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Research Article
Morphology and phylogeny reveal two new species and host records of hyphomycetous fungi on Areca species from marine habitats in Thailand
expand article infoRaheleh Asghari‡, Chayanard Phukhamsakda‡, E. B. Gareth Jones§, Ali Bahkali§, Carlo Chris S. Apurillo|‡, Omid Karimi‡, Pattana Kakumyan‡, Kevin D. Hyde§‡
‡ Mae Fah Luang University, Chiang Rai, Thailand
§ King Saud University, Riyadh, Saudi Arabia
| Philippine Science High School-Eastern Visayas Campus, Palo, Philippines

Corresponding author: Chayanard Phukhamsakda ( chayanard.phu@mfu.ac.th )

Academic editor: Danushka Sandaruwan Tennakoon
© 2025 Raheleh Asghari, Chayanard Phukhamsakda, E. B. Gareth Jones, Ali Bahkali, Carlo Chris S. Apurillo, Omid Karimi, Pattana Kakumyan, Kevin D. Hyde.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Asghari R, Phukhamsakda C, Jones EBG, Bahkali A, Apurillo CCS, Karimi O, Kakumyan P, Hyde KD (2025) Morphology and phylogeny reveal two new species and host records of hyphomycetous fungi on Areca species from marine habitats in Thailand. MycoKeys 118: 179-206. https://doi.org/10.3897/mycokeys.118.147229
Open Access

Abstract

The marine ecosystem is the largest on Earth, supporting a wide variety of organisms. Fungi in this environment are diverse and play promising ecological roles. This study investigated fungi on submerged, decaying plant materials of Areca species trapped between rocks in seawater in Prachuap Khiri Khan Province, Thailand. Morphological and multi-gene phylogenetic analysis (LSU-ITS-tub2-SSU for Tetraploa, ITS-LSU-tub2-rpb2 for Rosellinia, LSU-ITS-tef1-α-rpb2 for Musicillium and LSU-ITS-act for Sarocladium) revealed two new species: Tetraploa maritima, characterized by the presence of a hilum and elongating appendages, and Rosellinia maritima, distinguished by a brown to black conidial-like structure composed of interwoven, irregular cells without distinct conidiophores produced in sterile conditions, unlike other Rosellinia asexual morphs. Additionally, two new host records (Musicillium theobromae and Sarocladium gamsii) are documented, with detailed descriptions of their in vivo and in vitro morphologies. Detailed morphological descriptions and illustrations are provided. This study contributes to the understanding of fungal diversity in marine environments.

Key words:

2 new species, marine fungi, molecular phylogeny, palm fungi, rocky shore, saprobe

Introduction

Fungi live in a variety of ecosystems, from terrestrial to aquatic habitats on various substrates such as plant materials (Hyde et al. 2019, 2020, 2024; Bhunjun et al. 2022). They have a high diversity in marine environments and are known for their important roles as well as for their potential use in medicine, bioremediation, biotechnology and industry (Hyde et al. 1998; Bonugli-Santos et al. 2015; Pang et al. 2016; Jones et al. 2019). Despite their importance, marine fungi have not received much attention as compared to terrestrial fungi (Kohlmeyer and Kohlmeyer 1979; Hyde et al. 2000; Jones et al. 2006, 2015, 2019; Abdel-Wahab et al. 2020; Dayarathne et al. 2020). Currently, more than 2000 marine fungal species have been discovered belonging to 11 different phyla (www.marinefungi.org, accessed in April 2025), with Ascomycota as the most recorded phylum (Jones et al. 2019; Calabon et al. 2023). They are mostly reported in their sexual stages, followed by lesser reports of coelomycetous and hyphomycetous asexual morphs (Hyde et al. 2000; Chatmala et al. 2004; Jones et al. 2009).

Several studies have been conducted on saprobic, endophytic and pathogenic fungi on palms (Arecaceae), showing the high diversity of fungi on this host family worldwide (Hyde 1988; Taylor et al. 1999; Taylor and Hyde 2003; Liu et al. 2010; Marin-Felix et al. 2019; Chen et al. 2020; Konta et al. 2020, 2021, 2023; Karimi et al. 2024; Zhang et al. 2024). Arecaceae is an ancient plant family dating back 66–100 million years (Phukhamsakda et al. 2022). It has a global distribution, predominantly in tropical and subtropical regions, across diverse terrestrial and aquatic habitats (Pereira and Phillips 2023). The palm family comprises around 2600 species within 181 genera (Baker and Dransfield 2016). Areca is the type genus of Arecaceae, described by Linnaeus (1753), and mostly distributed in tropical islands (www.palmweb.org). Areca species thrive in humid tropical environments and are utilized for various purposes, including ornamental and cultural uses (Dransfield et al. 2008; Heatubun et al. 2012).

Fungal association with Areca has been studied predominantly with the focus of pathogenic fungi on the economically important species A. catechu (betel nut palm) (To-Anun et al. 2009; Phengsintham et al. 2013; Balanagouda et al. 2021; Harahap 2022). Asghari et al. (2023) introduced a new fungal species found on decaying wood of Areca species submerged in marine water. In this study, we proceed with the investigation of hyphomycetous saprobic marine fungi on decaying Areca in marine habitats in Thailand. Two new species and two new host records are introduced, supported by morphology and molecular data.

Materials and methods

Sampling and morphological studies

Decaying submerged woody specimens were collected from the rocky shore at Pranburi, Prachuap Khiri Khan Province (Thailand) in October 2022. The specimens were placed in moist plastic bags and incubated in sterile seawater moist chambers after transferring to the laboratory. The moist chambers were treated with sterilized natural seawater regularly. The samples were examined for fungal presence after returning to the laboratory and up to one month of incubation. The fungal macro-characters were examined using a stereomicroscope (Olympus SZX16, Japan) equipped with an Olympus SC180 digital camera (Olympus, Japan). The fungal microscopic features, such as conidiophores, conidiogenous cells and conidia, were examined by preparing microscopic slides using distilled water or lactoglycerol, followed by using a Nikon ECLIPSE Ni compound microscope (Nikon, Japan) equipped with a Nikon DS-Ri2 digital camera (Nikon, Japan). The Tarosoft (R) Image Framework program (Tarosoft, Thailand) was used to measure the fungal characters, and Adobe Photoshop CS6 Extended version 13.1.2 software (Adobe Systems Inc., The United States) was used to process the figures. The single spore isolation method was employed to obtain pure cultures (Senanayake et al. 2020) using potato dextrose agar (PDA). Specimens were deposited at the Mae Fah Luang University Herbarium, Chiang Rai, Thailand (MFLU), and the living cultures were deposited at the Mae Fah Luang University Culture Collection (MFLUCC). The new taxa were linked to the Facesoffungi (Jayasiri et al. 2015) and Index Fungorum databases.

DNA extraction, PCR amplification and sequencing

Fresh, pure cultures were selected for DNA extraction. Mycelium was scraped using sterilized surgical blades. Genomic DNA was extracted following the manufacturer’s protocol of a Mega Genomic DNA Extraction Kit (Omega Bio-tek Inc., The United States). Polymerase chain reactions (PCR) were conducted to amplify the internal transcribed spacer region rDNA (ITS), the 28S large subunit rDNA (LSU), the 18S small subunit rDNA (SSU), the RNA polymerase II second largest subunit (rpb2) gene, the translation elongation factor 1-alpha (tef1-α) gene, and the partial β-tubulin II (tub2). Primers and the PCR condition were described as in Table 1. The total volume of the PCR mixture (25 µL) contained double distilled water (ddH2O) (9.5 µL), 2X GoTaq® green PCR master mix (PROMEGA, Madison, WI, USA) (12.5 µL), DNA template (1 µL), and 1 µL of each primer (20 µM). The PCR products were checked on 1.7% agarose electrophoresis gels stained with 4S Green Stain. The amplified PCR products were sequenced at SolGent Co. (South Korea).

Table 1.
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Primers and PCR conditions used in the study.

Gene Regions Primers PCR conditions References
ITS ITS5/ITS4 95 °C for 5 min, 35 cycles of 94 °C for 45 s, 53 °C for 45 s, and 72 °C for 2 min, 72 °C for 10 min White et al. (1990)
LSU LR0R/LR5 94 °C for 4 min, 35 cycles of 95 °C for 45 s, 56 °C for 45 s, and 72 °C for 1 min, 72 °C for 10 min Vilgalys and Hester (1990);
Rehner and Samuels (1994)
SSU NS1/NS4 94 °C for 4 min, 35 cycles of 94 °C for 30 s, 55 °C for 50 s, and 72 °C for 1.30 min, 72 °C for 10 min White et al. (1990)
rpb2 fRPB2-5f/fRPB2-7cR 95 °C for 5 min, 40 cycles of 95 °C for 1 min, 54 °C, 90 s, and 72 °C for 90 s, 72 °C for 10 min Liu et al. (1999)
tef1-α EF1-983F/EF1-2218R 95 °C for 5 min, 40 cycles of 95 °C for 1 min, 54 °C, 90 s, and 72 °C for 90 s, 72 °C for 10 min Rehner (2001)
tub2 T1/T22 94 °C for 4 min, 35 cycles of 94 °C for 30 s, 55 °C for 50 s, and 72 °C for 1.30 min, 72 °C for 10 min Glass and Donaldson (1995)

Sequence alignment and phylogenetic analyses

The generated sequences were assembled using SeqMan software version 7.1.0. The assembled sequences were searched on the Nucleotide BLAST search (https://blast.ncbi.nlm.nih.gov) to identify the closest taxa. The reference sequences of closely related taxa were obtained from previous publications (Tibpromma et al. 2018; Giraldo and Crous 2019; Hyde et al. 2020; Phukhamsakda et al. 2020; Hou et al. 2023; Li et al. 2024; Zhang et al. 2024; Zhao et al. 2024) and GenBank. The sequences were aligned automatically in MAFFT version 7 (Katoh et al. 2019) and edited manually and/or on the NGPhylogeny.fr website (Lemoine et al. 2019) using the trimAl tool. The datasets were concatenated according to previous research on each genus, using Mesquite version 3.10 (Maddison and Maddison 2016). Maximum likelihood (ML) and Bayesian inference analyses were carried out for both individual loci and combined datasets on the CIPRES Science Gateway portal (CIPRES) (Miller et al. 2010). Maximum likelihood analysis was carried out using RAxMLHPC2 on the XSEDE (v. 8.2.10) tool (Stamatakis 2014) under the GTRCAT substitution model and 1000 bootstrap (B) iterations. Bayesian inference was performed using MrBayes 3.2.2 in CIPRES (Ronquist et al. 2012) to evaluate posterior probabilities (BPP) with four simultaneous Markov chains run for 10,000,000 generation. Trees were sampled every 1000th generations, and the consensus tree and posterior probability were determined after removing the first 25% of the sampled trees in burn-in phase (Larget and Simon 1999). Phylograms were visualized using FigTree version 1.4.0 (Rambaut 2015), and they were adjusted using Adobe Photoshop CS6 Extended version 13.1.2 and Incscape 1.3.3.

Results

Phylogenetic analyses of Tetraploa

The combined analysis of LSU, ITS, tub2 and SSU sequence data of Tetraploa was performed with Muritestudina chiangraiensis (MFLUCC 17-2551) and Montanitestudina hydei (SQUCC 15173) as the outgroup taxa (Fig. 1). The combined dataset contained 3298 characters (LSU = 1080 bp, ITS = 569 bp, tub2 = 638, SSU = 1011) after alignments, including the gaps. Analysis of the combined dataset yielded a best-scoring tree with an optimal log-likelihood value of -12408.104565 (Fig. 1). The alignment had 41 sequences with 812 distinct patterns. Estimated base frequencies were as follows: A = 0.247947, C = 0.249495, G = 0.273749, T = 0.228808, Rate parameters: AC = 3.650751; AG = 3.979381; AT = 2.302782; CG = 1.799753; CT = 9.619686; GT = 1.000000; Gamma shape alpha = 0.122858. After discarding the first 25% of generations in the Bayesian analyses, 7500 trees were remained, from which the 50% consensus tree and posterior probabilities were determined.

Figure 1. 

Phylogram generated from maximum likelihood analysis based on combined LSU, ITS, tub2 and SSU sequence data of Tetraploa. Maximum likelihood bootstrap support ≥ 65% and Bayesian posterior probabilities (BPP) ≥ 0.90 are given near the nodes, respectively. The tree is rooted to Muritestudina chiangraiensis (MFLUCC 17-2551) and Montanitestudina hydei (SQUCC:15173). The newly generated sequences are indicated in red, and the type strains are indicated in bold.

Phylogenetic analyses of Xylariaceae

The combined analysis of ITS, LSU, tub2 and rpb2 sequence data of Xylariaceae was performed with Hypoxylon rickii (MUCL 53309) and H. fragiforme (MUCL 51264) as the outgroup taxa (Fig. 2). The combined dataset contained 5441 characters (ITS = 560 bp, LSU = 860 bp, tub2 = 2900, rpb2 = 1121) after alignments, including the gaps. Analysis of the combined dataset yielded a best-scoring tree with an optimal log-likelihood value of -37958.515421 (Fig. 2). The alignment had 52 sequences with 2206 distinct patterns. Estimated base frequencies were as follows: A = 0.222688, C = 0.282274, G = 0.251124, T = 0.243914, Rate parameters: AC = 1.130042; AG = 4.802871; AT = 1.154978; CG = 0.881707; CT = 6.304393; GT = 1.000000; Gamma shape alpha = 0.238462. After discarding the first 25% of generations in the Bayesian analyses, 7500 trees remained, from which the 50% consensus tree and posterior probabilities were determined.

Figure 2. 

Phylogram generated from maximum likelihood analysis based on combined ITS, LSU, tub2 and rpb2 sequence data of Xylariaceae. Maximum likelihood bootstrap support ≥ 65% and Bayesian posterior probabilities (BPP) ≥ 0.90 are given near the nodes, respectively. The tree is rooted to Hypoxylon rickii (MUCL 53309) and H. fragiforme (MUCL 51264). The newly generated sequences are indicated in red, and the type strains are indicated in bold.

Phylogenetic analyses of Plectosphaerellaceae

The combined analysis of LSU, ITS, tef1-α and rpb2 sequence data of Plectosphaerellaceae was performed with Cylindrotrichum clavatum (CBS 125296, CBS 125297) as the outgroup taxa (Fig. 3). The combined dataset contained 3,269 characters (LSU = 870 bp, ITS = 544 bp, tef1-α = 858, rpb2 = 997) after alignments, including the gaps. Analysis of the combined dataset yielded a best-scoring tree with an optimal log-likelihood value of -18111.606810 (Fig. 4). The alignment had 31 sequences with 1017 distinct patterns. Estimated base frequencies were as follows: A = 0.224951, C = 0.295999, G = 0.282023, T = 0.197027, Rate parameters: AC = 0.696864; AG = 1.667863; AT = 1.053040; CG = 0.729378; CT = 4.900764; GT = 1.000000; Gamma shape alpha = 0.194519. After discarding the first 25% of generations in the Bayesian analyses, 7500 trees were remained from which the 50% consensus tree and posterior probabilities were determined.

Figure 3. 

Phylogram generated from maximum likelihood analysis based on combined LSU, ITS, tef1-α and rpb2 sequence data of Plectosphaerellaceae. Maximum likelihood bootstrap support ≥ 65% and Bayesian posterior probabilities (BPP) ≥ 0.90 are given near the nodes, respectively. The tree is rooted to Cylindrotrichum clavatum (CBS 125296, CBS 125297). The newly generated sequence is highlighted in yellow and the type strains are indicated in bold.

Figure 4. 

Phylogram generated from maximum likelihood analysis based on combined LSU, ITS and act sequence data of Sarocladium. Maximum likelihood bootstrap support ≥ 65% and Bayesian posterior probabilities (BPP) ≥ 0.90 are given near the nodes, respectively. The tree is rooted to Kiflimonium curvulum (CBS 430.66, CBS 384.70A). The newly generated sequence is highlighted in yellow, and the type strains are indicated in bold.

Phylogenetic analyses of Sarocladium

The combined analysis of LSU, ITS and act sequence data of Sarocladium was performed with Kiflimonium curvulum (CBS 430.66, CBS 384.70A) as the outgroup taxa (Fig. 4). The combined dataset contained 2,319 characters (LSU = 894 bp, ITS = 580 bp, act = 845) after alignments, including the gaps. Analysis of the combined dataset yielded a best-scoring tree with an optimal log-likelihood value of -9162.577439 (Fig. 4). The alignment had 22 sequences with 604 distinct patterns. Estimated base frequencies were as follows: A = 0.227911, C = 0.281575, G = 0.275350, T = 0.215164, Rate parameters: AC = 1.495411; AG = 2.342956; AT = 2.214487; CG = 0.793329; CT = 7.315308; GT = 1.000000; Gamma shape alpha = 0.146342. After discarding the first 25% of generations in the Bayesian analyses, 7500 trees were remained, from which the 50% consensus tree and posterior probabilities were determined.

Taxonomy

Tetraploa maritima R. Asghari, Phukhams. & K.D. Hyde, sp. nov.

Index Fungorum: IF903274
Facesoffungi Number: FoF17211

Etymology.

The epithet “maritima” refers to the marine habitat where the holotype was collected.

Holotype.

MFLU 24-0455.

Description.

Saprobic on decaying branches of Areca sp. Sexual morph: Undetermined. Asexual morph: Hyphomycetous (in vivo, Fig. 5). Mycelia on natural substrates superficial, branched, septate, sometimes with swollen cells, black or dark brown. Conidiophores not observed. Conidiogenous cells 4.5−17 × 1−3 µm (x– = 11 × 2 μm, n = 5) monoblastic, brown, thick walled, integrated. Conidia 14−28 × 13−18.5 µm (x– = 21.5 × 15 μm, n = 20), solitary, short cylindrical, thick-walled, verrucose to granular, brown, mostly with a subhyaline to pale brown hilum up to 13 µm (–17), with 3–4 columns 5.5−9 µm wide (x– = 7, n = 20), columns are compact and connected almost all their length, 2–4 septa in each column, constricted at septa. Appendages 19−48 × 2−3 µm (x– = 30 × 2.7 μm, n = 15), 3.5−4.5 µm at the base, setose, unbranched, thick-walled, thin-walled at the apex, up to four septa, finely verrucose, brown, pale brown to subhyaline at the apex, sometimes continuing to grow, forming a subcylindrical narrower paler cell up to 13 µm (n = 10) length.

Figure 5. 

Tetraploa maritima (MFLU 24-0455, holotype) a colonies on the host substrate b close -up of colonies c conidiogenous cell and conidium d–j conidia (arrows show the ongoing growth of appendages) k colony on PDA after 10 days (above) l colony on PDA after 10 days (reverse). Scale bares: 100 µm (b); 10 µm (c–f); 20 µm (g–j).

Culture characteristics.

Conidia germinating on PDA within 12 h. Colonies on PDA reaching 1.5 cm diam. after 10 d at 25 ± 2 °C, velvety to floccose, raised, round, entire margin, grayish yellow, with whitish edge, reverse reddish brown with pale buff edge.

Material examined.

Thailand, Prachuap Khiri Khan Province, Pranburi, on decaying Areca wood submerged in seawater and trapped between rocks, 25 October 2022, K.D. Hyde, R6g (MFLU 24-0455, holotype), ex-type living culture (MFLUCC 24-0565).

GenBank numbers.

Tetraploa maritima MFLUCC 24-0565 (ex-type): ITS = PQ778934, LSU = PQ778930, SSU = PQ778940, tub2 = PQ885482.

Notes.

The reconstruction of phylogenies from LSU, ITS, tub2 and SSU sequence data showed that Tetraploa maritima (MFLUCC 24-0565) formed a distinct clade with T. pseudoaristata (NFCCI 4624, NFCCI 4625, NFCCI 4626) in both maximum likelihood and Bayesian analyses with 95% ML/0.94 BPP statistical support. Both maximum likelihood and Bayesian analyses showed the same topology. The closest match of the ITS sequence of Tetraploa maritima (MFLUCC 24-0565) was 98.92% similar across 100% of the query sequence to T. pseudoaristata (NFCCI 4626). In a BLAST search in GenBank, the closest match of the LSU sequence of T. maritima (MFLUCC 24-0565) was 99.88% similar to T. pseudoaristata (NFCCI 4624, NFCCI 4625, NFCCI 4626). Tetraploa maritima (MFLU 24-0455) has similar morphology to T. pseudoaristata (NFCCI 4624) by having verrucose conidia with four apical appendages but differs in having a distinguishable hilum and shorter and narrower appendages (19−48 × 2−3 µm vs. 23–107.5 × 2.4–5.2 µm), which are sometimes elongate above the apex, unlike T. pseudoaristata (NFCCI 4624). Conidial columns in T. maritima (MFLU 24-0455) are compact and connected almost fully along the length, while in T. pseudoaristata (NFCCI 4624) they are partly split, with the two columns totally separating at the upper part (Hyde et al. 2020). Tetraploa maritima (MFLUCC 24-0565) showed 2.7% (16 out of 595), 0.2% (2 out of 874) and 2.4% (10 out of 423) base pair differences with T. pseudoaristata (NFCCI 4624) in ITS, LSU and tub2 respectively, without gaps. Therefore, T. maritima (MFLU 24-0455) is introduced as a novel species based on morphological and molecular evidence.

Rosellinia maritima R. Asghari, Phukhams. & K.D. Hyde, sp. nov.

Index Fungorum: IF903275
Facesoffungi Number: FoF17212

Etymology.

The epithet “maritima” refers to the marine habitat where the holotype was collected.

Holotype.

MFLU 24-0452.

Description.

Saprobic on decaying branches of Areca sp. Sexual morph: Undetermined. Asexual morph: Hyphomycetous (in vivo, Fig. 6a–e). Colonies on natural substrates superficial, effuse, scattered, black. Mycelia on natural substrates superficial to immersed, branched, septate, subhyaline to brown. Sterile globules 52−74 × 47−69 µm (x– = 62 × 56 μm, n = 20), conidial-like, produced laterally or terminally on the hyphae, solitary, globose to subglobose or irregular, muriform appearance, composed of interwoven, unevenly arranged cells with irregular shapes, thick-walled, dark brown to black, darker in the center. Hyphomycetous (in vitro, Fig. 6f–x). Colonies on PDA. Vegetative hyphae subhyaline, pale brown to brown, branched, septate, simple or sometimes irregularly moniliform. Conidiophores not seen or micronematous. Conidiogenous cells integrated, up to 13 μm in length, subcylindrical, subhyaline to brown. Conidia 8−13 × 5−10 µm (x– = 10.5 × 7.5 μm, n = 15), aleurioconidia, solitary, obovoid, truncate base, hyaline to brown, thick-walled, guttulate, intercalary, laterally or terminally, initiating by hyphal inflammation and separating from hyphae by forming septa. Sterile globules 37−145 × 31−66 µm (x– = 79 × 49 μm, n = 20), conidial-like, produced laterally or terminally on the hyphae, solitary, globose to subglobose, oblong or irregular, muriform appearance, agglomerations of hyphal structures, composed of interwoven, unevenly arranged cells with irregular shapes, thick wall, dark brown to black, darker in the center.

Figure 6. 

Rosellinia maritima (MFLU 24-0452, holotype) a colonies on substrate b close-up of colonies on substrate c–e sterile globules (conidial-like elements) f–x Rosellinia maritima (MFLUCC 24-0562, ex-type) f, g colony on PDA (above and reverse) h, i close-up of colonies in culture j sterile globules in culture k, l mycelia m–o developing aleurioconidia p aleurioconidia q–t developing sterile hyphal elements u–x developing sterile globules. Scale bars: 500 µm (b); 250 (h); 100 (i); 50 (j); 20 µm (c–e, k–l, w,x); 10 µm (m–v).

Culture characteristics.

Conidia germinating on PDA within 12 h. Colonies on PDA reaching 5 cm diam. after one month at 25 ± 2 °C, felty, flat, round, entire margin, dull, gray and white concentric rings, with dirty whitish edge, reverse dark brown with olivaceous gray edge.

Material examined.

Thailand, Prachuap Khiri Khan Province, Pranburi, on decaying Areca wood submerged in seawater and trapped between rocks, 25 October 2022, K.D. Hyde, R6c (MFLU 24-0452, holotype), isotype (MFLU 24-0453), ex-type living culture (MFLUCC 24-0562), ex-isotype living culture (MFLUCC 24-0564).

GenBank numbers.

Rosellinia maritima MFLUCC 24-0562 (ex-type): ITS = PQ778938, LSU = PQ778933, tub2 = PQ885479, rpb2 = PQ885483; Rosellinia maritima MFLUCC 24-0564 (ex-isotype): ITS = PQ778939.

Notes.

The construction of phylogenies from combined ITS, LSU, tub2 and rpb2 sequence data showed that Rosellinia maritima (MFLUCC 24-0562) formed a subclade to a group of Rosellinia species: R. sanctae-cruciana (HAST 90072903), R. nectrioides (CBS 449.89), R. abscondita (CBS 447.89), R. marcucciana (MUCL 51704), R. mammiformis (CBS 445.89) and R. britannica (MFLU 17-0987, HKAS 102349) in both maximum likelihood and Bayesian analyses with 85% ML/0.95 BPP statistical support showing the same topology. In a BLAST search in GenBank, the closest match of the ITS sequence of R. maritima (MFLUCC 24-0562) was 95% similar across 66% of the query sequence to Rosellinia sp. isolate (OTU1178), which was obtained using Illumina sequencing, and its morphology is not available (Gao et al. 2020). In a BLAST search in GenBank, the closest match of the LSU sequence of R. maritima (MFLUCC 24-0562) was 98.48% similar across 94% of the query sequence to R. sanctae-cruciana strain BB. Rosellinia maritima (MFLU 24-0452) is not comparable with Rosellinia sanctae-cruciana as the morphology of its asexual morph is not available. Asexual morphs of Rosellinia have been recorded to be geniculosporium-like, dematophora-like, or nodulisporium-like (Petrini 2013), which is different from R. maritima (MFLU 24-0452) in having distinguished conidiophores, monoblastic geniculate conidiogenous cells, and ellipsoidal conidia. Rosellinia maritima (MFLUCC 24-0562) has morphological similarities with R. truncatispora in culture, producing pale brown, thick-walled, septate hyphal sterile elements. However, it differs by having sterile globules and aleurioconidia in culture (Fournier et al. 2017). Therefore, we introduced our collection of R. maritima (MFLU 24-0452) as a novel species based on its distinct morphology and phylogenetic evidence.

Musicillium theobromae (Turconi) Zare & W. Gams

Index Fungorum: IF510697

Description.

Saprobic on decaying branches of Areca sp. Sexual morph: Undetermined. Asexual morph: Hyphomycetous (in vivo, Fig. 7). Colonies on natural substrates hairy, black, with glistering white conidial masses. Conidiophores 108–503 × 3–7 µm (x– = 252 × 4.5 μm, n = 15), solitary or in groups, macronematous, mononematous, mostly straight or slightly flexuous or bent, wider at the base, unbranched or branched, septate, thick-walled, smooth-walled or slightly asperulate, brown, pale brown to subhyaline or hyaline near the apex, bearing whorls of 3–6(–7) phialides. Conidiogenous cells 13.5–27 × 2–3 µm (x– = 19 × 2.5 μm, n = 20), monophialidic, subhyaline to hyaline, cylindrical, wider at the base and tapering towards the tips, sometimes with an inconspicuous collarette, asperulate. Conidia 3.1−4.8 × 1.4−2.1 µm (x– = 3.9 × 1.8 μm, n = 40), forming in slimy heads, hyaline, cylindrical, obovoid or ellipsoidal, thick-walled, guttulate, verrucose to granular. Hyphomycetous (in vitro, Fig. 8). Colonies on PDA. Vegetative hyphae subhyaline, pale brown to brown, sometimes aggregating into microsclerotium-like structures, moniliform hyphae are present. Conidiophores 178–375 × 2.5–4 µm (x– = 255 × 3 μm, n = 15), arising from vegetative hyphae, mostly straight or slightly flexuous or bent, slightly tapering toward the upper part, unbranched or branched, thick-walled, septate, smooth-walled, sometimes slightly asperulate, brown to pale brown, subhyaline or hyaline at upper part, bearing phialides. Conidiogenous cells 21–35(–43) × 2–4 µm (x– = 29 × 2 μm, n = 20), phialidic, in whorls of 1–6 phialides, smooth-walled, sometimes slightly asperulate, subhyaline to hyaline, cylindrical, wider at the base and tapering towards the tips. Conidia 3−6(−7.5) × 2−4 µm (x– = 4.8 × 2.8 μm, n = 40), forming in slimy heads, hyaline, obovoid or ellipsoidal to cylindrical, thick-walled, guttulate, smooth-walled.

Figure 7. 

Musicillium theobromae (MFLU 24-0454, a new host record) a appearance of colony on the host b, c a close appearance of colony on the host d, e conidiophores f conidiogenous cells and developing conidia g close up of the phialide with an inconspicuous collarette h, i conidia j germinated conidium k colony on PDA after two months. Scale bars: 100 µm (b,e); 200 µm (c); 50 µm (d); 20 µm (f); 5 µm (g–i); 10 µm (j).

Figure 8. 

Musicillium theobromae (MFLUCC 24-0563, a new host record) a colony on PDA after two weeks (above) b colony on PDA after two weeks (below) c, d appearance of conidiophores and conidia on the culture e conidiophore and conidia f conidiogenous cells bearing conidia g conidia h moniliform hyphae. Scale bars: 200 µm (c); 100 µm (d, e); 50 µm (f,h); 10 µm (g).

Culture characteristics.

Conidia germinating on PDA within 12 h. Colonies on PDA reaching 2 cm diam. after two weeks and 4 cm diam. after two months at 25 ± 2 °C, finely floccose, round, entire margin to finely rhizoid, white and gray brown at the center, becoming raised and brown with age. Reverse white with gray brown at the center, becoming darker with age.

Material examined.

Thailand, Prachuap Khiri Khan Province, Pranburi, on decaying Areca wood submerged in seawater and trapped between rocks, 25 October 2022, K.D. Hyde, R6a (MFLU 24-0454), living culture (MFLUCC 24-0563).

GenBank numbers.

Musicillium theobromae MFLUCC 24-0563: ITS = PQ778936, LSU = PQ778931, rpb2 = PQ885480, tef1-α = PQ885481.

Notes.

The reconstruction of phylogenies from LSU-ITS-tef1-α-rpb2 sequences showed that Musicillium theobromae (MFLUCC 24-0563) formed a sister clade with M. theobromae (CBS 968.72) in both maximum likelihood and Bayesian analyses with 100% ML/1.00 BPP statistical support (Fig. 3). In a BLAST search in GenBank, the closest match of the ITS sequence of M. theobromae (MFLUCC 24-0563) was 99.33% similar across 99% of the query sequence to M. theobromae (NZ62). In a BLAST search in GenBank, the closest match of the LSU sequence of M. theobromae (MFLUCC 24-0563) was 99.89% similar across 98% of the query sequence to M. theobromae (CBS 243.74). Musicillium theobromae (MFLUCC 24-0563) has similar morphology to the neotype species of M. theobromae (CBS 968.72), in having slightly asperulate brown septate conidiophores bearing whorls of up to six phialides, almost the same length (21–35(–43) µm vs. 17−35 µm) of phialide, which are tapering towards the tips, hyaline smooth-walled conidia forming in a slimy head (Zare et al. 2007). Therefore, M. theobromae (MFLUCC 24-0563) is introduced as a new host record on Areca based on morphology and phylogenetic evidence.

Sarocladium gamsii A. Giraldo, Gené & Guarro

Index Fungorum: IF807944

Description.

Saprobic on decaying branches of Areca sp. Sexual morph: Undetermined. Asexual morph: Hyphomycetous (in vivo, Fig. 9). Colonies on natural substrates hairy, effuse, scattered, grayish, shiny, with black conidiophores. Synemata, tree-like, 240–280 µm height (x– = 328 μm, n = 10), wider at the base 25–62 µm wide (x– = 44 μm, n = 10), narrower in the middle 17–30 µm wide (x– = 24 μm, n = 10), divergent in the upper part, composed of 8–20 conidiophores at the base of each stipe. Conidiophores 193–298 × 1.5–3 µm (x– = 259 × 2.5 μm, n = 15), solitary or in groups, macronematous, synnematous, twisted in the stipe, parallel in the upper part, irregularly branched, straight or slightly flexuous, septate, smooth, brown, subhyaline to hyaline at the upper part. Conidiogenous cells 10–39 × 1.5–2.5 µm (x– = 21 × 2 μm, n = 15), polyphialidic, integrated or terminal, cylindrical, straight to slightly curved or sympodial at the upper part, acropetally proliferating, subhyaline to hyaline, finely aculeate, denticle at the attachment site of conidia, without collarette. Conidia 2−6.5 × 1.5−2.5 µm (x– = 4 × 2 μm, n = 30), unicellular, fusiform, truncate at base, rounded apex, hyaline, thick-walled, smooth-walled, guttulate, bud scars or disjunctors present at the site of attachment. Hyphomycetous (in vitro, Fig. 10). Colonies on PDA. Vegetative hyphae hyaline, smooth-walled, thin-walled, septate. Conidiophores 8.5–47(–76) µm (x– = 29 μm, n = 15) height, arising from vegetative hyphae or ropes of hyphae, straight, flexuous or slightly bent, slightly tapering toward the apex, unbranched to rarely branched, smooth-walled to asperulate, hyaline, aseptate or uniseptate at the base. Conidiogenous cells 5.5–45 × 1–1.75 µm (x– = 27 × 1.3 μm, n = 15), monophialidic, acicular, with apical periclinal thickening, finely asperulate, hyaline. Conidia 3−5 × 1−2 µm (x– = 4.5 × 1.5 μm, n = 30), solitary or forming in slimy heads or chains, hyaline, unicellular, fusiform, thick-walled, guttulate, finely asperulate. Chlamydospores not observed.

Figure 9. 

Sarocladium gamsii (MFLU 24-0451, a new host record) a colonies on substrate b close-up of colonies on substrate c synnemata d base of synnemata e apex of synemata f, g conidiogenous cells and conidia h conidiogenous cell and conidia, indicating denticles at the attachment site of conidia i conidia j colony on PDA after one month (above and below). Scale bars: 100 µm (b, c); 20 µm (d, e); 10 µm (f, g); 5 µm (h, i).

Figure 10. 

Sarocladium gamsii (MFLUCC 24-0561, a new host record) a, b colonies on PDA after two months c–e conidiophores and conidia f conidia arranged in chains g conidia arranged in slimy heads h, i conidia. Scale bars: 20 µm (c–e); 5 µm (h, i).

Culture characteristics.

Conidia germinating on PDA within 12 h. Colonies on PDA reaching 2.5 cm diam. after one month and 3 cm diam. after two months at 25 ± 2 °C, umbonate, radially folded, lobate to irregular, yellowish white. Reverse buff.

Material examined.

Thailand, Prachuap Khiri Khan Province, Pranburi, on decaying Areca wood submerged in seawater and trapped between rocks, 25 October 2022, K.D. Hyde, R6d (MFLU 24-0451), living culture (MFLUCC 24-0561).

GenBank numbers.

Sarocladium gamsii MFLUCC 24-0561: ITS = PQ778937, LSU = PQ778932.

Notes.

The reconstruction of phylogenies from combined LSU, ITS and act sequences showed that Sarocladium gamsii (MFLUCC 24-0561) formed a sister clade with S. gamsii (CBS 707.73 in both maximum likelihood and Bayesian analyses with 100% ML/1 BPP statistical support (Fig. 4). In a BLAST search in GenBank, the closest match of the ITS sequence of S. gamsii (MFLUCC 24-0561) was 99% similar across 95% of the query sequence to S. gamsii (PUPYF263_ANAMORPH). In a BLAST search in GenBank, the closest match of the LSU sequence of Sarocladium gamsii (MFLUCC 24-0561) was 99% similar across 100% of the query sequence to Monocillium sp. (CBS 187.80). Sarocladium gamsii (MFLUCC 24-0561) has similar morphology to S. gamsii (CBS 707.73) in having straight or slightly bent, unbranched or rarely branched conidiophores arising from vegetative hyphae or ropes of hyphae, monophialidic conidiogenous cells with apical periclinal thickening, fusiform, hyaline, aseptate conidia forming in chains or slimy heads, but it differs in having finely asperulate conidiogenous cells and conidia (Giraldo et al. 2015). Sarocladium gamsii (MFLUCC 24-0561) is introduced as a new host record based on morphology and phylogenetic evidence.

Discussion

Marine fungi are a taxonomically diverse group, playing crucial ecological roles as saprobes, endophytes, symbionts, and pathogens (Hyde and Jones 1988; Hyde and Lee 1998; Jones et al. 2019; Calabon et al. 2023). Saprobic lignicolous fungi are the most extensively studied group of marine fungi (Jones et al. 2019). Ascomycota is the dominantly documented marine fungal group, including the major classes Dothideomycetes and Sordariomycetes, which are suggested to have evolved from terrestrial ancestors (Vijaykrishna et al. 2006). In this study, we continue our previous investigations into fungal presence on decaying Areca branches submerged in seawater. Species identification was performed using both morphological and molecular data, as both methods are essential in fungal taxonomy (Chethana et al. 2021; Maharachchikumbura et al. 2021). This research has led to novel discoveries of dothideomycetous and sordariomycetous fungi in this habitat, which are discussed herein.

Tetraploa (Tetraplosphaeriaceae, Pleosporales, Dothideomycetes) was introduced by Berkeley and Broome (1850). Initially, it was known by its 3–4 columnar hyphomycetous conidia. Later, it was linked to a massarina-like sexual morph (Tanaka et al. 2009; Hyde et al. 2013). The genus comprises 38 species isolated from various plant materials (Species Fungorum, accessed in April 2025). There are records of Tetraploa from palm hosts, such as Tetraploa palmae (Zhang et al. 2024), but without any records on Areca (Xiong et al. 2024). Tetraploa is mostly found in terrestrial and freshwater habitats, with a few records from marine environments, such as Tetraploa aristata on salt marsh hosts (Calabon et al. 2021; Zhang et al. 2024). We introduce our collection of Tetraploa as a novel species, T. maritima, found on Areca in a marine environment in Thailand. Phylogenetic analyses of combined LSU, ITS, tub2 and SSU sequence data showed separation of our collection (MFLUCC 24-0565) from other Tetraploa species, forming a subclade to T. pseudoaristata (NFCCI 4624, NFCCI 4625, NFCCI 4626). The topology of our phylogenetic tree is generally consistent with previous studies (Zhang et al. 2024; Zhao et al. 2024), although there are some differences that may be attributed to the use of different sequence combinations and taxon sampling. The molecular result is further supported by morphological differences.

Rosellinia (Xylariaceae, Xylariales, Sordariomycetes) was described by De Notaris (1844) to accommodate its sexual type species, R. aquila. The genus comprises around 150 species (Li et al. 2024), with some reported from palms, such as R. palmae, R. rhopalostylidicola, and R. calami, though there are no records for Areca species (Xiong et al. 2024). Rosellinia has been reported from various habitats, including marine environments (Gessner and Goos 1973). The asexual morph of Rosellinia is known to be geniculosporium-, dematophora-, or nodulisporium-like with hyphomycetous synnemata or conidiophores forming around stromata on their natural substrates. These synnemata or conidiophores may be sterile or fertile, with the fertile parts developing geniculate holoblastic conidiogenous cells that produce ellipsoidal conidia with truncate bases (Tang et al. 2009; Petrini 2013; Fournier et al. 2017). Based on phylogenetic analyses (Fig. 2), our collection of Rosellinia maritima (MFLUCC 24-0562) forms a distinct clade related to R. sanctae-cruciana (HAST 90072903), but its morphology differs from other known asexual morphs within the genus. This distinction is seen in its production of brown to black sterile globules with a conidial-like structure composed of interwoven irregular hyphal-like cells, that mostly attach directly to the mycelium, without distinct conidiophores. The mycelium of R. maritima in cultures is similar to R. truncatispora in having brown, thick-walled, hyphal sterile elements; however, it differs by forming dark-pigmented globose structures and aleurioconidia (Fournier et al. 2017). Due to the lack of molecular data, R. truncatispora cannot be compared phylogenetically. Producing intertwined or moniliform, dark brown, thick-walled hyphal sterile elements in culture is also observed in Dematophora acutispora and D. asperata, indicating the presence of such structures within the Xylariaceae (Fournier et al. 2017). However, since the Rosellinia (MFLU 24-0452) collection in this study was distinctly different from other asexual morphs in this genus, pictures of ex-type R. maritima (MFLUCC 24-0562) are provided showing the conidial-like structure, resembling those in the natural environment (Fig. 6h–x). Additionally, to minimize any possibility of contamination during the molecular process, the specimen was carefully re-examined and single spore isolation was repeated. An isotype (MFLU 24-0453) and ex-isotype (MFLUCC 24-0564) were provided. The results confirmed the morphology and sequences of the holotype (MFLU 24-0452) and ex-type (MFLUCC 24-0562).

Musicillium (Plectosphaerellaceae, Glomerellales, Sordariomycetes) was introduced by Zare et al. (2007) to accommodate a verticillium-like species, with M. theobromae designated as the type species. There are three other species included in this genus: M. elettariae, M. pandanicola and M. tropicale (Species Fungorum, accessed April 2025). Musicillium theobromae is a widespread species known as a pathogenic agent of cigar-end rot, frequently reported on banana (Zare et al. 2007; Youssef et al. 2020). Based on morphological and molecular evidence, our collection of M. theobromae (MFLUCC 24-0563) resembles M. theobromae (CBS 968.72). Although there have been other reports of this species on members of the palm family (Capdet and Romero 2010), in this study, we introduce our collection of M. theobromae (MFLUCC 24-0563), as a new host record on Areca (Xiong et al. 2024). Additionally, this is the first collection of Musicillium in a marine environment (www.marine.org; Jones et al. 2019; Calabon et al. 2023). We also provide an illustration and description of M. theobromae (MFLU 24-0454) from its natural substrate, which is similar to its morphology in culture, with some differences in the measurements of conidiophores, conidiogenous cells and conidia.

Sarocladium (Sarocladiaceae, Hypocreales, Sordariomycetes) is a hyphomycetous genus introduced by Gams and Hawksworth (1976). The genus includes 35 species reported as pathogens, endophytes, and saprobes from diverse environments (Species Fungorum, accessed April 2025; Gams and Hawksworth 1976; Yeh and Kirschner 2014; Phukhamsakda et al. 2020). A few species, such as S. strictum and S. kiliense, have been found in marine habitats (Jones et al. 2019). Sarocladium gamsii was introduced by Giraldo et al. (2015) for saprobic isolates characterized by conidial arrangements in chains and slimy heads. Our collection of Sarocladium (MFLUCC 24-0561) clustered with S. gamsii (CBS 707.73) with strong molecular support, and its morphological similarity—forming conidia in chains and slimy heads—further confirms its identification as S. gamsii. Sarocladium gamsii is presented here as a new host record on Areca, submerged in seawater. We also provide an illustration and description of S. gamsii (MFLU 24-0451), from its natural substrate, which differs from its morphology in culture. The synematous conidiomata observed match the synnematous morphology of S. clematidis (MFLU 17–1507), supporting the identification of this genus in its natural habitat (Phukhamsakda et al. 2020).

The four genera examined in this study—Tetraploa, Rosellinia, Musicillium, and Sarocladium—along with the previously examined genus Myrmecridium (Asghari et al. 2023), represent fungal groups with diverse ecological roles, ranging from saprobes to pathogens. Although these genera are mostly found in terrestrial or freshwater habitats, their presence on decaying Areca material submerged in seawater shows their adaptability to marine environments, which have high salinity. The discovery of three novel species and two new host records on Areca further demonstrates the diversity of marine fungi and suggests their ability to colonize palm substrates. Further research of this habitat may reveal more novel species and extend our knowledge of fungal diversity and their ecological functions.

Acknowledgments

Raheleh Asghari would like to thank the Mae Fah Luang University Partial Scholarship for the doctoral degree program (GR-ST-PS-65-21), Mushroom Research Foundation, and the National Research Council of Thailand (NRCT) grant “Total fungal diversity in a given forest area with implications towards species numbers, chemical diversity, and biotechnology” (grant no. N42A650547) for support. Permission for surveys, studies, research or academic experiments (No. 0907.4/23579) in the National Parks (book no. 02/020) and the Forest Parks, Botanic Gardens and Arboreta (book no. 02/001) is acknowledged. Kevin D. Hyde would like to thank the Flexible Talent Introduction Program (E16441) and the Chinese Research Fund, grant number E1644111K1, titled “Flexible introduction of the high-level expert program, Kunming Institute of Botany, Chinese Academy of Sciences” for support. Gareth Jones and Kevin D. Hyde are supported under the Distinguished Scientist Fellowship Program (DSFP), King Saud University, Kingdom of Saudi Arabia. Chayanard Phukhamsakda would like to thank the Basic Research Fund (Fundamental Fund) supported by the National Science, Research, and Innovation Fund under Grant No. 682A16032. Carlo Chris S. Apurillo would like to thank the Mae Fah Luang University for the partial scholarship grant (GR-ST-PS-65-21), Mushroom Research Foundation for financial support and the Department of Science and Technology-Philippine Science High School-EVC for the grant of study leave. Dr. Shaun Pennycook is thanked for advice regarding the new epithet. We would like to thank Dr. Martin van de Bult for the identification of the host.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work is grateful to the Basic Research Fund (Fundamental Fund) supported by the National Science, Research and Innovation Fund under Grant No. 682A16032 for full financial support.

Author contributions

Supervision and review: ChP, EBGJ, PK, KDH, AB. Writing – original draft: RA, OK. Molecular analyses: RA, OK, CCSA.

Author ORCIDs

Raheleh Asghari https://orcid.org/0009-0006-4897-5327

Chayanard Phukhamsakda https://orcid.org/0000-0002-1033-937X

E. B. Gareth Jones https://orcid.org/0000-0002-7286-5471

Ali Bahkali https://orcid.org/0000-0003-3936-0436

Carlo Chris S. Apurillo https://orcid.org/0000-0003-4348-0887

Omid Karimi https://orcid.org/0000-0001-9652-2222

Pattana Kakumyan https://orcid.org/0000-0001-8281-1082

Kevin D Hyde https://orcid.org/0000-0002-2191-0762

Data availability

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

References

  • Baker WJ, Dransfield J (2016) Beyond Genera Palmarum: Progress and prospects in palm systematics. Botanical Journal of the Linnean Society 182: 207–233. https://doi.org/10.1111/boj.12401
  • Balanagouda P, Sridhara S, Shil S, Hegde V, Naik MK, Narayanaswamy H, Balasundram SK (2021) Assessment of the spatial distribution and risk associated with fruit rot disease in Areca catechu L. Journal of Fungi 7(10): 797. https://doi.org/10.3390/jof7100797
  • Bhunjun CS, Niskanen T, Suwannarach N, Wannathes N, Chen YJ, McKenzie EHC, Maharachchikumbura SS, Buyck B, Zhao CL, Fan YG, Zhang JY, Dissanayake AJ, Marasinghe DS, Jayawardena RS, Kumla J, Padamsee M, Chen YY, Liimatainen K, Ammirati JF, Phukhamsakda C, Liu JK, Phonrob W, Randrianjohany E, Hongsanan S, Cheewangkoon R, Bundhun D, Khuna S, Yu WJ, Deng LS, Lu YZ, Hyde KD, Lumyong S (2022) The numbers of fungi: are the most speciose genera truly diverse? Fungal Diversity 27: 1–76. https://doi.org/10.1007/s13225-022-00501-4
  • Bonugli-Santos RC, dos Santos Vasconcelos MR, Passarini MR, Vieira GA, Lopes VC, Mainardi PH, Dos Santos JA, de Azevedo Duarte L, Otero IV, da Silva Yoshida M, Feitosa VA, Pessoa Jr A, Sette LD (2015) Marine-derived fungi: diversity of enzymes and biotechnological applications. Frontiers in Microbiology 6: 128060. https://doi.org/10.3389/fmicb.2015.00269
  • Calabon MS, Jones EBG, Pang KL, Abdel-Wahab MA, Jin J, Devadatha B, Sadaba RB, Apurillo CC, Hyde KD (2023) Updates on the classification and numbers of marine fungi. Botanica Marina 66: 213–238. https://doi.org/10.1515/bot-2023-0032
  • Chen YJ, Jayawardena RS, Bhunjun CS, Harishchandra DL, Hyde KD (2020) Pseudocer­cospora dypsidis sp. nov. (Mycosphaerellaceae) on Dypsis lutescens leaves in Thailand. Phytotaxa 474(3): 218–234. https://doi.org/10.11646/phytotaxa.474.3.2
  • Chethana KWT, Manawasinghe IS, Hurdeal VG, Bhunjun CS, Appadoo MA, Gentekaki E, Raspé O, Promputtha I, Hyde KD (2021) What are fungal species and how to delineate them? Fungal Diversity 109(1): 1–25. https://doi.org/10.1007/s13225-021-00483-9
  • Dayarathne MC, Jones EBG, Maharachchikumbura SSN, Devadatha B, Sarma VV, Khongphinitbunjong K, Chomnunti P, Hyde KD (2020) Morpho-molecular characterization of microfungi associated with marine-based habitats. Mycosphere 11: 1–188. https://doi.org/10.5943/mycosphere/11/1/1
  • De Notaris G (1844) Cenni sulla Tribù de’ Pirenomiceti Sferiacei e descrizione di alcuni nuovi generi. Giornale Botanico Italiano 1.1(2): 322–335.
  • Gao C, Montoya L, Xu L, Madera M, Hollingsworth J, Purdom E, Singan V, Vogel J, Hutmacher RB, Dahlberg JA, Coleman-Derr D (2020) Fungal community assembly in drought-stressed sorghum shows stochasticity, selection, and universal ecological dynamics. Nature Communications 11(1): 34. https://doi.org/10.1038/s41467-019-13913-9
  • Giraldo A, Gené J, Sutton DA, Madrid H, de Hoog GS, Cano J, Decock C, Crous PW, Guarro J. (2015) Phylogeny of Sarocladium (Hypocreales). Persoonia 34(1): 10–24. https://doi.org/10.3767/003158515X685364
  • Glass NL, Donaldson GC (1995) Development of primer sets designed for use with the PCR to amplify conserved genes from filamentous ascomycetes. Applied and Environmental Microbiology 61(4): 1323–1330. https://doi.org/10.1128/aem.61.4.1323-1330.1995
  • Heatubun CD, Dransfield J, Flynn T, Tjitrosoedirdjo SS, Mogea JP, Baker WJ (2012) A monograph of the betel nut palms (Areca: Arecaceae) of East Malesia. Botanical Journal of the Linnean Society 168(2): 147–173. https://doi.org/10.1111/j.1095-8339.2011.01199.x
  • Hou LW, Giraldo A, Groenewald JZ, Rämä T, Summerbell RC, Huang GZ, Cai L, Crous PW (2023) Redisposition of acremonium-like fungi in Hypocreales. Studies in Mycology 105: 23. https://doi.org/10.3114/sim.2023.105.02
  • Hyde KD, Lee SY (1998) Ecology of mangrove fungi and their role in nutrient cycling: What gaps occur in our knowledge? Hydrobiologia 295: 107–118. https://doi.org/10.1007/BF00029117
  • Hyde KD, Jones EBG, Leaño E, Pointing SB, Poonyth AD, Vrijmoed LL (1998) Role of fungi in marine ecosystems. Biodiversity and Conservation 7: 1147–1161. https://doi.org/10.1023/A:1008823515157
  • Hyde KD, Sarma VV, Jones EBG (2000) Morphology and taxonomy of higher marine fungi. In: Hyde KD, Pointing SB (Eds) Marine Mycology: A Practical Approach. Fungal Diversity Research Series 1: 172–204.
  • Hyde KD, Jones EBG, Liu JK, Ariyawansa H, Boehm E, Boonmee S, Braun U, Chomnunti P, Crous PW, Dai DQ, Diederich P, Dissanayake A, Doilom M, Doveri F, Hongsanan S, Jayawardena R, Lawrey JD, Li YM, Liu YX, Lücking R, Monka J, Muggia L, Nelsen MP, Pang KL, Phookamsak R, Senanayake IC, Shearer CA, Suetrong S, Tanaka K, Thambugala KM, Wijayawardene NN, Wikee S, Wu HX, Zhang Y, Begoña AH, Alias SA, Aptroot A, Bahkali AH, Bezerra JL, Bhat DJ, Camporesi E, Chukea E, Gueidan C, Hawksworth DL, Hirayama K, Hoog SD, Kang JK, Knudsen K, Li WJ, Li XH, Liu ZY, Mapook A, McKenzie EHC, Miller AN, Mortimer PE, Phillips AJL, Raja HA, Scheuer C, Schumm F, Taylor JE, Tian Q, Tibpromma S, Wanasinghe DN, Wang Y, Xu JC, Yacharoen S, Yan JY, Zang M (2013) Families of Dothideomycetes. Fungal Diversity 63: 1–313. https://doi.org/10.1007/s13225-013-0263-4
  • Hyde KD, Xu JC, Rapior S, Jeewon R, Lumyong S, Niego AGT, Abeywickrama PD, Aluthmuhandiram JVS, Brahamanage RS, Brooks S, Chaiyasen A, Chethana KWT, Chomnunti P, Chepkirui C, Chuankid B, de Silva NI, Doilom M, Faulds C, Gentekaki E, Gopalan V, Kakumyan P, Harishchandra D, Hemachandran H, Hongsanan S, Karunarathna A, Karunarathna SC, Khan S, Kumla J, Jayawardena RS, Liu JK, Liu N, Luangharn T, Macabeo APG, Marasinghe DS, Meeks D, Mortimer PE, Mueller P, Nadir S, Nataraja KN, Nontachaiyapoom S, O’Brien M, Penkhrue W, Phukhamsakda C, Ramanan US, Rathnayaka AR, Sadaba RB, Sandargo B, Samarakoon BC, Tennakoon DS, Siva R, Sriprom W, Suryanarayanan TS, Sujarit K, Suwannarach N, Suwunwong T, Thongbai B, Thongklang N, Wei DP, Wijesinghe SN, Winiski J, Yan J, Yasanthika E, Stadler M (2019) The amazing potential of fungi: 50 ways we can exploit fungi industrially. Fungal Diversity 97: 1–136. https://doi.org/10.1007/s13225-019-00430-9
  • Hyde KD, Jeewon R, Chen YJ, Bhunjun CS, Calabon MS, Jiang HB, Lin CG, Norphanphoun C, Sysouphanthong P, Pem D, Tibpromma S, Zhang Q, Doilom M, Jayawardena RS, Liu JK, Maharachchikumbura SS, Phukhamsakda C, Phookamsak R, Al-Sadi AM, Thongklang N, Wang Y, Gafforov Y, Jones EBG, Lumyong S (2020) The numbers of fungi: is the descriptive curve flattening? Fungal Diversity 103: 219–271. https://doi.org/10.1007/s13225-020-00458-2
  • Hyde KD, Noorabadi MT, Thiyagaraja V, He MQ, Johnston PR, Wijesinghe SN, Armand A, Biketova AY, Chethana KWT, Erdoğdu M, Ge ZW, Groenewald JZ, Hongsanan S, Kušan I, Leontyev DV, Li DW, Lin CG, Liu NG, Maharachchikumbura SSN, Matočec N, May TW, McKenzie EHC, Mešić A, Perera RH, Phukhamsakda C, Piątek M, Samarakoon MC, Selcuk F, Senanayake IC, Tanney JB, Tian Q, Vizzini A, Wanasinghe DN, Wannasawang N, Wijayawardene NN, Zhao RL, Abdel-Wahab MA, Abdollahzadeh J, Abeywickrama PD, Abhinav , Absalan S, Acharya K, Afshari N, Afshan NS, Afzalinia S, Ahmadpour SA, Akulov O, Alizadeh A, Alizadeh M, Al-Sadi AM, Alves A, Alves VCS, Alves-Silva G, Antonín V, Aouali S, Aptroot A, Apurillo CCS, Arias RM, Asgari B, Asghari R, Assis DMA, Assyov B, Atienza V, Aumentado HDR, Avasthi S, Azevedo E, Bakhshi M, Bao DF, Baral HO, Barata M, Barbosa KD, Barbosa RN, Barbosa FR, Baroncelli R, Barreto GG, Baschien C, Bennett RM, Bera I, Bezerra JDP, Bhunjun CS, Bianchinotti MV, Błaszkowski J, Boekhout T, Bonito GM, Boonmee S, Boonyuen N, Bortnikov FM, Bregant C, Bundhun D, Burgaud G, Buyck B, Caeiro MF, Cabarroi-Hernández M, Cai M Feng, Cai L, Calabon MS, Calaça FJS, Callalli M, Câmara MPS, Cano-Lira J, Cao B, Carlavilla JR, Carvalho A, Carvalho TG, Castañeda-Ruiz RF, Catania MDV, Cazabonne J, Cedeño-Sanchez M, Chaharmiri-Dokhaharani S, Chaiwan N, Chakraborty N, Cheewankoon R, Chen C, Chen J, Chen Q, Chen YP, Chinaglia S, Coelho-Nascimento CC, Coleine C, CostaRezende DH, Cortés-Pérez A, Crouch JA, Crous PW, Cruz RHSF, Czachura P, Damm U, Darmostuk V, Daroodi Z, Das K, Das K, Davoodian N, Davydov EA, da Silva GA, da Silva IR, da Silva RMF, da Silva Santos AC, Dai DQ, Dai YC, de Groot Michiel D, De Kesel A, De Lange R, de Medeiros EV, de Souza CFA, de Souza FA, dela Cruz TEE, Decock C, Delgado G, Denchev CM, Denchev TT, Deng YL, Dentinger BTM, Devadatha B, Dianese JC, Dima B, Doilom M, Dissanayake AJ, Dissanayake DMLS, Dissanayake LS, Diniz AG, Dolatabadi S, Dong JH, Dong W, Dong ZY, Drechsler-Santos ER, Druzhinina IS, Du TY, Dubey MK, Dutta AK, Elliott TF, Elshahed MS, Egidi E, Eisvand P, Fan L, Fan X, Fan XL, Fedosova AG, Ferro LO, Fiuza PO, Flakus A, W. Fonseca EO, Fryar SC, Gabaldón T, Gajanayake AJ, Gannibal PB, Gao F, GarcíaSánchez D, García-Sandoval R, Garrido-Benavent I, Garzoli L, Gasca-Pineda J, Gautam AK, Gené J, Ghobad-Nejhad M, Ghosh A, Giachini AJ, Gibertoni TB, Gentekaki E, Gmoshinskiy VI, GóesNeto A, Gomdola D, Gorjón SP, Goto BT, Granados-Montero MM, Griffith GW, Groenewald M, Grossart H-P, Gu ZR, Gueidan C, Gunarathne A, Gunaseelan S, Guo SL, Gusmão LFP, Gutierrez AC, Guzmán-Dávalos L, Haelewaters D, Haituk H, Halling RE, He SC, Heredia G, HernándezRestrepo M, Hosoya T, Hoog SD, Horak E, Hou CL, Houbraken J, Htet ZH, Huang SK, Huang WJ, Hurdeal VG, Hustad VP, Inácio CA, Janik P, Jayalal RGU, Jayasiri SC, Jayawardena RS, Jeewon R, Jerônimo GH, Jin J, Jones EBG, Joshi Y, Jurjević Ž, Justo A, Kakishima M, Kaliyaperumal M, Kang GP, Kang JC, Karimi O, Karunarathna SC, Karpov SA, Kezo K, Khalid AN, Khan MK, Khuna S, Khyaju S, Kirchmair M, Klawonn I, Kraisitudomsook N, Kukwa M, Kularathnage ND, Kumar S, Lachance MA, Lado C, Latha KPD, Lee HB, Leonardi M, Lestari AS, Li C, Li H. Li J, Li Q, Li Y, Li YC, Li YX, Liao CF, Lima JLR, Lima JMS, Lima NB, Lin L, Linaldeddu BT, Linn MM, Liu F, Liu JK, Liu JW, Liu S, Liu SL, Liu XF, Liu XY, Longcore JE, Luangharn T, Luangsa-ard JJ, Lu L, Lu YZ, Lumbsch HT, Luo L, Luo M, Luo ZL, Ma J, Madagammana AD, Madhushan A, Madrid H, Magurno F, Magyar D, Mahadevakumar S, Malosso E, Malysh JM, Mamarabadi M, Manawasinghe IS, Manfrino RG, Manimohan P, Mao N, Mapook A, Marchese P, Marasinghe DS, Mardones M, Marin-Felix Y, Masigol H, Mehrabi M, MehrabiKoushki M, Meiras-Ottoni A de, Melo RFR, Mendes-Alvarenga RL, Mendieta S, Meng QF, Menkis A, Menolli Jr N, Mikšík M, Miller SL, Moncada B, Moncalvo JM, Monteiro JS, Monteiro M, Mora-Montes HM, Moroz EL, Moura JC, Muhammad U, Mukhopadhyay S, Nagy GL, Najam ul Sehar A, Najafiniya M, Nanayakkara CM, Naseer A, Nascimento ECR, Nascimento SS, Neuhauser S, Neves MA, Niazi AR, Nie Yong, Nilsson RH, Nogueira PTS, Novozhilov YK, Noordeloos M, Norphanphoun C, Nuñez Otaño N, O’Donnell RP, Oehl F, Oliveira JA, Oliveira Junior I, Oliveira NVL, Oliveira PHF, Orihara T, Oset M, Pang KL, Papp V, Pathirana LS, Peintner U, Pem D, Pereira OL, Pérez-Moreno J, Pérez-Ortega S, Péter G, Pires-Zottarelli CLA, Phonemany M, Phongeun S, Pošta A, Prazeres JFSA, Quan Y, Quandt CA, Queiroz MB, Radek R, Rahnama K, Raj KNA, Rajeshkumar KC, Rajwar Soumyadeep , Ralaiveloarisoa AB, Rämä T, Ramírez-Cruz V, Rambold G, Rathnayaka AR, Raza M, Ren GC, Rinaldi AC, Rivas-Ferreiro M, Robledo GL, Ronikier A, Rossi W, Rusevska K, Ryberg M, Safi A, Salimi F, Salvador-Montoya CA, Samant B, Samaradiwakara NP, Sánchez-Castro I, Sandoval-Denis M, Santiago ALCMA, Santos ACDS, Santos LA dos, Sarma VV, Sarwar S, Savchenko A, Savchenko K, Saxena RK, Schoutteten N, Selbmann L, Ševčíková H, Sharma A, Shen HW, Shen YM, Shu YX, Silva HF, Silva-Filho AGS, Silva VSH, Simmons DR, Singh R, Sir EB, Sohrabi M, Souza FA, Souza-Motta CM, Sriindrasutdhi V, Sruthi OP, Stadler M, Stemler J, Stephenson SL, Stoyneva-Gaertner MP, Strassert JFH, Stryjak-Bogacka M, Su H, Sun YR, Svantesson S, Sysouphanthong P, Takamatsu S, Tan TH, Tanaka K, Tang C, Tang X, Taylor JE, Taylor PWJ, Tennakoon DS, Thakshila SAD, Thambugala KM, Thamodini GK, Thilanga D, Thines M, Tiago PV, Tian XG, Tian WH, Tibpromma S, Tkalčec Z, Tokarev YS, Tomšovský M, Torruella G, Tsurykau A, Udayanga D, Ulukapı M, Untereiner WA, Usman M, Uzunov BA, Vadthanarat S, Valenzuela R, Van den Wyngaert S, Van Vooren N, Velez P, Verma RK, Vieira LC, Vieira WAS, Vinzelj JM, Tang AMC, Walker A, Walker AK, Wang QM, Wang Y, Wang XY, Wang ZY, Wannathes N, Wartchow F, Weerakoon G, Wei DP, Wei X, White JF, Wijesundara DSA, Wisitrassameewong K, Worobiec G, Wu HX, Wu N, Xiong YR, Xu B, Xu JP, Xu R, Xu RF, Xu RJ, Yadav S, Yakovchenko LS, Yang HD, Yang X, Yang YH, Yang Y, Yang YY, Yoshioka R, Youssef Noha H, Yu FM, Yu ZF, Yuan LL, Yuan Q, Zabin DA, Zamora JC, Zapata CV, Zare R, Zeng M, Zeng XY, Zhang JF, Zhang JY, Zhang S, Zhang XC, Zhao CL, Zhao H, Zhao Q, Zhao H, Zhao HJ, Zhou HM, Zhu XY, Zmitrovich IV, Zucconi L, Zvyagina E (2024) The 2024 Outline of Fungi and fungus-like taxa. Mycosphere 15: 5146–6239. https://doi.org/10.5943/mycosphere/15/1/25
  • Jayasiri SC, Hyde KD, Ariyawansa HA, Bhat J, Buyck B, Cai L, Dai YC, Abd-Elsalam KA, Ertz D, Hidayat I, Jeewon R, Jones EBG, Bahkali AH, Karunarathna SC, Liu JK, Luangsa-ard JJ, Lumbsch HT, Maharachchikumbura SSN, McKenzie EHC, Moncalvo JM, Ghobad-Nejhad M, Nilsson H, Pang KL, Pereira OL, Phillips AJL, Raspé O, Rollins AW, Romero AI, Etayo J, Selçuk F, Stephenson SL, Suetrong S, Taylor JE, Tsui CKM, Vizzini A, Abdel-Wahab MA, Wen TC, Boonmee S, Dai DQ, Daranagama DA, Dissanayake AJ, Ekanayaka AH, Fryar SC, Hongsanan S, Jayawardena RS, Li WJ, Perera RH, Phookamsak R, de Silva NI, Thambugala KM, Tian Q, Wijayawardene NN, Zhao RL, Zhao Q, Kang JC, Promputtha I (2015) The Faces of Fungi database: fungal names linked with morphology, phylogeny, and human impacts. Fungal Diversity 74: 3–18. https://doi.org/10.1007/s13225-015-0351-8
  • Jones EBG, Sakayaroj J, Suetrong S, Somrithipol S, Pang KL (2009) Classification of marine Ascomycota, anamorphic taxa and Basidiomycota. Fungal Diversity 35: 1–187.
  • Jones EBG, Suetrong S, Sakayaroj J, Bahkali AH, Abdel-Wahab MA, Boekhout T, Pang KL (2015) Classification of marine Ascomycota, Basidiomycota, Blastocladiomycota and Chytridiomycota. Fungal Diversity 73: 1–72. https://doi.org/10.1007/s13225-015-0339-4
  • Jones EBG, Pang KL, Abdel-Wahab MA, Scholz B, Hyde KD, Boekhout T, Ebel R, Rateb ME, Henderson L, Sakayaroj J, Suetrong S, Dayarathne MC, Kumar V, Raghukumar S, Sridhar KR, Bahkali AHA, Gleason FH, Norphanphoun C (2019) An online resource for marine fungi. Fungal Diversity 96(1): 347–433. https://doi.org/10.1007/s13225-019-00426-5
  • Karimi O, Chethana KT, de Farias AR, Asghari R, Kaewchai S, Hyde KD, Li Q (2024) Morphology and multigene phylogeny reveal three new species of Distoseptispora (Distoseptisporales, Distoseptisporaceae) on palms (Arecaceae) from peatswamp areas in southern Thailand. MycoKeys 102: 55. https://doi.org/10.3897/mycokeys.102.112815
  • Katoh K, Rozewicki J, Yamada KD (2019) MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics 20: 1160–1166. https://doi.org/10.1093/bib/bbx108
  • Kohlmeyer J, Kohlmeyer E (1979) Marine mycology. The higher fungi. Academic Press, New York.
  • Konta S, Hyde KD, Eungwanichayapant PD, Doilom M, Tennakoon DS, Senwanna C, Boonmee S (2020) Fissuroma (Aigialaceae: Pleosporales) appears to be hyperdiverse on Arecaceae: evidence from two new species from Southern Thailand. Acta Botanica Brasílica 34(2): 384–393. https://doi.org/10.1590/0102-33062020abb0021
  • Konta S, Hyde KD, Karunarathna SC, Mapook A, Senwanna C, Dauner LAP, Nanayakkara CM, Xu J, Tibpromma S, Lumyong S (2021) Multigene phylogeny and morphology reveal Haplohelminthosporium gen. nov. and Helminthosporiella gen. nov. associated with palms in Thailand and a checklist for Helminthosporium reported worldwide. Life (Basel, Switzerland) 11(5): 454. https://doi.org/10.3390/life11050454
  • Konta S, Tibpromma S, Karunarathna SC, Samarakoon MC, Steven LS, Mapook A, Boonmee S, Senwanna C, Balasuriya A, Eungwanichayapant PD, Hyde KD (2023) Morphology and multigene phylogeny reveal ten novel taxa in Ascomycota from terrestrial palm substrates (Arecaceae) in Thailand. Mycosphere 14(1): 107–152. https://doi.org/10.5943/mycosphere/14/1/2
  • Lemoine F, Correia D, Lefort V, Doppelt-Azeroual O, Mareuil F, Cohen-Boulakia S, Gascuel O (2019) NGPhylogeny.fr: new generation phylogenetic services for non-specialists. Nucleic Acids Research 47: W260–W265. https://doi.org/10.1093/nar/gkz303
  • Li QR, Habib K, Long SH, Wu YP, Zhang X, Hu HM, Wu QZ, Liu LL, Zhou SX, Shen XC, Kang JC (2024) Unveiling fungal diversity in China: new species and records within the Xylariaceae. Mycosphere 15(1): 275–364. https://doi.org/10.5943/mycosphere/15/1/2
  • Maddison WP, Maddison DR (2016) Mesquite: a modular system for evolutionary analysis. Version 3.10.
  • Maharachchikumbura SSN, Chen Y, Ariyawansa HA, Hyde KD, Haelewaters D, Perera RH, Samarakoon MC, Wanasinghe DN, Bustamante DE, Liu JK, Lawrence DP (2021) Integrative approaches for species delimitation in Ascomycota. Fungal Diversity 109: 155–179. https://doi.org/10.1007/s13225-021-00486-6
  • Marin-Felix Y, Hernández-Restrepo M, Iturrieta-González I, García D, Gené J, Groenewald JZ, Cai L, Chen Q, Quaedvlieg W, Schumacher RK, Taylor PWJ, Ambers C, Bonthond G, Edwards J, Krueger-Hadfield SA, Luangsa-ard JJ, Morton L, Moslemi A, Sandoval-Denis M, Tan YP, Thangavel R, Vaghefi N, Cheewangkoon R, Crous PW (2019) Genera of phytopathogenic fungi: GOPHY 3. Studies in Mycology 94(1): 1–124. https://doi.org/10.1016/j.simyco.2019.05.001
  • Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for Inference of Large Phylogenetic Trees. SC10 Workshop on Gateway Computing Environments (GCE10). https://doi.org/10.1109/GCE.2010.5676129
  • Pang KL, Overy DP, Jones EBG, da Luz Calado M, Burgaud G, Walker AK, Johnson JA, Kerr RG, Cha HJ, Bills GF (2016) ‘Marine fungi’ and ‘marine-derived fungi’ in natural product chemistry research: toward a new consensual definition. Fungal Biology Reviews 30: 163–175. https://doi.org/10.1016/j.fbr.2016.08.001
  • Phengsintham P, Braun U, McKenzie EHC, Chukeatirote E, Cai L, Hyde KD (2013) Monograph of cercosporoid fungi from Thailand. Plant Pathology & Quarantine 3(2): 67–138. https://doi.org/10.5943/ppq/3/2/2
  • Phukhamsakda C, McKenzie EHC, Phillips AJL, Jones EBG, Bhat DJ, Stadler M, Bhunjun CS, Wanasinghe DN, Thongbai B, Camporesi E, Ertz D, Jayawardena RS, Perera RH, Ekanayake AH, Tibpromma S, Doilom M, Xu JC, Hyde KD (2020) Microfungi associated with Clematis (Ranunculaceae) with an integrated approach to delimiting species boundaries. Fungal Diversity 102(1): 1–203. https://doi.org/10.1007/s13225-020-00448-4
  • Phukhamsakda C, Nilsson RH, Bhunjun CS, de Farias ARG, Sun YR, Wijesinghe SN, Hyde KD (2022) The numbers of fungi: contributions from traditional taxonomic studies and challenges of metabarcoding. Fungal Diversity 114: 1–60. https://doi.org/10.1007/s13225-022-00502-3
  • Rehner SA (2001) Primers for elongation factor 1‐alpha (EF1‐alpha). Insect Biocontrol Laboratory, USDA, ARS, PSI, Beltsville, MD.
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
  • Senanayake IC, Rathnayaka AR, Marasinghe DS, Calabon MS, Gentekaki E, Lee HB, Hurdeal VG, Pem D, Dissanayake LS, Wijesinghe SN, Bundhun D, Nguyen TT, Goonasekara ID, Abeywickrama PD, Bhunjun CS, Jayawardena RS, Wanasinghe DN, Jeewon R, Bhat DJ, Xiang MM (2020) Morphological approaches in studying fungi: Collection, examination, isolation, sporulation and preservation. Mycosphere 11: 2678–2754. https://doi.org/10.5943/mycosphere/11/1/20
  • Tanaka K, Hirayama K, Yonezawa H, Hatakeyama S, Harada Y, Sano T, Shirouzu T, Hosoya T (2009) Molecular taxonomy of bambusicolous fungi: Tetraplosphaeriaceae, a new pleosporalean family with Tetraploa-like anamorphs. Studies in Mycology 64: 175–209. https://doi.org/10.3114/sim.2009.64.10
  • Taylor JE, Hyde KD (2003) Microfungi of tropical and temperate palms. Fungal Diversity Press, Hong Kong, 459 pp.
  • Taylor JE, Hyde KD, Jones EBG (1999) Endophytic fungi associated with the temperate palm, Trachycarpus fortunei, within and outside its natural geographic range. New Phytologist 142: 335–346. https://doi.org/10.1046/j.1469-8137.1999.00391.x
  • Tibpromma S, Hyde KD, McKenzie EHC, Bhat DJ, Phillips AJL, Wanasinghe DN, Samarakoon MC, Jayawardena RS, Dissanayake AJ, Tennakoon DS, Doilom M, Phookamsak R, Tang AMC, Xu JC, Mortimer PE, Promputtha I, Maharachchikumbura SSN, Khan S, Karunarathna SC (2018) Fungal diversity notes 840–928: Micro-fungi associated with Pandanaceae. Fungal Diversity 93(1): 1–160. https://doi.org/10.1007/s13225-018-0408-6
  • To-Anun C, Nguenhom J, Meeboon J, Hidayat I (2009) Two fungi associated with necrotic leaflets of areca palms (Areca catechu). Mycological Progress 8: 115–121. https://doi.org/10.1007/s11557-009-0583-7
  • Vijaykrishna D, Jeewon R, Hyde KD (2006) Molecular taxonomy, origins and evolution of freshwater ascomycetes. Fungal Diversity 23: 351–390.
  • Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172(8): 4238–4246. https://doi.org/10.1128/jb.172.8.4238-4246.1990
  • White TJ, Bruns T, Lee SJWT, Taylor JL (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ (Eds) PCR protocols, a guide to methods and applications 18. Academic Press, San Diego, CA, 315–322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1
  • Xiong Y, Ishara MS, Hyde KD, Taylor JE, Phillips A, Pereira DS, Lu L, Zhang SN, Mapook A, Xu B (2024) Introducing palm fungi. org, an integrated fungal-host data platform. Biodiversity Data Journal 12: e126553. https://doi.org/10.3897/BDJ.12.e126553
  • Yeh YH, Kirschner R (2014) Sarocladium spinificis, a new endophytic species from the coastal grass Spinifex littoreus in Taiwan. Botanical Studies 55: 1–6. https://doi.org/10.1186/1999-3110-55-25
  • Youssef K, Mustafa ZM, Kamel MA, Mounir GA (2020) Cigar end rot of banana caused by Musicillium theobromae and its control in Egypt. Archives of Phytopathology and Plant Protection 53(3–4): 162–177. https://doi.org/10.1080/03235408.2020.1735139
  • Zare R, Gams W, Starink-Willemse M, Summerbell RC (2007) Gibellulopsis, a suitable genus for Verticillium nigrescens, and Musicillium, a new genus for V. theobromae. Nova Hedwigia 85(3–4): 463–489. https://doi.org/10.1127/0029-5035/2007/0085-0463
  • Zhang SN, Hyde KD, Jones EBG, Yu XD, Cheewangkoon R, Liu JK (2024) Current insights into palm fungi with emphasis on taxonomy and phylogeny. Fungal Diversity 127(1): 55–301. https://doi.org/10.1007/s13225-024-00536-9
  • Zhao HJ, Doilom M, Mapook A, Wang G, Hyde KD, Dong W (2024) New insights into Tetraplosphaeriaceae based on taxonomic investigations of bambusicolous fungi and freshwater fungi. Journal of Fungi 10(5): 319. https://doi.org/10.3390/jof10050319
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