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. 2026 Sep 21;140:241–259. doi: 10.3897/mycokeys.140.206566

Phylogenetic analyses and morphological studies reveal two new species of Biscogniauxia (Xylariales, Ascomycota) from Hainan Tropical Rainforest National Park, China

Si-Yu Ma 1,2, Shuang Liu 1,2, Zhi Qu 1, Huan Jiang 1,2, An-Hong Zhu 1,3,4,✉, Hai-Xia Ma 1,5,✉
PMCID: PMC13620813  PMID: 42812803

Abstract

The species diversity of the family Graphostromataceae is poorly understood in tropical regions. The present work concerns a survey of wood-inhabiting macrofungi from the Hainan Tropical Rainforest, which is located in south-central Hainan Province. This study introduces two new species of Biscogniauxia, viz., B. diaoluoshanensissp. nov. and B. jianfengensissp. nov., which are herein described based on morphological and molecular evidence. Morphologically, B. diaoluoshanensis is characterized by applanate stromata, ovoid to pyriform perithecia, and ascospores with a conspicuous straight, almost spore-length germ slit. Biscogniauxia jianfengensis is distinguished by applanate to slightly pulvinate stromata, tubular perithecia, ascospores with an inconspicuous, unilateral, straight, spore-length germ slit, bearing an opaque cellular appendage on one side. The phylogenetic analysis inferred from ITS, RPB2 and TUB2 sequence data confirmed that the two new species are distinct within the genus Biscogniauxia. Detailed descriptions, illustrations, and phylogenetic analysis results are provided for each new species, and a dichotomous identification key is presented for the genus as well.

Key words: Graphostromataceae , phylogeny, species diversity, tropical rainforest, wood-inhabiting fungi

Introduction

Kuntze (1891) established Biscogniauxia Kuntze with B. nummularia (Bull.) Kuntze as the type species. It is characterized by corticolous, carbonaceous bipartite stromata developing underneath the bark of the host and breaking when mature, unitunicate asci with an amyloid apical apparatus, and ellipsoid brown ascospores with germ slits and usually lacking a slimy sheath or slimy appendages (Ju et al. 1998; Fournier et al. 2017). The genus Biscogniauxia was resurrected and amended by Pouzar (1979, 1986) and accepted the taxa formerly placed in Nummularia Tul. & C. Tul. (Nom. illegit., Art. 53.1 fideIndexFungorum 2026). Previously, the generic concept of Biscogniauxia was based only on morphological characteristics and was assigned to Xylariaceae Tul & C. Tul (Ju et al. 1998; Vasilyeva 2007). Based on multigene phylogenetic analyses, Wendt et al. (2018) segregated the genus Biscogniauxia from Xylariaceae and transferred it to Graphostromataceae with Graphostroma Piroz., Camillea Fr., Obolarina Pouzar, and Vivantia J.D. Rogers, Y.M. Ju & Cand.

The genus Biscogniauxia is the largest genus in the Graphostromataceae, and has been recorded worldwide, including North America (San Martín González and Rogers 1993), Africa (Mugambi et al. 2009; Linnakoski et al. 2012), Europe (Cannon et al. 1985; Rappaz 1995; Ragazzi 2009; Ragazzi et al. 2012), Asia (Whalley et al. 2012; Jantaharn et al. 2021; Samarakoon et al. 2022; Chen et al. 2024; Qiao et al. 2024; Liu et al. 2025) and Oceania (Tan et al. 2025a, 2025b). Species of Biscogniauxia are widely known as saprobes, but some are plant pathogens causing diseases of dicotyledonous angiosperm trees including Acacia, Acer, Alnus, Artocarpus, Carya, Celtis, Coprosma, Cydonia, Eucalyptus, Fagus, Fraxinus, Gluta, Lithocarpus, Padus, Phyllirea, Pisonia, Populus, Prunus, Psidium, Pyrus, Quercus, Rhamnus, Rubus, and Tilia (e.g. Rappaz 1995; Ju and Rogers 1996, 2001; Rogers et al. 2000; Whalley et al. 2000; Granata and Sidoti 2004; Nugent et al. 2005; Vasilyeva 2007; Mugambi et al. 2009; Vasilyeva et al. 2012; Raimondo et al. 2016), and some are endophytes which have the potential to become pathogens when their host plants are old and stressed (Helaly et al. 2018). In addition, some species are important producers of novel bioactive compounds and secondary metabolites (Purbaya et al. 2023). Therefore, Biscogniauxia species are of great interest due to their ecological and economic significance.

One hundred and twenty records of Biscogniauxia are listed in Index Fungorum (http://www.indexfungorum.org/names/Names.asp/, accessed on 5 August 2026), and so far, more than 65 species are widely accepted (Hyde et al. 2020; Ma et al. 2020; Perera 2020; Vujanovic et al. 2020; Jantaharn et al. 2021; Samarakoon et al. 2022; Chen et al. 2024; Qiao et al. 2024; Liu et al. 2025; Tan et al. 2025a, 2025b; Ren et al. 2026; Wongkanoun et al. 2026). Among these, 17 species and 4 varieties (the formal infraspecific taxonomic rank, abbreviated as var.) have been reported and described from China, including B. ambiens Y.M. Ju & J.D. Rogers, B. betulae S.H. Long, K. Habib & Q.R. Li, B. capnodes (Berk.) Y.M. Ju & J.D. Rogers, B. citriformis (Whalley, Hammelev & Taligoola) Van der Gucht & Whalley, B. cylindrispora Y.M. Ju & J.D. Rogers, B. dendrobii X.Y. Ma, K.D. Hyde & J.C. Kang, B. dicranopteridis X.L. Ding, Chang Xia Qiao & D.W. Li, B. effusa Q.R. Li, J.C. Kang & K.D. Hyde, B. formosana Y.M. Ju & J.D. Rogers, B. glaucae Q.R. Li & J.C. Kang, B. latirima Y.M. Ju & J.D. Rogers, B. petrensis Z.F. Zhang, F. Liu & L. Cai, B. philippinensis (Ricker) Whalley & Læssøe, B. plana (Petch) Y.M. Ju & J.D. Rogers, B. sinensis X.L. Ding, Chang Xia Qiao & D.W. Li, B. uniapiculata (Penz. & Sacc.) Y.M. Ju & J.D. Rogers, B. xishuangbannaensis G.C. Ren & Wanas., B. albosticta var. orientalis Y.M. Ju & J.D. Rogers, B. capnodes (Berk.) Y.M. Ju & J.D. Rogers var. rumpens (Cooke) Y.M. Ju & J.D. Rogers, B. formosana var. kentingensis Y.M. Ju & J.D. Rogers, and B. philippinensis var. microspora Y.M. Ju & J.D. Rogers (Ju et al. 1998; Ju and Rogers 2001; Ariyawansa et al. 2015; Zhang et al. 2017; Ma et al. 2020; Li et al. 2021; Chen et al. 2024; Qiao et al. 2024; Liu et al. 2025; Ren et al. 2026). However, collections from tropical regions of China are few, and many undescribed novel species probably exist and are expected to be discovered. During the investigations of the diversity of wood-decaying fungi in Hainan Tropical Rainforest National Park, several specimens of Xylariales were collected. Two novel species of Biscogniauxia are identified using morphological evidence and molecular phylogenetic analyses, and a dichotomous identification key to Biscogniauxia taxa in China is proposed in this survey. The study makes a contribution to our understanding of the species diversity of Graphostromataceae in tropical China and reveals the phylogenetic relationships amongst Biscogniauxia species.

Materials and methods

Specimen collection and morphological study

The National Park of Hainan Tropical Rainforest lies in south-central Hainan Province, covering the coordinate range of 18°33'16"– 19°14'16"N and 108°44'32"– 110°04'43"E, with a tropical monsoon climate. The study area harbours 4,367 higher plant species belonging to 1,368 genera and 289 families. The dead branches bearing fungal stromata were collected from the National Park. Fresh specimens were dried, labeled, and transported for morphological and molecular studies. The studied specimens are deposited in the Fungarium of the Institute of Tropical Bioscience and Biotechnology, Chinese Academy of Tropical Agricultural Sciences (FCATAS).

The macro-morphological descriptions were observed from field notes and photos captured in the field. The stromatal surface and perithecia were observed and measured using a VHX-6000 3D microscope of the Keyence Corporation (Osaka, Japan). The micro-morphological data were obtained from dried specimens observed under an Olympus IX73 inverted fluorescence microscope (Tokyo, Japan) with a 10 × 100 oil immersion objective (Ma et al. 2022a). Sections mounted in water, Melzer’s reagent, and KOH were used to observe micro-morphological characters, following the method described by Zhu et al. (2024a). At least 30 asci and 50 ascospores from each specimen were measured randomly. The following abbreviations are used: x̄ = arithmetical average length of all ascospores × arithmetical average width of all ascospores, and n = a number of ascospores measured from a number of specimens. When presenting the variation in ascospore sizes, 5% of measurements were excluded from each end of the range and given in parentheses.

DNA extraction, PCR amplification, and sequencing

Genomic DNA was extracted from the dried specimens using a Plant DNA Rapid Extraction Kit (Magen) according to the manufacturer’s instructions. Primer pairs used for amplification were nuclear rDNA region encompassing the internal transcribed spacer (ITS), along with the 5.8S rDNA with ITS5/ITS4 (White et al. 1990; Gardes and Bruns 1993), the portion of the β-tubulin gene (TUB2) with Bt2a/Bt2b (Glass and Donaldson 1995), and the RNA polymerase II subunit (RPB2) with RPB2-5F/RPB2-7cR (Liu et al. 1999). The PCR procedure was as follows: initial denaturation at 95 °C for 2 min, followed by 35 cycles of 95 °C for 30 s, annealing at 52 °C (ITS), 61 °C (TUB2), and 54 °C (RPB2) for 45 s each, extension at 72 °C for 1 min, and a final extension at 72 °C for 10 min. DNA sequencing was performed at Beijing Tsingke Biotechnology Co., Ltd., and all the newly generated sequences were uploaded to NCBI GenBank and listed in Table 1.

Table 1.

List of taxa used for phylogenetic reconstruction. GenBank accession numbers, specimen numbers, host, origin, and reference studies are given. Holotype specimens are labelled with T. Species highlighted in bold were derived from this study. NA: not available.

Species Sample No. Host Locality ITS RPB2 TUB2 Reference
Biscogniauxia anceps YMJ 123 Corylus avellana France NA JX507777 AY951671 Hsieh et al. 2010
B. arima YMJ 122T Unknown Mexico EF026150 GQ304736 AY951672 Crous et al. 2023
B. atropunctata YMJ 128 Unknown USA JX507799 JX507778 AY951673 Hsieh et al. 2010
B. australiana BRIP 66979aT Croton insularis Australia PV074480 NA NA Tan et al. 2025a
B. atropunctata var. intermedia B70M Quercus sp. Costa Rica AJ390412 NA NA Sánchez-Ballesteros et al. 2000
B. atropunctata var. intermedia ATCC 38987 Unknown Unknown AF201705 NA NA Pinto 2001
B. bartholomaei ATCC 38992 Unknown USA AF201719 NA NA Pinto 2001
B. betulae GMB 0711T Unknown Unknown PQ884683 PQ893615 NA Liu et al. 2025
B. capnodes YMJ 142 Nothofagus solandri New Zealand NA NA AY951674 Hsieh et al. 2010
B. capnodes YMJ 138 Unknown China NA JX507779 AY951675 Mirabolfathy et al. 2013
B. citriformis YMJ 88113012 Unknown China JX507800 JX507780 AY951677 Hsieh et al. 2010
B. citriformis YMJ 129 Unknown USA JX507801 JX507781 AY951678 Mirabolfathy et al. 2013
B. cylindrispora YMJ 89092701T Bark of Cinnamomum China EF026133 JX507782 AY951679 Hsieh et al. 2010
B. dendrobii MFLUCC 172607T Dendrobium aphyllum China NR172733 NA MK975826 Ma et al. 2020
B. dicranopteridis KUNCC 23-12692T Dendrobium aphyllum China OR001813 OR001855 NA Chen et al. 2024
B. diaoluoshanensis FCATA12071T Dead branch China PX060162 PX092165 PX092167 This study
B. diaoluoshanensis FCATA12659 Dead branch China PX251968 PX092164 PX092166 This study
B. formosana YMJ 89032201T Bark China JX507802 JX507783 AY951680 Hsieh et al. 2010
B. glaucae GMBC0007T Quercus glauca China MT624046 MT622652 MT622654 Li et al. 2021
B. glaucae GMBC0029 Quercus glauca China MT624047 MT622653 MT622655 Li et al. 2021
B. granmo YMJ 135 Bark Austria JX507803 JX507784 AY951681 Hsieh et al. 2010
B. jianfengensis FCATA12140T Dead branch China PX058858 PX092168 PX092170 This study
B. jianfengensis FCATA12339 Dead branch China PX251984 PX092169 PX092171 This study
B. latirima YMJ 89101101 Bark China JX507804 JX507785 NA Hsieh et al. 2010
B. latirima YMJ 90080703 Bark China EF026135 JX507786 AY951683 Ju and Rogers 2001
B. magna MFLU 18-0850T Unknown Thailand MW240616 MW342620 MW775577 Samarakoon et al. 2022
B. mangiferae MFLU 18-0827T Mangifera indica Thailand MN337232 MN366247 MN509782 Samarakoon et al. 2022
B. marginata MFLUCC 12-0740 Unknown France KJ958407 KJ958409 KJ958406 Daranagama et al. 2015
B. marginata CBS 124505 Quercus sp. USA KU684016 KU684310 KU6841249 Chen et al. 2024
B. mediterranea YMJ 147T Unknown France EF026134 NA AY951684 Hsieh et al. 2010
B. mediterranea AZ0703 Unknown USA HM123416 KU684217 KU684122 U’Ren et al. 2016
B. nummularia MUCL 51395T Fagus sylvatica France NR 153649 NA KX271241 Wendt et al. 2018
B. nummularia CBS 969.70 Unknown Unknown MH860015 KU684281 KU684125 Li et al. 2021
B. papillata BCC 36828T Unknown Unknown PQ586363 PQ604605 NA Wongkanoun et al. 2026
B. petrensis LC5698 Unknown China KU746670 NA KU746763 Blumenstein et al. 2021
B. petrensis MFLUCC 14-0151 Dendrobium harveyanum Thailand MK944283 MK975825 MK975827 Ma et al. 2020
B. petrensis HKAS 102388T Unknown Unknown MW240615 MW342619 MW775576 Hsieh et al. 2010
B. philippinensis var. microspora YMJ 89041101 Bark China EF026136 JX507787 AY951685 Hsieh et al. 2010
B. pezizoides ATCC 62606 Unknown USA KY610383 KY624237 KX271242 Brenken and Riebesehl 2025
B. repanda KAS-JKI-GF-2023-009 Unknown Czech Republic PQ227787 NA NA Brenken and Riebesehl 2025
B. rosacearum Bx63 Quercus pubescens Italy KT253501 NA KT253535 Raimondo et al. 2016
B. rosacearum Bx26T Prunus domestica Italy KT253493 NA KT253527 Raimondo et al. 2016
B. sinensis SC3T Pinus thunbergii China OR803755 OR832189 OR832179 Qiao et al. 2024
B. simplicior YMJ 136 Rhamnus cathartica France EF026130 JX507788 AY951686 Hsieh et al. 2010
B. uniapiculata YMJ 90080608 Bark China JX507805 JX507789 AY951687 Hsieh et al. 2010
B. xishuangbannaensis HKAS 122774 Dead twig China PX583847 PX521740 NA Ren et al. 2026
Camillea broomeana GMBC0218 Unknown. China MW854657 MW855488 MW855491 Li et al. 2021
C. obularia ATCC 28093 Unknown USA KY610384 KY624238 KX271243 Wendt et al. 2018
C. tinctor MFLU 18-0786 Unknown Unknown MW240614 MW342618 MW775575 Samarakoon et al. 2022
C. tinctor YMJ 363 Unknown Martinique JX507806 JX507790 JX507795 Hsieh et al. 2010
Cryptostroma corticale CBS 216.52 Acer sp. United Kingdom HG934110 HG934116 HG934102 Li et al. 2021
Cr. corticale CBS 218.52 Acer sp. United Kingdom HG934112 HG934118 HG934104 Li et al. 2021
Graphostroma guizhouensis GMBC0008 Unknown China MW854658 MW855486 MW855489 Li et al. 2021
G. guizhouensis GMBC0219T Unknown China MW854659 MW855487 MW855490 Li et al. 2021
G. platystomum CBS.270.87T Unknown France HG934115 KY624296 HG934108 Li et al. 2021
G. platystomum CPC:37153 Lindera benzoin USA GQ428316 NA NA Li et al. 2021
Hypoxylon rubiginosum YMJ 24 Quercus brantii Iran EF026143 JX507791 AY951751 Hsieh et al. 2010
Jackrogersella cohaerens YMJ 310 Fagus sp. Unknown EF026140 GQ844766 AY951655 Hsieh et al. 2010
Obolarina dryophila 8409 Unknown Unknown GQ428316 NA GQ428321 Li et al. 2021
O. dryophila H76 Quercus sp. France GQ428317 NA GQ428323 Li et al. 2021
O. persica YMJ 1461T Unknown Unknown JX507807 JX507792 JX507796 Li et al. 2021
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Phylogenetic analyses

Phylogenetic analyses for Biscogniauxia and related genera, including Graphostroma, Camillea, Cryptostroma, and Obolarina, were performed with Bayesian inference (BI) and Maximum likelihood (ML) analyses based on the combined ITS-RPB2-TUB2 dataset (Table 1). Jackrogersella cohaerens (Pers.) L. Wendt, Kuhnert & M. Stadler and Hypoxylon rubiginosum (Pers.) Fr. were selected as outgroups (Stadler et al. 2013; Wendt et al. 2018; Li et al. 2021; Crous et al. 2023). The sequences were aligned using the online analysis website MAFFT (MAFFT alignment and NJ/UPGMA phylogeny). The aligned sequences were trimmed using BioEdit software (Hall 1999), and individual gene datasets were concatenated using MEGA 6.0 (Tamura et al. 2011). The maximum likelihood (ML) phylogenetic tree was constructed using the RAxML GUI v1.3.1 software, employing the GTRGAMMA model and the ML+rapid bootstrap procedure with 1,000 replicates to assess the confidence of each branch (Stamatakis et al. 2008). The trimmed sequences were converted to NEXUS format, and manual adjustments were made using ClustalX 1.83 (Thompson et al. 1997). Bayesian inference (BI) analyses were performed using MrBayes v.3.2.6 (Ronquist et al. 2012). The best-fit substitution models for each gene were determined using jModelTest 2 (Darriba et al. 2012). Six Markov chains were run for 3,000,000 generations, with trees sampled every 100 generations. The first 25% of the sampled trees were discarded as burn-in, and the remaining trees were used to calculate posterior probabilities (Rannala and Yang 1996). The phylogenetic tree was visually edited using FigTree v1.4.4 (http://tree.bio.ed.ac.uk/software/figtree/).

Results

Phylogenetic inference

The phylogenetic analysis was based on a total of 148 sequences from three DNA loci (56 ITS, 44 RPB2, and 48 TUB2) derived from 61 strains representing 43 species, including two newly described species, 31 known species of Biscogniauxia, 3 species of Camillea, one species of Cryptostroma, 2 species of Graphostroma, 2 species of Obolarina, and the outgroups Jackrogersella cohaerens (YMJ 310) and Hypoxylon rubiginosum (YMJ 24).

The dataset contained a total of 3,233 aligned characters, of which 1,683 were constant characters, 1,326 were phylogenetically informative characters, and 224 were variable characters that lacked parsimony information. Phylogenetic trees generated from ML and BI analyses of the combined dataset of ITS-RPB2-TUB2 were highly similar in topology. Only the ML tree is represented in Fig. 1, with ML bootstrap values ≥ 80% and Bayesian posterior probabilities ≥0.90 labelled along the branches. The phylogram based on the combined ITS-RPB2-TUB2 sequences (Fig. 1) showed that our two species were assigned to the genus Biscogniauxia, in which B. jianfengensis grouped with B. philippinensis var. microspora Y.M. Ju & J.D. Rogers (YMJ 89041101) with high support value (BS = 100, BPP = 1), and B. diaoluoshanensis resolved as a basal lineage of the other Biscogniauxia taxa included in this study. Based on phylogenetic and morphological evidence (described below), we propose two new species, namely B. diaoluoshanensis sp. nov. and B. jianfengensis sp. nov., to accommodate our specimens.

Figure 1.

Figure 1.

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Phylogram generated from Maximum Likelihood (RAxML) analyses based on ITS-RPB2-TUB2 matrix. Branches are labelled with maximum likelihood bootstrap ≥80% and Bayesian posterior probabilities ≥0.90 at the nodes. New species in this study are indicated in bold. Holotype specimens are labelled with T.

Taxonomy

Biscogniauxia diaoluoshanensis

Hai X. Ma, A.H. Zhu, S. Liu & S.Y. Ma sp. nov.

0C65A224-C1F5-5013-8AAC-835D471D14A1

864450

Fig. 2

Figure 2.

Figure 2.

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Biscogniauxia diaoluoshanensis. (FCATAS 12071, Holotype). a, b. Stromata on the surface of the branch; c. Surface of stromata; d. Vertical section through stroma showing ostioles and perithecia; e, f. Asci in water; g. Apical ring of the asci; h. Ascal ring bluing in Melzer’s reagent; i. Ascospore with germ slit; j. Ascospore in water; k. Ascospore in KOH. Scale bars: 0.5 mm (c, d); 25 μm (e, f); 10 μm (g–k).

Holotype.

China • Hainan Province, Lingshui Li Autonomous County, Hainan Tropical Rainforest National Park, Diaoluo Mountain Branch, Fengguoshan Waterfall, saprobic on a dead branch of an unidentified plant, alt. 530 m, 27 May 2024, Hai X. Ma, DLS058 (FCATAS 12071, Holotype). GenBank accession numbers: ITS (PX060162); TUB2 (PX092167); RPB2 (PX092165).

Etymology.

Named after Diaoluo Mountain Branch of Hainan Tropical Rainforest National Park, the type locality in Hainan Province, China.

Description.

Sexual morph: Stromata applanate, orbicular to irregularly lobed, 4–12 × 1.5–5 cm, with slightly sloping margins, somewhat confluent; mature surface black, shiny, with conspicuous depressions, tissue beneath perithecia 0.03–0.04 mm thick, carbonaceous, underlying bark tissue black. Perithecia ovoid to pyriform, frequently laterally and basally flattened, 340–430 × 70–115 µm (x̄ = 385 × 92 µm, n = 20), opening centrally or laterally through individual ostioles. Ostioles papillate, apically obtuse or truncate, black, conspicuously shiny, at times punctate with a low rim. Asci cylindrical with 8 ascospores, 89–95 µm total length, short-stipitate to subsessile, the spore-bearing parts 76–80 × 6–8 µm (x̄ = 78 × 6 µm, n = 60/2), the stipes 12–14 µm; apical apparatus 2.6–5.7 × 5.3–5.7 µm (x̄ = 1.7 × 5.1 µm, n = 60/2), short-cylindrical, slightly trapezoid, bluing in Melzer’s reagent (J+). Paraphyses sparse, hyphal, unbranched, thin-walled, septate, 4–5 µm wide, tapering above asci. Ascospores (10.1–) 10.4–12.7 (–13.1) × (4.8–) 5.1–6.1(–6.3) µm (x̄ = 11.6 × 5.5 µm, n = 60), unicellular, light brown to brown, ellipsoid almost equilateral, one end slightly broader and rounded, the other slightly narrower and obliquely truncate, smooth-walled, with a conspicuous straight germ slit almost spore-length or slightly less than spore-length. Asexual morph: Undetermined.

Additional specimens examined.

China • Hainan Province, Lingshui Li Autonomous County, Hainan Tropical Rainforest National Park, Diaoluo Mountain Branch, Fengguoshan Waterfall, saprobic on a dead branch of an unidentified plant, alt. 527 m, 27 May 2024, Hai X. Ma, DLS061 (FCATAS 12659, Paratype).

Remarks.

Biscogniauxia diaoluoshanensis is characterized by slightly concave stromata with a blackish-brown surface and shiny black papillate ostioles, and nearly equilateral brown ascospores with a conspicuous straight, almost spore-length germ slit. Biscogniauxia granmoi Lar. N. Vassiljeva resembles the new species in its stromatal morphology and ascospore shape but possesses larger perithecia (0.6–0.8 mm high × 0.3–0.5 mm broad) and larger asci (119–142 µm total length), and occurs on trunks and branches of Prunus (Vasilyeva 1998; Læssøe et al. 1999). Another similar species, B. nummularia (Bull.: Fr.) Kuntze, resembles the new species in stromatal appearance, but can be distinguished by its larger obovoid perithecia (0.3–0.5 × 0.5–0.7 mm), longer asci (95–120 µm total length), and wider ascospores (7.5–8.5 µm) (Ju et al. 1998; Ju and Rogers 2001). In addition, it occurs on wood of Fagus (Ju et al. 1998; Zíbarová and Kout 2017). Phylogenetically, the new species falls within Biscogniauxia clade 4 yet exhibits no obvious affinity to other taxa in this clade. Clade 4 comprises the type species B. nummularia, ten additional Biscogniauxia species, and our new collection. Biscogniauxia nummularia (MUCL 51395, Holotype) differs from B. diaoluoshanensis by 168 base pairs in TUB2 alignment, representing 11.52% of the total aligned TUB2 region. Strain CBS 969.70 of B. nummularia shows 95-base-pair divergence in RPB2 and 164-base-pair divergence in TUB2 compared with the new species, corresponding to 9.27% and 11.28% of aligned nucleotide positions of RPB2 and TUB2 regions, respectively. Biscogniauxia capnodes (YMJ 138) differs from B. diaoluoshanensis by 83 base pairs in RPB2, accounting for 7.57% of the aligned RPB2 region. For other congeneric species, sequence similarity to the new species is below 91% for both RPB2 and TUB2. All previously known species display ITS sequence similarity below 86% relative to this novel taxon.

Biscogniauxia jianfengensis

Hai X. Ma, A.H. Zhu, S.Y. Ma & S. Liu sp. nov.

6C64DE9F-A69F-5118-BE2B-B7C1AD4E7340

864451

Fig. 3

Figure 3.

Figure 3.

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Biscogniauxia jianfengensis (FCATAS 12140, Holotype). a, b. Stromata on the surface of the branch; c. Surface of stromata; d. Vertical section through stroma showing ostioles and perithecia; e, f. Asci in water; g. Ascal ring bluing in Melzer’s reagent; h. Apical ring of the asci in water; i, j. Ascospore in water. Scale bars: 1 mm (b); 0.5 mm (c, d); 50 μm (e–g); 10 μm (h–j).

Holotype.

China • Hainan Province, Ledong Li Autonomous County, Hainan Tropical Rainforest National Park, Jianfengling Branch, Yulin Valley, alt. 650 m, saprobic on a dead branch of an unidentified plant, 29 May 2024, Hai X. Ma, JFL0041 (FCATAS 12140, Holotype). GenBank accession numbers: ITS (PX058858); TUB2 (PX092170); RPB2 (PX092168).

Etymology.

Named after Jianfengling Branch of Hainan Tropical Rainforest National Park, the type locality in Hainan Province, China.

Description.

Sexual morph: Stromata applanate to slightly pulvinate, orbicular to irregularly ellipsoid-lobed, 2–6 × 1–3 cm, coalescent into widely effused compound stromata, with slightly sloping margins lacking remnants of the outer dehiscing layer; mature stromatal surface dull black, matte, even, and carbonaceous immediately beneath the surface and between perithecia; tissue beneath perithecia 0.05–0.07 mm thick, carbonaceous, underlying bark tissue black. Perithecia tubular to short-tubular, 450–570 × 210–480 µm (x̄ = 506 × 289 µm, n = 20), opening centrally or laterally through individual ostioles. Ostiole umbilicate or punctate with a slightly to conspicuously raised rim, often surrounded by a darker halo, evenly distributed or in small clusters. Asci cylindrical with 8 ascospores, 125–186 µm total length, short-stipitate to subsessile, the spore-bearing parts 121–148 × 9–11 µm (x̄ = 136 × 10 µm, n = 60/2), the stipes 10–25 µm; apical apparatus 2.6–5.7 × 6.6–9.2 µm (x̄ = 4.1 × 7.6 µm, n = 60/2), short-cylindrical, apically flared, bluing in Melzer’s reagent (J+). Paraphyses copious, hyphal, unbranched, thin-walled, septate, 4–6 µm wide, tapering above asci. Ascospores (13.4–) 14.8–18.8 (–20.8) × (8.3–) 8.5–10.6 (–11.4) μm (x̄ = 17.1 × 9.6 μm, n = 50), unicellular, dark brown to blackish brown, ellipsoid-inequilateral with broadly rounded upper end, bearing an opaque, thick-walled broadly conical, obtusely-ended cellular appendage, 1.6–2.8 µm high × 2.3–3.8 µm broad (x̄ = 2.2 × 3.2 µm, n = 30), with an inconspicuous, unilateral, straight spore-length or slightly less germ slit on the most convex side. Asexual morph: Undetermined.

Additional specimens examined.

China • Hainan Province, Ledong Li Autonomous County, Hainan Tropical Rainforest National Park, Jianfengling Branch, alt. 660 m, saprobic on a dead branch of an unidentified plant, 29 May 2024, Hai X. Ma, (FCATAS 12339, Paratype). GenBank accession numbers: ITS (PX251984); TUB2 (PX092171); RPB2 (PX092169).

Remarks.

Phylogenetically, Biscogniauxia jianfengensis forms a sister lineage to the tropical variety B. philippinensis var. microspore, with this clade itself sister to B. latirima Y.M. Ju & J.D. Rogers. Biscogniauxia philippinensis var. microspore resembles the new species in stromatal morphology and ascospore shape but differs in having a larger perithecia (1.4–2.4 × 0.2–0.3 mm vs. 450–570 × 210–480 µm) and longer asci with the spore-bearing parts (asci 190–220 µm total length vs. 125–186 µm total length) (Ju et al. 1998). In contrast, B. latirima shares applanate stromata and similar perithecia shape but diverges in smaller apical apparatus (4–5 × 5–6 µm vs. 2.6–5.7 × 6.6–9.2 µm), and larger ascospores (18–21.5 × 10–12 µm vs. 14.8–18.8 × 8.5–10.6 μm) lacking a visible cellular appendage (Ju and Rogers 2001). Morphologically, the new species B. jianfengensis is similar to the tropical taxon B. philippinensis in stromatal morphology, but the latter has larger ascospores (22–27 × 11.5–13 µm) with a straight, faint spore-length germ slit (Ju et al. 1998). Unfortunately, no molecular sequences are available for B. philippinensis, so nucleotide difference comparison could not be implemented.

Discussion

In the present study, two new species of Biscogniauxia from Hainan Tropical Rainforest National Park in China are described based on morphological characteristics and phylogenetic analyses. Phylogenetic analyses on the species of Biscogniauxia presented confirmed that the genus Biscogniauxia is polyphyletic, as in previous studies (Ma et al. 2020; Li et al. 2021; Chen et al. 2024; Liu et al. 2025; Liu et al. 2026). The species analyzed appeared mainly distributed in six separate clades (except B. sinensis, B. atropunctata (Schwein.) Pouzar, B. uniapiculata, and B. bartholomaei (Peck) Lar.N. Vassiljeva) in our study. Biscogniauxia diaoluoshanensis and B. jianfengensis were clearly separated from other Biscogniauxia species and from each other in Biscogniauxia clade 4 and clade 2 respectively. Biscogniauxia nummularia, the type species of the genus, was included in clade 4. The phylogenetic tree indicates that the classification status of the genus Biscogniauxia remains ambiguous. In our phylogenetic analyses, Camillea species nested within the Biscogniauxia clade, which is consistent with recent findings reported by Wongkanoun et al. (2026). Accordingly, further studies on the diversity, phylogeny, and evolution of Biscogniauxia require expanded sampling and molecular data from broader geographic regions.

A number of macrofungi belonging to Basidiomycota and Ascomycota have been extensively studied in recent years, and numerous new taxa have been described from Hainan Province (Ma et al. 2022b; Pan et al. 2022; Deng et al. 2023, 2024; Huang et al. 2023; Gao et al. 2024; Zhu et al. 2024b; Yu et al. 2025; Zhang et al. 2025; Li et al. 2025). Members of Graphostromataceae are widely distributed across tropical regions worldwide; nevertheless, records from tropical Asia remain relatively sparse (Samarakoon et al. 2022; Liu et al. 2025; Ren et al. 2026; Wongkanoun et al. 2026). To date, only three species of Graphostromataceae have been documented from Hainan Province (Ju et al. 1998; Ariyawansa et al. 2015). Therefore, the species diversity of Graphostromataceae is still largely underexplored on this tropical island, and further surveys across diverse forest types in Hainan are likely to yield additional new species.

Dichotomous key to Biscogniauxia taxa known from China

Note that taxa with only anamorph reported are not included in this key.

1 Mature ascospores bearing a cellular appendage 2
– Mature ascospores lacking a cellular appendage 5
2 Ascospores with a conspicuous straight germ slit, (9–)10–14 × 5–7 µm B. uniapiculata
– Ascospores with an unconspicuous germ slit 3
3 Ascospores 22–27 × 11.5–13 µm B. philippinensis
– Ascospores smaller than 21 µm 4
4 Perithecia 0.45–0.57 mm high × 0.21–0.48 mm broad; ascospores (13.4–) 14.8–18.8 (–20.8) × (8.3–) 8.5–10.6 (–11.4) μm B. jianfengensis sp. nov.
– Perithecia 1.4–2.4 mm high × 0.2–0.3 mm broad; ascospores 13.5–19 × 8–10 µm B. philippinensis var. microspora
5 Ascospores inequilateral to strongly inequilateral 6
– Ascospores more or less equilateral 9
6 Ascospores 8.5–15 × 5–7.5 μm B. citriformis
– Ascospores greater than 15 μm long 7
7 Ascospores (16–)18–21.5 × 10–12 μm, with a broad germ-slit on more convex side B. latirima
– Ascospores with a narrow germ-slit on less convex side 8
8 Ascospores dark brown, (18–)19–24(–25) × 8.5–10 μm B. formosana
– Ascospores pale brown, narrow, 17–23.5 × 10–12 μm B. formosana var. kentingensis
9 Ascospores light brown to brown 10
– Ascospores dark brown to blackish brown 11
10 Ascospores (10.1–) 10.4–12.7 (–13.1) × (4.8–) 5.1–6.1(–6.3) µm B. diaoluoshanensis sp. nov.
– Ascospores 25–31 × 8–10 µm B. effusa
11 Stromata widely effused, with raised margins 12
– Stromata applanate 13
12 Ascospores 13.5–16 × 7.5–9.5 μm B. glaucae
– Ascospores 10–12.5 × 6.5–10 μm B. betulae
13 Mature stromatal surface dark brown with olivaceous tinge; ascospores 10–13 × 4–5 (–6) μm B. ambiens
– Mature stromatal surface blackish 14
14 Ascospores mostly greater than 16 μm long 15
– Ascospores mostly less than 16 μm long 16
15 Perithecia narrow, 0.05–0.1 mm diam; ascospores 20–26 × 10–14(–16) µm B. cylindrispora
– Perithecia broad, 0.2–0.4 mm diam; ascospores (14– )15–22. 5(–24) × 8–11(–12) µm B. capnodes var. rumpens
16 Perithecia less than 0.15 mm diam 17
– Perithecia more than 0.2 mm diam 18
17 Ascospores 11–14 × 5.5–7.5 µm, with narrowly or occasionally broadly rounded ends B. plana
– Ascospores 11–15 × 5.7–7 µm, with broadly rounded ends B. petrensis
18 Germ-slit on both sides of the ascospore, ascospores 13–16 × 5–7 µm B. albosticta var. orientalis
– Germ-slit on one side of the ascospore 19
19 Ascospores 8.5–15 × 5–7.5 µm B. capnodes
– Ascospores 8–11 × 4.4–6 µm B. xishuangbannaensis
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Supplementary Material

XML Treatment for Biscogniauxia diaoluoshanensis
XML Treatment for Biscogniauxia jianfengensis

Acknowledgements

We thank the Public Technology Research and Sharing Center of the Institute of Tropical Bioscience and the Biotechnology Chinese Academy of Tropical Agricultural Sciences for equipment sharing and technical support.

Citation

Ma S-Y, Liu S, Qu Z, Jiang H, Zhu A-H, Ma H-X (2026) Phylogenetic analyses and morphological studies reveal two new species of Biscogniauxia (Xylariales, Ascomycota) from Hainan Tropical Rainforest National Park, China. MycoKeys 140: 241–259. https://doi.org/10.3897/mycokeys.140.206566

Footnotes

Si-Yu Ma and Shuang Liu contributed equally to this work.

Contributor Information

An-Hong Zhu, Email: 18289679317@163.com.

Hai-Xia Ma, Email: mahaixia@itbb.org.cn.

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

The research was supported by National Natural Science Foundation of China (32470016), Hainan Province Science and technology Special Fund (ZDYF2023RDYL01, ZDYF2026SXLH033), Hainan Institute of National Park, HINP, KY-24ZK02.

Author contributions

Data curation: SYM, SL, ZQ, HJ. Investigation: SYM, SL, AHZ, HXM. Methogology: SYM, SL. Resources: HXM. Validation: SYM, SL, ZQ. Writing—original draft: SYM, SL. Writing—review and editing: AHZ, HXM.

Author ORCIDs

S.-Y. Ma https://orcid.org/0009-0000-2974-6277

S. Liu https://orcid.org/0009-0008-8121-6394

Z. Qu https://orcid.org/0000-0002-0909-0853

H. Jiang https://orcid.org/0009-0006-6430-5788

A.-H. Zhu https://orcid.org/0000-0002-2812-8108

H.-X. Ma https://orcid.org/0000-0001-6699-7454

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 Biscogniauxia diaoluoshanensis
mycokeys.140.206566-treatment1.xml (17.1KB, xml)
XML Treatment for Biscogniauxia jianfengensis
mycokeys.140.206566-treatment2.xml (13.5KB, 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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