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Morphology of Cytospora nivea from Salix psammophila (BJFU-S979). a, d: Habit of conidiomata on twig. b, c: Longitudinal sections through conidiomata. e, f: Transverse sections through conidiomata. g: Conidiophores. h: Conidia. i: Colonies on PDA at 3 days (left) and 30 days (right). j: Habit of ascomata on twig. k: Longitudinal sections through ascomata. l: Disc of ascomata on twig. m: Transverse sections through ascomata. n: Asci. o: ascospores. p: Colonies on PDA at 3 days (left) and 30 days (right). Scale bars: a, j=1 mm; b–f, k–m=0.5 mm; g=20 μm; h=5 μm; n–o=10 μm. 

Morphology of Cytospora nivea from Salix psammophila (BJFU-S979). a, d: Habit of conidiomata on twig. b, c: Longitudinal sections through conidiomata. e, f: Transverse sections through conidiomata. g: Conidiophores. h: Conidia. i: Colonies on PDA at 3 days (left) and 30 days (right). j: Habit of ascomata on twig. k: Longitudinal sections through ascomata. l: Disc of ascomata on twig. m: Transverse sections through ascomata. n: Asci. o: ascospores. p: Colonies on PDA at 3 days (left) and 30 days (right). Scale bars: a, j=1 mm; b–f, k–m=0.5 mm; g=20 μm; h=5 μm; n–o=10 μm. 

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Cytospora species are important phytopathogens causing severe canker disease with a worldwide distribution and broad host range. However, identification of taxa to species level is difficult due to poor phylogenetic understanding and lack of sequenced type species. Morphological and phylogenetic studies have been carried out on several important ho...

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... nivea (Hoffm.) Sacc., Michelia 2: 264 (1881) (Fig. 7) = Sphaeria nivea Hoffm., Veg. Crypt mm; b-f, k-m=0.5 mm; g=20 μm; h=5 μm; n-o=10 ...

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... Colonies on PDA flat, with flocculent aerial mycelium and entire edge, initially white, becoming luteous and reaching 80 mm diameter after 10 days at 25 °C, sterile. (Farr 1973;Fan et al. 2015;Gafforov 2017). This fungus is distinguished from the other Cytospora species by its 4-ascospored asci and undiscovered asexual state . ...
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During our biodiversity investigations in Tibet, China, typical Cytospora canker symptoms were observed on branches of hosts Myricaria paniculate, Prunus cerasifera and Sibiraea angustata. Samples were studied, based on morphological features coupled with multigene phylogenetic analyses of ITS, act, rpb2, tef1 and tub2 sequence data, which revealed two new species (Cytospora myricicolasp. nov. and C. sibiraeicolasp. nov.) and a known species (C. populina). In addition, Cytospora populina is newly discovered on the host Prunus cerasifera and in Tibet.
... Species of Cytospora were primarily identified and distinguished by their morphology and host [5][6][7]. However, recent studies employing molecular phylogeny revealed many cryptic species with similar morphology on the same host of known species of this genus [11][12][13][14][15]. For example, up to 28 Cytospora species were discovered from Eucalyptus spp. in South Africa with the help of DNA sequence evidence [2], eight from willow (Salix spp.) trees in China [16], six from Castanea mollissima in China [17], six from Populus hosts in China [18] and six from apple trees in Iran [19]. ...
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Cytospora (Cytosporaceae, Diaporthales) is a fungal genus that usually inhabits plants as endophytes, saprobes, as well as pathogens. Species of this genus are characterized by possessing allantoid hyaline conidia and ascospores. Samples with typical Cytospora canker symptoms on Prunus davidiana, P. padus and Salix sp. were collected in Tibet and Xinjiang, China. Species were identified using both morphological and molecular approaches of combined loci of internal transcribed spacer region rDNA (ITS), the partial actin (act) region, RNA polymerase II second largest subunit (rpb2), the translation elongation factor 1-alpha (tef1) gene and the partial be-ta-tubulin (tub2) gene. Six isolates in the present study formed three distinct clades from previously known species. Cytospora hejingensis sp. nov. from Salix sp., C. jilongensis sp. nov. from P. davidiana and C. kunsensis from P. padus were proposed herein. The current study improves the understanding of species concept in Cytospora.
... 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. ...
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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.
... Cytospora chrysosperma was subsequently selected as the type species [11]. Species in this genus are morphologically similar in having allantoid hyaline conidia and ascospores [12][13][14]. During species identification, host association and spore morphology were important than molecular phylogeny [15][16][17]. ...
... The present study revealed a new collection of C. euonymina from E. japonicus in Tibet, which is associated with branch and twig canker disease. Notes: Cytospora gigaspora has been recorded from Salix psammophila in Shaanxi Province and Juniperus procumbens in Shanxi Province, which are in North China [9,14]. In the present study, we isolated a pure culture of C. gigaspora from diseased branches of Larix gmelinii in Tibet. ...
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Branch canker diseases are important in forest ecosystems, causing economic and ecological losses. Members of Cytospora are common on cankered branches and associated with the diseases. Investigations on Cytospora cankers were conducted in Tibet, China, during 2022 and 2023. Samples were collected from Euonymus japonicus, Larix gmelinii, Malus pumila, M. spectabilis and Rosa omeiensis f. pteracantha, and cultures were obtained and identified by morphological features and molecular phylogeny of a combination of internal transcribed spacer region rDNA (ITS), the partial actin (act) region, RNA polymerase II second largest subunit (rpb2) gene, the translation elongation factor 1-alpha (tef1) gene and the partial beta-tubulin (tub2) gene. As a result, a new species is proposed herein named Cytospora lhasaensis and four known species are described for the first time from Tibet, viz. C. euonymina, C. gigaspora, C. mali and C. schulzeri. The current research enhances our understanding of the Cytospora species associated with woody host diseases in Tibet, China.
... Ostiole circular to ovoid, olivaceous buff to dark mouse grey, at the same level or above as disc surface, 66-98 µ Notes: Cytospora zhaitangensis is phylogenetically most closely related to C. euonymicola and C. gigalocus. Morphologically, C. zhaitangensis can be differentiated by the wider conidia from C. euonymicola (L/W ratio = 3.48 vs. L/W ratio = 4.5) and C. gigalocus (L/W ratio = 4.36) [32,86]. Additionally, C. zhaitangensis has smaller discs (166-224 µ m) than C. euonymicola (240-350 µ m) and C. gigalocus (330-620 µ m) [32,86]. ...
... Morphologically, C. zhaitangensis can be differentiated by the wider conidia from C. euonymicola (L/W ratio = 3.48 vs. L/W ratio = 4.5) and C. gigalocus (L/W ratio = 4.36) [32,86]. Additionally, C. zhaitangensis has smaller discs (166-224 µ m) than C. euonymicola (240-350 µ m) and C. gigalocus (330-620 µ m) [32,86]. Cytospora euonymi and C. euonymella were also recorded to host Euonymus [92,93]. ...
... 115°34′15′′E, 39°51′56′′N, from branches of Euonymus japonicus, M. Pan, Y.K. Bai and X.L. Fan, 21 August 2021 (BJFC CF20220127, living culture CFCC 56287).Notes: Cytospora elaeagni has been reported from Elaeagnus angustifolia in China, German, and the USA[87][88][89]. Fan et al.[86] provided its morphological descriptions and molecular data. In this study, five isolates are identified as Cytospora elaeagni based on phylogenetic analyses. ...
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Euonymus japonicus tolerates the dry and frigid climate of Beijing, China, and effectively filters out particles during the winter. However, fungal infestation frequently causes extreme illness and can even lead to shrub death. In this study, 104 diseased E. japonicus specimens were collected from seven districts in Beijing. Seventy-nine isolates were identified as 22 fungal species in seven genera. The species were Aplosporella hesperidica, A. javeedii, A. prunicola, Botryosphaeria dothidea, Colletotrichum aenigma, Co. euonymi, Co. euonymicola, Co. gloeosporioides, Cytospora ailanthicola, C. albodisca, C. diopuiensis, C. discotoma, C. elaeagni, C. euonymicola, C. euonymina, C. haidianensis, C. leucostoma, C. sophorae, C. zhaitangensis, Diaporthe eres, Dothiorella acericola, and Pestalotiopsis chaoyangensis. On the basis of morphological and phylogenetic analyses, Colletotrichum euonymi, Co. euonymicola, Cytospora zhaitangensis, and Pestalotiopsis chaoyangensis were introduced as novel species. Colletotrichum euonymi, Co. euonymicola, and Pestalotiopsis chaoyangensis were subsequently confirmed as pathogens of E. japonicus leaves by pathogenicity testing. This study provides an important assessment of the fungi associated with diseases of E. japonicus in Beijing, China.
... Cytospora hippophaes was reported as a pathogen on sea buckthorn twigs and branches associated with canker (Osipan and Martirosian, 1984;Fan et al., 2015). In northern China, C. hippophaes is associated with stem canker, and also it has been found in Australia, Spain, and Armenia (Osipan and Martirosian, 1984;Fan et al., 2015). ...
... Cytospora hippophaes was reported as a pathogen on sea buckthorn twigs and branches associated with canker (Osipan and Martirosian, 1984;Fan et al., 2015). In northern China, C. hippophaes is associated with stem canker, and also it has been found in Australia, Spain, and Armenia (Osipan and Martirosian, 1984;Fan et al., 2015). Cytospora ambiens has been reported as the cause of canker in Armenia (Osipan and Martirosian, 1984). ...
... Cytospora chrysosperma was diagnosed as the cause of stem canker on sea buckthorn in Canada (Manitoba Agriculture, Food and Rural Initiatives Crop Diagnostic Centre) (Desjardins et al., 2005). Cytospora species are important plant pathogens causing severe canker diseases worldwide and having a broad host range (Osipan and Martirosian, 1984;Fan et al., 2015). Several Cytospora species are known as causes of cankers on woody hosts, including various fruit crops (Fan et al., 2015;Kepley and Jacobi, 2000). ...
Article
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Sea buckthorn (Hippophae rhamnoides L.) is a native species in various regions of Asia and Europe. It is cultivated as a multipurpose horticultural species in northern temperate regions of Europe, Asia, and North America with large economic potential used for food, pharmacology,cosmetics, and environmental conservation. Diseases in natural populations and managed land-scapes have increased, endangering sea buckthorn growth and cultivation worldwide. This reviewarticle focuses on sea buckthorn canker, wilt and decline diseases caused by pathogenic fungi, their distribution, hosts of involved pathogenic fungi and symptoms. Published information on seabuckthorn fungal diseases is available only about a few diseases, such as wilt (Verticillium dahliae), the dried-shrink disease caused by various fungi and abiotic factors, and stem canker (Hymenopleella hippophaeicola, Cytospora spp., Stigmina sp.). Some fungi reported on seabuckthorn are poorly studied, or the sea buckthorn is a newly discovered host, as in the case of Eutypa spp. The most often reported symptoms of these diseases are cankers and cracks ontrunks and main branches, dead buds and leaves, necrosis of various tissues on branches, androot necrosis, resulting in the death of the shrubs. In general, the fungal diseases on seabuckthorn are not sufficiently addressed, and more research is needed.
... accessed on 3 August 2022). Cytospora is the asexual morph of Valsa (type genus of Valsaceae) in Diaporthales (Adams et al., 2005;Ariyawansa et al., 2015;Fan et al., 2015aFan et al., , 2015bHyde et al., 2016;Maharachchikumbura et al., 2015Maharachchikumbura et al., , 2016Wang et al., 2011). Cytospora has also been considered as the asexual morph of Leucocytospora, Leucostoma, Valsella, and Valseutypella (Adams et al., 2002(Adams et al., , 2005Bulgakov 2010;Castlebury et al., 2002;Fries 1823;Gvritishvili 1982;Hyde et al., 2016;Li et al., 2016;Maharachchikumbura et al., 2015Maharachchikumbura et al., , 2016Rossman et al., 2015;Saccardo 1884;Spielman 1985;Yang et al., 2015). ...
... However, Adams et al. (2006) synonymized all of these sexual genera under Valsa. In accordance with the recent changes in fungal nomenclature and the application of a single name for a fungus (Wingfield et al., 2012), the name Cytospora (Ehrenberg, 1818) has been preferred over Valsa (Fries, 1849) and Cytosporaceae has priority over Valsaceae (Fan et al., 2015a(Fan et al., , 2015bRossman et al., 2015). ...
... Even though Cytospora species are principally known as weak pathogens, they negatively affect the longevity and productivity of fruit trees worldwide and cause chronic wood infections on fruit trees such as Malus spp. (Adams et al., 2005(Adams et al., , 2006Azizi et al., 2020a;Biggs 1989;Bills 1996;Christensen 1940;Fan et al., 2014aFan et al., , 2014bFan et al., , 2015aFan et al., , 2015bGonz alez and Tello 2011;Lawrence et al., 2018;Mehrabi et al., 2011;Pan et al., 2020;Sinclair et al., 1987;Wang et al., 2011Wang et al., , 2020Zhu et al., 2018). Disease symptoms initially appear on trunk and branches as slightly sunken areas on bark with brown discoloration in the wood that might result in cracking and scaling-off of the bark with disease progress. ...
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In recent years, canker and die-back diseases have become a growing threat for the productivity and longevity of apple orchards in Iran. In this study, 131 Cytospora isolates were recovered from symptomatic tissues of apple trees in apple orchards of Iran. Multigene phylogenetic inference based on combined sequence data of ITS, act, rpb2, and tef1-α loci, supplemented with morphological characteristics and pathogenicity assay revealed four novel Cytospora species which were described as C. avicennae, C. azerbaijanica, C. ershadii, and C. iranica, and four known species, namely C. chrysosperma, C. parasitica, C. paratranslucens, and C. sorbicola. Also, C. sorbicola is newly reported on apple trees worldwide. Koch's postulates were fulfilled to confirm that all eight species in this study were pathogenic on apple trees in Iran, among which C. sorbicola was the most intensive species. The results of this study further highlight rich diversity among Cytospora species occurring on apple trees, revealing several novel Cytospora species on this host. The host range, geographical distribution, and economic significance of novel species on apple industry remains to be studied.
... The genus Cytospora (Cytosporaceae, Diaporthales) was established by Ehrenberg [21], which comprises important phytopathogenic, saprobic, and endophytic fungi [4,18]. Species of Cytospora inhabit a wide variety of hosts that include economically and ecologically important trees (e.g., Elaeagnaceae, Juglandaceae, Rosaceae, Salicaceae, Ulmaceae) [22][23][24][25][26]. About 150 species of Cytospora in total have been discovered on dieback and stem canker in over 130 species of woody hosts [18,[27][28][29][30][31][32][33]. ...
... Recently, Fan et al. [4] revisited ten related Cytospora species as belonging to the C. chrysosperma complex based on multigene analyses. Notes: Cytospora gigaspora was originally observed on twigs and branches of Salix psammophila in Shaanxi Province [24]. After that, Fan et al. [4] added specimens from Shanxi. ...
... In previous studies, five species (C. atrocirrhata, C. fugax, C. leucosperma, C. populina, and C. translucens) associated with willow canker disease have also been reported in China [3,4,13,24,51,57]. Cytospora atrocirrhata was first reported on branches of willow in Georgia [58], which was described as erumpent conidiomata with distinct conceptacles and single locules and ostiole [24]. ...
Article
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Species of Cytospora are considered important plant pathogens of a wide range of plant hosts, especially Salicaceae plants. Salix (Salicaceae, Malpighiales) has been widely cultivated in China because of its strong ecological adaptability, fast growth, and easy reproduction. In this study, a total of eight species of Cytospora were discovered on Salix in China, including C. ailanthicola, C. alba, C. chrysosperma, C. gigaspora, C. nivea, C. paracinnamomea, C. rostrata, and C. sophoriopsis. Among them, C. alba and C. paracinnamomea were identified as novel species based on morphology and phylogenetic analyses of ITS, act, rpb2, tef1-α, and tub2 gene sequences and were confirmed as pathogens of willow canker disease by pathogenicity tests. The mycelial growth rates of strains from these two novel species (C. alba and C. paracinnamomea) had optimum temperatures of 21 to 22 °C and an optimum pH value of 5 to 6. The effectiveness of six carbon sources on the mycelial growth showed that fructose and maltose had the highest influence. Cytospora species richness was significantly positively correlated with dry and wet areas. This study represents a significant evaluation of Cytospora associated with willow canker disease in China and provides a theoretical basis for predicting the potential risk of willow canker disease.
... Excluding the ambiguously aligned sequences from analyses. Reference sequences were retrieved from recent publications (Fan et al., 2014a;Fan et al., 2014b;Fan et al., 2015a;Fan et al., 2015b;Lawrence et al., 2017;Fan et al., 2020;Pan et al., 2020;Zhu et al., 2020;Pan et al., 2021). Diaporthe vaccinii (CBS 160.32) was included as the outgroup in Cytospora analysis (Table S2) and Diaporthella corylina (CBS 121124) was included in Diaporthe analysis (Table S3). ...
... However, this strain could be distinguished from the five most related species by the conidiomata (C. gigaspora with a flat locule, C. mali-spectabilis with a column lenticular tissue in the center, and C. nivea with a dark conceptacle) (Fan et al., 2015b;Pan et al., 2020). In addition, C. corylina has a smaller conidia size (3.5-7.5 × 1-1.7 µm) than C. gigaspora (8.9-12.1 × 1.9-2.9 ...
... × 1.5-2 µm), C. nivea (6.2-9.2 × 1.7-2.4 µm), and C. paratranslucens (6.5-7.3 × 1.3-1.5 µm) (Adams et al., 2006;Fan et al., 2015b;Norphanphoun et al., 2017;Pan et al., 2020). Therefore, we describe this species as novel based on DNA sequence data and morphology. ...
Article
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Hazelnut (Corylus heterophylla Fisch.) is an important nut crop in China but has been declining owing to the destructive effects of fungal branch canker and dieback. The identification and management of these pathogens are difficult because of the lack of attention to branch canker, insufficient understanding of phylogenetic, and overlapping morphological characteristics of the pathogens. In total, 51 strains were isolated from Chinese wild hazelnut in this study, and three species of Cytospora and two of Diaporthe were identified through morphological observation and multi-locus phylogenetic analyses (ITS, act, rpb2, tef1-α, and tub2 for Cytospora; ITS, cal, his3, tef1-α, and tub2 for Diaporthe). Three new species, Cytospora corylina, C. curvispora, and Diaporthe corylicola, and two known species, Cytospora leucostoma and Diaporthe eres, grew at 5–30°C and a pH of 3.0–11.0, with optimum growth at approximately 25°C and pH 4.0–7.0. Additionally, the effects of six carbon sources on mycelial growth were investigated. This study explored the main pathogenic fungi species of Corylus heterophylla, completed the corresponding database of pathogenic fungi information, and clarified their biological characteristics. Moreover, the results of this study provided a theoretical basis for Corylus heterophylla disease management and prevention in China.
... Notes: Cytospora gigalocus was recorded from Salix psammophila and Juniperus procumbens (Fan et al., 2015b, which was similar with C. nivea regarded as the pathogen for poplar and willow canker (Saccardo, 1884;Teng, 1963;Tai, 1979;Wei, 1979;Zhuang, 2005;Fan et al., 2014b). However, C. gigalocus differs from other Cytospora species by the flat locules and larger conidia size (10.4 × 2.2 µm) (Fan et al., 2015b). ...
... Notes: Cytospora gigalocus was recorded from Salix psammophila and Juniperus procumbens (Fan et al., 2015b, which was similar with C. nivea regarded as the pathogen for poplar and willow canker (Saccardo, 1884;Teng, 1963;Tai, 1979;Wei, 1979;Zhuang, 2005;Fan et al., 2014b). However, C. gigalocus differs from other Cytospora species by the flat locules and larger conidia size (10.4 × 2.2 µm) (Fan et al., 2015b). ...
... Lawrence et al. (2018) reported that Cytospora included generalist and specialist pathogens by taking C. chrysosperma and C. punicae as examples; however, a clear elucidation of the host ranges and distribution of Cytospora species will require a more exhaustive sampling of other coniferous trees from other regions of the world. Consistent with previous findings, some Cytospora species isolated from coniferous trees occurred on different hosts (i.e., C. ampulliformis, C. gigaspora, C. melnikii) (Fan et al., 2015bNorphanphoun et al., 2017Norphanphoun et al., , 2018Lawrence et al., 2018) rather than specific hosts. In addition, it cannot be denied that some species of Cytospora have a preference for certain hosts (i.e., C. japonica with chiefly Rosaceae host record, C. mali with apple host record and C. pini with pine host record) (Teng, 1963;Tai, 1979;Wei, 1979;Zhuang, 2005;Wang et al., 2011). ...
Article
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Cytospora species are widely distributed and often occur as endophytes, saprobes or phytopathogens. They primarily cause canker and dieback diseases of woody host plants, leading to the growth weakness or death of host plants, thereby causing significant economic and ecological losses. In order to reveal the diversity of Cytospora species associated with canker and dieback diseases of coniferous trees in China, we assessed 11 Cytospora spp. represented by 28 fungal strains from symptomatic branches or twigs of coniferous trees, i.e., Juniperus procumbens, J. przewalskii, Picea crassifolia, Pinus armandii, P. bungeana, Platycladus orientalis in China. Through morphological observations and multilocus phylogeny of ITS, LSU, act, rpb2, tef1-α, and tub2 gene sequences, we focused on four novel Cytospora species (C. albodisca, C. discostoma, C. donglingensis, and C. verrucosa) associated with Platycladus orientalis. This study represented the first attempt to clarify the taxonomy of Cytospora species associated with canker and dieback symptoms of coniferous trees in China.