New species and additional records of saprobic microfungi associated with Macadamia in China and Thailand and their lignocellulitic enzyme production

Xian Zhang; Samantha C Karunarathna; Saowaluck Tibpromma; Chanokned Senwanna; Jaturong Kumla; Tian-Ye Du; Abdallah M Elgorban; Alanoud T Alfagham; Faten Zubair Filimban; Dong-Qin Dai; Nakarin Suwannarach

doi:10.3897/mycokeys.141.209886

. 2026 Oct 7;141:217–281. doi: 10.3897/mycokeys.141.209886

New species and additional records of saprobic microfungi associated with Macadamia in China and Thailand and their lignocellulitic enzyme production

Xian Zhang ^1,^2,^3,⁴, Samantha C Karunarathna ^2,^3,⁴, Saowaluck Tibpromma ^2,³, Chanokned Senwanna ^4,⁵, Jaturong Kumla ^1,^4,⁵, Tian-Ye Du ⁶, Abdallah M Elgorban ⁷, Alanoud T Alfagham ⁸, Faten Zubair Filimban ⁹, Dong-Qin Dai ^2,³, Nakarin Suwannarach ^1,^4,^5,^✉

¹Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand, Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai, Thailand, https://ror.org/05m2fqn25

²Center for Yunnan Plateau Biological Resources Protection and Utilization & Yunnan International Joint Laboratory of Fungal Sustainable Utilization in South and Southeast Asia, College of Biology and Food Engineering, Qujing Normal University, Qujing 655099, China, Center for Yunnan Plateau Biological Resources Protection and Utilization & Yunnan International Joint Laboratory of Fungal Sustainable Utilization in South and Southeast Asia, College of Biology and Food Engineering, Qujing Normal University, Qujing, China, https://ror.org/02ad7ap24

³Key Laboratory of Yunnan Provincial Department of Education of the Deep-Time Evolution on Biodiversity from the Origin of the Pearl River, Qujing Normal University, Qujing 655011, China, Key Laboratory of Yunnan Provincial Department of Education of the Deep-Time Evolution on Biodiversity from the Origin of the Pearl River, Qujing Normal University, Qujing, China, https://ror.org/02ad7ap24

⁴Center of Excellence in Microbial Diversity and Sustainable Utilization, Chiang Mai University, Chiang Mai 50200, Thailand, Center of Excellence in Microbial Diversity and Sustainable Utilization, Chiang Mai University, Chiang Mai, Thailand, https://ror.org/05m2fqn25

⁵Office of Research Administration, Chiang Mai University, Chiang Mai 50200, Thailand, Office of Research Administration, Chiang Mai University, Chiang Mai, Thailand, https://ror.org/05m2fqn25

⁶Shenzhen Key Laboratory of Microbial Genetic Engineering, College of Life Science and Oceanography, Shenzhen University, Shenzhen 518060, China, Shenzhen Key Laboratory of Microbial Genetic Engineering, College of Life Science and Oceanography, Shenzhen University, Shenzhen, China, https://ror.org/01vy4gh70

⁷Center of Excellence in Biotechnology Research, DSR, King Saud University, Riyadh, 12372, Saudi Arabia, Center of Excellence in Biotechnology Research, DSR, King Saud University, Riyadh, Saudi Arabia, https://ror.org/02f81g417

⁸Department of Botany and Microbiology, College of Science, King Saud University, Riyadh 1l45l, Saudi Arabia, Department of Botany and Microbiology, College of Science, King Saud University, Riyadh, Saudi Arabia, https://ror.org/02f81g417

⁹Division of Plant Sciences, Department of Biological Sciences, Faculty of Sciences, King Abdulaziz University, Jeddah, Saudi Arabia, Division of Plant Sciences, Department of Biological Sciences, Faculty of Sciences, King Abdulaziz University, Jeddah, Saudi Arabia, https://ror.org/02ma4wv74

^✉

Corresponding author: Nakarin Suwannarach (suwan.462@gmail.com)

Academic editor: Xing-Guo Tian

^✉

Corresponding author.

Roles

Xian Zhang: Writing - original draft, Data curation, Formal analysis, Investigation, Methodology, Software, Visualization

Samantha C Karunarathna: Writing - original draft, Writing - review and editing, Data curation, Funding acquisition, Methodology, Validation, Visualization

Saowaluck Tibpromma: Conceptualization, Writing - original draft, Writing - review and editing, Data curation, Investigation, Methodology, Project administration, Software, Supervision, Validation, Visualization

Chanokned Senwanna: Writing - review and editing, Formal analysis, Investigation, Methodology, Resources, Visualization

Jaturong Kumla: Writing - original draft, Data curation, Formal analysis, Investigation, Methodology, Resources, Software, Visualization

Tian-Ye Du: Writing - review and editing, Investigation, Methodology, Visualization

Abdallah M Elgorban: Writing - review and editing

Alanoud T Alfagham: Writing - review and editing

Faten Zubair Filimban: Writing - review and editing

Dong-Qin Dai: Writing - review and editing, Visualization

Nakarin Suwannarach: Conceptualization, Writing - original draft, Writing - review and editing, Data curation, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization

PMCID: PMC13669986 PMID: 42889678

Abstract

Macadamia, a tropical and subtropical plant, is one of the most popular and nutritious nuts worldwide. Previous studies on macadamia-associated fungi have mainly focused on pathogenic and endophytic species, but the diversity and taxonomy of saprobic fungi associated with Macadamia remain poorly characterized. In this study, 18 fungal taxa associated with Macadamia spp. were collected from China and Thailand. Through morphological and multilocus phylogenetic analyses, five new species (Dothiorella lignicola, Lasiodiplodia macadamiae, Palmiascoma macadamiae, Xylochrysis macadamiae, and X. yunnanensis) are introduced, along with 13 previously known species across nine genera that are newly recorded from macadamia. In addition, 20 isolates representing these 18 taxa were evaluated for their potential to produce lignocellulolytic enzymes. Among the four enzymes evaluated, xylanase was the most frequently detected, followed by cellulase and mannanase, whereas laccase production was observed in only a limited number of isolates. These findings expand the known diversity of saprobic microfungi associated with macadamia and highlight their potential ecological roles in lignocellulose decomposition and possible applications in biotechnology.

Key words: Lignocellulose-degrading enzyme, morphology, new taxa, phylogenetic analysis, saprobes

Introduction

Macadamia F. Muell. belongs to an early-diverging eudicot family and is cultivated as a perennial crop, and its nuts are widely known as “the king of nuts” because of their high nutritional value and economic importance (Wrona et al. 2020; Li et al. 2022; Tao et al. 2022; Zhang et al. 2025c). Beyond the production of edible nuts, macadamia has considerable commercial value because its various components and by-products can be utilized in a wide range of applications, including the production of adsorbents, edible oils, beverages, personal care products, cleaning products, and other value-added food products (Mojapelo et al. 2023; Alasalvar et al. 2025). In the last century, macadamia was found in tropical forest fringes from southeastern Queensland to northeastern New South Wales, Australia (Carpenter et al. 2015; Nock et al. 2019; Akinsanmi and Carvalhais 2020). The genus Macadamia includes four species, viz. M. integrifolia, M. jansenii, M. ternifolia, and M. tetraphylla (Tao et al. 2022; Zhang et al. 2025c). Among these four species, M. integrifolia and M. tetraphylla produce edible nuts; however, the other two species (M. jansenii and M. ternifolia) produce small and inedible kernels (Prasannath et al. 2021; Zhang et al. 2025c). Macadamia integrifolia and hybrid cultivars of M. integrifolia and M. tetraphylla are widely planted worldwide (Hardner 2016; Nock et al. 2019). Currently, macadamia nuts are mainly grown in Australia, Brazil, China, Kenya, Malawi, New Zealand, South Africa, South America, Thailand, and the United States (Nock et al. 2019; Twiddy et al. 2024). In China, macadamia was first introduced and planted in the 1970s (Shuai et al. 2022; Tao et al. 2022). In recent years, China has accounted for more than 60% of the world’s cultivated area, with the main plantations located in Guangxi, Guangdong, and Yunnan provinces (Tao et al. 2022; Gong et al. 2024; Zhang et al. 2025b). Among them, Yunnan Province has the largest macadamia plantation area, accounting for 88%, and its harvested area accounts for more than 90% of China’s total (Gong et al. 2024). Meanwhile, Yunnan also has the world’s largest cultivated area (Gong et al. 2024). In Thailand, macadamia was introduced for experimental cultivation in 1953 and was subsequently cultivated in northern Thailand, particularly in Chiang Mai Province. In recent years, macadamia cultivation has expanded considerably in northern Thailand and has become an increasingly important cash crop, driven by growing domestic consumption and its potential for international export (Nilpanich et al. 2025).

Macadamia-growing regions provide favorable habitats for diverse organisms, including fungi, which are among the most prevalent microorganisms associated with macadamia from cultivation through postharvest handling and processing (Wrona et al. 2020; Twiddy et al. 2024; Zhang et al. 2025b). Microfungi, including saprobic, endophytic, and pathogenic forms, are widely distributed worldwide. Saprobic fungi play a very important role as decomposers; they contribute to the carbon, nitrogen, and phosphorus cycles by decomposing organic matter (Treseder and Lennon 2015; Tian et al. 2024a; Du et al. 2025b). Several publications have reported fungi associated with Macadamia spp. (Prasannath et al. 2020, 2021; Li et al. 2022; Tao et al. 2022). For example, Trichoderma hamatum has been isolated as an endophyte from healthy macadamia trees (Lewis et al. 1996; Rabeendran et al. 2006; Mao et al. 2020; Li et al. 2022). In contrast, numerous pathogenic fungi, predominantly belonging to the genera Colletotrichum, Diaporthe, Neopestalotiopsis, and Pseudocercospora, have been reported to cause diseases affecting various parts of macadamia plants, including husks, flowers, leaves, and fruits (Sutton 1991; Denman et al. 2003; Miles et al. 2009; Akinsanmi et al. 2016a, 2016b; Liddle et al. 2018; Jeff-Ego and Akinsanmi 2019; Tan et al. 2019; Prasannath et al. 2020, 2021; Wrona et al. 2020; Li et al. 2022; Mohankumar et al. 2022; Twiddy et al. 2024). However, saprobic fungi associated with Macadamia spp. remain poorly documented compared with endophytic and pathogenic fungi. Only a limited number of saprobic species, including Fusarium polyphialidicum, Melomastia puerensis, Neopalmiascoma macadamiae, and Phaeoacremonium chiangmaiense, have been reported (Ruiz et al. 1997; Zhang et al. 2003; Prasannath et al. 2021; Zhang et al. 2024, 2025b). The fungal life mode of several species has not been reported (Prasannath et al. 2021; CABI 2026).

Saprobic microfungi, particularly those associated with woody substrates, play an essential role in organic matter decomposition and nutrient cycling in ecosystems (Sánchez 2009; Sharma and Arora 2015; Eliopoulos et al. 2025). By secreting extracellular hydrolytic and oxidative lignocellulolytic enzymes, including cellulases, xylanases, laccases, mannanases, manganese peroxidases, and lignin peroxidases, these fungi contribute to the degradation of complex organic compounds, particularly lignocellulosic materials, thereby facilitating nutrient release (Manavalan et al. 2015; Benatti and Polizeli 2023). In addition, saprobic microfungi are of considerable biotechnological interest because they produce lignocellulolytic enzymes with diverse applications in biomass conversion, biofuel production, the food and feed industries, pulp and paper processing, textile production, and environmental bioremediation (Chukwuma et al. 2020; Kumla et al. 2020). However, the diversity and ecological roles of saprobic fungi associated with Macadamia spp. remain insufficiently explored, particularly in tropical and subtropical regions. Therefore, this study aimed to investigate saprobic microfungi associated with macadamia in China and Thailand using morphological and multilocus phylogenetic analyses. Eighteen fungal taxa were obtained, including five new species and 13 species newly recorded from macadamia (Table 1). Moreover, all obtained microfungal strains were evaluated for their ability to produce lignocellulolytic enzymes.

Table 1.

Saprobic microfungi obtained in this study.

*Phylum: Ascomycota* Caval.-Sm**
*Subphylum: Pezizomycotina* O.E. Erikss. & Winka**
Class: *Sordariomycetes* O.E. Erikss. & Winka
Subclass: *Diaporthomycetidae* Senan., Maharachch. & K.D. Hyde
Family: *Woswasiaceae* Huang Zhang, K.D. Hyde & Maharachch
Genus: *Xylochrysis* Réblová
1.	**Xylochrysis macadamiae** X. Zhang, Tibpromma & Karun., sp. nov.
2.	**Xylochrysis yunnanensis** X. Zhang, Tibpromma & Karun., sp. nov.
Subclass: *Pleosporomycetidae* C.L. Schoch, Spatafora, Crous & Shoemaker
Order: *Sporidesmiales* Crous
Family: *Sporidesmiaceae* Summa veg. Scand., Sectio Post
Genus: *Sporidesmium* Link
3.	**Sporidesmium tropicale** M.B. Elli (new host record)
Class: *Dothideomycetes* O.E. Erikss. & Winka
Subclass: *Pleosporomycetidae* C.L. Schoch, Spatafora, Crous & Shoemaker
Order: *Botryosphaeriales* C.L. Schoch, Crous & Shoemaker
Family: *Botryosphaeriaceae* Theiss. & Syd
Genus: *Dothiorella* Sacc.
4.	**Dothiorella lignicola** X. Zhang & N. Suwannar., sp. nov.
Genus: *Lasiodiplodia* Ellis & Everh.
5.	**Lasiodiplodia macadamiae** X. Zhang, Tibpromma & Karun., sp. nov.
6.	**Lasiodiplodia ponkanicola** X.E. Xiao, Crous & H.Y. Li (new host record and geographic record in Thailand)
7.	**Lasiodiplodia endophytica** N.I. de Silva, A.J.L. Phillips & K.D. Hyde (new host record and geographic record in Thailand)
Order: *Pleosporales* Luttr. ex M.E. Barr
Family: *Bambusicolaceae* D.Q. Dai & K.D. Hyde
Genus: *Palmiascoma* Phookamsak & K.D. Hyde
8.	**Palmiascoma macadamiae** X. Zhang, Tibpromma & Karun., sp. nov.
9.	**Palmiascoma chiangraiense** N. Afshari, Lumyong & K.D. Hyde (new host record and geographic record in China)
Genus: *Crassiparies* Matsumura, K. Hiray & Kaz.
10.	**Crassiparies quadrisporus** M. Matsum., K. Hiray. & Kaz. (new host record and geographic record in China)
Family: *Didymosphaeriaceae* Munk
Genus: *Chromolaenicola* Mapook & K.D. Hyde
11.	**Chromolaenicola ananasi** X.G. Tian, K.D. Hyde & Tibpromma (new host record)
Genus: *Montagnula* Berl.
12.	**Montagnula donacina** (Niessl) Wanas., E.B.G. Jones & K.D. Hyde (new host record)
13.	**Montagnula thailandica** Mapook & K.D. Hyde (new host record)
Family: *Macrodiplodiopsidaceae* Voglmayr, Jaklitsch & Crous
Genus: *Pseudochaetosphaeronema* Punith.
14.	**Pseudochaetosphaeronema chiangraiense** Wijes., Boonmee & K.D. Hyde (new host record and geographic record from China)
15.	**Pseudochaetosphaeronema puerense** R.F. Xu, K.D. Hyde & Tibpromma (new host record)
16.	**Pseudochaetosphaeronema siamense** Jayasiri, E.B.G. Jones & K.D. Hyde (new host record and geographic record from China)
Family: *Massarinaceae* Munk
Genus: *Helminthosporium* Link
17.	**Helminthosporium puerense** L. Lu & Tibpromma (new host record)
18.	**Helminthosporium nanjingense** Meng Zhang, Xiao J. Wang & H.Y. Wu (new host record)

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Materials and methods

Sample collection, fungal isolation, and morphological study

Dead and decaying twigs and stems of Macadamia sp. bearing fungal fruiting bodies were collected from macadamia plantations in Yunnan Province, China, and Chiang Mai Province, Thailand, during 2023–2024. Specimens were placed in separate plastic bags and transported to the mycology laboratories at Chiang Mai University, Thailand, and Qujing Normal University, China, respectively. Each bag was labeled with collection details, including altitude, date, site, and global positioning system (GPS) coordinates (Rathnayaka et al. 2025). The fruiting body features of the specimens were observed using an Olympus LEICA S8 APO optical microscope (Tokyo, Japan) and a Nikon SMZ800N stereomicroscope. Micromorphological structures were observed, and photographs were taken using an OLYMPUS BX53 compound microscope (Olympus, Tokyo, Japan) connected to an OLYMPUS DP74 digital camera (Olympus, Tokyo, Japan) or using a Nikon DS-Ri2 camera connected to a Nikon ECLIPSE Ni compound microscope (Tokyo, Japan). Adobe Photoshop CS3 Extended version 10.0 (Adobe Systems, USA) was used to prepare photographic plates, and Tarosoft (R) Image Frame Work (v. 0.9.0.7) was used to measure the microstructures. Pure fungal isolates were obtained by single-spore isolation following the method described by Senanayake et al. (2020). Germinated spores on potato dextrose agar (PDA) were transferred to new PDA plates, and culture characteristics were observed after 2–4 weeks of incubation at 25–28 °C in an incubator.

Specimens were deposited in the Herbarium of Guizhou Medical University (GMB-W) and the Herbarium of the Department of Biology (CMUB), Faculty of Science, Chiang Mai University, Thailand. Living cultures were deposited in the Guizhou Medical University Culture Collection (GMBCC), China, and the Sustainable Development of Biological Resources (SDBR-CMU), Faculty of Science, Chiang Mai University, Thailand. MycoBank numbers (MB) were registered as outlined in MycoBank (2026).

DNA extraction, PCR amplification, and sequencing

Molecular studies were carried out according to the method described by Dissanayake et al. (2020). Genomic DNA was extracted from two-week-old mycelium cultivated on PDA using the DNA Extraction Kit-BSC14S1 (BioFlux, Hangzhou, P.R. China) and the FavorPrep Tissue Genomic DNA Extraction Mini Kit (FAVORGEN Biotech Corp., Taiwan, P.R. China) according to the manufacturers’ instructions. Polymerase chain reactions (PCR) were performed to amplify partial gene regions using the primers and conditions listed in Table 2. The 25 μL PCR mixtures contained the following components: ddH₂O (9.5 μL), 2× Master Mix (12.5 μL) (Biotek Corporation, Beijing, China), DNA template (1 μL), and each forward and reverse primer (1 μL) (Tibpromma et al. 2018; Senwanna et al. 2023; Zhang et al. 2025b). PCR products were checked on 1% agarose gels and sent to Sangon Biotech (Shanghai Co., Ltd.), China, and 1^st BASE Company (Kembangan, Malaysia).

Table 2.

Loci, primers, and PCR conditions used in this study.

Loci	Primers	PCR conditions	References
ITS	ITS4/ITS5	95 °C 3 min, (94 °C 30 s, 55 °C 50 s, 72 °C 90 s) × 35 cycles, 72 °C 10 min	White et al. (1990)
nrLSU	LR5/LR0R	95 °C 3 min, (94 °C 30 s, 55 °C 50 s, 72 °C 90 s) × 35 cycles, 72 °C 10 min	Vilgalys and Hester (1990)
nrSSU	NS1/NS4	95 °C 3 min, (94 °C 30 s, 55 °C 50 s, 72 °C 90 s) × 35 cycles, 72 °C 10 min	White et al. (1990)
tef1-α	983F/2218R	95 °C 3 min, (95 °C 30 s, 55 °C 50 s, 72 °C 90 s) × 35 cycles, 72 °C 10 min	Rehner and Buckley (2005)
tef1-α	728F/986R	95 °C 3 min, (95 °C 30 s, 55 °C 30 s, 72 °C 60 s) × 40 cycles, 72 °C 10 min	Carbone and Kohn (1999)
tub2	T1/Bt2b	94 °C 4 min, (94 °C 40 s, 52 °C 30 s, 72 °C 60 s) × 35 cycles, 72 °C 10 min	Glass and Donaldson (1995); O’Donnell and Cigelnik (1997)
rpb2	fRPB2-5F/fRPB2-7cR	94 °C 5 min, (94 °C 45 s, 55 °C 120 s, 72 °C 180 s) × 40 cycles, 72 °C 10 min	Liu et al. (1999)

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Phylogenetic analyses

All sequences were assembled using Geneious 9.1.8 (https://www.geneious.com), and the newly generated sequences (Table 3) were submitted to BLASTn to search nucleotide databases and identify the most similar taxa. The sequences used in the phylogenetic analyses were downloaded from GenBank, as described in recent publications. Multiple sequence alignments were conducted using the online program MAFFT v. 7.110 (https://mafft.cbrc.jp/alignment/server/). TrimAl.v1.2rev59 (http://trimal.cgenomics.org) was used to automatically modify the alignments (Capella-Gutiérrez et al. 2009). All sequence data were combined using SequenceMatrix v. 1.7.8 (Vaidya et al. 2011). AliView v. 1.28 was used to convert the FASTA files to NEXUS files (Larsson 2014). Multilocus phylogenetic analyses of the combined sequences were conducted using randomized accelerated maximum likelihood (RAxML) and Bayesian inference (BI).

Table 3.

. Voucher specimen data and GenBank accession numbers for sequences generated from this study. “T” isolates represent ex-type, “-” indicates no sequence data in GenBank.

Species name	Strains	Host	Location	GenBank Accession No.
Species name	Strains	Host	Location	ITS	nrLSU	nrSSU	tef1-α	tub2	rpb2
Xylochrysis macadamiae	GMBCC 2440T	Macadamia sp.	Dehong, Yunnan, China	PX921755	PX921759	PX921934	PZ224681	-	PZ210148
Xylochrysis macadamiae	GMBCC 2441	Macadamia sp.	Dehong, Yunnan, China	PX921756	PX921760	PX921935	PZ224682	-	PZ210149
Xylochrysis yunnanensis	GMBCC 2442T	Macadamia sp.	Dehong, Yunnan, China	PX921757	PX921761	-	PZ224679	-	PZ210150
Xylochrysis yunnanensis	GMBCC 2443	Macadamia sp.	Dehong, Yunnan, China	PX921758	PX921762	-	PZ224680	-	PZ210151
Sporidesmium tropicale	GMBCC 2444	Macadamia sp.	Pu’er, Yunnan, China	PZ147879	PZ147883	-	-	-	PZ150683
Sporidesmium tropicale	GMBCC 2445	Macadamia sp.	Pu’er, Yunnan, China	PZ147880	PZ147884	-	-	-	PZ150684
Dothiorella lignicola	SDBR-CMU873T	Macadamia sp.	Doi Saket, Chiang Mai, Thailand	PZ162690	-	-	PZ167135	PZ167137	-
Dothiorella lignicola	SDBR-CMU874	Macadamia sp.	Doi Saket, Chiang Mai, Thailand	PZ162691	-	-	PZ167136	PZ167138	-
Lasiodiplodia macadamiae	GMBCC 2446T	Macadamia sp.	Pu’er, Yunnan, China	PZ158349	-	-	PZ150673	PZ150677	-
Lasiodiplodia macadamiae	GMBCC 2447	Macadamia sp.	Pu’er, Yunnan, China	PZ158350	-	-	PZ150674	PZ150678	-
Lasiodiplodia ponkanicola	CMUB 40122	Macadamia sp.	Samoeng Tai, Chiang Mai, Thailand	PZ158351	-	-	PZ150675	PZ150679	-
Lasiodiplodia ponkanicola	SDBR-CMU875	Macadamia sp.	Samoeng Tai, Chiang Mai, Thailand	PZ158352	-	-	PZ150676	PZ150680	-
Lasiodiplodia endophytica	CMUB 40123	Macadamia sp.	Samoeng Tai, Chiang Mai, Thailand	PZ158353	-	-	QB037503	PZ150681	-
Lasiodiplodia endophytica	SDBR-CMU876	Macadamia sp.	Samoeng Tai, Chiang Mai, Thailand	PZ158354	-	-	QB037504	PZ150682	-
Palmiascoma macadamiae	GMBCC 2448T	Macadamia sp.	Dehong, Yunnan, China	PX929868	PX929872	PX929876	-	-	PX931730
Palmiascoma macadamiae	GMBCC 2449	Macadamia sp.	Dehong, Yunnan, China	PX929869	PX929873	PX929877	-	-	PX931731
Palmiascoma chiangraiense	GMBCC 2450	Macadamia sp.	Pu’er, Yunnan, China	PX929870	PX929874	PX929878	-	-	PX931728
Palmiascoma chiangraiense	GMBCC 2451	Macadamia sp.	Pu’er, Yunnan, China	PX929871	PX929875	PX929879	-	-	PX931729
Crassiparies quadrisporus	GMBCC 2452	Macadamia sp.	Dehong, Yunnan, China	PX931083	PX931114	PX931117	PX931734	-	PX931732
Crassiparies quadrisporus	GMBCC 2453	Macadamia sp.	Dehong, Yunnan, China	PX931084	PX931115	PX931118	PX931735	-	PX931733
Chromolaenicola ananasi	CMUB 40120	Macadamia sp.	Doi Saket, Chiang Mai, Thailand	PZ151946	PZ151950	PZ151952	PZ210152	-	-
Chromolaenicola ananasi	SDBR-CMU872	Macadamia sp.	Doi Saket, Chiang Mai, Thailand	PZ151947	PZ151951	PZ151953	PZ210153	-	-
Montagnula donacina	GMBCC 2455	Macadamia sp.	Honghe, Yunnan, China	PX945742	PX945744	PX945756	PX957010	-	-
Montagnula thailandica	GMBCC 2454	Macadamia sp.	Honghe, Yunnan, China	PX945741	PX945743	PX945755	PX957009	-	-
Pseudochaetosphaeronema chiangraiense	GMBCC 2456	Macadamia sp.	Dehong, Yunnan, China	PX946142	PX946148	PX946154	PX957003	-	-
Pseudochaetosphaeronema chiangraiense	GMBCC 2457	Macadamia sp.	Dehong, Yunnan, China	PX946143	PX946149	PX946155	PX957004	-	-
Pseudochaetosphaeronema puerense	GMBCC 2458	Macadamia sp.	Dehong, Yunnan, China	PX946144	PX946150	PX946156	PX957005	-	-
Pseudochaetosphaeronema puerense	GMBCC 2459	Macadamia sp.	Dehong, Yunnan, China	PX946145	PX946151	PX946157	PX957006	-	-
Pseudochaetosphaeronema siamense	GMBCC 2460	Macadamia sp.	Dehong, Yunnan, China	PX946146	PX946152	PX946158	PX957007	-	-
Pseudochaetosphaeronema siamense	GMBCC 2461	Macadamia sp.	Dehong, Yunnan, China	PX946147	PX946153	PX946159	PX957008	-	-
Helminthosporium puerense	GMBCC 2462	Macadamia sp.	Pu’er, Yunnan, China	PX946836	PX946845	PX946841	PZ150669	-	PZ150665
Helminthosporium puerense	GMBCC 2464	Macadamia sp.	Pu’er, Yunnan, China	PX946837	PX946846	PX946842	PZ150670	-	PZ150666
Helminthosporium nanjingense	GMBCC 2466	Macadamia sp.	Pu’er, Yunnan, China	PX946838	PX946847	PX946843	PZ150671	-	PZ150667
Helminthosporium nanjingense	GMBCC 2467	Macadamia sp.	Pu’er, Yunnan, China	PX946839	PX946848	PX946844	PZ150672	-	PZ150668

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Maximum likelihood analyses were conducted on the online CIPRES Science Gateway portal (https://www.phylo.org/portal2/login!input.action; Miller et al. 2010) using the RAxML-HPC2 on XSEDE (8.2.12) tool with 1,000 bootstrap (BS) replicates (Stamatakis 2006, 2014; Stamatakis et al. 2008) and IQ-TREE v. 3.1.4 (https://www.hiv.lanl.gov/content/sequence/IQTREE/iqtree.html) (Nguyen et al. 2015). BI was performed using the MrBayes on XSEDE (3.2.7a) tool (Ronquist et al. 2012) and MrBayes v. 3.2.6 (Ronquist et al. 2012), and the best models of evolution were estimated using MrModelTest v. 2.3 (Nylander et al. 2008). Six simultaneous Markov chains were run for 2,000,000–100,000,000 generations (depending on the fungal genera), and trees were sampled every 100 generations (Lu et al. 2025b). The run was automatically terminated when the standard deviation of split frequencies dropped below 0.01, with a 25% burn-in. The phylogenetic trees were visualized using FigTree v. 1.4.2 (Rambaut 2012) and further edited in Microsoft PowerPoint 2021 and Adobe Photoshop CS3 Extended version 22.0.0 (Adobe Systems, California, USA). Maximum likelihood bootstrap support (MLBS) ≥ 60% and Bayesian posterior probabilities (BYPP) ≥ 0.90 are shown above the nodes.

Evaluation of lignocellulolytic enzyme production

The fungal isolates obtained in this study were used to investigate lignocellulolytic enzyme production using agar plate assays. The production of four lignocellulolytic enzymes, namely cellulase, xylanase, mannanase, and laccase, was investigated following the methods described in previous studies (Kasana et al. 2008; Kalim and Ali 2016). Carboxymethyl cellulose (CMC) agar (10 g/L CMC, 2 g/L NaNO₃, 1 g/L K₂HPO₄, 0.5 g/L MgSO₄, 0.5 g/L KCl, 0.2 g/L peptone, 17 g/L agar, pH 5.5), xylan agar (10 g/L xylan, 3 g/L yeast extract, 1.5 g/L peptone, 3.5 g/L NaCl, 1 g/L NaNO₃, 1 g/L KH₂PO₄, 0.3 g/L MgSO₄, 20 g/L agar, pH 5.5), locust bean gum (LBG) agar (10 g/L LBG, 2 g/L NaNO₃, 1 g/L K₂HPO₄, 0.5 g/L MgSO₄, 0.5 g/L KCl, 0.2 g/L peptone, 17 g/L agar, pH 5.5), and guaiacol agar (0.02% guaiacol, 2 g/L NaNO₃, 1 g/L K₂HPO₄, 0.5 g/L MgSO₄, 0.5 g/L KCl, 0.2 g/L peptone, 17 g/L agar, pH 5.5) were used to screen for cellulase, xylanase, mannanase, and laccase production, respectively. A 5-mm-diameter mycelial plug taken from a 5-day-old PDA culture of each fungal isolate was inoculated onto the center of each tested agar medium. After 1 week of incubation at 30 °C, the colony diameter was measured. The plates were then flooded with 0.1% Congo red solution for 1 min and washed three times with 15 mL of 1 M NaCl for 15 min each. The diameter of the clear halo zone surrounding the fungal colony was measured as an indicator of positive cellulase, xylanase, and mannanase production (Kasana et al. 2008; Kalim and Ali 2016; Nirmalasari et al. 2022). In contrast, the appearance of a brownish-red zone surrounding the fungal colony on guaiacol agar was used as an indicator of positive laccase production (Yuliana et al. 2024). The production of all four lignocellulolytic enzymes was evaluated using the enzyme potency index (EPI), calculated as the difference between the halo zone diameter or brownish-red zone diameter and the colony diameter, divided by the colony diameter (Sánchez-Corzo et al. 2021; Luong et al. 2025). All experiments were performed in triplicate.

Statistical analysis

Statistical analyses were performed using one-way analysis of variance (ANOVA) with SPSS Statistics version 27.0.1 (IBM Corp., Armonk, NY, USA). Differences among treatment means were considered statistically significant at p ≤ 0.05 and were compared using Duncan’s multiple range test.

Results

Based on molecular and morphological analyses, five new species and 13 new records of saprobic microfungi associated with macadamia in China and Thailand were identified. Multilocus phylogenetic analyses were conducted to determine the placement of species within the appropriate orders, families, and genera. Full descriptions and photographic plates illustrating the morphological characteristics were provided for all new species. For new records, descriptions and photographic plates were provided only for taxa exhibiting additional morphological characteristics compared with those reported in previous studies.

Phylogenetic analyses

Analysis 1: The phylogenetic analysis of the genus Xylochrysis was conducted based on a combined dataset of nrLSU, nrSSU, ITS, tef1-α, and rpb2 sequence data following Xiao and Liu (2025). A total of 28 strains were included in the combined sequence analysis, which comprised 4,380 characters with gaps. Pleurostoma repens (CBS 294.39) was used as the outgroup taxon. The ML tree topology was similar to the BI topology. The best-scoring RAxML tree with a final likelihood value of −22144.489 is presented. The matrix contained 1,714 distinct alignment patterns, with 39.82% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.242119, C = 0.259806, G = 0.279643, T = 0.218433; substitution rates AC = 1.000000, AG = 1.888192, AT = 1.000000, CG = 1.000000, CT = 4.993208, GT = 1.000000; proportion of invariable sites I = 0.642117; gamma distribution shape parameter α = 0.738000. The ML and BI trees showed identical topologies. Therefore, only the ML tree is presented in Fig. 1. The phylogenetic tree showed that the Xylochrysis clade was strongly supported (100% MLBS, 1.00 BYPP) and was clearly separated from the other genera. The four Xylochrysis isolates obtained in this study formed two distinct monophyletic clades. The two isolates, GMBCC 2442 and GMBCC 2443, formed a distinct and well-supported lineage (100% MLBS, 1.00 BYPP), representing a new species, X. yunnanensis. The two isolates GMBCC 2440 and GMBCC 2441 formed a distinct lineage closely related to X. hydei and X. chiangraiensis (100% MLBS, 1.00 BYPP) and are introduced herein as a new species, X. macadamiae.

Figure 1. — Phylogram generated from maximum likelihood analysis based on a combined dataset of nrLSU, nrSSU, ITS, *tef1-α*, and *rpb2* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown at the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 2: The phylogenetic analysis of the species of Sporidesmium was conducted based on a combined dataset of ITS, nrLSU, and rpb2 sequence data following Zhang et al. (2025a). A total of 30 strains were included in the combined sequence analysis, which comprised 3,481 characters with gaps. Tracylla aristata (CBS 141404) and Tracylla aristata (CPC 25500) were used as the outgroup taxa. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 2. The best-scoring RAxML tree with a final likelihood value of −15496.102001 is presented. The matrix contained 1,133 distinct alignment patterns, with 40.82% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.242664, C = 0.253299, G = 0.292907, T = 0.211130; substitution rates AC = 1.542893, AG = 2.733518, AT = 1.615427, CG = 0.991797, CT = 7.194628, GT = 1.000000; proportion of invariable sites I = 0.345021; gamma distribution shape parameter α = 0.533110. The phylogenetic tree showed that the two isolates GMBCC 2444 and GMBCC 2445 were nested within S. tropicale (GMB 5105, GMB 5109, DLUCC 1689, MFLU 17-0850, MFLUCC 17-0344) with 89% MLBS statistical support. Therefore, these two isolates are identified as S. tropicale.

Figure 2. — Phylogram generated from maximum likelihood analysis based on a combined dataset of ITS, nrLSU, and *rpb2* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 3: The phylogenetic analysis of the genus Dothiorella was conducted based on a combined dataset of ITS, tef1-α, and rpb2 sequence data following Zhou et al. (2025). A total of 30 strains were included in the combined sequence analysis, which comprised 1,439 characters with gaps. Neofusicoccum luteum (CBS 562.92) and Neofusicoccum parvum (CMW 9081) were used as the outgroup taxa. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 3. The best-scoring RAxML tree with a final likelihood value of −9878.969843 is presented. The matrix contained 745 distinct alignment patterns, with 23.50% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.209667, C = 0.306778, G = 0.250584, T = 0.232970; substitution rates AC = 1.108033, AG = 2.427500, AT = 1.299719, CG = 1.089085, CT = 4.472302, GT = 1.000000; proportion of invariable sites I = 0.464341; gamma distribution shape parameter α = 0.665543. The phylogenetic analysis showed that the two strains, SDBR-CMU873 and SDBR-CMU874, clustered within the genus Dothiorella and formed a distinct lineage with strong statistical support (100% MLBS, 0.97 BYPP). They were closely related to a clade comprising D. albiziae (MFLUCC 22-0057), D. asteracearum (MFLUCC 25-0275), D. dulcispinae (CBS 121764, CBS 130413), D. oblonga (CBS 121765, CBS 121766), and D. thailandica (CBS 133991), with strong statistical support (99% MLBS, 1.00 BYPP). Therefore, these two isolates are described as a new species, D. lignicola.

Figure 3. — Phylogram generated from maximum likelihood analysis based on a combined dataset of ITS, *tef1-α*, and *rpb2* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 4: The phylogenetic analysis of the genus Lasiodiplodia was conducted using a combined dataset of ITS, tef1-α, and tub2 sequence data following Zhou et al. (2024) and Zhang et al. (2026). A total of 127 strains were included in the combined sequence analysis, which comprised 1,628 characters with gaps. Diplodia mutila (CBS 136014) was used as the outgroup taxon. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 4. The best-scoring RAxML tree with a final likelihood value of −7361.068753 is presented. The matrix contained 532 distinct alignment patterns, with 27.44% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.210377, C = 0.306283, G = 0.256092, T = 0.228248; substitution rates AC = 1.000000, AG = 2.987354, AT = 1.000000, CG = 1.000000, CT = 4.262328, GT = 1.000000. A total of six fungal isolates obtained in this study were placed within the genus Lasiodiplodia. The two isolates, CMUB 40122 and SDBR-CMU875, clustered with L. ponkanicola and formed a well-supported lineage (78% MLBS) and were therefore identified as L. ponkanicola. In addition, the two isolates, CMUB 40123 and SDBR-CMU876, clustered with L. endophytica strain MFLUCC 18-1121 with 73% MLBS statistical support and were therefore identified as L. endophytica. The remaining two isolates, GMBCC 2446 and GMBCC 2447, formed a distinct, well-supported lineage (98% MLBS, 1.00 BYPP) and are proposed as a new species, L. macadamiae.

Figure 4. — Phylogram generated from maximum likelihood analysis based on a combined dataset of ITS, *tef1-α*, and *tub2* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 5: The phylogenetic analysis of the species of Palmiascoma was conducted based on a combined dataset of nrLSU, ITS, nrSSU, and rpb2 sequence data following Sun et al. (2025). A total of 30 strains were included in the combined sequence analysis, which comprised 3,298 characters with gaps. Murilentithecium clematidis (MFLUCC 14-0562) and Murilentithecium lonicerae (MFLUCC 18-0675) were used as the outgroup taxa. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 5. The best-scoring RAxML tree with a final likelihood value of −13933.604 is presented. The matrix contained 951 distinct alignment patterns, with 13.73% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.250562, C = 0.250730, G = 0.250256, T = 0.250453; substitution rates AC = 1.372348, AG = 3.498633, AT = 1.000000, CG = 1.372347, CT = 7.158491, GT = 1.000000; proportion of invariable sites I = 0.589309; gamma distribution shape parameter α = 0.546238. The phylogenetic tree showed that the Palmiascoma clade was strongly supported (86% MLBS, 0.99 BYPP) and was clearly separated from the other genera. The four Palmiascoma isolates obtained in this study formed two distinct monophyletic clades. The two isolates GMBCC 2448 and GMBCC 2449 formed a distinct and well-supported lineage (99% MLBS, 1.00 BYPP), representing a new species, P. macadamiae. The other two isolates, GMBCC 2450 and GMBCC 2451, formed a well-supported lineage with P. chiangraiense (MFLUCC 24-0546) and represent a new host record of P. chiangraiense.

Figure 5. — Phylogram generated from maximum likelihood analysis based on a combined dataset of nrLSU, ITS, nrSSU, and *rpb2* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 6: The phylogenetic analysis of the genus Crassiparies was conducted using a combined dataset of ITS, nrLSU, nrSSU, tef1-α, and rpb2 sequences following Lu et al. (2022). A total of 43 strains were included in the combined sequence analysis, which comprised 4,432 characters with gaps. Cyclothyriella rubronotata (CBS 141486) was used as the outgroup taxon. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 6. The best-scoring RAxML tree with a final likelihood value of −21428.969985 is presented. The matrix contained 1,524 distinct alignment patterns, with 42.07% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.249122, C = 0.237069, G = 0.268948, T = 0.244860; substitution rates AC = 1.714269, AG = 3.905301, AT = 1.953992, CG = 1.108375, CT = 7.859984, GT = 1.000000; proportion of invariable sites I = 0.479072; gamma distribution shape parameter α = 0.580554. The phylogenetic tree showed that the two isolates obtained in this study (GMBCC 2452 and GMBCC 2453) were nested within C. quadrisporus strains (MAFF 245408, MAFF 245408, MFLUCC 18-0298, and MFLUCC 21-0148) and formed a well-supported lineage (100% MLBS, 1.00 BYPP). Therefore, these two isolates are identified as C. quadrisporus.

Figure 6. — Phylogram generated from maximum likelihood analysis based on a combined dataset of ITS, nrLSU, nrSSU, *tef1-α*, and *rpb2* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 7: The phylogenetic analysis of species within the genus Chromolaenicola was conducted using a combined dataset of ITS, nrLSU, nrSSU, tef1-α, and rpb2 sequence data following Du et al. (2025a). A total of 30 strains were included in the combined sequence analysis, which comprised 3,351 characters with gaps. Paramassariosphaeria anthostomoides (CBS 615.86) and Paramassariosphaeria clematidicola (MFLU 16-0172) were used as the outgroup taxa. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 7. The best-scoring RAxML tree with a final likelihood value of −7173.931452 is presented. The matrix contained 382 distinct alignment patterns, with 25.28% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.232658, C = 0.257851, G = 0.270185, T = 0.239307; substitution rates AC = 1.197274, AG = 1.631475, AT = 1.105977, CG = 0.799163, CT = 5.824551, GT = 1.000000; proportion of invariable sites I = 0.674366; gamma distribution shape parameter α = 0.709915. The multilocus phylogenetic analysis showed that the two isolates obtained in this study (CMUB 40120 and SDBR-CMU872) were nested with Ch. ananasi (MFLU 23-0167 and MFLUCC 23-0111) in a well-supported lineage (100% MLBS, 1.00 BYPP). Therefore, they are identified as Ch. ananasi.

Figure 7. — Phylogram generated from maximum likelihood analysis based on a combined dataset of ITS, nrLSU, nrSSU, *tef1-α*, and *rpb2* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 8: The phylogenetic analysis of Montagnula species was conducted using a combined dataset of ITS, nrLSU, nrSSU, and tef1-α sequences following Wanasinghe et al. (2024). Seventy-one strains were included in the combined sequence analysis, which comprised 3,324 characters with gaps. Neokalmusia jonahhulmei (KUMCC 21-0818T) and Neokalmusia jonahhulmei (KUMCC 21-0819) were used as the outgroup taxa. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 8. The best-scoring RAxML tree with a final likelihood value of −11807.011024 is presented. The matrix contained 800 distinct alignment patterns, with 16.94% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.239518, C = 0.258465, G = 0.270946, T = 0.231072; substitution rates AC = 1.857369, AG = 3.055749, AT = 1.470808, CG = 1.117958, CT = 9.363964, GT = 1.000000; proportion of invariable sites I = 0.593192; gamma distribution shape parameter α = 0.662679. Multilocus phylogenetic analysis showed that one isolate, GMBCC 2454, was nested within M. thailandica in a well-supported lineage (84% MLBS, 0.96 BYPP); another isolate, GMBCC 2455, clustered with M. donacina with 81% ML bootstrap support. Therefore, they are identified as M. thailandica and M. donacina.

Figure 8. — Phylogram generated from maximum likelihood analysis based on a combined dataset of ITS, nrLSU, nrSSU, and *tef1-α* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 9: The phylogenetic analysis of Pseudochaetosphaeronema species was conducted using a combined dataset of ITS, nrLSU, nrSSU, and tef1-α sequences following Apurillo et al. (2025). Forty-five strains were included in the combined sequence analysis, which comprised 3,387 characters with gaps. Sulcatispora berchemiae (KUMCC 21-0823) and Sulcatispora berchemiae (KUMCC 21-0824) were used as the outgroup taxa. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 9. The best-scoring RAxML tree with a final likelihood value of −11641.241405 is presented. The matrix contained 758 distinct alignment patterns, with 12.56% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.237749, C = 0.251987, G = 0.269533, T = 0.240732; substitution rates AC = 1.091671, AG = 3.413617, AT = 2.115619, CG = 1.229077, CT = 8.686784, GT = 1.000000; proportion of invariable sites I = 0.632918; gamma distribution shape parameter α = 0.512730. The phylogenetic tree showed that the Pseudochaetosphaeronema clade was strongly supported (93% MLBS, 1.00 BYPP). Six Pseudochaetosphaeronema isolates from this study fell into three distinct clades corresponding to Pse. siamense, Pse. puerense, and Pse. chiangraiense, each with strong statistical support. Therefore, they are identified as Pse. siamense (GMBCC 2460 and GMBCC 2461), Pse. puerense (GMBCC 2458 and GMBCC 2459), and Pse. chiangraiense (GMBCC 2456 and GMBCC 2457).

Figure 9. — Phylogram generated from maximum likelihood analysis based on a combined dataset of ITS, nrLSU, nrSSU, and *tef1-α* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Analysis 10: The phylogenetic analysis of Helminthosporium species was conducted using a combined dataset of nrSSU, nrLSU, ITS, rpb2, and tef1-α sequences following Liao et al. (2025). One hundred and twenty-five strains were included in the combined sequence analysis, which comprised 5,111 characters with gaps. Periconia pseudodigitata (CBS 139699) was used as the outgroup taxon. The ML tree topology was similar to the BI topology. Therefore, only the ML tree is presented in Fig. 10. The best-scoring RAxML tree with a final likelihood value of −36400.855198 is presented. The matrix contained 2,133 distinct alignment patterns, with 38.91% of characters undetermined or missing. Estimated base frequencies were as follows: A = 0.239473, C = 0.252773, G = 0.270386, T = 0.237368; substitution rates AC = 1.540445, AG = 4.461885, AT = 1.674575, CG = 0.965087, CT = 7.862411, GT = 1.000000; proportion of invariable sites I = 0.491607; gamma distribution shape parameter α = 0.496098. The multilocus phylogenetic analysis showed that four isolates obtained in this study belonged to the genus Helminthosporium. Phylogenetically, isolates GMBCC 2462 and GMBCC 2464 clustered with H. puerense (KUNCC 24-18347) and are identified herein as H. puerense. Isolates GMBCC 2466 and GMBCC 2467 clustered with H. nanjingense (HHAUF020380) in a highly supported lineage (89% MLBS, 0.99 BYPP) and are identified herein as H. nanjingense.

Figure 10. — Phylogram generated from maximum likelihood analysis based on a combined dataset of nrSSU, nrLSU, ITS, *rpb2*, and *tef1-α* sequence data. The ML tree topology was similar to the BI topology. ML bootstrap support values of 60% or greater and BYPP values of 0.90 or greater are shown above the nodes. All type strains are shown in bold black, and newly generated sequences are shown in red.

Taxonomy

Xylochrysis macadamiae

X. Zhang, Tibpromma & Karun. sp. nov.

0B8AEBC2-AEA6-5F46-9E73-16A5E1219FA8

863022

Figs 1 , 11

Figure 11. — *Xylochrysis* *macadamiae* (GMB-W 1561, holotype). **a–d**. Appearance of ascomata on substrate; **e, f**. Vertical sections of ascomata; g. Peridium; h. Hamathecium; **i–l**. Asci; **m–o**. Ascospores; p. Germinated ascospore; **q, r**. Culture on PDA from above (q) and below (r). Scale bars: 200 μm (e); 100 μm (f); 50 μm (i); 20 μm (**g, h, j–l**); 10 μm (p); 5 μm (**m–o**); 2 cm (**q, r**).

Etymology.

“macadamiae” refers to the host plant genus “Macadamia.”

Diagnosis.

Xylochrysis macadamiae differs from X. chiangraiensis and X. hydei by its rough-walled ascospores, distinct nrLSU, nrSSU, ITS, tef1-α, and rpb2 sequences, and distinct phylogenetic position.

Holotype.

GMB-W 1561.

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morph: Ascomata (including necks) 435–695 × 235–360 µm (x̄ = 528 × 307 µm, n = 10), scattered or aggregated or confluent into small groups, semi-immersed, globose to subglobose to pyriform, uniloculate, nonstromatic, glabrous at neck, not surrounded by a pigmented layer of cells. Ostioles 95–165 × 25–70 µm (x̄ = 133 × 50 µm, n = 10), black, glabrous, papillate, shiny, filled with hyaline periphyses. Peridium multi-layered, with outer layers 20–70 µm thick, carbonized, composed of brown to dark brown cells of textura prismatica, with inner layers 12–34 µm, thin, composed of flattened, hyaline cells of textura prismatica. Hamathecium 3.5–6.0 µm wide (x̄ = 4.6 μm, n = 40), composed of abundant, narrowly cylindrical to filiform, hyaline, branched, septate, guttulate, paraphyses slightly constricted at septa, not embedded in a gelatinous matrix. Asci (65–) 80–135 (–215) × 3.5–4.5 µm (x̄ = 106 × 4 µm, n = 40), 8-spored, unitunicate, cylindrical, long pedicellate, clear apical apparatus, tapering to the bulbous base, with a bulbous long pedicel, apically rounded. Ascospores 4.5–6.5 × 2–3.5 µm (x̄ = 5.6 × 2.6 µm, n = 40), uniseriate, hyaline, oval to ellipsoid, straight, aseptate, 2-guttulate when mature, rough-walled, without mucilaginous sheath or appendages. Asexual morph: Not observed.

Culture characteristics.

Ascospores germinating on PDA within 24 h at 28 °C, with colonies on PDA reaching 3 cm in diam. after 3 weeks at 28 °C, with a rough surface, filamentous, hairy, slightly raised in the middle, cottony, yellow-brown at the edge, becoming pale yellow toward the center; on the reverse side, as above, light yellow at the margins, turning light brown toward the center, wrinkled, and rarely longitudinally wrinkled in the first circle.

Material examined.

China • Yunnan Province, Dehong City, on dead twigs of Macadamia sp., 16 April 2024, Xian Zhang, MDR1 (GMB-W 1561), holotype; ex-type living culture, GMBCC 2440, other living culture GMBCC 2441.

Notes.

In the phylogenetic analyses, the isolates of Xylochrysis macadamiae clustered within Xylochrysis, sister to X. hydei (GMB-W 1199, holotype, and GMB-W 1200) and X. chiangraiensis (MFLUCC 24-0550, ex-type), with 100% MLBS and 1.00 BYPP statistical support (Fig. 1). Morphologically, X. macadamiae resembles X. chiangraiensis (MFLU 24-0470, holotype) and X. hydei (GMB-W 1199, holotype) in having 8-spored, unitunicate, long-pedicellate asci tapering to a bulbous base and hyaline ascospores. However, X. macadamiae differs from X. chiangraiensis by having larger (435–695 × 235–360 µm vs. 138–282 × 130–216 μm), semi-immersed ascomata, a peridium composed of textura prismatica cells, and rough-walled ascospores, whereas X. chiangraiensis has smaller, immersed ascomata, a peridium composed of textura angularis cells, and smooth- and thin-walled ascospores (Afshari et al. 2025b). Xylochrysis macadamiae differs from X. hydei by having globose to subglobose to pyriform ascomata with long ostioles and oval to ellipsoid, rough-walled ascospores, whereas X. hydei has apically rounded asci with an apical ring and ellipsoidal ascospores with thin and smooth walls (Hongsanan et al. 2025). Based on nucleotide comparisons, X. macadamiae differs from X. hydei (GMB-W 1199) by 12/824 bp (1.5%, without gaps) in nrLSU, 38/545 bp (7.0%, without gaps) in ITS, 110/933 bp (11.8%, without gaps) in tef1-α, and 113/1112 bp (10.2%, without gaps) in rpb2, and differs from X. chiangraiensis (MFLUCC 24-0550) by 15/825 bp (1.82%, without gaps) in nrLSU, 48/526 bp (9.13%, without gaps) in ITS, and 139/1112 bp (12.5%, without gaps) in rpb2. Therefore, Xylochrysis macadamiae is introduced here as a new species in Xylochrysis, collected from Macadamia sp. in China.

Xylochrysis yunnanensis

X. Zhang, Tibpromma & Karun. sp. nov.

94E13853-5A17-5D35-9ECF-B195E4494EA6

863023

Figs 1 , 12

Figure 12. — *Xylochrysis* *yunnanensis* (GMB-W 1562, holotype). **a–c**. Appearance of ascomata on substrate; d. Vertical section of ascoma; e. Peridium; f. Hamathecium; **g–k**. Asci; **l–n**. Ascospores; o. Germinated ascospores; **p, q**. Culture on PDA from above (p) and below (q). Scale bars: 100 μm (d); 30 μm (**f–k**); 10 μm (**l–o**); 1 cm (**p, q**).

Etymology.

The species epithet refers to the type locality, Yunnan, China.

Diagnosis.

Xylochrysis yunnanensis differs from X. aquatica, X. guttulata, and X. lucida by its oval to ellipsoid ascospores, distinct nrLSU, nrSSU, ITS, tef1-α, and rpb2 sequences, and distinct phylogenetic position.

Holotype.

GMB-W 1562.

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morph: Ascomata 150–225 × 150–320 µm (x̄ = 193 × 238 µm, n = 10), scattered, semi-immersed, subglobose to irregular, glabrous, uniloculate, nonstromatic, short ostiolate. Peridium 15–55 µm (x̄ = 31 × 20 µm, n = 10), multi-layered, with outer layers thick, carbonized, composed of brown to dark brown cells of textura angularis, with inner layers thin, composed of flattened, hyaline cells of textura prismatica. Hamathecium 2.5–6.0 µm wide (x̄ = 4.2 μm, n = 30), composed of abundant, narrowly cylindrical to filiform, hyaline, unbranched, septate with slightly constricted, guttulate, paraphyses not embedded in a gelatinous matrix. Asci 90–140 × 5.5–7.5 µm (x̄ = 113 × 6.3 µm, n = 30), 8-spored, unitunicate, cylindrical, long pedicellate, tapering to a bulbous base, apically rounded, with a clear apical apparatus. Ascospores 6–8.5 (–11) × 3.5–5.0 µm (x̄ = 7.3 × 4 µm, n = 45), uniseriate, hyaline, oval to ellipsoid, aseptate or 1- septate, 1–2-guttulate, straight, without mucilaginous sheath or appendages. Asexual morph: Not observed.

Culture characteristics.

Ascospores germinating on PDA within 24 h at 28 °C, producing colonies on PDA reaching 1.5 cm in diam. after 3 weeks at 28 °C, with occasional orange pigmentation diffusing into the medium. Colonies with a rough surface, filamentous, hairy, irregular, raised, cottony, and white-yellow from above; light yellow to light brown and irregular in reverse.

Materials examined.

China • Yunnan Province, Dehong City, on dead twigs of Macadamia sp., 24 April 2024, Xian Zhang, MYD5 (GMB-W 1562), holotype; ex-type living culture, GMBCC 2442, other living culture GMBCC 2443.

Notes.

In the phylogenetic analyses, the isolates of Xylochrysis yunnanensis were placed as an independent clade within Xylochrysis, with 100% MLBS and 1.00 BYPP statistical support (Fig. 1). The collection shares similar morphology with X. guttulata, X. lucida, and X. aquatica in having semi-immersed ascomata, 8-spored, cylindrical asci, and hyaline, 2-guttulate ascospores (Réblová et al. 2014; Song et al. 2023; Xiao and Liu 2025). However, X. yunnanensis can be distinguished from X. guttulata (HKAS 135169, holotype) by having unbranched, septate paraphyses slightly constricted at the septa, wider (90–140 × 5.5–7.5 µm vs. 70–110 × 3.5–4.5 μm) asci, and aseptate or 1-septate, oval to ellipsoid ascospores, whereas X. guttulata has a hamathecium with branched paraphyses and lanceolate to oval ascospores (Xiao and Liu 2025). It differs from X. lucida (PRM 922621, holotype) by having wider paraphyses (2.5–6.0 µm vs. 2.5–3.0 µm), bulbous, long-pedicellate asci, and oval to ellipsoid, aseptate or 1-septate ascospores, whereas X. lucida has ellipsoidal and unicellular ascospores (Réblová et al. 2014). It also differs from X. aquatica (HKAS 124019, holotype) by having larger (6–8.5(–11) × 3.5–5.0 μm vs. 4.5–6.5 × 2.5–4.0 μm), oval to ellipsoid, aseptate or 1-septate ascospores and slightly constricted paraphyses, whereas X. aquatica has ellipsoidal, aseptate, 2-guttulate ascospores (Song et al. 2023). Based on both morphological characteristics and phylogenetic evidence, Xylochrysis yunnanensis is introduced as a novel species.

Sporidesmium tropicale

M.B. Elli, Mycology Pap. 70: 58 (1958)

444F299C-B58D-5A6D-A729-C6D22CEF3990

306326

Fig. 2

Description.

For the original description, see Bao et al. (2021).

Known hosts.

Hevea brasiliensis (Senwanna et al. 2021), Macadamia sp. (this study), Pinus yunnanensis (Zhang et al. 2025a), Psidium guajava (Ellis 1971), unknown submerged decaying wood (Yang et al. 2018; Bao et al. 2021), and Camellia sinensis (Johnston 1960).

Known distribution.

China (Yunnan Province; Bao et al. 2021; Zhang et al. 2025a; this study), Sierra Leone (Ellis 1971), Thailand (Yang et al. 2018; Senwanna et al. 2021), and Malaysia (Johnston 1960).

Materials examined.

China • Yunnan Province, Pu’er City, on dead twigs of Macadamia sp., 1 August 2024, Xian Zhang, MYP22 (GMB-W 1563); living culture, GMBCC 2444, other living culture GMBCC 2445.

Notes.

In the phylogenetic analyses, the isolates (GMBCC 2444 and GMBCC 2445) were nested with Sporidesmium tropicale strains (GMB 5105, GMB 5109, DLUCC 1689, MFLU 17-0850, and MFLUCC 17-0344) with 89% ML bootstrap support (Fig. 2). Ellis (1971) introduced Sporidesmium tropicale from decaying wood in Sierra Leone based on its asexual morph. The morphology of the collection is similar to that of S. tropicale (MFLUCC 17-0344), with terminal and intercalary conidiophores and verruculose, obclavate, rostrate conidia that are truncate at the base. Later, Sporidesmium tropicale was found on Hevea brasiliensis, Pinus yunnanensis, and unknown wood in China and Thailand (Yang et al. 2018; Bao et al. 2021; Senwanna et al. 2021; Zhang et al. 2025a). In this study, S. tropicale is introduced as a new host record collected from Macadamia sp. in China.

Dothiorella lignicola

X. Zhang & N. Suwannar. sp. nov.

52BF62D0-53B9-5142-BB1B-2D1CB9BABC64

863024

Figs 3 , 13

Figure 13. — *Dothiorella* *lignicola* (*CMUB* 40121, holotype). **a, b**. Appearance of ascomata on substrate; c. Vertical section of ascoma; d. Peridium; e. Hamathecium; **f–h**. Asci; **i–l**. Ascospores (arrows indicate mucilaginous polar appendages); m. Germinated ascospore; **n, o**. Culture on PDA from above (n) and below (o); p. Sporulating colonies on PDA; **q, r**. Chlamydospores; **s, t**. Conidiogenous cells with conidia; **u–z**. Conidia. Scale bars: 200 μm (c); 100 μm (h); 50 μm (**d, f, g**); 30 μm (**m, t**); 20 μm (**e, q–s**); 10 μm (**i–l, u, w–z**); 5 μm (v); 3 cm (**n, o**).

Etymology.

The name refers to the lignicolous habit of the species.

Diagnosis.

Dothiorella lignicola differs from D. albiziae, D. dulcispinae, and D. thailandica by its ovoid conidia; from D. oblonga by its aseptate conidia; and by distinct ITS, tub2, and tef1-α sequences and a distinct phylogenetic position.

Holotype.

CMUB 40121.

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morph: Ascomata 80–120 × 160–220 µm (x̄ = 100 × 193 µm, n = 10), immersed, visible as dark dots on host surface, dark brown to black, ampulliform, papillate, without ostiole. Peridium 50–190 µm wide (x̄ = 95 µm, n = 30), composed of three layers, outer layer dark brown to black cells of textura prismatica, middle layer light brown to brown cells of texture angularis, and inner hyaline cells of texture angularis. Hamathecium composed of 4.5–8.5 µm wide (x̄ = 6.5 μm, n = 30), hyaline, septate, pseudoparaphyses slightly constricted at septum. Asci 110–235 × 23–38 µm (x̄ = 173 × 32 µm, n = 30), 8-spored, bitunicate, clavate to cylindric-clavate, pedicel, straight or slightly curved, with well-developed ocular chamber, rounded at apex, irregularly biseriate. Ascospores (27–)30–37(–40) × 14–19 µm (x̄ = 33 × 17 µm, n = 60), irregularly overlapping, subglobose to elliptical, 0–1-septate, slightly or constricted at septum, subhyaline to pale brown, brown to dark brown on maturity with a dark brown band at septa, smooth-walled, cells obtuse towards ends, cells often slightly thickened at midpoint, with or without mucilaginous polar appendages at one or both ends. Asexual morph: Conidiomata produced in culture on PDA after seven weeks at room temperature, solitary, superficial, brown to dark brown, hairy, subglobose to globose, covered with hyphal hairs, unilocular. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, discrete, cylindrical to subcylindrical, sometimes slightly tapering towards apex, hyaline, smooth, terminal. Conidia 19–26.5 × 10–13.5 µm (x̄ = 22.6 × 11.2 µm, n = 50), ellipsoid to oblong to ovoid, with rounded ends, initially hyaline and aseptate, becoming brown to dark brown and 1-septate when mature, slightly constricted or not constricted at septum, smooth-walled.

Culture characteristics.

Ascospores germinating on PDA within 24 h at 28 °C, with colonies on PDA reaching 9 cm in diam. after 1 week at 28 °C, with a rough surface, filamentous, hairy, cottony, dark gray to black, with indistinct concentric zonation from above and dull gray to black from below.

Materials examined.

Thailand • Chiang Mai Province, Doi Saket District, 18°52'43"N, 99°13'15"E, 384 m elevation, on a dead twig of Macadamia sp., 24 November 2023, Xian Zhang, TCMM51 (CMUB40121), holotype; ex-type living culture, SDBR-CMU873, other living culture SDBR-CMU874.

Notes.

In the phylogenetic analyses, the isolates of Dothiorella lignicola clustered within Dothiorella, sister to D. albiziae (MFLUCC 22-0057), D. asteracearum (MFLUCC 25-0275), D. dulcispinae (CBS 121764 and CBS 130413), D. oblonga (CBS 121765 and CBS 121766), and D. thailandica (CBS 133991), with 99% MLBS and 1.00 BYPP statistical support (Fig. 3). Morphologically, D. lignicola differs from D. albiziae (MFLU 22-0093, holotype) by having larger (19–26.5 × 10–13.5 µm vs. 14–18 × 6–8 μm), ellipsoid to oblong to ovoid conidia, whereas D. albiziae has solitary or gregarious, immersed conidiomata (Rathnayaka et al. 2022). The isolates (CMUB 40121, holotype) differ from D. dulcispinae (PREM 60706, holotype) by having terminal conidiogenous cells and longer and narrower (19–26.5 × 10–13.5 µm vs. (6–)7–10(–11) × (14–)16–22(–24) µm), ellipsoid to oblong to ovoid conidia, whereas D. dulcispinae has percurrently proliferating conidiogenous cells near the apex and ovoid conidia (Jami et al. 2012). Dothiorella lignicola differs from D. oblonga (PREM 59628, holotype) by having initially hyaline and aseptate conidia, whereas D. oblonga has honey-colored to brown conidia that become blackish brown at maturity (Slippers et al. 2014). Dothiorella lignicola differs from D. thailandica (MFLU 12-0751, holotype) by having larger (19–26.5 × 10–13.5 µm vs. 15–20 × 6.5–8 μm), granular, oblong to ovoid conidia that are slightly constricted or not constricted at the septum, whereas D. thailandica has oblong to ellipsoidal conidia with a smaller lower cell (Liu et al. 2012; Phillips et al. 2013). Dothiorella albiziae, D. dulcispinae, D. oblonga, and D. thailandica have been recorded in their asexual morphs; therefore, the sexual morphological characteristics of D. lignicola could not be compared with those of the four previously described species. Regarding the sexual morphological characteristics, the isolates (CMUB 40121, holotype) differ from D. asteracearum (MFLU 25-0274, holotype) by having immersed ascomata, a peridium comprising three layers of cells of textura angularis and textura prismatica, and irregularly overlapping ascospores with the apex often slightly thickened, whereas D. asteracearum has immersed to semi-immersed ascomata and ascospores without a gelatinous sheath (Htet et al. 2025). Dothiorella asteracearum has only been recorded from its sexual morph; therefore, the asexual morphological characteristics could not be compared. Based on nucleotide comparisons, the species (SDBR-CMU873) was compared for ITS, tef1-α, and tub2 with seven strains (Table 4), and the tef1-α gene showed more than 20 bp differences. Therefore, Dothiorella lignicola is introduced here as a new species in Dothiorella, collected from Thailand.

Table 4.

Nucleotide comparisons of D. lignicola (SDBR-CMU873, ex-type) with D. albiziae (MFLUCC 22-0057, ex-type), D. asteracearum (MFLUCC 25-0275), D. dulcispinae (CBS 121764 and CBS 130413, ex-type), D. oblonga (CBS 121765, ex-type, and CBS 121766), and D. thailandica (CBS 133991), based on ITS, tef1-α, and tub2. All comparisons excluded gaps; “-” indicates unavailable data.

Taxa	ITS	*tef1-α*	*tub2*
D. albiziae (MFLUCC 22-0057)	3/471 bp (0.6%)	21/241 bp (8.7%)	1/341 bp (0.3%)
D. asteracearum (MFLUCC 25-0275)	12/502 bp (2.4%)	–	–
D. dulcispinae (CBS 121764)	3/500 bp (0.6%)	25/260 bp (9.6%)	5/431 bp (1.2%)
D. dulcispinae (CBS 130413)	3/478 bp (0.6%)	26/263 bp (9.9%)	5/428 bp (1.2%)
D. oblonga (CBS 121765)	2/500 bp (0.4%)	26/257 bp (10.1%)	5/431 bp (1.2%)
D. oblonga (CBS 121766)	2/500 bp (0.4%)	26/257 bp (10.1%)	5/431 bp (1.2%)
D. thailandica (CBS 133991)	5/531 bp (0.9%)	31/298 bp (10.4%)	4/380 bp (1.1%)

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Dothiorella lignicola was introduced as D. macadamiae X. Zhang & N. Suwannar. in Zhang et al. (2025b). However, the name D. macadamiae is rejected under Article 52.1 of the International Code of Nomenclature for algae, fungi, and plants (ICN) because the description was based on an incorrect host, species name, etymology, and DNA sequences. Therefore, D. lignicola is proposed as a replacement name for D. macadamiae.

Lasiodiplodia macadamiae

X. Zhang, Tibpromma & Karun. sp. nov.

B7228FE3-20C0-52F2-A898-DE65D1D6DD41

863025

Figs 4 , 14

Figure 14. — *Lasiodiplodia* *macadamiae* (GMB-W 1564, holotype). **a–c**. Appearance of ascomata on substrate; **d, e**. Vertical sections of ascomata; f. Peridium; g. Hamathecium; **h–k**. Asci; **l–q**. Ascospores; r. Germinated ascospore; **s, t**. Culture on PDA from above (s) and below (t). Scale bars: 200 μm (**d, e**); 100 μm (h); 50 μm (**f, i–k**); 20 μm (**g, o–r**); 10 μm (**l, m**); 2 cm (**s, t**).

Etymology.

“macadamiae” refers to the host plant genus Macadamia.

Diagnosis.

Lasiodiplodia macadamiae differs from L. chinensis by having wider asci and ascospores; from L. gonubiensis, L. lignicola, and L. pseudotheobromae by having hyaline ascospores; and from L. theobromae by having overlapping bi- to tri-seriate ascospores. It also has distinct ITS, tef1-α, and tub2 sequences and a distinct phylogenetic position.

Holotype.

GMB-W 1564.

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morph: Ascomata (including ostioles) 220–320 × 340–580 µm (x̄ = 277 × 451 µm, n = 10), solitary or in small groups, immersed to erumpent through the host surface, appearing as black spots on the host surface, subglobose to flask-shaped, sometimes ascomata gregarious, forming small clusters, dark brown to black, ostiolate, with a papilla. Ostioles up to 55–90 µm wide, central at neck, cylindrical to papillate. Peridium 30–90 µm thin to thick-walled, with outer layers composed of dark brown cells of textura angularis, with inner layers composed of light brown cells of textura prismatica. Hamathecium composed of 3–7.5 µm wide (x̄ = 4.7 μm, n = 40), abundant, dense, narrowly cylindrical to filiform, hyaline, septate, branched, broad cellular pseudoparaphyses sometimes constricted at the septum. Asci 95–180 × 25–50 µm (x̄ = 131.7 × 35.5 µm, n = 40), 8-spored, bitunicate, fissitunicate, clavate to broadly clavate, straight to slightly curved, rounded at apex, short furcate or obtuse at pedicel, with an apical chamber. Ascospores 30–40 × 8.5–17 µm (x̄ = 34.8 × 14 µm, n = 40), overlapping uni- to bi-seriate, fusiform to broadly fusiform, with rounded ends, wider at middle, aseptate, hyaline, guttulate, smooth-walled. Asexual morph: Not observed.

Culture characteristics.

Ascospores germinating on PDA within 24 h at 28 °C. Colonies on PDA reaching 6 cm in diam. after 3 weeks at 28 °C, with a rough surface, filamentous, hairy, circular, and cottony; from above, gray at the margin, becoming light gray to gray and dark gray at the center; gray to black in reverse.

Material examined.

China • Yunnan Province, Pu’er City, on dead twigs of Macadamia sp., 1 August 2024, Xian Zhang, MYP9 (GMB-W 1564), holotype; ex-type living culture, GMBCC 2446, other living culture GMBCC 2447.

Notes.

In the phylogenetic analyses, the isolates (GMBCC 2446 and GMBCC 2447) clustered within Lasiodiplodia, sister to L. clavispora (CGMCC 3.19594 and CGMCC 3.19595) and L. gonubiensis (CBS 115812 and CBS 116355), with 97% MLBS and 1.00 BYPP statistical support (Fig. 4). Based on base-pair comparisons, GMBCC 2446 was compared for ITS, tef1-α, and tub2 with L. clavispora (CGMCC 3.19594 and CGMCC 3.19595) and L. gonubiensis (CBS 116355 and CBS 115812) (Table 5), and the tef1-α gene showed more than 25 bp differences from L. clavispora and L. gonubiensis strains. Morphologically, L. clavispora has only been recorded in its asexual morph, whereas L. macadamiae is recorded from its sexual morph; therefore, comparing these species morphologically was not possible. In this genus, only five species (viz. L. chinensis, L. gonubiensis, L. lignicola, L. pseudotheobromae, and L. theobromae) have been recorded from their sexual morphs (Phillips et al. 2013; Trakunyingcharoen et al. 2015; Dou et al. 2017; Tennakoon et al. 2017). Lasiodiplodia macadamiae resembles other Lasiodiplodia species in having dark brown to black ascomata, bitunicate, clavate asci, and overlapping ascospores. The isolates can be distinguished from L. chinensis (HMAS247143, holotype) by having a peridium composed of cells of textura angularis and textura prismatica, fissitunicate and wider asci without a furcate pedicel (95–180 × 25–50 µm vs. 53–90(–116) × 14–23(–25) µm), and overlapping uni- to bi-seriate, wider ascospores (30–40 × 8.5–17 µm vs. 19–25 × 6–9 μm), whereas L. chinensis has solitary ascomata or ascomata in small groups of up to four, a peridium composed of textura angularis cells, asci with a thick endotunica, and overlapping bi- to tri-seriate ascospores (Dou et al. 2017).

Table 5.

Nucleotide comparisons of L. macadamiae (GMBCC 2446, ex-type) with L. clavispora (CGMCC 3.19594, ex-type, and CGMCC 3.19595) and L. gonubiensis (CBS 116355 and CBS 115812, ex-type), based on ITS, tef1-α, and tub2. All comparisons excluded gaps.

Taxa	ITS	*tef1-α*	*tub2*
L. clavispora (CGMCC 3.19594)	0/485 bp	31/245 bp (12.65%)	3/377 bp (0.8%)
L. clavispora (CGMCC 3.19595)	0/485 bp	31/245 bp (12.65%)	3/377 bp (0.8%)
L. gonubiensis (CBS 116355)	0/528 bp	29/242 bp (12%)	5/422 bp (1.2%)
L. gonubiensis (CBS 115812)	0/528 bp	29/242 bp (12%)	5/422 bp (1.2%)

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Lasiodiplodia macadamiae differs from L. gonubiensis (PREM 58127, holotype) by having 8-spored, short-stalked asci and aseptate, hyaline ascospores, whereas L. gonubiensis has asci containing (4–)6–8 ascospores and ascospores that rarely become 1–2-septate with age and turn pale brown, becoming 1–2-septate within the ascoma or shortly after discharge (Trakunyingcharoen et al. 2015). It differs from L. lignicola (MFLU 12-0750, holotype) by having immersed to erumpent ascomata, asci with a short, furcate or obtuse pedicel, longer asci (95–180 × 25–50 µm vs. 80–90 × 15–25 μm), and overlapping uni- to bi-seriate, hyaline ascospores, whereas L. lignicola has semi-immersed ascomata, asci with a short and narrow pedicel, and uniseriate or partially overlapping, reddish-brown to dark brown ascospores (Phillips et al. 2013). It differs from L. pseudotheobromae (CBS-H 19916, holotype) by having a branched hamathecium, short-stalked asci, and hyaline, overlapping bi- to tri-seriate ascospores, whereas L. pseudotheobromae has cylindric-clavate asci and ellipsoidal to fusiform ascospores that are initially hyaline to pale yellowish and become golden to dark brown at maturity (Tennakoon et al. 2017). It differs from L. theobromae (CBS H-21411, neotype) by having clavate to broadly clavate, non-stipitate asci and overlapping bi- to tri-seriate ascospores, whereas L. theobromae has stipitate asci and irregularly biseriate ascospores (Phillips et al. 2013). Therefore, Lasiodiplodia macadamiae, collected from Macadamia sp. in China, is introduced here as a new species in Lasiodiplodia.

Lasiodiplodia ponkanicola

X.E. Xiao, Crous & H.Y. Li, in Xiao, Wang, Crous, Wang, Jiao, Huang, Pu, Zhu & Li, Persoonia 47: 128 (2021)

FDD48B20-70D0-52F8-9C39-7B74FD9DBC21

840685

Fig. 4

Description.

For the original description, see Xiao et al. (2021).

Known hosts.

Citrus reticulata (Xiao et al. 2021), Macadamia sp. (this study).

Known distribution.

China (Zhejiang Province; Xiao et al. 2021), Thailand (this study).

Material examined.

Thailand • Chiang Mai Province, Samoeng Tai, on dead twigs of Macadamia sp. (18°85'38"N, 98°73'0"E, 512 m), 24 May 2024, Xian Zhang, TCSM58 (CMUB 40122); living culture, SDBR-CMU875.

Notes.

Xiao et al. (2021) introduced Lasiodiplodia ponkanicola as a pathogen associated with the trunk of Citrus reticulata cv. Ponkan and confirmed its pathogenicity with a pathogenicity test. The morphological characteristics of the isolate (CMUB 40122) resemble those of L. ponkanicola (ZJUE H-0044, holotype), with holoblastic, hyaline conidiogenous cells and hyaline, aseptate, ellipsoid to ovoid conidia (Xiao et al. 2021). In the phylogenetic analysis, the isolate clustered with L. ponkanicola (CGMCC 3.20388), with 78% ML bootstrap support (Fig. 4). Based on nucleotide comparisons, the isolate (SDBR-CMU875) showed similarities to L. ponkanicola (CGMCC 3.20388), with 1/433 bp (0.23%, without gaps) differences in tub2, 2/292 bp (0.68%, without gaps) differences in tef1-α, and identical ITS sequences. Therefore, based on morphological and phylogenetic analyses, the saprobic isolate of Lasiodiplodia ponkanicola is reported here as a new life mode (saprobic), a new host record, and a new country record, collected from Macadamia sp. in Thailand.

Lasiodiplodia endophytica

N.I. de Silva, A.J.L. Phillips & K.D. Hyde, in de Silva, Scientific Reports 9 (no. 14355): 4 (2019)

62FFE32C-19B7-58C9-8BF5-5ACCE0917AED

556218

Figs 4 , 15

Figure 15. — *Lasiodiplodia* *endophytica* (*CMUB* 40123, new host record and country record). **a–c**. Conidiomata on a decaying twig of *Macadamia* sp.; **d, e**. Vertical sections of conidioma; f. Conidiomata wall; g. Paraphyses; **h–j**. Conidiogenous cells with conidia; **k–n**. Conidia; **o, p**. Culture on PDA from above (o) and below (p). Scale bars: 100 μm (d); 50 μm (**e, f**); 20 μm (**g–n**); 2 cm (**o, p**).

Description.

Saprobic dead twigs of Macadamia sp. Asexual morph: Conidiomata 400–565 × 150–250 µm (x̄ = 483 × 195 µm, n = 10), pycnidial, immersed to semi-immersed, scattered to aggregated, unilocular or multilocular, brown to dark brown, globose to depressed globose, ostiolate. Conidiomata walls 20–60 µm wide, composed of hyaline to brown to dark brown cells of textura angularis. Paraphyses 3–6.5 μm wide arising amongst the conidiogenous cells, hyaline, cylindrical, the basal cells often slightly swollen, rounded at apex. Conidiophores absent. Conidiogenous cells 2.0–5.5 µm diam., hyaline, discrete, cylindrical, holoblastic. Conidia 19.5–32.5 × 10–17 (x̄ = 26.5 × 13.6 µm, n = 30) μm, hyaline, aseptate, ellipsoidal to broadly ellipsoidal, occasionally oblong, both ends broadly rounded, smooth- and thick-walled, contents granular, small guttules.

Known hosts.

Macadamia sp. (this study), Magnolia candolii (de Silva et al. 2019).

Known distribution.

China (Yunnan Province; de Silva et al. 2019), Thailand (this study).

Material examined.

Thailand • Chiang Mai Province, Samoeng Tai, on dead twigs of Macadamia sp. (18°85'38"N, 98°73'0"E, 512 m), 24 May 2024, Xian Zhang, TCSM53 (CMUB 40123); living culture, SDBR-CMU876.

Notes.

de Silva et al. (2019) introduced Lasiodiplodia endophytica. Lasiodiplodia endophytica has been reported as an endophyte on fresh leaves of Magnolia candolii (de Silva et al. 2019). The morphological characteristics of the isolates resemble those of L. hyalina (HMAS 255216, holotype) and Lasiodiplodia thailandica (CBS-H 21933, holotype) in having solitary conidiomata, holoblastic, discrete, hyaline conidiogenous cells, and thick-walled, aseptate conidia with both ends broadly rounded. However, they can be distinguished from L. hyalina (HMAS 255216, holotype) by having unilocular or multilocular conidiomata and ellipsoidal to broadly ellipsoidal conidia, and from L. thailandica (CBS-H 21933, holotype) by having immersed to semi-immersed conidiomata and ellipsoidal to broadly ellipsoidal conidia (Trakunyingcharoen et al. 2015; Dou et al. 2017). In the phylogenetic analysis, the isolates clustered with L. endophytica (MFLUCC 18-1121), with 73% ML bootstrap support (Fig. 4). Based on nucleotide comparisons, the isolates showed similarity to L. hyalina strain CGMCC 3.17975, with 1/353 bp (0.28%, without gaps) difference in tub2 and identical ITS and tef1-α sequences. Therefore, based on morphological and phylogenetic analyses, Lasiodiplodia endophytica is introduced here as a new host record and a new country record, collected from Macadamia sp. in Thailand.

Palmiascoma macadamiae

X. Zhang, Tibpromma & Karun. sp. nov.

5FC70F32-675B-5304-A219-A42A9220C165

863026

Figs 5 , 16

Figure 16. — *Palmiascoma* *macadamiae* (GMB-W 1565, holotype). **a–c**. Appearance of ascomata on substrate; **d, e**. Vertical sections of ascomata; f. Peridium; g. Hamathecium; **h–k**. Asci; **l–p**. Ascospores; q. Germinated ascospore; **r, s**. Culture on PDA from above (r) and below (s). Scale bars: 100 μm (**d, e**); 20 μm (**f–k**); 10 μm (q); 5 μm (**l–p**); 2 cm (**r, s**).

Etymology.

“macadamiae” refers to the host plant genus Macadamia.

Diagnosis.

Palmiascoma macadamiae differs from P. acericola by having overlapping 1–2-seriate ascospores, from P. gregariascomum by lacking a mucilaginous sheath or appendages on the ascospores, and from P. qujingense by having non-echinulate, slightly shorter ascospores. It also has distinct nrLSU, nrSSU, ITS, and rpb2 sequences and a distinct phylogenetic position.

Holotype.

GMB-W 1565.

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morph: Ascomata 115–205 × 145–265 µm (x̄ = 159 × 198 µm, n = 10), scattered to clustered, semi-immersed, slightly erumpent, dark spots on the host surface, globose to subglobose or irregular, glabrous, uniloculate, centrally ostiolate, papillate. Ostioles 40–150 µm long, 25–60 µm diam. (x̄ = 80 × 41 µm, n = 10), brown to dark brown, glabrous, papillate, shiny, lined with hyaline periphyses. Peridium 10–30 µm thin to thick-walled, of unequal thickness, slightly thick at sides, composed of brown to dark brown cells of textura angularis. Hamathecium 1.5–3.0 µm wide (x̄ = 2.2 μm, n = 30), abundant, dense, narrowly cylindrical to filiform, hyaline, branched, pseudoparaphyses; no distinct septation was observed, anastomosing above the asci, embedded in a mucilaginous matrix. Asci 40–60 × 7.5–11.5 µm (x̄ = 49.5 × 8.7 µm, n = 30), 8-spored, bitunicate, fissitunicate, cylindric-clavate, short pedicellate, with a furcate base, apically rounded, with an ocular chamber. Ascospores 8.5–12 × 4–5 µm (x̄ = 10.5 × 4.5 µm, n = 30), overlapping 1–2-seriate, fusiform to ellipsoid, with rounded ends, hyaline to slight brown to brown, straight to slightly curved, uniseptate, with slightly constricted at the septum, upper part slight wider, with or without guttules, without mucilaginous sheath or appendages, rough-walled. Asexual morph: Not observed.

Culture characteristics.

Ascospores germinating on PDA within 24 h at 28 °C, with colonies on PDA reaching 3.6 cm in diam. after 3 weeks at 28 °C, with a rough surface, filamentous, hairy, circular, cottony, and white from above; light yellow at the margins, turning light brown toward the center in reverse.

Materials examined.

China • Yunnan Province, Dehong City, on dead twigs of Macadamia sp., 24 April 2024, Xian Zhang, MYD1 (GMB-W 1565), holotype; ex-type living culture, GMBCC 2448, other living culture GMBCC 2449.

Notes.

In the phylogenetic analyses, the isolates (GMBCC 2448 and GMBCC 2449) clustered within Palmiascoma, sister to P. acericola (CGMCC 3.24963 and UESTCC 23.0018), P. gregariascomum (MFLUCC 11-0175 and MFLUCC 11-0424), and P. qujingense (KKUMCC 19-0201), with 100% MLBS and 1.00 BYPP statistical support (Fig. 5). Morphologically, Palmiascoma macadamiae resembles P. acericola, P. gregariascomum, and P. qujingense in having glabrous, uniloculate, papillate ascomata, a hamathecium containing hyaline, septate, branched pseudoparaphyses, 8-spored, bitunicate, fissitunicate, cylindric-clavate, apically rounded asci, and fusiform to ellipsoid, uniseptate, rough-walled ascospores (Liu et al. 2015; Monkai et al. 2021; Li et al. 2023). However, P. macadamiae can be distinguished from P. acericola (HKAS 127160, holotype) by having scattered to clustered, semi-immersed, subglobose or irregular ascomata, pseudoparaphyses without distinct septation, short-pedicellate asci, and overlapping 1–2-seriate, slightly shorter (8.5–12 × 4–5 µm vs. 12–14 × 3.5–4.5 μm) ascospores, whereas P. acericola has immersed to erumpent ascomata and elongated, cylindrical, pedicellate asci (Li et al. 2023). It can be distinguished from P. gregariascomum (MFLU 11-0211, holotype) by having a hamathecium without distinctly observed septation and ascospores that are hyaline when young, slightly constricted at the septum, and without a mucilaginous sheath or appendages, whereas P. gregariascomum has a hamathecium with distinct septa and echinulate ascospores surrounded by a mucilaginous sheath (Liu et al. 2015). It can also be distinguished from P. qujingense (KUN-HKAS109581, holotype) by having semi-immersed ascomata, pseudoparaphyses without distinctly observed septation, slightly shorter (40–60 × 7.5–11.5 µm vs. 60–80 × 9–10 μm) asci, and non-echinulate, slightly shorter (8.5–12 × 4–5 µm vs. 12–16 × 4.5–5.5 μm) ascospores, whereas P. qujingense has immersed to semi-immersed ascomata and clavate to ellipsoidal, echinulate ascospores (Monkai et al. 2021). Therefore, Palmiascoma macadamiae is introduced here as a new species in Palmiascoma, collected from Macadamia sp. in China.

Palmiascoma chiangraiense

N. Afshari, Lumyong & K.D. Hyde, in Afshari et al., Mycosphere 16: 4812 (2025)

97364F88-B55D-5468-96F6-EEB50A357EA5

904739

Figs 5 , 17

Figure 17. — *Palmiascoma* *chiangraiense* (GMB-W 1566, new host record and country record). **a–c**. Conidiomata on a decaying branch of *Macadamia* sp.; d. Vertical section of conidioma; e. Vertical section through conidioma showing the ostiolar neck; f. Conidiomata wall; **g–i**. Conidiogenous cells with conidia; j. Germinated conidium; **k, l**. Conidia; **m, n**. Culture on PDA from above (m) and below (n). Scale bars: 50 μm (**d–f**); 20 μm (j); 10 μm (**g, k, l**); 5 μm (**h, i**); 2 cm (**m, n**).

Holotype.

MFLU 24-0459.

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morph: Not observed. Asexual morph: Conidiomata 225–380 × 335–900 µm (x̄ = 299 × 634 µm, n = 10), pycnidial, immersed to semi-immersed or erumpent, solitary or aggregated, globose to subglobose, irregular, glabrous, ostiole, uniloculate, scattered, papillate. Conidiomata walls 25–75 µm (x̄ = 45 µm, n = 30) thin to thick walled, composed of several layers of hyaline to dark brown, outer layers composed of 13–37 µm (x̄ = 22.6 µm, n = 15), brown to dark brown cells of textura angularis, inner layers composed of 18–47 µm (x̄ = 30.8 µm, n = 15), hyaline cells, organized in a textura angularis. Conidiophores arising from basal cavity of conidiomata, mostly reduced to conidiogenous cells. Conidiogenous cells holoblastic, phialidic, ampulliform to cylindrical, hyaline, aseptate, smooth-walled, guttulate. Conidia 4–5.5 × 1–3 µm (x̄ = 4.8 × 2.3 µm, n = 30), solitary, oblong to ellipsoidal, aseptate, with rounded or obtuse ends, hyaline when young, becoming brown with maturity, smooth-walled, guttulate.

Culture characteristics.

Conidia germinating on PDA within 24 h at 28 °C, with colonies on PDA reaching 6 cm in diam. after 3 weeks at 28 °C, with a rough surface, filamentous, hairy, circular, cottony, and white from above; light brown at the margins, turning yellow-brown to brown toward the center in reverse.

Known hosts.

Dipterocarpus alatus (Afshari et al. 2025b), Macadamia sp. (this study).

Known distribution.

China (Yunnan Province; this study), Thailand (Afshari et al. 2025b).

Material examined.

China • Yunnan Province, Pu’er City, on dead twigs of Macadamia sp., 1 August 2024, Xian Zhang, MYP1 (GMB-W 1566); living culture, GMBCC 2450, other living culture GMBCC 2451.

Notes.

Palmiascoma chiangraiense was introduced by Afshari et al. (2025b), isolated from dead wood of Dipterocarpus alatus based on its sexual morph. In this study, the asexual morph of this species was discovered for the first time. In the phylogenetic analyses, the isolate (GMBCC 2450) clustered with P. chiangraiense strain MFLUCC 24-0546, with 100% MLBS and 1.00 BYPP statistical support (Fig. 5). Morphologically, the isolate (GMBCC 2450) resembles P. gregariascomum in having erumpent, pycnidial conidiomata, phialidic, discrete, ampulliform to cylindrical conidiogenous cells, and solitary, oblong to ellipsoidal conidia, but can be distinguished by having larger conidiomata (225–380 µm high × 335–900 µm diam. vs. 140–200 μm high × 130–220 μm diam.) (Liu et al. 2015). In addition, phylogenetic analyses show that these species form distinct lineages (Fig. 5). Palmiascoma acericola, P. macadamiae, and P. qujingense were recorded only from their sexual morphs. Therefore, the asexual morphological characteristics of P. chiangraiense could not be compared with those of the three species. Based on nucleotide comparisons, the isolate (GMBCC 2450) showed similarity to P. chiangraiense strain MFLUCC 24-0546, with 3/644 bp (0.47%, without gaps) differences in ITS, 2/836 bp (0.24%, without gaps) differences in nrLSU, 4/1033 bp (0.39%, without gaps) differences in rpb2, and identical nrSSU sequences. Therefore, the collection is introduced here as a new host record and a new country record of P. chiangraiense, collected from Macadamia sp. in China.

Crassiparies quadrisporus

M. Matsum., K. Hiray. & Kaz. Tanaka, in Li et al., Fungal Diversity 78: 63 (2016)

F83AB3CF-223F-5C4A-949B-7BD9D5F5A746

559359

Fig. 6

Description.

For the original description, see Li et al. (2016).

Known hosts.

Acer sp. (Li et al. 2016), Macadamia sp. (this study), Machilus japonica (Tanaka et al. 2017), Hevea brasiliensis (Senwanna et al. 2021).

Known distribution.

China (Yunnan Province; this study), Japan (Li et al. 2016; Tanaka et al. 2017), Thailand (Senwanna et al. 2021).

Material examined.

China • Yunnan Province, Dehong City, on dead twigs of Macadamia sp., 24 April 2024, Xian Zhang, MYD3 (GMB-W 1567); living culture, GMBCC 2452, • ibid., MYD9 other living culture GMBCC 2453.

Notes.

Li et al. (2016) introduced Crassiparies quadrisporus, isolated from dead twigs of Acer sp. The morphological characteristics of the isolates resemble those of Crassiparies quadrisporus (HHUF 30409, holotype) in having bitunicate, fissitunicate, pedicellate, cylindric-clavate or clavate asci and overlapping biseriate, hyaline, septate, fusiform ascospores (Li et al. 2016; Tanaka et al. 2017; Senwanna et al. 2021). In the phylogenetic analysis, the isolates clustered with Crassiparies quadrisporus strains (MAFF 245408, MAFF 246250, MFLUCC 18-0298, and MFLUCC 21-0148) with 100% MLBS and 1.00 BYPP statistical support (Fig. 6). Based on nucleotide comparisons, the isolate GMBCC 2452 was highly similar to Crassiparies quadrisporus (MAFF 245408), with only 3/927 bp (0.3%, without gaps) differences in tef1-α, and was identical in ITS, nrLSU, nrSSU, and rpb2. Therefore, based on morphological and phylogenetic analyses, Crassiparies quadrisporus is introduced here as a new host and country record, collected from Macadamia sp. in China.

Chromolaenicola ananasi

X.G. Tian, K.D. Hyde & Tibpromma, in Tian et al., Mycosphere 15(1): 31 (2024)

C7E18628-CD3F-5C41-B2F8-997643FAA8A7

900971

Figs 7 , 18

Figure 18. — *Chromolaenicola* *ananasi* (*CMUB* 40120, new host record and country record). **a, b**. Appearance of ascomata on substrate; c. Vertical section of ascomata; d. Peridium; e. Hamathecium; **f–j**. Asci; **k–q**. Ascospores; r. Germinated ascospore; **s, t**. Culture on PDA from above (s) and below (t). Scale bars: 200 μm (c); 50 μm (**d, e**); 30 μm (**f–j**); 20 μm (r); 10 μm (**k–q**); 2 cm (**s, t**).

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morphs: Ascomata 230–415 µm high × 200–415 µm diam. (x̄ = 332 × 309 µm, n = 10, including ostioles), solitary to scattered, immersed or erumpent, scattered, dark brown to black, globose to sub-globose, rarely irregular, unilocular to bilocular, with a central ostiole. Peridium 18–40 µm wide, thin-walled, composed of several layers, inner layers comprising hyaline cells of textura epidermoidea, outer layers comprising brown to pale brown cells of textura angularis. Hamathecium composed of 1.5–3.0 µm wide (x̄ = 1.9 μm, n = 40), cylindrical to filiform, hyaline, septate, branched, pseudoparaphyses. Asci 70–110 × 5.5–9 µm (x̄ = 90 × 7.0 µm, n = 40), 8-spored, bitunicate, cylindrical-clavate, straight to slightly curved, rounded at the apex, short pedicellate with club-shaped base. Ascospores 10–14 × 5.0–6.5 µm (x̄ = 12.5 × 5.4 µm, n = 40), slightly overlapping, uniseriate, fusiform to broadly fusiform, 1-septate at center, constricted at septum, with wider upper cell, straight or slightly curved, initially hyaline to pale brown, becoming reddish brown to brown at maturity with thickened septum, with or without guttules, without a gelatinous sheath. Asexual morph: See Tian et al. (2024b).

Known hosts.

Ananas comosus (Tian et al. 2024b), Macadamia sp. (this study).

Known distribution.

Thailand (Tian et al. 2024b; this study).

Material examined.

Thailand • Chiang Mai Province, Doi Saket District, 18°52'43"N, 99°13'15"E, 384 m elevation, on dead twigs of Macadamia sp., 24 November 2023, Xian Zhang, TCMM49 (CMUB 40120); living culture, SDBR-CMU872.

Notes.

Chromolaenicola ananasi was introduced by Tian et al. (2024b), isolated from dead leaves of Ananas comosus based on its asexual morph. In the present study, the sexual morph of this species was discovered for the first time. In the phylogenetic analysis, the isolate SDBR-CMU872 grouped with Chromolaenicola ananasi (MFLU 23-0167 and MFLUCC 23-0111) with 100% MLBS and 1.00 BYPP statistical support (Fig. 7). Morphologically, the collection is almost identical to Chromolaenicola sapindi (HKAS 122789, holotype) in having immersed-to-erumpent ascomata, 8-spored, bitunicate, cylindric-clavate asci, and uniseriate ascospores that are initially hyaline to pale brown, becoming reddish brown to brown at maturity with a thickened septum, constricted at the central septum, and lacking a gelatinous sheath (Ren et al. 2022). However, the asci and ascospores of the collection are slightly different from those of C. sapindi. The collection has a peridium composed of cells of textura epidermoidea and textura angularis, short-pedicellate asci with a club-shaped base, and ascospores that are 1-septate at the center from immature to mature stages, whereas C. sapindi has ampulliform or obovoid ascomata and ellipsoidal, aseptate or 1-septate ascospores; the phylogenetic tree also shows that they are distinct species (Ren et al. 2022). Based on nucleotide comparisons, the isolate SDBR-CMU872 showed similarity to C. ananasi (KUMCC 21-0564 and KUMCC 21-0594), with only 7/902 bp (0.8%, without gaps) differences in tef1-α, and was identical in ITS, nrLSU, and nrSSU. The collection is recognized as C. ananasi because it shares almost identical molecular data with other collections (Tian et al. 2024b). Therefore, based on morphological and phylogenetic analyses, Chromolaenicola ananasi is introduced here as a new host and country record, collected from Macadamia sp. in Thailand. In addition, this is the first report of Chromolaenicola ananasi associated with Macadamia sp.

Montagnula donacina

(Niessl) Wanas., E.G. Jones & D. Hyde, Fungal Biology 120 (11): 1365 (2016)

681B352B-5D79-5C2D-BBEB-0B9B72E7ADAC

626421

Fig. 8

Description.

For description, see Sun et al. (2023).

Known hosts.

Acacia reficiens (Aptroot 1995), Acacia sp. (Aptroot 1995), Adhatoda vasica (Aptroot 1995), Ailanthus altissima (Aptroot 1995), Althaea rosea (Aptroot 1995), Annona squamosa (Aptroot 1995), Arundo donax (Aptroot 1995), Bambusoideae sp. (Aptroot 1995), Cajanus cajan (Aptroot 1995), Calamus australis (Hyde et al. 1999), Careya arborea (Aptroot 1995), Chromolaena odorata (Aptroot 1995), Citrus aurantiifolia (Aptroot 1995), Citrus sp. (Aptroot 1995), Clerodendrum infortunatum (Aptroot 1995), Clerodendrum multiflorum (Aptroot 1995), Coffea arabica var. catimor (Aptroot 1995), Coffea arabica (Aptroot 1995), Coffea robusta (Aptroot 1995), Craterellus odoratus (Zhao et al. 2018), Duranta repens (Aptroot 1995), Ficus glomerata (Aptroot 1995), Funtumia africana (Aptroot 1995), Hevea brasiliensis (Aptroot 1995), Hibiscus sp. (Aptroot 1995), Ipomoea carnea (Aptroot 1995), Lagerstroemia sp. (Aptroot 1995), Litchi litchi (Thaung 2008), Macadamia sp. (this study), Mallotus philippinensis (Aptroot 1995), Morus alba (Aptroot 1995), Nerium odorum (Aptroot 1995), Paeonia suffruticosa (Li et al. 2023), Phyllostachys bambusoides (Wang et al. 2004), Pistacia sp. (Aptroot 1995; Wang et al. 2004), Premna cumingiana (Aptroot 1995), Pseudosasa japonica (Aptroot 1995), Saccharum officinarum (Aptroot 1995), Tectona grandis (Aptroot 1995), Terminalia tomentosa (Aptroot 1995), Trachycarpus fortune (Hyde et al. 1999), unidentified twig (Aptroot 1995), Vitis vinifera (Pitt et al. 2014), Wikstroemia sp. (Aptroot 1995), Zea mays (Aptroot 1995), dead wood (Ren et al. 2022; Sun et al. 2023; Wanasinghe et al. 2024).

Known distribution.

Australia (Pitt et al. 2014), Brazil (Aptroot 1995), Central African Republic (Aptroot 1995), China (Aptroot 1995; Hong Kong, Hyde et al. 1999; Sichuan Province, Li et al. 2023; Yunnan Province, Zhao et al. 2018; Wanasinghe et al. 2024; this study), Colombia (Aptroot 1995), France (Aptroot 1995), Georgia (Aptroot 1995), India (Aptroot 1995), Japan (Wang et al. 2004), Myanmar (Thaung 2008), Namibia (Aptroot 1995), Papua New Guinea (Aptroot 1995), Paraguay (Aptroot 1995), Philippines (Aptroot 1995), Portugal (Aptroot 1995), Sierra Leone (Aptroot 1995), Thailand (Ren et al. 2022), USA (Aptroot 1995; Wang et al. 2004).

Material examined.

China • Yunnan Province, Honghe City, on a dead branch of Macadamia sp., 16 July 2024, Xian Zhang, MYH4 (GMB-W 1570); living culture, GMBCC 2455.

Notes.

Sun et al. (2023) synonymized Montagnula chromolaenicola, M. puerensis, M. saikhuensis, and M. thailandica under M. donacina based on morphological and phylogenetic analyses. However, Wanasinghe et al. (2024) subsequently suggested that these taxa represent distinct species based on combined gene analyses. In the phylogenetic analysis, the isolate GMBCC 2455 was sister to M. donacina (KUNCC 23-14428 and KUNCC 23-14429), with 81% ML bootstrap support (Fig. 8). Morphological analysis also supports the identification of the isolate as M. donacina (HKAS 124552), with semi-immersed ascomata, bitunicate, clavate asci with a bulbous, long pedicel, and 1-septate ascospores constricted at the septum (Sun et al. 2023). Therefore, based on this evidence, the isolate is confirmed as Montagnula donacina, and this species is reported for the first time from Macadamia sp. in China.

Montagnula thailandica

Mapook & K.D. Hyde, in Mapook, Hyde, McKenzie, Jones, Bhat, Jeewon, Stadler, Samarakoon, Malaithong, Tanunchai, Buscot, Wubet & Purahong, Fungal Diversity 101: 35 (2020)

9736D64B-96BF-5FAD-8F69-BDB042F6AF7C

557299

Fig. 8

Description.

For description, see Mapook et al. (2020).

Known hosts.

Chromolaena odorata (Mapook et al. 2020), Coffea arabica var. catimor (Lu et al. 2022), Hevea brasiliensis (Senwanna et al. 2021), Macadamia sp. (this study), unidentified twig (Boonmee et al. 2021).

Known distribution.

Material examined.

China • Yunnan Province, Pu’er City, on dead twigs of Macadamia sp., 1 August 2024, Xian Zhang, MYP5 (GMB-W 1569); living culture, GMBCC 2454.

Notes.

Mapook et al. (2020) introduced Montagnula thailandica, isolated from dead stems of Chromolaena odorata, based on morphological and phylogenetic analyses. In the phylogenetic analysis, the isolate GMBCC 2454 was sister to M. thailandica (MFLUCC 17-1508) (Fig. 8). Morphologically, the collection fits well with M. thailandica (MFLU 20-0325, holotype), with immersed to erumpent ascomata and bitunicate, fissitunicate asci with long pedicels and 1-septate ascospores constricted at the septum (Mapook et al. 2020). Therefore, based on morphological and phylogenetic analyses, the isolate is confirmed as M. thailandica and is introduced as a new host and country record from Macadamia sp. in China.

Pseudochaetosphaeronema chiangraiense

Wijes., Boonmee & K.D. Hyde, in Boonmee et al., Fungal Diversity 111: 75 (2021)

BEBE40D8-1BDF-5020-A5B3-47E6CB7F00A4

558549

Fig. 9

Description.

For the original description, see Boonmee et al. (2021).

Known hosts.

Macadamia sp. (this study), Tamarindus sp. (Boonmee et al. 2021).

Known distribution.

China (Yunnan Province; this study), Thailand (Boonmee et al. 2021).

Materials examined.

China • Yunnan Province, Dehong City, on a dead branch of Macadamia sp., 24 April 2024, Xian Zhang, MYD19 (GMB-W 1571); living culture, GMBCC 2456, other living culture GMBCC 2457.

Notes.

Boonmee et al. (2021) introduced Pseudochaetosphaeronema chiangraiense, isolated from a terrestrial branch of Tamarindus sp. In the phylogenetic analysis, the isolate GMBCC 2456 clustered with Pse. chiangraiense strains (MFLUCC 21-0083 and UESTCC 23-0065) with 62% ML bootstrap support (Fig. 9). Based on nucleotide comparisons, the isolate GMBCC 2456 showed similarity to Pse. chiangraiense (MFLUCC 21-0083), with only 1/842 bp (0.1%, without gaps) difference in tef1-α, and was identical in ITS, nrLSU, and nrSSU. Morphologically, the collection fits well with the description of Pse. chiangraiense (MFLU 21-0064, holotype) in having 8-spored, bitunicate asci with rounded ends and overlapping, hyaline, fusiform, septate ascospores (Boonmee et al. 2021). Therefore, based on morphological and phylogenetic analyses, the isolates are confirmed as Pse. chiangraiense and are introduced as a new host and country record from Macadamia sp. in China.

Pseudochaetosphaeronema puerense

R.F. Xu, K.D. Hyde & Tibpromma [as ‘ puerensis’], in Manawasinghe et al., Fungal Diversity 130: 40 (2024) [2025]

42F633E9-E10D-5059-B55F-D6EF1C017D6D

901412

Fig. 9

Description.

For the original description, see Manawasinghe et al. (2025).

Known hosts.

Hevea brasiliensis (Manawasinghe et al. 2025), Macadamia sp. (this study).

Known distribution.

China (Yunnan Province; Manawasinghe et al. 2025; this study).

Material examined.

China • Yunnan Province, Dehong City, on a dead branch of Macadamia sp., 24 April 2024, Xian Zhang, MYD14 (GMB-W 1572); living culture, GMBCC 2458, other living culture GMBCC 2459.

Notes.

Manawasinghe et al. (2025) introduced Pseudochaetosphaeronema puerense, isolated from a dead branch of Hevea brasiliensis in Yunnan Province, China. In the phylogenetic analysis, the isolates GMBCC 2458 and GMBCC 2459 clustered with Pse. puerense strains (ZHKUCC 22-0288 and ZHKUCC 22-0289) with 100% MLBS and 1.00 BYPP statistical support (Fig. 9). Based on nucleotide comparisons, the isolate GMBCC 2458 showed similarity to Pseudochaetosphaeronema puerense (ZHKUCC 22-0288, ex-type), with only 2/561 bp (0.4%, without gaps) differences in ITS, 1/967 bp (0.1%, without gaps) differences in nrSSU, 2/750 bp (0.3%, without gaps) differences in nrLSU, and identical tef1-α sequences. Morphologically, the collection fits well with the description of Pse. puerense (MHZU 22-0153, holotype) in having solitary, immersed ascomata, 8-spored, cylindric-clavate to obovoid asci, and overlapping, hyaline, fusiform, septate ascospores constricted at the septa (Manawasinghe et al. 2025). Therefore, based on morphological and phylogenetic analyses, the collection is introduced here as a new host record of Pse. puerense, collected from Macadamia sp. in China.

Pseudochaetosphaeronema siamense

Jayasiri, E.B.G. Jones & K.D. Hyde [as ‘siamensis’], in Jayasiri et al., Mycosphere 10(1): 88 (2019)

4A83EDED-C85D-585D-A04B-F7DA9A22B0B8

648070

Figs 9 , 19

Figure 19. — *Pseudochaetosphaeronema* *siamense* (GMB-W 1573, new host record and country record). **a, b**. Appearance of ascomata on substrate; **c–e**. Vertical sections of ascomata; f. Peridium; g. Hamathecium; **h–k**. Asci; **l–o**. Ascospores; p. Germinated ascospore; **q, r**. Culture on PDA from above (q) and below (r). Scale bars: 200 μm (**d, e**); 50 μm (**f, h–k**); 30 μm (p); 20 μm (g); 10 μm (**l–o**); 2 cm (**q, r**).

Description.

Saprobic on a dead branch of Macadamia sp. Sexual morph: Ascomata 255–420 × 380–470 µm (x̄ = 327 × 355 µm, n = 10, including ostioles), solitary, immersed to erumpent, scattered, uni-loculate, dark brown to black, globose to subglobose, ostiolate, papillate. Ostioles up to 50–80 µm wide, neck, cylindrical to papillate. Peridium 15–50 µm wide, thin-walled, composed of several layers, brown to light brown cells of textura prismatica. Hamathecium composed of 1.5–3.0 µm wide (x̄ = 2.1 μm, n = 40), abundant, dense, narrowly cylindrical to filiform, hyaline, branched, pseudoparaphyses. Asci 80–130 × 15–25 µm (x̄ = 100 × 17.5 µm, n = 40), 4–8-spored, bitunicate, cylindric-clavate obovoid, straight to slightly curved, rounded at apex, pedicellate with club shaped base, with an apical chamber. Ascospores 23–33 × 7.5–12.5 µm (x̄ = 26.8 × 9.6 µm, n = 40), sometimes overlapping, fusiform to broadly fusiform, 1-septate at center, with pointed ends, with wider upper cell, hyaline, containing up to four guttules, smooth-walled, without mucilaginous sheath or appendages. Asexual morph: See Jayasiri et al. (2019).

Known hosts.

Macadamia sp. (this study), Tamarind sp. (Jayasiri et al. 2019).

Known distribution.

China (Yunnan Province; this study), Thailand (Jayasiri et al. 2019).

Material examined.

China • Yunnan Province, Dehong City, on a dead branch of Macadamia sp., April 2024, Xian Zhang, MYD6 (GMB-W 1573); living culture, GMBCC 2460, other living culture GMBCC 2461.

Notes.

Jayasiri et al. (2019) introduced Pseudochaetosphaeronema siamense from a fallen pod of Tamarind sp. (Fabaceae) in Thailand, based only on the asexual morph. In the phylogenetic analysis, the isolates GMBCC 2460 and GMBCC 2461 clustered with Pse. siamense (MFLUCC 17-2287) with 100% MLBS and 1.00 BYPP statistical support (Fig. 9). Based on nucleotide comparisons, the isolate GMBCC 2460 showed similarity to Pse. siamense (MFLUCC 17-2287), with 1/516 bp (0.1%, without gaps) difference in ITS, 1/830 bp (0.1%, without gaps) difference in nrLSU, and 1/1005 bp (0.1%, without gaps) difference in nrSSU. Morphologically, the collection revealed the sexual morph, which is almost identical to that of other species in the genus Pseudochaetosphaeronema, with immersed ascomata, bitunicate, fissitunicate asci with rounded ends, and fusiform, 1-septate, guttulate ascospores (Boonmee et al. 2021; Xu et al. 2024). Therefore, based on the phylogenetic analysis, the isolate is introduced here as a new host and country record of Pse. siamense, collected from Macadamia sp. in China. In addition, this is the first report of Pseudochaetosphaeronema siamense associated with Macadamia sp.

Helminthosporium puerense

L. Lu & Tibpromma, IMA Fungus 16(e144874): 22 (2025)

89E046CB-D5BF-56D8-8C38-D9D4BD48C9C4

901003

Figs 10 , 20

Figure 20. — *Helminthosporium* *puerense* (GMB-W 1574, new host record). **a–c**. Appearance of ascomata on substrate; d. Vertical section of ascoma; e. Ostiole; f. Peridium; g. Hamathecium; **h–k**. Asci; **l–o**. Ascospores; p. Germinated ascospore; **q, r**. Culture on PDA from above (q) and below (r). Scale bars: 200 μm (d); 100 μm (e); 50 μm (**f, i–k**); 30 μm (**g, h**); 10 μm (**l–o**); 2 cm (**q, r**).

Holotype.

HKAS 137606

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morph: Ascomata 250–530 × 330–500 µm (x̄ = 403 × 381 µm, n = 10, including ostioles), solitary to scattered, immersed, slightly erumpent, dark spots with black hair on the host surface, globose to subglobose, uniloculate, centrally ostiolate, papillate. Ostioles neck central, cylindrical to papillate, surrounded by a dark brown clypeus-like structure, without periphyses. Peridium 35–75 µm thin to thick-walled, composed of hyaline to brown to dark brown cells of textura angularis. Hamathecium composed of 1.5–3.0 µm wide (x̄ = 2.3 μm, n = 40), abundant, dense, narrowly cylindrical to filiform, hyaline, septate, branched, pseudoparaphyses. Asci 90–160 × 20–30 µm (x̄ = 118 × 23 µm, n = 40), 8-spored uni- to biseriate, bitunicate, fissitunicate, clavate, straight to slightly curved, rounded at apex, short-stalked with a club-shaped base, with a narrow apical chamber. Ascospores 25–32 × 7–11 µm (x̄ = 27.5 × 9 µm, n = 40), ellipsoidal to oblong, mostly straight, 1–3-septate with constriction at the septum, hyaline, with subacute to rounded end cells, asymmetric, with wider upper cell, guttulate, smooth-walled, with a thick sheath. Asexual morph: Not observed.

Culture characteristics.

Ascospores germinating on PDA within 24 h at 28 °C, turning light gray after germination. Colonies on PDA reaching 3.6 cm in diam. after 3 weeks of incubation at 28 °C, with a rough surface; mycelium with filamentous, hairy hyphae, circular, cottony, light yellow from above, surrounded near the center by a yellow to green to light gray ring, and light yellow to brown from the edge to the center; colonies with cracks in the middle.

Known hosts.

Coffea arabica (Lu et al. 2025a), Macadamia sp. (this study).

Known distribution.

China (Yunnan Province; Lu et al. 2025a; this study).

Materials examined.

China • Yunnan Province, Pu’er City, on dead twigs of Macadamia sp., 1 August 2024, Xian Zhang, MYP16 (GMB-W 1574); living culture, GMBCC 2462, • ibid., MYP24 (GMB-W 1575), living culture, GMBCC 2464.

Notes.

In the phylogenetic analyses, the isolates GMBCC 2462 and GMBCC 2464 clustered within Helminthosporium, sister to H. puerense (KUNCC 24-18347), with 82% MLBS and 1.00 BYPP statistical support (Fig. 10). Morphologically, the isolates resemble H. puerense (HKAS 137606, holotype) in having immersed ascomata, fissitunicate, bitunicate asci, and 1–3-septate ascospores with wider upper cells. However, the isolates differ from H. puerense (HKAS 137606, holotype) by having ascospores with subacute to rounded end cells and a conspicuous sheath and by having hyaline ascospores that turn light gray after germination (Lu et al. 2025a). Based on nucleotide comparisons, the isolates differ from H. puerense by 8/549 bp (1.46%, without gaps) in ITS, 2/834 bp (0.2%, without gaps) in nrLSU, no differences in nrSSU, 2/1043 bp (0.2%, without gaps) in rpb2, and 3/744 bp (0.4%, without gaps) in tef1-α. Therefore, Helminthosporium puerense is introduced here as a new host record, collected from Macadamia sp. in China.

Helminthosporium nanjingense

Meng Zhang, Xiao J. Wang & H.Y. Wu, Mycotaxon 127: 2 (2014)

C2579CD2-F5A1-5A2D-AA40-CA79AE10B934

804655

Figs 10 , 21

Figure 21. — *Helminthosporium* *nanjingense* (GMB-W 1576, new host record). **a–c**. Appearance of ascomata on substrate; d. Vertical section of ascoma; e. Peridium; f. Hamathecium; **g–k**. Asci; **l–q**. Ascospores (q: stained with Indian ink); r. Germinated ascospore; **s, t**. Culture on PDA from above (s) and below (t). Scale bars: 200 μm (d); 50 μm (e); 30 μm (**g–k, r**); 20 μm (**f, n–q**); 10 μm (**l, m**); 2 cm (**s, t**).

Description.

Saprobic on dead twigs of Macadamia sp. Sexual morph: Ascomata 200–310 µm high × 200–375 µm diam. (x̄ = 251 × 292 µm, n = 15, including ostioles), solitary to scattered, immersed, visible as dark spots with a small amount of black hairs on host surface, subglobose to pyriform, uniloculate, ostiolate, with a papilla. Ostioles neck central, cylindrical to papillate, surrounded by a dark brown clypeus-like structure, without periphyses. Peridium 25–50 µm thin to thick-walled, composed of hyaline to brown to dark brown cells of textura angularis. Hamathecium composed of abundant 2–3.5 µm wide (x̄ = 2.5 μm, n = 40), hyaline, septate, branched, dense, narrowly cylindrical to filiform, pseudoparaphyses. Asci 75–130 × 17–27 µm (x̄ = 99.6 × 20.5 µm, n = 30), 2–8-spored, bitunicate, fissitunicate, straight to slightly curved, clavate, rounded at apex, short-stalked, with a narrow apical chamber. Ascospores 25–33 × 6.5–11 µm (x̄ = 28.7 × 8.3 µm, n = 40), oblong-ellipsoidal to fusiform, straight to slight curved, 1–3-septate, strongly constricted at middle septum, slightly or not constricted at both side septa, asymmetric, with wider upper cell, hyaline, with subacute to rounded end cells, smooth-walled, with a sheath on surface, guttulate. Asexual morph: See Wang et al. (2014).

Culture characteristics.

Ascospores germinating on PDA within 24 h at 28 °C, turning light gray after germination. Colonies on PDA reaching 3.3 cm in diam. after 3 weeks at 28 °C, with a rough surface; mycelium composed of filamentous, hairy hyphae, circular with an irregular margin, cottony, white from above, surrounded near the center by a slightly yellowish to pale cream coloration, and irregularly light yellow to brown from the edge to the center; colonies with light cracks in the middle.

Known hosts.

Macadamia sp. (this study), on dead branches of an unidentified tree (Wang et al. 2014).

Known distribution.

China (Jiangsu Province; Wang et al. 2014, Yunnan Province; this study).

Material examined.

China • Yunnan Province, Pu’er City, on dead twigs of Macadamia sp., 1 August 2024, Xian Zhang, MYP10 (GMB-W 1576); living culture, GMBCC 2466, other living culture GMBCC 2467.

Notes.

In the phylogenetic tree, the isolates GMBCC 2466 and GMBCC 2467 were nested with Helminthosporium nanjingense (HHAUF020380) with 89% MLBS and 1.00 BYPP statistical support (Fig. 10). Based on pairwise nucleotide comparisons, the isolates differ from H. nanjingense (HHAUF020380) by 4/454 bp (0.9%, without gaps) in ITS. Morphologically, Wang et al. (2014) described the asexual morph of Helminthosporium nanjingense. In this study, the isolates exhibited the sexual morph. Helminthosporium chinense (HKAS 124017, holotype) has only been recorded from its asexual morph; therefore, morphological comparison with this species was not possible. The isolates can be distinguished from H. puerense (HKAS 137606, holotype) by having subglobose to pyriform ascomata, 2–8-spored, straight to slightly curved asci, and ascospores with conspicuous sheaths, whereas H. puerense has globose to subglobose ascomata, 2–8-spored asci, ascospores with inconspicuous sheaths, and sometimes long stipes (Lu et al. 2025a). The phylogenetic tree also shows that they are distinct species (Fig. 10). Therefore, based on the phylogenetic analysis, the isolates are introduced as a new host record of H. nanjingense, collected from Macadamia sp. in China. This is the first report of Helminthosporium nanjingense associated with Macadamia sp.

Evaluation of lignocellulitic enzyme production

In this study, 20 pure cultures representing 18 species of saprobic fungi obtained from Macadamia specimens collected in China and Thailand were evaluated for their ability to produce lignocellulolytic enzymes, including xylanase, cellulase, mannanase, and laccase. The results showed that the lignocellulolytic enzyme production profiles of 17 fungal isolates varied considerably among the four tested enzymes, whereas three isolates, Dothiorella lignicola SDBR-CMU873, Lasiodiplodia ponkanicola SDBR-CMU875, and Lasiodiplodia endophytica SDBR-CMU876, showed no detectable production of any of the four tested enzymes. A total of 17 isolates representing 15 species exhibited positive xylanase production, 14 isolates representing 12 species exhibited positive cellulase production, and 10 isolates representing nine species exhibited positive mannanase production, as determined by the Congo red assay (Fig. 22). In addition, five isolates representing five species exhibited positive laccase activity, as indicated by the formation of a brownish coloration in the culture medium.

Figure 22. — Plate assay for lignocellulolytic enzyme production by saprobic fungi isolated from *Macadamia* sp. in this study. a. Xylanase production detected by the formation of clear hydrolysis zones after Congo red staining (indicated by arrowheads); b. Cellulase production indicated by the formation of clear hydrolysis zones after Congo red staining (indicated by arrowheads); c. Mannanase production indicated by the formation of clear hydrolysis zones after Congo red staining (indicated by arrowheads); d. Laccase production indicated by reddish-brown coloration on guaiacol-containing medium. Representative photographs of the three isolates showing the highest enzyme production in each assay are presented. Scale bars: 10 mm.

The enzyme potency index (EPI) was calculated to quantify the relative enzyme-producing ability of the fungal isolates, and the results are presented in Fig. 23. Xylanase production was detected in 17 cultures representing 15 species. The highest xylanase activity was observed in Xylochrysis macadamiaeGMBCC 2440 (EPI = 3.50), followed by Sporidesmium tropicaleGMBCC 2444 (EPI = 3.10) and Pseudochaetosphaeronema chiangraienseGMBCC 2456 (EPI = 2.62). Cellulase production was detected in 14 cultures representing 12 species, with the highest activity observed in Helminthosporium puerenseGMBCC 2451 (EPI = 2.25), followed by H. nanjingenseGMBCC 2466 (EPI = 1.92) and H. puerenseGMBCC 2450 (EPI = 1.66). Additionally, mannanase production was detected in 10 cultures representing nine species, with the highest activity observed in Pseudochaetosphaeronema puerenseGMBCC 2458 (EPI = 1.98), followed by Pse. siamenseGMBCC 2460 (EPI = 1.96) and Crassiparies quadrisporusGMBCC 2453 (EPI = 1.37). For laccase production, five isolates representing four species exhibited positive activity, with the highest activity observed in Sporidesmium tropicaleGMBCC 2444 (EPI = 2.20), followed by Pseudochaetosphaeronema chiangraienseGMBCC 2456 (EPI = 1.70) and Crassiparies quadrisporusGMBCC 2452 (EPI = 1.42). Overall, four isolates, Crassiparies quadrisporusGMBCC 2452 and GMBCC 2453, Pseudochaetosphaeronema chiangraienseGMBCC 2456, and Sporidesmium tropicaleGMBCC 2444, exhibited the ability to produce all four tested lignocellulolytic enzymes.

Figure 23. — Heat map showing the enzyme potency index for xylanase, cellulase, mannanase, and laccase production by saprobic fungi isolated from macadamia. Data are presented as means ± standard deviation from three replicates. Different letters within each fungal isolate indicate significant differences among enzyme activities (p ≤ 0.05), as determined by Duncan’s multiple range test (DMRT).

Discussion

Saprobic fungi play a crucial role in ecosystem nutrient cycling by decomposing organic matter and releasing nutrients (Sharma and Arora 2015; Corbu et al. 2023; Eliopoulos et al. 2025). However, research on macadamia-associated saprobic fungi remains limited. In this study, saprobic fungi associated with macadamia were collected from Yunnan Province, China, and Chiang Mai Province, Thailand. Based on a combination of multilocus phylogenetic analyses and morphological characteristics, 18 fungal taxa were identified, comprising five newly introduced species (Dothiorella lignicola, Lasiodiplodia macadamiae, Palmiascoma macadamiae, Xylochrysis macadamiae, and X. yunnanensis) and 13 previously known species (Chromolaenicola ananasi, Crassiparies quadrisporus, Helminthosporium nanjingense, H. puerense, Lasiodiplodia ponkanicola, L. endophytica, Montagnula donacina, M. thailandica, Palmiascoma chiangraiense, Pseudochaetosphaeronema chiangraiense, Pse. puerense, Pse. siamense, and Sporidesmium tropicale). The identified fungi belonged to seven families and 10 genera: Chromolaenicola, Crassiparies, Dothiorella, Helminthosporium, Lasiodiplodia, Montagnula, Palmiascoma, Pseudochaetosphaeronema, Sporidesmium, and Xylochrysis. Among these, eight genera (Chromolaenicola, Crassiparies, Helminthosporium, Montagnula, Palmiascoma, Pseudochaetosphaeronema, Sporidesmium, and Xylochrysis) are reported from macadamia for the first time, whereas Lasiodiplodia and Dothiorella have previously been documented from this host. All 13 known species represent new host records on macadamia. Notably, four species are reported here as new geographical records. Furthermore, the sexual morphs of three species, Chromolaenicola ananasi, Pseudochaetosphaeronema siamense, and Helminthosporium nanjingense, were observed for the first time in this study.

This study shows that saprobic fungi associated with Macadamia spp. constitute a diverse source of lignocellulolytic enzymes, although their enzymatic capabilities differ markedly among taxa. Such variation in enzyme production is expected because the degradation of plant biomass requires a complex enzymatic system, and different fungal species have evolved distinct strategies for utilizing lignocellulosic substrates according to their ecological niches and nutritional requirements (Sánchez-Corzo et al. 2021; Bhadra et al. 2022; Khan et al. 2025; Luong et al. 2025;). Among the enzymes evaluated, xylanase was the most frequently detected, followed by cellulase and mannanase, whereas only a few isolates showed laccase activity. These results are consistent with similar trends reported for numerous saprobic ascomycetes, in which xylanase production exceeds cellulase and lignin-degrading enzyme activities during the early stages of plant litter decomposition (Afshari et al. 2025a; Wang et al. 2026). The predominance of xylanase production suggests that hemicellulose degradation represents an important initial step during the decomposition of macadamia tissues. Hemicellulose is generally more accessible and less crystalline than cellulose, allowing fungi to utilize it before attacking the more recalcitrant cellulose and lignin components (Scheller and Ulvskov 2010; Marriott et al. 2016; Petridis and Smith 2018; Sun et al. 2023). The high xylanase activity exhibited by Xylochrysis macadamiaeGMBCC 2440, Sporidesmium tropicaleGMBCC 2444, and Pseudochaetosphaeronema chiangraienseGMBCC 2456 indicates that these species possess considerable potential for hemicellulose degradation. Likewise, the strong cellulase activities observed in Helminthosporium puerense and H. nanjingense suggest that members of Helminthosporium may play an important role in cellulose decomposition within macadamia ecosystems. Mannanase activity was less common than xylanase and cellulase but was particularly high in Pseudochaetosphaeronema puerense and Pse. siamense. Laccase production was detected in only five isolates, Crassiparies quadrisporusGMBCC 2453, Crassiparies quadrisporusGMBCC 2452, Montagnula donacinaGMBCC 2455, Pseudochaetosphaeronema chiangraienseGMBCC 2456, and Sporidesmium tropicaleGMBCC 2444. Similarly, previous studies have demonstrated that mannanase and laccase activities are generally less prevalent than xylanase and cellulase activities. This pattern may be attributed to the relatively lower abundance of mannan compared with cellulose and xylan in plant cell walls, together with the complex structure of lignin, which is highly resistant to degradation and requires specialized oxidative enzymes for its breakdown (Scheller and Ulvskov 2010; Janusz et al. 2017; Petridis and Smith 2018; Liu et al. 2022; Curry et al. 2023). In contrast, three isolates, Dothiorella lignicola SDBR-CMU873, Lasiodiplodia ponkanicola SDBR-CMU875, and Lasiodiplodia endophytica SDBR-CMU876, showed no detectable activity for any of the four enzymes under the experimental conditions. This lack of detectable activity may reflect enzyme production below the detection limit of the agar plate assay or preferential utilization of readily available simple sugars in the substrate, reducing the need to produce extracellular lignocellulolytic enzymes (Benocci et al. 2017; Saroj and Narasimhulu 2018; Peng et al. 2021). Moreover, extracellular enzyme production depends heavily on substrate composition, nutrient availability, pH, temperature, and incubation time (Loeppmann et al. 2016; Liu et al. 2019; Sharma et al. 2024).

Previous studies on fungi associated with Macadamia have primarily focused on pathogenic fungi, with relatively few investigations of endophytic fungi, whereas saprobic fungi have received comparatively little attention (Akinsanmi and Drenth 2013; Wrona et al. 2020; Zhang et al. 2025c). This study expands the current knowledge of saprobic fungi associated with Macadamia spp. in China and Thailand by documenting their diversity and evaluating their lignocellulolytic enzyme-producing potential. Most isolates produced at least one lignocellulolytic enzyme, although enzyme production varied considerably among species. Xylanase was the most frequently detected enzyme, followed by cellulase and mannanase, whereas laccase activity was restricted to a limited number of isolates. Notably, Crassiparies quadrisporus, Pseudochaetosphaeronema chiangraiense, and Sporidesmium tropicale exhibited the ability to produce all four tested enzymes, highlighting their potential as promising sources of lignocellulolytic enzymes for biotechnological applications. These findings not only improve understanding of the ecological roles of saprobic fungi in lignocellulose decomposition but also demonstrate that macadamia-associated saprobic fungi represent an underexplored reservoir of industrially relevant enzymes. Further studies should focus on quantitative enzyme assays, optimization of culture conditions, purification and biochemical characterization of the enzymes, and evaluation of their potential for industrial biomass conversion and other biotechnological applications.

Supplementary Material

XML Treatment for Xylochrysis macadamiae

mycokeys.141.209886-treatment1.xml^{(19.7KB, xml)}

XML Treatment for Xylochrysis yunnanensis

mycokeys.141.209886-treatment2.xml^{(17.8KB, xml)}

XML Treatment for Sporidesmium tropicale

mycokeys.141.209886-treatment3.xml^{(13.4KB, xml)}

XML Treatment for Dothiorella lignicola

mycokeys.141.209886-treatment4.xml^{(33.7KB, xml)}

XML Treatment for Lasiodiplodia macadamiae

mycokeys.141.209886-treatment5.xml^{(28.9KB, xml)}

XML Treatment for Lasiodiplodia ponkanicola

mycokeys.141.209886-treatment6.xml^{(9.5KB, xml)}

XML Treatment for Lasiodiplodia endophytica

mycokeys.141.209886-treatment7.xml^{(14.9KB, xml)}

XML Treatment for Palmiascoma macadamiae

mycokeys.141.209886-treatment8.xml^{(20.5KB, xml)}

XML Treatment for Palmiascoma chiangraiense

mycokeys.141.209886-treatment9.xml^{(16.9KB, xml)}

XML Treatment for Crassiparies quadrisporus

mycokeys.141.209886-treatment10.xml^{(13.3KB, xml)}

XML Treatment for Chromolaenicola ananasi

mycokeys.141.209886-treatment11.xml^{(15.8KB, xml)}

XML Treatment for Montagnula donacina

mycokeys.141.209886-treatment12.xml^{(27.7KB, xml)}

XML Treatment for Montagnula thailandica

mycokeys.141.209886-treatment13.xml^{(12.4KB, xml)}

XML Treatment for Pseudochaetosphaeronema chiangraiense

mycokeys.141.209886-treatment14.xml^{(10.5KB, xml)}

XML Treatment for Pseudochaetosphaeronema puerense

mycokeys.141.209886-treatment15.xml^{(11.8KB, xml)}

XML Treatment for Pseudochaetosphaeronema siamense

mycokeys.141.209886-treatment16.xml^{(16.1KB, xml)}

XML Treatment for Helminthosporium puerense

mycokeys.141.209886-treatment17.xml^{(14.1KB, xml)}

XML Treatment for Helminthosporium nanjingense

mycokeys.141.209886-treatment18.xml^{(15.3KB, xml)}

Acknowledgements

The authors thank Dr. Shaun Pennycook for suggestions regarding the Latin names of the new taxa.

Citation

Zhang X, Karunarathna SC, Tibpromma S, Senwanna C, Kumla J, Du T-Y, Elgorban AM, Alfagham AT, Filimban FZ, Dai D-Q, Suwannarach N (2026) New species and additional records of saprobic microfungi associated with Macadamia in China and Thailand and their lignocellulitic enzyme production. MycoKeys 141: 217–281. https://doi.org/10.3897/mycokeys.141.209886

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Artificial Intelligence (AI) use

The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.

No AI tools were used in the preparation of this manuscript.

Funding

Nakarin Suwannarach and Samantha C. Karunarathna thank the CMU Visiting Researcher 2025 program, Chiang Mai University, Thailand (Grant No. CMUVP048/2568). This work was partially supported by the Yunnan Revitalization Talents Support Plan (Young Talents Program and High-End Foreign Experts Program), the Yunnan Provincial Department of Science and Technology “Zhihui Yunnan” Plan (202403AM140023), the National Natural Science Foundation of China (Nos. 32260004 and 3246002), Yunnan Fundamental Research Projects (Grant No. 202201AU070017), the Special Basic Cooperative Research Programs of Yunnan Provincial Undergraduate Universities (Grant Nos. 202101BA070001-209 and 202101BA070001-279), Mee-mann Chang Academician Workstation in Yunnan Province (Grant No. 202205AF150002), the Ongoing Research Funding program (ORF-RICSP-2026-6), King Saud University, Riyadh, Saudi Arabia, and the Key Laboratory of Yunnan Provincial Department of Education of the Deep-Time Evolution on Biodiversity from the Origin of the Pearl River.

Author contributions

Conceptualization: NS, ST. Data curation: XZ, SCK, JK, ST, NS. Formal analysis: XZ, CS, JK, T-YD. Funding acquisition: NS, SCK. Investigation: XZ, CS, JK, T-YD, ST, NS. Methodology: XZ, SCK, CS, JK, T-YD, ST, NS. Project administration: NS, ST. Software: XZ, JK, ST, NS. Supervision: ST, NS. Validation: SCK, ST, NS. Visualization: XZ, SCK, CS, JK, T-YD, D-QD, ST, NS. Writing – original draft: XZ, SCK, JK, ST, NS. Writing – review and editing: XZ, SCK, CS, JK, T-YD, AME, ATA, FZF, D-QD, ST, NS.

Author ORCIDs

X. Zhang https://orcid.org/0000-0001-6097-8922

S.C. Karunarathna https://orcid.org/0000-0001-7080-0781

S. Tibpromma https://orcid.org/0000-0002-4706-6547

C. Senwanna https://orcid.org/0000-0002-1008-4514

J. Kumla https://orcid.org/0000-0002-3673-6541

T.-Y. Du https://orcid.org/0000-0003-2105-1803

A.M. Elgorban https://orcid.org/0000-0003-3664-7853

A.T. Alfagham https://orcid.org/0000-0001-9008-3679

F.Z. Filimban https://orcid.org/0000-0001-7527-1313

D.-Q. Dai https://orcid.org/0000-0001-8935-8807

N. Suwannarach https://orcid.org/0000-0002-2653-1913

Data availability

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

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Xylochrysis macadamiae

mycokeys.141.209886-treatment1.xml^{(19.7KB, xml)}

XML Treatment for Xylochrysis yunnanensis

mycokeys.141.209886-treatment2.xml^{(17.8KB, xml)}

XML Treatment for Sporidesmium tropicale

mycokeys.141.209886-treatment3.xml^{(13.4KB, xml)}

XML Treatment for Dothiorella lignicola

mycokeys.141.209886-treatment4.xml^{(33.7KB, xml)}

XML Treatment for Lasiodiplodia macadamiae

mycokeys.141.209886-treatment5.xml^{(28.9KB, xml)}

XML Treatment for Lasiodiplodia ponkanicola

mycokeys.141.209886-treatment6.xml^{(9.5KB, xml)}

XML Treatment for Lasiodiplodia endophytica

mycokeys.141.209886-treatment7.xml^{(14.9KB, xml)}

XML Treatment for Palmiascoma macadamiae

mycokeys.141.209886-treatment8.xml^{(20.5KB, xml)}

XML Treatment for Palmiascoma chiangraiense

mycokeys.141.209886-treatment9.xml^{(16.9KB, xml)}

XML Treatment for Crassiparies quadrisporus

mycokeys.141.209886-treatment10.xml^{(13.3KB, xml)}

XML Treatment for Chromolaenicola ananasi

mycokeys.141.209886-treatment11.xml^{(15.8KB, xml)}

XML Treatment for Montagnula donacina

mycokeys.141.209886-treatment12.xml^{(27.7KB, xml)}

XML Treatment for Montagnula thailandica

mycokeys.141.209886-treatment13.xml^{(12.4KB, xml)}

XML Treatment for Pseudochaetosphaeronema chiangraiense

mycokeys.141.209886-treatment14.xml^{(10.5KB, xml)}

XML Treatment for Pseudochaetosphaeronema puerense

mycokeys.141.209886-treatment15.xml^{(11.8KB, xml)}

XML Treatment for Pseudochaetosphaeronema siamense

mycokeys.141.209886-treatment16.xml^{(16.1KB, xml)}

XML Treatment for Helminthosporium puerense

mycokeys.141.209886-treatment17.xml^{(14.1KB, xml)}

XML Treatment for Helminthosporium nanjingense

mycokeys.141.209886-treatment18.xml^{(15.3KB, xml)}

Data Availability Statement

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

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PERMALINK

New species and additional records of saprobic microfungi associated with Macadamia in China and Thailand and their lignocellulitic enzyme production

Xian Zhang

Samantha C Karunarathna

Saowaluck Tibpromma

Chanokned Senwanna

Jaturong Kumla

Tian-Ye Du

Abdallah M Elgorban

Alanoud T Alfagham

Faten Zubair Filimban

Dong-Qin Dai

Nakarin Suwannarach

Roles

Abstract

Introduction

Table 1.

Materials and methods

Sample collection, fungal isolation, and morphological study

DNA extraction, PCR amplification, and sequencing

Table 2.

Phylogenetic analyses

Table 3.

Evaluation of lignocellulolytic enzyme production

Statistical analysis

Results

Phylogenetic analyses

Figure 1.

Figure 2.

Figure 3.

Figure 4.

Figure 5.

Figure 6.

Figure 7.

Figure 8.

Figure 9.

Figure 10.

Taxonomy

Xylochrysis macadamiae

Figure 11.

Etymology.

Diagnosis.

Holotype.

Description.

Culture characteristics.

Material examined.

Notes.

Xylochrysis yunnanensis

Figure 12.

Etymology.

Diagnosis.

Holotype.

Description.

Culture characteristics.

Materials examined.

Notes.

Sporidesmium tropicale

Description.

Known hosts.

Known distribution.

Materials examined.

Notes.

Dothiorella lignicola

Figure 13.

Etymology.

Diagnosis.

Holotype.

Description.

Culture characteristics.

Materials examined.

Notes.

Table 4.

Lasiodiplodia macadamiae

Figure 14.

Etymology.

Diagnosis.

Holotype.

Description.

Culture characteristics.

Material examined.