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2021 Volume 6
Article Contents
Abstract
Introduction
Bioactive secondary metabolites
Prospective medicinal properties of ramaria species
Antimicrobial activity
Antioxidant activity
Anticancer activity
Other biological activities of ramaria species
Clavaria sp.
Clavaria sp. biological activities
Conclusion
Funding
Conflicts of interest
Rights and permissions
References
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ARTICLE   Open Access    

New host and geographic records of five pleosporalean hyphomycetes associated with Musa spp. (Banana)

  • 1.

    The State Phosphorus Resource Development and Utilization Engineering Technology Research Centre, Yunnan Phosphate Chemical Group Co. Ltd, Kunming 650201, China

  • 2.

    Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand

  • 3.

    School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand

  • 4.

    CAS Key Laboratory for Plant Biodiversity and Biogeography of East Asia (KLPB), Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China

  • 5.

    East and Central Asia Regional Office, World Agroforestry Centre (ICRAF), Kunming 650201, Yunnan, China

  • 6.

    Centre for Mountain Futures (CMF), Kunming Institute of Botany, Kunming 650201, Yunnan, China

  • 7.

    Honghe Center for Mountain Futures, Kunming Institute of Botany, Chinese Academy of Sciences, Honghe County 654400, Yunnan, People's Republic of China

  • 8.

    Department of Health Sciences, Faculty of Medicine and Health Sciences, University of Mauritius, Reduit, Mauritius

  • 9.

    Research Center of Microbial Diversity and Sustainable Utilization, Faculty of Sciences, Chiang Mai University, Chiang Mai 50200, Thailand

More Information
  • A study was undertaken to collect and identify saprobic fungi associated with Musa spp. (banana) from Taiwan (China), and Thailand. Samples were collected during the dry season and their morpho–molecular relationships were investigated. Five brown pleosporalean hyphomycetous taxa in Periconiaceae and Torulaceae viz. Periconia cortaderiae, P. delonicis, Torula chromolaenae, T. fici, and T. masonii were identified for the first time from Musa spp. (Musaceae). Phylogenetic analyses of a combined SSU, LSU, ITS, RPB2 and TEF DNA sequence dataset further justified the taxonomic placements of these five taxa in the above mentioned families. Periconia delonicis is reported for the first time on a monocotyledonous host and T. masonii is the first geographical record from Taiwan (China).

  • Introduction

    Wild mushrooms have been consumed since ancient times due to their good taste and nutritional values. Also, mushrooms have the advantage of being rich in vital components such as proteins, vitamins, chitin, fibers, iron, zinc, selenium, sodium, etc. (Breene 1990, Chang & Miles 1992, Elkhateeb et al. 2019, 2020a, b, Daba et al. 2020, El-Hagrassi et al. 2020). Beside these nutritional values, many mushrooms have gained importance due to their therapeutical potentials (Bobek et al. 1995, Bobek & Galbavý 1999, Vaz et al. 2011, Khatun et al. 2012, Elkhateeb & Daba 2020, 2021a, b). The extracts of fruit bodies and mycelium obtained from various fungi have shown a crucial role in the prevention and/or the treatment of many diseases (Wasser & Weis 1999, Sharma et al. 2015). Nowadays, some immunomodulating molecules that have been obtained from different medicinal mushrooms have found great importance in improving the immune function in cancer patients while they are receiving chemo or radiotherapy (Mizuno 1995). Other edible mushrooms have gained medicinal importance due to their effectiveness in adjusting the blood pressure and reducing the free plasma cholesterol, as well as due to antimicrobial, antioxidant, and antitumor properties (Kabir & Kimura 1989, Smânia et al. 1995, Mau et al. 2005, Elkhateeb 2020, Thomas et al. 2020). The genus Ramaria is found all over the world, in which various species of Ramaria have been consumed by people for nutritional purposes. However, more studies are required to be conducted on these genera in order to detect the nutraceutical and nutritional compounds of these mushrooms. The genus Ramaria belonging to Basidiomycota, Class; Agaricomycetes, Order; Gomphales, Family; Gomphaceae, which includes approximately 200 species of coral fungi (Sharma et al. 2015). Several taxa are edible and picked in Europe and also they are easily confused with several mildly poisonous species capable of causing nausea, vomiting and diarrhea, in which among the most unusually shaped mushrooms of all, Ramaria are appropriately referred to as the coral mushrooms. Ramaria are recognizable by their variably thickly branched fruit bodies, fleshy and tough textures sometimes partly gelatinous or jellylike. Although many Ramaria species are very brightly colored, just as many are dull, and all have a tendency to become rather drab in age. Similar genera include Phaeoclavulina, Lentaria, Artomyces, and Ramariopsis, which differ in various aspects of color, consistency, substrate, microscopic features, or ecology (Sharma & Gautam 2017).

    Ramaria basidiocarps are either lignicolous or terricolous (white, yellow, orange, red, brilliant purple, brown and sometimes green hues) with many species forming a characteristic mycelial mat in the soil beneath the sporocarps (Nouhra et al. 2004). The colour may fade or change after collection in the field. Species of this genus often show a color change on bruising. The flesh may change to green (virescent), red brown (rubribrunnescent), brown to yellow brown (brunnescent), or a red wine color (vinescent) due to age or environmental conditions (Exeter et al. 2006).

    Clavaria is belonging to Basidiomycota, class Agaricomycetes, order Agaricales, and family Clavariaceae. Species of Clavaria produce basidiocarps (fruitbodies) that are cylindrical to club-shaped, branched and coral-like. They are often grouped with similar looking species from other genera, and they are collectively known as the clavarioid fungi (Olariaga et al. 2015).

    The most common species of this genus is Ramaria stricta (Kuo 2009), which grows on wood debris, which features branches that are usually strictly oriented, so that they are mostly straight and ascending. When fresh, its branch tips are yellow and its branches are dull yellowish buff, but its surfaces bruise and discolor purplish brown. Under the microscope it features roughened spores, clamp connections, and thick-walled hyphae. Several very similar species have been separated by mycologists, and the name Ramaria stricta should probably represent a group of potential species awaiting contemporary study. Ecology: Uncertain; while most ramarias are thought to be mycorrhizal, the wood-inhabiting species could be mycorrhizal or saprobic; growing from the dead (but sometimes buried) wood of conifers; appearing alone, scattered, or gregariously; early summer through fall; apparently widely distributed in North America, but more common from the Rocky Mountains westward. Fruitbody: 4−14 cm high; 4−10 cm wide; base well developed or nearly absent; branching repeatedly. Branches: Vertically oriented and elongated; often flattened; smooth; yellowish buff, becoming orangish buff as the spores mature; bruising and discoloring purplish brown; tips yellow when fresh and young. Base: Nearly absent, or fairly well developed; to 2 cm wide; white below; colored like the branches above; attached to numerous white rhizomorphs. Flesh: Whitish; fairly tough. Odor and Taste: Odor not distinctive, or sweet and fragrant; taste bitter. Spore color: Rusty yellowish. Microscopic Features: Spores are yellow-brown to rusty-brown in mass deposit 7.5−10.5 × 3.5−5µm; stretched-elliptical; smooth to roughened. Clamp connections present. Thick-walled hyphae present (Kuo 2009) (Figs 1, 2).

    Figure 1.  Fruitbody of Ramaria stricta in the field. The photograph was taken at the Sewickley Heights Borough Park, PA, USA by Fluffberger (https://www.inaturalist.org/observations/3596638). The photo was used under the CC BY-NC 4.0 non-commercial use license. Scale bar = 10 cm.
    DownLoad: Full-Size Img PowerPoint
    Figure 2.  Fruitbody of Ramaria araiospora in the field. The photograph was taken at the Millersylvania State Park, Olympia, WA, USA by Yay4john (https://www.inaturalist.org/observations/62355314). The photo was used under the CC BY-NC 4.0 non-commercial use license. Scale bar = 10 cm.
    DownLoad: Full-Size Img PowerPoint

    Few reports have been found about the discovery of these genera in the Northwestern Himalayas. The current surveys to Northwestern Himalayas have shown the existence of six species of coral mushrooms belonging to the genus Ramaria viz., R. aurea, R. botrytis, R. flava, R. flavescens, R. rubripermanens, and R. stricta (Sharma & Gautam 2017). Details about their culinary status were obtained from native inhabitants. Moreover, studies have been conducted on these species to get detailed biochemical profiling as well as exploring their various biological activities (Sharma & Gautam 2017).

    Bioactive secondary metabolites

    Coral mushrooms gain great therapeutical and nutritional importance due to their valuable components. Sharma & Gautam (2017), reported that 12 edible species of Ramaria occurring in Northwestern Himalayas contain proximal components, amino acids, and minerals, fatty acids. Ramaria aurea is an edible mushroom that is considered a promising dietary supplement as it consists of carbohydrates, amino acids, proteins, fibers, and minerals (Rai et al. 2013). Ramaria botrytis, another edible mushroom, consists of unsaturated fatty acids, tocopherol as well as other bioactive compounds (e.g. ascorbic acid, carotene, flavonoids, β- and lycopene) (Barros et al. 2008). Ramaria subalpina also contains significant amounts of ascorbic acid, β-carotene, flavonoids, and lycopene (Acharya et al. 2017). Also, some Ramaria species show the presence of alkaloids, saponins, terpenoids, coumarins, and cardiac glycosides (Aldred 2008, Dattaraj et al. 2020). Due to the existence of these bioactive molecules, these mushrooms possess various medicinal importance against various diseases. Among these medicinal benefits are antimicrobial, antiviral, anti-parasitic, antioxidant, radical scavenger, anticancer, anti-inflammatory, immune system enhancer, and anti-hyperlipidemia (Wasser 2017).

    Prospective medicinal properties of Ramaria species

    Antimicrobial activity

    Although there is a vast diversity of synthetic antibacterial compounds, the development of bacterial resistance has been substantially increasing (Alves et al. 2012). Thus, discovering novel antibiotics especially from natural sources is urgently required. Since ancient times, several natural resources have exerted potent antimicrobial activities. Among these natural resources are mushrooms that might be valuable alternative sources of novel antimicrobials (Gebreyohannes et al. 2019, Ghosh et al. 2020). In this context, various Ramaria sp. have gained great importance due to their potential antimicrobial activities against various pathogens (Table 1). The ethanolic extract of Ramaria flava was detected by the agar-well diffusion method, and the result showed positive activity against some Gram-positive bacteria such as Staphylococcus aureus ATCC 25923, Staphylococcus aureus Cowan I, Micrococcus luteus NRRL B-4375, Micrococcus flavus, Bacillus subtilis ATCC 6633, Bacillus cereus RSKK 863. Also, R. flava ethanolic extract showed potent activity against the tested Gram-negative bacteria including Salmonella enteritidis RSKK 171, Yersinia enterecolitica RSKK 1501, Klebsiella pneumoniae ATCC 27736 (Gezer et al. 2006). Additionally, Ramaria cystidiophora showed antimicrobial activity against Mycobacterium smegmatis (MIC = 8 µg/mL), and against Mycobacterium tuberculosis (MIC 64−128 µg/mL) (Hassan et al. 2019).

    Table 1.  Antimicrobial activities of various Ramaria species
    Ramaria species Tested extract Employed methods Tested bacteria Tested fungi References
    Gram-positive Gram-negative
    R. formosa Ethyl acetate, methanol, and water Percentage of inhibition B. subtili, S. aureus E. coli, K. pneumonia, Proteus vulgaris, P. aeruginosa Not tested Pala et al. 2019
    R. formosa Methanol Inhibition zone assay S. aureus P. aeruginosa Candida albicans Ramesh & Pattar 2010
    R. zippellii Ethanol, water 96-well microplate bioassay S. aureus E. coli No activity Bala et al. 2011
    R. aurea Ethanol Inhibition zone assay S. aureus E. coli, P. aeruginosa, P. vulgari Candida albicans Rai et al. 2013
    R. flava Ethanol Inhibition zone assay B. subtili, S. aureus E. coli Fusarium auenaceum, F. graminearum and Cercosporella albomaculans Liu et al. 2013
    R. flava Ethanol Agar-well diffusion method S. aureus, Micrococcus luteus, M. flavus, B. subtilis, B. cereus Salmonella enteritidis, Yersinia enterecolitica and K. pneumoniae No activity Gezer et al. 2006
    R. botrytis Ethanol Methanol Acetone Ethyl acetate Inhibition zone diameter S. aureus P. aeruginosa, E. coli, Enterobacter cloacae No activity Han et al. 2016
     | Show Table
    DownLoad: CSV

    The studies also reported that the ethanolic extract of Ramaria flava exhibited antifungal activities against three pathogenic fungi which are C. albomaculans, Fusarium graminearum and Fusarium auenaceum. The extract showed the highest efficacy against F. auenaceum, where 36.64% reduction in fungal growth was observed at the concentration of 2 mg/mL. The extract also inhibited the growth of F. graminearum and C. albomaculans by 19.99 and 30.03% respectively, at the same concentration of 2 mg/mL (Liu et al. 2013). Additionally, Ramaria mushrooms show their effectiveness to cure various viral infections. Zhang et al. (2015) isolated a novel ribonuclease from Ramaria formosa fruiting bodies and tested its antiviral activity against HIV-1 reverse transcriptase enzyme. The results showed that the ribonuclease showed 93% inhibition at a concentration of 30 µM (maximum tested concentration) and with an IC50 value of 3 µM. It was interesting to report that this enzyme exhibits unique features; these include its unique N-terminal sequences, optimum acidic pH value, and temperature resistance. Taken together these features suggest that this ribonuclease could play a vital role in HIV diseases prevention.

    Antioxidant activity

    Many reports regarding the antioxidative properties of various Ramaria sp. have been published, which showed the activity of these fungi as effective antioxidants (Table 2). Linoleic acid oxidation was compared with those of α- tocopherol, BHA, and R. flava ethanol extract. The results revealed that the inhibition values of both the standards and R. flava ethanol extract increased gradually with increasing the concentrations. 80 µg/ml BHA, α -tocopherol, and R. flava ethanolic extract exhibited 96.4, 98.6, and 73.3% inhibition, respectively. Whereas increasing the concentrations to 160 µg/ml concentrations resulted in 98.9, 99.2 and 94.7% inhibition, respectively (Gezer et al. 2006). Also, the ethanolic extract of that mushroom exhibited promising OH and DPPH radical-scavenging activities with low IC50 values of 18.08 and 5.86 μg/mL, respectively (Liu et al. 2013). The high inhibition value of R. flava ethanolic extract could be related to the high concentration of phenolic compounds (Komali et al. 1999).

    Table 2.  Antioxidant activities of various Ramaria species isolated from different places
    Ramaria species Place of collection Methods used Activity (EC 50 % of inhibition) References
    R. flava Turkey DPPH scavenging assay 94.78% at 12 mg/mL Gursoy et al. 2010
    β-carotene linoleic acid assay 95.02% at 20 mg/mL
    Reducing power 95.02% at 20 mg/mL
    Metal chelating effect 96.75 at 2 mg/mL
    R. flava Turkey DPPH scavenging assay 276 µg/mL Gezer et al. 2006
    β-carotene linoleic acid assay 94.7% at 160 µg/mL
    R. formosa India DPPH radical scavenging activity 5.8 mg/ml Ramesh & Pattar 2010
    R. patagonica Argentina DPPH scavenging assay 770 µg/mL Toledo et al. 2016
    β-carotene linoleic acid assay 610 µg/mL
    Reducing power 170 µg/mL
    TBARS inhibition activity 60 µg/mL
    R. Formosa Korea DPPH scavenging assay 117 AsA/mg/mL at 500 µg/mL Kim et al. 2016
    Reducing power 36% copper ion inhibition at 20 µg/mL concentration
    Peroxyl radical scavenging activity 7.8 µM trolox equivalent at 20 µg/mL
    R. stricta Ukraine Total antioxidant status 4.223±0.054 mmol/l Krupodorova & Sevindik 2020
    Oxidative stress index 0.194±0.001
    Total oxidant status 8.201 ± 0.095 μmol/l
     | Show Table
    DownLoad: CSV

    ABTS assay is another assay employed to evaluate the free radical scavenging ability resulted from hydrogen-donating ability (Re et al. 1999). A study reported that R. largentii extract was able to scavenge ABTS radical cation in a concentration-dependent manner. At an extract concentration of 250 mg/mL the scavenging activity was recorded to be 93.53 ± 0.03% (Aprotosoaie et al. 2017). A polyphenol-rich extract of R. aurea exhibited superoxide and DPPH radicals scavenging ability with EC50 of about 0.283 and 0.384 mg/mL, respectively (Khatua et al. 2015). Thus, Ramaria species extract seems to be a reasonably free radical scavenger agent.

    Anticancer activity

    Cancer is one of the main causes of death worldwide. Thus, finding new molecules to prevent and/or treat cancer, especially from natural sources with lower toxicity, is considered a goal for scientists nowadays (Wang et al. 2012). As mentioned previously, mushrooms are sources of powerful pharmaceutical products. About 651 higher basidiomycetes species exhibit antitumor activities (Fan et al. 2006). In general, fungal anticancer substances are divided into two main groups which are low and high molecular-weight compounds. The low-molecular-weight secondary compounds comprise mainly sesquiterpenes, steroids and sterols, triterpenes, and polyketides (Mahajna et al. 2008). These molecules can penetrate the cell membrane and work on the targeted signal-transduction pathways (Zaidman et al. 2005). However, the high-molecular-weight compounds include polysaccharides or protein-bound polysaccharides (Kidd 2000). In 1982, a study reported by Yoo et al. (1982), examined the antitumor activity of a high molecular weight protein-bound polysaccharide fraction obtained from Ramaria formosa. These molecules were able to inhibit 66% of a tumor when administered at the dosage of 50 mg/kg/day and the tumor was wholly degenerated in two out of eight mice.

    The growth inhibitory effect of the ethanol extract obtained from Ramaria flava was tested on human cancer cell lines (BGC-803, NCI-H520, and MDA-MB-231) and the results showed a promising inhibition activity on the three human cancer cells with inhibition percentages of 33.83%, 54.63%, and 71.66% respectively, at mushroom extract concentration of 200 µg/mL. A major antitumor sterol found in many edible mushrooms is ergosterol peroxide is (Lindequist et al. 2005). Previous studies reported the antitumor properties of ergosterol peroxide in various cancer cells including SCC4 (head and neck squamous cell carcinoma), U266 (multiple myeloma), DU145 (prostate cancer), as well as MDA-MB-231 (breast cancer) cells (Rhee et al. 2012). Ergosterol peroxide inhibited cell growth and induced apoptosis in human prostate cancer cells DU-145 and LNCaP (Russo et al. 2010). Moreover, ergosterol peroxide showed a cytotoxic effect against Hep 3B cells (Chen et al. 2009). Six sterols comprising ergosterol peroxide were isolated from R. flava (Liu et al. 2012). And interestingly, their ethanolic extract exerted promising growth inhibitory activity on MDA-MB-231(human breast cancer cell line) which suggests that R. flava could find a great application as an antitumor agent.

    Other biological activities of Ramaria species

    To date, a few numbers of other biological activities have been determined in Ramaria species. In a previous study, the hepatoprotective activity of methanol extract of R. botrytis towards liver toxicity induced by benzo(α)pyrene in mice was evaluated (Kim & Lee 2003). The results reported that the methanolic extract significantly reduced the elevated enzyme activities including glutathione S-transferase and r-glutamylcysteine synthetase that resulted from the induction of benzo(α)pyrene. The hepatoprotective activity of this mushroom could be related to its high antioxidant potential which was revealed by the low EC50 value (0.109 mg/ml) for DPPH radical scavenging assay. Ramaria botrytis collected from hilly areas of Darjeeling exerted potential immunostimulatory effect in a murine macrophage cell line (RAW264.7 cell) as well as in thymocyte cells and splenocytes. This activity was related to a water-soluble glucan obtained from fresh fruiting bodies of such mushrooms and which consisted of (1→6)-linked-β-D glucopyranosyl residues with the branching of (1→3)-linked-β-D-glucopyranosyl at O-3 position. This glucan enhanced the nitric oxide level and also resulted in the stimulation of thymocyte and splenocyte proliferation rates. Thus, this glucan can be employed as a potent immunostimulatory (Bhanja et al. 2013).

    Ramarin A and B are two novel sesquiterpene derivatives that were obtained and purified from the methanol extract of Ramaria formosa that is a very rare mushroom and that inhibits human neutrophil elastase, and thus can be employed for the treatment of skin aging (Kim et al. 2016). A recent study reported by Bhanja et al. (2020), prepared bio metallic composite nanoparticles from polysaccharides isolated from Ramaria botrytis. The nanoparticles exerted antibacterial activity against Pseudomonas aeruginosa. Moreover, these nanoparticles showed potential antioxidant activities towards DPPH radical, hydrogen peroxide, and nitric oxide radicals. They also catalyzed the p-nitrophenol reduction, indicating a new direction in the field of biomedicine mediated by nanotechnology (Bhanja et al. 2020).

    Clavaria sp.

    All Clavaria species are terrestrial and most are believed to be saprotrophic (Decaying dead plant material). In North America and elsewhere, they are more commonly found in woodlands. Clavaria fruit bodies are simple (cylindrical to club-shaped) or more rarely branched, sometimes with a distinct stem. Several of the species with simple fruit bodies form them in dense clusters. The fruit bodies themselves are smooth to grooved and typically brittle. Depending on species, they vary in color from white or cream to yellow, pink, violet, brown, or black. The hyphal system of Clavaria species is always monomitic. The context hyphae are inflated, thin-walled, and lack clamp connections (Olariaga et al. 2015). The basidia are two to four spored, in some species with an open, loop-like clamp connection at the base. Spores are smooth or spiny and color is white. Most Clavaria species are thought to be saprotrophic, decomposing leaf litter and other organic materials on the woodland floor. In Europe, species are more frequently found in old, unimproved grasslands, where they are presumed to be decomposers of dead grass and moss. Species of Clavaria occur in suitable habitats throughout the temperate regions and the tropics (Figs 3, 4) (Acharya 2012, Kautmanová et al. 2012).

    Figure 3.  Fruitbody of Clavaria zollingeri in the field. The photograph was taken at Ohio, USA by Rcurtis (https://www.inaturalist.org/observations/1693040). The photo was used under the CC BY-NC 4.0 non-commercial use license. Scale bar = 10 cm.
    DownLoad: Full-Size Img PowerPoint
    Figure 4.  Fruitbodies of Clavaria rubicundula in the field. The photograph was taken at Hewitt, West Milford, NJ 07421, USA by Tombigelow (https://www.inaturalist.org/observations/16044756). The photo was used under the CC BY-NC 4.0 non-commercial use license. Scale bar = 10 cm.
    DownLoad: Full-Size Img PowerPoint

    Clavaria sp. biological activities

    Ramaria and Clavaria are the two major genera of coral mushrooms within families Gomphaceae and Clavariaceae, respectively. Besides having important role in forest ecology, some species of these are reported to possess high nutraceutical and bioactive potential (Sharma & Gautam 2017). Many current studies describe the detailed biochemical profiling and antioxidant, and antibacterial activities of twelve coral mushroom (Ramaria and Clavaria) species (Vidović et al. 2014). Antioxidant activities were calculated using EC50 values from mushroom extracts. Antibacterial activities were checked on six pathogenic bacterial strains through minimum inhibition concentrations. All the species were found to be rich in protein, macro and micro minerals, carbohydrates, unsaturated fatty acids, essential amino acids, phenolics, tocopherols, anthocynadins and carotenoids. All the species showed significant antioxidant and antibacterial activities. These species are reported to free from heavy toxic metals. Sharma & Gautam (2017), reported that these Ramaria and Clavaria species will open the way for their large-scale commercial exploitations and use in pharmaceutical industries as antioxidant, antibacterial and nutraceutical constituents.

    Conclusion

    Many studies conducted on Ramaria and Clavaria species are represented in the current review and showed that coral mushrooms exhibit the potential as a vital therapeutic food. However, more studies for profound exploration are still required. Ramaria and Clavaria species exerted some vital biological activities such as antimicrobial, antioxidant, antitumor, hepatoprotective, and anti-skin aging activity, etc. Further investigation is needed to explain the different mechanisms of action of these wild mushrooms. Although the nutritional values of the genus Ramaria and Clavaria have been also studied, there is a lack of information about their bioavailability, vitamin and mineral, and compositions. Thus, the current review recommends further exploration to get a full profile of the active components obtained from various Ramaria species and Clavaria species in nutrition and mycomedicine fields.

    Funding

    This research does not receive any external funding.

    Conflicts of Interest

    The authors declare no conflict of interest.

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  • Cite this article

    BC Samarakoon, R Phookamsak, SC Karunarathna, R Jeewon, P Chomnunti, JC Xu, YJ Li. 2021. New host and geographic records of five pleosporalean hyphomycetes associated with Musa spp. (Banana). Studies in Fungi 6(1):92−115 doi: 10.5943/sif/6/1/5
    BC Samarakoon, R Phookamsak, SC Karunarathna, R Jeewon, P Chomnunti, JC Xu, YJ Li. 2021. New host and geographic records of five pleosporalean hyphomycetes associated with Musa spp. (Banana). Studies in Fungi 6(1):92−115 doi: 10.5943/sif/6/1/5
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New host and geographic records of five pleosporalean hyphomycetes associated with Musa spp. (Banana)

  • 1.

    The State Phosphorus Resource Development and Utilization Engineering Technology Research Centre, Yunnan Phosphate Chemical Group Co. Ltd, Kunming 650201, China

  • 2.

    Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand

  • 3.

    School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand

  • 4.

    CAS Key Laboratory for Plant Biodiversity and Biogeography of East Asia (KLPB), Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China

  • 5.

    East and Central Asia Regional Office, World Agroforestry Centre (ICRAF), Kunming 650201, Yunnan, China

  • 6.

    Centre for Mountain Futures (CMF), Kunming Institute of Botany, Kunming 650201, Yunnan, China

  • 7.

    Honghe Center for Mountain Futures, Kunming Institute of Botany, Chinese Academy of Sciences, Honghe County 654400, Yunnan, People's Republic of China

  • 8.

    Department of Health Sciences, Faculty of Medicine and Health Sciences, University of Mauritius, Reduit, Mauritius

  • 9.

    Research Center of Microbial Diversity and Sustainable Utilization, Faculty of Sciences, Chiang Mai University, Chiang Mai 50200, Thailand

  • Received: 08 January 2021
  • Accepted: 02 February 2021
  • Published online: 23 February 2021
Studies in Fungi  6,  Article number: 5  (2021)  |  Cite this article

Abstract: A study was undertaken to collect and identify saprobic fungi associated with Musa spp. (banana) from Taiwan (China), and Thailand. Samples were collected during the dry season and their morpho–molecular relationships were investigated. Five brown pleosporalean hyphomycetous taxa in Periconiaceae and Torulaceae viz. Periconia cortaderiae, P. delonicis, Torula chromolaenae, T. fici, and T. masonii were identified for the first time from Musa spp. (Musaceae). Phylogenetic analyses of a combined SSU, LSU, ITS, RPB2 and TEF DNA sequence dataset further justified the taxonomic placements of these five taxa in the above mentioned families. Periconia delonicis is reported for the first time on a monocotyledonous host and T. masonii is the first geographical record from Taiwan (China).

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Introduction
Materials and methods

    Fungal collections, isolation and morphological characterization

  • Dead leaves of Musa spp. were collected from China and Thailand. Specimens were transferred to the laboratory in cardboard boxes. Samples were examined with a Motic SMZ 168 Series microscope. Powder like masses of conidia of hyphomycetous taxa on Musa samples were mounted in water for microscopic studies and photomicrography. The hyphomycetous taxa were examined using a Nikon ECLIPSE 80i compound microscope and photographed with a Canon 550D digital camera fitted to the microscope. Measurements were made with the Tarosoft (R) Image Frame Work program and images used for figures processed with Adobe Photoshop CS6 Extended version 12.0 software (Adobe, USA).

    Single spore isolation was carried out following the method described in Senanayake et al. (2020). Germinated spores were individually transferred to potato dextrose agar (PDA) plates and grown at 25℃ in normal light. Colony characteristics were observed and measured after 3 weeks. The specimens are deposited at the Mae Fah Luang University (MFLU) Herbarium, Chiang Rai, Thailand. Living cultures are deposited at the Culture Collection of Mae Fah Luang University (MFLUCC). Faces of fungi numbers are registered as in Jayasiri et al. (2015).

    DNA extraction and PCR amplification

  • Fungal isolates derived from single spore cultures were grown on potato dextrose agar (PDA) for 4 weeks at 25℃ and the axenic mycelia (50–100 mg) of each isolate were scrapped off for DNA extraction purposes. Mycelia were ground to a fine powder with liquid nitrogen and fungal DNA was extracted using the Biospin Fungus Genomic DNA Extraction Kit–BSC14S1 (BioFlux®, P.R. China) according to the instructions of the manufacturer. Five gene regions were used for the polymerase chain reaction (PCR) amplification, including partial 18S small subunit rDNA (SSU), partial 28S large subunit rDNA (LSU), internal transcribed spacer (ITS), RNA polymerase II second largest subunit (RPB2) and partial translation elongation factor 1–alpha gene (TEF) using the primers NS1/NS4 (White et al. 1990), LR0R/LR5 (Vilgalys & Hester 1990), ITS5/ITS4 (White et al. 1990), fRPB2–5f/fRPB2–7cR (Liu et al. 1999) and EF1–983F/EF1–2218R (Rehner 2001), respectively.

    The final volume of the PCR reaction was 25 μl, consisting of 2 μl of DNA template, 1 μl of each forward and reverse primer, 12.5 μl of 2×Easy Taq PCR SuperMix (mixture of EasyTaqTM DNA Polymerase, dNTPs, and optimized buffer, Beijing TransGen Biotech Co., Ltd., Beijing, P.R. China) and 8.5 μl of the sterilized double–distilled water (ddH2O). The thermal cycle programs were set up following the procedures described by Samarakoon et al.(2019, 2020b) for the respective genes. The amplified PCR fragments were sent to a commercial sequencing provider (TsingKe Biological Technology (Beijing) Co., Ltd, China) for PCR purification and sequencing. The Sanger DNA sequences obtained from this study were deposited in GenBank (Tables 1, 2).

    Table 1.  Taxa used in the phylogenetic analyses of Periconiaceae with the corresponding GenBank accession numbers. Type strains are superscripted with "T" and newly generated strains are indicated in black bold.

    Taxa Culture collection/Voucher no. GenBank accession numbers
    LSU SSU ITS TEF
    Flavomyces fulophazii CBS 135761T KP184040 KP184082 KP184001 NA
    F. fulophazii MF09 MN515261 NA MN537663 MN535259
    Helminthosporium H 4628 AB807521 AB797231 LC014555 AB808497
    dalbergiae
    Massarina cisti CBS 266.62T AB807539 AB797249 NA AB808514
    Noosia banksiae CPC: 17282 JF951167 NA JF951147 NA
    N. banksiae CBS 129526 MH878062 NA NA NA
    Periconia aquatica MFLUCC 16–0912T KY794705 NA KY794701 KY814760
    P. byssoides MFLUCC 17–2292 MK347968 MK347858 MK347751 MK360069
    P. byssoides MFLUCC 18–1548 MK348013 MK347902 MK347794 MK360070
    P. caespitosa LAMIC_110_16 MH051907 NA MH051906 NA
    P. cortaderiae MFLUCC 15–0457T NG_068238 NG_068373 NR_165853 KY310703
    P. cortaderiae MFLUCC 15–0451 KX954403 KX986346 KX965734 KY429208
    P. cortaderiae MFLUCC 20–0236 MW406971 MW406969 MW406973 MW422156
    P. cyperacearum CPC: 32138T NG_064549 NA NR_160357 MH327882
    P. delonicis MFLUCC 17–2584T MK347941 MK347832 NA MK360071
    P. delonicis MFLUCC 20–0235 MW406970 MW406968 MW406972 MW422155
    P. digitata CBS 510.77 AB807561 AB797271 NA AB808537
    P. epilithographicola CBS 144017T NA NA NR_157477 NA
    P. epilithographicola PL5–1B NA NA MF422162 NA
    P. homothallica HHUF 29105 NG_059397 NG_064851 NR_153446 AB808541
    P. homothallica KT 916 AB807565 AB797275 AB809645 NA
    P. igniaria CBS 379.86 AB807566 AB797276 LC014585 AB808542
    P. igniaria CBS 845.96 AB807567 AB797277 LC014586 AB808543
    P. macrospinosa CBS 135663 KP184038 KP184080 KP183999 NA
    P. neobrittanica CPC 37903T NG_068342 NA NR_166344 NA
    P. palmicola MFLUCC 14–0400T NG_068917 MN648319 NA MN821070
    P. pseudobyssoides DLUCC 0850 MG333494 NA MG333491 MG438280
    P. pseudobyssoides H 4151 AB807568 AB797278 LC014587 AB808544
    P. pseudobyssoides H 4790 AB807560 AB797270 LC014588 AB808536
    P. pseudodigitata KT 644 AB807562 AB797272 LC014589 AB808538
    P. pseudodigitata KT 1195A AB807563 AB797273 LC014590 AB808539
    P. pseudodigitata KT 1395 AB807564 AB797274 LC014591 AB808540
    P. salina MFLU 19–1235T MN017846 MN017912 MN047086 NA
    P. submersa MFLUCC 16–1098T KY794706 NA KY794702 KY814761
    P. thailandica MFLUCC 17–0065T KY753888 KY753889 KY753887 NA

    Table 2.  Taxa used in the phylogenetic analyses of Torulaceae with the corresponding GenBank accession numbers. Type strains are superscripted with "T" and newly generated strains are indicated in black bold.

    Taxa Culture collection/Voucher no. ITS LSU SSU RPB2 TEF
    Dendryphion aquaticum MFLUCC 15–0257T KU500566 KU500573 KU500580 NA NA
    D. comosum CBS 208.69T MH859293 MH871026 NA NA NA
    D. europaeum CPC 23231 KJ869145 KJ869202 NA NA NA
    D. hydei KUMCC 18–0009T MN061343 MH253927 MH253929 NA NA
    Neotorula aquatica MFLUCC 15–0342T KU500569 KU500576 KU500583 NA NA
    N. submersa HKAS 92660 NR_154247 KX789217 NA NA NA
    Rostriconidium aquaticum KUMCC 15–0297 MG208165 MG208144 NA MG207975 MG207995
    R. aquaticum MFLUCC 161113T MG208164 MG208143 NA MG207974 MG207994
    R. pandanicola KUMCC 17–0176T MH275084 MH260318 MH260358 MH412759 MH412781
    Roussoella nitidula MFLUCC 11–0182T KJ474835 KJ474843 NA KJ474859 KJ474852
    R. scabrispora MFLUCC 11–0624T KJ474836 KJ474844 NA KJ474860 KJ474853
    Roussoellopsis tosaensis KT 1659 NA AB524625 AB524484 AB539104 AB539117
    Rutola graminis CPC 33267 MN313814 MN317295 NA NA NA
    R. graminis CPC 33695 MN313815 MN317296 NA NA NA
    R. graminis CPC 33715T MN313816 MN317297 NA NA NA
    Sporidesmioides thailandica MFLUCC 13–0840 MN061347 NG_059703 NG_061242 KX437761 KX437766
    S. thailandica KUMCC 16–0012T MN061348 KX437758 KX437760 KX437762 KX437767
    Thyridaria broussonetiae TB1 KX650569 NA KX650515 KX650586 KX650539
    Thyridariella mahakoshae NFCCI 4215 MG020435 MG020438 MG020441 MG020446 MG023140
    Th. mangrovei PUFD 17–98T MG020434 MG020437 MG020440 MG020445 MG020443
    Torula acaciae CPC 29737T NR_155944 NG_059764 NA KY173594 NA
    T. aquatica DLUCC 0550 MG208166 MG208145 NA MG207976 MG207996
    T. aquatica MFLUCC 16–1115T MG208167 MG208146 NA MG207977 NA
    T. breviconidiophora KUMCC 18–0130T MK071670 MK071672 MK071697 NA MK077673
    T. camporesii KUMCC 19–0112T MN507400 MN507402 MN507401 MN507404 MN507403
    T. chiangmaiensis KUMCC 16–0039T MN061342 KY197856 KY197863 NA KY197876
    T. chromolaenae MFLUCC 20–0237 MW412524 MW412518 MW412515 MW422161 MW422158
    T. fici CBS 595.96T KF443408 KF443385 KF443387 KF443395 KF443402
    T. fici KUMCC 15–0428 MG208172 MG208151 NA MG207981 MG207999
    T. fici KUMCC 16–0038 MN061341 KY197859 KY197866 KY197872 KY197879
    T. fici MFLUCC 20–0238 MW412525 MW412519 MW412516 NA MW422159
    T. gaodangensis MFLUCC 17–0234T MF034135 NG_059827 NG_063641 NA NA
    T. goaensis MTCC 12620T NR_159045 NG_060016 NA NA NA
    T. herbarum CPC 24414 KR873260 KR873288 NA NA NA
    T. hollandica CBS 220.69 NR_132893 NG_064274 KF443389 KF443393 KF443401
    T. hydei KUMCC 16–0037T MN061346 MH253926 MH253928 NA MH253930
    T. mackenziei MFLUCC 13–0839T MN061344 KY197861 KY197868 KY197874 KY197881
    T. masonii CBS 245.57T NR_145193 NG_058185 NA NA NA
    T. masonii DLUCC 0588 MG208173 MG208152 NA MG207982 MG208000
    T. masonii MFLUCC 20–0239 MW412523 MW412517 MW412514 MW422160 MW422157
    T. pluriseptata MFLUCC 14–0437T MN061338 KY197855 KY197862 KY197869 KY197875
    T. polyseptata KUMCC 18–0131T MK071671 MK071673 MK071698 NA MK077674
    Abbreviations of culture collections: CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, Netherlands. CPC: Working collection of Pedro Crous housed at CBS. DLUCC: Dali University Culture Collection, China. H: University of Helsinki, Helsinki, Finland. HKAS: Herbarium of Cryptogams, Kunming Institute of Botany, Academia Sinica, China. KT: K. Tanaka. LAMIC: Laboratorio Asociaciones suelo, planta microorganismos, Pontificia Universidad Javeriana, Bogotá, D.C., Colombia. MTCC: Microbial Type Culture Collection, CSIR–Institute of Microbial Technology, Sector 39–A, Chandigarh – 160036, India. KUMCC: Kunming Institute of Botany Culture Collection, China. MFLU: Mae Fah Luang University, Chiang Rai, Thailand. MFLUCC: Mae Fah Luang University Culture Collection, Chiang Rai, Thailand. NFCCI: National Fungal Culture Collection of India. NA: DNA sequence data are not available in GenBank.

    Sequence alignment

  • Obtained sequences were subjected to BLASTn search tool in GenBank (https://blast.ncbi.nlm.nih.gov/Blast.cgi) for finding the closely related taxa. BLASTn search results and initial morphological studies have supported that our isolates belong to Periconiaceae and Torulaceae. Other sequences used in the analyses were obtained from GenBank (Tables 1, 2) based on recently published data (Jayasiri et al. 2019, Hongsanan et al. 2020, Hyde et al. 2020a, Li et al. 2020). The single gene alignments were automatically analysed by MAFFT v. 7.036 (http://mafft.cbrc.jp/alignment/server/large.html; Katoh et al. 2019) using the default settings and refined where necessary, using BioEdit v. 7.0.5.2 (Hall 1999). The single gene matrixes were prior analyzed by maximum likelihood (ML) criterion for checking the congruence of the tree topologies and if the tree topologies were congruent, the concatenated sequence dataset were performed for further analyses.

    Phylogenetic analyses

  • Phylogenetic analyses were preformed based on maximum likelihood (ML) and Bayesian inference (BI) criteria. The phylogenetic trees showing relationships of taxa in Periconiaceae and Torulaceae were generated separately. Data matrixes used in these analyses were followed as; Periconiaceae (Analysis 1): the combined SSU, LSU, ITS and TEF data matrix comprised 35 sequences of representative taxa in Periconiaceae. Helminthosporium dalbergiae (MAFF 243853) and Massarina cisti (CBS 266.62) were selected as outgroup taxa. Torulaceae (Analysis 2): the combined SSU, LSU, ITS, TEF and RPB2 matrix comprised 40 sequences of selected genera in Torulaceae. Taxa in Roussoellaceae were selected as the outgroup taxa viz. Roussoella nitidula (MFLUCC 11–0182), R. scabrispora (MFLUCC 11–0624) and Roussoellopsis tosaensis (KT 1659).

    Maximum likelihood (ML) trees were generated using the RAxML–HPC2 on XSEDE (8.2.8) (Stamatakis et al. 2008, Stamatakis 2014) in the CIPRES Science Gateway platform (Miller et al. 2010) using GTR+I+G model of evolution and 1, 000 replicates of rapid bootstrap. Bayesian inference (BI) analysis was conducted with MrBayes v. 3.1.2 (Huelsenbeck & Ronquist 2001) to evaluate posterior probabilities (PP) (Rannala & Yang 1996, Zhaxybayeva & Gogarten 2002) by Markov Chain Monte Carlo sampling (BMCMC). Two parallel runs were conducted, using the default settings, but with the following adjustments: four simultaneous Markov chains were run for 2, 000, 000 generations (Analysis 1, 2) and trees were sampled every 100th generation and 20, 000 trees were obtained. The first 4, 000 trees, representing the burn–in phase of the analyses were discarded. The remaining 16, 000 trees were used for calculating PP in the 50% majority rule consensus tree.

    Phylograms were visualized with FigTree v1.4.0 program (Rambaut 2011) and reorganized in Microsoft PowerPoint (2007, USA) and converted to jpeg file in Adobe Photoshop CS6 Extended version 12.0 software (Adobe, USA). The final trees and data matrixes were submitted in TreeBASE (https://www.treebase.org/), submission ID: 27468 for Periconiaceae and 27469 for Torulaceae.

Results

    Phylogenetic analyses

  • Analysis 1: Periconiaceae

    The best scoring RAxML tree resulted from the combined gene analysis of SSU, LSU, ITS and TEF sequence data is shown in Fig. 1 with a final ML optimization likelihood value of – 9645.86. The matrix had 561 distinct alignment patterns, with 26.15% of undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.233866, C = 0.256179, G = 0.274088, T = 0.235866; substitution rates AC = 2.126081, AG = 2.703293, AT = 1.718216, CG = 1.412442, CT = 10.292987, GT = 1.000000; proportion of invariable sites I =0.624461; gamma distribution shape parameter α = 0.615919. Bayesian posterior probabilities (BYPP) from MCMC were evaluated with a final average standard deviation of split frequencies was 0.008. All trees (ML & BI) were similar in topology and did not differ significantly with respects to generic placements and this is in agreement with previous studies based on multi–gene phylogeny (Jayasiri et al. 2019, Hongsanan et al. 2020, Hyde et al. 2020a). Our MFLUCC 20–0235 strain is sister to Periconia delonicis (MFLUCC 17-2584) and grouped with P. palmicola (MFLUCC 14-0400) and P. verrucosa (MFLUCC 17-2158) with a strong statistical support (ML = 98%, BYPP =1.00; Fig. 1). The MFLUCC 20–0236 strain grouped with P. cortaderiae (MFLUCC 15-0457, MFLUCC 15-0451) with high support (ML = 100%, BYPP = 1.00), and has close phylogenetic affinity to Noosia banksiae (CPC: 17282, CBS 129526) and Periconia homothallica (KT 916, HHUF 29105). Therefore, we identify MFLUCC 20–0235 as P. delonicis and MFLUCC 20–0236 as P. cortaderiae, respectively.

    Figure 1. 

    Maximum likelihood tree revealed by RAxML from an analysis of a concatenated SSU, LSU, ITS and TEF sequence dataset of the species in Periconiaceae, showing the phylogenetic position of Periconia delonicis (MFLUCC 20–0235) and P. cortaderiae (MFLUCC 20–0236). ML bootstrap supports (≥ 60%) and Bayesian posterior probabilities (≥ 0.95 BYPP) are given above the branches as ML/BYPP. The tree is rooted with Helminthosporium dalbergiae (MAFF 243853) and Massarina cisti (CBS 266.62). Strains generated in this study are indicated in red bold. Ex–type strains are indicated in black bold. The scale bar 0.02 represents the expected number of nucleotide substitutions per site.

    Analysis 2: Torulaceae

    The best scoring RAxML tree (Fig. 2) has a final ML optimization likelihood value of – 19754.05. The matrix had 1189 distinct alignment patterns, with 31.37% of undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.241559, C = 0.263242, G = 0.273563, T = 0.221637; substitution rates AC = 1.375308, AG = 2.974871, AT = 1.345285, CG = 0.922655, CT = 6.841524, GT = 1.000000; proportion of invariable sites I = 0.486989; gamma distribution shape parameter α = 0.482115. Bayesian posterior probabilities (BYPP) from MCMC were evaluated with a final average standard deviation of split frequencies was 0.005. All the phylogenetic trees (ML & BI) were similar in topology and no considerable difference at the generic level and similar with the phylogenetic analyses of Li et al.(2017, 2020), Su et al. (2018) and Hyde et al. (2020a). Our isolate (MFLUCC 20–0237) formed a clade with Torula chromolaenae (KUMCC 16–0036) with high support (ML = 100%, BYPP = 1.00), and clustered with T. breviconidiophora (KUMCC 18–0130) and T. mackenziei (MFLUCC 13-0839). The MFLUCC 20–0238 strain clustered with T. fici (CBS 595.96, KUMCC 15–0428, KUMCC 16–0038) with moderate support in ML analysis (ML = 80%) and not significant support in BI analysis and has a close relationship with T. hydei (KUMCC 16–0037; ML 98%, BYPP = 1.00). The MFLUCC 20–0239 strain clustered with T. masonii (CBS 245.57, DLUCC 0588, KUMCC 16–0033) with high support (ML = 100%, BYPP = 1.00), and also close to T. acaciae (CPC 29737). We therefore identify MFLUCC 20–0237 as T. chromolaenae, MFLUCC 20–0238 as T. fici and MFLUCC 20–0239 as T. masonii, respectively.

    Figure 2. 

    Maximum likelihood tree (RAxML) revealed by an analysis of a concatenated SSU, LSU, ITS, TEF and RPB2 sequence dataset of the species in Torulaceae, showing the phylogenetic position of Torula chromolaenae (MFLUCC 20–0237), T. fici (MFLUCC 20–0238) and T. masonii (MFLUCC 20–0239). ML bootstrap supports (≥ 60%) and Bayesian posterior probabilities (≥ 0.95 BYPP) are given above the branches as ML/BYPP. The tree is rooted with Roussoella nitidula (MFLUCC 11–0182), R. scabrispora (MFLUCC 11–0624) and Roussoellopsis tosaensis (KT 1659). Strains generated in this study are indicated in red bold. Ex–type strains are indicated in black bold. The scale bar 0.04 represents the expected number of nucleotide substitutions per site.

    Taxonomy

  • Periconia delonicisJayasiri, E.B.G. Jones & K.D. Hyde, in Jayasiri et al., Mycosphere 10(1): 9 (2019)                  Fig. 3

    Figure 3. 

    Periconia delonicis (MFLU 20-0696). a-c Conidiophores bearing conidia on host. d-f Conidia and conidiophores. g, h Conidiogenous cells bearing conidia. i Base of the conidiophore. j-s Mature and immature conidia. t Colonies on PDA after 14 days. Scale bars: a, b = 500 μm, c, d = 100 μm, e, f, i = 20 μm, g, h = 10 μm, j-s = 5 μm.

    Index Fungorum number: IF555562; Facesoffungi number: FoF 05268

    Saprobic on a dead leaf of Musa sp. Sexual morph: Undetermined. Asexual morph: hyphomycetous. Conidiophores 320-400 × 8-12 μm (x = 365 × 9.5 μm, n = 10) macronematous, rarely micronematous. Macronematous conidiophores and conidia resembled a stipe and a globular head. Stipe of the Conidiophores unbranched, straight or flexuous from the middle, septate, pale to dark brown, often appearing black, smooth, globular heads shining by reflected light. Conidiogenous cells 4-7.5 × 3.4-7.4 μm (x = 5.5 × 5.3 μm, n = 15) monoblastic or polyblastic, discrete on the stipe, determinate, ellipsoidal, pale brown to brown. Conidia 4-8 × 4-6 μm (x = 5.6 × 5.4 μm, n =15), catenate, in chains, arising at one or more points on the curved surface of the conidiogenous cell, simple, usually spherical or subspherical, pale to dark brown, smooth to minutely veruculose, aseptate.

    Culture characteristics – Conidia germinated on PDA within 48 hrs, reaching 40 mm diam. in 2 weeks at 25℃. Colonies on PDA with sparse, pinkish white mycelia on the surface, circular and flattened. Surface is smooth, with small, brown granular–like, powdery masses at maturity. The reverse of the colony is dark brown and yellow in the center with a white margin. Conidia and conidiophores are not observed in mature colonies.

    Material examined – THAILAND, Chiang Mai Province, Mae Taeng District, on a dead leaf of Musa sp. (Musaceae), 22 September 2018, B.C. Samarakoon, BNS012 (MFLU 20–0696), living culture MFLUCC 20–0235.

    Known hosts & distribution – on pods of Delonix regia (Fabaceae, Dicotyledon) from Thailand (Jayasiri et al. 2019); on dead leaf of Musa sp. (Musaceae, Monocotyledon) from Thailand (this study).

    Notes – Based on BLASTn search results of SSU, LSU, ITS, TEF sequence data, our strain (MFLUCC 20-0235) showed high identity to the taxa in GenBank as follows; SSU = 99.75% similarity to Periconia palmicola (MFLUCC 14-0400), LSU = 99.88% similarity to P. delonicis (MFLUCC 17-2584), ITS = 99.20% similarity to P. palmicola (MFLUCC 14-0400). Multi–loci phylogenetic analysis (Fig. 1) showed that the new strain MFLUCC 20–0235 forms a clade together with P. palmicola (MFLUCC 14-0400) and P. verrucosa (MFLUCC 17-2158) with high statistical support (ML = 98%, BYPP = 1.00).

    Our strain (MFLUCC 20–0235) shares similar morphology to the type of Periconia delonicis in having macronematous, unbranched conidiophores, with monoblastic, terminal conidiogenous cells and brown, globose to subglobose, aseptate, veruculose conidia (Jayasiri et al. 2019). Our strain (MFLUCC 20-0235) also shares similar size range of conidiophores (320-400 × 8-12 μm vs. 360-420 × 8-12 μm) and conidia (4-8 × 4-6 μm vs. 5.5-7 μm diam.). Based on recommendations by Jeewon & Hyde (2016), we also compare the nucleotide bases of TEF region for our new strain and the type strain of P. delonicis. MFLUCC 20-0235 differs from P. delonicis (MFLUCC 17-2584) in 5/746 bp (0.67%).

    In our phylogenetic analyses, MFLUCC 20-0235 and ex–type stain of Periconia delonicis (MFLUCC 17-2584) showed a close phylogenetic affinity to P. palmicola (MFLUCC 14-0400) and P. verrucosa (MFLUCC 17-2158). The conidiophores, conidiogenous cells and the conidia of MFLUCC 20-0235 shares similar morphology with the holotype illustration of P. palmicola in Hyde et al. (2020a) (i.e. dark brown to black conidiophores, hyaline conidiogenous cells, subglobose to globose conidia). However, P. palmicola differs from MFLUCC 20-0235 in having comparatively short conidiophores (151–188 × 5.6–8 μm vs. 320-400 × 8-12 μm) which were branched at apex. The conidiophores of MFLUCC 20-0235 are unbranched and comparatively long with respect to P. palmicola. A nucleotide base pair comparison of TEF region showed that MFLUCC 20-0235 differs from P. palmicola (MFLUCC 14-0400) in 4/746 (0.53%). MFLUCC 20-0235 also has similar morphology to P. verrucosa (MFLUCC 17-2158) but differs in having longer conidiophores (320-400 × 8-12 μm vs. 170–296 × 10–12 μm) and smaller conidia (4-8 × 4-6 μm. vs. 7–15 μm diam.) (Phukhamsakda et al. 2020). A nucleotide base comparison of ITS and TEF regions showed that MFLUCC 20-0235 differs from P. verrucosa (MFLUCC 17-2158) in 7/501 bp (1.39%) of ITS and 4/746 bp (0.53%) of TEF. Based on morphological comparison with the types of P. delonicis, P. palmicola and P. verrucosa, a nucleotide base comparison of ITS and TEF regions and phylogenetic evidence, we thus identify our new collection as P. delonicis. In this study, we report P. delonicis on Musa sp. (Musaceae, monocotyledon) for the first time.

    Periconia cortaderiaeThambugala & K.D. Hyde, in Thambugala et al., Mycosphere 8(4): 734 (2017)                  Fig. 4

    Figure 4. 

    Periconia cortaderiae (MFLU 20-0697). a-c Colonies on host substrate. d, f Conidiophores bearing conidia. e, g, h Conidial masses. i, j, m-o Conidiogenesis from terminal mother cells. l Monoblastic, annellidic conidiogenous cells bearing conidial chains. k, p, q, r Conidial chains. s Conidia. Scale bars: a = 3 mm, b = 3.5 mm, c = 50 μm, d, f = 50 μm, e, g = 20 μm, h-s = 5 μm.

    Index Fungorum number: IF553165; Facesoffungi number: FoF 03226

    Saprobic on a dead leaf of Musa sp. Sexual morph: Undetermined. Asexual morph: hyphomycetous. Colonies on host effuse, powdery, gregarious, with a black, conidial masses on the host. Mycelium composed of cottony, branched, hyphae forming dark clusters with conidia scattered on the host surface. Conidiophores 40–120 × 1-6 μm (x = 58.5 × 3.5 μm, n = 15), macronematous, mononematous, single or rarely 2–3 together, erect, thick–walled, brown to dark brown, septate, rough, 4–5 μm wide at base, and 1–2 μm wide at apex; base shoe–shaped, distinct. Conidiogenous cells 2.5–4.5 μm × 2.5–3.5 μm (x = 3.5 × 2.8 μm, n = 10), annellidic, monoblastic, discrete on stipe, percurrent proliferations present as scars at the apex of conidiophore. Conidia 4–7.6 × 4–6.5 μm (x = 6.5 × 5.9 μm, n = 40), catenate, globose, one–celled, hyaline to pale brown when immature, becoming brown to dark brown, smooth or minutely verruculose.

    Culture characteristics – Conidia germinating on PDA within 48 hrs. Colonies growing on MEA, reaching a diameter of 20 mm after 14 days at 25℃, flat when immature, raised at maturity with unevenly distributed radial furrows or linear marks, surface is smooth at immature stage and notably rough at maturity with crenulate to crenate margin. Colony is initially white, moderately dense and completely black at maturity with the sporulation, reverse white to black.

    Material examined – THAILAND, Nan Province, Nai Wiang, on a dead leaf of Musa sp. (Musaceae), 1 March 2019, B.C. Samarakoon, BNS100 (MFLU 20-0697), living culture MFLUCC 20-0236.

    Known hosts & distribution – on Cortaderia sp. (Poaceae, Monocotyledon) from Thailand (Thambugala et al. 2017); on Caragana arborescens (Fabaceae, Dicotyledon) from Yunnan, China (Phookamsak et al. 2019); on Musa sp. (Musaceae, Monocotyledon) from Thailand (this study).

    Notes – Based on BLASTn search results of SSU, LSU, ITS sequence data, our strain (MFLUCC 20-0236) showed high identity to the taxa in GenBank as follows; SSU = 99.76%, LSU = 99.88%, ITS = 99.80% similarities to Periconia cortaderiae (MFLUCC 15-0457). The morphology of our strain is similar to the holotype of P. cortaderiae (Thambugala et al. 2017), except the length of the conidiophores. Our strain has short conidiophores with respect to the holotype (40–120 × 1-6 μm vs. 400–800 × 4–9.4 μm). A nucleotide base comparison of ITS and TEF regions showed that MFLUCC 20-0236 differs from P. cortaderiae (MFLUCC 15-0457) in 1/493 bp (0.2%) of ITS and 1/829 bp (0.12%) of TEF. In addition, our MFLUCC 20-0236 strain differs from P. cortaderiae (MFLUCC 15-0451) in 2/493 bp (0.4%) of ITS and 2/872 bp (0.2%) of TEF. Therefore, we identify our new collection as P. cortaderiae, and this is reported from Musa sp. (monocotyledon) from Thailand for the first time.

    Torula chromolaenaeJun F. Li, Phook., Mapook & K.D. Hyde, in Li et al., Mycol. Progr. 16(4): 454 (2017)                  Fig. 5

    Figure 5. 

    Torula chromolaenae (MFLU 20–0698). a-c Colonies on leaf vein of Musa sp. d, e Conidia arranged in multi planes. f Conidiophore and conidiogenous cell. g-p Conidia and the formation of immature conidia. q Colonies on PDA after 21 days. Scale bars a = 1000 μm, b = 500 μm, c = 50 μm, d-h, k, o, p = 10 μm, l-n, i, j = 5 μm.

    Index Fungorum number: IF819536; Facesoffungi number: FoF 02713

    Saprobic on a dead leaf vein of Musa sp. (Musaceae). Sexual morph: Undetermined. Asexual morph: hyphomycetous. Colonies effuse, black, powdery, thread–like on host. Mycelium slightly immersed, septate, unbranched, smooth, pale brown hyphae. Conidiophores 2–5 × 2–4 μm (x = 3.8 × 3.3 μm, n = 10), micronematous or semi–macronematous, unbranched, straight or flexuous, subhyaline or pale brown, smooth or minutely verruculose. Conidiogenous cells 2–6 × 2–4.5 μm (x = 4.5 × 3.7 μm, n =10), polyblastic or sometimes monoblastic, integrated, terminal, discrete, determinate, usually spherical, smooth, distal fertile part thin–walled, subhyaline to pale brown, proximal sterile part dark brown, thick–walled, produced conidia in multiple planes. Conidia 10–15 × 5–7 μm (x = 11.4 × 6.2 μm, n = 40) dry, in simple or branched chains arising from the surface of the upper half of the characteristic conidiogenous cells, cylindrical with rounded ends, ellipsoidal or subspherical, brown or dark brown, minutely verruculose, 1–3 transverse septa, usually strongly constricted at the septa; conidial chains arranged in multiple planes.

    Culture characteristics – Conidia germinating on PDA within 18 hrs and germ tubes produced from the tip cell. Colonies growing on PDA, reaching 50 mm diam. in 14 days at 25℃. Mycelium partly immersed to superficial, slightly effuse, and hairy, with dentate margin, pale pink at periphery golden brown in the middle with whitish hairy mycelial clumps. Sporulation was not observed in mature cultures.

    Material examined – THAILAND, Chiang Mai Province, Mae Taeng District, on a dead leaf vein of Musa sp. (Musaceae), 15 February 2019, B.C. Samarakoon, BNS083 (MFLU 20-0698), living culture MFLUCC 20-0237.

    Known hosts & distribution – on Chromolaena odorata (Asteraceae, Dicotyledon) from Thailand (Li et al. 2017, Mapook et al. 2020); on Clematis fulvicoma, (Ranunculaceae, Dicotyledon) from Thailand (Phukhamsakda et al. 2020); on Pandanus tectorius (Pandanaceae, Monocotyledon) from Yunnan, China (Tibpromma et al. 2018); on herbaceous litter (Dicotyledon) from Yunnan, China (Hyde et al. 2020a); Musa sp. (Monocotyledon) from Thailand (this study).

    Notes – The BLASTn search results of SSU, LSU, ITS and TEF sequence data, indicated that our strain (MFLUCC 20-0237), showed a high identity to Torula chromolaenae (KUMCC 16–0036) as follows; SSU = 100.00%, LSU = 100.00%, ITS = 100.00%, TEF = 99.5% similarities. Morphological comparison with the type specimen of T. chromolaenae showed that our new collection (MFLU 20-0698/ MFLUCC 20-0237) is typical to T. chromolaenae in having brown or dark brown, minutely verruculose, 1–3–septate conidia. However, our collection (MFLU 20-0698) has slightly smaller conidiophores (2–5 × 2–4 μm vs. 5–6.3 × 3.5–4.6 μm) and conidia (10–15 × 5–7 μm vs. 2.1–16.5 × (3.6–) 4.1–5 μm) with respect to Li et al. (2017). A nucleotide base comparison of ITS, TEF and RPB2 regions also showed that MFLUCC 20–0237 is conspecific with T. chromolaenae (KUMCC 16–0036) (0/502 bp of ITS, 0/787 bp of TEF, and 0/817 bp of RPB2). We thus identify our new collection (MFLU 20–0698) as T. chromolaenae. Previously, T. chromolaenae was reported as a saprobe on dicotyledonous and monocotyledonous hosts from China and Thailand, indicating that the species is not specific on hosts and normally found in tropical region.

    Torula ficiCrous [as 'ficus'], in Crous et al., IMA Fungus 6(1): 192 (2015)     Fig. 6

    Figure 6. 

    Torula fici (MFLU 20-0699). a-c Colonies on dead leaf of Musa sp. d-n Conidiogenesis. o, p Conidial masses. q Conidia (in chain). r-u Disposed conidia. v Colonies on PDA after 28 days. Scale bars: a = 500 μm, b = 100 μm, c = 20 μm, d-u = 5 μm.

    Index Fungorum number: IF816154; Facesoffungi number: FoF 02712

    Saprobic on dead leaf vein of Musa sp. (Musaceae). Sexual morph: Undetermined. Asexual morph: hyphomycetous. Colonies superficial, black, appearing as powdery masses on the host. Mycelium slightly immersed, septate, smooth, pale brown. Conidiophores 1–6 × 1.5–3 μm (x = 4.8 × 1.8 μm, n = 10), micronematous or semi–macronematous, unbranched, straight or slightly flexuous, subhyaline or pale grayish brown, verruculose. Conidiogenous cells 3.5–4 × 2.5–3.5 μm (x = 3.2 × 2.7 μm, n = 10) polyblastic or sometimes monoblastic, terminal, discrete, determinate, brown to dark brown, usually spherical, thin–walled, veruculose, having rough distal fertile part and a notably thick–walled, proximal sterile part. Base of sterile part of the conidiogenous cells that arise from mycelium is notably flat. Conidia (4–) 10–25(–75) × 3.5–4.5 μm (x = 24.8 × 3.8 μm, n = 40) dry, in simple or branched chains arising from the surface of the upper half of the characteristic conidiogenous cells, cylindrical with rounded ends, ellipsoidal or subspherical, brown or dark brown, notably verruculose, 1–3 transverse septa, slightly constricted at the septa, immature conidia hyaline to subhyaline, comparatively thin–walled, mature conidia notably thick–walled.

    Culture characteristics – Conidia germinating on PDA within 20 hrs and germ tubes produced from the tip cell. Colonies growing on PDA, reaching 50 mm diam. in 14 days at 25℃, mycelium partly immersed to superficial, slightly effuse, hairy, with dentate to irregular margin, pinkish white.

    Material examined – CHINA, Yunnan, Xishuangbanna, on a dead leaf vein of Musa sp. (Musaceae), 20 December 2018, D.N Wanasinghe, BNSWN5 (MFLU 20–0699), living culture MFLUCC 20–0238.

    Known hosts & distribution – on Chromolaena odorata (Asteraceae, Dicotyledon) from Thailand (Li et al. 2017, Mapook et al. 2020); on Ficus religiosa (Moraceae, Dicotyledon) from Cuba (Crous et al. 2015); on Garcinia sp. (Clusiaceae, Dicotyledon) from Thailand (Jayasiri et al. 2019); on Magnolia grandiflora (Magnoliaceae, Dicotyledon) from Yunnan, China (Jayasiri et al. 2019); on Olea europaea (Oleaceae, Dicotyledon) from South Africa (Spies et al. 2020); on Pandanus sp. (Pandanaceae, Monocotyledon) from Thailand (Tibpromma et al. 2018); on submerged decaying wood in Yunnan, China (Su et al. 2018); on Musa sp. (Musaceae, Monocotyledon) from Yunnan, China (this study).

    Notes – The BLASTn search results of SSU, LSU, ITS and TEF sequence data, indicated that our strain (MFLUCC 20–0238), has a high identity to Torula fici (KUMCC 16–0038) as follows; SSU = 99.88%, LSU = 100.00%, ITS = 98.99%, TEF = 98.53% similarities. Our new collection (MFLU 20–0699) is morphologically similar to T. fici in having short conidiophores, or reduced to conidiogenous cells, mono– to polyblastic, brown, verruculose conidiogenous cells and brown or dark brown, 1–3–septate conidia (Crous et al. 2015). However, our collection has slightly smaller conidiogenous cells (3.5–4 × 2.5–3.5 μm vs. (5–) 6(–8) × 5(–7) μm) and overlapped size of conidia (4–16 × 3.5–4.5 μm vs. (12–) 13–14(–15) (–19) × 5(–6) μm; Crous et al. 2015). A nucleotide base comparison of ITS and TEF regions also showed that MFLUCC 20–0238 differs from T. fici (CBS 595.96, type strain) in 7/467 bp (1.49%) of ITS and 16/776 bp (2.06%) of TEF. In addition, our strain MFLUCC 20–0238 differs from the strains KUMCC 15–0428 and KUMCC 16–0038 in 3/429 bp (0.69%) and 5/494 bp (1.01%) of ITS, and 15/808 bp (1.8%) and 11/813 bp (1.35%) of TEF, respectively. Nevertheless, our new strain (MFLUCC 20–0238) shows a high variation of TEF sequence with the type strain of T. fici (CBS 595.96; 2.06%) and strain KUMCC 15–0428 (1.8%). Based on current phylogenetic results (Fig. 2), we thus identify our new collection (MFLU 20–0699/ MFLUCC 20–0238) as T. fici. The conspecific of these strains is needed to be clarified in further studies. In this study, we report T. fici on Musa sp. (Monocotyledon) from Yunnan, China for the first time.

    Torula masoniiCrous, in Crous et al., IMA Fungus 6(1): 195 (2015)          Fig. 7

    Figure 7. 

    Torula masonii (MFLU 20–0700). a Colonies on dead leaf of Musa sp. b, c Masses of conidia on host surface. d–h, j–o Conidia. i Conidiophore, conidiogenous cell and budding of conidia. p Colonies on PDA after 28 days. Scale bars: a = 50 μm, b = 500 μm, c = 1000 μm, d, f–o = 10 μm, e = 5 μm.

    Index Fungorum number: IF812806; Facesoffungi number: FoF 02711

    Saprobic on a dead leaf of Musa sp. (Musaceae). Sexual morph: Undetermined. Asexual morph: hyphomycetous. Colonies superficial, appearing as black, powdery masses on the host. Mycelium slightly immersed, septate, smooth, grayish brown. Conidiophores 3–6.5 × 4–5 μm (x = 5.4 × 4.3 μm, n = 10), micronematous or semi–macronematous, unbranched, straight or slightly flexuous, pale grayish brown to brown, subcylindrical, septate, or reduced to conidiogenous cells. Conidiogenous cells 5.5–6.5 × 4.5–6 μm (x = 6.3 × 5.3 μm, n = 10) polyblastic or sometimes monoblastic, terminal, discrete, determinate, grayish brown to dark brown, darker than the conidiophores or conidia, spherical, cup–shaped, or inverted cup–shaped, thin–walled, smooth, pale brown of distal fertile part and subhyaline or pale grayish brown of proximal sterile part. Conidia 8–25 × 4–6.5 μm (x = 18.3 × 5.6 μm, n = 40), dry, in simple or branched chains arising from the surface of the upper half of the characteristic conidiogenous cells, cylindrical with rounded ends, ellipsoidal or subspherical in each cell, grayish brown to dark brown, (2–) 3–5(–12) transverse septa, notably constricted at the septa, verruculose, arranged in different planes with respect to the main axis.

    Culture characteristics – Conidia germinating on PDA within 14 hrs and germ tubes produced from the apex. Colonies growing on PDA, reaching 50 mm diam. in 15 days at 25℃, mycelium partly immersed to superficial, slightly effuse and hairy. Colonies medium dense, circular, flattened, slightly raise, surface smooth, with edge entire to sinuate margin, hairy at the center and margin, with dense, floccose to cottony at the middle, three distinct colour zones present in the culture. Periphery pinkish white, middle gray and grayish white next.

    Material examined – Taiwan (China), Alishan, on a dead leaf of Musa sp. (Musaceae), 20 September 2018, D.S. Tennakoon, BNS023 (MFLU 20–0700), living culture MFLUCC 20–0239.

    Known host & distribution – on Brassica sp. (Brassicaceae, Dicotyledon) from United Kingdom (Crous et al. 2015); Iris germanica (Iridaceae, Monocotyledon) from Italy (Li et al. 2017); on submerged decaying wood (Dicotyledon) from Yunnan, China (Su et al. 2018); on Musa sp. (Musaceae, Monocotyledon) from Taiwan (China, this study).

    Notes – The BLASTn search results of SSU, LSU, ITS, TEF and RPB2 sequence data, showed that our strain (MFLUCC 20-0239), is similar to taxa in GenBank as follows; SSU = 99.87% similarity to Torula chromolaenae (MFLUCC 17-1514), LSU = 100.00% similarity to T. masonii (CBS 245.57), ITS = 99.88% similarity to T. masonii (KUMCC 16–0033), TEF = 100% and RPB2 = 99.75% similarities to T. masonii (DLUCC 0588). Comparing with the type specimen, our new isolate (MFLU 20–0700) is morphologically similar to T. masonii; however, the type of T. masonii has slightly larger size of conidiophores and conidia. The conidia of the type of T. masonii are predominantly 6–septate, but also vary in conidial septation from 2– to 12–septate (Crous et al. 2015), whereas the conidia of our new isolate are predominantly 3–5–sepate, occasionally 2– to 12–septate. A nucleotide base pair comparison of ITS showed that our strain (MFLUCC 20-0239) differs from other T. masonii strains (CBS 245.57, DLUCC 0588 and KUMCC 16–0033) in 8/509 (1.57%), 2/441 (0.45%) and 1/508 (0.19%). Our strain MFLUCC 20-0239 also differs from the strains DLUCC 0588 and KUMCC 16–0033 in 2/798 bp (0.25%) and 2/830 (0.34%) of RPB2 but has no different nucleotide base in TEF gene region. We identify our new isolate as T. masonii, based on phylogenetic evidence, although there is more than 1.5% variation in the ITS DNA regions as compared to the type strain (CBS 245.57). We document T. masonii on Musa sp. for the first time. This is the first geographical record of T. masonii from Taiwan (China).

Discussion

  • Many Torula and Periconia species have been reported on Musa spp. worldwide. Torula herbarum has been reported on Musa spp. from Papua New Guinea, Somalia, Taiwan (China), Thailand, and Zambia (Castellani & Ciferri 1937, Riley 1956, Matsushima 1971, 1980, Photita et al. 2001a, 2003b). Periconia byssoides (Venezuela, Cuba, and Somalia) (Matsushima 1971, Urtiaga 1986, Delgado–Rodriguez & Mena–Portales 2004), P. digitata (Thailand, Malaysia) (Williams & Liu 1976, Photita et al. 2001b), P. lateralis (Thailand) (Photita et al. 2001b) and P. minutissima (Ghana) (Hughes 1953) have also been reported on Musa spp. The identification of the latter taxa on Musa spp. was solely based on morphology and molecular data were not integrated. Therefore, more taxon sampling of saprobic and endophytic fungi on Musa spp. should be carried out by integrating morpho–molecular data in taxonomy.

    Some Periconia species have been reported as plant pathogens (i.e., P. cicirnata, P. digitata and P. macrospinosa) on leaves, roots and stems of economically important crops such as maize, sorghum and pointed gourd (Stojkov et al. 1996, Sarkar et al. 2019). In addition, P. keratitis has been reported as a human pathogen from India (Gunasekaran et al. 2020). Periconia produce some economically important bioactive compounds with antimicrobial activities (Kim et al. 2004, Bhilabutra et al. 2007, Hongsanan et al. 2020). It is interesting to note that P. delonicis and P. cortaderiae may also produce bioactive compounds which were discovered from taxa in the same genus. In addition, some species of Torula also produce chemically active compounds (i.e., Dichlorinated Aromatic Lactones and erythritol) which have a wide range of applications in the food industry (Chunyu et al. 2018). Mapook et al. (2020) also reported that T. chromolaenae, T. fici and T. polyseptata showed antimicrobial activity against Bacillus subtilis, Escherichia coli and Mucor plumbeus on their preliminary screening of antimicrobial activity of fungi on Chromolaena odorata. Therefore, it will be interesting to know whether T. chromolaenae and T. fici from Musa spp. will have the same biological ability.

    Recent taxonomic studies integrated DNA sequence data on the introduction of novel taxa in Periconia and Torula (Crous et al. 2015, Su et al. 2016, 2018, Li et al. 2017, 2020, Liu et al. 2017, Jayasiri et al. 2019, Hyde et al. 2020a, Mapook et al. 2020, Phukhamsakda et al. 2020). Protein– coding genes revealed to be good phylogenetic markers in species delineation of Periconia and Torula (Su et al. 2016, 2018, Li et al. 2017, 2020, Jayasiri et al. 2019, Hyde et al. 2020a, Mapook et al. 2020, Phukhamsakda et al. 2020). Corresponding protein–coding sequences (RPB2 and TEF) with ribosomal DNA (SSU, LSU and ITS) sequence dataset can provide well–resolved tree topologies for the taxa in Periconiaceae and Torulaceae. However, several taxa of Periconiaceae and Torulaceae lack protein–coding DNA sequences in GenBank. Therefore, many Periconia and Torula taxa need to be recollected so that valid sequence data are provided to GenBank for better taxonomic resolutions.

    Phylogenetic tree of Periconiaceae (Fig. 1) showed that Periconia delonicis does not form a well–separated clade with P. palmicola (MFLUCC 14-0400) and P. verrucosa (MFLUCC 17-2158). The tree topology showed that they are conspecific, and this result is also supported by a nucleotide base comparison of ITS and TEF regions. However, P. delonicis is phylogenetically well–resolved and is distinct from P. verrucosa in Phukhamsakda et al. (2020), indicating that ITS and TEF regions are not good phylogenetic markers for some species in Periconiaceae. Periconia delonicis was not included in phylogenetic analyses of Periconiaceae when Hyde et al. (2020a) introduced P. palmicola as a new species. The conspecific of P. delonicis and P. palmicola is therefore questionable and should be reinvestigated in future studies.

    Phylogenetic tree of Torulaceae (Fig. 2) showed that Torula chromolaenae, T. fici and T. masonii form well–resolved subclades within Torulaceae. However, the four strains (including type strain) of T. fici and T. masonii formed insignificantly separated branch lengths in this analysis and this phylogenetic result is also supported by Hongsanan et al. (2020), Hyde et al. (2020a), Li et al. (2020), Mapook et al. (2020) and Phukhamsakda et al. (2020). This may be the result of a high variation (> 1.5%) in the TEF region (see notes under T. fici and T. masonii). Further studies on the conspecific or complexity of these species are needed for their clarification based on the reliable protein coding genes.

    Documentation of fungi from new hosts and geographical locations supports the accurate estimates and taxonomic establishments of fungal diversity and distribution. In addition, new occurrences of fungi from various hosts and habitats further provide insights to determine host jumping patterns, host shift speciation and the adaptations of fungi during their life cycle (Hyde et al. 2020c). Taxonomy and phylogeny of fungal pathogens on Musa spp. (i.e., Colletotrichum, Fusarium, Mycosphaerella, Neocordana and Phyllosticta) have been well–studied worldwide (Giatgong 1980, Wulandari et al. 2010, Churchill 2011, Guarnaccia et al. 2017, Marin–Felix et al. 2019, Maryani et al. 2019). The detailed taxonomic works on endophytic fungi in Musa spp. were previously conducted by Brown et al. (1998), Photita et al.(2001b, 2004), Zakaria & Aziz (2018) and Samarakoon et al. (2019). Still the saprobic fungal niches on Musa spp. remain unrevealed and many more taxa are yet to be discovered. Hence the morpho–molecular data of this study will be further useful in future taxonomic works of fungi.

    Acknowledgments

    • This study is supported by Key Research Project "Agroforestry Systems for restoration and bio–industry technology development (grant no. 2017YFC0505101)". We also thank Biology Experimental Center, Germplasm Bank of Wild Species, Kunming Institute of Botany, Chinese Academy of Sciences for providing the facilities of molecular laboratory. Binu C. Samarakoon is grateful to Danushka Tennakoon for collecting the specimens from Taiwan (China), Dr. Dhanushka N. Wanasinghe and Junfu Li for the valuable comments and suggestions on the morphological studies of Periconia and Torula. Rungtiwa Phookamsak thanks CAS President's International Fellowship Initiative (PIFI) for young staff (grant no. Y9215811Q1), the National Science Foundation of China (NSFC) project code 31850410489 (grant no. Y81I982211) and Chiang Mai University for financial support. Samantha C. Karunarathna thanks CAS President's International Fellowship Initiative (PIFI) young staff under the grant number: 2020FYC0002 and the National Science Foundation of China (NSFC) for funding this work under the project code 31851110759. Jianchu Xu thanks Key Research Program of Frontier Sciences of the Chinese Academy of Sciences (grant no. QYZDY–SSW–SMC014) and the Strategic Priority Research Program of Chinese Academy of Sciences for supporting this research.

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    BC Samarakoon, R Phookamsak, SC Karunarathna, R Jeewon, P Chomnunti, JC Xu, YJ Li. 2021. New host and geographic records of five pleosporalean hyphomycetes associated with Musa spp. (Banana). Studies in Fungi 6(1):92−115 doi: 10.5943/sif/6/1/5
    BC Samarakoon, R Phookamsak, SC Karunarathna, R Jeewon, P Chomnunti, JC Xu, YJ Li. 2021. New host and geographic records of five pleosporalean hyphomycetes associated with Musa spp. (Banana). Studies in Fungi 6(1):92−115 doi: 10.5943/sif/6/1/5
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    • Introduction
    • Materials and methods

    • Fungal collections, isolation and morphological characterization

    • Dna extraction and pcr amplification

    • Sequence alignment

    • Phylogenetic analyses

    • Results

    • Phylogenetic analyses

    • Taxonomy

    • Discussion
    • Acknowledgments
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