Species diversity in Pseudocercospora

Fungal Systematics and Evolution is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License © 2024 Westerdijk Fungal Biodiversity Institute 29 Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Fungal Systematics and Evolution doi.org/10.3114/fuse.2024.13.03 Species diversity in Pseudocercospora J.Z. Groenewald1#, Y.Y. Chen2#*, Y. Zhang3, J. Roux4, H.-D. Shin5, R.G. Shivas6, B.A. Summerell7, U. Braun8, A.C. Alfenas9, A.H. Ujat10, C. Nakashima10, P.W. Crous1,4,11* 1Westerdijk Fungal Biodiversity Institute, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands 2College of Forestry, Henan Agricultural University, Zhengzhou, Henan, China 3Institute of Microbiology, School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China 4Department of Biochemistry, Genetics and Microbiology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, South Africa 5Division of Environmental Science and Ecological Engineering, Korea University, Seoul 02841, Korea 6Centre for Crop Health, University of Southern Queensland, Toowoomba 4350, Queensland, Australia 7Botanic Gardens of Sydney, Mrs Macquaries Rd, Sydney, NSW 2000, Australia 8Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, 06099 Halle (Saale), Germany 9Departmento de Fitopatologia/Instituto de Biotecnologia Aplicada à Agropecuária (BIOAGRO), Universidade Federal de Viçosa, Viçosa, MG, 36570900, MG, Brazil 10Graduate school of Bioresources, Mie University, Kurima-machiya 1577, Tsu, Mie 514-8507, Japan 11Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands *Corresponding authors: [email protected]; [email protected] #These authors contributed equally. Citation: Groenewald JZ, Chen YY, Zhang Y, Roux J, Shin H-D, Shivas RG, Summerell BA, Braun U, Alfenas AC, Ujat AH, Nakashima C, Crous PW (2024). Species diversity in Pseudocercospora. Fungal Systematics and Evolution 13: 29–89. doi: 10.3114/fuse.2024.13.03 Received: 6 February 2024 ; Accepted: 4 March 2024; Effectively published online: 11 April 2024 Corresponding editor: A.J.L. Phillips VOLUME 13 JUNE 2024 PAGES 29–89 Key words: Multi-gene phylogeny Mycosphaerellaceae new taxa plant pathogen taxonomy Abstract: Species of Pseudocercospora are commonly associated with leaf and fruit spots on diverse plant hosts in sub-tropical and tropical regions. Pseudocercospora spp. have mycosphaerella-like sexual morphs, but represent a distinct genus in Mycosphaerellaceae (Mycosphaerellales, Dothideomycetes). The present study adds a further 29 novel species of Pseudocercospora from 413 host species representing 297 host genera occurring in 60 countries and designates four epitypes and one lectotype for established names. This study recognises 329 species names, with an additional 69 phylogenetic lineages remaining unnamed due to difficulty in being able to unambiguously apply existing names to those lineages. To help elucidate the taxonomy of these species, a phylogenetic tree was generated from multi-locus DNA sequence data of the internal transcribed spacers and intervening 5.8S nuclear nrRNA gene (ITS), partial actin (actA), and partial translation elongation factor 1-alpha (tef1), as well as the partial DNA-directed RNA polymerase II second largest subunit (rpb2) gene sequences. Novel species described in this study include those from various countries as follows: Australia, Ps. acaciicola from leaf spots on Acacia sp., Ps. anopter from leaf spots on Anopterus glandulosus, Ps. asplenii from leaf spots on Asplenium dimorphum, Ps. australiensis from leaf spots on Eucalyptus gunnii, Ps. badjensis from leaf spots on Eucalyptus badjensis, Ps. erythrophloeicola from leaf spots on Erythrophleum chlorostachys, Ps. grevilleae from leaf spots on Grevillea sp., Ps. lophostemonigena from leaf spots on Lophostemon confertus, Ps. lophostemonis from leaf spots on Lophostemon lactifluus, Ps. paramacadamiae from leaf spots on Macadamia integrifolia, Ps. persooniae from leaf spots on Persoonia sp., Ps. pultenaeae from leaf spots on Pultenaea daphnoides, Ps. tristaniopsidis from leaf spots on Tristaniopsis collina, Ps. victoriae from leaf spots on Eucalyptus globoidea. Brazil, Ps. musigena from leaf spots on Musa sp. China, Ps. lonicerae-japonicae from leaf spots on Lonicera japonica, Ps. rubigena leaf spots on Rubus sp. France (Réunion), Ps. wingfieldii from leaf spots on Acacia heterophylla. Malaysia, Ps. musarum from leaf spots on Musa sp. Netherlands, Ps. rhododendri from leaf spots on Rhododendron sp. South Africa, Ps. balanitis from leaf spots on Balanites sp., Ps. dovyalidicola from leaf spots on Dovyalis zeyheri, Ps. encephalarticola from leaf spots on Encephalartos sp. South Korea, Ps. grewiana from leaf spots on Grewia biloba, Ps. parakaki from leaf spots on Diospyros kaki, Ps. pseudocydoniae from leaf spots on Chaenomeles lagenaria, Ps. paracydoniae from leaf spots on Chaenomeles speciosa. Thailand, Ps. acerigena from leaf spots on Acer sp., Ps. tectonigena from leaf spots on Tectona grandis. Epitypes are designated for Cercospora bonjeaneae-rectae, Cercospora halleriae, Ps. eucleae, and an epitype as well as a lectotype for Ps. macadamiae. Results obtained in the present study contribute to a better understanding of the host specificity and distribution in Pseudocercospora spp., many of which represent important pathogens of food or fibre crops, or organisms of quarantine concern. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 30 INTRODUCTION In his monograph, Chupp (1954) placed all cercosporoid species in the genus Cercospora that he treated in a very wide sense (Mycosphaerellaceae, Mycosphaerellales, Dothideomycetes). Subsequent morphological studies have shown however that several cercosporoid genera can be distinguished based on a combination of characteristics such as conidial pigmentation, and the structure of conidiogenous loci (scars) and hila (Deighton 1976, 1979, 1983, 1987, 1990, Pons & Sutton 1988, Braun 1995, Crous & Braun 2003). The separation of these genera was further corroborated in molecular studies (Crous et al. 2013a, Nakashima et al. 2016, Chen et al. 2022), which led to them being widely accepted among mycologists and plant pathologists (Braun et al. 2013a, 2014, 2015a, b, 2016). With the end of dual nomenclature, several names were again reduced to synonymy, such as Mycosphaerella under Ramularia (Videira et al. 2015, 2016), while genera with mycosphaerella-like sexual morphs, such as Passalora, Pseudocercospora and Zasmidium, were recognised as distinct (Videira et al. 2017). With several hundred species associated with leaf spot diseases on a wide global host range, Pseudocercospora (Ps.) is a well-known genus of cercosporoid fungi that contains numerous important plant pathogenic species. Well-known pathogens include Ps. angolensis causing fruit and leaf spot disease on Citrus (Pretorius et al. 2003), Ps. pini-densiflorae causing brown spot needle blight of Pinus (Quaedvlieg et al. 2012, Braun et al. 2013a), Ps. griseola causing angular leaf spot of Phaseolus (Crous et al. 2006), Ps. ulei causing South American leaf blight of Hevea spp. (Hora Júnior et al. 2014), and Ps. fijiensis, causing Black Sigatoka disease on Musa spp. (Churchill 2011, Chang et al. 2016, Crous et al. 2021). To place these taxa into a broader evolutionary context, Crous et al. (2013a) published a phylogenetic analysis of 146 Pseudocercospora spp. based on multi-locus sequence data of the nuclear ribosomal RNA (nrRNA) gene (LSU; 28S), the internal transcribed spacers and intervening 5.8S nuclear nrRNA gene (ITS), partial actin (actA), and translation elongation factor 1-alpha (tef1) gene regions. To further improve the backbone resolution of the genus, Nakashima et al. (2016) added data of the partial DNA-directed RNA polymerase II second largest subunit (rpb2) gene, which proved effective in delineating species and genera in the Mycosphaerellaceae (Videira et al. 2015). Based on these results most species of Pseudocercospora were shown to be highly host or genus specific (Bakhshi et al. 2014, Shivas et al. 2015, Guatimosim et al. 2016, Silva et al. 2016), while the genus Pseudocercospora was resolved as monophyletic (Nakashima et al. 2016). Furthermore, these studies also concluded that Pseudocercospora spp. on the same host species or family that are morphologically similar, frequently represented distinct species on different continents, and that European or American names could not readily be applied to taxa from other continents. These previous studies were however hampered by the non-availability of sequence data from (ex-)type material or at least authentic material, a situation that has not greatly improved in recent years. Given the huge species diversity within Pseudocercospora, the present study further expands the generic phylogeny generated by Crous et al. (2013a) and Nakashima et al. (2016), and treats 329 species of Pseudocercospora, introducing 29 novel species of Pseudocercospora from 413 host species representing 297 host genera occurring in 60 countries, and designates four epitypes and one lectotype. A further 69 phylogenetic lineages remain unnamed due to difficulty in being able to unambiguously apply existing names to those lineages, and an existing name is tentatively applied to 14 lineages pending future neoor epitypification. MATERIALS AND METHODS Isolates Isolations were made from conidiophores with conidia on leaves, twigs, and fruits (Suppl. Table S1). Some samples were incubated in moist chambers for 2–3 d to enhance sporulation before single conidial colonies were established on 2 % malt extract agar (MEA) (Crous et al. 2019b). Leaf spots bearing ascomata were soaked in water for approximately 2 h, after which they were attached to the inner surface of Petri dish lids over plates containing MEA for 24–48 h (Crous et al. 1991). Colonies were sub-cultured onto synthetic nutrient-poor agar (SNA), potato-dextrose agar (PDA), oatmeal agar (OA), and MEA (Crous et al. 2019b), 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 and fungarium (CBS and CBS H) of the Westerdijk Fungal Biodiversity Institute (WI), Utrecht, the Netherlands; the culture collection (MUCC) and the fungarium (TSU-MUMH) of the Phytopathology Lab., Mie University, Tsu, Mie, Japan; the research center of genetic resources (MAFF), National agriculture and food research organization, Tsukuba, Japan; and the fungarium (TFM), Forestry and Forest products research institute, Tsukuba, Japan. DNA extraction, PCR amplification and sequencing Fungal mycelium (Suppl. Table S1) was scraped from the surface of actively growing agar cultures with a sterile scalpel and the genomic DNA was extracted using either a DNeasy ® UltraClean® Microbial DNA isolation kit (Qiagen, Hilden, Germany) or the Wizard® Genomic DNA Purification Kit (Promega Corporation, WI, USA), following the manufacturer’s instructions. Four loci were amplified and sequenced as described in Nakashima et al. (2016), namely the internal transcribed spacers and intervening 5.8S nuclear nrRNA (ITS), the partial actin (actA), translation elongation factor 1-alpha (tef1), and DNA-directed RNA polymerase II second largest subunit (rpb2) gene regions. The resulting amplicons were sequenced in both directions using the respective PCR primers and the BigDyeTM Terminator Cycle Sequencing Kit v. 3.1 (Applied Biosystems, Life Technologies, Carlsbad, CA, USA). DNA sequencing amplicons were subsequently purified through Sephadex G-50 Superfine columns (Sigma-Aldrich, St. Louis, MO) in MultiScreen HV plates (Millipore, Billerica, MA) and 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 Prime v. 2022.0.2 (http://www. geneious.com, Kearse et al. 2012). Phylogenetic analysis The sequences for each gene region were subjected to megablast searches in the NCBI’s GenBank nucleotide database © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 31 (Zhang et al. 2000), and also supplemented with sequences of ex-type strains of more distant published species. Sequence alignments were generated per locus using the online version of MAFFT v. 7 (https://mafft.cbrc.jp/alignment/server/index.html; Katoh et al. 2019) with default settings. Leading and trailing gaps were removed as far as possible without removing too much potential phylogenetic signal, after which the alignments were concatenated using SequenceMatrix v. 1.9 (Vaidya et al. 2011). An initial guide tree was constructed with IQ-TREE v. 2.1.3 (Nguyen et al. 2015), after which the sequences in the alignment were sorted according to the tree topology using Mesquite v. 3.70 (Maddison & Maddison 2023), and the local alignment of adjacent sequences improved by eye where necessary using Geneious Prime v. 2023.2.1. Maximum-likelihood (ML) phylogenetic trees were constructed using IQ-TREE v. 2.1.3 and branch support values were calculated with 1 000 nonparametric bootstrap replicates and optimal modelfinding using the TESTNEW option of ModelFinder (Kalyaanamoorthy et al. 2017) as implemented in IQ-TREE. RAxML v. 8.0.0.0 (Stamatakis 2014) was used with default parameters and 1 000 non-parametric bootstrap replicates to provide additional ML support values for the multigene phylogeny. All resulting trees were printed with Geneious Prime v. 2023.2.1 and layout of the tree was done with Adobe Illustrator 2024 v. 28.0. Sequences derived in this study were deposited in GenBank (Suppl. Table S1), and the alignment and phylogenetic trees in figshare.com (doi: 10.6084/m9.figshare.23447345). Morphology Slide preparations were mounted in clear lactic acid or Shear’s mounting fluid. Descriptions were chiefly based on fungarium specimens. In cases where this was not possible, descriptions were based on colonies sporulating on MEA, PDA, SNA 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. 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). Taxonomic novelties were deposited in MycoBank (www.MycoBank.org; Crous et al. 2004a). RESULTS Phylogeny The concatenated alignment consists of 723 strains, including the outgroup (Trochophora simplex) and the basal relative (Parapallidocercospora colombiensis). The alignment contained a total of 2 489 characters with the following partitions: 1–520 (ITS), 521–760 (actA), 761–1 813 (tef1) and 1 814–2 489 (rpb2). The phylogeny resulting from the IQ-TREE maximum likelihood analysis is presented in Fig. 1, with the bootstrap support values from both the IQ-TREE and RAxML analyses plotted on the branches. Statistics for the IQ-TREE analysis are 1 842 distinct patterns, 1 242 parsimony-informative, 277 singleton, and 970 constant sites; additional statistical measures for the different analyses are provided in Suppl. Table S2. Blocks (loosely applied to correspond to phylogenetic clades or groups of clades for most instances) were numbered to facilitate referencing to the position of a species in the phylogenetic tree in the Taxonomy section below. The majority of terminal (species) clades are virtually the same between the two analyses. In many instances where a single strain moved to a different position between the analyses, this was due to the fact that the given strain has only an ITS sequence available. Forty-five strains representing 43 species are represented by ITS sequences only in this dataset (Suppl. Table S3). In Suppl. Table S3, the two strains each for Ps. microlepiae and “Ps. platylobii (block 7)” were collapsed to a single row as each set of two strains functioned as a unit in the phylogenetic trees. Two other strains represent species for which additional strains are available with complete datasets (Ps. platanigena and Ps. punctata); these two strains also clusters with the other strains in both the IQ-TREE and RAxML analyses. Of the 45 strains, 18 did not have conflicting positions in the phylogenetic trees obtained from the IQ-TREE (Fig. 1) and RAxML (see Figshare) analyses. The remaining 27 strains did cluster differently between the phylogenetic trees obtained from the IQ-TREE and RAxML analyses and these differences are highlighted in Suppl. Table S3 and in the species notes below. For example, Ps. camelliicola (BCRC FU30031) is located towards the bottom of block 11 in Fig. 1 as close relative of Ps. trichogena, but in the RAxML phylogeny (see Figshare) it is located on a long branch between the two strains of Ps. lonicerae-japonicae, which is located in block 9 of Fig. 1 based on the IQ-TREE phylogeny. Strains represented by only an ITS sequence in the dataset are shown in Fig. 1 in dark blue font if there are no conflicts in their position between the IQ-TREE and RAxML analyses, and in an orange font if there are conflicts in their position. Only the bootstrap support values from the RAxML analysis are plotted on Fig. 1; the obtained RAxML phylogenetic tree is deposited in Figshare. Overall, the obtained RAxML bootstrap support values were much lower than the obtained IQ-TREE bootstrap support values. This resulted in many branches not having a RAxML bootstrap support value as the value was below the threshold value chosen for plotting even though the clustering of the isolates was the same between the two analyses. TAXONOMY Pseudocercospora Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires, Ser. 3, 20: 437. 1910. Synonyms: See Crous & Braun (2003), Braun et al. (2013a), Crous et al. (2013a), Videira et al. (2017). Foliicolous, chiefly phytopathogenic, but also endophytic; commonly associated with leaf spots, but also occurring on fruits and twigs. Mycelium internal and external, consisting of smooth, septate, subhyaline to brown branched hyphae. Stroma absent to well-developed. Conidiophores in vivo arranged in loose to dense fascicles, sometimes forming distinct synnemata or sporodochia, emerging through stomata or erumpent through the cuticle, often arising from substomatal or subcuticular to intraepidermal stromata, or occurring singly on superficial hyphae, short to long, septate or continuous, i.e., conidiophores may be reduced to conidiogenous cells, simple to branched and straight to geniculate-sinuous, subhyaline, pale to dark olivaceous to brown, smooth to finely verruculose. Conidiogenous cells integrated, © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 32 Fig. 1. Consensus phylogram (50 % majority rule) obtained from the maximum likelihood analysis with IQ-TREE v. 2.1.3 of the concatenated nucleotide alignment. Bootstrap support values (> 74 %) from 1 000 non-parametric bootstrap replicates are shown at the nodes, followed by RAxML 1 000 replicate bootstrap support values (> 74 %). Culture collection or voucher numbers are followed by the country and host information where available. Sequences derived from material with a type status are indicated with a superscript HT (from (ex-)type), IT (from (ex-)isotype), PT (from (ex-)paratype), NT (from (ex-)neotype, AUT (from authentic), REF (from reference) and ET (from (ex-)epitype). Strains in dark blue font represent strains with only an ITS sequence in the dataset but with no conflict in their positions between the IQ-TREE and RAxML analyses, whereas an orange font indicates such a strain with a conflict in its position between the two analyses. Numbered coloured blocks are provided to facilitate referencing to the position of a species in the phylogenetic tree. Taxa for which it was not possible to assign the correct name due to lack of type material are indicated together with their block number between parentheses in the phylogenetic tree, Suppl. Tables S1 and S3 and the Taxonomy section. Novel species described in this study are highlighted with bold font. The tree was rooted to Trochophora simplex with Parapallidocercospora colombiensis as internal distant genus. The scale bar indicates the expected number of changes per site. 0.1 100/100 2x 100/- 100/86 100/100 100 /96 100/- 100/100 81/- 83/- 96/- 82/- 90/- 98/- 100/- 100/- 100/89 100/100 100/100 100/99 100/- 97/91 96/- 100/97 100/- 100/- 100/- 92/- 85/- 86/- 100/97 100/99 100/100 99/97 100/100 100/100 100/98 100/100 99/97 98100 /100 87/100 100/100 100/- 84/83 100/- Trochophora simplex CBS 124744 Parapallidocercospora colombiensis CBS 110968HT Ps. angolensis CBS 112933ET Zimbabwe: Citrus sp. Ps. angolensis CBS 149.53HT Angola: Citrus sinensis Ps. chamaecristae COAD 1973ET Brazil: Chamaecrista sp. Ps. exilis COAD 1501ET Brazil: Chamaecrista orbiculata Ps. smilacicola VSC-F0001015 USA: Smilax rotundifolia Ps. pistacina CBS 135841 Türkiye: Pistacia vera Ps. pistacina CPC 32979 Türkiye: Pistacia vera Ps. pistacina CPC 32978 Türkiye: Pistacia vera Ps. pistacina CPC 32985 Türkiye: Pistacia vera Ps. pistacina CPC 32974 Türkiye: Pistacia vera Ps. pistacina CPC 32986 Türkiye: Pistacia vera Ps. pistacina CPC 32981 Türkiye: Pistacia vera Ps. pistacina CPC 32984 Türkiye: Pistacia vera Ps. pistacina CPC 32977 Türkiye: Pistacia vera Ps. pistacina CPC 32983 Türkiye: Pistacia vera Ps. pistacina CPC 32980 Türkiye: Pistacia vera Ps. pistacina CBS 135840 Türkiye: Pistacia vera Ps. pistacina CPC 32976 Türkiye: Pistacia vera Ps. sophoricola CBS 136020HT Iran: Sophora alopecuroides Ps. ocimi-basilici CBS 114646 Fiji: Unknown Ps. ocimi-basilici CPC 10283HT Mexico: Ocimum basilicum Ps. heteropyxidicola CBS 146082HT South Africa: Heteropyxis natalensis Ps. heteropyxidicola CPC 31133 Unknown: Litter of Heteropyxis sp. Ps. heteropyxidicola CPC 31359 Unknown: Heteropyxis sp. Ps. udagawana CBS 131931 South Korea: Hovenia dulcis Pseudocercospora sp. 001 CPC 21998 Thailand: Moringa oleifera Ps. zambiensis CBS 136423HT Zambia: Terminalia sp. Ps. punctata CBS 132116 Madagascar: Syzygium sp. Ps. punctata CPC 14737 Madagascar: Syzygium sp. Ps. punctata CPC 19814 South Africa: Syzygium cordatum Ps. punctata CPC 14743 Madagascar: Syzygium sp. Ps. punctata CPC 14740 Madagascar: Syzygium sp. Ps. punctata CPC 39344ET South Africa: Syzygium cordatum Ps. ulei GCL012 Colombia: Hevea brasiliensis Ps. fici Cerf63-10 India: Ficus religiosa Ps. nodosa CBS 554.71HT Romania: Psoralea bituminosa Ps. vitis CBS 132012 South Korea: Vitis vinifera Ps. vitis CPC 14668 South Korea: Vitis vinifera Ps. clematidis CPC 11657 USA: Clematis sp. Ps. pseudostigmina-platani CPC 14775 South Korea: Platanus occidentalis Ps. pseudostigmina-platani CBS 131588HT South Korea: Platanus occidentalis Ps. pseudostigmina-platani CPC 14816 South Korea: Platanus occidentalis Ps. platanigena CBS 125773 Iran: Platanus orientalis Ps. platanigena CBS 125774 Iran: Platanus orientalis Ps. platanigena DAR 84735 Australia: Platanus orientalis cv. Digitata Ps. platanigena CBS 336.33 Unknown: Platanus orientalis Ps. platanigena CBS 110755 India: Unknown Ps. cladrastidis MUCC 1494ET Japan: Maackia amurensis Ps. sambucigena CBS 126000ET Netherlands: Sambucus nigra Ps. sambucigena CPC 14798 South Korea: Sambucus williamsii Ps. sambucigena CPC 21573 UK: Fraxinus sp. Ps. griseola f. mesoamericana CBS 119113HT South Africa: Phaseolus vulgaris Ps. griseola f. mesoamericana CPC 16119 Mexico: Legiminosae sp. Ps. griseola f. griseola CBS 119906ET Tanzania: Phaseolus vulgaris Ps. griseola f. griseola CBS 111825 South Africa: Unknown Ps. griseola f. griseola CBS 194.47 Portugal: Phaseolus vulgaris 1 2 3 100/- 100/99 93/- 96 92 90 97 100/97 92 100/99 100 85/- 97/- 92/- 100/100 100/- 75/- 98/- 85/- 75/- 78/99 75/- Ps. macrospora CBS 114696 Brazil: Bertholletia excelsa Ps. tecomicola COAD 1585 Brazil: Tecoma stans Ps. angularis COAD 2073HT Brazil: Prunus persica Ps. manihotis COAD 1534HT Brazil: Manihot sp. Ps. wulffiae COAD 1976HT Brazil: Wulffia stenoglossa Ps. musigena sp. nov. CBS 149357HT Brazil: Musa sp. Ps. norchiensis CBS 118413 New Zealand: Rubus sp. Ps. norchiensis CBS 114641 New Zealand: Rubus sp. Ps. norchiensis CBS 120738HT Italy: Eucalyptus sp. Ps. nogalesii CBS 115022 New Zealand: Chamaecytisus proliferus Ps. nogalesii CBS 122501 New Zealand: Virgilia divaricata Pseudocercospora sp. 005 CBS 111374 Uruguay: Eucalyptus globulus Pseudocercospora sp. 005 CPC 15102 Brazil: Eucalyptus sp. Ps. brackenicola COAD 1991HT Brazil: Pteridium arachnoideum Pseudocercospora sp. 006 CPC 15830 Mexico: Prunus sp. Ps. xylopiae COAD 1469HT Brazil: Xylopia aromatica Ps. purpurea CBS 114163 Mexico: Persea americana Ps. purpurea CBS 111427 Mexico: Persea americana Ps. purpurea CPC 19701 South Africa: Persea americana Ps. styracina COAD 2369HT Brazil: Styrax sp. Ps. bixae COAD 1563ET Brazil: Bixa orellana Pseudocercospora sp. 007 CPC 31710 Colombia: Eucalyptus pellita Pseudocercospora sp. 008 CBS 115495 Unknown: Unknown Ps. sordida MUCC 913 Japan: Campsis radicans Ps. luzardii COAD 1505ET Brazil: Hancornia speciosa Ps. luzardii CPC 2556 Brazil: Hancornia speciosa Ps. convoluta CBS 113377HT Costa Rica: Chromolaena odorata 4 © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 33 Fig 1. (Continued). 97/- 100/95 83/- 84/- 84/- 98/- 100/100 98/- 100/96 Pseudocercospora sp. 003 CBS 113099 Panama: Smilacina paniculata Ps. viburnigena CBS 125998ET Netherlands: Viburnum davidii Ps. leandrae-fragilis COAD 1977HT Brazil: Leandra fragilis Pseudocercospora sp. 002 CPC 17800 Australia: Eucalyptus sp. Ps. sphaerulinae CBS 112621HT Chile: Eucalyptus nitens Ps. plunkettii COAD 1548 Brazil: Mikania sp. Ps. basitruncata CBS 114664ET Colombia: Eucalyptus grandis Ps. paranaensis COAD 1180 Brazil: Cyathea atrovirens Ps. paranaensis COAD 1987HT Brazil: Cyathea atrovirens Ps. richardsoniicola COAD 1568ET Brazil: Richardia brasiliensis Ps. borreriae 1091023-1 Taiwan: Spermacoce latifolia Pseudocercospora sp. 004 CPC 18640 Thailand: Eupatorium coelestinum 5 0.1 100/- 100/- 95/- 95/- 100/- 100/- 100/- 100/80 89/- 89/75 98/81 89/- 85/- 84/- 91/- 100/76 100/92 100/100 86/- 95/- 90/- 100/75 100/92 95/- 100/98 100/- 95/- 91/- 100/99 100/99 95/- 100/- 100/75 98/- 100/83 100/- 99/- 99/- 100/- 100/- 100/- 100/- 84/- 99/94 80/- 76/- 77/- 85/- 86/- Ps. cryptomeriicola MAFF 240073ET Japan: Cryptomeria japonica Ps. leucadendri CPC 1869 South Africa: Leucadendron sp. Ps. encephalarticola sp. nov. CBS 149351HT South Africa: Encephalartos sp. Pseudocercospora sp. 067 CPC 18337 South Africa: Freylinia lanceolata Pseudocercospora sp. 067 CPC 32677 South Africa: Diospyros whyteana Ps. myrticola CPC 20421 USA: Myrtus communis Ps. myrticola CPC 20422 USA: Myrtus communis Ps. myrticola MUCC 632 Japan: Myrtus communis Ps. proteae CBS 131587HT South Africa: Protea mundii Ps. symploci CBS 142471ET Taiwan: Symplocos paniculata Ps. sarcocephali CBS 118759 Taiwan: Uncaria sp. Ps. theae CBS 128.30 Italy: Camellia sinensis Ps. elaeocarpicola MAFF 237189HT Japan: Elaeocarpus japonicus Ps. lyoniae MAFF 237775ET Japan: Lyonia ovalifolia var. elliptica Pseudocercospora sp. 009 CPC 32989 USA: Rhododendron sp. Ps. natalensis CBS 111069HT South Africa: Eucalyptus nitens Ps. fori CBS 113285HT South Africa: Eucalyptus grandis Ps. fori CMW 9096 South Africa: Eucalyptus grandis Ps. cymbidiicola CBS 115132ET New Zealand: Cymbidium sp. Ps. rhododendri sp. nov. CBS 149360HT Netherlands: Rhododendron sp. Ps. curcumicola MUCC 733HT Japan: Curcuma longa Ps. ixorae CBS 118760 Taiwan: Ixora sp. Ps. parakaki sp. nov. CBS 132019 South Korea: Diospyros lotus Ps. parakaki sp. nov. CBS 149388HT South Korea: Diospyros kaki Ps. cladosporioides CBS 113866 Spain: Olea europaea Ps. cladosporioides CBS 114079 Spain: Olea europaea Ps. cladosporioides CBS 159.48 Papua New Guinea: Liliaceae sp. Ps. cladosporioides CBS 113867 Spain: Olea europaea Ps. cladosporioides CBS 117482ET Tunisia: Olea europaea Ps. robusta CBS 111175HT Malaysia: Eucalyptus robusta Ps. australiensis sp. nov. CPC 32446 Australia: Eucalyptus bicostata Ps. australiensis sp. nov. CBS 149422HT Australia: Eucalyptus gunnii Ps. australiensis sp. nov. CPC 29068 Australia: Eucalyptus caesia “Ps. platylobii (block 7)” VPRI 21698AUT Australia: Platylobium formosum “Ps. platylobii (block 7)” VPRI 22656 Australia: Platylobium formosum Ps. gracilis CBS 111189 Indonesia: Eucalyptus urophylla Ps. gracilis CBS 149390 Malaysia: Leaf litter of Eucalyptus sp. Ps. gracilis CPC 20852 Malaysia: Leaf litter of Eucalyptus sp. Ps. gracilis CBS 243.94HT Indonesia: Eucalyptus urophylla Ps. gracilis CBS 111372 Indonesia: Eucalyptus urophylla Ps. gracilis CPC 19804 Indonesia: Eucalyptus sp. Ps. eucalyptorum CPC 29991 Australia: Eucalyptus sp. Ps. eucalyptorum CPC 32444 Australia: Eucalyptus globulus Ps. eucalyptorum CPC 31628 Ethiopia: Eucalyptus sp. Ps. eucalyptorum CPC 31545 USA: Eucalyptus sp. Ps. eucalyptorum CPC 31630 Ethiopia: Eucalyptus sp. Ps. eucalyptorum CPC 23271 Ethiopia: Eucalyptus sp. Ps. eucalyptorum CPC 23263 Unknown: Unknown Ps. eucalyptorum CPC 23267 Ethiopia: Eucalyptus globulus Ps. eucalyptorum CPC 23269 Ethiopia: Eucalyptus globulus Ps. eucalyptorum CBS 110777HT South Africa: Eucalyptus nitens Ps. eucalyptorum CBS 114866HT South Africa: Eucalyptus nitens Ps. eucalyptorum CBS 132308 Unknown: Unknown Ps. eucalyptorum CPC 31100 UK: Eucalyptus sp. Ps. eucalyptorum CPC 31602 Portugal: Eucalyptus sp. Ps. eucalyptorum CBS 112509 Australia: Eucalyptus sp. Ps. eucalyptorum CPC 31089 Germany: Tilia platyphyllos Ps. eucalyptorum CPC 26541 France (Réunion): Eucalyptus robusta Ps. eucalyptorum CBS 116358 Australia: Eucalyptus sp. Ps. eucalyptorum CPC 25613 Brazil: Litter of Eucalyptus sp. 6 7 100/86 100/87 100/96 100/93 100/79 Pseudocercospora sp. 010 CBS 115116 New Zealand: Rhopalostylis sapida Ps. arecacearum CBS 118406HT New Zealand: Rhopalostylis sapida Ps. arecacearum CPC 32790 New Zealand: Rhopalostylis sp. Ps. coprosmae CPC 32788 New Zealand: Coprosma robusta Ps. coprosmae CBS 114639ET New Zealand: Coprosma robusta Ps. coprosmae CPC 32769 New Zealand: Coprosma robusta 8 © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 34 Fig 1. (Continued). 0.1 89/- 100/- 100/- 99/- 92/- 96/- 75/- 81/- 94/- 96/- 90/- 80/- 100/- 83/- 85/75 100/95 93/- 92/- 100/- 92/- 100/99 90/- 100/- 94/82 98/91 79/77 86/- 93/- 80/- 93/- 88/- 100/100 94/- 99/- 97/- 100/100 93/93 99/- 98/- 99/- 96/- 100/- 100/99 95/- 95/90 Ps. kadsurae MUCC 752ET Japan: Kadsura japonica Ps. yakushimensis MAFF 237025ET Japan: Hydrangea kawagoeana var. kawagoeana Ps. paederiae MAFF 239161 Japan: Paederia foetida Ps. rubigena sp. nov. CBS 149424HT China: Rubus sp. Ps. hibbertiae-asperae VPRI 22652AUT Australia: Hibbertia aspera Ps. iwakiensis MUCC 1736HT Japan: Ilex crenata Ps. rhododendri-indici CBS 131591HT South Korea: Rhododendron indicum Ps. lonicerae-japonicae sp. nov. CBS 149389HT China: Lonicera japonica Ps. lonicerae-japonicae sp. nov. CPC 14801 South Korea: Lonicera japonica Ps. palleobrunnea CBS 124771HT Australia: Syzygium sp. Pseudocercospora sp. 011 CPC 22525 Australia: Ceratopetalum apetalum Ps. pultenaeae sp. nov. CBS 149419HT Australia: Pultenaea daphnoides Ps. ackamae CBS 122626 New Zealand: Ackama rosifolia Ps. aristoteliae CBS 114638 New Zealand: Aristotelia serrata Ps. crousii CBS 119487 New Zealand: Eucalyptus sp. Ps. metrosideri CBS 114294ET New Zealand: Metrosideros excelsa Ps. metrosideri CPC 32785 New Zealand: Metrosideros excelsa Ps. badjensis sp. nov. CBS 149427HT Australia: Eucalyptus badjensis Ps. badjensis sp. nov. CPC 28316 Australia: Eucalyptus sp. Ps. subulata CBS 118489 New Zealand: Eucalyptus botryoides Ps. victoriae sp. nov. CBS 149429HT Australia: Eucalyptus globoidea Ps. victoriae sp. nov. CBS 149428 Australia: Eucalyptus radiata Ps. victoriae sp. nov. CPC 16385 Australia: Eucalyptus dives Ps. victoriae sp. nov. CPC 28347 Australia: Eucalyptus sp. Ps. melicyti CBS 115023HT New Zealand: Melicytus macrophyllus Pseudocercospora sp. 012 CBS 122627 Unknown: Unknown Pseudocercospora sp. 012 CPC 28692 New Zealand: Musa sp. Ps. persooniae sp. nov. CBS 149421 Australia: Unidentified tree host Ps. persooniae sp. nov. CBS 149430HT Australia: Persoonia sp. Ps. tristaniopsidis sp. nov. CBS 149418HT Australia: Tristaniopsis collina Ps. tristaniopsidis sp. nov. CPC 16373 Australia: Eucalyptus approximans Pseudocercospora sp. 014 CPC 17375 Australia: Eucalyptus sp. Ps. christellae MFLU 10-0405HT Thailand: Christella parasitica Ps. lophostemonigena sp. nov. CBS 149352HT Australia: Lophostemon confertus Ps. anopteri sp. nov. CBS 149359HT Australia: Anopterus glandulosus Pseudocercospora sp. 015 CPC 29921 Australia: Cissus antarctica “Ps. platylobii (block 9)” CPC 28317 Australia: Platylobium formosum “Ps. platylobii (block 9)” CPC 20164 Australia: Grevillea ilicifolia “Ps. platylobii (block 9)” CPC 28321 Australia: Platylobium formosum Ps. paramacadamiae sp. nov. CBS 129066 Australia: Macadamia integrifolia Ps. paramacadamiae sp. nov. CBS 129067HT Australia: Macadamia integrifolia Ps. libertiae CBS 114643AUT New Zealand: Libertia ixioides Pseudocercospora sp. 013 CPC 28296 Australia: Platylobium obtusangulum Ps. macadamiae CBS 129069 Australia: Macadamia integrifolia Ps. macadamiae CPC 28320 Australia: Hibbertia aspera Ps. macadamiae CPC 28319 Australia: Hibbertia aspera Ps. macadamiae CPC 28322 Australia: Hibbertia aspera Ps. macadamiae CBS 129068 Australia: Macadamia integrifolia Ps. macadamiae CBS 129071 Australia: Macadamia integrifolia Ps. macadamiae CBS 129070 Australia: Macadamia integrifolia Ps. macadamiae CBS 133432ET Australia: Macadamia integrifolia Ps. correae CPC 28336 Australia: Correa reflexa var. nummulariifolia Ps. correae CPC 28323 Australia: Correa reflexa Ps. correae CPC 28337 Australia: Correa pulchella Ps. correae CPC 28335 Australia: Correa alba Ps. correae CPC 28324 Australia: Correa pulchella Ps. correae CPC 28325 Australia: Correa cv. Mannii 9 96/- 100/- 99/- 100/83 93/- 99/- 95/- 100/92 77/- 78/- 100/85 Ps. hachijokibushi MAFF 238479 HT Japan: Stachyurus praecox var. matsuzakii Ps. boehmeriigena COAD 1562 ET Brazil: Boehmeria nivea Ps. boehmeriigena MAFF 238121 ET of Ps. fukuii Japan: Boehmeria nivea var. concolor f. nipononivea Ps. lygodiicola COAD 1745 Brazil: Lygodium volubile Ps. macarangae P564 Laos: Macaranga denticulata Ps. tectariae BCRC FU30377 AUT Taiwan: Tectaria harlandii Ps. farfugii MUCC 978 HT Japan: Farfugium japonicum Ps. humulicola CBS 131585 HT South Korea: Humulus scandens Ps. rhabdothamni CBS 114872 IT New Zealand: Rhabdothamnus solandri Ps. crocea CBS 126004 HT South Korea: Pilea hamaoi Ps. fatouae CPC 11648 South Korea: Fatoua villosa Ps. profusa CBS 132306 South Korea: Acalypha australis Ps. profusa CPC 10713 ET South Korea: Acalypha australis Ps. dianellae CBS 117746 New Zealand: Dianella caerulea Ps. pteridophytophila R. Kirschner 3602 Taiwan: Cyclosorus parasiticus Ps. pteridicola R. Kirschner 4166 Taiwan: Pteris semipinnata Ps. balsaminae CBS 131882 South Korea: Impatiens textorii Ps. balsaminae CBS 132020 South Korea: Impatiens textorii Ps. plectranthi CBS 131586 HT South Korea: Plectranthus sp. Pseudocercospora sp. 016 CPC 14813 New Zealand: Rubus x barkeri Ps. cyatheicola CBS 129520 HT Australia: Cyathea australis Ps. rumohrae CBS 117747 New Zealand: Marattia salicina 10 © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 35 Fig 1. (Continued). 76/- 100/- 88/- 100/100 89/- 100/89 89/- 95/- 94/- 87/- 100/79 100/95 100/99 83/- 100/100 84/- 76 82/- 81/- 95/- 100/100 Ps. platyceriicola MUCC 2876 HT Malaysia: Platycerium sp. Ps. actinidiae HJWB1 China: Actinidia chinensis Ps. actinidiae HJWB2 China: Actinidia chinensis Ps. actinidiae HJWB3 China: Actinidia chinensis Ps. pouzolziae CBS 122280 Taiwan: Gonostegia hirta Ps. pouzolziae NCHUPP L1608 Taiwan: Pouzolzia sp. Ps. nephrolepidis CBS 119121 AUT Taiwan: Nephrolepis auriculata Ps. microlepiae BCRC FU30353 HT Taiwan: Microlepia speluncae Ps. microlepiae BCRC FU30378 Taiwan: Microlepia trichocarpa Ps. lythri CBS 132115 ET South Korea: Lythrum salicaria Ps. perrottetiae CBS 147382 HT Taiwan: Perrottetia arisanensis Ps. perrottetiae CBS 147383 HT Taiwan: Perrottetia arisanensis Ps. tectonigena sp. nov. CBS 149361 HT Thailand: Tectona grandis Ps. tectonigena sp. nov. CPC 21861 Unknown: Curcuma sp. Ps. biophyti CPC 20020 REF Benin: Biophytum petersianum Ps. houttuyniae CBS 149366 South Korea: Lycopus lucidus Ps. houttuyniae MAFF 238071 ET Japan: Houttuynia cordata Pseudocercospora sp. 017 CBS 118761 Taiwan: Quercus sp. Ps. jussiaeae CBS 132117 South Korea: Ludwigia prostrata Ps. jussiaeae CPC 14822 South Korea: Ludwigia prostrata Ps. microsori CBS 139476 HT Australia: Microsorum pustulatum Ps. dendrobii MUCC 596 Japan: Dendrobium sp. Ps. araliae MAFF 238192 ET Japan: Aralia elata Pseudocercospora sp. 018 CBS 122098 Taiwan: Smilacaceae sp. Ps. ebulicola CBS 147387 ET Taiwan: Sambucus sp. Ps. angiopteridis CBS 147385 ET Thailand: Angiopteris evecta Pseudocercospora sp. 019 CPC 22742 Zambia: Harungana madagascariensis Ps. camelliicola BCRC FU30031 Taiwan: Camellia japonica Ps. trichogena COAD 1087 HT Brazil: Macrothelypteris torresiana Ps. trichogena COAD 1088 Brazil: Deparia petersenii 11 0.1 9998/- 100/- 95/- 82/- 95/- 86/- 100/91 100/- 100/- 100/- 100/90 97/- 92/- 81/- 100/90 100/86 86/- 93/- 100/84 100/100 97/- 100/- 100/91 99/86 87/- 80/- 98/- 100/98 98/75 92/- 100/- 97/- 99/- 100/91 94/- 100/89 100/84 “Ps. cornicola (block 12)” KACC 48076 South Korea: Cornus officinalis Ps. cyatheae BCRC FU30106 Taiwan: Sphaeropteris lepifera Ps. athyrii BCRC FU30236 Taiwan: Deparia longipes Ps. pruni-grayanae MUCC 1715 ET Japan: Padus grayana Ps. halleriae CBS 149362 ET South Africa: Halleria lucida Ps. xanthocercidis CBS 131593 HT South Africa: Xanthocercis zambesiaca Ps. madagascariensis CBS 124155 HT Madagascar: Eucalyptus camaldulensis Pseudocercospora sp. 044 CPC 30395 Brazil: Vigna sp. “Ps. cruenta (block 13)” CBS 117232 Taiwan: Phaseolus vulgaris Ps. tabei YMM220 HT Benin: Vigna unguiculata Ps. wuchienshiungiae BRIP 72387b HT Australia: Persoonia falcata Pseudocercospora sp. 050 CPC 22026 Thailand: Unknown Ps. balanitis sp. nov. CBS 149363 HT South Africa: Balanites sp. Ps. bonjeaniae-rectae CBS 149393 ET Greece: Dorycnium rectum Ps. dovyalidis CBS 126002 ET South Africa: Dovyalis zeyheri Ps. breonadiae CBS 143489 HT South Africa: Breonadia microcephala Ps. kiggelariae CBS 132016 NT South Africa: Kiggelaria africana Ps. kiggelariae CPC 19873 South Africa: Kiggelaria africana Ps. kiggelariae CPC 21128 South Africa: Kiggelaria africana Ps. kiggelariae CPC 18286 South Africa: Protea sp. Ps. kiggelariae CPC 18306 South Africa: Kiggelaria africana Ps. neriicola CBS 138010 HT Italy: Nerium oleander Ps. parapseudarthriae CBS 137996 HT South Africa: Pseudarthria hookeri Ps. stahlii CPC 17316 Laos: Passiflora foetida Ps. stahlii CBS 117549 Taiwan: Passiflora foetida Ps. stahlii CPC 19399 Australia: Unknown Ps. stahlii CPC 27586 Malaysia: Passiflora foetida Ps. crispans CBS 125999 HT South Africa: Eucalyptus sp. Pseudocercospora sp. 043 CBS 110995 Unknown: Populus sp. Pseudocercospora sp. 043 CBS 110996 Unknown: Populus sp. Ps. eucleae CBS 149431 ET South Africa: Euclea sp. Ps. destructiva MUCC 870 Japan: Euonymus japonicus Ps. chiangmaiensis CBS 123244 HT Thailand: Eucalyptus camaldulensis Ps. chiangmaiensis CPC 15450 HT Thailand: Eucalyptus camaldulensis Ps. lindericola BJFC ZYP141005.43 China: Lindera reflexa Ps. lindericola BJFC ZYP141005.40 China: Lindera reflexa Ps. lindericola BJFC ZYP141005.41 China: Lindera reflexa Ps. ampelopsidis CBS 131583 HT South Korea: Ampelopsis glandulosa var. heterophylla Pseudocercospora sp. 045 CPC 22043 Thailand: Acer sp. Ps. carbonacea CPC 19213 Thailand: Dioscorea bulbifera Pseudocercospora sp. 046 CBS 117233 Taiwan: Liana sp. Ps. assamensis CBS 122467 HT India: Musa cv. Nanderan Ps. indonesiana CBS 122473 HT Indonesia: Musa cv. Buai Pseudocercospora sp. 047 CPC 22001 Thailand: Fragaria ananassa Ps. wusaulaniae BRIP 72389f HT Australia: Senna alata Ps. jagerae BRIP 58549 HT Australia: Jagera pseudorhus var. pseudorhus Ps. erythrophleicola sp. nov. CBS 149391 HT Australia: Erythrophleum chlorostachys Ps. cotini MAFF 410088 HT Japan: Cotinus coggygria Ps. securinegae CBS 131930 South Korea: Flueggea suffruticosa Ps. zanthoxyli CPC 10065 South Korea: Zanthoxylum ailanthoides Ps. morigena NFCCI 4877 HT India: Morus alba Ps. mori CBS 122101 Taiwan: Morus sp. Ps. mori CPC 14695 South Korea: Morus alba Ps. snelliana CPC 14781 South Korea: Broussonetia papyrifera Ps. snelliana CBS 131592 South Korea: Morus bombycis Ps. snelliana CPC 14674 South Korea: Morus bombycis 12 13 14 15 16 17 18 19 © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 36 Fig 1. (Continued). 0.1 100/- 93/- 100/- 100/99 82/- 97/- 96/87 100/100 96/- 83/- 83/94 82/- 98/- 96/- 98/- 95/- 100/- 91/- 100/- 100/- 100/86 100/76 84/- 91/- 83/- 96/- 89/- 100/95 78/- 100/- 82/- 82/- 96/- 77/- Ps. cydoniae CBS 131923 South Korea: Chaenomeles speciosa Pseudocercospora sp. 068 CBS 113096 Panama: Acanthaceae sp. Ps. photiniae MUCC 1661NT Japan: Photinia glabra Ps. mazandaranensis CBS 136115HT Iran: Nerium oleander Ps. mazandaranensis CCTU 1146 Iran: Nerium oleander Ps. naitoi MAFF 237906ET Japan: Ilex serrata f. argutidens “Ps. cornicola (block 20a)” MAFF 237773 Japan: Cornus alba var. sibirica Pseudocercospora sp. 049 CPC 24466 China: Citrus grandis Ps. euonymi-japonici CGMCC 3.18576HT China: Euonymus japonicus Ps. eupatoriella CBS 113372HT Jamaica: Chromolaena odorata Ps. rizhaoensis CFCC 57581HT China: Ligustrum japonicum Ps. rizhaoensis CFCC 57582 China: Ligustrum japonicum Ps. paracydoniae sp. nov. CBS 149367HT South Korea: Chaenomeles speciosa Ps. ligustri JS1201 China: Ligustrum japonicum Ps. ligustri JS1202 China: Ligustrum japonicum Ps. ligustri JS1203 China: Ligustrum japonicum Ps. lilacis CBS 132031 USA: Ligustrum japonicum Pseudocercospora sp. 048 CPC 21996 Thailand: Nymphaea lotus Ps. eupatorii-formosani NCHUPP L1606ET Taiwan: Eupatorium sp. Pseudocercospora sp. 069 CBS 149423 China: Buxus megistophylla Ps. eupatorii-formosani CPC 25040 South Africa: Solanum mauritianum Ps. pruni-persicicola KACC 47019 South Korea: Prunus avium Ps. eupatorii-formosani CBS 112825 Unknown: Unknown Ps. eupatorii-formosani CBS 115494 Unknown: Unknown Ps. ginkgoana R. Kirschner 3563AUT for Ps. ginkgoana Taiwan: Ginkgo biloba Ps. eupatorii-formosani CPC 17319 Laos: Chromolaena odorata Ps. eupatorii-formosani CBS 115493 USA: Unknown Ps. eupatorii-formosani CPC 19195 Laos: Chromolaena odorata Ps. eupatorii-formosani CPC 20054 Taiwan: Unknown Ps. duabangae CPC 19201 Laos: Duabanga grandiflora Pseudocercospora sp. 021 CPC 19218 Thailand: Radermachera ignea Ps. cf. destructiva CGMCC 3.18591 China: Euonymus japonicus Pseudocercospora sp. 020 CPC 24507 China: Citrus reticulata cv. Ponkan Ps. kenyirana MUCC 2873HT Malaysia: Trigoniastrum sp. Ps. ixorana MUCC 2608ET Malaysia: Ixora chinensis Ps. ranjita CBS 126005 Indonesia: Gmelina sp. Ps. ranjita CPC 15985 Mexico: Gmelina arborea Ps. robertsiorum BRIP 72515bHT Australia: Senna tora Ps. xenosyzygiicola MAFF 237986ET Japan: Syzygium samarangense Pseudocercospora sp. 023 CPC 17270 Australia: Unidentified red berry tree Pseudocercospora sp. 022 CBS 149432 Mexico: Ornamental plant Ps. avicenniae CBS 146479HT Australia: Avicennia marina Ps. avicenniae CPC 17295 Australia: Avicennia marina Ps. avicenniae CPC 17304 Australia: Avicennia marina Ps. hardenbergiae CBS 147381HT Australia: Hardenbergia violacea Pseudocercospora sp. 042 CPC 24763 Malaysia: Unidentified plant “Ps. puerariicola (block 22)” CPC 19193 Laos: Pueraria phaseoloides Ps. stizolobii COAD 1532 Brazil: Mucuna aterrima Ps. lophostemonis sp. nov. CBS 149350HT Australia: Lophostemon lactifluus Pseudocercospora sp. 025 CPC 14634 Australia: Corymbia dunlopiana Ps. vernoniae NFCCI 4441HT India: Vernonia cineria Ps. holarrhenae P540 Laos: Holarrhena curtisii Ps. planaltinensis COAD 1495HT Brazil: Chamaecrista sp. Ps. tereticornis CPC 13008 Australia: Eucalyptus tereticornis Ps. tereticornis CBS 125214HT Australia: Eucalyptus tereticornis Ps. tereticornis CBS 124996 Australia: Eucalyptus nitens Ps. tereticornis CPC 13315HT Australia: Eucalyptus tereticornis Ps. flavomarginata CBS 126001 China: Eucalyptus sp. Ps. flavomarginata CBS 118824HT Thailand: Eucalyptus camaldulensis Ps. flavomarginata CBS 118823HT Thailand: Eucalyptus camaldulensis Ps. flavomarginata CBS 124990 Thailand: Eucalyptus camaldulensis Ps. flavomarginata CBS 112745 Vietnam: Unknown Ps. flavomarginata CBS 112747 Thailand: Unknown 20b 20c 21 22 20a 80/- 77/- 92/- 100/100 99/- 89/- 100/100 77/- 94/- 87/- 100/100 Ps. sennicola YMM12HT Benin: Senna occidentalis Pseudocercospora sp. 024 CPC 17369 Australia: Eucalyptus sp. Ps. grewiana sp. nov. CBS 149368HT South Korea: Grewia parviflora Ps. dodonaeae CBS 114647ET New Zealand: Dodonaea viscosa Ps. dovyalidicola sp. nov. CBS 149364HT South Africa: Dovyalis zeyheri Ps. dovyalidicola sp. nov. CBS 149365 South Africa: Dovyalis zeyheri Ps. grevilleae sp. nov. CBS 149356HT Australia: Grevillea sp. Ps. clerodendri-hastati NBRC 105401HT Indonesia: Clerodendrum hastatum Ps. raybouldiae BRIP 75777aHT Australia: Croton phebalioides Ps. raybouldiae BRIP 75778a Australia: Croton phebalioides Ps. airliensis BRIP 58550HT Australia: Polyalthia nitidissima Ps. dypsidis Lu17 Thailand: Dypsis lutescens Ps. hakeae CBS 112226 Australia: Grevillea sp. Ps. hakeae CBS 144520 Australia: Hakea sp. Ps. hakeae CBS 149426 Australia: Acacia sp. Ps. hakeae CBS 112225 Australia: Grevillea sp. Ps. solanicola India: Solanum nigrum Ps. hakeae CBS 149425 Australia: Hakea sp. Ps. hakeae CBS 114184 Australia: Grevillea sp. Ps. hakeae CBS 114185 Australia: Grevillea sp. 23 24 © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 37 Fig 1. (Continued). 0.1 83/- 100/95 91/- 86/- 100/- 100/89 93/- 95/- 100/- 97/- 97/89 99/- 100/95 100/100 100100100/- 100/97 98/- 91/- 94/- 95/- 97/- 97/- 97/- 100/- 99/- 88/- 100/99 100/93 98/- Ps. acaciicola sp. nov. CBS 149354HT Australia: Acacia sp. Ps. acaciicola sp. nov. CBS 149355 Australia: Acacia sp. Ps. ceratoniae CBS 147386ET Italy: Ceratonia siliqua Ps. piperis COAD 1111 Brazil: Piper aduncum Ps. subtorulosa CBS 117230 Taiwan: Melicope sp. Ps. bruceae MUCC 2875 Malaysia: Brucea javanica Ps. diospyriphila KACC 47650 South Korea: Diospyros kaki Ps. casuarinae CBS 128218HT Australia: Casuarina cunninghamiana Ps. glochidii MAFF 237000ET Japan: Glochidion zeylanicum Ps. proiphydis CBS 146481HT Australia: Proiphys infundibularis Ps. delonicicola MUCC 2869HT Malaysia: Delonix sp. Pseudocercospora sp. 053 CPC 22037 Thailand: Solanaceae Pseudocercospora sp. 054 CPC 17365 Laos: Tabernaemontana divaricata “Ps. puerariicola (block 25)” KUS F33308 South Korea: Pueraria montana var. lobata “Ps. puerariicola (block 25)” KUS F33294 South Korea: Pueraria montana var. lobata “Ps. puerariicola (block 25)” KUS F33343 South Korea: Pueraria montana var. lobata Ps. acerigena sp. nov. CBS 149358HT Thailand: Acer sp. Ps. sphendamnophila CBS 122279HT Taiwan: Acer albopurpurascens Pseudocercospora sp. 052 CBS 122094 Taiwan: Mangifera indica Ps. sawadae MAFF 239714 Japan: Psidium guajava Ps. mali MUCC 886 Japan: Malus sieboldii Ps. fukuokaensis MAFF 237768ET Japan: Styrax japonicus Ps. imazekii MUCC 1668ET Japan: Kolkwitzia amabilis Ps. fuliginosa MAFF 237710 Japan: Diospyros kaki Ps. cercidis-chinensis CBS 132109ET South Korea: Cercis chinensis Ps. sennae-multijugae COAD 1519HT Brazil: Senna multijuga Ps. trinidadensis COAD 1756ET Brazil: Croton urucurana Ps. solani-pseudocapsicicola COAD 1974HT Brazil: Solanum pseudocapsicum Ps. nymphaeacea KACC 47784 South Korea: Nymphaea tetragona Ps. tinea NCHUPP L1603ET Taiwan: Viburnum sp. Ps. subsynnematosa CMM 2892HT Brazil: Tibouchina herbacea Ps. glauca CBS 131884 South Korea: Albizia julibrissin Ps. chrysanthemicola CBS 131888 South Korea: Chrysanthemum sp. Ps. wingfieldii sp. nov. CBS 149369HT France (Réunion): Acacia heterophylla Ps. aleuritis CBS 281.62 USA: Aleurites fordii Ps. aleuritis MAFF 237174NT Japan: Aleurites montanus Ps. marginalis CBS 131582HT South Korea: Fraxinus rhynchophylla Ps. chionanthi-retusi KACC 47790 South Korea: Chionanthus retusus Ps. chionanthi-retusi NCHUPP L1605ET Taiwan: Chionanthus retusus 25 26a 97/- 82/- 98/- 100/- 100/- 92/- 98/- 92/- 92/- 96/- 97/79 100/100 85/- 86/- 100/- 96/- 78/- 93/- 90/- 85/- 100/96 94/- 100/84 100/83 100/80 95/- 88/- 78/- 99/- 81/- 90/- 94/- 100/100 Ps. vignae MFLUCC 21-0117 HT Thailand: Vigna sp. Pseudocercospora sp. 061 CPC 20722 Thailand: Centrosema pubescens Ps. serpocaulonicola COAD 1866 HT Brazil: Serpocaulon triseriale Pseudocercospora sp. 066 CBS 110994 South Africa: Eucalyptus grandis Pseudocercospora sp. 066 CBS 110997 South Africa: Eucalyptus grandis Ps. pothomorphes COAD 1450 HT Brazil: Pothomorphe umbellata Ps. cordiana CBS 114685 HT Brazil: Cordia goeldiana Ps. fraxinites NCHUPP L1607 Taiwan: Fraxinus formosana Pseudocercospora sp. 062 CPC 19237 Brazil: Cariniana legalis Ps. paraguayensis CBS 111286 Brazil: Eucalyptus nitens Ps. paraguayensis CBS 111375 Brazil: Eucalyptus nitens Ps. nandinae MAFF 237633 ET Japan: Nandina domestica Pseudocercospora sp. 063 CPC 24511 China: Citrus grandis Ps. pini-densiflorae MUCC 534 ET Japan: Pinus strobus Ps. pini-densiflorae CBS 125139 Japan: Pinus thunbergii Ps. pini-densiflorae CBS 125140 Philippines: Pinus kesiya Ps. malloticola CPC 19209 Laos: Mallotus thorelii Ps. schizolobii CBS 120029 HT Ecuador: Schizolobium parahyba Ps. pyracanthigena CBS 131589 HT South Korea: Pyracantha angustifolia Pseudocercospora sp. 064 CPC 24482 China: Citrus grandis Ps. euphorbiacearum COAD 1537 ET Brazil: Dalechampia sp. Ps. beijingensis BJFC WQ1808291 China: Euonymus japonicus Ps. beijingensis CGMCC 3.19020 HT China: Euonymus japonicus Ps. rhoina CBS 131891 South Korea: Rhus chinensis Ps. dingleyae CBS 114645 New Zealand: Haloragis erecta Ps. atromarginalis CPC 14649 South Korea: Solanum nigrum Ps. atromarginalis CBS 132010 South Korea: Solanum nigrum Ps. fuligena CBS 109729 Taiwan: Solanum lycopersicum Ps. fuligena PF001 USA: Solanum lycopersicum Ps. tabernaemontanae CPC 19198 ET Laos: Tabernaemontana coronaria Pseudocercospora sp. 059 CBS 149417 Australia: Santalum lanceolatum Pseudocercospora sp. 059 CBS 149420 Australia: Santalum acuminatum Pseudocercospora sp. 060 CPC 28350 Australia: Solanum sp. Pseudocercospora sp. 060 CBS 114642 Unknown: Unknown Pseudocercospora sp. 060 CBS 115122 New Zealand: Gunnera tinctoria Ps. atromarginalis COAD 1975 Brazil: Solanum americanum Ps. chengtuensis CBS 131924 South Korea: Lycium chinense Ps. fuligena CBS 132017 Thailand: Solanum lycopersicum Ps. atromarginalis CBS 114640 New Zealand: Solanum sp. Ps. atromarginalis CCTU 1193 Iran: Solanum nigrum Ps. atromarginalis CBS 136112 Iran: Solanum nigrum Ps. atromarginalis CCTU 1052 Iran: Solanum nigrum Ps. atromarginalis CCTU 1091 Iran: Solanum nigrum Ps. rauvolfiicola NFCCI 4586 HT India: Rauvolfia serpentina “Ps. cruenta (block 28)” CBS 132021 Trinidad and Tobago: Vigna sp. Ps. musarum sp. nov. CBS 149516 HT Malaysia: Musa sp. Ps. mangiferae CRC180 Thailand: Mangifera indica Ps. mangiferae CRC314 HT Thailand: Mangifera indica 26b 27 28 © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 44 yellow green to white; reverse olivaceous, Colonies reaching 34 mm diam. Typus: Australia, Tasmania, from leaf spots on Anopterus glandulosus (Escalloniaceace), 11 Dec. 2011, W. Quaedvlieg (holotype CBS H-25038, culture ex-type CPC 20152 = CBS 149359). Note: Pseudocercospora anopteri (Fig. 1, block 9) is introduced here as a new species, and represents the only species of Pseudocercospora known from the genus Anopterus, and family Escalloniaceace. Pseudocercospora asplenii Crous, R.G. Shivas & Yuan Yuan Chen, sp. nov. MycoBank MB 852263. Fig. 7. Etymology: Name refers to the host genus it was collected from, Asplenium. Description in vitro (SNA; CPC 17011): Mycelium subhyaline, smooth, delicate, uniform in width, 1.5–2.5 μm. Conidiophores microto macronematous, emerging from hyphae or conidia, pale brown, solitary, smooth or finely roughened, unbranched, cylindrical, straight to curved in segments, geniculate-sinuous at the apex, 1–2-septate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 3.8–25 × 2.3–4 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown, monoor polyblastic, proliferating sympodially, conidiogenous loci at the apex and shoulders, protruding and conically truncate, 2.8–10 × 2.3–4 μm, scars inconspicuous, 0.5– 1.5 μm diam. Conidia solitary, pale brown, guttulate, narrowly obclavate, apex obtuse to subobtuse, base obconically truncate to long obconically truncate, straight to slightly curved, (24–)30– 80 × (1.5–)2–3 μm, (1–)3–6-septate; hila neither thickened nor darkened-refractive, 0.5–1.5 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of ridges, grey olivaceous to smoke grey with patches of white; reverse mouse grey to vinaceous buff, Colonies reaching 37 mm diam. Typus: Australia, Queensland, Noosa, from leaf spots on Asplenium dimorphum (Aspleniaceae), 14 Jul. 2007, P.W. Crous (holotype CBS H-25031, culture ex-type CPC 17011 = CBS 149353). Additional material examined: Thailand, Chiang Rai, on Musa sp., 4 Dec. 2010, R. Cheewangkoon, culture CPC 19014. Notes: Pseudocercospora asplenii (Fig. 1, block 46) is introduced here as a new species, and represents the only species of Pseudocercospora known from the genus Asplenium, and family Aspleniaceae. For notes on Cercospora asplenii, see Braun et al. (2013a). Pseudocercospora australiensis Crous, Summerell & Yuan Yuan Chen, sp. nov. MycoBank MB 852264. Fig. 8. Etymology: Name refers to the country where it was collected, Australia. Leaf spots amphigenous, angular, delimited by leaf veins, 1–5 mm diam, medium brown with raised pale brown border. Fig. 6. Pseudocercospora anopteri (CPC 20152, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C. Conidiogenous cells giving rise to conidia. D. Conidia. Scale bars = 10 μm. Fig. 7. Pseudocercospora asplenii (CPC 17011, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C. Conidiogenous cells giving rise to conidia. D. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 45 Mycelium internal and external, pale brown, consisting of septate, branched, smooth hyphae, 2–3 µm diam. Caespituli fasciculate, hypophyllous, grey brown on leaves, up to 100 µm diam and 60 µm high. Conidiophores aggregated in loose to dense fascicles arising from the upper cells of a brown stroma, 40–70 µm diam; conidiophores medium brown, smooth, 2–4-septate, subcylindrical, straight to geniculate-sinuous, unbranched, 30– 45 × 4–6 µm. Conidiogenous cells terminal, unbranched, medium brown, smooth, tapering to flat-tipped apical loci, 2.5–4 µm diam, proliferating sympodially or percurrently near apex, 12–20 × 3–5 µm; scars inconspicuous, 3–3.5 µm diam. Conidia solitary, pale to medium brown, smooth, guttulate, subcylindrical, apex subobtuse, base truncate, straight to geniculate, 3–6-septate, (55–)58–65(–75) × (3.5–)4(–5) µm; hila neither thickened nor darkened-refractive with marginal frill, 3.5–4 µm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; Surface irregularly folded, with a prominent network of ridges, mouse grey to white; reverse dark mouse grey to pale mouse grey, Colonies reaching 11 mm diam. Typus: Australia, Western Australia, Mount Barker, from leaf spots on Eucalyptus gunnii (Myrtaceae), 23 Aug. 2015, P.W. Crous, HPC 606 (holotype CBS H-25053, culture ex-type CPC 29218 = CBS 149422). Additional material examined: Australia, Western Australia, Mount Barker, from leaf spots on E. caesia, 23 Aug. 2015, P.W. Crous, HPC 607, culture CPC 29068; Victoria, La Trobe State Forest, on leaves and twig cankers of E. bicostata, Nov. 2017, P.W. Crous, HPC 1874, culture CPC 32446. Notes: Pseudocercospora australiensis (Fig. 1, block 7) is phylogenetically distinct from all Pseudocercospora spp. occurring on Eucalyptus (Crous et al. 2019c), including those not known from culture (Braun & Dick 2002). It is presently known from three different eucalypt host species and was observed to also cause twig cankers on E. bicostata, which is a rarely observed disease symptom in the genus Pseudocercospora. Morphologically, it is characterised by loose to dense fascicles that give rise to short, wide, 3–6-septate, subcylindrical conidia. Pseudocercospora badjensis Crous, Summerell & Yuan Yuan Chen, sp. nov. MycoBank MB 852265. Figs 9, 10. Etymology: Name refers to the host species on which it occurs, Eucalyptus badjensis. Leaf spots amphigenous, sub-circular to angular, 3–5 mm diam, medium brown with raised dark brown border. Mycelium internal and external, pale to medium brown, consisting of septate, branched, smooth to finely verruculose hyphae, 2.5– 3.5 µm diam. Caespituli fasciculate, hypophyllous, grey brown on leaves, up to 50 µm diam and 220 µm high. Conidiophores arising singly from superficial mycelium, or aggregated in loose Fig. 8. Pseudocercospora australiensis (CPC 29218, ex-type culture). A. Leaf spots on lower and upper leaf surface. B. Close-up of lesion. C–F. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. G. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 46 fascicles arising from the upper cells of a brown stroma up to 40 µm diam and 30 µm high; conidiophores medium brown, finely verruculose, 4–10-septate, subcylindrical, straight to slightly curved, unbranched or branched above, 80–200 × 5–6 µm. Conidiogenous cells terminal or intercalary, unbranched, medium brown, finely verruculose, tapering to flat-tipped apical loci, proliferating sympodially and percurrently near apex, 25–35 × 4–5 µm; scars inconspicuous, 2–3 µm diam. Conidia solitary, medium brown, smooth, guttulate, narrowly obclavate, apex subobtuse, base long obconically truncate, straight to slightly curved, 4–10-septate, (45–)75–100(–120) × (3.5–)4(–5) µm; hila neither thickened nor darkened-refractive, 2.5–3 µm diam. Description in vitro (SNA; CPC 32376): Mycelium subhyaline to pale brown, smooth, delicate, 1.5–2 μm diam. Conidiophores macronematous, emerging from hyphae or conidia, pale brown to medium brown, solitary, smooth or roughened, unbranched to branched, cylindrical, straight to slightly curved in segments, geniculate-sinuous at the apex, 1–4-septate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 8.5–78 × 3–6 μm. Conidiogenous cells integrated, terminal or intercalary, pale to medium brown, monoor polyblastic, proliferating sympodially, unbranched, conidiogenous loci at the apex and shoulders, apex conically truncate, 8.5–31.5 × 2.5–5 μm; scars inconspicuous, 1–2 μm diam. Conidia solitary, pale brown, Fig. 9. Pseudocercospora badjensis (CPC 32376, ex-type culture). A. Leaf spots on upper and lower leaf surface. B. Close-up of lesion. C–H. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. I. Conidia. Scale bars = 10 μm. Fig. 10. Pseudocercospora badjensis (CPC 32376, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 47 guttulate, obclavate, apex obtuse, base obconically truncate, straight to slightly curved, 34–87 × 3–4 μm, 3–8-septate; hila neither thickened nor darkened-refractive, 1–2 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, with sparse to absent aerial mycelium, and smooth, rounded margins. Surface mouse grey to olivaceous grey with patches of dirty white; reverse dark mouse grey with olivaceous grey at centre. Colonies reaching 20 mm diam. Typus: Australia, Victoria, Mount Best, Tin Mine road, from leaf spots on Eucalyptus badjensis (Myrtaceae), 2015, P.W. Crous, HPC 1799 (holotype CBS H-25060, culture ex-type CPC 32376 = CBS 149427). Additional material examined: Australia, Victoria, inland from head of Wingan Inlet, Croajingolong National Park, on leaf spots of Eucalyptus sp. in natural habitat, 8 Nov. 2001, V. Beilharz, culture CPC 28316. Notes: Pseudocercospora badjensis (Fig. 1, block 9) is closely related to Ps. subulata and Ps. victoriae. It is morphologically distinct from species known from Eucalyptus in having superficial mycelium with solitary, long conidiophores, and narrowly obclavate, 4–10-septate conidia that can be up to 120 µm long (Braun & Dick 2002, Crous et al. 2019c). Pseudocercospora balanitis Crous, Jol. Roux & Yuan Yuan Chen, sp. nov. MycoBank MB 852266. Figs 11, 12. Etymology: Name refers to host genus on which it occurs, Balanites. Leaf spots amphigenous, sub-circular, 4–8 mm diam, grey brown with dark brown margin. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2.5–3 µm diam. Caespituli fasciculate, epiphyllous, brown on leaves, up to 120 µm diam and 160 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 100 µm diam and 50 µm high; conidiophores medium brown, smooth, 0–2-septate, subcylindrical, straight to slightly curved, unbranched, 15–30 × 4–5 µm. Conidiogenous cells terminal, pale to medium brown, smooth, tapering to flat-tipped apical loci, proliferating inconspicuously percurrently near apex, 7–15 × 4–5 µm; scars inconspicuous, 2–2.5 µm diam. Conidia solitary, pale brown, smooth, guttulate, subcylindrical, apex obtuse, base truncate, straight to curved, 1–3-septate, (50–)60–75(–80) × (3–)3.5(–4) µm; hila neither thickened nor darkened-refractive, 2–2.5 µm diam. Description in vitro (SNA; CPC 25271): Mycelium subhyaline to pale brown, smooth to verruculose, 1.5–4 μm diam. Conidiophores microto macronematous, emerging from hyphae, pale brown to brown, solitary, smooth or slightly roughened, unbranched, cylindrical, straight to slightly curved in segments, geniculate-sinuous at the apex, 1–3-septate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 16.5–50.5 × 5–7 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown to brown, monoor polyblastic, proliferating sympodially, conidiogenous loci at the apex and shoulders, apex conically truncate, 4.5–25 Fig. 11. Pseudocercospora balanitis (CPC 25271, ex-type culture). A. Leaf spots on lower and upper leaf surface. B. Close-up of lesion. C, D. Fascicles of conidiophores. E–G. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. H. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 48 × 4–6.5 μm; scars inconspicuous, 1–2 μm diam. Conidia solitary, pale brown, obclavate to filiform, apex obtuse to subobtuse, base long obconically truncate to truncate, straight to slightly curved, 35–95 × 3–4.5 μm, 4–10-septate; hila neither thickened nor darkened-refractive, 1–2.5 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, spreading, with sparse to absent aerial mycelium, and smooth, rounded margins. Surface smoke grey with patches of greyish sepia; reverse olivaceous. Colonies reaching 22 mm diam. Typus: South Africa, Limpopo Province, Louis Trichardt, from leaf spots on Balanites sp. (Zygophyllaceae), 2 Oct. 2014, J. Roux (holotype CBS H-25043, culture ex-type CPC 25271 = CBS 149363). Notes: Pseudocercospora balanitis (Fig. 1, block 15) is introduced here as a new species, and represents the only species of Pseudocercospora known from the genus Balanites, and family Zygophyllaceae. It is phylogenetically closely related to Ps. bonjeaneae-rectae on Lotus rectus and Ps. dovyalidis on Dovyalis zeyheri. Pseudocercospora bonjeaneae-rectae (Caball.) U. Braun, Schlechtendalia 40: 278. 2023. Fig. 13. Basionym: Cercospora bonjeaneae-rectae Cabal., Fac. Sci. Univ. Barcelona Publ. Secc. Ci. Nat. 12: 104. 1920. Synonyms: Cercospora bonjeaneae Maire [as ‘bonjeaniae’], Bull. Soc. Hist. Nat. Afrique N. 8: 193. 1917. nom. inval. Art. 36.1(a). Pseudocercospora bonjeaneae (Maire) U. Braun & Crous, in Crous & Braun, CBS Diversity Ser. (Utrecht) 1: 85. 2003, nom. inval. Art. 40.1 (Shenzhen). Leaf spots amphigenous, irregular to sub-circular, 7–12 mm diam, brown on adaxial and abaxial surface, with light brown borders. Mycelium internal to rarely external, pale brown, consisting of septate, branched, verruculose hyphae, 2–5 µm diam. Caespituli fasciculate, amphigenous, grey brown to olivaceous brown on leaves, up to 155 µm diam and 115 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 75 µm diam and 50 µm high; conidiophores medium brown, smooth, 0–1-septate, clavate, straight to slightly curved, unbranched, usually reduce to conidiogenous cell, 14– 32.5 × 5–6.5 µm. Conidiogenous cells terminal, unbranched, medium brown, smooth, tapering to obconically truncate apical loci, proliferating percurrently near apex, 15–25 × 4.5–6.5 µm; scars inconspicuous, 1–2.5 µm diam. Conidia solitary, pale to medium brown, smooth, guttulate, narrowly obclavate, apex obtuse, base long obconically truncate, straight to curved, slightly sinuous, 3–9-septate, 30–125 × 4.5–6.5 µm; hila neither thickened nor darkened-refractive, 1.5–3 µm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, spreading, with sparse aerial mycelium, surface irregularly folded, with a prominent network of ridges, grey white becoming pale mouse grey at margin; reverse dark mouse grey. Colonies reaching 28 mm diam. Typus: Spain, Barcelona, La Planas, on Lotus rectus (≡ Bonjeanea recta), Oct. 1919, A. Caballero 4315 (MA-FunHist 6514). Greece, Rhodos, Fig. 12. Pseudocercospora balanitis (CPC 25271, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. Fig. 13. Pseudocercospora bonjeaneae-rectae (CPC 31698, ex-epitype culture). A–C. Fascicles with conidiogenous cells giving rise to conidia. D. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 49 ca. 3.3 km W Archangelos, creek at the road to Malona, 36°13’06’’N, 28°04’49’’E, ca. 85 m alt, from leaf spots on Lotus rectus (≡ Dorycnium rectum), 19 Sep. 2016, V. Kummer, HPC 1369, U. Braun: Fungi selecti exsiccati 233 (epitype designated here CBS H-25057, MBT 10018174, culture ex-epitype CPC 31698 = CBS 149393). Notes: Pseudocercospora bonjeaneae-rectae (Fig. 1, block 15) was distributed as Fungi selecti exsiccati ex Herbario Universitatis Halensis No. 233 (duplicated in BPI, BRIP, GZU, HMAS, K, KR, KUS, LE, M, PDD) (Braun et al. 2017), which was also used to derive this ex-epitype culture. Because the name C. bonjeaniae was invalidly published, the synonym, C. bonjeaneae-rectae, was chosen (see Braun 2023 for explanation). Pseudocercospora bonjeaneae-rectae is phylogenetically closely related to Ps. balanitis on Balanites sp. and Ps. dovyalidis on Dovyalis zeyheri. Pseudocercospora dovyalidicola Crous & Yuan Yuan Chen, sp. nov. MycoBank MB 852267. Figs 14, 15. Etymology: Name refers to host genus on which it occurs, Dovyalis. Leaf spots amphigenous, sub-circular to circular, 2–7 mm diam, pale brown in centre, darker brown toward raised border. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2.5–3 µm diam. Caespituli fasciculate, amphigenous, brown on leaves, up to 120 µm diam and 100 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 80 µm diam and 40 µm high; conidiophores medium brown, finely verruculose to verrucose, 1–3-septate, subcylindrical, straight to geniculatesinuous, unbranched, 30–60 × 5–7 µm. Conidiogenous cells terminal, unbranched, medium brown, finely verruculose, tapering to flat-tipped apical loci, proliferating percurrently near apex, with rough, cercostigmina-like annellations, 10–22 × 5–7 µm; scars inconspicuous, 3.5–4.5 µm diam. Conidia solitary, medium brown, finely verruculose, guttulate, obclavate Fig. 14. Pseudocercospora dovyalidicola (CPC 25273, ex-type culture). A. Leaf spots on upper and lower leaf surface. B, C. Close-up of lesions. D–I. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. J. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 50 to subcylindrical, apex subobtuse, base obconically truncate, straight to slightly curved, (1–)5–8(–10)-septate, (35–)55– 75(–95) × (5–)6–7 µm; hila neither thickened nor darkenedrefractive, (2.5–)3–5 µm diam. Description in vitro (SNA; CPC 25275): Mycelium pale brown, smooth to slightly verruculose, 1.8–6 μm diam. Conidiophores microto macronematous, emerging from hyphae, pale brown to brown, solitary or fasciculate, smooth or slightly roughened, unbranched, cylindrical, straight to curved in segments, geniculate-sinuous at the apex, multiseptate, 10–213 × 4.5–6 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown to brown, monoor polyblastic, proliferating sympodially, conidiogenous loci at the apex and shoulders, protruding and conically truncate, apex conically truncate, 8–37 × 3.8–6 μm, scars inconspicuous, 1.3–2.5 μm diam. Conidia solitary, pale brown, guttulate, obclavate, apex obtuse to subobtuse, base obconically truncate to long obconically truncate, straight to slightly curved, 33.5–62 × 4–5.5 μm, 3–5-septate; hila neither thickened nor darkened-refractive, 1.5–2.5 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, spreading, with moderate aerial mycelium, and rough, rounded margins. Surface white to mouse grey; reverse olivaceous to mouse grey. Colonies reaching 12 mm diam. Typus: South Africa, KwaZulu-Natal Province, Drakensberg, The Hedges, from leaf spots on Dovyalis zeyheri (Salicaceae), 2014, J. Roux (holotype CBS H-25044, culture ex-type CPC 25273 = CBS 149364). Additional material examined: South Africa, KwaZulu-Natal Province, Drakensberg, The Hedges, from leaf spots on D. zeyheri, 2014, J. Roux (CBS H-25045, culture CPC 25275 = CBS 149365). Notes: Pseudocercospora dovyalidicola (Fig. 1, block 23) needs to be compared to Ps. dovyalidis (ex-epitype CBS 126002; Fig. 1, block 15) a foliar pathogen of D. zeyheri occurring in Gauteng Province of South Africa (conidiophores 12–34 × 3–6 µm, conidia smooth, pale brown or subhyaline, 1–10-septate, subcylindrical, (20–)30–70(–84) × (3–)3.5–5(–6) μm; Crous et al. 2013a). Although the two species are phylogenetically distinct, they show considerable morphological overlap, with Ps. dovyalidicola distinct in having longer conidiophores, wider conidia, and rougher percurrent proliferations on its conidiogenous cells. Pseudocercospora encephalarticola Crous & Yuan Yuan Chen, sp. nov. MycoBank MB 852268. Fig. 16. Etymology: Name refers to host genus on which it occurs, Encephalartos. Description in vitro (SNA; CPC 15278): Mycelium pale brown, smooth, 1.5–4 μm diam. Conidiophores microto macronematous, emerging from hyphae, pale brown to brown, solitary, smooth to slightly verruculose, unbranched to branched, cylindrical, straight to slightly curved in segments, geniculatesinuous at the apex, multiseptate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 20– 155 × 2–4.5 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown to brown, monoor polyblastic, proliferating sympodially or percurrently, conidiogenous loci at the apex and shoulders, protruding and conically truncate, apex truncate, 3–51.5 × 1.5–4.5 μm, scars inconspicuous, 1–2.5 μm diam. Conidia solitary, pale brown to medium brown, guttulate, cylindrical, apex rounded, base truncate, straight to slightly curved, 33–97 × 2.5–3 μm, 3–10-septate; hila neither thickened nor darkened-refractive, 1–2.5 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface folded with a prominent network of ridges, erumpent, spreading, with sparse aerial mycelium, and smooth, lobate margins. Surface smoke grey to white, with patches of dark mouse grey; reverse olivaceous to cinnamon. Colonies reaching 22 mm diam. Typus: South Africa, Western Cape, from leaf spots on Encephalartos sp. (Zamiaceae), 22 May 2008, A.R. Wood (holotype CBS H-25029, culture ex-type CPC 15278 = CBS 149351). Notes: Pseudocercospora encephalarticola (Fig. 1, block 7) needs to be compared to Ps. encephalarti (on leaves of Encephalartos barteri, from Benin), which is distinct in that it has obclavate to subacicular conidia, that are also longer and wider, (55–)90– 167.5(–190) × 3.5–4.5(–5) μm, 3–8-septate (Meswaet et al. 2019). Pseudocercospora erythrophleicola Crous, R.G. Shivas & Yuan Yuan Chen, sp. nov. MycoBank MB 852269; Fig. 17. Etymology: Name refers to the host genus from which it was collected, Erythrophleum. Fig. 15. Pseudocercospora dovyalidicola (CPC 25275). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 51 Leaf spots amphigenous, sub-circular, 2–20 mm diam, light brown to dark brown on adaxial and abaxial surface, with dark brown margin. Mycelium internal and rarely external, pale brown, consisting of septate, branched, smooth hyphae, 2–4 µm diam. Caespituli fasciculate, amphigenous, grey brown on leaves, up to 88 µm diam and 140 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 65 µm diam and 50 µm high; conidiophores medium brown, smooth, 0–3-septate, clavate to subcylindrical, straight to variously curved, unbranched, 13–21.5 × 4.5–6 µm. Conidiogenous cells terminal, unbranched, medium brown, smooth, tapering to flat-tipped apical loci, proliferating percurrently near apex, 7–25 × 3.5–6 µm; scars inconspicuous, 3–4 µm diam. Conidia solitary, pale brown, smooth, guttulate, cylindric to obclavate, apex obtuse to subobtuse, base long obconically truncate to truncate, straight to slightly curved, contract at septate, 3–8-septate, 17.5–60(–65) × 4–5.5 µm; hila neither thickened nor darkened-refractive, 2.5–3.5 µm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of ridges, pale mouse grey becoming lavender grey at margin; reverse dark mouse grey becoming mouse grey at margin. Colonies reaching 25 mm diam. Typus: Australia, Queensland, from leaf spots on Erythrophleum chlorostachys (Fabaceae), 10 Aug. 2009, P.W. Crous (holotype CBS H-25034, culture ex-type CPC 17241 = CBS 149391). Notes: Pseudocercospora erythrophleicola (Fig. 1, block 19) is distinguished from Ps. erythrophlei (on Erythrophleum chlorostachys, across northern Australia) by having large stromata and smaller conidia (stromata absent to small, conidia 44–100 × 3–4.5 µm in Ps. erythrophlei; Yuan 1996). Pseudocercospora eucleae Crous & B. Sutton, J. S. African Bot. 63: 283. 1997. Fig. 18. Leaf spots amphigenous, irregular to angular, 2–8 mm diam, dark brown with raised dark brown border. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2–2.5 µm diam. Caespituli fasciculate, hypophyllous, brown on leaves, up to 60 µm diam and 50 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 30 µm diam and 20 µm high; conidiophores medium brown, smooth, 1–4-septate, subcylindrical, straight to variously curved, unbranched or branched below, 20–40 × 5–7 µm. Conidiogenous cells terminal, pale to medium brown, smooth, tapering to flattipped apical loci, proliferating sympodially and percurrently, 7–16 × 3–4 µm; scars inconspicuous, 1.5–2 µm diam. Conidia solitary, pale brown, smooth, guttulate, narrowly obclavate, apex obtuse, base long obconically truncate, straight to slightly curved, (1–)5–7(–9)-septate, (50–)60–85(–120) × 3.5(–4) µm; hila neither thickened nor darkened-refractive, 1.5–2 µm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, spreading, with moderate/sparse to absent aerial mycelium, and smooth, lobate margins. Surface pale olivaceous grey; reverse olivaceous grey. Colonies reaching 13 mm diam. Fig. 16. Pseudocercospora encephalarticola (CPC 15278, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. Fig. 17. Pseudocercospora erythrophleicola (CPC 17241, ex-type culture). A–C. Fascicles with conidiogenous cells giving rise to conidia. D. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 52 Typus: South Africa, Gauteng Province, Warmbaths Rd. beyond Pienaars River, Euclea undulata (Ebenaceae), 20 Mar. 1950, P.H.B. Talbot (holotype PREM 39020); KwaZulu-Natal Province, Champagne resort, from leaf spots on Euclea sp., 16 Jan. 2017, P.W. Crous, HPC 1512 (epitype designated here CBS H-25064,MBT 10018178, culture ex-epitype CPC 32792 = CBS 149431). Notes: Pseudocercospora eucleae (Fig. 1, block 17) was described as a foliar pathogen of Euclea undulata from Gauteng Province in South Africa, characterised by irregular, grey to brown leaf spots up to 7 mm diam, amphigenous caespituli, 1–3-septate conidiophores, 15–45 × 3–6 µm, with inconspicuous percurrent proliferation, and subcylindrical to narrowly obclavate conidia, 1–6-septate, 35–90 × 3–4 µm (Crous & Sutton 1997). The present collection matches well with the morphology of the type specimen, and is herewith designated as epitype in order to obtain a culture and ex-type reference sequence data which are essential for taxonomic-phylogenetic purposes. Pseudocercospora gracilis Crous & Alfenas, Mycologia 87: 123. 1995. Fig. 19. Synonym: Mycosphaerella gracilis Crous & Alfenas, Mycologia 87: 123. 1995. Leaf spots absent, pseudothecial ascomata occurring on leaf litter. Pseudothecia amphigenous, predominantly hypophyllous, black, subepidermal, erumpent to superficial, globose, 40–70 µm diam; apical ostiole 10 µm diam; wall consisting of 2–3 layers of medium brown textura angularis. Asci aparaphysate, fasciculate, bitunicate, subsessile, obovoid to broadly ellipsoid, straight to slightly curved, 8-spored, (30–)35–45(–50) × (8–)9(– 10) µm. Ascospores multiseriate, overlapping, hyaline, guttulate, thin-walled, straight to slightly curved, narrowly fusoidellipsoidal with subobtuse ends, widest just above septum, medianly 1-septate, slightly constricted at the septum, tapering towards both ends, (14–)15–17(–20) × (2–)2.5(–3) µm. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of ridges, glaucous grey to mouse grey with patches of brown; reverse dark mouse grey to fawn; folds appearing cinnamon. Colonies reaching 20 mm diam. Typus: Indonesia, N. Sumatra, on Eucalyptus urophylla, 22 Nov. 1993, A.C. Alfenas (holotype of Mycosphaerella gracilis PREM 51718, holotype of Pseudocercospora gracilis PREM 51719, culture ex-type CBS 243.94 = CMW 14455 = CPC 730 = STE-U 730). Fig. 18. Pseudocercospora eucleae (CPC 32792, ex-epitype culture). A. Leaf spots on lower and upper leaf surface. B. Close-up of lesions. C–F. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. G. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 53 Additional materials examined: Indonesia, on Eucalyptus urophylla, 12 Mar. 1996, M.J. Wingfield, culture CPC 1314 = CBS 111372; on Eucalyptus urophylla, 12 Mar. 1996, M.J. Wingfield, culture CPC 1315 = CBS 111189; on Eucalyptus sp., Jan. 2011, M.J. Wingfield, culture CPC 19804. Malaysia, from leaf litter of Eucalyptus sp. (Myrtaceae), 22 Jun. 2012, M.J. Wingfield (holotype CBS H-25039, culture ex-type CPC 20850 = CBS 149390; CPC 20852) Notes: The isolates studied here are sterile in culture. Phylogenetically, they fall within the variation accepted for Ps. gracilis, a foliar pathogen of E. urophylla in Indonesia (Fig. 1, block 7). It is characterised by narrowly fusoid-ellipsoidal ascospores, (10–)15–18(–20) × (2–)2.5–3 μm, and uniformly cylindrical, pale olivaceous conidia (Crous et al. 2019c). Pseudocercospora grevilleae Crous & Yuan Yuan Chen, sp. nov. MycoBank MB 852270. Figs 20, 21. Etymology: Name refers to host genus on which it occurs, Grevillea. Leaf spots amphigenous, irregular to elongated, 2–3 mm diam, medium brown. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2–3 µm diam. Caespituli fasciculate, hypophyllous, dark brown on leaves, up to 110 µm diam and 160 µm high. Conidiophores aggregated in loose fascicles arising from the upper cells of a brown stroma up to 90 µm diam and 50 µm high; conidiophores medium brown, finely verruculose, 2–12-septate, subcylindrical, straight to variously curved, thick-walled, unbranched, 45–110 × 4–7 µm. Conidiogenous cells terminal, medium brown, finely verruculose, tapering to flat-tipped apical loci, proliferating percurrently near apex with rough annellations, 10–35 × 4–7 µm; scars inconspicuous, 2.5–3 µm diam. Conidia solitary, medium brown, finely verruculose, guttulate, obclavate to subcylindrical, apex obtuse, base obconically truncate, straight to curved, (3–)5–7(–8)-septate, (30–)45–70(–80) × (5–)6(–7) µm; hila neither thickened nor darkened-refractive, 2.5–3 µm diam. Description in vitro (SNA; CPC 17585): Mycelium subhyaline to pale brown, smooth, delicate, 1.5–5 μm diam. Conidiophores microto macronematous, emerging from hyphae, pale brown to brown, solitary, smooth, unbranched, cylindrical, straight to sinuous, geniculate-sinuous at the apex, 1–3-septate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 8–35 × 4–6.3 μm. Conidiogenous cells integrated, terminal, pale brown, monoor polyblastic, proliferating sympodially or percurrently, conidiogenous loci at the apex and shoulders, protruding and conically truncate, apex conically truncate, 7.5–20 × 4–6 μm, scars inconspicuous, 1–2 μm diam. Conidia solitary, pale brown, guttulate, obclavate, apex rounded, base obconically truncate to long obconically truncate, straight to strongly curved, 57–108 × 3.5–5 μm, 4–9-septate; hila neither thickened nor darkened-refractive, 1–2 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of Fig. 19. Pseudocercospora gracilis (CPC 20850). A. Leaf spots on lower and upper leaf surface. B. Close-up of lesion with ascomata. C–E. Asci and ascospores. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 60 Mpumalanga Province, South Africa (Chupp & Doidge 1948). The present collection correlates well with Ps. halleriae, which causes angular leaf spots and has hypophyllous caespituli, short conidiophores, 10–35 × 2–4 µm, and 1–5-septate, obclavate conidia, 15–70 × 1.5–3 µm. The present collection is herein designated as epitype. Pseudocercospora lonicerae-japonicae Crous, Y. Zhang ter. & Yuan Yuan Chen, sp. nov. MycoBank MB 852272. Fig. 30. Etymology: Name refers to the host genus from which it was collected, Lonicera. Leaf spots amphigenous, irregular to sub-circular, delimited by leaf veins, 1–8 mm diam, sometimes confluent up to 20 mm, purple brown to brown on adaxial surface, yellowish or pale to medium brown on abaxial surface, without definite margin. Mycelium internal and external, pale brown, consisting of septate, branched, smooth hyphae, 1.5–3 µm diam. Caespituli fasciculate, inconspicuous, pale brown on abaxial surface, up to 45 µm diam and 100 µm high. Conidiophores arranged in small loose fascicles, arising from internal hyphae, emerging from very small stromata composed of a few brown hyphal cells, or arising from superficial hyphae; conidiophores pale to medium Fig. 30. Pseudocercospora lonicerae-japonicae (CPC 30538, ex-type culture). A–D. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. Fig. 31. Pseudocercospora lophostemonigena (CPC 16409, ex-type culture). A. Leaf spots on lower and upper leaf surface. B. Close-up of lesion. C–F. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. G. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 61 brown, smooth, 0–7-septate, subcylindrical, straight to variously curved, branched below, 10–80 × 3–5.5 µm. Conidiogenous cells terminal, unbranched, pale brown, smooth, tapering to flattipped apical loci, proliferating percurrently near apex, 10–27 × 3–5 µm; scars inconspicuous, 1.5–3 µm diam. Conidia solitary, pale brown, smooth, guttulate, narrowly obclavate, apex obtuse to subobtuse, base long obconically truncate, straight to slightly curved, 3–9-septate, (30–)40–85(–90) × 2.5–4 µm; hila neither thickened nor darkened-refractive, 1.5–2.5 µm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of ridges, mouse grey becoming white at margin; reverse dark mouse grey becoming smoke grey at margin. Colonies reaching 25 mm diam. Typus: China, Guizhou Province, from leaf spots on Lonicera japonica (Caprifoliaceae), 2015, J.J. Gan, HPC 1112 = GJJ 160212-31 (holotype CBS H-25054, culture ex-type CPC 30538 = CBS 149389). Additional material examined: South Korea, from leaf spots on Lonicera japonica (Caprifoliaceae), 7 Nov. 2007, H.D Shin, culture CPC 14801. Notes: Pseudocercospora lonicerae-japonicae (Fig. 1, block 9) differs from Ps. lonicerae and Ps. lonicerigena in having smaller to absent stromata, and smaller conidia (40–120 × 3.2–6 µm in Ps. lonicerae; 20–100 × 2–4 µm in Ps. lonicerigena; Guo 1994, Braun & Crous 2007). Phylogenetically, these taxa are distinct from Ps. lonicericola, conidia 2–6-septate, 25–64 × 2.5–5 μm (on Lonicera gracilipes var. glabra, Japan) (Fig. 1, block 35) (see Nakashima et al. 2016). Pseudocercospora lophostemonigena Crous, Summerell & Yuan Yuan Chen, sp. nov. MycoBank MB 852274. Fig. 31. Etymology: Epithet composed of the name of the host genus on which it occurs, Lophostemon, + -genus (produced in a certain place). Fig. 32. Pseudocercospora lophostemonis (CPC 14517, ex-type culture). A. Leaf spots on lower and upper leaf surface. B. Close-up of lesion. C. Closeup of fascicles. D–I. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. J. Conidia. Scale bars = 10 μm © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 62 Leaf spots amphigenous, irregular to sub-circular, 1–8 mm diam, grey brown with raised red-purple border. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2.5–3 µm diam. Caespituli fasciculate to sporodochial, amphigenous, grey brown on leaves, up to 110 µm diam and 120 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 85 µm diam and 50 µm high; conidiophores medium brown, smooth, 1–9-septate, subcylindrical, straight to variously curved, unbranched, 25–90 × 4–5 µm. Conidiogenous cells terminal, unbranched, medium brown, smooth, tapering to flat-tipped apical loci, proliferating sympodially and percurrently near apex, 10–16 × 3–4 µm; scars inconspicuous, 2–2.5 µm diam. Conidia solitary, medium brown, smooth, guttulate, narrowly obclavate, apex subobtuse, base truncate, straight to slightly curved, 3–6-septate, (35–)45–55(– 65) × 3(–3.5) µm; hila neither thickened nor darkened-refractive, 2–2.5 µm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of ridges, olivaceous grey with patches of dirty white; reverse grey sepia becoming smoke grey at margin; folds appearing dark slate blue, Colonies reaching 28 mm diam. Typus: Australia, New South Wales, Washpool National Park, S29°11’201 E152°25’70”, 789 m, from leaf spots on Lophostemon confertus (Myrtaceae), Mar. 2009, B.A. Summerell (holotype CBS H-25030, culture ex-type CPC 16409 = CBS 149352). Notes: Pseudocercospora lophostemonigena (Fig. 1, block 9) needs to be compared to Ps. sawadae (on Psidium guajava, Taiwan; Fig. 1, block 26a) and Ps. lophostemonicola (on Lophostemon, New Zealand). It can be easily distinguished from both species by having 3–6-septate, narrowly obclavate conidia that are shorter than 65 µm (see Braun et al. 2013b). See also the species notes of Ps. lophostemonis. Pseudocercospora lophostemonis Crous, Summerell & Yuan Yuan Chen, sp. nov. MycoBank MB 852275. Figs 32, 33. Etymology: Name refers to the host genus on which it occurs, Lophostemon. Leaf spots amphigenous, irregular to sub-circular, 3–8 mm diam, pale to medium brown with diffuse red-brown border. Mycelium internal pale brown, consisting of septate, branched, smooth hyphae, 2–2.5 µm diam. Caespituli fasciculate, amphigenous, grey brown on leaves, up to 130 µm diam and 60 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 110 µm diam and 50 µm high; conidiophores medium brown, finely verruculose, 1–3-septate, subcylindrical, straight to geniculate-sinuous, unbranched or branched below or above, 20–45 × 4–6 µm. Conidiogenous cells terminal or intercalary, medium brown, finely verruculose, tapering to flat-tipped apical loci, proliferating sympodially and percurrently near apex, 15–25 × 3.5–4 µm; scars inconspicuous, 2–2.5 µm diam. Conidia solitary, medium brown, smooth, guttulate, subcylindrical to narrowly obclavate, apex obtuse, base obconically truncate, straight, (1–)3-septate, (30–)35–40(– 45) × (3.5–)4 µm; hila neither thickened nor darkened-refractive, 2 µm diam. Description in vitro (SNA; CPC 14517): Mycelium hyaline to pale brown, smooth, delicate, 1.5–2.5 μm. Conidiophores microto macronematous, emerging from hyphae, pale brown, solitary, smooth, unbranched, cylindrical, straight to slightly curved in segments, geniculate-sinuous at the apex, 1–3-septate, sometimes reduced to conidiogenous cells, straight, holoblastic, 7–47 × 2.5–5 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown, monoor polyblastic, proliferating sympodially, aseptate, unbranched, conidiogenous loci at the apex and shoulders, apex conically truncate, 6–30 × 2.5–5 μm, scars inconspicuous, 1–1.5 μm diam. Conidia solitary, pale brown, guttulate, narrowly obclavate to filiform, apex obtuse to subobtuse, base obconically truncate to long obconically truncate, straight to slightly curved, 45–90(–95) × 2.5–3.5 μm, 5–10-septate; hila neither thickened nor darkened-refractive, 1 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface folded with a prominent network of ridges, with smooth, lobate margins. Surface olivaceous grey to pale olivaceous grey; reverse olivaceous to mouse grey. Colonies reaching 26 mm diam. Typus: Australia, Northern Territory, Robin Falls, from leaf spots on Lophostemon lactifluus (Myrtaceae), 23 Sep. 2007, B.A. Summerell (holotype CBS H-25026, culture ex-type CPC 14517 = CBS 149350). Notes: Pseudocercospora lophostemonis (Fig. 1, block 22) needs to be compared to Ps. lophostemonicola (on Lophostemon confertus, New Zealand, conidiophores solitary on superficial Fig. 33. Pseudocercospora lophostemonis (CPC 14517, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 63 mycelium, conidia 3–10-septate, 30–105 × 3–5 μm), from which it is distinguished based on its dense fasciculate conidiophores, and smaller, (1–)3-septate conidia. It is also distinct from Ps. lophostemonigena (Lophostemon confertus, Australia; Fig. 1, block 9) which has longer conidia with more septa, 3–6-septate, (35–)45–55(–65) × 3(–3.5) µm. For notes on Ps. sawadae (Fig. 1, block 26a), see Braun et al. (2013b). Pseudocercospora macadamiae Beilharz et al., Australas. Pl. Path. 32: 280. 2003. Typus: Australia, Queensland, Glasshouse Mountains, on living husks of Macadamia integrifolia var. Hinde H2, 5 Feb. 1999, P. Mayers (holotype VPRI 21900, specimen confirmed lost); lectotype here designated, Australas. Pl. Path. 32: 281. 2003, fig. 1, MBT 10018454. Australia, Queensland, Glasshouse Mountains, on living husks of Macadamia integrifolia var. Hinde H2, 12 Nov. 2011, O.A. Akinsanmi (designated as epitype of Ps. macadamiae Beilharz et al. here: BRIP 55526, MBT 10018192, preserved as metabolically inactive culture, and as specimen, culture ex-epitype BRIP 55526 = CBS 133432). Notes: Pseudocercospora macadamiae (Fig. 1, block 9) causes husk spot of Macadamia in Australia, resulting in premature fruit abscission, and giving rise to nuts with low oil content (Beilharz et al. 2003). To help understand the mechanism of disease development, its spread, and subsequent role in fruit abscission, Akinsanmi & Carvalhais (2020) published a draft genome sequence of a culture they accepted as ex-epitype of Ps. macadamiae (BRIP 55526), but this typification was never formally introduced, and thus it is done here. The holotype of Ps. macadamiae has the same collection details as the proposed epitype specimen BRIP 55526 collected from the same farm 12 years later. Pseudocercospora musarum Crous & Yuan Yuan Chen, sp. nov. MycoBank MB 852276. Figs 34, 35. Etymology: Name refers to host genus on which it occurs, Musa, in genitive plural. Leaf spots amphigenous, lens-shaped to irregular, 1–4 mm diam, grey brown with raised border and dark brown margin. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 1.5–2 µm diam. Caespituli fasciculate, amphigenous, pale brown on leaves, up to 50 µm diam and 50 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 60 µm diam and 20 µm high; conidiophores pale brown, smooth, reduced to conidiogenous cells, ampulliform to subcylindrical, proliferating percurrently near apex, 10–15 × 5–8 µm; scars inconspicuous, 2 µm diam. Conidia solitary, pale olivaceous, smooth, guttulate, subcylindrical, apex obtuse, base truncate, straight to variously curved, 3–6-septate, (55–)60–80(–90) × (2.5–)3(–3.5) µm; hila neither thickened nor darkened-refractive, 2–2.5 µm diam. Fig. 34. Pseudocercospora musarum (CPC 32815, ex-type culture). A. Leaf spots on upper and lower leaf surface. B. Close-up of lesions. C. Close-up of stroma. D–G. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. H. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 64 Description in vitro (SNA; CPC 32815): Mycelium hyaline to pale brown, smooth, delicate, 1.5–2.3 μm diam. Conidiophores macronematous, emerging from hyphae or conidia, pale brown, solitary, smooth, unbranched, cylindrical, straight to curved in segments, geniculate-sinuous at the apex, 1-septate, sometimes reduced to conidiogenous cells, straight to curved in segments, unbranched, holoblastic, 7.5–11 × 2.7–3.5 μm. Conidiogenous cells integrated, terminal, pale brown, monoor polyblastic, proliferating sympodially, unbranched, conidiogenous loci at the apex and shoulders, protruding and conically truncate, apex conically truncate, 4.5–8 × 2.5–3.5 μm, scars inconspicuous, 1 μm diam. Conidia solitary, hyaline to pale, guttulate, filiform, apex obtuse, base truncate to long obconically truncate, straight to slightly curved, 50–100 × 1.52.5 μm, 3–5-septate; hila neither thickened nor darkened-refractive, 1–1.5 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, spreading, with moderate aerial mycelium, and smooth, lobate margins. Surface pale olivaceous grey; reverse saffron to smoke grey. Colonies reaching 35 mm diam. Typus: Malaysia, from leaf spots on Musa sp. (Musaceae), 2010, P.W. Crous, HPC 1610 (holotype CBS H-25065, culture ex-type CPC 32815 = CBS 149516). Notes: Pseudocercospora musarum (Fig. 1, block 28) is phylogenetically distinct from all Pseudocercospora spp. presently recognised on Musa (Crous et al. 2021). Using the key provided by Braun et al. (2014), it is similar to Ps. assamensis, though phylogenetically distinct (Fig. 1, block 18). Pseudocercospora musigena Crous, Alfenas & Yuan Yuan Chen, sp. nov. MycoBank MB 852277; Figs 36, 37. Etymology: Epithet composed of the name of the host genus on which it occurs, Musa, + -genus (produced in a certain place). Leaf spots amphigenous, elongated angular, 1–2 mm diam, 4–20 mm long, pale brown with dark brown raised margin. Fig. 35. Pseudocercospora musarum (CPC 32815, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. Fig. 36. Pseudocercospora musigena (CPC 18460, ex-type culture). A. Leaf spots on lower and upper leaf surface. B. Close-up of lesion. C. Close-up of stroma. D. Spermatogonium. E, F. Fascicles with conidiogenous cells giving rise to conidia. G. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 65 Spermatogonia amphigenous, chiefly epiphyllous, dark brown, substomatal, erumpent, globose, 50–90 µm diam, with central ostiole; wall consisting of 2–3 layers of brown textura angularis. Mycelium internal pale brown, consisting of septate, branched, smooth hyphae, 2–2.5 µm diam. Sporulation sparse. Conidiophores arising from the upper cells of spermatogonia, pale to medium brown, smooth, reduced to conidiogenous cells, straight, unbranched, holoblastic, 10–30 × 3–4 µm; scars inconspicuous, 2–2.5 µm diam. Conidia solitary, medium brown, smooth, guttulate, subcylindrical to narrowly obclavate, apex obtuse, base narrowly obclavate, straight to slightly curved, (1–)3(–5)-septate, (25–)35–50(–60) × (4–)5(–6.5) µm; hila neither thickened nor darkened-refractive, 3–3.5 µm diam. Description in vitro (SNA; CPC 18460): Mycelium subhyaline, smooth, delicate, 1–2.5 μm diam. Conidiophores macronematous, pale brown, emerging from hyphae or conidia, solitary, cylindrical, smooth, unbranched, 1–2-septate, straight to curved in segments, geniculate-sinuous at the apex, sometimes reduced to conidiogenous cells, straight, holoblastic, 5–21 × 2–3.5 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown, monoor polyblastic, proliferating sympodially, aseptate, unbranched, conidiogenous loci at the apex and shoulders, protruding and conically truncate, apex conically truncate, 4–15 × 2–3.5 μm; scars inconspicuous, 1–1.5 μm diam. Conidia solitary, subhyaline to pale brown, guttulate, obclavate to filiform, apex obtuse to subobtuse, base obconically truncate to truncate, straight to slightly curved, 38–110(–135) × 2–3 μm, 3–9(–11)-septate; hila neither thickened nor darkenedrefractive, 1–1.5 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, aerial mycelium absent, with smooth, lobate margins. Surface irregularly folded, pale mouse grey with patches of rosy buff; reverse olivaceous to mouse grey. Colonies reaching 27 mm diam. Typus: Brazil, Minas Gerais, Viçosa, from leaf spots on Musa sp. (Musaceae), 1990, P.W. Crous (holotype CBS H-25036, culture ex-type CPC 18460 = CBS 149357). Notes: Pseudocercospora musigena (Fig. 1, block 4) is phylogenetically distinct from all Pseudocercospora spp. presently recognised on Musa (Crous et al. 2021; Fig. 1, blocks 4, 9, 18, 28, 30, 31, 33, and 46), and is closely related to Ps. norchiensis obtained from Rubus from New Zealand. Pseudocercospora paracydoniae Crous, H.D. Shin & Yuan Yuan Chen, sp. nov. MycoBank MB 852278. Figs 38, 39. Etymology: Name refers to its morphological similarity to Ps. cydoniae. Leaf spots amphigenous, irregular to sub-circular, 2–8 mm diam, medium brown with raised border. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2–2.5 µm diam. Caespituli fasciculate, epiphyllous, iron-grey on leaves, up to 60 µm diam and 70 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 50 µm diam and 50 µm high; conidiophores medium brown, smooth, 1–3-septate, subcylindrical, straight to variously curved, unbranched, 25–35 × 4–5 µm. Conidiogenous cells terminal, medium brown, smooth, tapering to flat-tipped apical loci, proliferating sympodially and percurrently near apex, 12–20 × 4–5 µm; scars inconspicuous, 2–2.5 µm diam. Conidia solitary, medium brown, smooth, guttulate, narrowly obclavate, apex subobtuse, base obconically truncate, straight to curved, (1–)3(–6)-septate, (35–)45–55(–65) × (3–)4 µm; hila neither thickened nor darkened-refractive, 2.5–3 µm diam. Description in vitro (SNA; CPC 25478): Mycelium subhyaline to pale brown, smooth to verruculose, delicate, 1.3–3.5 μm diam. Conidiophores micronematous, emerging from hyphae, pale brown, solitary, smooth or roughened, unbranched, cylindrical, straight to slightly curved in segments, geniculate-sinuous at the apex, 1–3-septate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 13–37 × 3–5 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown, monoor polyblastic, proliferating sympodially, conidiogenous loci at the apex and shoulders, protruding and conically truncate, apex conically truncate, 6–15.5 × 3–4 μm; scars inconspicuous, 1–1.5 μm diam. Conidia solitary, pale brown, guttulate, obclavate to filiform, apex obtuse to subobtuse, base obconically truncate to long obconically truncate, straight to slightly curved, 40–99 × 2.5–3.5 μm, 3–10-septate; hila neither thickened nor darkenedrefractive, 1.5–2 μm diam. Culture characteristics: Colonies after 2 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of ridges, smoke grey to mouse grey with patches of dirty white; reverse olivaceous. Colonies reaching 28 mm diam. Fig. 37. Pseudocercospora musigena (CPC 18460, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 66 Typus: South Korea, Seoul, from leaf spots on Chaenomeles speciosa (Rosaceae), 28 Oct. 2014, P.W. Crous, HPC 8 (holotype CBS H-25046, culture ex-type CPC 25478 = CBS 149367). Notes: Pseudocercospora paracydoniae (Fig. 1, block 20) needs to be compared to Ps. cydoniae (on Cydonia japonica, USA; conidia 1–3-septate, 30–45 × 2.5 µm; Fig. 1, block 20). The latter fungus has been reported from Asia [China, Japan (MUCC 1184) and Korea] (Hsieh & Goh 1990, Shin & Kim 2001), and placed into phylogenetic context by Crous et al. (2013a) based on a Korean isolate (CPC 10678). Shin & Kim (2001) also discuss morphological variation within the Asian collections, but still accepted it as one species. As we have shown here, however, there are several distinct phylogenetic species in Asia, with overlapping morphology. Fresh collections are required from the USA, to determine if the fungus on C. japonica is conspecific with material studied from Asia on C. speciosa. Pseudocercospora parakaki Crous, H.D. Shin & Yuan Yuan Chen, sp. nov. MycoBank MB 852280. Fig. 40. Etymology: Name refers to its morphological similarity to Pseudocercospora kaki. Leaf spots amphigenous, irregular to sub-circular, delimited by leaf veins, 1–10 mm diam, red brown on adaxial surface, medium brown on abaxial surface, with raised dark brown borders. Mycelium internal to rarely external, pale brown, consisting of septate, branched, smooth hyphae, 1.5–3.5 µm diam. Caespituli fasciculate, amphigenous, grey brown on leaves, up to 130 µm diam and 145 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 70 µm diam and 60 µm high; conidiophores medium brown, smooth, 1–6-septate, subcylindrical, straight to variously curved, unbranched, 30–62 × 3–4.5. Conidiogenous cells terminal, unbranched, medium brown, smooth, tapering to flat-tipped apical loci, proliferating percurrently near apex, 13.5–25.5 × 3–4.5 µm; scars inconspicuous, 1.5–2.5 µm diam. Conidia solitary, pale brown, smooth, guttulate, narrowly obclavate, apex obtuse to subobtuse, base long obconically truncate, straight to slightly curved, 3–9-septate, (30–)40–75(– 85) × 3–4 µm; hila neither thickened nor darkened-refractive, 1.5–2.5 µm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of ridges, pale mouse grey; reverse dark mouse grey becoming vinaceous buff at margin. Colonies reaching 38 mm diam. Typus: South Korea, from leaf spots on Diospyros kaki (Ebenaceae), 30 Oct. 2007, H.D. Shin (holotype CBS H-25028, culture ex-type CPC 14686 = CBS 149388). Additional material examined: South Korea, Gongju, on D. lotus, 28 Oct. 2003, H.D. Shin, CBS H-20871, cultures CBS 132019 = CPC 10837, CPC 10838, and CPC 10839. Fig. 38. Pseudocercospora paracydoniae (CPC 25478, ex-type culture). A. Leaf spots on upper and lower leaf surface. B. Close-up of lesion. C, D. Fascicles with conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. Fig. 39. Pseudocercospora paracydoniae (CPC 25478, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 67 Notes: The Pseudocercospora spp. occurring on Diospyros were treated by Braun et al. (2020). Crous et al. (2013a) reported a collection from D. lotus from South Korea (CPC 10837–10839) as Ps. kaki. Braun et al. (2020) subsequently treated the Korean material from D. lotus as a new species, Ps. diospyriphila (Fig. 1, block 25). In the present study we add an additional collection, namely from D. kaki, and show this taxon treated as ‘Ps. kaki’ by Crous et al. (2013a) to represent a new species, Ps. parakaki (Fig. 1, block 7) distinct from Ps. kaki (Fig. 1, block 37) and Ps. diospyriphila (Fig. 1, block 25). Conidia of Ps. diospyriphila are shorter, 15–55 × 2.5–3.5 μm, 0–3-septate, while those of Ps. kaki are similar, (15–)25–80(–100) × 2–4 μm, 1–9-septate, although these species are phylogenetically distinct (Braun et al. 2020). Pseudocercospora paramacadamiae Crous & Yuan Yuan Chen, sp. nov. MycoBank MB 852281. Fig. 41. Etymology: Name refers to its similarity to Pseudocercospora macadamiae. Description in vitro (SNA; CPC 19150): Mycelium hyaline to pale brown, smooth to slightly sinuous, delicate, 1.5–3 μm diam. Conidiophores microto macronematous, emerging from hyphae or conidia, pale brown, solitary, smooth, unbranched to branched, cylindrical, straight to curved, geniculate-sinuous at the apex, 1–3-septate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 12–47 × 3–4.5 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown to medium brown, monoor polyblastic, proliferating sympodially, conidiogenous loci at the apex and shoulders, apex conically truncate, 7.5–29 × 3–3.5 μm, scars inconspicuous, 1–1.5 μm diam. Conidia solitary, pale brown, guttulate, obclavate, apex obtuse to subobtuse, base obconically truncate, straight to slightly curved, 34–61.5 × 2.5–3.5 μm, 3–5-septate; hila neither thickened nor darkened-refractive, 1–1.5 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, spreading, with sparse to absent aerial mycelium, and smooth, lobate margins. Surface dirty white with patches of pale mouse grey; reverse vinaceous grey. Colonies reaching 29 mm diam. Typus: Australia, New South Wales, Valla, Sullivans Road, Gunder plantation, from leaf spots on Macadamia integrifolia (Proteaceae), 14 Oct. 1997, B. Maier (holotype CBS H-25179, culture ex-type CPC 19150 = CBS 129067). Additional material examined: Australia, New South Wales, Valla, Sullivans Road, Gunder plantation, from leaf spots on Macadamia integrifolia (Proteaceae), 14 Oct. 1997, B. Maier, culture CPC 19149 = CBS 129066. Notes: Pseudocercospora paramacadamiae represents a sister clade (Fig. 1, block 9) to Ps. macadamiae (conidia subcylindrical, (0–)5–9-septate, (17–)45–69 × 2–2.5 μm; Beilharz et al. 2003), having wider conidia that on average have fewer septa. Isolates of Ps. paramacadamiae were also isolated from nuts with husk spot disease symptoms, suggesting that two species are associated with the disease in Australia. Pseudocercospora persooniae Crous, Summerell & Yuan Yuan Chen, sp. nov. MycoBank MB 852282. Figs 42, 43. Fig. 40. Pseudocercospora parakaki (CPC 14686, ex-type culture). A–D. Fascicles with conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. Fig. 41. Pseudocercospora paramacadamiae (CPC 19150, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 68 Etymology. Name refers to host genus on which it occurs, Persoonia. Leaf spots amphigenous, associated with tip blight, medium brown with raised red-purple border. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2–2.5 µm diam. Caespituli fasciculate to sporodochial, hypophyllous, grey brown on leaves, up to 160 µm diam. Conidiophores aggregated in dense fascicles arising from the upper cells of a brown stroma up to 100 µm diam and 40 µm high; conidiophores medium brown, smooth to finely verruculose, 1–4-septate, subcylindrical, straight to geniculate-sinuous, unbranched or branched above, 25–50 × 4–6 µm. Conidiogenous cells terminal or intercalary, unbranched, pale brown, smooth to finely roughened, tapering to flat-tipped apical loci, proliferating sympodially to rarely percurrently near apex, 12–20 × 3–5 µm; scars inconspicuous, 2–3 µm diam. Conidia solitary, pale brown, smooth, guttulate, subcylindrical to narrowly obclavate, apex obtuse, base truncate, straight to curved to once geniculate, (3–)5–7-septate, (35–)45–65(–70) × (3–)3.5–4 µm; hila neither thickened nor darkened-refractive, 2–2.5 µm diam. Description in vitro (SNA; CPC 32456): Mycelium subhyaline to pale brown, smooth to slightly verruculose, delicate, 1.5–3.5 μm diam. Conidiophores microto macronematous, emerging from hyphae, pale brown, solitary, smooth or slightly roughened, Fig. 42. Pseudocercospora persooniae (CPC 32456, ex-type culture). A. Leaf spot on lower leaf surface. B. Close-up of lesion. C. Close-up of fascicles. D–H. Fascicles with conidiogenous cells giving rise to conidia. I. Conidia. Scale bars = 10 μm. Fig. 43. Pseudocercospora persooniae (CPC 32456, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 69 unbranched, cylindrical, straight to curved in segments, geniculate-sinuous at the apex, multiseptate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 7–110 × 3–6 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown, monoor polyblastic, proliferating sympodially, unbranched to branched above, conidiogenous loci at the apex and shoulders, protruding truncate, apex conically truncate or truncate, 6.5–23 × 3–6 μm, scars inconspicuous, 1–2 μm diam. Conidia solitary, pale brown, guttulate, obclavate, apex obtuse, base obconically truncate, straight to slightly curved, (38–)45– 75 × (3–)3.5–4.5 μm, 4–10-septate; hila neither thickened nor darkened-refractive, 1–2 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, aerial mycelium absent, with smooth, lobate margins. Surface irregularly folded, grey olivaceous becoming pale lavender grey to dirty white at margin; reverse vinaceous grey to olivaceous. Colonies reaching 22 mm diam. Typus: Australia, Victoria, Royal Botanical Gardens Melbourne, from leaf spots on Persoonia sp. (Proteaceae), 2015, P.W. Crous, HPC 1920 (holotype CBS H-25063, culture ex-type CPC 32456 = CBS 149430). Additional material examined: Australia, Victoria, Sardine Creek, from leaves of unidentified tree host, 7 Nov. 2006, P.W. Crous, CBS H-25052, culture CPC 28349 = CBS 149421. Notes: Pseudocercospora persooniae (Fig. 1, block 9) represents the second Pseudocercospora species described from this host. It is phylogenetically distinct from the first species, Ps. wuchienshiungiae (Fig. 1, block 14). Several species have to date been recorded from Proteaceae (Crous et al. 2013b), which all differ from Ps. persooniae phylogenetically. It is closely related to Ps. christellae (from Christella parasitica (Thelypteridaceae), Thailand; Fig. 1, block 9), but morphologically distinct in that the latter has longer conidia (3–9-septate, 53–105 × 2–4 μm), and shorter conidiophores (0–1-septate, 9–14 × 2–4 µm; Phengsintham et al. 2013a). Pseudocercospora christellae is currently only known from an ITS sequence and its position in the phylogenetic tree is not stable between different analyses; in the IQ-TREE analysis (Fig. 1) it clusters with Pseudocercospora sp. 014 in block 9 and in the RAxML analysis it clusters among strains of Ps. badjensis (see Figshare; equivalent of clustering in Fig. 1, block 9) (Suppl. Table S3). The ITS sequence of Ps. christellae is 99 % (identity 339/340, including one gap) similar to the ITS sequences of Pseudocercospora sp. 014 and Ps. badjensis. Pseudocercospora pseudocydoniae Crous, H.D. Shin & Yuan Yuan Chen, sp. nov. MycoBank MB 852283. Fig. 44. Etymology: Name refers to the epithet of Pseudocercospora cydoniae, with which it was confused in the past, expressed by adding the prefix pseudo-. Leaf spots amphigenous, irregular to angular, delimited by leaf veins, 1–10 mm diam, sometimes confluent up to 20 mm, dark brown with grey center on adaxial surface, dark brown on abaxial surface, without definite margin. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2–4.5 µm diam. Caespituli fasciculate to sporodochial, amphigenous, grey on leaves, up to 115 µm diam and 150 µm high. Conidiophores Fig. 44. Pseudocercospora pseudocydoniae (CPC 14665, ex-type culture). A–C. Fascicles with conidiogenous cells giving rise to conidia. D. Conidia. Scale bars = 10 μm. Fig. 45. Pseudocercospora pultenaeae (CPC 28318, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 76 Phylogenetically it is closely related to Ps. persooniae (conidia subcylindrical, (3–)5–7-septate, (35–)45–65(–70) × (3–)3.5–4 µm), thus being morphologically clearly distinct. It is also phylogenetically distinct from species known from Eucalyptus. Pseudocercospora victoriae Crous, Summerell & Yuan Yuan Chen, sp. nov. MycoBank MB 852289. Figs 53–55. Etymology: Name refers to the state in Australia where it was collected, Victoria. Leaf spots amphigenous, irregular to angular, 2–8 mm diam, medium brown with raised border and red purple margin. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2.5–3 µm diam. Caespituli fasciculate, amphigenous, pale grey brown on leaves, up to 80 µm diam and 70 µm high. Conidiophores aggregated in loose fascicles arising from the upper cells of a brown stroma up to 25 µm diam and 20 µm high; conidiophores medium brown, smooth, 1–3-septate, subcylindrical, straight to slightly curved, unbranched, 35–70 × 4–5 µm. Conidiogenous cells terminal, unbranched, pale Fig. 54. Pseudocercospora victoriae (CPC 32428). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. Fig. 55. Pseudocercospora victoriae (CPC 32442, ex-type culture). A, B. Leaf spots on upper leaf surface. C–G. Fascicles with conidiogenous cells giving rise to conidia. H. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 77 brown, smooth, tapering to flat-tipped apical loci, proliferating sympodially and percurrently near apex, 18–30 × 3.5–4 µm; scars inconspicuous, 2–2.5 µm diam. Conidia solitary, medium brown, smooth, guttulate, narrowly obclavate, apex subobtuse, base long obconically truncate, straight to slightly curved, (3–)5–7-septate, (45–)60–75(–80) × (3–)4(–5) µm; hila neither thickened nor darkened-refractive, 2–2.5 µm diam. Description in vitro (SNA; CPC 32428): Mycelium subhyaline to pale brown, smooth to slightly verruculose, 1.5–3.5 μm diam. Conidiophores microto macronematous, emerging from hyphae or germinated conidia, pale brown, solitary, smooth or slightly roughened, unbranched, cylindrical, straight to slightly curved in segments, geniculate-sinuous at the apex, multiseptate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 19–193 × 4.5–6 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown, monoor polyblastic, proliferating sympodially, unbranched, conidiogenous loci at the apex and shoulders, protruding and truncate, apex conically truncate, 8.5–19 × 4–7 μm, scars inconspicuous, 1–1.5 μm diam. Conidia solitary, pale brown, guttulate, obclavate, apex obtuse, base obconically truncate, straight to slightly curved, 33.5–67 × 3.5–5.5 μm, 5–10-septate; hila neither thickened nor darkenedrefractive, 1–2 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; erumpent, with moderate to absent aerial mycelium, and roughed, lobate margin. Surface pale mouse grey to white with patches of cinnamon; reverse smoke grey to hazel. Colonies reaching 23 mm diam. Typus: Australia, Victoria, Mount Best, Tin Mine road, from leaf spots on Eucalyptus globoidea (Myrtaceae), 2015, P.W. Crous (holotype CBS H-25062, culture ex-type CPC 32442 = CBS 149429). Additional materials examined: Australia, Victoria, from leaf spots on Eucalyptus radiata (Myrtaceae), 2015, P.W. Crous, HPC 1828 = CBS H-25061, culture CPC 32428 = CBS 149428; New South Wales, Wingecarribee Shire, Paddy’s River, on leaves of E. dives, Mar. 2009, B.A. Summerell, culture CPC 16385; Queensland, on leaves of Eucalyptus sp. on cultivated forest tree, Aug. 2005, A. Carnegie, culture VPRI 40611 = CPC 28347. Notes: Pseudocercospora victoriae (Fig. 1, block 9) is described as a new foliar pathogen of Eucalyptus dives, E. globoidea and E. radiata, being closely related to Ps. subulata (percurrent annellations conspicuous, conidia broadly acicular to obclavatecylindrical, 3–10-septate, 10–100 × 4–7 µm; Braun & Dick 2002; Crous et al. 2019c). Pseudocercospora wingfieldii Crous & Yuan Yuan Chen, sp. nov. MycoBank MB 852290. Fig. 56, 57. Etymology: Named in honour of Michael J. Wingfield, who collected this fungus, and many others, with PWC during a collection trip to Réunion. Leaf spots amphigenous, irregular to sub-circular, 2–3 mm diam, medium brown with raised dark brown border. Mycelium internal, subhyaline, consisting of septate, branched, smooth hyphae, 1.5–2 µm diam. Caespituli fasciculate, hypophyllous, Fig. 56. Pseudocercospora wingfieldii (CPC 26287, ex-type culture). A. Leaf spots on upper and lower leaf surface. B. Close-up of lesion. C–F. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. G. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 78 pale olivaceous on leaves, up to 60 µm diam and 70 µm high. Conidiophores aggregated in dense fascicles arising from the upper cells of a pale brown stroma up to 45 µm diam and 20 µm high; conidiophores pale olivaceous, smooth, 1–2-septate, subcylindrical, straight to slightly curved, unbranched, 15– 35 × 3–5 µm. Conidiogenous cells terminal, unbranched, pale olivaceous, smooth, tapering to flat-tipped apical loci, proliferating sympodially and inconspicuously percurrently near apex, 7–15 × 2.5–4 µm; scars inconspicuous, 1.5–2 µm diam. Conidia solitary, pale olivaceous, smooth, guttulate, subcylindrical, apex obtuse, base truncate, straight to curved, 5–10(–22)-septate, (65–)75–100(–160) × (2.5–)3(–3.5) µm; hila neither thickened nor darkened-refractive, 2–2.5 µm diam. Description in vitro (SNA; CPC 26287): Mycelium subhyaline to pale brown, smooth to slightly verruculose, delicate, uniform in width, 1–3 μm. Conidiophores micronematous, emerging from hyphae or conidia, pale brown, solitary, smooth, unbranched, cylindrical, straight, geniculate-sinuous at the apex, 1-septate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 5–15 × 2.5–4.5 μm. Conidiogenous cells integrated, terminal, pale brown, monoor polyblastic, proliferating sympodially, conidiogenous loci at the apex and shoulders, protruding and conically truncate, apex conically truncate, 5–10.5 × 3–4 μm; scars inconspicuous, 1–1.5 μm diam. Conidia solitary, pale brown, cylindrical to filiform, apex obtuse to subobtuse, base obconically truncate to truncate, slightly curved to curved, 84.5–118(–130) × 2.5–3 μm, 8–11-septate; hila neither thickened nor darkened-refractive, 1–2 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface irregularly folded, with a prominent network of ridges, greyish green and becoming dirty white at margin; reverse fawn to olivaceous. Colonies reaching 25 mm diam. Typus: France, Réunion, S21°5’45.7” E55°33’3.6 from leaf spots on Acacia heterophylla (Fabaceae), 7 Mar. 2015, P.W. Crous & M.J. Wingfield, HPC 257 (holotype CBS H-25048, culture ex-type CPC 26287 = CBS 149369). Notes: See discussion under Ps. acaciicola above (Fig. 1, block 25). Pseudocercospora wingfieldii (Fig. 1, block 26a) is distinct from other species of Pseudocercospora known from this host genus based on its long, narrow, multiseptate, pale olivaceous, subcylindrical conidia. Unresolved species complexes and synonymies Pseudocercospora diplusodonis Meir. Silva et al. [as ‘diplusodonii’], Persoonia 37: 153. 2016. Hosts and distribution: Brazil, on Diplusodon sp. (Lythraceae), culture ex-type COAD 1476. South Korea, on Diospyros lotus (Ebenaceae), cultures CPC 14778 and CPC 14804; on Ilex verticillata (Aquifoliaceae), culture CPC 25429; on Rosa multiflora (Rosaceae), culture CPC 25419. New synonym: Pseudocercospora ershadii M. Bakhshi et al., Fungal Syst. Evol. 6: 105. 2020. Hosts and distribution: Iran, on Diospyros lotus (Ebenaceae), IRAN 16456F – holotype, CCTU 1206 = CBS 136114 – ex-type cultures, additional cultures IRAN 3431C, IRAN 3429C, IRAN 3428C, IRAN 3430C. Note: Although Ps. diplusodonis (Fig. 1, block 43) was described from Diplusodon sp. in Brazil, it is here recorded from several unrelated hosts in South Korea and Iran. Pseudocercospora nogalesii (Urries) U. Braun & M.A. Dick, Australas. Pl. Path. 32: 91. 2003. Hosts and distribution: New Zealand, Chamaecytisis proliferus (Fabaceae), culture CBS 115022; Virgilia divaricara (Fabaceae), culture CBS 122501. Spain (Canary Islands), Cytisus proliferus (original description). Note: Originally described from leaves of Cytisus proliferus (Fabaceae), Canary Islands, both strains in the present study were also isolated from Fabaceae hosts (Fig. 1, block 4). Pseudocercospora norchiensis Crous, Fungal Diversity 26: 172. 2007. Hosts and distribution: Italy, Eucalyptus sp. (Myrtaceae; original description). New Zealand, Eucalyptus sp. (Myrtaceae), culture CBS 120738 = CPC 13049, Rubus sp. (Rosaceae), cultures CBS 114641 and CBS 118413. Note: Originally described from Eucalyptus leaves in Italy, and here reported on an unrelated host family, Rosaceae, in New Zealand (Fig. 1, block 4). Pseudocercospora trichogena Guatim. et al., Persoonia 37: 135. 2016. Fig. 57. Pseudocercospora wingfieldii (CPC 26287, ex-type culture). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 79 Hosts and distribution: Brazil, on leaves of Deparia petersenii (Athyriaceae), culture COAD 1088; on leaves of Macrothelypteris torresiana (Thelypteridaceae), culture ex-type COAD 1087. Note: The two involved host families of the Polypodiopsida (ferns) are allied and pertain to the Polypodiales, Aspleniineae, so that it is not surprising that the two isolates appear phylogenetically identical (similarities 504/504 nt ITS, 198/198 nt actA, 471/471 nt tef1) (Fig. 1, block 11). Pseudocercospora wuchienshiungiae Y.P. Tan et al. [as ‘wuchienshiung’], Index of Australian Fungi 6: 11. 2023. New synonym: Pseudocercospora blackwoodiae Y.P. Tan et al., Persoonia 50: 283. 2023. Notes: Pseudocercospora blackwoodiae was published as a Fungal Planet description sheet in Persoonia with an effective online publication date of 29 June 2023 (Crous et al. 2023), while Ps. wuchienshiungiae (as ‘wuchienshiung’) (Fig. 1, block 14) was published as an Index of Australian Fungi entry with an effective online publication date of 9 May 2023 (Tan & Shivas 2023). The two species were described from Queensland on Persoonia falcata. The metabolically inactive culture as holotype and GenBank numbers of derived sequences are identical between the two publications and therefore the later published Ps. blackwoodiae is reduced to synonymy under the first published name. Pseudocercospora wusaulaniae Y.P. Tan et al. [as ‘wusaulan’], Index of Australian Fungi 6: 12. 2023. New synonym: Pseudocercospora dalyelliae Y.P. Tan et al., Persoonia 50: 285. 2023. Notes: Pseudocercospora dalyelliae was published as a Fungal Planet description sheet in Persoonia with an effective online publication date of 29 June 2023 (Crous et al. 2023), while Ps. wusaulaniae (as ‘wusaulan’) (Fig. 1, block 19) was published as an Index of Australian Fungi entry with an effective online publication date of 9 May 2023 (Tan & Shivas 2023). The two species were described from Queensland on Senna alata. The metabolically inactive culture as holotype and GenBank numbers of derived sequences are identical between the two publications and therefore the later published Ps. dalyelliae is reduced to synonymy under the first published name. Block 4 Pseudocercospora luzardii Furlan. & Dianese, Mycol. Res. 103: 1207. 1999. Host and distribution: Brazil, on leaves of Hancornia speciosa (Apocynaceae), culture ex-epitype COAD 1505 = CPC 25196 and culture CPC 2556 = STE-U 2556. Pseudocercospora convoluta (Crous & Den Breeÿen) U. Braun et al., Stud. Mycol. 87: 312. 2017. Basionym: Passalora convoluta Crous & Den Breeÿen, Fungal Diversity 23: 96. 2006. Host and distribution: Costa Rica, on leaves of Chromolaena odorata (Asteraceae), ex-type CBS 113377. Note on Block 4 : Although phylogenetically similar (ITS: 485/487 nt; additional genes needed) (Fig. 1, block 4), they occur on different host families, and are morphologically distinct (Den Breeÿen et al. 2006, Silva et al. 2016). Block 10 Pseudocercospora balsaminae (Syd.) Deighton, Mycol. Pap. 140: 139. 1976. Basionym: Cercoseptoria balsaminae Syd., Ann. Mycol. 33: 69. 1935. Hosts and distribution: India, Uttar Pradesh, on leaves of Impatiens balsamina (Balsaminaceae) (type). South Korea, on leaves of Impatiens textorii, cultures CBS 131882 = CPC 10044 and CBS 132020 = CPC 10699. Pseudocercospora boehmeriigena U. Braun, in Braun & Mel’nik, Trudy Botanicheskogo Instituta im. V.L. Komarova 20: 42. 1997. Replaced synonym: Cercospora boehmeriae Peck, Ann. Rep. N.Y. St. Mus. nat. Hist. 34: 48. 1883. [1881]. Host and distribution: Brazil, on leaves of Boehmeria nivea (Urticaceae), culture ex-epitype COAD 1562 = CPC 25243. New synonym: Cercospora fukuii W. Yamam. [as ‘fukui’], Jour. Soc. Trop. Agric. Formosa 6: 601. 1934. Pseudocercospora fukuii (W. Yamam.) W.H. Hsieh & Goh, Trans. Mycol. Soc. R.O.C. 2: 115. 1987. Host and distribution: Japan, Tokyo, Akiruno, Itsukaichi, on leaves of Boehmeria nivea var. concolor f. nipononivea (Urticaceae), culture ex-epitype MAFF 238121 = MUCC 1297. Pseudocercospora crocea Crous et al., Stud. Mycol. 75: 83. 2012. [2013]. Host and distribution: South Korea, on leaves of Pilea hamaoi (Urticaceae), culture ex-type CBS 126004 = CPC 11668. Pseudocercospora cyatheicola Crous & R.G. Shivas [as ‘cyathicola’], Crous et al., Persoonia 26: 121. 2011. Host and distribution: Australia, on fronds of Cyathea australis (Cyatheaceae), culture ex-type CBS 129520 = CPC 17047. Pseudocercospora dianellae U. Braun & C.F. Hill, Australas. Pl. Path. 32: 89. 2003. Hosts and distribution: New Zealand, on leaves of Dianella nigra (Asphodelaceae) (type); on leaves of Dianella caerulea, culture CBS 117746. Pseudocercospora farfugii C. Nakash., I. Araki & Ai Ito, in Chen et al., Stud. Mycol. 101: 540. 2022. Host and distribution: Japan, on leaves of Farfugium japonicum (Asteraceae), culture ex-type MUCC 978. Notes: The present species formed a clade with Ps. textariae on Tectaria harlandii (Tectariaceae). However, the sequencing data of the latter species is only known as an rDNA ITS, and its accurate phylogenetic position is unclear. The morphological characteristics are also different from each other in the size and shape of conidia and conidiogenesis. See Chen et al. (2022) and Kirschner & Wang (2015). Pseudocercospora fatouae Goh & W.H. Hsieh, in Hsieh & Goh, Cercospora and Similar Fungi from Taiwan (Taipei): 236. 1990. Hosts and distribution: Taiwan, on leaves of Fatoua villosa (Moraceae) (type). South Korea, on leaves of Fatoua villosa, culture CPC 11648. Pseudocercospora hachijokibushi C. Nakash., H. Horie & Tak. Kobay., Mycoscience 45(1): 50. 2004. Host and distribution: Japan, on leaves of Stachyurus praecox var. matsuzakii (Stachyuraceae), culture ex-type MAFF 238479 = MUCC 1337. Pseudocercospora humulicola Crous et al., Stud. Mycol. 75: 89. 2012. [2013]. Host and distribution: South Korea, on leaves of Humulus scandens (Cannabaceae), culture ex-type CBS 131585 = CPC 11358. Pseudocercospora lygodiicola Y.L. Guo & U. Braun, in Braun, © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 80 Nakashima & Crous, IMA Fungus 4(2): 317. 2013. Hosts and distribution: China, on leaves of Lygodium japonicum (Lygodiaceae) (type). Brazil, on fronds of Lygodium volubile, culture COAD 1745 = CPC 25755. Pseudocercospora macarangae (Syd. & P. Syd.) Deighton, Mycol. Pap. 140: 47. 1976. Basionym: Cercospora macarangae Syd. & P. Syd., Annls mycol. 12(6): 575. 1914. Hosts and distribution: Philippines, on leaves of Macaranga tanarius (Euphorbiaceae) (type). Laos, on leaves of Macaranga denticulata, culture P564. Pseudocercospora plectranthi G.C. Hunter et al., Stud. Mycol. 75: 96. 2012. [2013] Host and distribution: South Korea, from leaves of Plectranthus sp. (Lamiaceae), culture ex-type CBS 131586 = CPC 11462. Pseudocercospora profusa (Syd. & P. Syd.) Deighton, Trans. Brit. Mycol. Soc. 88: 388. 1987. Basionym: Cercospora profusa Syd. & P. Syd., Ann. Mycol. 7: 175. 1909. Host and distribution: South Korea, on leaves of Acalypha australis (Euphorbiaceae), culture ex-epitype CPC 10713 and culture CBS 132306 = CPC 10055. Pseudocercospora pteridicola U. Braun & Y.L. Guo, in Braun & Mel’nik, Trudy Botanicheskogo Instituta im. V.L. Komarova 20: 84. 1997. Hosts and distribution: China, Sichuan, on leaves of Pteris vittata (Pteridaceae) (type). Taiwan, on leaves of Pteris semipinnata, specimen R. Kirschner 4166. Pseudocercospora pteridophytophila Goh & W.H. Hsieh, Trans. Mycol. Soc. Rep. China 4(2–3): 25–38. 1989. Host and distribution: Taiwan, on leaves of Cyclosorus acuminatus (Thelypteridaceae) (type); on leaves of Cyclosorus parasiticus, voucher TNM 3602. Pseudocercospora rhabdothamni U. Braun & C.F. Hill, Australas. Pl. Path. 33: 489. 2004. Host and distribution: New Zealand, on leaves of Rhabdothamnus solandri (Gesneriaceae), culture ex-type CBS 114872 = ICMP 15289. Pseudocercospora rumohrae Goh & W.H. Hsieh, Trans. Mycol. Soc. Rep. China 4: 29. 1989. Hosts and distribution: Taiwan, on leaves of Rumohra adiantiformis (Dryopteridaceae) (type). New Zealand, on leaves of Ptisana (=Marattia) salicina (Marattiaceae), culture CBS 117747. Pseudocercospora sp. 016 Host and distribution: New Zealand, on Rubus × barkeri (Rosaceae), culture CPC 14813. Pseudocercospora tectariae R. Kirschner, in Kirschner & Wang, Mycol. Progr. 14(65): 7. 2015. Host and distribution: Taiwan, on living sterile fronds of Tectaria harlandii (Tectariaceae), from type and from authentic culture BCRC FU30377. Notes on Block 10 : Although similar in DNA phylogeny (Fig. 1, block 10), these taxa are morphologically different, and mostly occur on unrelated host families, some of which are restricted Fig. 58. Pseudocercospora sp. 069 (CPC 30540). A. Leaf spots on lower and upper leaf surface. B. Close-up of lesion. C–F. Fascicles with conidiophores and conidiogenous cells giving rise to conidia. G. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 81 in their distribution, suggesting that the genes used here lack the resolution to properly distinguish these species. See also Ps. farfugii. Although Ps. fatouae, Ps. macarangae, Ps. pteridicola, Ps. pteridophytophila and Ps. tectariae are represented by only an ITS sequence each in the current dataset, their phylogenetic association stays the same between the IQ-TREE and RAxML analyses (Suppl. Table S3). Block 20c - Pseudocercospora pruni-persicicola complex Pseudocercospora sp. 069 on Buxus megistophylla in China. Figs 58, 59. (CBS H-25055): Leaf spots amphigenous, sub-circular to circular, 4–10 mm diam, medium brown with raised border and redbrown margin. Mycelium internal, pale brown, consisting of septate, branched, smooth hyphae, 2.5–3 µm diam. Caespituli fasciculate, chiefly hypophyllous, brown on leaves, up to 100 µm diam and 120 µm high. Conidiophores in dense fascicles arising from the upper cells of a brown stroma up to 65 µm diam and 40 µm high; conidiophores brown, verrucose, 1–2-septate, subcylindrical, straight to variously curved, unbranched, 20–35 × 4–5 µm. Conidiogenous cells terminal, pale to medium brown, verruculose, proliferating percurrently near apex, with rough annellations, 10–15 × 4–5 µm; scars inconspicuous, 2.5–3 µm diam. Conidia solitary, pale brown, verruculose with longitudinal striations, guttulate, narrowly obclavate, apex subobtuse, base obclavate, straight to slightly curved, (2–)3–5(–8)-septate, (27–) 45–60(–80) × (2.5–)3 µm; hila neither thickened nor darkened, 2 µm diam. Description in vitro (SNA; CPC 30540): Mycelium subhyaline to pale brown, smooth to slightly verruculose, delicate, 1.5–3.5 μm diam. Conidiophores microto macronematous, emerging from hyphae or conidia, pale brown, solitary, smooth or slightly roughened, unbranched, cylindrical, straight to slightly curved in segments, geniculate-sinuous at the apex, 1–2-septate, sometimes reduced to conidiogenous cells, straight, unbranched, holoblastic, 12–26.5 × 3.5–5 μm. Conidiogenous cells integrated, terminal or intercalary, pale brown, monoor polyblastic, proliferating sympodially, conidiogenous loci at the apex and shoulders, protruding, apex conically truncate, 5–16.5 ×3–4 μm; scars inconspicuous, 1–1.5 μm diam. Conidia solitary, pale brown, obclavate, apex obtuse to subobtuse, base long obconically truncate, straight to slightly curved, 25–75 × 2.5–3 μm, (1–)3–8-septate; hila neither thickened not darkened, 1–1.5 μm diam. Culture characteristics: Colonies after 3 wk at 25 °C in the dark on MEA; surface folded with a prominent network of ridges, surface smoke grey with patches of lavender blue; reverse mouse grey becoming olivaceous at the margin. Colonies reaching 28 mm diam. Material examined: China, Guizhou, from leaf spots on Buxus megistophylla (Buxaceae), 2015, J.J. Gan, HPC 1113 = GJJ 160210-01 = CBS H-25055, culture CPC 30540 = CBS 149423. Species involved in this complex based on Fig. 1: Pseudocercospora eupatorii-formosani U. Braun & Bagyan., Sydowia 51: 8. 1999. Synonyms: Cercospora eupatorii-formosani Sawada, Rep. Dept. Agric. Gov. Res. Inst. Formosa 86: 169. 1943, nom. inval. (Art. 39.1). Pseudocercospora eupatorii-formosani J.M. Yen [as ‘(Sawada) J.M. Yen’], Gard. Bull. Singapore 33: 175. 1980, nom. inval. (Art. 39.1). Pseudocercospora eupatorii-formosanae J.M. Yen ex Y.L. Guo & W.H. Hsieh [as ‘(Sawada ex Y.L. Guo & W.H. Hsieh) J.M. Yen’], Mycosystema Monographicum Series 2: The genus Pseudocercospora in China: 67. 1995, nom. inval. (Art. 40.1, Art. 40.4). Host and distribution: Laos, on Chromolaena odorata (Asteraceae), cultures CPC 17319 = NOUL p.9 and CPC 19195 = LC 0395. South Africa, on leaves of Solanum mauritianum (Solanaceae), culture CPC 25040. Taiwan, on leaves of Eupatorium sp. (Asteraceae), culture ex-epitype NCHUPP_ L1606, and from unknown host, culture CPC 20054. Unknown, no collection information, cultures CBS 112825 = C 439 = CPC 4087 = STE-U 4087 and CBS 115494 = C 444 = CPC 4088 = STE-U 4088. USA, unknown host, culture CBS 115493 = C 448 (a) = CPC 4073 = STE-U 4073. Pseudocercospora ginkgoana R. Kirschner, Pl. Pathol. Quarant. 8: 12. 2018. Host and distribution: Taiwan, on dead margin of living leaves of Ginkgo biloba (Ginkgoaceae), culture R. Kirschner 3563. Pseudocercospora pruni-persicicola (J.M. Yen) J.M. Yen, Bull. Soc. Mycol. Fr. 97: 94. 1981. Synonym: Cercospora pruni-persicicola J.M. Yen, Rev. Mycol. (Paris): 63. 1978. Host and distribution: South Korea, on leaves of Prunus avium (Rosaceae), culture KACC 47019. Fig. 59. Pseudocercospora sp. 069 (CPC 30540). A. Colony on MEA. B. Sporulation on SNA. C, D. Conidiogenous cells giving rise to conidia. E. Conidia. Scale bars = 10 μm. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 82 Pseudocercospora sp. 069 Host and distribution: China, on leaves of Buxus megistophylla (Buxaceae), culture CPC 30540 = CBS 149423. Notes on Block 20c: Because there are presently no records of Pseudocercospora spp. known from Buxus, the Chinese material was initially assumed to represent a new species (Fig. 1, block 20c). However, based on the phylogenetic data, the Pseudocercospora on Chinese Buxus pertains to a complex (clade; Block 20c) within Block 20 comprised of sequences retrieved from several species, including Ps. pruni-persicicola, Ps. eupatorii-formosani, and Ps. ginkgoana, with insufficient resolution based on the currently applied markers. The involvement of a single plurivorous species cannot be excluded with absolute certainty, but it is highly unlikely that we are dealing with a single species. The hosts involved in this complex are completely unrelated, ranging from Ginkgo (gymnosperms) to Chromolaena odorata (Compositae, i.e., advanced dicots). Furthermore, the morphology of the taxa involved is strongly different from each other. The Pseudocercospora on Buxus is, in contrast to the other involved species, characterised by having internal mycelium in vivo, distinctly verruculose conidiophores and conidia, and percurrently proliferating conidiogenous cells with conspicuous rough annellations. This type of proliferation and annellation is not present in all other species of this complex, and there are additional significant morphological differences in the formation of stromata, external mycelium, and in the dimension and shape of the conidiophores and conidia. Pseudocercospora pruni-persicicola is currently represented by a single isolate with only an ITS sequence in the dataset and its phylogenetic position varies depending on the phylogenetic analysis used, namely clustering among isolates of Ps. eupatoriiformosani in block 20 (Fig. 1, IQ-TREE analysis) or sister to Ps. sphendamnophila (see Figshare; equivalent of clustering in Fig. 1, block 25) in the RAxML analysis (Suppl. Table S3). The ITS sequence of Ps. pruni-persicicola is 99 % (identities 466/469 and 473/474, both including one gap) similar to the ITS sequences of Ps. eupatorii-formosani and Ps. sphendamnophila. Pseudocercospora ginkgoana is also represented only by an ITS sequence, but it clusters with the same association between the IQ-TREE and RAxML analyses (Suppl. Table S3). The Pseudocercospora pruni-persicicola complex requires the application of additional makers for a better resolution at species level, and further pathogenicity studies are needed to confirm the phylogenetic-taxonomic all species involved in this clade (see Choi et al. 2014). Block 26a Pseudocercospora chionanthi-retusi Goh & W.H. Hsieh, in Hsieh & Goh, Cercospora and Similar Fungi from Taiwan (Taipei): 249. 1990. Host and distribution: Taiwan, on leaves of Chionanthus retusus var. serrulatus (Oleaceae), culture ex-epitype NCHUPP L1605. Pseudocercospora marginalis G.C. Hunter et al., Stud. Mycol. 75: 92. 2012. [2013]. Host and distribution: South Korea, on leaves of Fraxinus rhynchophylla (Oleaceae), ex-type CBS 131582 = CPC 12497. Note on Block 26a: Although phylogenetically similar (ITS: 475/477 nt; actA: 220/220 nt; tef1: 311/313 nt) (Fig. 1, block 26), the host genera and fungal morphology are distinct. Block 28 – Pseudocercospora atromarginalis complex Pseudocercospora atromarginalis (G.F. Atk.) Deighton, Mycol. Pap. 140: 139. 1976. Basionym: Cercospora atromarginalis G.F. Atk., J. Elisha Mitchell scient. Soc. 8: 59. 1892. Synonyms: See Braun (2017). Hosts and distribution: Brazil, on leaves of Solanum americanum (Solanaceae), culture COAD 1975 = CPC 25230. Iran, on leaves of Solanum nigrum (Solanaceae), cultures CBS 136112 = CCTU 1056, CCTU 1052, CCTU 1091 and CCTU 1193. New Zealand, on Solanum sp. (Solanaceae), culture CBS 114640. South Korea, on leaves of Solanum nigrum (Solanaceae), cultures CPC 14649 and CBS 132010 = CPC 11372. USA, on Solanum nigrum (Solanaceae) (type). Pseudocercospora chengtuensis (F.L. Tai) Deighton, Mycol. Pap. 140: 141. 1976. Basionym: Cercospora chengtuensis F.L. Tai, Lloydia 11: 40. 1948. Hosts and distribution: China, on Lycium chinense (Solanaceae) (type). South Korea, on Lycium chinense (Solanaceae), culture CBS 131924 = CPC 10696. Pseudocercospora cruenta (Sacc.) Deighton, Mycol. Pap. 140: 142. 1976. Basionym: Cercospora cruenta Sacc., Michelia 2(no. 6): 149. 1880. Hosts and distribution: USA, on leaves of Phaseolus (Fabaceae) (type). Trinidad and Tobago, on Vigna sp. (Leguminosae), culture CBS 132021 = CPC 10846 (block 28). Pseudocercospora dingleyae U. Braun & C.F. Hill [as ‘dingleyi’], Mycol. Progr. 1(1): 23. 2002. Competing homonym: non Pseudocercospora haloragis (Hansf.) U. Braun [as ‘haloragidis’], Monogr. Cercosporella, Ramularia Allied Genera (Phytopath. Hyphom.) 1: 109. 1995. Replaced synonym: Cercospora haloragis Dingley [as ‘haloragi’], N.Z. Jl agric. Res. 8(4): 913. 1965. Host and distribution: New Zealand, on Haloragis erecta (Haloragaceae), type as well as culture CBS 114645. Pseudocercospora fuligena (Roldan) Deighton, Mycol. Pap. 140: 144. 1976. Basionym: Cercospora fuligena Roldan, Philipp. J. Sci. 66: 8. 1938. Hosts and distribution: Philippines, on leaves of Solanum lycopersicum (Solanaceae) (type). Thailand, on Solanum lycopersicum, culture CBS 132017 = CPC 12296. Taiwan, on Solanum lycopersicum, culture CBS 109729. USA, on Solanum lycopersicum, culture PF001. Pseudocercospora mangiferae Tamakaew & Cheew., Chiang Mai J. Sci. 49(3): 687. 2022. Host and distribution: Thailand, on leaves of Mangifera indica (Anacardiaceae), culture SDBR-CMU401 and culture ex-type SDBR-CMU403. Pseudocercospora musarum Crous & Yuan Yuan Chen, sp. nov. Present study. Host and distribution: Malaysia, on living leaves of Musa sp. (Musaceae), culture ex-type CBS 149516 = CPC 32815. Pseudocercospora rauvolfiicola A. Singh, P.N. Singh & N.K. Dubey, Phytotaxa 545(2): 131. 2022. Host and distribution: India, on living leaves of Rauvolfia serpentina (Apocynaceae), culture ex-type NFCCI 4586. Pseudocercospora solanicola Arch. Singh, et al., Turk. J. Bot. 46: 508. 2022. Host and distribution: India, on leaves of Solanum nigrum (Solanaceae), culture ex-type NFCCI 5015. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 83 Pseudocercospora sp. 059 Hosts and distribution: Australia, on Santalum acuminatum (Santalaceae), culture CBS 149420 = CPC 28327; on Santalum lanceolatum (Santalaceae), culture CBS 149417 = CPC 27618. Pseudocercospora sp. 060 Hosts and distribution: Australia, on Solanum sp. (Solanaceae), culture CPC 28350. New Zealand, on Gunnera tinctoria (Gunneraceae), culture CBS 115122. Unknown, unknown host, culture CBS 114642. Pseudocercospora tabernaemontanae (Syd. & P. Syd.) Deighton, Mycol. Pap. 140: 154. 1976. Basionyn: Cercospora tabernaemontanae Syd. & P. Syd., Philipp. J. Sci., C, Bot. 8(5): 507. 1913. Hosts and distribution: Laos, on leaves of Tabernaemontana coronaria (Apocynaceae), culture CPC 19198 (ex-epitype). Philippines, on leaves of Tabernaemontana pandacaqui (Apocynaceae) (type). Notes on Block 28 : Wang et al. (1995) and Braun (2017) proposed a synonymy between Ps. atromarginalis and Ps. fuligena based on overlapping host ranges, lack of distinguishing morphological features and lack of sufficient numbers of nucleotide differences to support the two species as distinct. However, Braun (2017) did state that the designation of epitypes for both species is required to finally clarify the taxonomic status of these species. Therefore the present study treats them as separate pending the collection and sequencing of material that can be designated as epitypes for both species. Currently there are only an ITS and LSU sequence available for Ps. solanicola (GenBank MZ474952 and MZ474953, respectively), of which the ITS sequence was used in the phylogenetic analyses. Interestingly, this sequence clustered with Ps. hakeae in the IQ-TREE analysis (Fig. 1, block 24), but intermingled with strains in the Ps. atromarginalis complex in the RAxML analysis (see Figshare; equivalent of clustering in Fig. 1, block 28). The ITS sequence differs by four gaps and one substitution (484/489 (99 %) identity; the region containing the three gaps was at the very end of the ITS sequence and excluded from the analyses as they appear to be consistent with basecalling problems) from that of Ps. hakeae culture CBS 112226 (GenBank GU269784) and with a single gap (477/478 (99 %) identity) from that of Ps. atromarginalis culture CBS 136112 (GenBank KM452851). Block 38 Pseudocercospora zelkovae X.J. Liu & Y.L. Guo [as ‘zelkowae’], Acta Mycol. Sin. 12: 33. 1993. Hosts and distribution: Japan, on leaves of Zelkova serrata (Ulmaceae), culture ex-neotype MAFF 410008 = MUCC 1398 and culture MAFF 238237 = MUCC 872. South Korea, on leaves of Zelkova serrata, cultures CBS 132106 = CPC 14484 and CBS 132118 = CPC 14717. Pseudocercospora riachueli var. horiana (Togashi & Katsuki) U. Braun & Crous, in Crous & Braun, CBS Diversity Ser. (Utrecht) 1: 354. 2003. Host and distribution: Japan, on leaves of Parthenocissus tricuspidate (Vitaceae), culture ex-epitype MUCC 2141. Notes on Block 38: Although occurring on unrelated hosts, the two species appear phylogenetically similar (ITS: 478/478 nt; actA: 219/219 nt; tef1: 307/308 nt; rpb2: 673/673 nt) (Fig. 1, block 38). The ex-type culture was located in the clade of Ps. zelkovae (Fig. 1, block 38). The epitypified specimen of Ps. riachueli var. horiana was collected from the fringe of woods near a populated area, which was planted with Zelkova trees. The morphological characteristics of both specimens are similar. Pseudocercospora zelkovae might temporarily colonise Parthenocissus. More detailed examinations with fresh samples are required. Block 51 Pseudocercospora amelanchieris C. Nakash. et al., Mycol. Progr. 15: 1097. 2016. Host and distribution: Japan, on leaves of Amelanchier canadensis (Rosaceae), culture ex-type MAFF 237782 = MUCC 885. Pseudocercospora corylopsidis (Togashi & Katsuki) C. Nakash. & Tak. Kobay., Mycoscience 40: 270. 1999. Basionym: Cercospora corylopsidis Togashi & Katsuki, Bot. Mag., Tokyo 65: 20. 1952. Host and distribution: Japan, on leaves of Corylopsis spicata (Hamamelidaceae), culture ex-epitype MAFF 237795 = MUCC 908. Pseudocercospora chibaensis Tak. Kobay. & Nagash., Trans. Mycol. Soc. Japan 32: 328. 1991. Host and distribution: Japan, on leaves of Nyssa sylvatica (Cornaceae), culture ex-epitype MUCC 1670. Pseudocercospora cotoneastri (Katsuki & Tak. Kobay.) Deighton [as ‘cotoneasteris’], Trans. Brit. Mycol. Soc. 88: 389. 1987. Basionym: Cercospora cotoneastri Katsuki & Tak. Kobay. [as ‘cotoneasteris’], Trans. Mycol. Soc. Japan 17: 276. 1976. Host and distribution: Japan, on leaves of Cotoneaster dammeri (Rosaceae), culture ex-type MAFF 410089 = MUCC 1416. Pseudocercospora daphniphylli (Katsuki & Tak. Kobay.) Deighton, Trans. Brit. Mycol. Soc. 88(3): 389. 1987. Basionym: Cercospora daphniphylli Katsuki & Tak. Kobay., Trans. Mycol. Soc. Japan 23: 44. 1982. Host and distribution: Japan, on leaves of Daphniphyllum macropodum (Daphniphyllaceae), culture ex-type MAFF 410009 = MUCC 1399. Pseudocercospora emmoticola Meir. Silva, R.W. Barreto & Crous [as ‘emmotunicola’], in Silva, Barreto, Pereira, Freitas, Groenewald & Crous, Persoonia 37: 153. 2016. Host and distribution: Brazil, on leaves of Emmotum nitens (Icacinaceae), culture ex-type COAD 1491 = CPC 25187. Pseudocercospora eriodendri (Racib.) U. Braun, Nova Hedwigia 55(1–2): 217. 1992. Basionym: Ramularia eriodendri Racib., Parasit. Alg. Pilze Java’s (Jakarta) 1: 35. 1900. Host and distribution: Java, on Eriodendron anfractuosum (Bombacaceae) (lectotype). Somalia, on Ceiba pentandra (Bombacaceae), authentic culture CBS 130.32. Pseudocercospora formosana (W. Yamam.) Deighton, Mycol. Pap. 140: 144. 1976. Basionym: Cercospora formosana W. Yamam., Journal of the Society of Tropical Agriculture, Formosa 6: 600. 1934. Host and distribution: Japan, on Lantana camara (Verbenaceae), culture ex-epitype MAFF 238239 = MUCC 879 and culture MUCC 855. Malaysia, on Lantana sp., culture MUCC 2612. Pseudocercospora forsythiae (Katsuki & Tak. Kobay.) Deighton, Trans. Brit. Mycol. Soc. 88: 389. 1987. Basionym: Cercospora forsythiae Katsuki & Tak. Kobay., Trans. Mycol. Soc. Japan 17: 273. 1976. Host and distribution: Japan, on leaves of Forsythia suspensa (Oleaceae), culture ex-type MAFF 410087 = MUCC 1414. © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] J.Z. Groenewald et al. 84 Pseudocercospora hiratsukana (Togashi & Katsuki) Deighton, Mycol. Pap. 140: 34. 1976. Basionym: Cercospora hiratsukana Togashi & Katsuki, J. Jap. Bot. 28: 286. 1953. Host and distribution: Japan, on leaves of Dioscorea tokoro (Dioscoreaceae), culture ex-epitype MAFF 238300 = MUCC 1105. Pseudocercospora izuohshimensis C. Nakash. et al., Mycoscience 45: 49. 2004. Host and distribution: Japan, on leaves of Helwingia japonica (Helwingiaceae), culture ex-type MAFF 238478 = MUCC 1336. Pseudocercospora mangifericola R.G. Shivas, A.J. Young & Grice, Persoonia 23: 197. 2009. Host and distribution: Australia, on leaves of Mangifera indica (Anacardiaceae), culture ex-type BRIP 52776b. Pseudocercospora mombin (Petr. & Cif.) Deighton, Mycol. Pap. 140: 148. 1976. Basionym: Cercospora mombin Petr. & Cif., Annls mycol. 30(3/4): 322. 1932. Hosts and distribution: Dominican Republic, on living leaves of Spondias mombin (Anacardiaceae) (lectotype). Thailand, on Spondias pinnata, culture P625. Pseudocercospora pallida (Ellis & Everh.) H.D. Shin & U. Braun, Mycotaxon 74(1): 114. 2000. Basionym: Cercospora pallida Ellis & Everh., J. Mycol. 3(2): 21. 1887. Hosts and distribution: USA, on living leaves of Tecoma radicans (Bignoniaceae) (type). South Korea, on Campsis grandiflora (Bignoniaceae), culture CBS 131889. Pseudocercospora perae Meir. Silva, R.W. Barreto & Crous, in Silva, Barreto, Pereira, Freitas, Groenewald & Crous, Persoonia 37: 158. 2016. Host and distribution: Brazil, on leaves of Pera glabrata (Euphorbiaceae), culture ex-type COAD 1465 = CPC 25171. Pseudocercospora prunicola (Ellis & Everh.) U. Braun, in Braun & Mel’nik, Trudy Botanicheskogo Instituta im. V.L. Komarova 20: 82 (1997) Basionym: Cercospora prunicola Ellis & Everh., J. Mycol. 3: 17. 1887. Hosts and distribution: USA, on leaves of Prunus americana (Rosaceae) (type). South Korea, on leaves of Prunus yedoensis, cultures CBS 132107 = CPC 14511, CPC 14792, and on leaves of Prunus sp., culture CPC 25431. Pseudocercospora pruni-yedoensis Sawada ex Goh & W.H. Hsieh, in Hsieh & Goh, Cercospora and Similar Fungi from Taiwan (Taipei): 282. 1990. Hosts and distribution: Taiwan, on leaves of Prunus yedoensis (Rosaceae) (type); on Prunus campanulate, voucher CEO13. Pseudocercospora pseudocydoniae sp. nov. (see elsewhere in this publication) Host and distribution: South Korea, on leaf spots on Chaenomeles lagenaria (Rosaceae), culture ex-type CBS 149392 = CPC 14665. Pseudocercospora puerariicola (W. Yamam.) Deighton, Mycol. Pap. 140: 151. 1976. Basionym: Cercospora puerariicola W. Yamam. [as ‘pueraricola’], Trans. Sapporo nat. Hist. Soc. 13(2–3): 142. 1934. Hosts and distribution: Japan, on living leaves of Pueraria thunbergiana (Leguminosae) (original description). Taiwan, on Pueraria montana, culture BCRC FU30030 (block 51). Pseudocercospora pyracanthae (Katsuki) C. Nakash. & Tak. Kobay., Ann. phytopath. Soc. Japan 63(4): 313. 1997. Basionym: Cercospora pyracanthae Katsuki [as ‘pyrecanthae’], Bull. agr. impr. Sect. Econ. Dept. Fukuoka Prefecture Japan 1: 19. 1949. Host and distribution: Japan, on Pyracantha angustifolia (Rosaceae), culture ex-epitype MAFF 237140 = MUCC 1226. Pseudocercospora rhododendrigena C. Nakash., in Nakashima, Oetari, Kanti, Saraswati, Widyastuti & Ando, Mycosphere 1(6): 318. 2011. Host and distribution: Indonesia, on leaves of Rhododendron sinense (Ericaceae), culture ex-type NBRC 105400 = BTCC F-62. Pseudocercospora sp. 039 Host and distribution: Thailand, on Prunus ceraseidos (Rosaceae), culture CPC 21670. Pseudocercospora sp. 040 Host and distribution: Brazil, unknown, culture CPC 18501. Pseudocercospora sp. 041 Host and distribution: China, on fruit of Citrus grandis (Rutaceae), culture CPC 24510. Pseudocercospora stephanandrae (Tak. Kobay. & H. Horie) C. Nakash. & Tak. Kobay., Mycoscience 41(1): 27 (2000) Basionym: Cercospora stephanandrae Tak. Kobay. & H. Horie, in Kobayashi, Horie & Sasaki, Trans. Mycol. Soc. Japan 20(3): 331. 1979. Host and distribution: Japan, on leaves of Stephanandra incisa (Rosaceae), culture ex-epitype MAFF 237799 = MUCC 914. Pseudocercospora stranvaesiae (Katsuki & Tak. Kobay.) Deighton, Trans. Brit. Mycol. Soc. 88: 389. 1987. Basionym: Cercospora stranvaesiae Katsuki & Tak. Kobay. [as ‘stranvasiae’], Trans. Mycol. Soc. Japan 17: 278. 1976. Host and distribution: Japan, on leaves of Photinia davidiana (Rosaceae), culture ex-type MAFF 410090 = MUCC 1417. Pseudocercospora tamarindi Goonas. & K.D. Hyde, in Liu et al., Fungal Diversity: 10.1007/s13225-015-0324-y, [131]. 2015. Host and distribution: Thailand, on living leaves of Tamarindus indica (Leguminosae), culture ex-type MFLUCC 14-0805. Pseudocercospora togashiana (S. Ito & Tak. Kobay.) C. Nakash. & Tak. Kobay., Stud. Mycol. 101: 548. 2022. Basionym: Mycosphaerella togashiana S. Ito & Tak. Kobay., Bull. Govt Forest Exp. Stn Meguro 59: 23. 1953. Host and distribution: Japan, on leaves of Populus simonii (Salicaceae), culture ex-type MAFF 410006. Pseudocercospora vassobiae Meir. Silva, R.W. Barreto & Crous, in Silva, Barreto, Pereira, Freitas, Groenewald & Crous, Persoonia 37: 167. 2016. Host and distribution: Brazil, on leaves of Vassobia breviflora (Solanaceae), culture ex-type COAD 1572 = CPC 25251. Pseudocercospora weigelae (Ellis & Everh.) Deighton [as ‘weigeliae’], Trans. Brit. Mycol. Soc. 88: 389. 1987. Basionym: Cercospora weigelae Ellis & Everh. [as ‘weigeliae’], Proc. Acad. Nat. Sci. Philad. 45: 170. 1893. Hosts and distribution: USA, on leaves of Weigela sp. (Caprifoliaceae) (type). Japan, on leaves of Weigela coraeensis, culture MAFF 237794 = MUCC 899. Notes on Block 51: Similar to the other clades containing diverse species, this clade (Fig. 1, block 51) is also unresolved. Two isolates deposited as Ps. formosana do not cluster with the ex-epitype culture and could represent an additional species on Lantana in Asia (ITS: 474–477/477 nt; actA: 189/190 and 192/198 nt; tef1: 302–305/305 nt; rpb2: 636–637/673 nt). Besides unrelated host families and species, there are also significant morphological differences between the species involved. Species belonging this block are share Rosaceae plants as a common ancestral host plant. The Rosaceae hosts, Ameranchier, Chaenomeles, Cotoneaster, Nyssa, Photinia, © 2024 Westerdijk Fungal Biodiversity Institute Editor-in-Chief Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail:[email protected] Species diversity in Pseudocercospora 85 Prunus, Pyracantha, and Stephanandra, are diffuse on this clade and species from various plants are mixed. The distribution of these Pseudocercospora species is mainly in Asian countries. The phylogeny of these species suggest that the genes used here lack the resolution to properly distinguish these species. They may be showing the host expansion and speciation at the terminal clade on Rosaceae hosts. Pseudocercospora mangifericola is only represented by an ITS sequence in the present analyses and either clusters in block 51 in the IQ-TREE analysis (Fig. 1; sister to Ps. pruni-yedoensis) or sister to Ps. kenyirana (see Figshare; equivalent of clustering in Fig. 1, block 21) in the RAxML analysis (Suppl. Table S3). The ITS sequence of Ps. mangifericola is 99 % (identities 467/470 and 468/470, both including one gap) similar to the ITS sequences of Ps. pruni-yedoensis and Ps. kenyirana. Likewise for Ps. mombin, which is sister on long branches to Ps. pyracanthae (Fig. 1, block 51) or clusters among strains of Ps. flavomarginata (see Figshare; equivalent of clustering in Fig. 1, block 22) in the RAxML analysis (Suppl. Table S3). The ITS sequence of Ps. mombin is 95 % (identities 392/414 and 393/415, including 21 and 22 gaps, respectively) similar to the ITS sequences of Ps. pyracanthae and Ps. flavomarginata. However, Ps. eriodendri and Ps. tamarindi are also represented by only an ITS sequence each but have a similar clustering in both the IQ-TREE and RAxML analyses (Suppl. Table S3). Although Ps. pruni-yedoensis is also represented by only an ITS sequence, in both analyses it clusters with species in block 51: sister to Ps. rhododendrigena in the IQ-TREE analysis and sister to Ps. vassobiae in the RAxML analysis. The ITS sequence of Ps. pruni-yedoensis is 99 % (identity 472/473, including one gap) or 100 % (identity 473/473, including one gap) similar to the ITS sequences of Ps. rhododendrigena and Ps. vassobiae. Also see the notes under Ps. puerariicola and Ps. rhododendrigena. Problematic application of names During the present study, numerous isolates were encountered for which it was impossible to unequivocally assign a species identification. This was due to several reasons, e.g., i) multiple species known from the substrate and/or locality but sequence data for the species are not available from (ex-)type or authentic material, ii) the loci commonly used for phylogenetic reconstruction in the genus do not provide sufficient resolution, iii) availability of limited sequence data, e.g. only ITS, for many species, iv) different interpretations of the application of a name by authors, and v) cultures proved to be sterile or the fungus could not be located on the original substrate material. For these reasons, we have chosen not to formally name a large number of the obtained cultures in this study (as Pseudocercospora sp. in Suppl. Table S1). A few problematic species for which a name could be or was applied to a culture or specimen in past studies or for which a culture was supplied under the given name in the present study, but of which the derived sequences show them not to be conspecific in the present study, are discussed below. Pseudocercospora catappae (Henn.) Y.L. Guo & X.J. Liu, Mycosystema 2: 230. 1989. Basionym: Cercospora catappae Henn., Bot. Jb. 34: 56. 1904. Notes: The two cultures included in the present study (Suppl. Table S1) (Fig. 1, blocks 34 and 48) both were isolated from Terminalia catappa in Japan and were published separately by Crous et al. (2013a) and Videira et al. (2017). The study of Videira et al. (2017) used ITS and rpb2 while that of Crous et al. (2013a) used ITS, actA and tef1. The ITS sequences of the two cultures are 99 % similar (478/481 nt). Unfortunately, no other markers are shared between the two cultures and further comparison on more informative loci are therefore not possible. The species was described from Terminalia catappa in Tanzania and needs to be recollected from the type locality to confirm its phylogenetic placement. Pseudocercospora cornicola (Tracy & Earle) Y.L. Guo & X.J. Liu, Mycosystema 2: 232. 1989. Basionym: Cercospora cornicola Tracy & Earle, Bull. Torrey bot. Club 23(5): 205. 1896. Notes: The two cultures included in the present study (Suppl. Table S1) (Fig. 1, blocks 12 and 20) were isolated from different host species and different countries, namely Cornus alba var. sibirica in Japan (Crous et al. 2013a) and Cornus officinalis in South Korea (Choi et al. 2022). In their publication, Choi et al. (2022) indicated that the sequences from their material do not match the earlier published sequences of Ps. cornicola (ITS: 478/500 nt including seven gaps; actA: 162/198 nt including 12 gaps; tef1: 236/318 nt including 32 gaps). Pathogenicity of their isolates on the host was confirmed, fulfilling Koch’s postulates. The species was described from Cornus florida in Mississippi, USA, and needs to be recollected from the type locality to confirm its phylogenetic placement. Pseudocercospora cruenta (Sacc.) Deighton, Mycol. Pap. 140: 142. 1976. Basionym: Cercospora cruenta Sacc., Michelia 2(no. 6): 149. 1880. Notes: The two cultures included in the present study (Suppl. Table S1) (Fig. 1, blocks 13 and 28) were isolated from different host genera and different countries, namely Vigna sp. in Trinidad and Tobago (CBS 132021; Crous et al. 2013a) and Phaseolus vulgaris in Taiwan (CBS 117232; Crous et al. 2013a). The authors treated CBS 117232 as “Pseudocercospora cf. cruenta” (ITS: 495/500 nt including one gap; actA: 209/231 nt; tef1: 276/317 nt including nine gaps). The species was described from Phaseolus in South Carolina, USA, and needs to be recollected from the type locality to confirm its phylogenetic placement. Pseudocercospora jahnii (Syd.) U. Braun & Crous, in Crous & Braun, CBS Diversity Ser. (Utrecht) 1: 230. 2003. Basionym: Cercospora jahnii Syd., Annls Mycol. 28(1/2): 214. 1930. Notes: The two cultures included in the present study (Suppl. Table S1) (Fig. 1, blocks 44 and 45) were isolated from different host species and different countries, namely Tabebuia pallida in Philippines (CBS 138757; Acabal Jr et al. 2014) and Tabebuia chrysotricha in Thailand (CPC 19214; present study). The two cultures are genetically distant (ITS: 446/448 nt including one gap; tef1: 438/513 nt including 17 gaps). In their publication, Acabal Jr et al. (2014) indicated that this is a morphologically variable taxon, with conidia from the type specimen (not known from sequence data) are larger than those from Asian collections. The species was described from Tabebuia rosea in Venezuela, and needs to be recollected from the type locality to confirm its phylogenetic placement.