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Research Article
Taxonomy and phylogenetic appraisal of Spegazzinia musae sp. nov. and S. deightonii (Didymosphaeriaceae, Pleosporales) on Musaceae from Thailand
expand article infoBinu C. Samarakoon§|, Rungtiwa Phookamsak|#¤, Dhanushka N. Wanasinghe|#, Putarak Chomnunti§, Kevin D. Hyde§|#, Eric H. C. Mckenzie«, Itthayakorn Promputtha», Jian-Chu Xu|, Yun-Ju Li˄
‡ The State Phosphorus Resource Development and Utilization Engineering Technology Research Centre, Chiang Rai, Thailand
§ Mae Fah Luang University, Chiang Rai, Thailand
| Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China
¶ World Agroforestry Centre, Kunming, China
# Zhongkai University of Agriculture and Engineering, Guang Dong, China
¤ Chiang Mai University, Chiang Mai, Thailand
« Landcare Research Manaaki Whenua, Christchurch, New Zealand
» Chiang Mai University, Chiang Mai, China
˄ The State Phosphorus Resource Development and Utilization Engineering Technology Research Centre, Kunming, China
Open Access

Abstract

Tropical plants host a range of fungal niches including endophytes, pathogens, epiphytes and saprobes. A study undertaken to discover the saprobic fungal species associated with Musa sp. (banana) from northern Thailand found two hyphomycetous taxa of Spegazzinia (Didymosphaeriaceae, Pleosporales). These were collected during the dry season and their morpho-molecular taxonomic relationships were investigated. Based on phylogenetic analysis of combined SSU, LSU, ITS and TEF1-α sequence data (77% ML, 0.99 BYPP) and contrasting morphological features to the sister taxon, we introduce Spegazzinia musae as a novel species from a decaying leaf of Musa sp. Details on the taxonomy, ecology and geographical distribution of Spegazzinia species are provided. In addition, we report S. deightonii as a new host record from Musa sp. Our data further validate the taxonomic placement of Spegazzinia in Didymosphaeriaceae.

Keywords

Ascomycota, Dothideomycetes, fungi on banana, Hyphomycetes, Thai mycobiota

Introduction

Several taxonomic studies have been conducted to assess the saprobic fungal diversity in Musa species (Ellis 1971, 1976; Matsushima 1971; Photita et al. 2001b; Somrithipol 2007; Hernández-Restrepo et al. 2015; Crous et al. 2016; Hyde et al. 2017). Ellis (1971) described several species on Musa (i.e. Arthrinium sacchari, Cladosporium musae, Cordana musae, Curvularia fallax, Deightoniella torulosa, Gliomastix elata, G. murorum var. polychroma, G. musicola, Gyrothrix hughesii, Haplobasidion musae, Memnoniella subsimplex, Periconia digitata, P. lateralis, Periconiella musae, Pithomyces sacchari, Pyriculariopsis parasitica, Spegazzinia tessarthra, Stachylidium bicolor, Tetraploa aristata, Zygosporium gibbum, Z. masonii and Z. minus). Ellis (1976) also described Bidenticula cannae, Chlamydomyces palmarum, Cordana johnstonii, Parapyricularia musae and Veronaea musae on Musa sp. Photita et al. (2001b) identified 46 saprobic fungal taxa from Musa acuminata in Hong Kong. Most of the saprobes reported by Photita et al. (2001b) belonged to the genera Anthostomella, Deightoniella, Durispora, Hansfordia, Memnoniella, Nigrospora, Pyriculariopsis, Pseudopithomyces, Verticillium and Zygosporium. In addition, Dictyoarthrinium (Somrithipol 2007) and Ramichloridium (Kirschner and Piepenbring 2014) were also recorded as saprobes on Musa sp. Considering the economic importance of Musa sp. there are not many studies on the saprobic fungal populations associated with this host. Few studies have molecular data for the identified strains. To address this research gap, we are investigating the saprobic fungal diversity of Musa sp. in the Asian region where the fungi are highly diverse (Hyde et al. 2018).

Spegazzinia was established by Saccardo (1880) based on S. ornata. Currently 17 taxa are listed in Species Fungorum (2020). Based on morphology, the genus was placed in Apiosporaceae (Sordariomycetes) by Hyde et al. (1998). Based on SSU, LSU, ITS and TEF1-α sequence data of S. deightonii and S. tessarthra, Tanaka et al. (2015) placed Spegazzinia in Didymosphaeriaceae (Dothideomycetes). This was supported by a phylogenetic analysis which placed Spegazzinia in a basal clade in Didymosphaeriaceae (Thambugala et al. 2017).

Hughes (1953) characterized Spegazzinia as a hypomycetous taxon with a unique basauxic conidiophore ontogeny (conidiophores that arise and elongate from a cup-shaped basal cell called a conidiophore mother cell). The conidia of Spegazzinia are brown to dark brown and dimorphic in most species, with a disc-shaped form and a stellate form (Ellis 1971; Manoharachary and Kunwar 2010). However, little molecular data for this genus is available in the GenBank (https://www.ncbi.nlm.nih.gov/). Therefore, for a better phylogenetic resolution of the genus in Didymosphaeriaceae, the previously identified taxa should be recollected to obtain DNA sequence data and morphological descriptions.

In this present study, we introduce Spegazzinia musae sp. nov. and report the first occurrence of Spegazzinia deightonii from Musa sp. in Thailand. We provide detailed morphological descriptions, illustrations and molecular justification for the introduction of Spegazzinia musae sp. nov. Our molecular analyses further support the phylogenetic placement of Spegazzinia in Didymosphaeriaceae.

Materials and methods

Sample collection, morphological studies and isolation

Dead plant materials of Musa sp. (banana) were collected from Thailand during the dry season of 2018 to 2019. Specimens were transferred to the laboratory in cardboard boxes. Samples were examined with a Motic SMZ 168 Series microscope. Powdery masses of conidia were mounted in water for microscopic studies and photomicrography. The 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 CS3 Extended version 10.0 software (Adobe Systems, USA).

Single spore isolation was carried out following the method described in Chomnunti et al. (2014). Germinated spores were individually transferred to potato dextrose agar (PDA) plates and grown at 25 °C in daylight. Colony characteristics were observed and measured after 3 weeks. Specimens were deposited in the Mae Fah Luang University (MFLU) Herbarium, Chiang Rai, Thailand. Living cultures were deposited in the Culture Collection of Mae Fah Luang University (MFLUCC).

DNA extraction and PCR amplification

Fungal isolates were grown on PDA for 4 weeks at 25 °C and total genomic DNA was extracted from 50 to 100 mg of axenic mycelium of the growing cultures according to Wanasinghe et al. (2018). The mycelium was 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. Four gene regions, the internal transcribed spacer (ITS), partial 18S small subunit (SSU), partial 28S large subunit (LSU), and partial translation elongation factor 1-alpha gene (TEF1-α) were amplified using ITS5/ITS4 (White et al. 1990), NS1/NS4 (White et al. 1990), LR0R/LR5 (Vilgalys and Hester 1990) and EF1-983F/EF1-2218R (Rehner 2001) primers, respectively.

Polymerase chain reaction (PCR) was conducted according to the following protocol. The total volume of the PCR reaction was 25 μL containing 12.5 μL of 2 × Power Taq PCR MasterMix (a premix and ready to use solution, including 0.1 Units/ μL Taq DNA Polymerase, 500 μm dNTP Mixture each (dATP, dCTP, dGTP, dTTP), 20 mM Tris-HCL pH 8.3, 100 mM KCl, 3 mM MgCl2, stabilizer and enhancer), 1 μL of each primer (10 pM), 2 μL genomic DNA template and 8.5 μL double distilled water (ddH2O). The reaction was conducted by running for 40 cycles. The annealing temperature was 56 °C for ITS and LSU, 57.2 °C for TEF1-α and 55 °C for SSU and initially 95 °C for 3 mins, denaturation at 95 °C for 30 seconds, annealing for 1 min, elongation at 72 °C for 30 seconds, and final extension at 72 °C for 10 mins for all gene regions. PCR amplification was confirmed on 1% agarose electrophoresis gels stained with ethidium bromide. The amplified PCR fragments were sent to a commercial sequencing provider (TsingKe Biological Technology (Beijing) Co., Ltd, China). The nucleotide sequence data acquired were deposited in GenBank.

Sequencing and sequence alignment

Obtained sequences were subjected to BLASTn search in GenBank (https://blast.ncbi.nlm.nih.gov/Blast.cgi). BLASTn search results and initial morphological studies supported that our isolates belonged to Didymosphaeriaceae. Other sequences used in the analyses were obtained from GenBank based on recently published data (Tanaka et al. 2015; Jayasiri et al. 2019) (Table 1). The single gene alignments were automatically done by MAFFT v. 7.036 (http://mafft.cbrc.jp/alignment/server/index.html, Katoh et al. 2019) using the default settings and later refined where necessary, using BioEdit v. 7.0.5.2 (Hall 1999). The finalized alignment and tree were submitted to TreeBASE (submission ID: 25686, http://www.treebase.org/).

Table 1.

Taxa used in the phylogenetic analysis of Spegazzinia with the corresponding GenBank accession numbers. Type strains are superscripted with T and newly generated strains are indicated in bold.

Species Strains * GenBank accession numbers References
LSU SSU ITS TEF1-α
Alloconiothyrium aptrootii CBS 980.95T JX496234 JX496121 Verkley et al. (2014)
Bimuria novae zelandiae CBS 107.79T AY016356 AY016338 Lumbsch and Lindemuth (2001)
Dendrothyrium variisporum CBS 121517T JX496143 JX496030 Vu et al. (2019)
Deniquelata barringtoniae MFLUCC 110422T JX254655 JX254656 JX254654 Ariyawansa et al. (2013)
Didymocrea sadasivanii CBS 438.65T DQ384103 DQ384066 Vu et al. (2019)
Didymosphaeria rubi ulmifolii MFLUCC 14-0023T KJ436586 KJ436588 Ariyawansa et al. (2014)
Kalmusia spartii MFLUCC 14-0560T KP744487 KP753953 KP744441 Liu et al. (2015)
Karstenula rhodostoma CBS 690.94 GU301821 GU296154 Schoch et al. (2009)
Laburnicola muriformis MFLUCC 16-0290T KU743198 KU743199 KU743197 KU743213 Wanasinghe et al. (2016)
Montagnula cirsii MFLUCC 13-0680T KX274249 KX274255 KX274242 KX284707 Hyde et al. (2016)
Montagnula graminicola MFLUCC 13-0352T KM658315 KM658316 KM658314 Liu et al. (2015)
Neokalmusia brevispora KT 2313T AB524601 AB524460 AB539113 Tanaka et al. (2009)
Neokalmusia scabrispora KT 2202 AB524594 AB524453 AB539107 Tanaka et al. (2009)
Paracamarosporium hawaiiense CBS 120025T JX496140 EU295655 JX496027 Verkley et al. (2014)
Paraconiothyrium cyclothyrioides CBS 972.95T JX496232 AY642524 JX496119 Verkley et al. (2014)
Paraconiothyrium estuarinum CBS 109850T JX496129 AY642522 JX496016 Verkley et al. (2014)
Paramassariosphaeria clematidicola MFLU 16-0172T KU743207 KU743208 KU743206 Wanasinghe et al. (2016)
Paraphaeosphaeria michotii MFLUCC 13-0349T KJ939282 KJ939285 KJ939279 Tennakoon et al. (2016)
Phaeodothis winteri AFTOL-ID 1590 DQ678073 DQ678021 DQ677917 Schoch et al. (2006)
Pleospora herbarum CBS 191.86T GU238160 GU238232 KC584731 Aveskamp et al. (2010)
Pseudocamarosporium cotinae MFLUCC 14-0624T KP744505 KP753964 KP744460 Liu et al. (2015)
Pseudocamarosporium propinquum MFLUCC 13-0544T KJ813280 KJ819949 KJ747049 Wijayawardene et al. (2014)
Pseudopithomyces chartarum UTHSC 04-678 HG518065 HG518060 Da Cunha et al. (2014)
Spegazzinia bromeliacearum URM 8084T MK809513 MK804501 Crous et al. (2019)
Spegazzinia deightonii yone 212 AB807582 AB797292 AB808558 Tanaka et al. (2015)
Spegazzinia deightonii MFLUCC 20-0002 MN956772 MN956770 MN956768 MN927133 This study
Spegazzinia deightonii yone 66 AB807581 AB797291 AB808557 Tanaka et al. (2015)
Spegazzinia intermedia CBS 249.89 MH873861 MH862171 Vu et al. (2019)
Spegazzinia lobulata CBS 361.58 MH869344 MH857812 Vu et al. (2019)
Spegazzinia musae MFLUCC 20-0001T MN930514 MN930513 MN930512 MN927132 This study
Spegazzinia neosundara MFLUCC 15–0456T KX954397 KX986341 KX965728 Thambugala et al. (2017)
Spegazzinia radermacherae MFLUCC 17-2285T NG_066308 MK347848 NR_163331 MK360088 Jayasiri et al. (2019)
Spegazzinia sp. yone 279 AB807583 AB797293 AB808559 Tanaka et al. (2015)
Spegazzinia tessarthra SH 287 AB807584 AB797294 AB808560 Tanaka et al. (2015)
Stemphylium botryosum CBS 714.68T KC584345 KC584603 KC584238 KC584729 Woudenberg et al. (2013)
Tremateia arundicola MFLU 16-1275T KX274248 KX274254 KX274241 KX284706 Tennakoon et al. (2016)
Tremateia guiyangensis GZAAS01T KX274247 KX274253 KX274240 KX284705 Tennakoon et al. (2016)
Xenocamarosporium acaciae CPC 24755T KR476759 KR476724 Tennakoon et al. (2016)

Phylogenetic analysis

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. Bootstrap support was obtained by running 1000 pseudo-replicates. Maximum likelihood bootstrap values (ML) equal or greater than 60% are given above each node in blue (Figure 1).

Figure 1. 

Maximum likelihood tree revealed by RAxML from an analysis of SSU, LSU and ITS and TEF1-α sequence data of selected genera of family Didymosphaeriaceae, showing the phylogenetic position of Spegazzinia musae (MFLUCC 20-0001) and S. deightonii (MFLUCC 20-0002). ML bootstrap supports (≥60 %) and Bayesian posterior probabilities (≥ 0.95 BYPP) are given above in the branches, respectively. The tree was rooted with Pleospora herbarum and Stemphylium botryosum (Pleosporaceae). Strains generated in this study are indicated in red-bold. Ex-type species are indicated in bold. The scale bar represents the expected number of nucleotide substitutions per site. A best scoring RAxML tree is shown with a final ML optimization likelihood value of -13516.66. The matrix had 795 distinct alignment patterns, with 33.60% of undetermined characters or gaps. Estimated base frequencies were: A = 0.239862, C = 0.245185, G = 0.277025, T = 0.237927; substitution rates AC = 1.626982, AG = 2.468452, AT = 1.211822, CG = 1.092437, CT = 6.295657, GT = 1.000000; proportion of invariable sites I = 0.484119; gamma distribution shape parameter α = 0.445929.

A Bayesian inference analysis was conducted with MrBayes v. 3.1.2 (Huelsenbeck and Ronquist 2001) to evaluate posterior probabilities (PP) (Rannala and Yang 1996; Zhaxybayeva and 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 and trees were sampled every 100th generation and 20,000 trees were obtained. The first 4,000 trees, representing the burning phase of the analyses were discarded. The remaining 16,000 trees were used for calculating PP in the majority rule consensus tree. Branches with Bayesian posterior probabilities (BYPP) greater than 0.95 are indicated above each node in blue (Figure 1). Phylograms were visualized with FigTree v1.4.0 program (Rambaut 2011) and reorganized in Microsoft Power Point.

Data resources

The data underpinning the analysis reported in this paper are deposited in the Dryad Data Repository at https://doi.org/10.5061/dryad.2ngf1vhk6.

Results

Phylogenetic analysis

The combined SSU, LSU, ITS, TEF1-α matrix comprised 38 sequences including selected genera in Didymosphaeriaceae. A best scoring RAxML tree is shown in Figure 1. All trees (ML and BYPP) were similar in topology and did not differ (data not shown) at the generic relationships which are in agreement with multi-gene phylogeny of Tanaka et al. (2015). All Spegazzinia strains analyzed here were clustered as a highly supported monophyletic clade (100% ML, 1.00 BYPP) in Didymosphaeriaceae (Figure 1) sister to Alloconiothyrium, Dendrothyrium, Laburnicola and Xenocamarosporium. Our new species, Spegazzinia musae (MFLUCC 20-0001) clustered with Spegazzinia sp. (yone 279) and S. deightonii (yone 66, MFLUCC 20-0002, yone 212) with significant statistical support (77% ML, 0.99 BYPP). Strain MFLUCC 20-0002 grouped with S. deightonii (yone 66, yone 212) with high statistical support (96% ML, 0.99 BYPP).

Taxonomy

Spegazzinia deightonii (S. Hughes) Subram., J. Indian bot. Soc. 35: 78 (1956)

Figure 2

Description

Saprobic on dead leaves of Musa sp. Sexual morph Undetermined. Asexual morph Hyphomycetous. Sporodochia powder like, dark, dense, dry, 1–3 mm diameter. Conidiophore mother cells 3.5–6.8 × 2.5–5.0 μm ( = 5.59 × 4.15 μm, n = 6), hyaline to light brown, subspherical or doliiform. Conidiophores long or short and give rise to two types of conidia referred here as α and β. Conidiophores of α conidia up to 48–120 × 1–2 μm ( = 95.3 × 1.6 μm, n = 20) long, erect or flexuous, narrow, verruculose, unbranched, hyaline to golden-brown. Conidiophores of β conidia initially hyaline, light brown to brown at maturity, very short and slightly bent, 1.6–2 × 2.5–3 μm ( = 1.8 × 2.6 μm, n =10). Conidiogenous cell development basauxic, forming a single, terminal holoblastic conidium at the apex of conidiophore. Conidial development holoblastic. Conidia two types: α conidia stellate, 18–28 × 17–29 μm ( = 25.1 × 23.3 μm, n = 25), solitary, globose to variously shaped, with spines 4–6 μm long, 4–8-celled, frequently 4- to 6-celled, deeply constricted at the septa. β conidia disc-shaped, initially hyaline, light brown to dark brown at maturity, 8-celled, 16–21 × 11–14 μm ( = 19.2 × 14.6 μm, n = 25), flat from both sides with short and blunt spines, frequently with attached conidiogenous cells when splitting from the conidiophores.

Culture characteristics

Conidia germinating on PDA within 13–14 h. Colonies growing on PDA, reaching a diameter of 55 mm after 14 d at 25 °C, raised, moderately dense, undulate margin, middle grey, periphery brownish grey and olive green at immature stage; reverse white to greyish white.

Material examined

Thailand, Chiang Rai Province, Doi Thun, on a dead leaf of Musa sp. (Musaceae), 7 December 2018, M.C. Samarakoon, BNS 072 (MFLU 19-2908), living culture MFLUCC 20-0002.

Notes

Spegazzinia deightonii MFLUCC 20-0002 clustered with S. deightonii (yone 66, yone 212) with significant statistical support (Figure 1). All the strains of S. deightonii described in Ellis (1971) and Tanaka et al. (2015) have similar morphological features with our strain such as dark brown, 8-celled, disked-shaped, spiny conidia. With morphological and multigene phylogenetic support, we report a new host record of S. deightonii from Musa sp.

Figure 2. 

Spegazzinia deightonii (MFLU 19-2908) a–c fungal colonies on host surface d conidiophore mother cell of α conidia e–g α conidia i a developmental stage of β conidia h, k conidia l colonies on PDA after 28 days showing sporulation j, m–p β conidia. Scale bars: 500μm (a), 200μm (b), 50 μm (c), 20μm (e–h), 10μm (d, k, m–p), 5 μm (i, j).

Spegazzinia musae Samarakoon, Phookamsak, Wanas., Chomnunti & K.D. Hyde, sp. nov.

MycoBank No: 835298
Figure 3

Etymology

The name reflects the host genus, Musa (Musaceae).

Holotype

MFLU 19-2907

Description

Saprobic on a dead leaf of Musa sp. Sexual morph Undetermined. Asexual morph Hyphomycetous. Sporodochia dark, dense, dry, powdery, velvety, 1–2 mm diameter. Conidiophore mother cells 3.4–5.8 × 3.7–4.7 μm ( = 4.6 × 4.1 μm, n = 10) subhyaline or light brown, doliiform or subspherical. Conidiophores usually short to long bearing two types of conidia referred to here as α and β. Conidiophores of α conidia up to 40–85 × 0.8–2.5 μm ( = 64 × 21.7 μm, n = 15), pale brown or dark golden brown, rough-walled, hyaline at bottom near the conidiophore mother cell, pale brown at middle, dark golden brown at top near conidial cells, erect or flexuous, narrow and long, generally unbranched, rarely branched. Conidiophores of β conidia 0.7–3.5 × 1.5–3 μm ( = 1.9 × 2.3 μm, n = 15) short, erect, unbranched, hyaline when immature, subhyaline or hyaline at maturity. Conidiogenous cell development basauxic, forming a single, terminal holoblastic conidium at the apex of conidiophore. Conidial development holoblastic. Conidia solitary, dry, two types: α conidia stellate, 15–22.7 × 14.5–20.5 μm ( = 18.8 × 17.8 μm, n = 15), 4–6 celled, each cell globose to subglobose, deeply constricted at the septa, conspicuously spinulate, 4–6 spines, each 2–8 μm long arise from surface of each cell. β conidia disc-shaped, initially hyaline, 4-celled, each cell slightly turbinate in shape, rough-walled, crossed septate, becoming brown to dark brown at maturity, each cell turbinate, crossed-septate, smooth-walled, light brown at the center near the septa, dark brown at periphery in constricted areas, 9.3–14.2 × 8.4–12.5 μm ( = 12.7 × 10.8 μm, n = 40), somewhat obovoid, deeply constricted at the septa, flat from side view, frequently with attached conidiogenous cells when splitting from the conidiophores.

Culture characteristics

Conidia germinating on PDA within 12–15 h, germ tubes produced from one or several cells. Colonies growing on PDA, reaching a diameter of 46 mm after 14 d at 25 °C, greyish white, unevenly raised, surface rough, moderately dense, radially striated at center, margin crenulate; reverse white to greyish white.

Material examined

Thailand, Nan Province, on a dead leaf of Musa sp. (Musaceae), 12 September 2018, B.C. Samarakoon, BNS 069 (MFLU 19-2907, holotype), ex-type living culture MFLUCC 20-0001.

Notes

Based on BLASTn search results of SSU, LSU, ITS and TEF1-α sequence data, Spegazzinia musae showed a high similarity (SSU = 98.24%, LSU = 98.92%, ITS = 96.91%, TEF1-α = 98.11%) to S. neosundara (MFLUCC 15-0456). In the multigene phylogeny, S. musae groups as a sister taxon to S. deightonii with strong statistical support (77% ML, 0.99 BYPP) (Figure 1). Also, ITS sequence comparison revealed 3.75% base pair differences between S. musae and S. deightonii, which is in agreement with the species concept outlined by Jeewon and Hyde (2016). Besides, S. musae has contrasting morphological features to S. deightonii in both kinds of conidia. The disk-shaped conidia of S. musae are 4-celled and do not bear spines at the periphery of cells, while the disc-shaped conidia of S. deightonii are 8-celled and spiny. Based on contrasting morphological differences and significant statistical support from our molecular phylogeny, Spegazzinia musae is introduced as a new species.

Figure 3. 

Spegazzinia musae (MFLU 19-2907, holotype) a–c fungal colonies on host surface d mature conidia e conidiophore of α conidia with the mother cell f, g α conidia h–q β conidia r colony on PDA after 28 days. Scale bars: 200 μm (a–c), 20 μm (d–g, j), 10 μm (h, i, k–q).

Discussion

Spegazzinia is ubiquitous in the environment. Several taxa of Spegazzinia occur as saprobes on dead material of tropical, subtropical and temperate vascular plants (Ellis 1971; Subramanian 1988; Caretta et al. 1999; Delgado-Rodríguez et al. 2002; Bhat 2010; Leão-Ferreira and Gusmão 2010; Manoharachary and Kunwar 2010). In addition, Spegazzinia was also recorded from soil (Ellis 1971), dredged sediments of marine and brackish estuaries (Borut and Johnson 1962) and grassland vegetation (Caretta et al. 1999). Spegazzinia tessarthra was recorded as an endophyte from lichens (Manish et al. 2014) and recently S. bromeliacearum was introduced as an endophyte from the leaves of Tilandsia catimbauensis (Crous et al. 2019). Damon (1953) considered S. tessarthra to be an important decomposer of monocotyledonous plants and other cellulose containing materials in tropical and subtropical areas. Spegazzinia deightonii was previously recorded on monocotyledons such as Areca catechu (China, Taiwan; Matsushima 1980), Cocos nucifera (China; Tianyu et al. 2009) and Panicum maximum (Hong Kong; Lu et al. 2000) (Farr and Rossman 2020). Our study presents the first report of Spegazzinia deightonii in Musaceae as a saprobe and introduces our new species, S. musae.

There does not appear to be any host-specificity as the genus is found on a wide range of hosts in various habitats and there are no records of a pathogenic lifestyle. Some Spegazzinia species (such as S. tessarthra) have been identified as saprobes and endophytes and therefore the genus may have the potential of switching nutritional modes during the degradation of plant material (Promputtha et al. 2007).

Spegazzinia is a unique taxon among other dematiaceous hyphomycetes due to its conidial morphology and basauxic conidiogenesis. Most Spegazzinia species have contrasting morphological features in the shapes of α and β conidia. Some taxa bear spines in both types of conidia while some taxa do not bear spines. Simultaneously, some species of Spegazzinia such as S. radermacherae, S. tessarthra show similar characters in morphology apart from dimensions of conidia. The length of conidiophores can be varied with the environmental stresses (Cole 1974). Therefore, the use of morphological data coupled with DNA sequence data (SSU, LSU, ITS and TEF-α) will be crucial for better taxonomic resolutions in this genus.

Dictyoarthrinium (Apiosporaceae) bears some similar morphological features to Spegazzinia such as basauxic conidiogenesis (Ellis 1971) and cross septate, 4-celled, dematiaceous conidia with warts (Rao and Rao 1964). However, generic placement of Dictyoarthrinium in Apiosporaceae was confirmed by Vu et al. (2019) based on the LSU sequence of D. sacchari strain CBS 529.73. Therefore, Dictyoarthrinium was treated as a distinct genus with Spegazzinia (Vu et al. 2019).

Microfungal studies in Musa sp. are mostly oriented towards pathogens and endophytes due to the economic value of the fruit crop. Most of the pathogenic species descriptively studied from Musa sp. are identified as Colletotrichum, Fusarium, Mycosphaerella, Neocordana and Phyllosticta (Giatgong 1980; Wulandari et al. 2010; Churchill 2011; Guarnaccia et al. 2017; Marin-Felix et al. 2019; Maryani et al. 2019). The endophytic fungal populations of Musa sp. were studied by Brown et al. (1998), Photita et al. (2001a, 2004) and Samarakoon et al. (2019). Few studies have documented the saprobic diversity of Musa sp. and as we believe that there are saprobic niches associated with Musa sp. that are still unrevealed, taxonomists should investigate this hidden diversity for conservation purposes.

Acknowledgements

Authors would like to acknowledge Mae Fah Luang University (grant No. DR256201012003) and the grant titled “Impact of climate change on fungal diversity and biogeography in the Greater Mekong Subregion” (grant number: RDG6130001) for financial support. R. 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). R. Phookamsak and I. Promputtha thank Chiang Mai University for their partial support of this research work. D. N. Wanasinghe would like to thank the CAS President’s International Fellowship Initiative (PIFI) for funding his postdoctoral research (number 2019PC0008), the National Science Foundation of China and the Chinese Academy of Sciences for financial support under the following grants: 41761144055, 41771063 and Y4ZK111B01. D.N. Wanasinghe also thanks the 64th batch of China Postdoctoral Science Foundation (grant no.: Y913083271). J.C. Xu thanks the Key Research Program of Frontier Sciences of the Chinese Academy of Sciences (grant no. QYZDY-SSW-SMC014). S.M.B.C. Samarakoon gives her sincere appreciation to Milan Samarakoon, Junfu Li, De-Ping Wei, Achala Jeevani, G. Samarakoon and Kaanchana Senadheera for the great support.

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