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Research Article
Taxonomic novelties and global biogeography of Montagnula (Ascomycota, Didymosphaeriaceae)
expand article infoDhanushka N. Wanasinghe, Thilina S. Nimalrathna§|#, Li Qin Xian, Turki Kh. Faraj¤, Jianchu Xu«, Peter E. Mortimer
‡ Kunming Institute of Botany, Chinese Academy of Sciences, Honghe County, China
§ University of Chinese Academy of Sciences, Beijing, China
| CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, China
¶ Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences & Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, China
# Yunnan International Joint Laboratory of Southeast Asia Biodiversity Conservation & Yunnan Key Laboratory for Conservation of Tropical Rainforests and Asian Elephants, Mengla, China
¤ King Saud University, Riyadh, Saudi Arabia
« CIFOR-ICRAF China Country Program, Kunming, China
Open Access

Abstract

Whilst conducting surveys of lignicolous microfungi in Yunnan Province, we collected a large number of taxa that resemble Montagnula (Didymosphaeriaceae, Pleosporales). Our phylogenetic study on Montagnula involved analysing sequence data from ribosomal RNA genes (nc18S, nc28S, ITS) and protein-coding genes (rpb2, tef1-α). We present a biphasic approach (morphological and molecular phylogenetic evidence) that supports the recognition of four new species in Montagnula viz., M. lijiangensis, M. menglaensis, M. shangrilana and M. thevetiae. The global diversity of Montagnula is also inferred from metabarcoding data and published records based on field observations. Metabarcoding data from GlobalFungi and field observations provided insights into the global diversity and distribution patterns of Montagnula. Studies conducted in Asia, Australia, Europe, and North America revealed a concentration of Montagnula species, suggesting regional variations in ecological preferences and distribution. Montagnula species were found on various substrates, with sediments yielding a high number of sequences. Poaceae emerged as a significant contributor, indicating a potential association between Montagnula species and grasses. Culture-based investigations from previously published data revealed Montagnula species associations with 105 plant genera (in 45 plant families), across 55 countries, highlighting their wide ecological range and adaptability. This study enhances our understanding of the taxonomy, distribution, and ecological preferences of Montagnula species. It emphasizes their role in the decomposition of organic matter in grasslands and savannah systems and suggests further investigation into their functional roles in ecosystem processes. The global distribution patterns and ecological interactions of Montagnula species underscore the need for continued research and conservation efforts.

Key words

Global distribution, microfungi, molecular phylogeny, taxonomy, Yunnan

Introduction

Fungi are the second largest group of eukaryotes, performing vital ecological functions such as decomposition, mutualism, and pathogenesis to plants and animals (Tedersoo et al. 2014). Ascomycota, which forms the largest phylum of Fungi, and includes the genus Montagnula, is an incredibly diverse group, with an estimated global species richness of ~154,500 species (Bánki et al. 2023). Despite their ecological and economic importance, many Ascomycota species remain undescribed, and their distribution and diversity have yet to be properly determined (Maharachchikumbura et al. 2021a, b; Wijayawardene et al. 2022). This is somewhat due to the fact that many Ascomycota species are microscopic and inconspicuous, making them difficult to find and subsequently study, or sometimes these smaller species can be overlooked with studies focussing on more charismatic species of macrofungi (Wanasinghe et al. 2022a). The investigation of taxonomic and phylogenetic systematics in Ascomycota is bridging crucial knowledge gaps and enhancing our understanding of this particular group of fungi. Montagnula (typified with M. infernalis), is an example of a relatively understudied genus within Ascomycota, and many species remain undescribed. Understanding the taxonomic, phylogenetic and host relationships between Montagnula species will help us better understand how they have diversified and adapted to different habitats in various ecological zones. These data are useful to make predictions about the ecology and biology of the genus and to guide future research into their interactions with other organisms and their roles in ecosystem processes. Understanding the taxonomy and phylogeny of Montagnula is also important for conservation purposes. With ongoing habitat destruction and climate change, it is more important than ever to understand the current diversity and distribution of fungi around the world (Wanasinghe et al. 2022a).

Therefore, our research group at the Center for Mountain Futures (CMF), has been conducting investigations into the microfungal diversity and biogeography in Yunnan Province, Southwest China. Specifically, we are focusing on various substrates such as leaf and woody litter, aiming to clarify the taxonomy of fungi on these substrates, using morphology in conjunction with multigene phylogeny. As a result, we have successfully isolated numerous anamorphic and teleomorphic Ascomycota species in Yunnan, and we have published our findings based on different themes, including their relationship with hosts, substrates, and localities (Thiyagaraja et al. 2019, 2020, 2021; Abeywickrama et al. 2020; Wanasinghe et al. 2020, 2021, 2022b, 2023; Yasanthika et al. 2020; Bundhun et al. 2021; Dissanayake et al. 2021; Gao et al. 2021; Monkai et al. 2021; Mortimer et al. 2021; Ren et al. 2021a, b, 2022a, b; Aluthmuhandiram et al. 2022; Maharachchikumbura et al. 2022; Wanasinghe and Mortimer 2022). The objectives of this study are (1) to identify the lignicolous Montagnula species collected from Yunnan using both morphological and phylogenetic approaches, and (2) to utilize metabarcoding data and published records based on field observations to infer the global diversity and biogeography of Montagnula. The analyses conducted in this study revealed four new species and four existing species of Montagnula, in Yunnan. The discovery of several previously undescribed Ascomycota species in the genus Montagnula in Yunnan Province is a significant advancement in our understanding of the diversity and distribution of this group of fungi. Furthermore, the utilization of metabarcoding data and published records based on field observations to infer the global diversity of Montagnula demonstrates the potential of these approaches in elucidating the biogeography of fungi on a large scale. By studying and documenting the diversity of Montagnula species, we can enhance our appreciation for the importance of conserving these fungi and their habitats, and take appropriate measures to mitigate the threats they face.

Materials and methods

Sample collecting

Fresh fungal materials were collected from dead woody twigs from Honghe, Kunming, Mengla, Shangri-La and Yulong Counties, all within Yunnan Province, China, during the dry season (January, March, April) and wet season (August, September). To preserve their integrity, the specimens were transported to the laboratory in Zip lock plastic bags during the dry season and in paper bags during the wet season.

Morphological observations

The morphology of external and internal macro-/micro-structures were observed as described in Wanasinghe et al. (2017, 2018a, 2020). Hand sections of the ascomata were mounted in distilled water and the following characteristics were evaluated and measured: ascomata diameter, height, color and shape; width of peridium; and height and diameter of ostioles. Length and width (at the widest point) of asci and ascospores. Images were captured with a Canon EOS 600D digital camera fitted to a Nikon ECLIPSE Ni compound microscope. Macroscopic images of colonies were documented using an iPhone XS Max (Apple Inc., Cupertino, CA, USA) with daylight. Measurements were made with the Tarosoft (R) Image Frame Work program, and images used for figures were processed with Adobe Photoshop CS5 Extended version 10.0 software (Adobe Systems, San José, CA, USA).

Isolation

Single spore isolation was conducted by following the methods described in Wanasinghe et al. (2018b). Germinated spores were individually transferred to potato dextrose agar (PDA: 39 g/L distilled water, Difco potato dextrose) plates and grown at 20 °C in the daylight.

Deposition of specimens, cultures and registering names

The living cultures were deposited at the Kunming Institute of Botany Culture Collection (KUNCC), Kunming, China. Dry herbarium materials were deposited in the herbarium of Cryptogams Kunming Institute of Botany, Academia Sinica (KUN-HKAS). MycoBank numbers have been obtained as outlined in MycoBank (http://www.MycoBank.org accessed on 21 September 2023) for the novel taxa.

DNA extraction, PCR amplifications and sequencing

Genomic DNA was extracted from the axenic mycelium as described by Phookamsak et al. (2017). Mycelia for DNA extraction from each isolate were grown on PDA for 3–4 weeks at 20 °C and total genomic DNA was extracted from approximately 150 ± 50 mg axenic mycelium scraped from the edges of the growing culture. Mycelium was ground to a fine powder with liquid nitrogen and DNA extracted using the Biospin Fungus Genomic DNA Extraction Kit-BSC14S1 (BioFlux, P.R. China) following the instructions of the manufacturer. When fungi failed to grow in culture, DNA extraction was carried out directly from fruiting bodies, adhering to the protocol outlined by Wanasinghe et al. (2018b). DNA to be used as templates for Polymerase Chain Reaction (PCR) were stored at 4 °C for use in regular work and duplicated at -20 °C for long-term storage.

We used primers ITS5/ITS4 (White et al. 1990), LR0R/LR5 (Vilgalys and Hester 1990; Rehner and Samuels 1994), NS1/NS4 (White et al. 1990), EF1-983F/EF1-2218R (Liu et al. 1999; Rehner and Buckley 2005), and fRPB2-5f/fRPB2-7cR (Sung et al. 2007) to amplify sequence data for a total of five markers: the internal transcribed spacers (ITS), partial 28S large subunit rDNA (LSU), partial 18S small subunit rDNA (SSU), translation elongation factor 1-α (tef1-α), and RNA polymerase II second largest subunit (rpb2). PCR amplifications were performed following the methods described in Wanasinghe et al. (2021). We sequenced complementary strands with the same primers used for PCR amplifications and sequencing was done from a commercial sequencing provider (BGI, Ltd Shenzhen, P.R. China). The nucleotide sequence data obtained were deposited in GenBank (Table 2).

Sequencing assembly and alignments

Sequences generated from different primers of the five genes were analysed with other sequences retrieved from GenBank (Table 2). Sequences with high similarity indices were determined from a BLAST search to find the closest matches with taxa in Didymosphaeriaceae, using recently published data (Du et al. 2021; Ren et al. 2022a; Sun et al. 2023). The multiple alignments of all consensus sequences, as well as the reference sequences were automatically generated with MAFFT v. 7 (Katoh et al. 2019), and manually corrected where necessary using BioEdit v. 7.0.5.2 (Hall 1999).

Phylogenetic inference

The single-locus datasets were examined for topological incongruence among loci for members of the analyses. The alignments were concatenated into a multi-locus alignment that was analyzed with maximum likelihood (ML) and Bayesian (BI) phylogenetic methods in the CIPRES Science Gateway (Miller et al. 2010). ML tree was obtained using RAxML-HPC2 on XSEDE v. 8.2.10 (Stamatakis 2014) with applying GTR+G+I model. Support values were obtained with 1,000 bp replicates (Felsenstein 1985). ML bootstrap values equal or greater than 75% are given above each node. The best-fit model was selected with respect to Bayesian Information Criterion (BIC) scores using the IQ-TREE web application at http://iqtree.cibiv.univie.ac.at (Trifinopoulos et al. 2016). For model selection, we restricted the pool of available models to JC, F81, HKY, SYM and GTR (Ronquist et al. 2011). BI were performed with two parallel runs of 2 M generations, using four chains in each, and retaining one tree every 100 generations. The dataset was partitioned by gene region, and a GTR + G + I model was applied to each partition, ending the run automatically when standard deviation of split frequencies dropped below 0.01 with a burn-in fraction of 0.25. A fifty percent majority rule consensus tree was obtained after discarding the first 25% of trees, and posterior probabilities were used as a measure of nodal support. The posterior probability in BI (BYPP) greater than 0.95 are given above each node. Phylograms were visualized with FigTree v1.4.0 program (Rambaut 2012) and reorganized in Microsoft power point (2019).

The biogeographical distribution of Montagnula

In our initial approach, we obtained detailed geographical distribution information for the Montagnula genus. This data was extracted from the GlobalFungi database (https://globalfungi.com, accessed on 04 December 2023), as outlined by Větrovský et al. (2020). The database provided information on the countries and precise geographical coordinates of recorded Montagnula occurrences. To visualize these occurrences, we employed a range of packages in R version 4.2.1 (R Core Team 2022), including ‘sf’ (Pebesma and Bivand 2023), ‘raster’ (Hijmans 2023), ‘rgdal’ (Bivand et al. 2022), and ‘ggplot2’ (Wickham 2011). In our map, each marker signifies an individual occurrence of Montagnula. These occurrences are visually distinguished by a color scheme, with each color denoting the specific biome from which the samples were collected, as illustrated in Fig. 2a. Additionally, we have developed two donut charts, showcased in Fig. 2b, c, which effectively illustrate the distribution of Montagnula sequences. These charts present the sequence abundance as a percentage of the total, segmented across various biomes and continents, providing a clear visual breakdown of their distribution. Furthermore, we have gathered Environmental DNA (eDNA) data from diverse sources in metabarcoding studies focusing on fungi, as found in the GlobalFungi database (Fig. 3). This dataset included specifics about eDNA sources, locations of the studies, and the sequence abundance of Montagnula sequences. It is important to note that the sequence abundance in metabarcoding studies might not always accurately represent the actual abundance of species in a habitat. Nonetheless, these data can provide valuable insights into the potential rarity or prevalence of the group in the eDNA source. We analyzed the sequence abundance in diverse eDNA samples from different continents. Before visualization, the abundance values were normalized via a logarithmic transformation to ensure a standardized and comparable presentation of Montagnula sequence abundance. Post-transformation abundance data were visualized using the ‘ggplot2’ package, aiding in highlighting the focus areas of metabarcoding and identifying the environmental sample types from which Montagnula sequences were derived across various continents (Figs 2, 3).

The host relations of Montagnula

To illustrate the host specificity of Montagnula species, we utilized detailed information regarding host species from the literature (Table 1). This enabled us to create informative bar plots displaying the host preferences of Montagnula species (Fig. 4). This information was visualized using the ‘ggplot2’ package in R.

Table 1.

Accepted species in Montagnula including their host and geographic location.

Species Host species Host family Country Reference
Montagnula acaciae Acacia auriculiformis Fabaceae Thailand Tennakoon et al. (2022) #
Montagnula aloes Aloe sp. Asphodelaceae South Africa Crous et al. (2012) #
Montagnula appendiculata Zea mays Poaceae China Aptroot (2004) #
Montagnula aquatica Submerged wood NA Thailand Sun et al. (2023) #
Dead woody litter NA China This study#
Montagnula aquilariae Aquilaria sinensis Thymelaeaceae China Hyde et al. (2023) #
Dead woody litter NA China This study#
Montagnula baatanensis Agave sp. Asparagaceae USA Crivelli (1983)
Montagnula bellevaliae Bellevalia romana Asparagaceae Italy Hongsanan et al. (2015) #
Montagnula camporesii Dipsacus sp. Caprifoliaceae Italy Hyde et al. (2020) #
Montagnula camarae Cytisus scoparius Fabaceae Portugal Checa (2004)
Montagnula chiangraiensis Chromolaena odorata Asteraceae Thailand Mapook et al. (2020) #
Montagnula chromolaenae Chromolaena odorata Asteraceae Thailand Mapook et al. (2020) #
Montagnula chromolaenicola Chromolaena odorata Asteraceae Thailand Mapook et al. (2020) #
Lagerstroemia sp. Lythraceae China This study#
Montagnula cirsii Cirsium sp. Asteraceae Italy Hyde et al. (2016) #
Montagnula cylindrospora Human skin## NA USA Crous et al. (2020) #
Montagnula dasylirionis Dasylirion sp. Asparagaceae USA Ramaley and Barr (1995)
Montagnula donacina Acacia reficiens Fabaceae Namibia Aptroot (1995)
Acacia sp. Fabaceae India Aptroot (1995)
Adhatoda vasica Acanthaceae India Aptroot (1995)
Ailanthus altissima Simaroubaceae India Aptroot (1995)
Althaea rosea Malvaceae China Aptroot (1995)
Annona squamosa Annonaceae India Aptroot (1995)
Arundo donax Poaceae Portugal Aptroot (1995)
Bambusoideae Poaceae Brazil Aptroot (1995)
Bambusoideae Poaceae Papua New Guinea Aptroot (1995)
Cajanus cajan Fabaceae India Aptroot (1995)
Calamus australis Arecaceae Australia Hyde et al. (1999)
Careya arborea Lecythidaceae India Aptroot (1995)
Citrus aurantiifolia Rutaceae India Aptroot (1995)
Clerodendrum infortunatum Lamiaceae India Aptroot (1995)
Clerodendrum multiflorum Lamiaceae India Aptroot (1995)
Coffea arabica Rubiaceae Paraguay Aptroot (1995)
Coffea robusta Rubiaceae Central African Republic Aptroot (1995)
Craterellus odoratus ## Cantharellaceae China Zhao et al. (2018) #
Duranta repens Verbenaceae India Aptroot (1995)
Ficus glomerata Moraceae India Aptroot (1995)
Funtumia africana Apocynaceae Sierra Leone Aptroot (1995)
Hibiscus sp. Malvaceae India Aptroot (1995)
Ipomoea carnea Convolvulaceae India Aptroot (1995)
Mallotus philippinensis Euphorbiaceae India Aptroot (1995)
Morus alba Moraceae India Aptroot (1995)
Litchi litchi Sapindaceae Myanmar Thaung (2008)
Montagnula donacina Nerium odorum Apocynaceae India Aptroot (1995)
Paeonia suffruticosa Paeoniaceae China Li et al. (2023) #
Phyllostachys bambusoides Poaceae Japan Wang et al. (2004)
Pistacia sp. Anacardiaceae India Aptroot (1995)
Platanus sp. Platanaceae USA Wang et al. (2004)
Premna cumingiana Lamiaceae Philippines Aptroot (1995)
Pseudosasa japonica Poaceae France Aptroot (1995)
Saccharum officinarum Poaceae Brazil Aptroot (1995)
Unknown stem NA India Aptroot (1995)
Tectona grandis Lamiaceae India Aptroot (1995)
Terminalia tomentosa Combretaceae India Aptroot (1995)
Trachycarpus fortunei Arecaceae China Hyde et al. (1999)
Unknown bark NA India Aptroot (1995)
Unknown branches NA Sierra Leone Aptroot (1995)
Unknown plant NA Colombia Aptroot (1995)
Dead wood NA China Sun et al. (2023) #
Dead wood NA Thailand Ren et al. (2022a) #
Dead wood NA China This study#
Vitis vinifera Vitaceae Australia Pitt et al. (2014) #
Wikstroemia sp. Thymelaeaceae USA Aptroot (1995)
Zea mays Poaceae Georgia Aptroot (1995)
Montagnula dura Aconitum septentrionale Ranunculaceae Sweden Eriksson (1992)
Lonicera etrusca Caprifoliaceae Spain Checa (2004)
Montagnula gilletiana Fraxinus ornus Oleaceae Bulgaria Fakirova (2004)
Retama sphaerocarpa Fabaceae Spain Checa (2004)
Ulex europaeus Fabaceae Spain Checa (2004)
Montagnula graminicola Poaceae Poaceae Italy Liu et al. (2015) #
Montagnula guiyangensis Helwingia himalaica Helwingiaceae China Sun et al. (2023) #
Montagnula hirtula Cerastium latifolium Caryophyllaceae Austria Leuchtmann (1984)
Cerastium sp. Caryophyllaceae Italy Leuchtmann (1984)
Epilobium parviflorum Onagraceae Switzerland Leuchtmann (1984)
Rubus idaeus Rosaceae Finland Leuchtmann (1984)
Rubus sp. Rosaceae Sweden Eriksson (1992)
Montagnula infernalis Agave americana Asparagaceae Portugal Checa (2004)
Agave americana Asparagaceae Spain Checa (2004)
Fourcroya sp. Asparagaceae Portugal Ariyawansa et al. (2014)
Furcraea gigantea Asparagaceae Portugal Checa (2004)
Furcraea gigantea Asparagaceae Spain Checa (2004)
Furcraea longaeva Asparagaceae Portugal Checa (2004)
Furcraea longaeva Asparagaceae Spain Checa (2004)
Montagnula infernalis Furcraea macrophylla Asparagaceae Bahamas Barr (1990)
Montagnula jonesii Fagus sylvatica Fagaceae Italy Tennakoon et al. (2016) #
Ficus benjamina Moraceae Thailand Tennakoon et al. (2022) #
Montagnula krabiensis Pandanus sp. Pandanaceae Thailand Tibpromma et al. (2018) #
Montagnula lijiangensis Quercus sp. Fagaceae China This study#
Montagnula longipes Agave americana Asparagaceae Algeria Aptroot (1995)
Montagnula melanorhabdos Agave sp. Asparagaceae Turkey Aptroot (2006)
Montagnula menglaensis Indocalamus tessellatus Poaceae China This study#
Montagnula mohavensis Yucca mohavensis Asparagaceae USA Ramaley and Barr (1995)
Montagnula obtusa Ilex sp. Aquifoliaceae USA French (1989)
Juglans sp. Juglandaceae USA French (1989)
Pinus pinaster Pinaceae Portugal Checa (2004)
Sorbus aucuparia Rosaceae Sweden Eriksson (1992)
Montagnula opaca Phalaris Poaceae Switzerland Crivelli (1983)
Montagnula opulenta Ammophila arenaria Poaceae France Aptroot (1995)
Ammophila arenaria Poaceae Germany Aptroot (1995)
Ammophila arenaria Poaceae Sweden Aptroot (1995)
Festuca brachyphylla Poaceae Canada Aptroot (1995)
Opuntia ficus-indica Cactaceae Canary Islands Aptroot (1995)
Opuntia ficus-indica Cactaceae France Aptroot (1995)
Opuntia ficus-indica Cactaceae Italy Aptroot (1995)
Opuntia ficus-indica Cactaceae Malta Aptroot (1995)
Opuntia ficus-indica Cactaceae Tunisia Aptroot (1995)
Opuntia sp. Cactaceae Cyprus Aptroot (1995)
Opuntia sp. Cactaceae Israel Aptroot (1995)
Opuntia sp. Cactaceae Italy Aptroot (1995)
Opuntia sp. Cactaceae Tunisia Aptroot (1995)
Opuntia tuna Cactaceae USA Aptroot (1995)
Poa abbreviata Poaceae Canada Aptroot (1995)
Puccinellia angustata Poaceae Greenland Aptroot (1995)
Stipa himalaica Poaceae India Aptroot (1995)
Montagnula opuntiae Opuntia lindheimeri Cactaceae USA Huhndorf (1992)
Montagnula palmacea Chamaerops humilis Arecaceae France Aptroot (1995)
Cocos capitata Arecaceae Spain Aptroot (1995)
Daviesia nudiflora Fabaceae Australia Aptroot (1995)
Phoenix dactylifera Arecaceae Egypt Aptroot (1995)
Phoenix dactylifera Arecaceae Greece Aptroot (1995)
Phoenix dactylifera Arecaceae Iraq Aptroot (1995)
Phoenix dactylifera Arecaceae Italy Aptroot (1995)
Phoenix dactylifera Arecaceae Pakistan Aptroot (1995)
Phoenix dactylifera Arecaceae Saudi Arabia Aptroot (1995)
Phoenix dactylifera Arecaceae Tunisia Aptroot (1995)
Phoenix sylvestris Arecaceae Pakistan Aptroot (1995)
Pitcairnia chrysantha Bromeliaceae Chile Aptroot (1995)
Unknown leaves NA USA Aptroot (1995)
Unknown petiole NA USA Aptroot (1995)
Montagnula perforans Calamagrostis arenaria Poaceae France Aptroot (2006)
Montagnula phragmospora Agave americana Asparagaceae Portugal Checa (2004)
Agave americana Asparagaceae Spain Checa (2004)
Agave hookeri Asparagaceae Portugal Checa (2004)
Agave hookeri Asparagaceae Spain Checa (2004)
Agave sp. Asparagaceae France Barr (1990)
Agave sp. Asparagaceae Portugal Checa (2004)
Agave sp. Asparagaceae Spain Checa (2004)
Montagnula phragmospora Yucca brevifolia Asparagaceae California Barr (1990)
Yucca sp. Asparagaceae Portugal Checa (2004)
Yucca sp. Asparagaceae Spain Checa (2004)
Montagnula puerensis Dead wood NA China Du et al. (2021) #
Montagnula rhodophaea Arundo donax Poaceae Italy Leuchtmann (1984)
Phragmites communis Poaceae Switzerland Leuchtmann (1984)
Montagnula saikhuensis Citrus sp. Rutaceae Thailand Wanasinghe et al. (2016) #
Montagnula scabiosae Scabiosa sp. Caprifoliaceae Italy Hongsanan et al. (2015) #
Montagnula shangrilana Rhododendron sp. Ericaceae China This study#
Montagnula sp. Carex fuliginosa Cyperaceae Austria Scheuer (1988)
Montagnula spartii Aeluropus littoralis Poaceae Russia Aptroot (1995)
Ammophila arenaria Poaceae Belgium Aptroot (1995)
Ammophila arenaria Poaceae Denmark Aptroot (1995)
Ammophila arenaria Poaceae Sweden Aptroot (1995)
Ammophila arenaria Poaceae United Kingdom Aptroot (1995)
Calamagrostis epigeios Poaceae Russia Aptroot (1995)
Calycotome spinosa Fabaceae France Aptroot (1995)
Calycotome spinosa Fabaceae Spain Aptroot (1995)
Calycotome villosa Fabaceae Italy Aptroot (1995)
Carex rostrata Cyperaceae Sweden Aptroot (1995)
Chamaerops humilis Arecaceae Spain Aptroot (1995)
Leymus arenarius Poaceae Russia Aptroot (1995)
Ephedra ciliata Ephedraceae Unknown country in Asia Aptroot (1995)
Ephedra sp. Ephedraceae Iran Aptroot (1995)
Festuca arenaria Poaceae France Aptroot (1995)
Festuca sulcata Poaceae Iran Aptroot (1995)
Genista aspalathoides Fabaceae Italy Aptroot (1995)
Gramineae Gramineae Austria Aptroot (1995)
Koeleria cristata Poaceae Germany Aptroot (1995)
Koeleria glauca Poaceae Denmark Aptroot (1995)
Linum austriacum Linaceae Germany Aptroot (1995)
Luzula spadicea Juncaceae Switzerland Aptroot (1995)
Lygeum spartum Poaceae Spain Aptroot (1995)
Melica ciliata Poaceae France Aptroot (1995)
Nardus stricta Poaceae Austria Aptroot (1995)
Puccinellia peisonis Poaceae Austria Aptroot (1995)
Sarothamnus scoparius Fabaceae Poland Mulenko et al. (2008)
Sarothamnus scoparius Fabaceae Switzerland Aptroot (1995)
Sesleria caerulea Poaceae Italy Aptroot (1995)
Montagnula spartii Spartium junceum Fabaceae Albania Aptroot (1995)
Spartium junceum Fabaceae France Aptroot (1995)
Spartium junceum Fabaceae Greece Aptroot (1995)
Spartium junceum Fabaceae Turkey Aptroot (1995)
Ulex sp. Fabaceae Spain Aptroot (1995)
Montagnula spinosella Abelia triflora Caprifoliaceae Spain Checa (2004)
Carex aterrima Cyperaceae Austria Scheuer (1988)
Montagnula spinosella Carex misandra Cyperaceae Norway Holm and Holm (1993, 1994)
Colpodium vahlianum Poaceae Norway Holm and Holm (1993, 1994)
Deschampsia caespitosa Poaceae Norway Holm and Holm (1993, 1994)
Juncus maritimus Juncaceae Spain Holm and Holm (1993), Checa (2004)
Luzula confusa Juncaceae Norway Holm and Holm (1993, 1994)
Montagnula stromatosa Phoenix hanceana Arecaceae China Lu et al. (2000)
Phoenix sp. Arecaceae China Zhuang (2001)
Trachycarpus fortunei Arecaceae China Hyde et al. (1999)
Trachycarpus fortunei Arecaceae United Kingdom Hyde et al. (1999)
Montagnula subsuperficialis Panicum grumosum Poaceae Argentina Shoemaker (1989)
Montagnula thailandica Chromolaena odorata Asteraceae Thailand Mapook et al. (2020) #
Hevea brasiliensis Euphorbiaceae Thailand Senwanna et al. (2021) #
Coffea arabica var. catimor Rubiaceae China Lu et al. (2022) #
Unidentified twig NA Thailand Boonmee et al. (2021) #
Montagnula thevetiae Thevetia peruviana Apocynaceae China This study#
Montagnula thuemeniana Yucca sp. Asparagaceae USA Barr (1990)
Montagnula triseti Trisetum distichophyllum Poaceae Switzerland Crivelli (1983)
Montagnula vakrabeejae Unidentified twig NA Andaman Niranjan and Sarma (2018)
Montagnula verniciae Vernicia fordii Euphorbiaceae China Li et al. (2023) #
Montagnula yuccigena Yucca baccata Asparagaceae Mexico Ramaley and Barr (1995)
Table 2.

GenBank accession numbers of sequences used for the phylogenetic analyses.

Taxon Strain number GenBank accession numbers Reference
ITS LSU SSU tef1-α rpb2
Montagnula acaciae MFLUCC 18-1636 ON117280 ON117298 ON117267 ON158093 NA Tennakoon et al. (2022)
NCYUCC 19-0087T ON117281 ON117299 ON117268 ON158094 NA Tennakoon et al. (2022)
Montagnula aloes CPC 19671 JX069863 JX069847 NA NA NA Crous et al. (2012)
CBS 132531T NR_111757 NG_042676 NA NA NA Crous et al. (2012)
Montagnula appendiculata CBS 109027T DQ435529 AY772016 NA NA NA Wanasinghe et al. (2016)
Montagnula aquatica MFLU 22-0171T OP605992 OP605986 OP600504 NA NA Sun et al. (2023)
Montagnula aquatica KUNCC 23-14425 OR583097 OR583116 OR583135 OR588088 OR588107 This study
KUNCC 23-14557 OR583099 OR583118 OR583137 OR588090 OR588109 This study
Montagnula aquilariae KUNCC 22-10815T OP452927 OP482265 OP482268 OP426318 NA Hyde et al. (2023)
KUNCC 22-10816 OP554219 OP482266 OP482269 OP426319 NA Hyde et al. (2023)
KUNCC 22-10815T OP452927 OP482265 OP482268 OP426318 NA Hyde et al. (2023)
KUNCC 22-10816 OP554219 OP482266 OP482269 OP426319 NA Hyde et al. (2023)
Montagnula aquilariae KUNCC 23-14430 OR583100 OR583119 OR583138 OR588091 OR588110 This study
KUNCC 23-14431 OR583101 OR583120 OR583139 OR588092 OR588111 This study
KUNCC 23-14432 OR583102 OR583121 OR583140 OR588093 OR588112 This study
Montagnula bellevaliae MFLUCC 14-0924T KT443906 KT443902 KT443904 NA NA Hongsanan et al. (2015)
Montagnula camporesii MFLUCC 16-1369T MN401746 NG_070946 NG_068418 MN397908 MN397909 Hyde et al. (2020)
Montagnula chiangraiensis MFLUCC 17-1420T NR_168864 NG_068707 NG_070155 NA NA Mapook et al. (2020)
Montagnula chromolaenae MFLUCC 17-1435T NR_168865 NG_068708 NG_070156 NA NA Mapook et al. (2020)
Montagnula chromolaenicola MFLUCC 17-1469T NR_168866 NG_070948 NG_070157 MT235773 MT235809 Mapook et al. (2020)
Montagnula chromolaenicola KUNCC 23-14426 OR583098 OR583117 OR583136 OR588089 OR588108 This study
KUNCC 23-14427 OR583103 OR583122 OR583141 OR588094 OR588113 This study
KUNCC 23-14558 OR583104 OR583123 OR583142 OR588095 OR588114 This study
Montagnula cirsii MFLUCC 13-0680 KX274242 KX274249 KX274255 KX284707 NA Hyde et al. (2016)
Montagnula cylindrospora CBS 146572T LT796834 LN907351 NA LT797074 LT796994 Crous et al. (2020)
Montagnula donacina HFG07004 MF967419 MF183940 NA NA NA Zhao et al. (2017)
HVVV01 KJ628375 KJ628377 KJ628376 NA NA Pitt et al. (2014)
HKAS 124552 OP605991 OP605987 NA NA NA Sun et al. (2023)
KUMCC 21-0653 OP058961 OP059052 OP059003 OP135938 NA Ren et al. (2021)
KUMCC 21-0579 OP058963 OP059054 OP059005 OP135940 NA Ren et al. (2021)
KUMCC 21-0631 OP058962 OP059053 OP059004 OP135939 NA Ren et al. (2021)
UESTCC 23.0030 OR253120 OR253279 OR253194 NA NA Unpublished
Montagnula donacina KUNCC 23-14428 OR583105 OR583124 OR583143 OR588096 OR588115 This study
KUNCC 23-14429 OR583106 OR583125 OR583144 OR588097 OR588116 This study
Montagnula graminicola MFLUCC 13-0352T KM658314 KM658315 KM658316 NA NA Liu et al. (2015)
Montagnula guiyangensis HKAS 124556T OP605989 OP600484 OP600500 NA NA Sun et al. (2023)
GUCC 22–0817 OP605990 OP600485 OP600501 NA NA Sun et al. (2023)
Montagnula jonesii MFLUCC 16-1448T KY313619 KY273276 KY313618 KY313620 NA Tennakoon et al. (2016)
MFLU 18-0084 ON117282 ON117300 ON117269 ON158095 NA Tennakoon et al. (2022)
Montagnula krabiensis MFLUCC 16-0250T NR168179 NG068826 NG068385 MH412776 NA Tibpromma et al. (2018)
Montagnula lijiangensis HKAS 126540 OR583107 OR583126 OR583145 OR588098 OR588117 This study
HKAS 126541T OR583108 OR583127 OR583146 OR588099 OR588118 This study
Montagnula menglaensis KUNCC 23-14422 OR583109 OR583128 OR583147 OR588100 OR588119 This study
KUNCC 23-14423 OR583110 OR583129 OR583148 OR588101 OR588120 This study
KUNCC 23-14424T OR583111 OR583130 OR583149 OR588102 OR588121 This study
Montagnula puerensis KUMCC 20-0225T MW567739 MW575866 MW575864 MW575859 NA Du et al. (2021)
KUMCC 20-0331 MW567740 MW575867 MW575865 MW575860 NA Du et al. (2021)
Montagnula saikhuensis MFLUCC 16-0315T KU743209 KU743210 KU743211 NA NA Wanasinghe et al. (2016)
Montagnula scabiosae MFLUCC 14-0954T KT443907 KT443903 KT443905 NA NA Hongsanan et al. (2015)
Montagnula shangrilana KUNCC 23-14433 OR583112 OR583131 OR583150 OR588103 OR588122 This study
KUNCC 23-14434T OR583113 OR583132 OR583151 OR588104 OR588123 This study
Montagnula thailandica MFLUCC 17-0363 OL782142 OL782059 OL780525 OL875102 OL828754 Senwanna et al. (2021)
MFLUCC 17-1508T MT214352 NG070949 NG070158 MT235774 MT235810 Mapook et al. (2020)
MFLUCC 21-0075 OP297807 OP297777 OP297791 OP321576 NA Lu et al. (2022)
ZHKUCC 22-0206 OP297808 OP297778 OP297792 OP321577 NA Lu et al. (2022)
ZHKUCC 22-0207 MZ538515 MZ538549 NA MZ567092 NA Boonmee et al. (2021)
Montagnula thevetiae HKAS 126963 OR583114 OR583133 OR583152 OR588105 OR588124 This study
HKAS 126964T OR583115 OR583134 OR583153 OR588106 OR588125 This study
Neokalmusia jonahhulmei KUMCC 21-0818T ON007043 ON007039 ON007048 ON009133 ON009137 Wanasinghe and Mortimer (2022)
Neokalmusia jonahhulmei KUMCC 21-0819 ON007044 ON007040 ON007049 ON009134 ON009138 Wanasinghe and Mortimer (2022)

Results

Phylogenetic analyses

In order to examine the evolutionary relationships of our new strains within Montagnula, phylogenetic analyses were performed based on the combined SSU, LSU, ITS, tef1-α, and rpb2 DNA sequences of 56 representatives of the genus and two strains from Neokalmusia jonahhulmei (KUMCC 21-0818, KUMCC 21-0819) as the outgroup taxon. The full dataset consisted of 4,268 characters including gaps (18S = 1,023 characters, 28S = 896, ITS = 508, tef1-α = 885, rpb2 = 956). The RAxML analysis of the combined dataset yielded a best-scoring tree with a final ML optimization likelihood value of -14,343.052271. The matrix had 1004 distinct alignment patterns, with 23.88% undetermined characters or gaps. Parameters for the GTR + I + G model of the combined amplicons were as follows: Estimated base frequencies; A = 0.244145, C = 0.256118, G = 0.269851, T = 0.229886; substitution rates AC = 1.815063, AG = 3.954334, AT = 1.414215, CG = 1.362941, CT = 10.779403, GT = 1.000; proportion of invariable sites I = 0.559204; and gamma distribution shape parameter α = 0.542439. The Bayesian analysis ran 1,675,000 generations before the average standard deviation for split frequencies reached below 0.01 (0.009994). The analyses generated 16,751 trees, from which we sampled 12,564 trees after discarding the first 25% as burn-in. The alignment contained a total of 1,005 unique site patterns. The BI and ML trees were not in conflict; the ML tree is shown in Fig. 1. Where applicable, the phylogenetic results obtained (Fig. 1) are discussed in the descriptive notes below.

Figure 1. 

Phylogenetic analysis of SSU, LSU, ITS, tef1-α, and rpb2 of the Montagnula. Species names given in bold are ex-type, ex-epitype and ex-paratype strains. Species names highlighted in blue are generated from this study. Branch support of nodes ≥75% ML BS and ≥0.95 PP is indicated above the branches. The genus Montagnula is depicted within a pale gray box, with new species highlighted in white, and the outgroup indicated by a blue box.

Figure 2. 

Geographical distribution of Montagnula species with known ITS sequence data. a the map summarizes data from the GlobalFungi database (shown by circles). Each circle symbolizes a unique sample, with each color representing the specific biome from which it has been collected b the distribution of Montagnula sequences as a percentage of total abundance across different biomes c the distribution of Montagnula sequences as a percentage of total abundance across different continents. See Suppl. material 1 for primary data.

We conducted a thorough study of a compilation of data derived from multiple metabarcoding studies, which documented the occurrence of Montagnula species worldwide, excluding Antarctica. Among the continents, the highest number of studies were recorded in Asia, Australia, Europe, and North America (Fig. 2). These studies encompassed a diverse range of 11 distinct sources, revealing that sediments and “other” sources yielded the highest number of sequences (Fig. 3). Across different continents, the sequences obtained from various sources exhibited moderate similarity. However, in regions such as Asia, Australia, Europe, and North America, studies revealed Montagnula species from a diverse array of sources, in contrast to other studies, which identified species from a more limited selection of sources. Furthermore, in culture-based investigations, the primary focus was on extracting Montagnula species from plant substrates originating from 45 distinct plant families (Fig. 4). Among these families, Poaceae yielded the most substantial number of isolated species, followed by Asparagaceae and Fabaceae. Additionally, two records were also detected in mushrooms and human skin samples.

Figure 3. 

The distribution of Montagnula occurrences across oceans, continents and various substrates, as documented in the existing literature. On the x-axis, the logarithmic abundance of each record for different sources is displayed.

Figure 4. 

The species richness of recorded Montagnula species across different plant families (Table 1).

Taxonomy

Pleosporales Luttr. ex M.E. Barr, Prodromus to class Loculoascomycetes: 67 (1987)

Didymosphaeriaceae Munk, Dansk botanisk Arkiv 15 (2): 128 (1953)

Montagnula Berl., Icones Fungorum. Pyrenomycetes 2: 68 (1896)

Notes

This study presents an updated and comprehensive phylogenetic classification of the genus Montagnula, incorporating SSU, LSU, ITS, tef1-α, and rpb2 DNA sequence analyses. By combining morphological and phylogenetic considerations, we have identified four new species, M. lijiangensis, M. menglaensis, M. shangrilana and M. thevetiae within the genus. Additionally, this research accounts for the existing species viz., M. aquatica, M. aquilariae, M. chromolaenicola and M. donacina. The note sections of this publication provide detailed information on these taxonomic accounts, including additional discussion and supporting evidence. Each newly identified species adds to the known biodiversity within the genus, expanding our knowledge of the ecological and morphological characteristics exhibited by Montagnula taxa.

Montagnula aquatica Y.R. Sun, Yong Wang bis & K.D. Hyde, Plants 12 (4, no. 738): 2 (2023)

MycoBank No: 900129

Descriptions and illustrations

See Sun et al. (2023).

Habitat and distribution

This species is found in freshwater habitats of Chiang Rai, Thailand, terrestrial habitats of Yunnan, China, inhabiting dead wood of deciduous hosts (Sun et al. 2023, this study).

Material examined

China, Yunnan Province, Honghe Hani and Yi Autonomous Prefecture, Honghe County, Dayangjiexiang (23.389965°N, 102.225552°E, 1194 m), on dead woody litter of an unidentified plant, 13 March 2023, D.N. Wanasinghe, DWHH23-51 (HKAS 130322), new country and habitat record, living culture KUNCC 23-14425. ibid. 23.388966°N, 102.224786°E, 1215 m, DWHH23-51-2 (HKAS 130323), living culture KUNCC 23-14557.

Notes

Based on our phylogenetic analyses, we have determined that the newly collected strains (i.e. KUNCC 23-14425 and KUNCC 23-14557) are monophyletic with the ex-type strain of Montagnula aquatica (MFLU 22-0171). Further morphological investigations comparing our isolate with the type species have revealed similarities in the size range of the ascomata, asci, and ascospores, as well as the ascospore septation (Sun et al. 2023). Therefore, we document KUNCC 23-14425 and KUNCC 23-14557 as new records of Montagnula aquatica in China, accompanied by protein sequence data (tef1-α and rpb2) for this species. It is worth noting that the holotype of Montagnula aquatica was previously reported on submerged decaying wood in a freshwater habitat in Thailand, while our collection was made from a terrestrial habitat in China. This observation suggests that this fungus exhibits adaptability to a wide range of habitats, although its exploration in diverse geographic locations remains limited. The inclusion of Montagnula aquatica as a new record in China expands our understanding of the distribution and ecological preferences of this species in both terrestrial and aquatic habitats. Additionally, the protein sequence data obtained for this strain contributes valuable information to the existing knowledge on Montagnula aquatica. Further studies exploring the ecological aspects of this fungus in different geographic locations will provide deeper insights into its adaptability and potential ecological roles.

Montagnula aquilariae T.Y. Du & Tibpromma, Mycosphere 14 (1): 705 (2023)

MycoBank No: 846332
Fig. 5

Description

Saprobic on dead woody litter of an unknown deciduous host. Teleomorph Ascomata 450–600 μm high × 480–550 μm diam., immersed to semi-erumpent, gregarious or rarely clustered, globose to subglobose, ostiolate. Ostiole 120–220 × 70–110 µm (x– = 139 × 89 μm, n = 5), papillate, central, straight, dark brown to black, filled with hyaline cells, periphyses are lacking. Peridium 20–40 μm thick on the sides and can reach up to 60 μm near the apex, with an outer layer consisting of heavily pigmented cells that have thick walls and exhibit a textura angularis to textura globulosa texture at the apex, textura angularis texture at the sides and base; the innermost layer consists of narrow, hyaline compressed rows of cells that merge with pseudoparaphyses. Hamathecium of 2–4 μm broad, dense, narrow, branched, cellular pseudoparaphyses. Asci 100–120 × 16–22 µm (x– = 110.8 × 18.4 μm, n = 20), bitunicate, fissitunicate, cylindrical-clavate to clavate, pedicel 30–50 μm long, 8-spored, biseriate, with a minute ocular chamber best seen in immature ascus. Ascospores 20–25 × 8.5–11 µm (x– = 21.8 × 9.6 μm, n = 30), ellipsoidal to narrowly oblong, straight or somewhat curved, ends conically rounded, golden-brown to dark brown, 1-septate, constricted at the septum, large guttules in each cell, verruculose, with a thin mucilaginous sheath. Anamorph Undetermined.

Habitat and distribution

This species is found in terrestrial habitats of Yunnan, China, specifically inhabiting dead woody twigs of deciduous hosts, including Aquilaria sinensis (Hyde et al. 2023, this study).

Material examined

China, Yunnan Province, Kunming City, Kunming Institute of Botany (25.141723°N, 102.750013°E, 1970 m), on dead woody litter of an unidentified plant, 24 April 2022, L. Qinxian, KIB22-17-1 (HKAS 126542), living culture KUNCC 23-14430; ibid. 25.141487°N, 102.748863°E, 1982 m, K2B22-17-3 (HKAS 126543), living culture KUNCC 23-14431; ibid. K2B22-17-4 (HKAS 126544), living culture KUNCC 23-14432.

Notes

Montagnula aquilariae was recently introduced by Hyde et al. (2023) based on samples obtained from Aquilaria sinensis in Xishuangbanna, Yunnan Province. In our new collections, three strains (KUNCC 23-14430, KUNCC 23-14431, KUNCC 23-14432) exhibited a monophyletic relationship with the previously known strains of Montagnula aquilariae (KUNCC 22-10815 [ex-type] and KUNCC 22-10816). Through further morphological, ecological, and nucleotide (SSU, LSU, ITS, tef1-α) comparisons, we have confirmed that these new strains indeed belong to Montagnula aquilariae. Our research also provides additional insights into the characteristics of Montagnula aquilariae. Specifically, we report the verruculose feature of the ascospores and present rpb2 sequence data for this fungus, advancing our knowledge of its morphological and genetic attributes.

Figure 5. 

Montagnula aquilariae (HKAS 126542) a, b ascomata on natural wood surface c vertical section through an ascoma d ostiolar neck e peridium cells at the apex f peridium cells at the side g pseudoparaphyses h–l asci m–r ascospores (see verruculose feature of the ascospore in r) s, t culture characters on PDA (s = above, t = reverse). Scale bars: 100 μm (c, d); 50 μm (e); 10 μm (e–g, m–r); 20 μm (h–l).

Montagnula chromolaenicola Mapook & K.D. Hyde, Fungal Diversity 101: 35 (2020)

MycoBank No: 557298

Descriptions and illustrations

See Mapook et al. (2020).

Habitat and distribution

This species was observed in terrestrial habitats in Mae Hong Son, Thailand, specifically on dead stems of Chromolaena odorata (Mapook et al. 2020). Additionally, it has also been found in terrestrial habitats in Yunnan, China, where it inhabits dead wood of deciduous hosts (this study).

Material examined

China, Yunnan Province, Honghe County, Honghe Hani and Yi Autonomous Prefecture, Dayangjiexiang (23.389965°N, 102.225552°E, 1201 m), on a dead woody climber of an unidentified host, 13 March 2023, D.N. Wanasinghe, DWHH23-17A (HKAS 130321), living culture KUNCC 23-14426. ibid. 23.389295°N, 102.224780°E, 1200 m, on dead twigs of Lagerstroemia sp. DWHH23-33-2 (HKAS 126543), living culture KUNCC 23-14427; ibid. DWHH23-33-3 (HKAS 130320), living culture KUNCC 23-14558.

Notes

Through our phylogenetic analyses, we have determined that the newly isolated strains HH33 and HH17A exhibit a monophyletic relationship with the ex-type strain of Montagnula chromolaenicola (MFLUCC 17-1469). Upon conducting further investigations and morphological comparison of our collection with the type species, we have discovered several similarities. These include the size range of the ascomata, asci, and ascospores, as well as the ascospore septation (Mapook et al. 2020). Consequently, we hereby document our new collections (i.e. HKAS 130321, HKAS 126543 and HKAS 130320) as new records of Montagnula chromolaenicola in China. In a recent study by Sun et al. (2023), Montagnula chromolaenicola, M. puerensis, M. saikhuensis, and M. thailandica were synonymized under the name M. donacina due to the absence of obvious branches in their phylogenetic tree and the close morphological resemblance between these species. However, it is important to note that most of these strains lack informative sequence data for tef1-α or rpb2. Our observations, on the other hand, have revealed that the inclusion of protein data in this group leads to the formation of distinct branches and independent lineages. Therefore, we propose retaining the older names for these species, except for Montagnula thailandica, until further research resolves this group using all available sequence data.

Montagnula donacina (Niessl) Wanas., E.B.G. Jones & K.D. Hyde, Index Fungorum 319: 1 (2017)

MycoBank No: 552762

Descriptions and illustrations

See Pitt et al. (2014).

Habitat and distribution

This species has been reported worldwide on various hosts within terrestrial habitats (see Table 2). Specifically, it has been documented in Australia (Calamus australis, Vitis vinifera), Brazil (Bambusoideae, Saccharum officinarum), Central African Republic (Coffea robusta), China (Althaea rosea, Craterellus odoratus, Trachycarpus fortunei), Colombia (unknown plant), France (Pseudosasa japonica), Georgia (Zea mays), India (Acacia sp., Adhatoda vasica, Ailanthus altissima, Annona squamosa, Cajanus cajan, Careya arborea, Citrus aurantiifolia, Clerodendrum infortunatum, C. multiflorum, Duranta repens, Ficus glomerata, Hibiscus sp., Ipomoea carnea, Mallotus philippinensis, Morus alba, Nerium odorum, Pistacia indica, Tectona grandis, Terminalia tomentosa), Japan (Phyllostachys bambusoides), Myanmar (Nephelium litchi), Namibia (Acacia reficiens), Papua New Guinea (Bambusoideae), Paraguay (Coffea arabica), Philippines (Premna cumingiana), Portugal (Arundo donax), Sierra Leone (Funtumia africana), Thailand (dead wood) and the USA (Platanus sp., Wikstroemia sp.).

Material examined

China, Yunnan Province, Honghe (23.424892°N, 102.231417°E, 600 m), on dead woody litter of an unidentified plant, 14 August 2022, D.N. Wanasinghe, DWHH22-23-1 (HKAS 126545), living culture KUNCC 23-14428. ibid. DWHH22-23-2 (HKAS 126546), living culture KUNCC 23-14429.

Notes

Wanasinghe et al. (2016) regarded Munkovalsaria as a synonym of Montagnula and established Montagnula donacina (=Munkovalsaria donacina). So far, Montagnula donacina stands as the most extensively distributed species within the genus. Despite its global presence, there is a scarcity of molecular data available for Montagnula donacina. A preliminary analysis revealed only 20 sequence data entries when searching for “ Montagnula donacina” in the NCBI database, originating from only seven strains: HFG07004, HKAS 124552, HVVV01, KUMCC 21-0579, KUMCC 21-0631, KUMCC 21-0653, and UESTCC:23.0030. Our phylogenetic analysis demonstrated a close relationship between two strains designated as Montagnula donacina (HVVV01 and HFG07004) and the type strain of Montagnula chromolaenicola (MFLUCC 17-1469). Additionally, we observed that the strains of Montagnula thailandica formed a monophyletic group alongside the remaining Montagnula donacina strains (HKAS 124552, KUMCC 21-0579, KUMCC 21-0631, KUMCC 21-0653, and UESTCC:23.0030). Furthermore, two newly generated sequences, KUNCC 23-14428 and KUNCC 23-14429, were also clustered with the strains of Montagnula donacina. We hereby introduce these two strains as belonging to Montagnula donacina and provide rpb2 sequence data for this species for the first time.

Montagnula lijiangensis Wanas., sp. nov.

MycoBank No: 850093
Fig. 6

Etymology

The specific epithet “lijiangensis” refers to Lijiang, Yunnan Province, where the holotype was collected.

Holotype

HKAS 126541.

Description

Saprobic on dead woody litter of Quercus sp. Teleomorph Ascomata 500–700 μm high × 500–600 μm diam., immersed, gregarious or rarely clustered, globose to subglobose, ostiolate. Ostiole 100–140 × 80–120 µm (x– = 125 × 96 μm, n = 5), apapillate, central, straight, filled with hyaline cells. Peridium 20–30 μm thin on the sides and can reach up to 70 μm near the apex, with an outer layer consisting of heavily pigmented cells that have thick walls and exhibit a textura angularis texture at the apex, textura angularis texture at the sides and base; the innermost layer consists of narrow, hyaline compressed rows of cells. Hamathecium of 3–7.5 μm broad, dense, narrow, branched, cellular pseudoparaphyses that are swollen at the base. Asci 130–160 × 20–26 µm (x– = 152.8 × 23.9 μm, n = 20), bitunicate, fissitunicate, cylindrical-clavate to clavate, pedicel 30–60 μm long, 8-spored, uni to biseriate, with a minute ocular chamber best seen in immature ascus. Ascospores 22–26 × 10–14 µm (x– = 24.8 × 11.8 μm, n = 30), ellipsoidal to narrowly oblong, mostly straight, with conically rounded ends at the immature stage that become rounded when mature, golden-brown to dark brown, 1-septate and constricted at the septum, with large guttules in each cell, verruculose, surrounded by a thick mucilaginous sheath. Anamorph Undetermined.

Figure 6. 

Montagnula lijiangensis (HKAS 126541, holotype) a, b ascomata on natural wood surface c vertical section through an ascoma d ostiolar neck and peridium cells at the apex e pseudoparaphyses f–i asci j–o ascospores (see verruculose feature of the ascospore in k). Scale bars: 100 μm (c); 20 μm (d, f–i); 10 μm (e–o).

Habitat and distribution

This species is found in terrestrial habitats of Yunnan, China, inhabiting dead woody twigs of deciduous hosts (this study).

Material examined

China, Yunnan Province, Lijiang, Yulong County (26.86389°N, 99.824738°E, 2725 m), on dead woody litter of Quercus sp. (Fagaceae), 17 August 2021, L. Qinxian, STX09-03-1 (holotype, HKAS 126541, ibid. 26.863484°N, 99.824548°E, 2706 m, STX09-03-3 (HKAS 126540).

Notes

The analysis of two newly generated sequences revealed a monophyletic clade in our phylogenetic analysis (Fig. 1), demonstrating a close phylogenetic relationship to Montagnula aquilariae. This relationship is further supported by morphological features such as asci and ascospores. However, a comparison of nucleotide differences (without gaps) between these two clades (KUNCC 22-10815 and KUNCC 23-14430 vs HKAS 126541) showed 12/508 (2.3%) differences in the ITS region, 15/885 (1.7%) differences in the tef1-α region, and 19/956 (2%) differences in the rpb2 region.

Montagnula menglaensis Wanas., sp. nov.

MycoBank No: 850094
Fig. 7

Etymology

The specific epithet “menglaensis” refers to Mengla County, Yunnan Province, where the holotype was collected.

Holotype

HKAS 130318.

Description

Saprobic on dead culms of Indocalamus tessellatus (Munro) Keng f. Teleomorph Ascomata 200–300 μm high × 240–320 μm diam., immersed, gregarious or rarely clustered, globose to subglobose. Peridium 10–25 μm thin with an outer layer consisting of heavily pigmented cells that have thick walls and exhibit a textura angularis texture at the sides and base; the innermost layer consists of narrow, hyaline compressed rows of cells. Hamathecium of 3–7.5 μm broad, dense, branched, cellular pseudoparaphyses that are swollen at some septa. Asci 60–80 × 9–11 µm (x– = 71 × 9.8 μm, n = 15), bitunicate, fissitunicate, cylindrical-clavate, pedicel 15–30 μm long, 8-spored, uni to biseriate, with a minute ocular chamber best seen in immature ascus. Ascospores 10.5–14 × 4.5–5.5 µm (x– = 12.6 × 5.1 μm, n = 20), ellipsoidal, mostly straight, with conically rounded ends, golden-brown to dark brown, 1-septate and constricted at the septum, upper cell wider than the lower cell, with large guttules in each cell, verruculose, and surrounded by a thin mucilaginous sheath which is thicker at both ends. Anamorph Coelomycetous on PDA. Conidiomata pycnidial, gregarious, immersed to superficial, globose to subglobose, dark brown to black. Pycnidial wall thin, composed of brown cells of textura angularis. Conidiogenous cells did not observed. Conidia 2.3–3.3 × 1.4–2 μm (x– = 3 × 1.7 μm, n = 30), hyaline, aseptate, round to oblong or ellipsoidal, with small guttules.

Figure 7. 

Montagnula menglaensis (HKAS 130318, holotype) a–c ascomata on natural wood surface d, e vertical section through ascomata f, g pseudoparaphyses h peridium i–k asci l, m ascospores (see verruculose feature of the ascospore in n) o, p culture characters on PDA (o = above, p = reverse) q, r conidiomata s pycnidial wall t conidia. Scale bars: 100 μm (d, e); 10 μm (f–h, l–n, s, t); 20 μm (i–k).

Culture characteristics

Ascospores germinated on PDA within 24 h. Following a two-week incubation period at 25 °C, the colonies on PDA medium reached a diameter of 5 cm. These colonies exhibited an undulate margin, initially appearing creamy whitish and transitioning to orange, raised in the center. The colonies were orange at the center and a creamy orange towards the periphery when observed from the reverse side.

Habitat and distribution

This species is found in terrestrial habitats of Yunnan, China, inhabiting dead woody twigs of deciduous hosts (this study).

Material examined

China, Yunnan Province, Xishuangbanna, Mengla County (21.588394°N, 101.435042°E, 776 m), on dead culms of Indocalamus tessellatus, 29 January 2022, L. Qinxian, ML23-7-3 (holotype, HKAS 130318), ex-type KUNCC 23-14424; ibid. 21.589178°N, 101.435752°E, 782 m, ML23-7-2 (HKAS 130316), living culture KUNCC 23-14422; ibid. ML23-7-5 HKAS 130317), living culture KUNCC 23-14423.

Notes

Montagnula menglaensis is described as a novel species based on its holomorph. The anamorph of Montagnula is rarely encountered; however, Crous et al. (2020) recently reported Montagnula cylindrospora based on its anamorphic features. The conidia of Montagnula menglaensis resemble to those of M. cylindrospora, although the latter fungus exhibits a more cylindrical shape.

Montagnula shangrilana Wanas., sp. nov.

MycoBank No: 850095
Fig. 8

Etymology

The specific epithet “shangrilana” refers to Shangri-La, Yunnan Province, where the holotype was collected.

Holotype

HKAS 126539.

Description

Saprobic on dead woody litter of Rhododendron sp. Teleomorph Ascomata 120–180 μm high × 150–210 μm diam., immersed to semi-erumpent, gregarious or rarely clustered, globose to subglobose, ostiolate. Ostiole 80–110 × 50–80 µm (x– = 100 × 64 μm, n = 6), papillate, central, straight, filled with hyaline cells. Peridium 10–20 μm thin on the sides and can reach up to 40 μm near the apex, with an outer layer consisting of heavily pigmented cells that have thick walls and exhibit a textura angularis arrangement at the apex, textura angularis texture at the sides; the innermost layer consists of hyaline compressed rows of cells. Hamathecium of 2–4.5 μm broad, dense, branched, cellular pseudoparaphyses. Asci 90–140 × 20–30 µm (x– = 116.2 × 24 μm, n = 10), bitunicate, fissitunicate, cylindrical-clavate, pedicel 25–40 μm long, 8-spored, uni to biseriate, with a minute ocular chamber best seen in immature ascus. Ascospores 48–60 × 17–22 µm (x– = 55.8 × 19.3 μm, n = 20), ellipsoidal to narrowly oblong, mostly straight, with conically rounded ends at the immature stage that become rounded when mature, golden-brown to dark brown, 3-septate, with large guttules in each cell, verruculose, surrounded by a thick mucilaginous sheath. Anamorph Undetermined.

Figure 8. 

Montagnula shangrilana (HKAS 126541, holotype) a ascomata on natural wood surface b vertical section through an ascoma c pseudoparaphyses d peridium cells e–h asci i–o ascospores (see verruculose feature of the ascospore in o). Scale bars: 100 μm (b); 10 μm (c, d, j–o); 20 μm (e–h).

Culture characteristics

Ascospores germinated on PDA within 24 h. Following a two-week incubation period at 25 °C, the colonies on PDA medium reached a diameter of 5 cm. These colonies exhibited a filiform margin, initially appearing whitish and transitioning to greenish gray, raised in the center. The colonies were grey at the center and a greenish gray towards the periphery and radiated when observed from the reverse side.

Habitat and distribution

This species is found in terrestrial habitats of Yunnan, China, inhabiting dead woody twigs of deciduous hosts, in a subalpine environment (this study).

Material examined

China, Yunnan Province, Diqing Tibetan Autonomous Prefecture, Shangri-La (27.289707°N, 100.034477°E, 2744 m), on dead woody litter of Rhododendron sp. (Ericaceae), 22 August 2021, L. Qinxian, WTS8-2-2 (holotype, HKAS 126539), ex-type KUNCC 23-14434; ibid. (27.290007°N, 100.035233°E, 2833 m, WTS8-2 (HKAS 126538), living culture KUNCC 23-14433.

Notes

In the combined SSU, LSU, ITS, tef1-α, and rpb2 phylogenetic analysis, two strains of Montagnula shangrilana (HKAS 126538, HKAS 126539) formed a monophyletic clade closely related to M. camporesii (MFLUCC 16-1369), M. cirsii (MFLUCC 13-0680), and M. scabiosae (MFLUCC 14-0954). While there were slight variations in size, shape, and color, all four species shared the common characteristic of 3-transversely septate ascospores. The sequence data of Montagnula camporesii, M. cirsii, and M. scabiosae showed no significant differences in their base pair comparisons, suggesting that they may be conspecific. Morphologically, these three species exhibited clavate asci and ellipsoid to fusiform, brown, overlapping, 3-septate ascospores. In contrast, our newly discovered species differed from these three species by 10/508 (1.96%) differences in the ITS region, 13/885 (1.5%) differences in the tef1-α region, and 15/956 (1.56%) differences in the rpb2 region (only M. camporesii possesses rpb2).

Montagnula thevetiae Wanas., sp. nov.

MycoBank No: 850096
Fig. 9

Etymology

The specific epithet “thevetiae” refers to the host Thevetia peruviana from which the holotype was isolated.

Holotype

HKAS 126964.

Description

Saprobic on dead twigs of Thevetia peruviana. Teleomorph Ascomata 140–160 μm high × 150–190 μm diam., immersed, gregarious or rarely clustered, globose to subglobose, ostiolate. Ostiole 40–65 × 50–90 µm (x– = 50 × 78 μm, n = 6), papillate, central, straight, filled with hyaline to brown cells. Peridium 10–20 μm thin on the sides and can reach up to 30 μm near the apex, with an outer layer consisting of heavily pigmented cells that have thick walls and textura angularis arrangement, the inner layer consists of hyaline compressed rows of cells. Hamathecium of 2–3.5 μm broad, dense, branched, cellular pseudoparaphyses. Asci 110–160 × 25–35 µm (x– = 126.4 × 30.3 μm, n = 12), bitunicate, fissitunicate, cylindrical-clavate, pedicel 25–35 μm long, 8-spored, uni to biseriate, with a minute ocular chamber best seen in immature ascus. Ascospores 30–40 × 11.5–14 µm (x– = 37.3 × 12.8 μm, n = 20), ellipsoidal to narrowly oblong, straight to curved, with conically rounded ends, brown to dark brown, 1-septate, constricted at the septum, with large guttules in each cell, verruculose, surrounded by a thin mucilaginous sheath. Anamorph Undetermined.

Figure 9. 

Montagnula thevetiae (HKAS 126564, holotype). a, b ascomata on natural wood surface c vertical section through an ascoma d closeup of ostiole e pseudoparaphyses f–h asci j–l ascospores m, n culture characteristics on PDA (m = above, n = reverse). Scale bars: 100 μm (c); 50 μm (d, f–h); 10 μm (e, i–l).

Culture characteristics

Ascospores germinated on PDA within 24 h. Following a two-week incubation period at 25 °C, the colonies on PDA medium reached a diameter of 4 cm. These colonies exhibit an irregular, flattened to slightly raised morphology and display various color sectors ranging from white, creamy orange to pale brown. The reverse side of the colonies appears creamy orange, with occasional dark patches that can be observed.

Habitat and distribution

This species is found in terrestrial habitats of Yunnan, China, inhabiting dead woody twigs of Thevetia peruviana (this study).

Material examined

China, Yunnan Province, Kunming city, Kunming Institute of Botany (25.142238°N, 102.750354°E, 1971 m), on dead twigs of Thevetia peruviana, 24 April 2022, L. Qinxian, K2B22-26-2 (holotype, HKAS 126964), ibid. (25.140859°N, 102.749045°E, 1968 m, K2B22-26 (HKAS 126963).

Notes

Montagnula thevetiae is isolated from the dead twigs of Thevetia peruviana. The newly obtained sequences of this fungus formed a monophyletic clade closely related to Montagnula menglaensis. Morphologically, they share similarities in having 1-septate ascospores, although Montagnula thevetiae exhibits a darker pigmentation. On the other hand, Montagnula thevetiae differs from M. menglaensis by 15/1023 (1.46%) differences in the SSU region, 19/895 (2.12%) differences in the LSU region, 32/508 (6.3%) differences in the ITS region, 27/885 (3%) differences in the tef1-α region, and 86/956 (9%) differences in the rpb2 region.

Discussion

Montagnula species in Yunnan Province

The study of lignicolous microfungi in Yunnan Province resulted in the collection of eight Montagnula species, including four novel species. This study contributes to our understanding of the diversity and distribution of Montagnula species and provides insight into the ecological roles played by these fungi in their respective habitats. Montagnula aquatica was previously documented as occurring on submerged decaying wood within a freshwater habitat in Thailand (Sun et al. 2023). However, our recent collection of this species was obtained from a terrestrial habitat in China. The holotype was collected in the Bandu District of the Chiang Rai Province, situated at an approximate elevation of 400–450 m and characterized by a tropical climate. The collection site was near to a waterfall (Sun et al. 2023). In contrast, our new collections were made in the Honghe region of Yunnan Province, which possesses an elevation of approximately 1200 m. The local environment in this region is characterized by poor, eroded soils, steep valleys, and a subtropical climate. This observation suggests that Montagnula aquatica may possess an adaptable nature, enabling it to thrive in a wide range of habitats across diverse geographic locations. Montagnula aquilariae, another species within the genus, has been identified in the terrestrial habitats of Yunnan, China. It specifically colonizes dead woody twigs of deciduous hosts, including Aquilaria sinensis (Hyde et al. 2023). The holotype of this species was collected from a hilly area in Nanmo, Menghai and Xishuangbanna, situated at an elevation of ~1100 m and characterized by a tropical climate. Additional collections were made from Kunming, located within the same province but at an elevation of ~2000 m, and characterized by a warm and temperate climate. Montagnula chromolaenicola has been observed in terrestrial habitats in Thailand, particularly on dead stems of Chromolaena odorata (Mapook et al. 2020). The holotype of this species was collected from the Mae Yen mountainous area of Mae Hong Son Province, at an elevation of ~900 m. The local environment of this area exhibits a tropical savanna climate. In our study, we collected this fungus from a terrestrial habitat within the steep valleys of subtropical Honghe, Yunnan, China. In this region, Montagnula chromolaenicola was found to inhabit the dead woody litter of deciduous hosts. Montagnula donacina has been reported across various terrestrial habitats worldwide, with the majority of records originating from India (Table 1). This species primarily associates with hosts from the Poaceae family. In our study, we collected Montagnula donacina from the subtropical Honghe region in China, specifically on decaying woody litter at an elevation of ~600 m. Montagnula lijiangensis was collected from terrestrial habitats at a high elevation of ~2725 m. This species was found on dead woody litter of Quercus sp. within an environment characterized by a mild subtropical highland climate. Montagnula menglaensis was discovered in the terrestrial habitats of Mengla County, Yunnan, China. It was observed colonizing dead culms of Indocalamus tessellatus. The local environment of this region exhibits a tropical savanna climate, with an elevation of ~800 m. Montagnula shangrilana was found in the terrestrial habitats of Shangri-La, Yunnan, China, where it inhabits dead woody twigs of Rhododendron sp. This species has also been observed at higher elevations, reaching ~2800 m, within an environment characterized by a humid continental climate. Montagnula thevetiae was discovered within the terrestrial habitat of the botanical garden at the Kunming Institute of Botany in Yunnan, China. This species was found colonizing dead woody twigs of Thevetia peruviana. The collection site is situated at an elevation of ~2000 m and experiences a warm and temperate climate.

Taxonomic reassessment and phylogenetic analysis of Montagnula species

In a recent study conducted by Sun et al. (2023), Montagnula chromolaenicola, M. puerensis, M. saikhuensis, and M. thailandica were regarded as the synonyms of M. donacina (Wanasinghe et al. 2016). This decision was based on the absence of clear branches in their phylogenetic tree and the close morphological resemblance between these species. However, upon further examination, it was observed in this study that only Montagnula donacina and M. thailandica appear to be conspecific, based on combined gene analyses (Fig. 1). When informative sequence data such as tef1-α or rpb2 were added to the analysis for Montagnula chromolaenicola, M. puerensis, M. saikhuensis, and M. thailandica, distinct branches and independent lineages were observed (Fig. 1). This suggests that these species are separate entities. Notably, two sequences of M. donacina (HVVV01 and HFG07004) were found to be monophyletic with the type strain of Montagnula chromolaenicola (MFLUCC 17-1469), indicating that they belong to the latter species. In the case of Montagnula camporesii (MFLUCC 16-1369), M. cirsii (MFLUCC 13-0680), and M. scabiosae (MFLUCC 14-0954), the type strains formed a monophyletic lineage as a single species. Nucleotide base pair comparison of LSU, SSU, and ITS between these three strains did not reveal any differences. Therefore, it is suggested that Montagnula camporesii and M. cirsii should be synonymized under M. scabiosae, as it is the oldest name. However, it is important to note that this taxonomic clarification was not within the scope of our study, and future studies should compare the morphology of the holomorphs to resolve any remaining taxonomic confusion. Apart from these two clades, all other species formed distinct lineages in the multi-gene phylogenetic analysis. Out of the accepted 54 species in this genus, sequence data are currently available for only 28 species, including the four new species introduced in this study. This leaves approximately 48% of the species in need of phylogenetic sorting. Hence, future studies based on taxonomy should prioritize obtaining DNA sequence data for the remaining species. They should aim to acquire informative sequence data, such as tef1-α and rpb2, for all strains, and focus on revising the taxonomy of all species within the genus Montagnula.

Morphological characterization of Montagnula species

The genus Montagnula exhibits rare reporting of its anamorphic features, with only one species, M. cylindrospora, described from its anamorph in addition to our study (Crous et al. 2020). This finding has helped confirm its phylogenetic placement within the genus. The teleomorph, rather than the anamorph, appears to be more commonly observed in the natural environment. The majority of Montagnula species produce immersed or semi-immersed ascomata, which are globose to subglobose in shape and possess a central papillate ostiole. However, there are a few exceptions, such as M. camporesii, M. cirsii, and M. longipes, which have been reported to have superficial ascomata. Upon closer examination, it becomes apparent that Montagnula camporesii and M. cirsii actually have semi-immersed ascomata, as illustrated in Hyde et al. (2016, 2020). It is worth mentioning that Aptroot (1995) did not illustrate the ascomata, and their orientation remains unclear. Additionally, only one species, Montagnula bellevaliae, has been reported to possess an eccentric ostiole (Hongsanan et al. 2015). The peridium cells of Montagnula species commonly exhibit a thick-walled arrangement with a textura angularis pattern. Notably, the cells near the apex are often thicker compared to those on the sides and base walls. A distinguishing characteristic for species within this genus is the presence of swollen cells in pseudoparaphyses. The asci, typically exhibit a cylindrical to clavate shape with a prominent pedicel. Ascospores in Montagnula are predominantly described as ellipsoidal to fusiform, pigmented, and septate. The majority of species (>15) have ascospores with a single septum, while some species, including M. dasylirionis, M. dura, M. infernalis, M. mohavensis, M. phragmospora, M. spinosella, and M. yuccigena, have been reported to possess muriform spores (Du et al. 2023). The remaining species have ascospores with either 3 or 5 septa. A distinct characteristic within the genus is the verruculose surface texture of the ascospores which is neglected by most of the studies. Only Montagnula appendiculata, M. chiangraiensis, and M. chromolaenae have been documented to possess polar appendages (Aptroot 2004; Mapook et al. 2020).

Ecological preferences and worldwide distribution of Montagnula species through culture-dependent studies

The information we gathered from our culture-based investigations revealed that Montagnula species were found on 105 genera in 45 distinct plant families, in 55 countries (Table 1). This highlights the wide ecological range and adaptability of Montagnula species across different hosts and geographic regions. Among the plant families, Poaceae emerged as the most significant contributor, yielding the highest number of isolated Montagnula species (Fig. 4). This finding suggests a potential association between Montagnula species and grasses, indicating the ecological importance of the Poaceae family in the life cycle and development of Montagnula species. Furthermore, Montagnula species were also detected in other plant families, such as Asparagaceae and Fabaceae, indicating their potential interactions with a diverse range of host plants. Among the more than 100 plant genera associated with Montagnula species, Agave (Asparagaceae), Opuntia (Cactaceae), Phoenix (Arecaceae), Ammophila (Poaceae), and Yucca (Asparagaceae) were found to have the greatest number of species, collectively representing 25% of the total count. This highlights the potential preference of Montagnula species for these specific plant genera within their respective families. The analysis of country-wise distribution revealed that India had the highest number of Montagnula entries (Table 1). The majority of these entries were attributed to Montagnula donacina, indicating a wide distribution of this species in India. Among the countries where Montagnula species were reported, China exhibited the highest diversity with nine different species, followed by Italy and the USA with seven different species each. This suggests regional variations in the diversity and distribution of Montagnula species. Interestingly, our study also detected Montagnula species in mushrooms and human skin samples, indicating their presence in alternative sources and potential interactions with other organisms. This highlights the need for further investigation into the ecological roles and potential impacts of Montagnula species in these non-traditional habitats. Except for Antarctica, Montagnula donacina has been reported from various countries across all six continents. Additionally, it has been identified in 25 different plant families. Investigating the reasons behind its wide distribution and adaptation to diverse ecological conditions would be intriguing. Future studies should focus on the morphological features, secondary metabolites, and gene data-based analyses of the species. To date, only six studies, including this one, have provided entries featuring both morphology and DNA-based sequence data evidence (Pitt et al. 2014; Zhao et al. 2018; Ren et al. 2022a; Li et al. 2023; Sun et al. 2023).

These findings elucidate the global distribution and ecological preferences of Montagnula species, highlighting the significance of different sources and plant families in their occurrence and potential ecological interactions. The wide range of sources from which species were identified suggests their adaptability and potential ecological roles in various ecosystems. The study also has important implications for our understanding of the ecology and biology of Montagnula fungi. All of the new species described in this study were found to be associated with dead wood, indicating the role that these fungi play in the decomposition of organic matter in forest ecosystems. We suggest that future studies could investigate the functional roles played by Montagnula fungi in ecosystem processes, such as carbon and nutrient cycling.

Global biogeography and ecological versatility of Montagnula based on metabarcoding data through culture-independent studies (NGS)

In addition to the taxonomic novelties, this study utilized metabarcoding data from the GlobalFungi database (Větrovský et al. 2020) to gain insights into the global diversity and distribution of Montagnula. Metabarcoding is a valuable tool that allows for the rapid identification of multiple species from complex environmental samples, providing confirmation of their presence in specific habitats. The analysis of multiple metabarcoding studies provided comprehensive information on the occurrence and distribution patterns of Montagnula species worldwide. The distribution of Montagnula across diverse biomes underscores their remarkable ecological adaptability and diversity. Forests, constituting 61% of their habitats, emerge as the predominant biome, indicating a strong preference or adaptation of the genus to forest ecosystems. Grasslands, accounting for 18%, also represent a significant habitat, suggesting the versatility in adapting to open and semi-open landscapes of them. Croplands (6%) and shrublands (7%) further exemplify the adaptability of Montagnula, thriving in both cultivated areas and natural, low-vegetation environments. Notably, woodlands and anthropogenic areas, representing 2% and 1% respectively, highlight the ability to exist in moderately wooded areas and regions significantly influenced by human activity. Additionally, their presence in aquatic environments, deserts, and wetlands, each accounting for 1% of their habitats, along with a notable 3% in mangroves, reflects the broad ecological niche of them. The marginal occurrence in tundras (0.1%) suggests a limited but notable ability to survive in extreme cold climates. The presence of Montagnula in such varied biomes underscores its ecological versatility and the importance of diverse habitats in understanding its biogeography.

The presence of Montagnula species has been documented in various regions of Africa, Arctic Ocean, Asia, Australia, Europe, Indian Ocean, North America, Pacific Ocean and South America indicating their widespread occurrence and ecological significance in these areas. In Asia, Montagnula species have been observed in multiple countries, including China, India, Indonesia, Iran, Japan, Malaysia, South Korea, Thailand and others (Suppl. material 1). The diverse range of habitats in these regions, such as freshwater habitats, terrestrial environments, and mountainous areas, offer suitable ecological niches for Montagnula colonization and growth. The detection of Montagnula species in different ecological contexts within Asia suggests their ability to adapt to various local conditions and substrates, contributing to their wide distribution across the continent. For example, in China, Montagnula species have been found in diverse habitats ranging from aquatic environments to forests and grasslands (Suppl. material 1), indicating their adaptability to different ecosystems. This adaptability may be attributed to their ability to utilize a wide range of organic materials as substrates, including decaying plant remains.

Australia also exhibits a notable presence of Montagnula species, indicating their occurrence in diverse habitats throughout the continent (Bissett et al. 2016; Luis et al. 2019; Turner et al. 2019; Gui et al. 2023). The unique ecosystems in Australia, including deserts, rainforests and grasslands, provide opportunities for Montagnula to establish themselves in different ecological niches. The metabarcoding studies were used for various biomes i.e. anthropogenic, aquatic, cropland, desert, forest, grassland, mangrove, shrubland, wetland and woodland (Fig. 3). This highlights the higher presence and distribution of Montagnula in different habitats within Australia. In Europe, Montagnula species have been recorded in several countries, including Austria, Belgium, Czech Republic (highest), Estonia, France, Germany, Italy, Netherlands Slovenia, Sweden Switzerland and Spain (Suppl. material 1). The presence of Montagnula in Europe suggests their ability to adapt to different climates and ecological conditions. This broad distribution across Europe indicates the need for further investigation into the ecological preferences and potential impacts of Montagnula species in this region. For instance, studies in Europe have identified Montagnula species in different habitats, such as anthropogenic, aquatic, cropland, desert, forest, grassland, shrubland, tundra, wetland and woodland (Suppl. material 1). Africa and North America also demonstrates a diverse distribution of Montagnula species, with the majority of records coming from the South Africa, Namibia, Botswana, Zambia, Mozambique, Kenya, Kenya and Ivory Coast in Africa respectively. United States was having the highest number of sampling locations in North America. Comparatively, the occurrences of Montagnula species using metabarcoding data in China, the USA, and European countries are relatively well-documented. However, the rest of the world remains a mystery in terms of Montagnula distribution. For example, the majority of Asia, including India and Russia, lacks metabarcoding data for Montagnula species. This emphasizes the need for more extensive research and data collection to better understand the global distribution of Montagnula and its ecological roles.

Conclusion

Our study on Montagnula species has provided valuable insights into their ecological preferences and global distribution patterns. The findings indicate that these fungi exhibit a wide range of climatic distribution, suggesting their adaptability to different temperature ranges and potentially reducing their vulnerability to climate change. The ability of Montagnula species to utilize a diverse range of organic materials as substrates, including decaying plant remains, contributes to their widespread distribution across various habitats. Our analysis revealed a diverse range of sources from which Montagnula species were detected, including freshwater and terrestrial habitats, further highlighting their ecological versatility. Sediments were found to be particularly rich in Montagnula sequences, suggesting their potential as suitable habitats for colonization and growth. Although moderate sequence similarity was observed across different sources and continents, regional variations in ecological preferences and distribution patterns were evident. The diverse host range observed in our field collections aligns with global meta-barcoding sources, emphasizing the ability of Montagnula species to thrive in various ecosystems. The ecological adaptability and versatility of Montagnula species underscore their success in colonizing diverse habitats. Further research and investigation into their biogeography will contribute to our understanding of their global distribution, ecological roles, and potential impacts on ecosystems. This knowledge is crucial for effective conservation efforts, understanding ecosystem dynamics, and managing ecological balance in different regions.

Acknowledgments

We gratefully thank the Chinese Academy of Sciences for providing molecular laboratory facilities.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

Dhanushka Wanasinghe thanks CAS President’s International Fellowship Initiative (number 2021FYB0005), the National Science Foundation of China (NSFC) under the project code 32150410362, Smart Yunnan Project (Young Scientists) under project code E13K281261 and the Postdoctoral Fund from Human Resources and Social Security Bureau of Yunnan Province. Thilina Nimalrathna expresses gratitude for the support provided by the Belt and Road Chinese Government Scholarship and The Alliance of International Science Organizations (ANSO) Ph.D. scholarship. We also extend our appreciation to the Researchers Supporting Project at King Saud University, Riyadh, Saudi Arabia, for funding this research project (Fund no. RSP2024R784). Jianchu Xu thanks National Natural Science Foundation of China (grant number: 31861143002), the Yunnan Provincial Science and Technology Department (grant number: 202101AS070045), Yunnan Provincial Science and Technology Department (grant number: 202205AM070007) and Yunnan Department of Sciences and Technology of China (grant number: 202302AE090023).

Author contributions

Conceptualization: DNW. Data curation: LQX, DNW. Formal analysis: TKF, DNW, TSN. Investigation: TSN, DNW. Methodology: TSN, DNW. Project administration: PEM, JX. Resources: JX. Supervision: JX, PEM. Writing – original draft: TSN, DNW. Writing – review and editing: PEM, TKF.

Author ORCIDs

Dhanushka N. Wanasinghe https://orcid.org/0000-0003-1759-3933

Thilina S. Nimalrathna https://orcid.org/0000-0002-2368-042X

Li Qin Xian https://orcid.org/0009-0006-4936-9409

Turki KH. Faraj https://orcid.org/0000-0002-6012-8474

Jianchu Xu https://orcid.org/0000-0002-2485-2254

Peter E. Mortimer https://orcid.org/0000-0003-3188-9327

Data availability

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

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Supplementary material

Supplementary material 1 

The biogeography, substrate and habitat affinity of Montagnula inferred from the GlobalFungi database

Dhanushka N. Wanasinghe, Thilina S. Nimalrathna, Li Qin Xian, Turki KH. Faraj, Jianchu Xu, Peter E. Mortimer

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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