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Research Article
Morpho-molecular characterisation of Arecophila, with A. australis and A. clypeata sp. nov. and A. miscanthi comb. nov.
expand article infoQi Rui Li‡§, Xu Zhang§, Yan Lin§, Milan C. Samarakoon|, Kevin David Hyde, Xiang Chun Shen§, Wan Qing Liao#, Anuruddha Karunarathna, Si Han Long§, Ying Qian Kang§, Ji Chuan Kang‡
‡ Guizhou University, Guizhou, China
§ Guizhou Medical University, Guizhou, China
| Chiang Mai University, Chiang Mai, Thailand
¶ Mae Fah Luang University, Chiang Rai, Thailand
# Department of Dermatology and Venereology, Changzheng Hospital, Shanghai, China

Corresponding author: Ying Qian Kang ( kangyingqian@gmc.edu.cn )

Corresponding author: Ji Chuan Kang ( jckang@gzu.edu.cn )

Academic editor: Sajeewa Maharachchikumbura
© 2022 Qi Rui Li, Xu Zhang, Yan Lin, Milan C. Samarakoon, Kevin David Hyde, Xiang Chun Shen, Wan Qing Liao, Anuruddha Karunarathna, Si Han Long, Ying Qian Kang, Ji Chuan Kang.
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: Li QR, Zhang X, Lin Y, Samarakoon MC, Hyde KD, Shen XC, Liao WQ, Karunarathna A, Long SH, Kang YQ, Kang JC (2022) Morpho-molecular characterisation of Arecophila, with A. australis and A. clypeata sp. nov. and A. miscanthi comb. nov. MycoKeys 88: 123-149. https://doi.org/10.3897/mycokeys.88.79475
Open Access

Abstract

Three arecophila-like fungal samples were collected on dead culms of gramineous plants in China. Morphological studies of our new collections and the herbarium specimen of Arecophila gulubiicola (generic type) were conducted and the morphological affinity of our new collections with Arecophila was confirmed. Maximum likelihood and Bayesian analyses using combined ITS, LSU, rpb2 and β-tubulin data from our collections revealed the phylogeny of Cainiaceae. The monospecific genus Alishanica (type species Al. miscanthi), which had been accepted in Cainiaceae, is revisited and synonymised under Arecophila. Based on morphology and phylogeny, Arecophila australis sp. nov. and A. clypeata sp. nov. are introduced as new species, while A. miscanthi is a new record for China. All the new collections are illustrated and described.

Keywords

Cainiaceae, gramineous plants, phylogeny, taxonomy

Introduction

The current study is a part of a series of papers on Xylariales (Sordariomycetes) from China (Long et al. 2019; Xie et al. 2019, 2020; Pi et al. 2020). Arecophila K.D. Hyde, which is typified by A. gulubiicola K.D. Hyde, was introduced by Hyde (1996) with five species. Arecophila is characterised by immersed, subglobose to lenticular ascomata, peridium with textura angularis cells, non- or poorly-developed clypeus, asci with a wedge-shaped, apical ring, J+ in Melzer’s reagent and 2-celled, brown ascospores with wall striations, surrounded by a mucilaginous sheath. Thanks to subsequently undertaken morphological studies of holotypes, several species have been transferred to Arecophila from genera such as Amphisphaeria Ces. & De Not., Cainia Arx & E. Müll., Didymosphaeria Fuckel and Schizostoma Ehrenb. ex Lév. (Hyde 1996; Umali et al. 1999; Wang et al. 2004).

Currently, there are 15 Arecophila epithets in Index Fungorum (http://www.indexfungorum.org/Names/Names.asp, May 2021), which have been introduced, based on morphology and lack sequence data (e.g. Hyde 1996; Umali et al. 1999; Wang et al. 2004). After searching for Arecophila in NCBI, there were only five hits of LSU, SSU and metagenomic sequences of A. bambusae and Arecophila sp. HKUCC 6487 in GenBank.

Arecophila was introduced as a genus of Amphisphaeriaceae (Hyde 1996), based on its unitunicate, cylindrical asci with a J+ apical ring and brown, 2-celled ascospores. Kang et al. (1999) reviewed the genus and accepted it in Cainiaceae and the occurrence on monocotyledons (palms and bamboo). The single and combined molecular analyses of LSU and SSU genes resulted in Arecophila grouping with Cainia in Xylariales (Smith et al. 2003). Based on analyses of partial LSU gene sequences, the generic placement of Arecophila within the Cainiaceae has been verified (Jeewon et al. 2003; Senanayake et al. 2015; Hyde et al. 2020; Wijayawardene et al. 2020). However, the available molecular data do not provide strong evidence of the phylogenetic affinity of Arecophila and related taxa.

During our continuous collecting of xylarialean taxa in China, we found some specimens that share a morphology resembling Arecophila. In this paper, two new species and a new record of Arecophila are provided with descriptions and illustrations. Furthermore, Alishanica is synonymised under Arecophila, based on morphology and phylogeny.

Materials and methods

Collection, isolation and morphology

Fresh samples were collected in Guizhou and Yunnan Provinces in China during the rainy season and taken to the laboratory in paper bags. Single-spore isolations were obtained following the method described in Chomnunti et al. (2014). The cultures on potato dextrose agar (PDA) were transferred to 2 ml screw cap centrifuge tubes filled with 10% glycerol and sterile water to deposit at –20 °C and 4 °C, respectively. Herbarium materials were deposited at the Herbarium of Guizhou Agricultural College (GACP) and the Herbarium of Guizhou University (GZUH). Cultures were deposited at the Culture Collection of Guizhou University (GZUCC).

The morphological examination of fresh and herbarium specimens was carried out as described by Hyde (1996). Macro-morphological characters were examined and photographed using a digital camera (Canon 700D) fitted to the Olympus SZ61 stereomicroscope. Materials mounted in water, Melzer’s reagent and Indian ink were examined. At least 30 ascospores, 30 asci and 20 apical rings were measured for each taxa with Tarosoft (R) Image Frame Work (v. 0.9.0.7) and photographed using a digital camera (Nikon 700D) fitted to a light microscope (Nikon Ni).

DNA extraction, polymerase chain reaction (PCR) amplification and sequencing

Total genomic DNA was extracted from fresh mycelium scraped off from pure cultures with the BIOMIGA fungus genomic DNA extraction kit (GD2416) (Wijayawardene et al. 2013) following the manufacturer’s instructions. Primers, LR0R/LR5 (Vilgalys and Hester 1990), ITS4/ITS5 (White et al. 1990), RPB2-5F/RPB2-7cR (Liu et al. 1999), Bt2a/Bt2b and ACT-512F/ACT-783R (Hsieh et al. 2005) were used for amplifying partial large-subunit ribosomal RNA (LSU), internal transcribed spacer (ITS), partial second-largest subunit of the RNA polymerase II (rpb2), β-tubulin (tub) and α-actin gene (Hsieh et al. 2005). The amplification conditions were carried out according to Liu et al. (2011) and Hsieh et al. (2005). Amplified products were examined and sent to the sequencing company, Sangon Biotech, Shanghai, China. The obtained sequences were checked, assembled and uploaded to GenBank.

Sequence alignment and phylogenetic analyses

Following the NCBI BLAST results and literature (e.g. Jeewon et al. 2003; Senanayake et al. 2015), relevant sequences from all families of Xylariomycetidae were downloaded from GenBank for the phylogenetic analyses (Table 1). Sequences of each segment were aligned using MAFFT (http://mafft.cbrc.jp/alignment/server/index.html, Katoh and Standley 2019) and improved manually in BioEdit 7.2.3 (Hall 1999). The combined alignment of ITS, LSU, rpb2 and β-tubulin was concatenated from individual datasets. Ambiguously aligned areas of each gene region were excluded and gaps were treated as missing data. The ALTER (http://sing.ei.uvigo.es/ALTER/) phylogeny website tool was used to obtain the phylip file for RAxML analysis and the nexus file for Bayesian analysis (Glez-Peña et al. 2010). Phylogenetic trees were visualised using FigTree v.1.4.0. and processed using Adobe Photoshop CS6 software (Adobe Systems, USA). The alignment for the tree in this paper was uploaded on the website (https://treebase.org/) with submission ID 26613.

Table 1.
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Sequences used for phylogenetic analyses in this study.

Species Strain number Status GenBank accession numbers References
ITS LSU rpb2 β-tubulin
Achaetomium macrosporum CBS 532.94 – KX976574 KX976699 KX976797 KX976915 Wang et al. (2016)
Amphibambusa bambusicola MFLUCC 11-0617 HT KP744433 KP744474 N/A N/A Senanayake et al. (2015)
Amphisphaeria acericola MFLU 16-2479 HT NR_171945 MK640424 N/A N/A Senanayake et al. (2019, submitted directly)
Amphisphaeria thailandica MFLU 18-0794 HT NR_168783 NG_068588 MK033640 MK033639 Samarakoon et al. (2019)
Amphisphaeria umbrina AFTOL-ID 1229 AF009805 N/A FJ176863 FJ238348 N/A Schoch (2008, submitted directly)
Apiospora bambusae ICMP 6889 – N/A DQ368630 DQ368649 N/A Tang et al. (2007)
Apiospora hyphopodii MFLUCC 15-0003 HT KR069110 KY356093 N/A N/A Dai et al. (2016)
Apiospora setosa ICMP 4207 – N/A DQ368631 DQ368650 DQ368620 Tang et al. (2007)
Apiospora yunnana MFLUCC 15-0002 HT KU940147 NG_057104 KU940177 MK291950 Dai et al. (2017)
Arecophila australis GZUCC0112 HT MT742126 MT742133 N/A MT741734 This study
Arecophila australis GZUCC0124 – MT742125 MT742132 N/A N/A This study
Arecophila bambusae HKUCC 4794 – N/A AF452038 N/A N/A Kang et al. (1999)
Arecophila clypeata GZUCC0110 HT MT742129 MT742136 MT741732 N/A This study
Arecophila clypeata GZUCC0127 – MT742128 MT742135 N/A N/A This study
Arecophila miscanthi GZUCC0122 – MT742127 MT742134 N/A N/A This study
Arecophila miscanthi MFLU 19-2333 HT NR_171235 MK503827 N/A N/A Hyde et al. (2020)
Arecophila sp. HKUCC 6487 – N/A AF452039 N/A N/A Jeewon et al. (2003)
Apiospora yunnana MFLUCC 15-0002 HT KU940147 NG_057104 KU940177 MK291950 Dai et al. (2017)
Atrotorquata spartii MFLUCC 13-0444 HT N/A KP325443 N/A N/A Thambugala et al. (2015)
Bagadiella lunata CBS 124762 HT NR_132832 NG_058637 N/A N/A Cheewangkoon et al. (2009)
Barrmaelia rappazii Cr2 = CBS 142771 HT MF488989 MF488989 MF488998 MF489017 Voglmayr et al. (2018)
Barrmaelia rhamnicola BR = CBS 142772 HT MF488990 MF488990 MF488999 MF489018 Voglmayr et al. (2018)
Bartalinia pondoensis CMW 31067 – MH863602 MH875078 MH554904 MH554663 Vu et al. (2019)
Beltrania pseudorhombica CBS 138003 HT MH554124 NG_058667 MH555032 N/A Liu et al. (2019)
Beltrania rhombica CBS 123.58 T MH857718 MH868082 MH554899 MH704631 Vu et al. (2019)
Beltraniopsis longiconidiophora MFLUCC 17-2139 HT NR_158353 NG_066200 N/A N/A Liu et al. (2017)
Biscogniauxia nummularia MUCL 51395 ET KY610382 KT281894 KY624236 KX271241 Senanayake et al. (2015)
Cainia anthoxanthis MFLUCC 15-0539 HT NR_138407 KR092777 N/A N/A Senanayake et al. (2015)
Cainia graminis CBS 136.62 – MH858123 AF431949 N/A N/A Vu et al. (2019)
Cainia graminis MFLUCC 15-0540 – KR092793 KR092781 N/A N/A Senanayake et al. (2015)
Camillea obularia ATCC 28093 – KY610384 KY610429 KY624238 KX271243 Wendt et al. (2018)
Castanediella acaciae CBS 139896 HT NR_137985 NG_067293 N/A N/A Crous et al. (2015)
Castanediella couratarii CBS 579.71 HT NR_145250 NG_066249 N/A N/A Vu et al. (2019)
Castanediella eucalypticola CPC 26539 HT KX228266 KX228317 N/A KX228382 Crous et al. (2013)
Chaetomium elatum CBS 374.66 – KC109758 KC109758 KF001820 KC109776 Wang et al. (2016)
Ciferriascosea fluctuatimura MFLUCC 15-0541 HT KR092789 KR092778 N/A N/A Senanayake et al. (2015)
Ciferriascosea rectimura MFLUCC 15-0542 HT NR_153905 KR092776 N/A N/A Senanayake et al. (2015)
Clypeophysalospora latitans CBS 141463 ET NR_153929 NG_058958 N/A N/A Giraldo et al. (2017)
Coniocessia maxima CBS 593.74 HT NR_137751 MH878275 N/A N/A Vu et al. (2019)
Coniocessia nodulisporioides CBS 281.77 IT MH861061 AJ875224 N/A N/A García et al. (2006)
Creosphaeria sassafras STMA 14087 – KY610411 KY610468 KY624265 KX271258 Wendt et al. (2018)
Cylindrium aeruginosum CBS 693.83 – KM231854 KM231734 KM232430 KM232124 Lombard et al (2014, submitted directly)
Cylindrium grande CBS 145655 HT NR_165557 NG_068656 MK876481 MK876502 Crous et al. (2019)
Cylindrium purgamentum CPC 29580 HT NR_155691 NG_067320 N/A N/A Koppel et al. (2017)
Daldinia concentrica CBS 113277 – AY616683 KT281895 KY624243 KC977274 Senanayake et al. (2015)
Delonicicola siamense MFLUCC 15-0670 HT MF167586 NG_059172 MF158346 N/A Perera et al. (2017)
Diatrype palmicola MFLUCC 11-0018 – KP744439 KP744481 N/A N/A Liu et al. (2015)
Diatrype whitmanensis ATCC MYA-4417 – FJ746656 FJ430587 N/A N/A Igo et al. (2009, direct submission)
Entosordaria perfidiosa EPE = CBS 142773 ET MF488993 MF488993 MF489003 MF489021 Voglmayr et al. (2018)
Entosordaria quercina RQ = CBS 142774 HT MF488994 MF488994 MF489004 MF489022 Voglmayr et al. (2018)
Eutypa flavovirens MFLUCC 13-0625 – KR092798 KR092774 N/A N/A Senanayake et al. (2015)
Eutypa laevata CBS 291.87 – HM164737 N/A HM164805 HM164771 Trouillas and Gubler (2010)
Eutypa lata CBS 208.87 NT MH862066 MH873755 KF453595 DQ006969 Vu et al. (2019)
Furfurella nigrescens CBS 143622 HT MK527844 MK527844 MK523275 MK523332 Voglmayr et al. (2019)
Furfurella stromatica CBS 144409 HT NR_164062 MK527846 MK523277 MK523334 Voglmayr et al. (2019)
Graphostroma platystomum AFTOL-ID 1249 HT HG934115 DQ836906 DQ836893 HG934108 Zhang et al. (2006)
Hyponectria buxi UME 31430 – - AY083834 N/A N/A Smith et al. (2002, submitted directly)
Hypoxylon fragiforme MUCL51264 ET KM186294 KM186295 KM186296 KM186293 Daranagama et al. (2015)
Iodosphaeria honghensis MFLU 19-0719 HT MK737501 MK722172 MK791287 N/A Marasinghe et al. (2019)
Iodosphaeria tongrenensis MFLU 15-0393 HT KR095282 KR095283 N/A N/A Li et al. (2015)
Jackrogersella multiformis CBS 119016 ET KC477234 KT281893 KY624290 KX271262 Wendt et al. (2018)
Kretzschmaria deusta CBS 163.93 – KC477237 KT281896 KY624227 KX271251 Senanayake et al. (2015)
Lepteutypa fuckelii CBS 140409 NT NR_154123 KT949902 MK523280 MK523337 Jaklitsch et al. (2016)
Leptosillia pistaciae CBS 128196 HT NR_160064 MH798901 MH791334 MH791335 Voglmayr et al. (2019)
Leptosillia wienkampii CBS 143630 ET NR_164067 MK527865 MK523297 MK523353 Voglmayr et al. (2019)
Longiappendispora chromolaenae MFLUCC 17-1485 HT NR_169723 NG_068714 N/A N/A Mapook et al. (2020)
Lopadostoma americanum LG8 HT KC774568 KC774568 KC774525 N/A Jaklitsch et al. (2014)
Lopadostoma dryophilum LG21 ET KC774570 KC774570 KC774526 MF489023 Jaklitsch et al. (2014)
Lopadostoma fagi LF1 HT KC774575 KC774575 KC774531 N/A Jaklitsch et al. (2014)
Lopadostoma quercicola LG27 HT KC774610 KC774610 KC774558 N/A Jaklitsch et al. (2014)
Lopadostoma turgidum LT2 ET KC774618 KC774618 KC774563 MF489024 Jaklitsch et al. (2014)
Melogramma campylosporum MBU – JF440978 JF440978 N/A N/A Jaklitsch and Voglmayr (2012)
Neophysalospora eucalypti CBS 111123 – KP031107 KP031109 N/A N/A Crous et al. (2014)
Neophysalospora eucalypti CBS 138864 HT KP004462 MH878627 N/A N/A Crous et al. (2014)
Oxydothis metroxylicola MFLUCC 15-0281 HT KY206774 KY206763 KY206781 N/A Konta et al. (2016)
Oxydothis palmicola MFLUCC 15-0806 HT KY206776 KY206765 KY206782 N/A Konta et al. (2016)
Oxydothis phoenicis MFLUCC 18-0269 HT MK088065 MK088061 N/A N/A Hyde et al. (2020)
Phlogicylindrium uniforme CBS 131312 HT JQ044426 JQ044445 MH554910 MH704634 Crous et al. (2011)
Podosordaria tulasnei CBS 128.80 – KT281902 KT281897 N/A N/A Senanayake et al. (2015)
Poronia punctata CBS 656.78 HT KT281904 KY610496 KY624278 KX271281 Wendt et al. (2018)
Pseudomassaria chondrospora MFLUCC 15-0545 – KR092790 KR092779 N/A N/A Senanayake et al. (2015)
Pseudomassaria sepincoliformis CBS 129022 – JF440984 JF440984 N/A N/A Jaklitsch and Voglmayr (2012)
Pseudosporidesmium knawiae CBS 123529 HT MH863299 MH874823 N/A N/A Crous et al. (2017, submitted directly)
Pseudosporidesmium lambertiae CBS 143169 HT NR_156656 NG_058506 N/A N/A Crous et al. (2017)
Pseudotruncatella arezzoensis MFLUCC 14-0988 HT NR_157489 NG_070426 N/A N/A Perera et al. (2018)
Pseudotruncatella bolusanthi CBS 145532 HT NR_165575 MK876448 N/A N/A Crous et al. (2019)
Robillarda roystoneae CBS 115445 HT NR_145251 NG_069287 MH554880 KR873317 Liu et al. (2019)
Sarcoxylon compunctum CBS 359.61 – MH858083 KY610462 KY624230 KX271255 Wendt et al.(2018)
Seiridium marginatum CBS 140403 ET NR_156602 MH554223 LT853149 MT853249 Liu et al. (2019)
Seynesia erumpens SMH 1291 – N/A AF279410 AY641073 N/A Bhattacharya et al. (2000)
Sordaria fimicola CBS 723.96 – MH862606 MH874231 DQ368647 DQ368618 Vu et al. (2019)
Sporocadus rotundatus CBS 616.83 HT NR_161091 NG_069584 MH554974 MH554737 Liu et al. (2019)
Subsessila turbinata MFLUCC 15-0831 HT NR_148122 NG_059724 N/A N/A Lin et al. (2017)
Vialaea insculpta DAOM 240257 – JX139726 JX139726 N/A N/A Hambleton et al. (2010, submitted directly)
Vialaea mangiferae MFLUCC 12-0808 HT NR_171903 NG_073594 N/A N/A Senanayake et al. (2021, submitted directly)
Vialaea minutella BRIP 56959 – KC181926 KC181924 N/A N/A McTaggart et al. (2013)
Xyladictyochaeta lusitanica CBS 143502 – MH107926 MH107972 N/A MH108053 Crous et al. (2013)
Xylaria hypoxylon CBS 122620 ET KY610407 KY610495 KY624231 KX271279 Wendt et al. (2018)
Xylaria obovata MFLUCC 13-0115 – KR049088 KR049089 N/A N/A Wendt et al. (2018)
Xylaria polymorpha MUCL 49884 – KY610408 KT281899 KY624288 KX271280 Wendt et al. (2018)

Note. Type specimens are labelled with HT (holotype), ET (epitype) and IT (isotype), T (Type). N/A: not available.

Maximum likelihood (ML) analysis was performed on the CIPRES Science Gateway v.3.3 (http://www.phylo.org/portal2; Miller et al. 2010) using RAxML v.8.2.8 as part of the ‘RAxML-HPC BlackBox’ tool (Stamatakis et al. 2008). All free model parameters were estimated by RAxML with ML estimates of 25 per-site rate categories. GTRGAMMA + I model was chosen for RAxML, based on the result of MrModeltest 2.2. The best-scoring tree was selected with a final likelihood value of –10720.566919.

A Bayesian analysis (BY) was performed using MrBayes v.3.2.2 (Ronquist et al. 2012). The best-fit model was selected with MrModeltest 2.2 (Nylander 2004). Posterior probabilities (PP) (Rannala and Yang 1996) were determined by Markov Chain Monte Carlo sampling (MCMC) (Ronquist and Huelsenbeck 2003). Six simultaneous Markov chains were initially run for 30 × 106 generations and for every 1000th generation, a tree was sampled (resulting in 30,000 total trees). The MCMC heated chain was set with a ‘temperature’ value of 0.15. All sampled topologies beneath the asymptote (20%) were discarded. The remaining 24,000 trees were used to calculate the posterior probability (PP) values in the majority rule consensus tree (Liu et al. 2011).

Results

Phylogenetic analyses

The resulted trees from ML and BY were similar in topology. Cainiaceae is a monophyletic group (Fig. 1) with 100%/1.00 (PP/BS) support. Arecophila species form two clades. Clade 1 consists of A. miscanthi (≡ Alishanica miscanthi), A. clypeata and A. australis, with high statistical support (100%/1.00 PP). In Clade 2, A. bambusae (HKUCC 4794) and Arecophila sp. (HKUCC 6487) display a close relationship with Amphibambusa bambusicola.

Figure 1. 

Phylogenetic tree, based on a combined ITS, LSU, rpb2 and β-tubulin gene dataset. Numbers close to each node represent Maximum Likelihood bootstrap values (≥ 75%) and Bayesian posterior probabilities (≥ 0.95). The hyphen (“–”) means a value lower than 75% (BS) or 0.95 (PP). New taxa are marked in red. Type materials are marked with T after the strains. The tree is rooted to Achaetomium macrosporum (CBS 532.94), Chaetomium elatum (CBS 374.66) and Sordaria fimicola (CBS 723.96).

Taxonomy

Arecophila K.D. Hyde, Nova Hedwigia 63(1–2): 82 (1996)

MycoBank No: 27653

≡ Alishanica Karun., C.H. Kuo & K.D. Hyde, in Hyde et al., Mycosphere 11(1): 460 (2020)

Sexual morph

Ascomata immersed, raised, blackened areas on the host surface, a central erumpent, short, cone-shaped or umbilicate papilla, subglobose to lenticular in vertical section. Clypeus present or not, comprising host cells and intracellular brown hyphae. Peridium comprising several layers of angular cells. Paraphyses hypha-like, filamentous, septate, hyaline. Asci 8-spored, unitunicate, cylindrical, with an apical ring bluing in Melzer’s reagent or not. Ascospores ellipsoidal, 2-celled, constricted at the septum, brown, with longitudinal striations or a verrucose wall and surrounded by a wide mucilaginous sheath (Hyde 1996).

Asexual morph

Undetermined.

Arecophila australis Q.R. Li, J.C. Kang & K.D. Hyde, sp. nov.

MycoBank No: 836166
Fig. 2

Diagnosis

Arecophila australis differs from similar species by its dimension of ascospores (22.5–29 × 8–11 µm) covered by striations and ascomata with a disc area surrounding the ostioles.

Figure 2. 

Arecophila australis (holotype) A material B ascoma on the surface of host C section of ascoma D peridium E paraphyses F, G ascus apex with a J+, apical ring (stained in Melzer’s reagent) H–K asci with ascospores L–O ascospores surrounded by a wide mucilaginous sheath (O stained in India ink). Scale bars: 300 μm (B); 50 μm (C); 5 μm (D–O).

Holotype. China, Guizhou Province, Guiyang City, Forest Park of Guiyang (26°32'55"N, 106°45'25"E), on dead culm of Phragmites australis (Cav.) Steud., 15 March 2014, Q.R. Li, GZ58 (GZUH0112, holotype, ex-type: GZUCC0112; GACP QR0152, isotype).

Additional sequences

ACT: MT741737

Etymology

In reference to the host, Phragmites australis (Cav.) Steud. australis

Description

Saprobic on dead culm of gramineous host. Sexual morph: Ascomata 420–560 × 290–380 µm (x̄= 495 × 325 µm, n = 10), immersed under a clypeus, solitary, slightly raised, blackened, dome-shaped areas, scattered or gregarious, globose to subglobose, with a central, erumpent, cone-shaped papilla in vertical section. Clypeus black, comprising host cells and intracellular brown hyphae. Ostioles papillate, black. Peridium 15–25 µm (x̄= 21 µm, n = 15) wide, comprising several layers, outer layer brown, thick-walled angular cells, inner layer hyaline. Paraphyses 3.3–5 μm (x̄= 3.5 µm, n = 15) wide, hyaline, unbranched, septate. Asci 140–230 × 15.5–24 µm (x̄= 183.5 × 19 µm, n = 30), 8-spored, unitunicate, long-cylindrical, short-pedicellate, apically rounded, with a 4–5 × 2.5–3 μm (x̄= 4.5 × 2.7 μm, n = 20), trapezoidal, J+, apical ring. Ascospores 22.5–29 × 8–11 µm (x̄= 25.5 × 9 µm, n = 30), overlapping uniseriate, 2-celled, light brown to brown, equilateral ellipsoidal, constricted at the septum, longitudinal with sulcate striations, along the entire spore length, surrounded by a mucilaginous sheath, lacking germ slits and appendages. Asexual morph: undetermined.

Culture characteristics

Colonies on PDA, reached 3 cm diam. after one week at 25 °C, white, cottony, flat, low, dense, with slightly wavy margin.

Known distribution

China

Additional material examined

China, Guizhou Province, Guiyang City, Leigongshan National Nature Reserve (26°21'39"N, 108°9'59"E), on dead culm of an unidentified gramineous plant, 13 June 2015, Q.R. Li, GY67 (GACP QR0124, GZUH 0136; living cultures, GZUCC0124).

Notes

Arecophila australis resembles A. serrulata (Ellis & Martin) K.D. Hyde and A. calamicola K.D. Hyde (Hyde 1996). However, A. serrulata has white ring surrounding ostioles of ascomata, narrower ascospores (17–26 × 7–9.5 µm vs. 22.5–29 × 8–11 µm), smaller asci and apical ring (3.2 × 2.4 µm vs. 4.5 × 2.7 μm) compared to A. australis (Hyde 1996). Arecophila calamicola differs from A. australis in lacking clypeus, ascospores covered by verrucose ornamentation and surrounding by a mucilaginous sheath attached at the poles. Molecular phylogeny, based on combined ITS, LSU, rpb2 and β-tubulin sequences, shows that A. australis clusters as a distinctive clade in Arecophila (Clade 1). Based on its distinct morphology and phylogeny, A. australis is introduced as a new species. Here, we need to explain the name of A. serrulata. Although Index Fungorum (02/07/2022) shows that the current name of A. serrulata is Roussoella serrulata (Ellis & G. Martin) K.D. Hyde & Aptroot, we have not found relevant literature. Hyde (1996) renamed Didymosphaeria serrulota Eltis & G. Martin and Roussoella serrulata as synonyms of A. serrulata (Ellis & G. Martin) K.D. Hyde. Arecophila serrulata was erected with the unitunicate asci with a blue-staining ring (Hyde 1996) which is clearly inconsistent with the morphological features of Roussoella Sacc. Therefore, we still compare with the original description of A. serrulata in this article.

Arecophila clypeata Q.R. Li, J.C. Kang & K.D. Hyde, sp. nov.

MycoBank No: 836167
Fig. 3

Diagnosis

Arecophila clypeata differs from similar species by its ascomata with clypeus and ascospores (18.5–22.5 × 6.5–9 µm).

Holotype. China, Yunnan Province, Kunming City, Kunming Botanical Garden (25°8'51"N, 102°44'57"E), on dead culm of gramineous plant, 20 March 2014, Q.R. Li, kib21 (holotype: GZUH0110; isotype: GACP QR0173; ex-type living cultures: GZUCC0110).

Figure 3. 

Arecophila clypeata (holotype) A material B ascomata on the surface of host C, D section of ascomata E peridium F, G ascus apex with a J+, apical ring (stained in Melzer’s reagent) H–K asci with ascospores L–O ascospores. Scale bars: 500 μm (B, C); 100 μm (D); 10 μm (E, H–K); 5 μm (F, G, L–O).

Etymology

In reference to the clypeus.

Description

Saprobic on dead stem of a gramineous. Sexual morph: Ascomata 367–448 × 278–363 µm (x̄= 403 × 323 µm, n = 8), immersed under a black clypeus, solitary, slightly raised, dome-shaped areas, scattered or gregarious, subglobose to globose, with a central, erumpent, cone-shaped papilla, in vertical section. Ostioles papillate on the centre, black. Peridium 15–30 µm (x̄= 25 µm, n = 10) wide, comprising several layers, outer layer brown, thick-walled angular cells, inner layer hyaline. Paraphyses 3–5 μm (x̄= 4 µm, n =15) wide, hyaline, unbranched, septate. Asci 180–245 × 10.5–14.5 µm (x̄= 215.5 × 12 µm, n=20), 8-spored, unitunicate, long-cylindrical, short-pedicellate, apically rounded, with a square-shaped, J+, apical ring, 3–4 × 3–4 μm. Ascospores 18.5–22.5 × 6.5–9 µm (x̄= 20.5 × 7.5 µm, n = 30), overlapping uniseriate, 2-celled, light brown to brown, equilateral ellipsoidal, constricted at the septum, longitudinal, sulcate along the entire spore length, faint, surrounded by a mucilaginous sheath, lacking germ slits and appendages. Asexual morph: undetermined.

Culture characteristics

Colonies on PDA, reached 3 cm diam. after one week at 25 °C, white, cottony, flat, low, dense, with slightly wavy margin; fructifications were not observed in culture.

Known distribution

China

Additional material examined

China, Guizhou Province, Buyi and Miao Autonomous Prefecture in southern Guizhou Province, Maolan National Nature Reserve (25°17'17"N, 107°59'1"E), on dead culm of an unidentified gramineous plant, 12 June 2015, Q.R. Li, GZ120 (GACP QR0129; GZUH0127; living cultures, GZUCC0127).

Additional sequences

ACT: MT741737

Notes

Arecophila clypeata has long and weakly striate ascospores similar to A. coronata (Rehm) Umali & K.D. Hyde, A. serrulata (Ellis & G. Martin) K.D. Hyde and A. bambusae (Hyde 1996; Umali et al. 1999). However, A. coronata does not have a prominent clypeus and has longer and fusiform ascospores. Arecophila clypeata differs from A. serrulata by the ascomata without a central papilla surrounded by a circle of white tissue, further in having ascospores with wide sheaths (Hyde 1996). Arecophila clypeata is similar to A. bambusae which, however, has narrower ascospores (19–22.5 × 5.5–7 µm) covered by the strong striations and has ascomata without a central papilla surrounded by a black corolla protuberance (Umali et al. 1999).

Arecophila gulubiicola K.D. Hyde, Nova Hedwigia 63(1–2): 91 (1996)

MycoBank No: 416041
Fig. 4

Description

Saprobic on dead trunk of Gulubia costata (Becc.) Becc. Sexual morph: Ascomata 290–400 × 140–190 μm (x̄ = 336 × 167 µm, n = 8), immersed under a clypeus, solitary or clustered, in vertical section, lenticular, with a central ostiole. Clypeus raised, oval, blackened areas on the host surface, dome-shaped, well-developed and black. Peridium 25–35 μm wide, dense, compressed layers of brown-walled, angular cells, tightly adhered to the host tissues. Paraphyses 2–2.5 μm wide, filamentous, hyaline, septate, branched, tapering distally. Asci 107–145 × 11–13.5 μm (x̄ = 114.3 × 12.4 μm, n = 15), 8-spored, unitunicate, cylindrical, short-pedicellate, apically rounded, wedge-shaped, J+, subapical ring, 3–4 × 1–2 μm (x̄ =3.5 × 1.5 μm, n = 15). Ascospores 14.5–18.5 × 6–9 μm (x̄ = 17.4 × 6.5 μm, n = 25), overlapping uniseriate, ellipsoidal, brown, 2-celled, septate at the centre, constricted at the septum, longitudinal, sulcate striations along the entire spore length, surrounded by a mucilaginous sheath. Asexual morph: Undetermined.

Figure 4. 

Arecophila gulubiicola (BRIP 23002a, holotype) A, B herbarium material with label C ascomata on the host D, E sections of ascomata F paraphyses G–J asci K peridium L, M wedge-shaped, J+ apical ring bluing in Melzer’s reagent N–Q ascospores. Scale bars: 50 μm (D, E); 5 μm (F–Q).

Material examined

Papua New Guinea, Central Province, 08°30'00"N, 147°24'35"E, on dead trunk of G. costate (Becc.) Becc. (Arecaceae), May 1992, K.D. Hyde, (BRIP 23002a, holotype).

Notes

Arecophila gulubiicola has deeply immersed, subglobose to lenticular ascomata with a small or lacking clypeus, cylindrical, short-pedicellate asci with a wedge-shaped, conical, apical ring and ellipsoidal, brown ascospores with wall striations and surrounded by a mucilaginous sheath (Hyde 1996). Alishanica has been introduced as a monospecific genus with the type species Al. miscanthi Karun. et al. on dead sheaths of Miscanthus sinensis (Poaceae) from Taiwan (Hyde et al. 2020). We re-examined both A. gulubiicola and Al. miscanthi herbarium specimens and observed that they are congeneric. Alishanica miscanthi has characters that immersed ascoma under a clypeus, unitunicate, cylindrical asci with a J+ apical ring and brown, 2-celled ascospores with longitudinal wall striations and a mucilaginous sheath which are consistent with the generic characteristics of Arecophila. The phylogeny of Al. miscanthi was mainly considered by the A. bambusae (HKUCC 4794) sequences (Hyde et al. 2020). However, HKUCC 4794 is not the type material of Arecophila and cannot be used to represent Arecophila. In our phylogeny, HKUCC 4794 forms a distinct clade (Fig. 1; Clade 2) from the Arecophila representing the clade. Based on morphology and phylogeny, we synonymise Alishanica under Arecophila and Al. miscanthi is accepted as an Arecophila species. Furthermore, A. bambusae needs to be recollected and provided with the phylogenetic affinity in future studies.

Arecophila miscanthi (Karun., C.H. Kuo & K.D. Hyde) Q.R Li & J.C. Kang, comb. nov.

MycoBank No: 839706

≡ Alishanica miscanthi Karun., C.H. Kuo & K.D. Hyde [as ‘miscanthii’], in Hyde et al., Mycosphere 11(1): 461 (2020)

Description

(MFLU 19-2333). Saprobic on dead sheaths of Miscanthus sinensis (Poaceae). Sexual morph: Ascomata 272–277 × 283–296 µm (x̄ = 275 × 291.5 µm, n = 8), immersed beneath blackened aggregated clypeus of the surface of dead sheath, loosely aggregated or rarely solitary; dark brown to black, globose to subglobose, slightly depressed, uniloculate. Ostiole 92–110 μm long, 52–56 μm diameter (x̄ = 101 × 54 μm, n = 5), centrally erumpent, with periphyses, surrounded by distinct shiny black flanges, the tissue spreading down along the papilla. Peridium 51–60 μm wide, comprising 4–5 cell layers of thin-walled, brown cells of textura angularis, inwardly lighter. Paraphyses filamentous, distinctly septate, embedded in a hyaline gelatinous matrix. Asci 147–189 × 10–13 μm (x̄ = 167 × 11 μm, n = 30), 8-spored, unitunicate, cylindrical, short pedicellate, slightly truncate at the apex, with a wedge-shaped J+, subapical ring, 3.5–4 µm broad, 2–2.5 µm high. Ascospores 20–24 × 6–8 μm (x̄ = 22 × 7 μm, n = 40), overlapping, uniseriate, ellipsoidal, slightly tapering at the ends, equally 2-celled and guttulate at both cells, constricted at the septum, brown with striations, surrounded by a thick, hyaline mucilaginous sheath, subglobose, parallel to the margin of the spore. Asexual morph: Undetermined.

Material examined

China, Taiwan, Chiayi Province, Ali Mountain, Kwang Hwa, on dead sheaths of Miscanthus sinensis (Poaceae), 5 May 2018, A. Karunarathna, AKTW 44 (MFLU 19-2333, holotype)

Additional material

China, Yunnan Province, Kunming City, Kunming Botanical Garden (25°8'45"N, 102°44'59"E), on dead culm of monocotyledon, 20 March 2014, Q.R. Li, GZ43 (GZUH0122, GACP QR0201; living cultures, GZUCC0122).

Note

The characteristics of the holotype specimen Arecophila miscanthi (≡ Alishanica miscanthi) were revised, re-measured and described. Alishanica miscanthi is similar to A. muroiana and A. serrulata (Wang et al. 2004, Hyde et al. 2020). However, no clypeus was observed for A. muroiana. Arecophila serrulata has larger ascomata (480–560 × 280–320 μm) with a central papilla surrounded by a circle of white tissue (Hyde 1996) which differs from those of A. miscanthi. One new collection (GZUH0122, Fig. 5) shows the same traits of A. miscanthi (MFLU 19-2333) in having immersed ascomata with clypeus, a wedge-shaped J+, ascus subapical ring, same dimensions of ascospores and here we provide it as a new geographical record from China.

Figure 5. 

Arecophila miscanthi (GZUH0122) A, B ascomata on the surface of host C paraphyses and asci D section of ascoma E peridium F, G apical rings H–K asci with ascospores M–P ascospores. Scale bars: 50 μm (C, D); 10 μm (E, F–K); 5 μm (L–P).

Discussion

Arecophila shares similar morphology to Atrotorquata, Cainia and Seynesia in having immersed ascomata and 2-celled ascospores (Hyde 1996). Arecophila, Atrotorquata, Cainia and Seynesia are accepted in Cainiaceae with newly-introduced genera, such as Amphibambusa and Longiappendispora (Mapook et al. 2020). Cainia has similar characteristics to Arecophila in its occurrence on monocotyledons, having asci with J+, apical rings and brown 2-celled ascospores (Kohlmeyer and Volkmann-Kohlmeyer 1993). The ascospores of Cainia are provided with several longitudinal germ slits and differ from those of Arecophila, where the ascospores are provided with ridges or a verrucose wall and lack germ slits. Seynesia produces ascospores that are smooth-walled and surrounded by mucilaginous sheaths that are drawn out at the poles with germ slits, which differ from Arecophila (Hyde 1995). Amphibambusa possesses hyaline ascospores pointed at both ends, which differs from that of Arecophila (Liu et al. 2015). The phylogenetic tree (Fig. 1) displays that Arecophila miscanthi (≡ Alishanica miscanthi) clusters in the Arecophila group with high support values (100%/1.00 PP). Longiappendispora possesses ascospores with longitudinal striations and bristle-like polar appendages at both ends, without a gelatinous sheath, which differentiates it from other genera in Cainiaceae. Ascospores of Atrotorquata are provided with several longitudinal germ slits and differ from those of Arecophila (Kohlmeyer and Volkmann-Kohlmeyer 1993). At present, 16 Arecophila species have been described and a summary of each species are given in the Table 2.

Table 2.
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Synopsis of the species of Arecophila.

Species Host Clypeus Ascomata Asci Ascal ring Ascospores Distribution
A. australis Phragmites australis Present 420–560 × 290–380 µm, globose to subglobose 140–230 × 15.5–24 µm 4–5 × 2.5–3 μm, trapezoidal, J+ 22.5–29 × 8–11 µm, wall striate, mucilaginous sheath China (Guizhou)
A. bambusae Bambusa sp. Absent 500–560 × 294–350 µm, globose to subglobose 132.5–140 × 7.5–8 µm 2.5–3 µm in diam., ca. 2.5 µm high, wedge-shaped, J+ 19–22.5 × 5.5–7 µm, slightly tapering at the ends, wall striate, mucilaginous sheath Hong Kong
A. calamicola Calamus sp. Absent 520 × 390 µm, subglobose 160–190 × 14–20 µm 4-4.8 µm diam., 3.2-4 µm high, wedge-shaped, J+ 24–33 × 5.5–9 µm, wall striate, verrucose, mucilaginous sheath Brunel, Indonesia
A. chamaeropis Chamaerops humilis Minute 400–700 × 300–400 µm, subglobose 150–190 × 9–10 µm 3.5–4.5 diam., 1.5–2 µm high, wedge-shaped, J+ 15–23 × 5.5–7 µm, wall striate, covered by pronounced verrucose ornamentation, mucilaginous sheath Spain
A. coronata Gigantochloa scribneriana, Bambusa sp. Present 90–100 × 42–105 µm, subglobose or ellipsoidal 132.5–157.5 × 7.5–9 µm 3.5–4 µm in diam., 2–2.5 µm high, wedge-shaped, J+, with a faint canal leading to the apex. 29–31 × 5–5.5 µm, wall faint striate, mucilaginous sheath Philippines, Hong Kong
A. clypeata A unknown gramineous plant Present 367‒448 × 278‒363 µm, subglobose to globose 180–245 × 10.5–14.5 µm 3–4 × 3–4 μm, square-shaped, J+ 18.5–22.5 × 6.5–9 µm, wall striate, mucilaginous sheath China (Guizhou)
A. deutziae Deutziae stamineae Absent 400–600 µm diam., globose 180–240 × 16–19 µm 3.5–4. 5 µm diam., 1.5–2 µm high, wedge-shaped, J+ 26–32 × 11–13 µm, wall striate India
A. eugeissonae Eugeissona tristis Absent 460–520 × 180–260 µm, Subglobose or ellipsoidal 175–220 × 11–16.5 µm 3–4 µm diam., 1.5–2.0 µm high, discoid, J+ 25-40 × 6.5–9 µm, wall weakly striate, verrucose, mucilaginous sheath Malaysia
A. foveata Nolinae sp. Present 300–400 ×400–500 µm, globose or ovoid 130–150 × 14–15 µm 3–4 µm wide, 4–5 µm high, tubular, J+ 16–20 × 8–10 µm, wall striate, foveate, surface aspect of numerous warts USA
A. gulubiicola Gulubia costate Present 290–400 × 140–190 μm, subglobose or lenticular 107–145 × 11–13.5 μm 3.2–4 µm diam., 2.4–3.2 µm high, cylindrical, J+ 14.5–18.5 × 6–9 μm with a minutely verrucose wall, mucilaginous sheath Papua New Guinea
A. miscanthi Miscanthus sinensis Present 283–296 × 272–277 µm, globose to subglobose 147–189 × 10–13 μm 3.5–4 µm broad, 2–2.5 µm high, wedge-shaped, J+ 20–24 × 6–8 μm, wall striate, mucilaginous sheath. China (Taiwan, Yunnan)
A. muroiana Phyllostachys bambusoides Absent 350–460 × 320–400 µm, globose 125–165 × 10–12 µm 3.5–4 µm diam., 2–2.5 µm high, wedge-shaped, J+ 20–25 × 6–7.5 µm, wall finely striate, mucilaginous sheath Japan
A. notabilis Calamus, Bamboo Present 400 × 360 µm, subglobose 180–220 × 11–14 µm 4–4.45 µm diam., 3–4.5 µm high, wedge-shaped, J+ 20–26 × 6–8 µm, wall striate, finely verrucose, mucilaginous sheath Brunei, Hong Kong, Indonesia
A. nypae Nypa fruticans Absent 400–500 µm diam., subglobose 140–205 × 11–13 µm 4.5 µm diam., 2.5–4 µm high, wedge-shaped, J+ 19–26 × 7–8 µm, wall striate, mucilaginous sheath Malaysia
A. saccharicola Sacchari officinarum Absent 420–525 × 350–420 µm high 140–16 × 7–10 µm Not blued by Melzer’s reagent 20–24 × 6–8 µm, wall smooth or striated Jamaica
A. serrulata Korthalsia sp., Sabal sp., Serenoa sp. Present 480–560 × 280–320 µm, conical with flattened base 110–112 × 10–12 µm, 3.2 µm diam., 2.4 µm high, wedge-shaped, J+ 17–26 × 7–9.5 µm, wall striate, mucilaginous sheath Brunei, USA, Florida

The combined ITS, LSU, rpb2 and β-tubulin phylogeny (Fig. 1) showed two clades of Arecophila as Clade 1 and Clade 2. The Arecophila differs from Amphibambusa and Cainia (see above). The sequence from the holotype of Atrotorquata spartii is noticeably clustered with Coniocessia spp. in Coniocessiaceae (Fig. 1). However, Atrotorquata spartii showed a close affinity with Cainia spp. in Cainiaceae, based on analysis of the combined LSU and ITS sequence alignment in Senanayake et al. (2015). Atrotorquata has similar characteristics to Arecophila and other genera of Cainiaceae (Hyde 1996). Hence, there should be more evidence to reassess Atrotorquata in the future. The unitunicate asci with a J+ apical ring in Melzer’s regent and brown ascospores covered with longitudinal wall striations, without germ slits can clearly distinguish Arecophila from its similar genera. In addition, a table including synopsis of the species of Arecophila is provided.

Acknowledgements

We would like to thank the curator of BRIP for the loan of fungal material. This research was supported by the National Natural Science Foundation of China (32000009 and 31960005); the Open Fund Program of Engineering Research Center of Southwest Bio-Pharmaceutical Resources, Ministry of Education, Guizhou University No. GZUKEY20160; the Fund of the Science and Technology Foundation of Guizhou Province ([2020]1Y059); Guizhou Provincial Academician Workstation of Microbiology and Health (No. [2020]4004); International Science and Technology Cooperation Base of Guizhou Province ([2020]4101); the Fund of High-Level Innovation Talents [No. 2015-4029], the Base of International Scientific and Technological Cooperation of Guizhou Province [No. [2017]5802].

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