<V 132 I MA FUNGUS The Genera of Fungi – G4 Hazslinszkyomyces (Fig. 2) and a new generic name is mosing, hyaline, smooth. Ascii"' ARTICL ;<VV\ ' [ [ Ascospores `' X+z]/A *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 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: