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Research Article
Multigene phylogeny and morphology reveal three new species of Cytospora isolated from diseased plant branches in Fengtai District, Beijing, China
expand article infoAoli Jia‡, Baoyue Chen§, Hongyan Lu§, Yu Xing§, Bin Li§, Xinlei Fan‡
‡ Beijing Forestry University, Beijing, China
§ Forestry Workstation, Fengtai District Bureau of Forestry and Parks of Beijing Municipality, Beijing, China

Corresponding author: Xinlei Fan ( xinleifan@bjfu.edu.cn )

Academic editor: Ning Jiang
© 2024 Aoli Jia, Baoyue Chen, Hongyan Lu, Yu Xing, Bin Li, Xinlei Fan.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Jia A, Chen B, Lu H, Xing Y, Li B, Fan X (2024) Multigene phylogeny and morphology reveal three new species of Cytospora isolated from diseased plant branches in Fengtai District, Beijing, China. MycoKeys 101: 163-189. https://doi.org/10.3897/mycokeys.101.116272
Open Access

Abstract

Members of Cytospora include saprobes, endophytes and important plant pathogens, which are widely distributed on various wood hosts and have a wide global distribution. In this study, the species definitions were conducted, based on multigene phylogeny (ITS, act, rpb2, tef1-α and tub2 genes) and comparisons of morphological characters. A total of 22 representative isolates obtained from 21 specimens in Fengtai District of Beijing City were identified as seven species of Cytospora, including four known species (C. albodisca, C. ailanthicola, C. euonymina, C. haidianensis) and three novel species (C. fengtaiensis, C. pinea, C. sorbariae). The results provide an understanding of the taxonomy of Cytospora species associated with canker and dieback diseases in Fengtai District, Beijing, China.

Key words

Canker disease, Diaporthales, pathogens, taxonomy

Introduction

The genus Cytospora was established by Ehrenberg (1818) and classified in Cytosporaceae, Diaporthales, Sordariomycetes (Wijayawardene et al. 2018; Fan et al. 2020). It includes numerous important pathogens associated with canker and dieback diseases of woody plants, with a worldwide distribution and broad host range (Sinclair et al. 1987; Adams et al. 2005, 2006; Lawrence et al. 2018; Fan et al. 2020; Lin et al. 2023a, b). Dieback and stem canker caused by Cytospora lead to the growth weakness or death of host plants, thereby causing significant economic and ecological losses (Sinclair et al. 1987; Adams et al. 2005). Currently, 695 species epithets of Cytospora have been listed in Index Fungorum (www.indexfungorum.org; accessed on 24 November 2023).

The taxonomy and correspondence between sexual and asexual morphs of Cytospora is quite confusing. Previous Cytospora species and their related sexual morphs viz. Leucostoma, Valsa, Valsella and Valseutypella were listed by old fungal literature for their identification (Fries 1823; Saccardo 1884; Kobayashi 1970; Barr 1978; Sutton 1980; Gvritishvili 1982; Spielman 1983, 1985). Adams et al. (2005) revised the genus Cytospora from Eucalyptus with 28 species and accepted all sexual genera combined under Valsa, either as subgenera or species without additional infrageneric rank, regarding the sexual genera (Leucocytospora, Leucostoma, Valsella and Valseutypella) as synonyms of Valsa. Based on the one fungus = one name initiative (Wingfield et al. 2012), Fan et al. (2015a, b) and Rossman et al. (2015) recommended to use Cytospora, the oldest name having priority over Valsa.

Cytospora canker symptoms initially appear on trunks and branches as slightly sunken bark with brown discolouration of the xylem, which may result in trunk and branch cracking (Adams et al. 2005). The asexual morph of Cytospora is characterised by the pycnidial stromata submerged in cortex with single or multiple locule(s), with or without conceptacle, filamentous conidiophores producing hyaline, allantoid, eguttulate and smooth conidia. The sexual morph is characterised by the ascomata submerged in the substrate with an erumpent pseudostroma, with or without necks. Asci are unitunicate, clavate to cylindrical with four or eight ascospores which are biseriate or multi-seriate, elongate-allantoid, thin-walled, hyaline and aseptate (Spielman 1983, 1985; Adams et al. 2005).

Currently, use of polyphasic approaches, such as morphological and phylogenetic analyses to define species of Cytospora has been proposed (Norphanphoun et al. 2017; Fan et al. 2020). In morphology, presence or absence of conceptacle, quantity and arrangement of locule(s), shape and size of conidiophores and conidial size are significantly taxonomic. In phylogeny, the current studies use the internal transcribed spacer (ITS), the partial actin (act), the RNA polymerase II subunit (rpb2), the translation elongation factor 1-α (tef1-α) and the beta-tubulin (tub2) genes to perform phylogenetic analysis.

Beijing is the capital city of China, located in the northern part of the North China Plain with more than 1,000 species of tree hosts (Liu et al. 2022). As more plant species were recorded in this city, the exploration of fungal diversity gradually increased as most fungi are often linked to particular host plants as pathogens or endophytes. With the modern taxonomic approaches applying, more than 30 Cytospora species have been reported in the last five years in Beijing (Fan et al. 2020; Pan et al. 2021; Lin et al. 2023a, b). Fengtai is one of the districts in Beijing with high forest cover and rich tree species which is located in the south-western suburbs of Beijing. However, there are few studies associated with fungal diversity in Fengtai District. A research to explore more hidden species of Cytospora in this region is considered imperative. Therefore, a survey on the diversity of Cytospora on diseased branches was conducted in Fengtai District from 2022 to 2023. The objectives of this study were to summarise the systematic study of Cytospora species in Fengtai District and to clarify the systematics and taxonomy of Cytospora species with detailed descriptions and illustrations and compare it to known species in the genus.

Materials and methods

Sample collection and isolation

Twenty-one fresh specimens with typical conidiomata and/or ascomata were collected from diseased branches or twigs of wood hosts which are distributed in Beigong National Forest Park, Century Forest Park, Garden Expo Park, Lotus Pond Park and Qianling Mountain in Fengtai District, Beijing City. Sampled trees expressed general symptoms and signs of canker diseases including elongate, slightly sunken and discoloured areas in the bark, several prominent dark conidiomata and/or ascomata immersed in bark, erumpent through the surface of bark when mature (Fig. 1). A total of 22 isolates were obtained by removing a mucoid spore mass from conidiomata and/or ascomata, spreading the suspension on the surface of 1.8% potato dextrose agar (PDA) (potato, 200 g; glucose, 20 g; agar, 20 g; distilled water, to complete 1000 ml) media in a Petri dish and incubating at 25 °C for up to 24 h. Hyphal tips were removed to a new PDA plate twice to obtain a pure culture. Specimens were deposited in the Museum of Beijing Forestry University (BJFC) and at the working Collection of X.L. Fan (CF), housed at the BJFU. Axenic cultures are maintained in the China Forestry Culture Collection Centre (CFCC).

Figure 1. 

Disease symptoms associated with Cytospora species collected from Fengtai District, Beijing, China A Acer palmatum ‘Atropurpureum’ B Acer pictum subsp. Mono. C Euonymus japonicus D Malus ‘American’ E Malus × micromalus F Pinus bungeanae G Salix babylonica H Sorbaria sorbifolia.

Morphological analyses

The identification of species was based on morphological characteristics of the ascomata or conidiomata formed on infected host materials. Macro-morphological features (structure and size of conidiomata and ascomata, ectostromatic disc and ostioles) were photographed using a Leica stereomicroscope (M205 FA) (Leica Microsystems, Wetzlar, Germany). Micromorphological features (conidiophores, conidiogenous cells, asci and conidia/ascospores) were photographed using a Nikon Eclipse 80i microscope (Nikon Corporation, Tokyo, Japan), equipped with a Nikon digital sight DS-Ri2 high resolution colour camera with differential interference contrast. Over 30 conidiomata were sectioned and 50 conidia were selected randomly to measure their lengths and widths. Colony diameters were measured and the colony colours described after 3 days and 14 days according to the colour charts of Rayner (1970).

DNA extraction, PCR amplification and sequencing

Mycelium used for DNA extraction was grown on PDA for three days and obtained from the cellophane surface by scraping. The genomic DNA was extracted using the modified CTAB method (Doyle and Doyle 1990). PCR amplifications and sequencing of five genes (ITS, act, rpb2, tef1-α and tub2) were performed. The primers and PCR conditions are listed in Table 1. PCR products were electrophoresed in 1% agarose gel and the DNA was sequenced by the Sino Geno Max Biotechnology Company Limited (Beijing, China). DNA sequences generated by the forward and reverse primers combination were used to obtain consensus sequences using Seqman v. 7.1.0 (DNASTAR Inc., Madison, WI, USA).

Table 1.
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Genes used in this study with PCR primers, primer DNA sequence, optimal annealing temperature and corresponding references.

Locus PCR primers PCR: thermal cycles: (Annealing temp. in bold) References of primers used
ITS ITS1 (95 °C: 30 s, 51 °C: 30 s, 72 °C: 1min) × 35 cycles White et al. (1990)
ITS4
act ACT-512F (95 °C: 45 s, 55 °C: 45 s, 72 °C: 1min) × 35 cycles Carbone and Kohn (1999)
ACT-783R
rpb2 RPB2-5F (95 °C: 30 s, 52 °C: 1 min, 72 °C:1 min) × 35 cycles Liu et al. (1999)
RPB2-7cR
tef1-α 728F (95 °C: 15 s, 55 °C: 20 s, 72 °C: 1min) × 35 cycles Rehner et al. (2005)
1567R
tub2 T1 (95 °C: 30 s, 55 °C: 30 s, 72 °C: 1min) × 35 cycles Glass and Donaldson (1995)
Bt2b

Phylogenetic analyses

The phylogenetic analyses were performed, based on the individual datasets of each gene region and combined five genes (ITS, act, rpb2, tef1-α and tub2) to compare Cytospora species from the current study with other sequences obtained from GenBank. The sequence datasets used in this study were based on Lin et al. (2023b). Sequence alignments of the individual gene were performed in MAFFT v. 6 (Katoh and Standley 2013) and adjusted by MEGA v. 6.0 (Tamura et al. 2013). Ambiguous regions were excluded from alignments. Phylogenetic analyses were conducted using the programme PhyML v. 3.0 (Guindon et al. 2010) for Maximum Likelihood (ML) analysis and MrBayes v. 3.1.2 (Ronquist and Huelsenbeck 2003) for Bayesian Inference (BI) analysis. For ML analysis, the substitution model (GTR+G+I model) for each dataset was selected following recent studies (Fan et al. 2020; Pan et al. 2020, 2021). Confidence levels for the nodes were determined using 1,000 replicates of bootstrapping methods (Hillis and Bull 1993). For BI analysis, the best-fit evolutionary models for each partitioned locus were estimated in MrModelTest v. 2.3 (Posada and Crandall 1998) with a Markov Chain Monte Carlo algorithm. Phylograms were plotted in FigTree v. 1.4.3 (http://tree.bio.ed.ac.uk/software/figtree) and edited in Adobe Illustrator CS6 v.16.0.0 (https://www.adobe.com/cn/products/illustrator.html). Sequence data were submitted to GenBank (https://www.ncbi.nlm.nih.gov) (Table 2). The multigene sequence alignments and the trees obtained were deposited in TreeBASE (https://treebase.org; study ID S30958).

Table 2.
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Strains of Cytospora used in the molecular analyses in this study.

Species Strain Host Origin GenBank accession numbers
ITS act rpb2 tef1-α tub2
Cytospora ailanthicola CFCC 89970 Ailanthus altissima Ningxia, China MH933618 MH933526 MH933592 MH933494 MH933565
Cytospora ailanthicola CFCC 59446 Salix matsudana Beijing, China OR826163 OR831996 OR832018 OR832040 OR832062
Cytospora albodisca CFCC 53161 Platycladus orientalis Beijing, China MW418406 MW422899 MW422909 MW422921 MW422933
Cytospora albodisca CFCC 54373 Platycladus orientalis Beijing, China MW418407 MW422900 MW422910 MW422922 MW422934
Cytospora albodisca CFCC 59467 Malus × micromalus Beijing, China OR826179 OR832012 OR832034 OR832056 OR832076
Cytospora albodisca CFCC 59537 Euonymus japonicus Beijing, China OR826180 OR832013 OR832035 OR832057 OR832077
Cytospora alba CFCC 55462T Salix matsudana Gansu, China MZ702593 OK303457 OK303516 OK303577 OK303644
Cytospora alba CFCC 55463T Salix matsudana Gansu, China MZ702594 OK303458 OK303517 OK303578 OK303645
Cytospora ampulliformis MFLUCC 16-0583T Sorbus intermedia Russia KY417726 KY417692 KY417794 NA NA
Cytospora ampulliformis MFLUCC 16-0629 Acer platanoides Russia KY417727 KY417693 KY417795 NA NA
Cytospora amydgali CBS 144233T Prunus dulcis California, USA MG971853 MG972002 NA MG971659 MG971718
Cytospora atrocirrhata CFCC 89615 Juglans regia Qinghai, China KR045618 KF498673 KU710946 KP310858 KR045659
Cytospora atrocirrhata CFCC 89616 Juglans regia Qinghai, China KR045619 KF498674 KU710947 KP310859 KR045660
Cytospora atrocirrhata CXY 1401 Populus sp. Inner Mongolia, China JX534242 NA NA NA KM034904
Cytospora atrocirrhata CXY 1402 Populus sp. Inner Mongolia, China JX534243 NA NA NA KM034903
Cytospora avicennae IRAN 4199CT Malus domestica Nahavand, Iran MW295650 MZ014511 MW824358 MW394145 NA
Cytospora avicennae IRAN 4625C Malus domestica Arak, Iran OM368648 NA NA OM372510 NA
Cytospora azerbaijanica IRAN 4201CT Malus domestica Urmia, Iran MW295526 MZ014513 MW824360 MW394147 NA
Cytospora azerbaijanica IRAN 4627C Malus domestica Miandoab, Iran OM368650 NA NA OM372512 NA
Cytospora beilinensis CFCC 50493T Pinus armandii Beijing, China MH933619 MH933527 NA MH933495 MH933561
Cytospora beilinensis CFCC 50494 Pinus armandii Beijing, China MH933620 MH933528 NA MH933496 MH933562
Cytospora berberidis CFCC 89927T Berberis dasystachya Qinghai, China KR045620 KU710990 KU710948 KU710913 KR045661
Cytospora berberidis CFCC 89933 Berberis dasystachya Qinghai, China KR045621 KU710991 KU710949 KU710914 KR045662
Cytospora bungeanae CFCC 50495T Pinus bungeanae Shanxi, China MH933621 MH933529 MH933593 MH933497 MH933563
Cytospora bungeanae CFCC 50496 Pinus bungeanae Shanxi, China MH933622 MH933530 MH933594 MH933498 MH933564
Cytospora calamicola MFLUCC 15-0397 Calamus Thailand NR_185736 NA NA ON734013 NA
Cytospora californica CBS 144234T Juglans regia California, USA MG971935 MG972083 NA MG971645 NA
Cytospora carbonacea CFCC 89947 Ulmus pumila Qinghai, China KR045622 KP310842 KU710950 KP310855 KP310825
Cytospora carpobroti CMW 48981T Carpobrotus edulis South Africa MH382812 NA NA MH411212 MH411207
Cytospora celtidicola CFCC 50497T Celtis sinensis Anhui, China MH933623 MH933531 MH933595 MH933499 MH933566
Cytospora celtidicola CFCC 50498 Celtis sinensis Anhui, China MH933624 MH933532 MH933596 MH933500 MH933567
Cytospora centrivillosa MFLUCC 16-1206T Sorbus domestica Italy MF190122 NA MF377600 NA NA
Cytospora centrivillosa MFLUCC 17-1660 Sorbus domestica Italy MF190123 NA MF377601 NA NA
Cytospora ceratosperma CFCC 89624 Juglans regia Gansu, China KR045645 NA KU710976 KP310860 KR045686
Cytospora ceratosperma CFCC 89625 Juglans regia Gansu, China KR045646 NA KU710977 KP31086 KR045687
Cytospora ceratospermopsis CFCC 89626T Juglans regia Shaanxi, China KR045647 KU711011 KU710978 KU710934 KR045688
Cytospora ceratospermopsis CFCC 89627 Juglans regia Shaanxi, China KR045648 KU711012 KU710979 KU710935 KR045689
Cytospora chrysosperma CFCC 89629 Salix psammophila Shaanxi, China KF765673 NA KF765705 NA NA
Cytospora chrysosperma CFCC 89981 Populus alba subsp. pyramidalis Gansu, China MH933625 MH933533 MH933597 MH933501 MH933568
Cytospora chrysosperma CFCC 89982 Ulmus pumila Tibet, China KP281261 KP310835 NA KP310848 KP310818
Cytospora cinnamomea CFCC 53178T Prunus armeniaca Xinjiang, China MK673054 MK673024 NA NA MK672970
Cytospora coryli CFCC 53162T Corylus mandshurica Beijing, China MN854450 NA MN850751 MN850758 MN861120
Cytospora corylina CFCC 54684T Corylus heterophylla Beijing, China MW839861 MW815951 MW815937 MW815886 MW883969
Cytospora corylina CFCC 54685 Corylus heterophylla Beijing, China MW839862 MW815952 MW815938 MW815887 MW883970
Cytospora corylina CFCC 54686 Corylus heterophylla Beijing, China MW839863 MW815953 MW815939 MW815888 MW883971
Cytospora corylina CFCC 54687 Corylus heterophylla Beijing, China MW839864 MW815954 MW815940 MW815889 MW883972
Cytospora cotini MFLUCC 14-1050T Cotinus coggygria Russia KX430142 NA KX430144 NA NA
Cytospora cotoneastricola CF 20197027 Cotoneaster sp. Tibet, China MK673072 MK673042 MK673012 MK672958 MK672988
Cytospora cotoneastricola CF 20197028 Cotoneaster sp. Tibet, China MK673073 MK673043 MK673013 MK672959 MK672989
Cytospora cotoneastricola CF 20197030 Cotoneaster sp. Tibet, China MK673074 MK673044 MK673014 MK672960 MK672990
Cytospora cotoneastricola CF 20197031T Cotoneaster sp. Tibet, China MK673075 MK673045 MK673015 MK672961 MK672991
Cytospora curvata MFLUCC 15-0865T Salix alba Russia KY417728 KY417694 KY417796 NA NA
Cytospora curvispora CFCC 54000T Corylus heterophylla Beijing, China MW839851 MW815931 MW815945 MW815880 MW883963
Cytospora curvispora CFCC 54001 Corylus heterophylla Beijing, China MW839853 MW815932 MW815946 MW815881 MW883964
Cytospora curvispora CFCC 54676 Corylus heterophylla Beijing, China MW839854 MW815933 MW815947 MW815882 MW883965
Cytospora curvispora CFCC 54677 Corylus heterophylla Beijing, China MW839855 MW815934 MW815948 MW815883 MW883966
Cytospora curvispora CFCC 54678 Corylus heterophylla Beijing, China MW839856 MW815935 MW815949 MW815884 MW883967
Cytospora curvispora CFCC 54679 Corylus heterophylla Beijing, China MW839857 MW815936 MW815950 MW815885 MW883968
Cytospora davidiana CXY 1350T Populus davidiana Inner Mongolia, China KM034870 NA NA NA NA
Cytospora diopuiensis MFLUCC 18-1419T Undefined wood Chiang Mai, Thailand MK912137 MN685819 NA NA NA
Cytospora diopuiensis CFCC55884 Kerria japonica f. pleniflora Beijing, China OK316819 NA OK358569 OK358471 OK358473
Cytospora diopuiensis CFCC55885 Kerria japonica f. pleniflora Beijing, China OK316820 NA OK358570 OK358472 OK358474
Cytospora diopuiensis CFCC 56961 Koelreuteria paniculata Beijing, China ON376918 ON390905 ON390908 ON390914 ON390923
Cytospora diopuiensis CFCC 56970 Koelreuteria paniculata Beijing, China ON376917 ON390904 ON390907 ON390913 ON390922
Cytospora diopuiensis CFCC 56971 Koelreuteria paniculata Beijing, China ON376919 ON390906 NA ON390915 NA
Cytospora discotoma CFCC 53137T Platycladus orientalis Beijing, China MW418404 MW422897 MW422907 MW422919 MW422931
Cytospora discotoma CFCC 54368 Platycladus orientalis Beijing, China MW418405 MW422898 MW422908 MW422920 MW422932
Cytospora donetzica MFLUCC 15-0864 Crataegus monogyna Russia KY417729 KY417695 KY417797 NA NA
Cytospora donetzica MFLUCC 16-0574T Crataegus monogyna Russia KY417731 KY417697 KY417799 NA NA
Cytospora donglingensis CFCC 53159T Platycladus orientalis Beijing, China MW418412 MW422903 MW422915 MW422927 MW422939
Cytospora donglingensis CFCC 53160 Platycladus orientalis Beijing, China MW418414 MW422905 MW422917 MW422929 MW422941
Cytospora donglingensis CFCC 54371 Platycladus orientalis Beijing, China MW418413 MW422904 MW422916 MW422928 MW422940
Cytospora donglingensis CFCC 54372 Platycladus orientalis Beijing, China MW418415 MW422906 MW422918 MW422930 MW422942
Cytospora elaeagni CFCC 89632 Elaeagnus angustifolia Ningxia, China KR045626 KU710995 KU710955 KU710918 KR045667
Cytospora elaeagni CFCC 89633 Elaeagnus angustifolia Ningxia, China KF765677 KU710996 KU710956 KU710919 KR045668
Cytospora elaeagnicola CFCC 52882T Elaeagnus angustifolia Xinjiang, China MK732341 MK732344 MK732347 NA NA
Cytospora elaeagnicola CFCC 52883 Elaeagnus angustifolia Xinjiang, China MK732342 MK732345 MK732348 NA NA
Cytospora elaeagnicola CFCC 52884 Elaeagnus angustifolia Xinjiang, China MK732343 MK732346 MK732349 NA NA
Cytospora ershadii IRAN 4197C Malus domestica Nahavand, Iran MW295510 NA NA MW394143 NA
Cytospora ershadii IRAN 4198CT Malus domestica Arak, Iran MW295523 MZ014510 MW824357 MW394144 NA
Cytospora erumpens CFCC 50022 Prunus padus Shanxi, China MH933627 MH933534 NA MH933502 MH933569
Cytospora erumpens MFLUCC 16-0580T Salix × fragilis Russia KY417733 KY417699 KY417801 NA NA
Cytospora erumpens CFCC 53163 Prunus padus Xinjiang, China MK673059 MK673029 MK673000 MK672948 MK672975
Cytospora eucalypti CBS 144241 Eucalyptus globulus California, USA MG971907 MG972056 NA MG971617 MG971772
Cytospora euonymicola CFCC 50499T Euonymus kiautschovicus Shaanxi, China MH933628 MH933535 MH933598 MH933503 MH933570
Cytospora euonymicola CFCC 50500 Euonymus kiautschovicus Shaanxi, China MH933629 MH933536 MH933599 MH933504 MH933571
Cytospora euonymina CFCC 89993T Euonymus kiautschovicus Shanxi, China MH933630 MH933537 MH933600 MH933505 MH933590
Cytospora euonymina CFCC 89999 Euonymus kiautschovicus Shanxi, China MH933631 MH933538 MH933601 MH933506 MH933591
Cytospora euonymina CFCC 59444 Salix babylonica Beijing, China OR826164 OR831997 OR832019 OR832041 NA
Cytospora euonymina CFCC 59479 Salix babylonica Beijing, China OR826165 OR831998 OR832020 OR832042 NA
Cytospora fengtaiensis CFCC 59442 Acer palmatum ‘Atropurpureum’ Beijing, China OR826166 OR831999 OR832021 OR832043 OR832063
Cytospora fengtaiensis CFCC 59449T Acer palmatum ‘Atropurpureum’ Beijing, China OR826167 OR832000 OR832022 OR832044 OR832064
Cytospora fengtaiensis CFCC 59525 Acer palmatum ‘Atropurpureum’ Beijing, China OR826168 OR832001 OR832023 OR832045 OR832065
Cytospora fengtaiensis CFCC 59526 Acer palmatum ‘Atropurpureum’ Beijing, China OR826169 OR832002 OR832024 OR832046 OR832066
Cytospora fengtaiensis CFCC 59527 Acer palmatum ‘Atropurpureum’ Beijing, China OR826170 OR832003 OR832025 OR832047 OR832067
Cytospora fraxinigena BBH 42442 Fraxinus ornus NA MF190133 NA NA NA NA
Cytospora fraxinigena MFLUCC 14-0868T Fraxinus ornus Italy MF190133 NA NA NA NA
Cytospora fugax CXY 1371 Populus simonii Jilin, China KM034852 NA NA NA KM034891
Cytospora fugax CXY 1381 Populus ussuriensis Heilongjiang, China KM034853 NA NA NA KM034890
Cytospora galegicola MFLUCC 18-1199T Galega officinalis Forlì-Cesena, Italy MK912128 MN685810 MN685820 NA NA
Cytospora gigalocus CFCC 89620T Juglans regia Qinghai, China KR045628 KU710997 KU710957 KU710920 KR045669
Cytospora gigalocus CFCC 89621 Juglans regia Qinghai, China KR045629 KU710998 KU710958 KU710921 KR045670
Cytospora gigaspora CFCC 50014 Juniperus procumbens Shanxi, China KR045630 KU710999. KU710959 KU710922 KR045671
Cytospora gigaspora CFCC 89634T Salix psammophila Shaanxi, China KF765671 KU711000 KU710960 KU710923 KR045672
Cytospora globosa MFLU 16-2054T Abies alba Italy MT177935 NA MT432212 MT454016 NA
Cytospora granati CBS 144237T Punica granatum California, USA MG971799 MG971949 NA MG971514 MG971664
Cytospora haidianensis CFCC 54056 Euonymus alatus Beijing, China MT360041 MT363978 MT363987 MT363997 MT364007
Cytospora haidianensis CFCC 54057T Euonymus alatus Beijing, China MT360042 MT363979 MT363988 MT363998 MT364008
Cytospora haidianensis CFCC 54184 Euonymus alatus Beijing, China MT360043 MT363980 MT363989 MT363999 MT364009
Cytospora haidianensis CFCC 59450 Euonymus japonicus Beijing, China OR826171 OR832004 OR832026 OR832048 OR832068
Cytospora haidianensis CFCC 59475 Malus ‘American’ Beijing, China OR826172 OR832005 OR832027 OR832049 OR832069
Cytospora haidianensis CFCC 59471 Acer pictum subsp. mono Beijing, China OR826173 OR832006 OR832028 OR832050 OR832070
Cytospora haidianensis CFCC 59536 Acer pictum subsp. mono Beijing, China OR826174 OR832007 OR832029 OR832051 OR832071
Cytospora hippophaës CFCC 89639 Hippophaë rhamnoides Gansu, China KR045632 KU711001 KU710961 KU710924 KR045673
Cytospora hippophaës CFCC 89640 Hippophaë rhamnoides Gansu, China KF765682 KF765730 KU710962 KP310865 KR045674
Cytospora huairouensis CFCC 56940 Prunus armeniaca Beijing, China ON188758 OR662079 OR662096 OR662113 OR662060
Cytospora huairouensis CFCC 56973 Prunus armeniaca Beijing, China ON188759 OR662080 OR662097 OR662114 OR662061
Cytospora huairouensis CFCC 57286 Prunus armeniaca Beijing, China ON188760 OR662081 OR662098 OR662115 OR662062
Cytospora iranica IRAN 4200CT Malus domestica Arak, Iran MW295652 MZ014512 MW824359 MW394146 NA
Cytospora iranica IRAN 4628C Malus domestica Nahavand, Iran OM368651 NA NA OM372513 NA
Cytospora japonica CFCC 89956 Prunus cerasifera Ningxia, China KR045624 KU710993 KU710953 KU710916 KR045665
Cytospora japonica CFCC 89960 Prunus cerasifera Ningxia, China KR045625 KU710994 KU710954 KU710917 KR045666
Cytospora joaquinensis CBS 144235 Populus deltoides California, USA MG971895 MG972044 NA MG971605 MG971761
Cytospora junipericola BBH 42444 Juniperus communis Italy MF190126 NA NA MF377579 NA
Cytospora junipericola MFLU 17-0882T Juniperus communis Italy MF190125 NA NA MF377580 NA
Cytospora juniperina CFCC 50501T Juniperus przewalskii Sichuan, China MH933632 MH933539 MH933602 MH933507 NA
Cytospora juniperina CFCC 50502 Juniperus przewalskii Sichuan, China MH933633 MH933540 MH933603 MH933508 MH933572
Cytospora juniperina CFCC 50503 Juniperus przewalskii Sichuan, China MH933634 MH933541 MH933604 MH933509 NA
Cytospora kantschavelii CXY 1383 Populus maximowiczii Jilin, China KM034867 NA NA NA NA
Cytospora kantschavelii CXY 1386 Populus maximowiczii Chongqing, China KM034867 NA NA NA NA
Cytospora kuanchengensis CFCC 52464T Castanea mollissima Hebei, China MK432616 MK442940 MK578076 NA NA
Cytospora kuanchengensis CFCC 52465 Castanea mollissima Hebei, China MK432617 MK442941 MK578077 NA NA
Cytospora longispora CBS 144236T Prunus domestica California, USA MG971905 MG972054 NA MG971615 MG971764
Cytospora longistiolata MFLUCC 16-0628 Salix × fragilis Russia KY417734 KY417700 KY417802 NA NA
Cytospora leucosperma CFCC 89622 Pyrus bretschneideri Gansu, China KR045616 KU710988 KU710944 KU710911 KR045657
Cytospora leucosperma CFCC 89894 Pyrus bretschneideri Qinghai, China KR045617 KU710989 KU710945 KU710912 KR045658
Cytospora leucostoma CFCC 50023 Cornus alba Shanxi, China KR045635 KU711003 KU710964 KU710926 KR045676
Cytospora leucostoma CFCC 50024 Prunus pseudocerasus Qinghai, China MH933640 MH933547 MH933605 NA MH933576
Cytospora leucostoma CFCC 53140 Prunus sibirica Beijing, China MN854445 MN850760 MN850746 MN850753 MN861115
Cytospora leucostoma CFCC 53141 Prunus sibirica Beijing, China MN854446 MN850761 MN850747 MN850754 MN861116
Cytospora leucostoma CFCC 53156 Juglans mandshurica Beijing, China MN854447 MN850762 MN850748 MN850755 MN861117
Cytospora leucostoma CFCC 53167 Prunus armeniaca Xinjiang, China MK673056 MK673026 MK672998 MK672946 MK672972
Cytospora leucostoma CFCC 53169 Prunus persica Beijing, China MK673080 MK673050 MK673020 MK672966 MK672996
Cytospora leucostoma CFCC 53170 Prunus persica Beijing, China MK673081 MK673051 MK673021 MK672967 MK672997
Cytospora leucostoma CFCC 54680 Corylus heterophylla Beijing, China MW839857 MW815941 MW815955 MW815890 MW883973
Cytospora leucostoma CFCC 54681 Corylus heterophylla Beijing, China MW839857 MW815942 MW815956 MW815891 MW883974
Cytospora leucostoma CFCC 54682 Corylus heterophylla Beijing, China MW839857 MW815943 MW815957 MW815892 MW883975
Cytospora leucostoma CFCC 54683 Corylus heterophylla Beijing, China MW839857 MW815944 MW815958 MW815893 MW883976
Cytospora lumnitzericola MFLUCC 17-0508T Lumnitzera racernosa Tailand MG975778 MH253457 MH253453 NA NA
Cytospora macropycnidia CBS 149338 Vitis vinifera USA OP038094 OP003977 OP095265 OP106954 OP079909
Cytospora mali CFCC 50028 Malus pumila Gansu, China MH933641 MH933548 MH933606 MH933513 MH933577
Cytospora mali CFCC 50029 Malus pumila Ningxia, China MH933642 MH933549 MH933607 MH933514 MH933578
Cytospora mali CFCC 50030 Malus pumila Shaanxi, China MH933643 MH933550 MH933608 MH933524 MH933579
Cytospora mali CFCC 50031 Crataegus sp. Shanxi, China KR045636 KU711004 KU710965 KU710927 KR045677
Cytospora mali CFCC 50044 Malus baccata Qinghai, China KR045637 KU711005 KU710966 KU710928 KR045678
Cytospora mali-spectabilis CFCC 53181T Malus spectabilis ‘Royalty’ Xinjiang, China MK673066 MK673036 MK673006 MK672953 MK672982
Cytospora melnikii CFCC 89984 Rhus typhina Xinjiang, China MH933678 MH933551 MH933609 MH933515 MH933580
Cytospora melnikii MFLUCC 15-0851 Malus domestica Russia KY417735 KY417701 KY417803 NA NA
Cytospora melnikii MFLUCC 16-0635 Populus nigra var. italica Russia KY417736 KY417702 KY417804 NA NA
Cytospora myrtagena CFCC 52454 Castanea mollissima Shaanxi, China MK432614 MK442938 MK578074 NA NA
Cytospora myrtagena CFCC 52455 Castanea mollissima Shaanxi, China MK432615 MK442939 MK578075 NA NA
Cytospora nivea MFLUCC 15-0860 Salix acutifolia Russia KY417737 KY417703 KY417805 NA NA
Cytospora nivea CFCC 89641 Elaeagnus angustifolia Ningxia, China KF765683 KU711006 KU710967 KU710929 KR045679
Cytospora nivea CFCC 89643 Salix psammophila Shaanxi, China KF765685 NA KU710968 KP310863 KP310829
Cytospora notastroma NE_TFR5 Populus tremuloides USA JX438632 NA NA JX438543 NA
Cytospora notastroma NE_TFR8 Populus tremuloides USA JX438633 NA NA JX438542 NA
Cytospora ochracea CFCC 53164T Cotoneaster sp. Xinjiang, China MK673060 MK673030 MK673001 MK672949 MK672976
Cytospora oleicola CBS 144248T Olea europaea California, USA MG971944 MG972098 NA MG971660 MG971752
Cytospora olivacea CFCC 53174 Prunus cerasifera Xinjiang, China MK673058 MK673028 MK672999 NA MK672974
Cytospora olivacea CFCC 53175 Prunus dulcis Xinjiang, China MK673062 MK673032 MK673003 NA MK672978
Cytospora olivacea CFCC 53176T Sorbus tianschanica Xinjiang, China MK673068 MK673038 MK673008 MK672955 MK672984
Cytospora olivacea CFCC 53177 Prunus virginiana Xinjiang, China MK673071 MK673041 MK673011 NA MK672987
C. olivarum UCD634-Oe CBS 145585 Olea europaea Ventura Co., CA, U.S.A. MK514094 MK509025 NA MK509030 MK509035
C. olivarum UCD644-Oe Olea europaea Ventura Co., CA, U.S.A. MK514095 MK509026 NA MK509031 MK509036
Cytospora palm CXY 1276 Cotinus coggygria Beijing, China JN402990 NA NA KJ781296 NA
Cytospora palm CXY 1280T Cotinus coggygria Beijing, China JN411939 NA NA KJ781297 NA
Cytospora paracinnamomea CFCC 55453T Salix matsudana Gansu, China MZ702594 OK303456 OK303515 OK303576 OK303643
Cytospora paracinnamomea CFCC 55455T Salix matsudana Gansu, China MZ702598 OK303460 OK303519 OK303580 OK303647
Cytospora parakantschavelii MFLUCC 15-0857T Populus × sibirica Russia KY417738 KY417704 KY417806 NA NA
Cytospora parakantschavelii MFLUCC 16-0575 Pyruspyraster Russia KY417739 KY417705 KY417807 NA NA
Cytospora parapistaciae CBS 144506T Pistacia vera California, USA MG971804 MG971954 NA MG971519 MG971669
Cytospora parasitica MFLUCC 15-0507T Malus domestica Russia KY417740 KY417706 KY417808 NA NA
Cytospora parasitica XJAU 2542-1 Malus sp. Xinjiang, China MH798884 NA NA MH813452 NA
Cytospora parasitica CFCC 53171 Malus pumila Xinjiang, China MK673061 MK673031 MK673002 MK672950 MK672977
Cytospora parasitica CFCC 53172 Malus pumila Xinjiang, China MK673069 MK673039 MK673009 MK672956 MK672985
Cytospora parasitica CFCC 53173 Berberis sp. Xinjiang, China MK673070 MK673040 MK673010 MK672957 MK672986
Cytospora paratranslucens MFLUCC 15-0506T Populus alba var. bolleana Russia KY417741 KY417707 KY417809 NA NA
Cytospora paratranslucens MFLUCC 16-0627 Populus alba Russia KY417742 KY417708 KY417810 NA NA
Cytospora paraplurivora FDS-439 Prunus armeniaca Canada OL640182 OL631586 NA OL631589 NA
Cytospora paraplurivora FDS-564 Prunus persica var. nucipersica Canada OL640183 OL631587 NA OL631590 NA
Cytospora paraplurivora FDS-623 Prunus persica var. persica Canada OL640181 OL631588 NA OL631591 NA
Cytospora phialidica MFLUCC 17-2498 Alnus glutinosa Italy MT177932 NA MT432209 MT454014 NA
Cytospora piceae CFCC 52841T Picea crassifolia Xinjiang, China MH820398 MH820406 MH820395 MH820402 MH820387
Cytospora piceae CFCC 52842 Picea crassifolia Xinjiang, China MH820399 MH820407 MH820396 MH820403 MH820388
Cytospora pinea CFCC 59521T Pinus bungeanae Beijing, China OR826181 OR832014 OR832036 OR832058 OR832078
Cytospora pinea CFCC 59522 Pinus bungeanae Beijing, China OR826182 OR832015 OR832037 OR832059 OR832079
Cytospora pinea CFCC 59523 Pinus bungeanae Beijing, China OR826183 OR832016 OR832038 OR832060 OR832080
Cytospora pinea CFCC 59524 Pinus bungeanae Beijing, China OR826184 OR832017 OR832039 OR832061 OR832081
Cytospora pingbianensis MFLUCC 18-1204T Undefined wood Yunnan, China MK912135 MN685817 MN685826 NA NA
Cytospora pistaciae CBS 144238T Pistacia vera California, USA MG971802 MG971952 NA MG971517 MG971667
Cytospora platanicola MFLU 17-0327 Platanus hybrida Italy MH253451 MH253449 MH253450 NA NA
Cytospora platyclada CFCC 50504T Platycladus orientalis Yunnan, China MH933645 MH933552 MH933610 MH933516 MH933581
Cytospora platyclada CFCC 50505 Platycladus orientalis Yunnan, China MH933646 MH933553 MH933611 MH933517 MH933582
Cytospora platyclada CFCC 50506 Platycladus orientalis Yunnan, China MH933647 MH933554 MH933612 MH933518 MH933583
Cytospora platycladicola CFCC 50038T Platycladus orientalis Gansu, China KT222840 MH933555 MH933613 MH933519 MH933584
Cytospora platycladicola CFCC 50039 Platycladus orientalis Gansu, China KR045642 KU711008 KU710973 KU710931 KR045683
Cytospora plurivora CBS 144239T Olea europaea California, USA MG971861 MG972010 NA MG971572 MG971726
Cytospora populicola CBS 144240 Populus deltoides California, USA MG971891 MG972040 NA MG971601 MG971757
Cytospora populina CFCC 89644T Salix psammophila Shaanxi, China KF765686 KU711007 KU710969 KU710930 KR045681
Cytospora populinopsis CFCC 50032T Sorbus aucuparia Ningxia, China MH933648 MH933556 MH933614 MH933520 MH933585
Cytospora populinopsis CFCC 50033 Sorbus aucuparia Ningxia, China MH933649 MH933557 MH933615 MH933521 MH933586
Cytospora predappioensis MFLUCC 17-2458T Platanus hybrida Italy MG873484 NA NA NA NA
Cytospora prunicola MFLU 17-0995T Prunus sp. Italy MG742350 MG742353 MG742352 NA NA
Cytospora pruni-mume CFCC 53179 Prunus armeniaca Xinjiang, China MK673057 MK673027 NA MK672947 MK672973
Cytospora pruni-mume CFCC 53180T Prunus mume Xinjiang, China MK673067 MK673037 MK673007 MK672954 MK672983
Cytospora prunina CFCC 58997 Prunus armeniaca Beijing, China OR578808 NA NA NA OR662077
Cytospora prunina CFCC 58998 Prunus armeniaca Beijing, China OR578809 NA NA NA OR662078
Cytospora pruinopsis CFCC 50034T Ulmus pumila Shaanxi, China KP281259 KP310836 KU710970 KP310849 KP310819
Cytospora pruinopsis CFCC 50035 Ulmus pumila Jilin, China KP281260 KP310837 KU710971 KP310850 KP310820
Cytospora pruinopsis CFCC 53153 Ulmus pumila Beijing, China MN854451 MN850763 MN850752 MN850759 MN861121
Cytospora pruinosa CFCC 50036 Syringa oblata Qinghai, China KP310800 KP310832 NA KP310845 KP310815
Cytospora pruinosa CFCC 50037 Syringa oblata Qinghai, China MH933650 MH933558 NA MH933522 MH933589
Cytospora pubescentis MFLUCC 18-1201T Quercus pubescens Forlì-Cesena, Italy MK912130 MN685812 MN685821 NA NA
Cytospora punicae CBS 144244 Punica granatum California, USA MG971943 MG972091 NA MG971654 MG971798
Cytospora quercicola MFLU 17-0881 Quercus sp. Italy MF190128 NA NA NA NA
Cytospora quercicola MFLUCC 14-0867T Quercus sp. Italy MF190129 NA NA NA NA
Cytospora ribis CFCC 50026 Ulmus pumila Qinghai, China KP281267 KP310843 KU710972 KP310856 KP310826
Cytospora ribis CFCC 50027 Ulmus pumila Qinghai, China KP281268 KP310844 NA KP310857 KP310827
Cytospora rosae MFLU 17-0885 Rosa canina Italy MF190131 NA NA NA NA
Cytospora rosicola CF 20197024T Rosa sp. Tibet, China MK673079 MK673049 MK673019 MK672965 MK672995
Cytospora rosigena MFLUCC 18-0921T Rosa sp. Russia MN879872 NA NA NA NA
Cytospora rostrata CFCC 89909 Salix cupularis Gansu, China KR045643 KU711009 KU710974 KU710932 KR045684
Cytospora rostrata CFCC 89910 Salix cupularis Gansu, China KR045644 KU711010 KU710975 KU710933 NA
Cytospora rusanovii MFLUCC 15-0853 Populus × sibirica Russia KY417743 KY417709 KY417811 NA NA
Cytospora rusanovii MFLUCC 15-0854T Salix babylonica Russia KY417744 KY417710 KY417812 NA NA
Cytospora salicacearum MFLUCC 15-0509 Salix alba Russia KY417746 KY417712 KY417814 NA NA
Cytospora salicacearum MFLUCC 15-0861 Salix × fragilis Russia KY417745 KY417711 KY417813 NA NA
Cytospora salicacearum MFLUCC 16-0587 Prunus cerasus Russia KY417742 KY417708 KY417810 NA NA
Cytospora salicacearum MFLUCC 16-0576 Populus nigra var. italica Russia KY417741 KY417707 KY417809 NA NA
Cytospora salicicola MFLUCC 14-1052T Salix alba Russia KU982636 KU982637 NA NA NA
Cytospora salicicola MFLUCC 15-0866 Salix sp. Thailand KY417749 KY417715 KY417817 NA NA
Cytospora salicina MFLUCC 15-0862 Salix alba Russia KY417750 KY417716 KY417818 NA NA
Cytospora salicina MFLUCC 16-0637 Salix × fragilis Russia KY417751 KY417717 KY417819 NA NA
Cytospora schulzeri CFCC 50042 Malus pumila Gansu, China KR045650 KU711014 KU710981 KU710937 KR045691
Cytospora sibiraeae CFCC 50045T Sibiraea angustata Gansu, China KR045651 KU711015 KU710982 KU710938 KR045692
Cytospora sibiraeae CFCC 50046 Sibiraea angustata Gansu, China KR045652 KU711015 KU710983 KU710939 KR045693
Cytospora sophorae CFCC 50047 Styphnolobium japonicum Shanxi, China KR045653 KU711017 KU710984 KU710940 KR045694
Cytospora sophorae CFCC 50048 Magnolia grandiflora Shanxi, China MH820401 MH820409 MH820397 MH820405 MH820390
Cytospora sophorae CFCC 89598 Styphnolobium japonicum Gansu, China KR045654 KU711018 KU710985 KU710941 KR045695
Cytospora sophoricola CFCC 89596 Styphnolobium japonicum var. pendula Gansu, China KR045656 KU711020 KU710987 KU710943 KR045697
Cytospora sophoricola CFCC 89595T Styphnolobium japonicum var. pendula Gansu, China KR045655 KU711019 KU710986 KU710942 KR045696
Cytospora sophoriopsis CFCC 55469 Salix matsudana Gansu, China MZ702583 OK303445 OK303504 OK303565 OK303632
Cytospora sophoriopsis CFCC 89600 Styphnolobium japonicum Gansu, China KR045623 KU710992 KU710951 KU710915 KP310817
Cytospora sorbariae CFCC 59443 Sorbaria sorbifolia Beijing, China OR826175 OR832008 OR832030 OR832052 OR832072
Cytospora sorbariae CFCC 59445T Sorbaria sorbifolia Beijing, China OR826176 OR832009 OR832031 OR832053 OR832073
Cytospora sorbariae CFCC 59529 Sorbaria sorbifolia Beijing, China OR826177 OR832010 OR832032 OR832054 OR832074
Cytospora sorbariae CFCC 59530 Sorbaria sorbifolia Beijing, China OR826178 OR832011 OR832033 OR832055 OR832075
Cytospora sorbi MFLUCC 16-0631T Sorbus aucuparia Russia KY417752 KY417718 KY417820 NA NA
Cytospora sorbicola MFLUCC 16-0584T Acer pseudoplatanus Russia KY417755 KY417721 KY417823 NA NA
Cytospora sorbicola MFLUCC 16-0633 Cotoneaster melanocarpus Russia KY417758 KY417724 KY417826 NA NA
Cytospora sorbina CF 20197660T Sorbus tianschanica Xinjiang, China MK673052 MK673022 NA MK672943 MK672968
Cytospora spiraeae CFCC 50049T Spiraeasalicifolia Gansu, China MG707859 MG708196 MG708199 NA NA
Cytospora spiraeae CFCC 50050 Spiraeasalicifolia Gansu, China MG707860 MG708197 MG708200 NA NA
Cytospora spiraeicola CFCC 53138T Spiraeasalicifolia Beijing, China MN854448 NA MN850749 MN850756 MN861118
Cytospora spiraeicola CFCC 53139 Tilia nobilis Beijing, China MN854449 NA MN850750 MN850757 MN861119
Cytospora tamaricicola CFCC 50507 Rosa multifolora Yunnan, China MH933651 MH933559 MH933616 MH933525 MH933587
Cytospora tamaricicola CFCC 50508T Tamarix chinensis Yunnan, China MH933652 MH933560 MH933617 MH933523 MH933588
Cytospora tanaitica MFLUCC 14-1057T Betula pubescens Russia KT459411 KT459413 NA NA NA
Cytospora thailandica MFLUCC 17-0262T Xylocarpus moluccensis Thailand MG975776 MH253459 MH253455 NA NA
Cytospora thailandica MFLUCC 17-0263T Xylocarpus moluccensis Thailand MG975777 MH253460 MH253456 NA NA
Cytospora tibetensis CF 20197026 Cotoneaster sp. Tibet, China MK673076 MK673046 MK673016 MK672962 MK672992
Cytospora tibetensis CF 20197029 Cotoneaster sp. Tibet, China MK673077 MK673047 MK673017 MK672963 MK672993
Cytospora tibetensis CF 20197032T Cotoneaster sp. Tibet, China MK673078 MK673048 MK673018 MK672964 MK672994
Cytospora tibouchinae CPC 26333T Tibouchina semidecandra France KX228284 NA NA NA NA
Cytospora translucens CXY 1351 Populus davidiana Inner Mongolia, China KM034874 NA NA NA KM034895
Cytospora translucens CXY 1359 Populus × Beijingensis Beijing, China KM034871 NA NA NA KM034894
Cytospora ulmi MFLUCC 15-0863T Ulmus minor Russia KY417759 NA NA NA NA
Cytospora verrucosa CFCC 53157 T Platycladus orientalis Beijing, China MW418408 NA MW422911 MW422923 MW422935
Cytospora verrucosa CFCC 53158 Platycladus orientalis Beijing, China MW418410 MW422901 MW422913 MW422925 MW422937
Cytospora verrucosa CFCC 54369 Platycladus orientalis Beijing, China MW418409 NA MW422912 MW422924 MW422936
Cytospora verrucosa CFCC 54370 Platycladus orientalis Beijing, China MW418411 MW422902 MW422914 MW422926 MW422938
Cytospora vinacea CBS 141585T Vitis interspecific hybrid ‘Vidal’ USA KX256256 NA NA KX256277 KX256235
Cytospora viridistroma CBS 202.36T Cercis canadensis USA MN172408 NA NA MN271853 NA
Cytospora viticola Cyt2 Vitis interspecific hybrid ‘Frontenac’ USA KX256238 NA NA KX256259 KX256217
Cytospora viticola CBS 141586T Vitis vinifera ‘CabernetFranc’ USA KX256239 NA NA KX256260 KX256218
Cytospora xinjiangensis CFCC 53182 Rosa sp. Xinjiang, China MK673064 MK673034 MK673004 MK672951 MK672980
Cytospora xinjiangensis CFCC 53183T Rosa sp. Xinjiang, China MK673065 MK673035 MK673005 MK672952 MK672981
Cytospora xinglongensis CFCC 52458T Castanea mollissima Hebei, China MK432622 MK442946 MK578082 NA NA
Cytospora xinglongensis CFCC 52459 Castanea mollissima Hebei, China MK432623 MK442947 MK578083 NA NA
Cytospora xylocarpi MFLUCC 17-0251T Xylocarpus granatum Thailand MG975775 MH253458 MH253454 NA NA
Cytospora yakimana CBS 149297 Vitis vinifera USA OM976602 ON012555 ON045093 ON012569 ON086750
Cytospora yakimana CBS 149298 Vitis vinifera USA OM976603 ON012556 ON045094 ON012570 ON086751
Cytospora zhaitangensis CFCC 56227T Euonymus japonicus Beijing, China OQ344750 OQ398760 OQ398789 OQ410623 OQ398733
Cytospora zhaitangensis CFCC 57537 Euonymus japonicus Beijing, China OQ344751 OQ398761 OQ398790 OQ410624 OQ398734
Diaporthe vaccinii CBS 160.32 Vaccinium macrocarpon USA KC343228 JQ807297 NA KC343954 KC344196

1Acronyms: ATCC: American Type Culture Collection, Virginia, USA; BBH: BIOTEC Bangkok Herbarium, National Science and Technology Development Agency, Thailand; CBS: Westerdijk Fungal Biodiversity Institute (CBS-KNAW Fungal Biodiversity Centre), Utrecht, The Netherlands; CFCC: China Forestry Culture Collection Centre, Beijing, China; CMW: Culture Collection of Michael Wingfield, University of Pretoria, South Africa; CPC: Culture Collection of Pedro Crous, The Netherlands; IMI: Culture Collection of the International Mycological Institute, CABI Bioscience, Egham, Surrey, UK; MFLU: Mae Fah Luang University herbarium, Thailand; MFLUCC: Mae Fah Luang University Culture Collection, Thailand; MUCC: Murdoch University Culture Collection, Perth, Australia; NE: Gerard Adams Collections, University of Nebraska, Lincoln NE; PPRI: Culture Collection of the Plant Protection Research Institute, Agriculture Research Center, Pretoria, South Africa; XJAU: Xinjiang Agricultural University, Xinjiang, China; NA: not applicable. All the new isolates used in this study are in bold and the type materials are marked with T.

Results

Phylogenetic analyses

Each gene region and the combined matrix of five gene sequences of Cytospora were both considered. The concatenated alignment comprised sequences from 296 strains and Diaporthe vaccinii CBS 160.32 was selected as the outgroup. Cytospora ingroup strains with a total of 3166 characters including gaps (615 characters for ITS, 344 for act, 731 for rpb2, 811 for tef1-α and 665 for tub2). ML bootstraps (ML BS ≥ 60%) and Bayesian posterior probabilities (BPP ≥ 0.90) have been shown above the branches (Fig. 2). For ML analysis, the substitution model (GTR+G+I model) for each dataset was selected following recent studies (Fan et al. 2020; Pan et al. 2020, 2021). Confidence levels for the nodes were determined using 1,000 replicates of bootstrapping methods (Hillis and Bull 1993). The matrix had 1992 distinct alignment patterns. Estimated base frequencies are as follows: A = 0.244402, C = 0.286560, G = 0.238889, T = 0.230150; substitution rates: AC = 1.282426, AG = 3.546575, AT = 1.431177, CG = 0.946427, CT = 6.172877, GT = 1.000000; gamma distribution shape parameter: α = 0.364165. For BI analysis, the best-fit model of nucleotide evolution was deduced on the AIC (ITS and act: GTR+I+G; rpb2 and tef1-α: TrN+I+G; and tub2: HKY+I+G).

Figure 2. 

Phylogram of Cytospora based on Maximum Likelihood (ML) analysis of the dataset of combined ITS, act, rpb2, tef1-a and tub2 genes. Numbers above the branches indicate ML bootstrap values (ML-BS ≥ 60%) and Bayesian Posterior Probabilities (BPP ≥ 0.9). Ex-type isolates are in bold. lsolates in this study marked with its hosts and highlighted in two different colours where the novel species are shown in dark blue and the known species are shown in light blue.

The topologies resulting from ML and BI analyses of the concatenated dataset were similar. In the present study, 22 isolates formed seven clades representing seven species, of which four clades were grouped with the strains of four known species (C. ailanthicola, C. albodisca, C. euonymina, C. haidianensis). Isolates in other three clades were separated from all other species and were also highly supported (ML/BI = 100/1) (Fig. 2), representing three new species (C. fengtaiensis, C. pinea, C. sorbariae), which have been described below.

Taxonomy

Cytospora ailanthicola X.L. Fan & C.M. Tian, Persoonia 45: 13 (2020)

Fig. 3

Description

Sexual morph : not observed. Asexual morph: Conidiomata pycnidial, immersed in the bark, scattered, producing black area on bark, circular to ovoid, with multiple locules, occasionally slightly erumpent through the surface. Conceptacle absent. Ectostromatic disc inconspicuous, grey to black, circular to ovoid, producing one ostiole per disc when mature. Ostiole in the centre of the disc, black, 50–110 µm in diam. Locules numerous, subdivided frequently by invaginations with common walls, circular to ovoid, 300–500 µm in diam. Conidiophores hyaline, unbranched, approximately cylindrical, 6.5–9 × 1–1.5 (av. = 8 ± 1.5 × 1.3 ± 0.2, n = 50) µm. Conidiogenous cells enteroblastic, phialidic. Conidia hyaline, elongate-allantoid, smooth, aseptate, 2.8–3 × 0.8–1.2 (av. = 3 ± 0.3 × 1 ± 0.2, n = 50) µm.

Figure 3. 

Cytospora ailanthicola from Salix matsudana (BJFC CF20230400) A, B habit of conidiomata on branch C transverse section through conidiomata D longitudinal section through conidiomata E, F conidiophores and conidiogenous cells G conidia. Scale bars: 1 mm (A); 200 µm (B); 100 µm (C, D); 10 µm (E–G)

Culture characteristics

Cultures on PDA are initially white, growing fast up to 5 cm after 3 d and entirely covering the 6 cm Petri dish after 7 d, with fluffy and whitish aerial mycelium, producing black pycnidia with cream to yellowish conidial drops exuding from the ostioles after 30 d. Pycnidia aggregated on surface.

Materials examined

China, Beijing City, Fengtai Distinct, Qianling Mountain scenic area, 39°51'12.28"N, 116°5'17.74"E, from branches of Salix matsudana, 12 Apr 2023, A.L. Jia & X.L. Fan (BJFC CF20230400, living culture CFCC 59446).

Notes

Cytospora ailanthicola was first observed on branches of Ailanthus altissima in China by Fan et al. (2020). Lin et al. (2022) confirmed this species was a pathogen with strong virulence caused by poplar canker disease. In this study, CFCC 59446 was isolated from symptomatic branches of Salix matsudana in Beijing, which clustered in a well-supported clade with C. ailanthicola ex-holotype CFCC 89970 (ML/BI = 100/1). Therefore, CFCC 59446 is identified as C. ailanthicola.

Cytospora albodisca M. Pan & X.L. Fan, Front. Plant Sci. 12 (636460): 3 (2021).

Fig. 4

Description

Sexual morph : not observed. Asexual morph: Conidiomata pycnidial, semi-immersed in the bark, scattered, producing black area on bark, circular to ovoid, with multiple locules, occasionally slightly erumpent through the surface. Conceptacle absent. Ectostromatic disc conspicuous, black, discoid, circular to ovoid, 680–1200 µm in diam., producing one ostiole per disc when mature. Ostiole grey to black, in the centre of the disc, 140–300 µm in diam. Locules numerous, subdivided frequently by invaginations with common walls, circular to ovoid, 500–1200 µm in diam. Conidiophores hyaline, unbranched, approximately cylindrical, 7–11× 0.8–2 (av. = 9 ± 2.2 × 1.3 ± 0.3, n = 50) µm. Conidiogenous cells enteroblastic, phialidic. Conidia hyaline, elongate-allantoid, smooth, aseptate, 5–7 × 1–2 (av. = 6 ± 0.5 × 1.5 ± 0.3, n = 50) µm.

Culture characteristics

Cultures on PDA are initially white, growing fast up to 5 cm in diam. after 3 d and entirely covering the 6 cm Petri dish after 5 d, becoming dark herbage green to dull green after 7–10 d. Colonies are sparse in the centre and compact to the margin. After 30 d, pycnidia distributed irregularly on surface.

Figure 4. 

Cytospora albodisca from Euonymus japonicus (BJFC CF20230402) A, B habit of conidiomata on branch C transverse section through conidiomata D longitudinal section through conidiomata E conidiophores and conidiogenous cells F conidia. Scale bars: 2 mm (A); 1 mm (B); 500 µm (C); 200 µm (D); 10 µm (E, F).

Materials examined

China, Beijing City, Fengtai Distinct, Qianling Mountain scenic area, 39°51'12.28"N, 116°5'17.74"E, from branches of Malus × micromalus, 12 Apr 2023, A.L. Jia & X.L. Fan (BJFC CF20230401, living culture CFCC 59467); Qianling Mountain scenic area, 39°51'12.28"N, 116°5'17.74"E, from branches of Euonymus japonicus, 12 Apr 2023, A.L. Jia & X.L. Fan (BJFC CF20230402, living culture CFCC 59537).

Notes

Cytospora albodisca was described by Pan et al. (2021) associated with canker disease of Platycladus orientalis in China. It can be identified by having ascostroma surrounded by a black conceptacle, producing allantoid, aseptate ascospores (8–14 × 2–3.5 µm). In this study, the asexual morph of Cytospora albodisca is characterised by the pycnidial stromata submerged in the cortex with multiple locules, filamentous conidiophores producing hyaline, allantoid, eguttulate and smooth conidia. Phylogenetically, the isolates (CFCC 59459 and 59537) clustered together with C. albodisca with high statistical support (ML/BI = 100/1) (Fig. 2). Therefore, the isolate in this study was confirmed to be C. albodisca.

Cytospora euonymina X.L. Fan & C.M. Tian, Persoonia 45: 21 (2020)

Fig. 5

Description

Sexual morph : not observed. Asexual morph: Conidiomata pycnidial, immersed in the bark, scattered, producing black area on bark, erumpent through the surface, with multiple locules. Conceptacle absent. Ectostromatic disc honey to dark mouse grey, conspicuous, circular to ovoid, 200–500µm in diam, with one ostiole per disc. Ostiole in the centre of the disc, black, conspicuous, 80–200 μm diam. Locules numerous, subdivided frequently by invaginations with common walls, 400–750 µm in diam. Conidiophores borne along the locules, hyaline, unbranched or occasionally branched at the base or in the middle, thin-walled, 8–12 × 1.5–2 (av. = 10 ± 2.1 × 1.8 ± 0.3, n = 50) μm, embedded in a gelatinous layer. Conidiogenous cells enteroblastic, phialidic. Conidia hyaline, elongate-allantoid, smooth, aseptate, 5–7 × 1–2 (av. = 6 ± 0.5 × 1.5 ± 0.3, n = 50) µm.

Culture characteristics

Cultures on PDA are initially white, irregular, lacking aerial mycelium, fast growing up to 5 cm diam. after 3 d. Colonies pale white to light salmon after 30 d, pycnidia distributed sparsely over the medium surface.

Figure 5. 

Cytospora euonymina from Salix babylonica (BJFC CF20230403) A, B habit of conidiomata on branch C transverse section through conidiomata D longitudinal section through conidiomata E, F conidiophores and conidiogenous cells G conidia. Scale bars: 500 µm (A); 200 µm (B); 100 µm (C, D); 10 µm (E–G).

Materials examined

China, Beijing City, Fengtai Distinct, Qianling Mountain scenic area, 39°51'12.28"N, 116°5'17.74"E, from branches of Salix babylonica, 12 Apr 2023, A.L. Jia & X.L. Fan (BJFC CF20230403, living culture CFCC 59444; BJFC CF20230404, living culture CFCC 59479).

Notes

Cytospora euonymina was isolated from Euonymus kiautschovicus in Shanxi Province, China (Fan et al. 2020). It is characterised by having pycnidia covered by the darkened cuticle. Lin et al. (2023b) reported this species from leaves of Euonymus japonicus. In this study, two isolates grouped together with C. euonymina in ML and BI trees (ML/BI = 98/1). Therefore, they were identified as C. euonymina. Additionally, CFCC 59444 and 59479 extends its host range which were isolated from branches of Salix babylonica in the current study.

Cytospora fengtaiensis A.L. Jia & X.L. Fan, sp. nov.

MycoBank No: 850894
Fig. 6

Etymology

Named after the place where it was first collected, Fengtai District, Beijing City.

Typification

China. Beijing City, Fengtai District, Qianling Mountain scenic area, 39°51'12.28"N, 116°5'17.74"E, from branches of Acer palmatum ‘Atropurpureum’, 7 Apr 2023, A.L. Jia & X.L. Fan (holotype BJFC CF20230405, ex-holotype living culture CFCC 59449); 39°51'12.51"N, 116°5'17.32"E, from branches of Acer palmatum ‘Atropurpureum’, 7 Apr 2023, A.L. Jia & X.L. Fan (paratype BJFC CF20230406, ex-paratype living culture CFCC 59442.

Description

Sexual morph : not observed. Asexual morph: Conidiomata pycnidial, immersed in the bark, scattered, producing black area on bark, circular to ovoid, with multiple locules, occasionally slightly erumpent through the surface. Conceptacle absent. Ectostromatic disc conspicuous, grey to black, discoid, circular to ovoid, 180–250 µm in diam., producing one ostiole per disc when mature. Ostiole grey to black, nearly at the same level as the disc surface, 70–105 µm in diam. Locules numerous, subdivided frequently by invaginations with common walls, circular to ovoid, 560–800 µm in diam. Conidiophores hyaline, unbranched, approximately cylindrical, 11–17 × 1.5–2 (av. = 14.7 ± 2.7 × 1.6 ± 0.3, n = 50) µm. Conidiogenous cells enteroblastic, phialidic. Conidia hyaline, elongate-allantoid, smooth, aseptate, 5–6 × 1–2 (av. = 5.5 ± 0.5 × 1.6 ± 0.2, n = 50) µm.

Culture characteristics

Cultures on PDA are initially white to pale vinaceous, growing slowly up to 3 cm after 3 d and entirely covering the 6 cm Petri dish after 7 d, becoming fawn after 14 d. Colonies are flat with a uniform texture, Colony margin irregular. After 30 d, pycnidia aggregated on surface.

Figure 6. 

Cytospora fengtaiensis from Acer palmatum ‘Atropurpureum’ (BJFC CF20230405) A, B habit of conidiomata on branch C transverse section through conidiomata D longitudinal section through conidiomata E conidiophores and conidiogenous cells F conidia. Scale bars: 1 mm (A); 200 µm (B–D); 10 µm (E, F).

Additional materials examined

China. Beijing City, Fengtai District, Qianling Mountain scenic area, 39°51'11.45"N, 116°5'15.36"E, from branches of Acer palmatum ‘Atropurpureum’, 7 Apr 2023, A.L. Jia & X.L. Fan (BJFC CF20230407, living culture CFCC 59525; BJFC CF20230408, living cultures CFCC 59526 and 59527).

Notes

Cytospora fengtaiensis is associated with canker disease of Acer palmatum ‘Atropurpureum’ in the current study. It can be identified by its conidiomata producing larger black areas on bark. Phylogenetically, five isolates in this study formed a distinct lineage in the phylogenetic trees of each individual gene (ITS, act, rpb2, tef1-α and tub2) and the combined gene dataset (Fig. 2).

Cytospora haidianensis X. Zhou & X.L. Fan, Forests 11: 524 (2020)

Fig. 7

Description

Sexual morph : not observed. Asexual morph: Conidiomata pycnidial, immersed in the bark, scattered, producing black area on bark, circular to ovoid, with multiple locules, occasionally slightly erumpent through the surface. Conceptacle absent. Ectostromatic disc isabelline to dark brick, conspicuous, circular to ovoid, 130–350 µm in diam, with one ostiole per disc. Ostiole in the centre of the disc, black, conspicuous, 90–180 µm in diam. Locules numerous, subdivided frequently by invaginations with common walls, circular to ovoid, 500–1200 µm in diam. Conidiophores hyaline, branched at the base or unbranched, approximately cylindrical, 12–19 × 1–1.5 (av. = 15.5 ± 4.3 × 1.1 ± 0.4, n = 50) µm. Conidiogenous cells enteroblastic, phialidic, subcylindrical to cylindrical. Conidia hyaline, elongate-allantoid, smooth, aseptate, thin-walled, 4.8–6 × 1.5–2 (av. = 5.3 ± 0.7 × 1.7 ± 0.3, n = 50) µm.

Cultural characteristics

Colonies on PDA are initially white after 3 d, becoming light brown after 14 d. The colonies are thin with a uniform texture, lacking aerial mycelium. Pycnidia were randomly observed on the surface of the colony after 30 d.

Figure 7. 

Cytospora haidianensis from Salix babylonica (BJFC CF20230411) A, B habit of conidiomata on branch C transverse section through conidiomata D longitudinal section through conidiomata E, F conidiophores and conidiogenous cells G conidia. Scale bars: 1 mm (A); 200 µm (B–D); 10 µm (E–G).

Materials examined

China, Beijing City, Fengtai Distinct, Beigong National Forest Park, 39°52'20.46"N, 116°7'47.60"E, from branches of Euonymus japonicus, 12 Apr 2023, A.L. Jia & X.L. Fan (BJFC CF20230409, living culture CFCC 59450); Beigong National Forest Park, 39°52'20.46"N, 116°7'47.60"E, from branches of Malus ‘American’, 12 Apr 2023, A.L. Jia & X.L. Fan (BJFC CF20230410, living culture CFCC 59475); Century Forest Park, 39°49'43"N, 116°14'27"E, from branches of Acer pictum subsp. mono, 18 May 2023, A.L. Jia & Y.X. Li (BJFC CF20230411, living culture CFCC 59471; BJFC CF20230412, living culture CFCC 59536).

Notes

Cytospora haidianensis was first introduced by Zhou et al. (2020) and which was isolated from Euonymus alatus in Beijing, China. This species has numerous locules with a central column of ostiolar tissue (Zhou et al. 2020). In this study, four isolates grouped together with C. haidianensis in ML and BI trees (ML/BI = 100/1). Therefore, they are identified as Cytospora haidianensis. The current study extends its host range to Buxus megistophylla, Malus ‘American’ and Acer pictum subsp. mono.

Cytospora pinea A.L. Jia & X.L. Fan, sp. nov.

MycoBank No: 850895
Fig. 8

Etymology

Named after the host genus on which it was collected, Pinus.

Typification

China, Beijing City, Fengtai Distinct, Lotus Pond Park, 39°53'27.64"N, 116°18'49.21"E, from branches of Pinus bungeanae, 9 Feb 2023, X.L. Fan (holotype BJFC CF20230413, ex-holotype living culture CFCC 59521; 39°53'27.21"N, 116°18'49.56"E, from branches of Pinus bungeanae, 9 Feb 2023, X.L. Fan (paratype BJFC CF20230415, ex-paratype living culture CFCC 59523).

Description

Sexual morph : not observed. Asexual morph: Conidiomata pycnidial, immersed in bark, scattered, nearly flat, slightly erumpent through the bark surface in a large area, with multiple locules. Conceptacle absent. Ectostromatic disc light brown to black, inconspicuous, circular to ovoid, with one ostiole per disc. Ostiole black, conspicuous, 150–200 μm diam. Locules numerous, irregular, subdivided frequently by invaginations with common walls, 980–1130 µm diam. Conidiophores borne along the locules, hyaline, branched at the base, in the middle or unbranched, thin-walled, 15–22 × 1.5–2.5 μm (av. = 18 ± 2.3 × 2 ± 0.3 μm, n = 30), embedded in a gelatinous layer. Conidiogenous cells enteroblastic, phialidic, sub-cylindrical, 3–7.5(–8) × 1–2 μm (av. = 4.5 ± 1.4 × 1.6 ± 0.3 μm, n = 50), tapering towards apices; arranged in rosettes. Conidia hyaline, allantoid, eguttulate, smooth, aseptate, thin-walled, 3.5–5 × 1–2 μm (av. = 4.3 ± 0.5 × 1.4 ± 0.2 μm, n = 50).

Culture characteristics

Cultures on PDA are initially white, growing slowly up to 2 cm in diam. after 3 d and becoming yellowish after 7–10 d. Colonies thin with a uniform texture, lacking aerial mycelium, entirely covering the 6 cm Petri dish after 14 d, with a regular edge. After 30 d, pycnidia irregularly distributed on culture surface.

Figure 8. 

Cytospora pinea from Pinus bungeanae (BJFC CF20230413) A, B habit of conidiomata on branch C transverse section through conidiomata D longitudinal section through conidiomata E conidiophores and conidiogenous cells F conidia. Scale bars: 2 mm (A); 200 µm (B, D); 100 µm (C); 10 µm (E, F).

Additional materials examined

China, Beijing City, Fengtai Distinct, Lotus Pond Park, 39°53'26.87"N, 116°18'43.46"E, from branches of Pinus bungeanae, 9 Feb 2023, X.L. Fan (BJFC CF20230414, living culture CFCC 59522; BJFC CF20230416, living culture CFCC 59524).

Notes

Cytospora pinea is associated with canker disease of Pinus bungeanae in China. Cytospora pinea is close to C. bungeanae in the phylogenetic diagram (Fig. 2) and was isolated from the same host species Pinus bungeanae (Fan et al. 2020). It can be distinguished from C. bungeanae by smaller conidiophores (3–7.5(–8) × 1–2 vs. 15–27(–30) × 1.5–2 μm in C. bungeanae) and smaller locules (980–1130 vs. (1150–)1220–1480(–1600) μm in C. bungeanae). Furthermore, Cytospora pinea has a black conspicuous ostiole per disc, whereas the ostiole of C. bungeanae is inconspicuous. Phylogenetically, there are differences of 76/344 in the act region and 7/811 in the tef1-α gene with gaps.

Cytospora sorbariae A.L. Jia & X.L. Fan, sp. nov.

MycoBank No: 850896
Fig. 9

Etymology

Named after the host genus on which it was collected, Sorbaria.

Typification

China. Beijing City, Fengtai District, Beijing Garden Expo, 39°52'35.65"N, 116°11'4.02"E, from branches of Sorbaria sorbifolia, 7 Apr 2023, A.L. Jia & X.L. Fan (holotype BJFC CF20230417, ex-holotype living culture CFCC 59445); 39°52'35.43"N, 116°11'4.62"E, from branches of Sorbaria sorbifolia, 7 Apr.2023, A.L. Jia & X.L. Fan (paratype BJFC CF20230419, ex-paratype living culture CFCC 59529).

Description

Sexual morph : not observed. Asexual morph: Conidiomata pycnidial immersed in the bark, scattered, erumpent through the surface of bark in a large area, with multiple locules. Conceptacle absent. Ectostromatic disc brown to black, circular to ovoid, erumpent through the surface of bark in a large area, conspicuous when mature, 160–300 µm in diam., with one or two ostioles per disc. Ostioles grey to black, at the same or slightly above the level of the disc surface, 50–85 µm in diam. Locules numerous, subdivided frequently by invaginations with common walls, circular to ovoid, 550–750 µm in diam. Conidiophores hyaline, unbranched, approximately cylindrical, 14–18 × 1–1.5 µm. Conidiogenous cells enteroblastic, phialidic. Conidia hyaline, elongate-allantoid, smooth, aseptate, 5.5–7.5 × 1.5–2.5 (av. = 6.5 ± 0.7 × 2 ± 0.3, n = 50) µm.

Culture characteristics

Cultures on PDA are initially white, growing fast up to cover the 5.5 cm Petri dish after 3 d, becoming vinaceous buff after 7–10 d. Colonies are flat with a uniform texture, lacking aerial mycelium. Colony margin regular. After 30 d, pycnidia distributed irregularly on surface.

Figure 9. 

Cytospora sorbariae from Sorbaria sorbifolia (BJFC CF20230417) A, B habit of conidiomata on branch C transverse section through conidiomata D longitudinal section through conidiomata E, F conidiophores and conidiogenous cells G conidia. Scale bars: 1 mm (A); 100 µm (B–D); 10 µm (E–G).

Additional materials examined

China. Beijing City, Fengtai District, Beijing Garden Expo, 39°52'35.10"N, 116°11'4.31"E, from branches of Sorbaria sorbifolia, 7 Apr 2023, A.L. Jia & X.L. Fan (BJFC CF20230418, living culture 59443; BJFC CF20230420, living culture 59530).

Notes

Cytospora sorbariae is associated with canker disease of Sorbaria sorbifolia in the current study. It can be identified by having conidiomata with a column lenticular tissue in the centre and its distinct disc of stromata on branches. Additionally, the four strains are phylogenetically separated from all other available strains included in this study. The clear multi-gene phylogram placed it in a distinct clade with high support (ML/BI = 100/1, Fig. 2).

Discussion

The present study identified seven Cytospora species (C. ailanthicola, C. albodisca, C. euonymina, C. fengtaiensis sp. nov., C. haidianensis, C. pinea sp. nov. and C. sorbariae sp. nov.) from symptomatic branches and twigs associated with canker and dieback disease. This study represents an investigation of Cytospora species associated with canker disease in Fengtai District, Beijing and included a comprehensive analysis of DNA sequence data to compare the novelties with known Cytospora species.

In recent years, the study of Cytospora species on a particular host has received much attention from experts. For example, Jiang et al. (2020) identified six Cytospora species on Chinese chestnut (Castanea mollissima) which proved that Cytospora canker is a common disease on chestnut trees. Lin et al. (2023a) revealed the presence of Cytospora species from Populus in China and confirmed Cytospora ailanthicola, C. chrysosperma, C. paratranslucens and C. sophoriopsis as pathogens by pathogenicity tests. In this study, Cytospora species has a high diversity on Malus spectabilis and Euonymus japonicus (Cytospora albodisca and C. haidianensis). There are many studies about Cytospora related to E. japonicus, while few studies on Malus spectabilis have been recorded (Lin et al. 2023b). Therefore, many varieties of Malus spectabilis associated with Cytospora species need a systematic study and their pathogenicity is required to be confirmed in the future.

Cytospora included both generalist pathogens and specialist pathogens (Lawrence et al. 2018). Most Cytospora species have been discovered in a wide range of hosts (Adams et al. 2005, 2006; Lawrence et al. 2018; Norphanphoun et al. 2018; Fan et al. 2020). In this study, Cytospora sorbariae and C. fengtaiensis were introduced as two new species from the single host species, so more exhaustive sampling from other regions of the world is needed in future studies for a clear elucidation of their host ranges and distribution.

In this article, seven species, associated with Cytospora disease, were identified in Fengtai District, Beijing. A targeted prevention and treatment strategy is needed to be drawn up. The occurrence of Cytospora canker and dieback diseases can be minimised by removing dead and dying branches in the dry season and maintaining susceptible trees as strong as possible. Moreover, the occurrence of Cytospora canker diseases is affected by the environment, distribution and transmission (Fan et al. 2015b), which may act as potential inoculum sources for other hosts in natural and artificial environments.

This study focused on Cytospora species in Fengtai District of Beijing, an attractive location with a high richness of fungal species (Zhu et al. 2018b, 2019). The descriptions and molecular data of Cytospora in this study could provide a resource for future studies in this genus and lay the foundation for the future investigation of canker disease caused by Cytospora species.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was funded by the National Natural Science Foundation of China (32101533), National Science and Technology Fundamental Resources Investigation Program of China (2021FY100900).

Author contributions

Conceptualisation: XF, AJ. Formal analysis: BC, AJ. Funding acquisition: XF. Investigation: XF, AJ, HL. Methodology: AJ. Resources: YX, BL, XF. Software: AJ, XF. Supervision: XF. Validation: AJ, HL. Visualisation: AJ. Writing - original draft: AJ. Writing - review and editing: XF.

Author ORCIDs

Aoli Jia https://orcid.org/0009-0004-0265-5454

Xinlei Fan https://orcid.org/0000-0002-4946-4442

Data availability

All of the data that support the findings of this study are available in the main text.

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