Three New Species and Three New Records of Arthoniaceae (Ascomycota, Arthoniales) from China
1
College of Agriculture and Biology, Liaocheng University, Liaocheng 252059, China
2
State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China
*
Authors to whom correspondence should be addressed.
J. Fungi 2026, 12(1), 42; https://doi.org/10.3390/jof12010042
Submission received: 21 November 2025
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Revised: 26 December 2025
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Accepted: 4 January 2026
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Published: 6 January 2026
(This article belongs to the Special Issue Lichenized Fungi: A Themed Issue Dedicated to Academician Jiang-Chun Wei)
Abstract
Based on morphological and phylogenetic analyses, during a study on the biodiversity of the lichenized fungi family Arthoniaceae in Yunnan, China, three new species of Arthonia, Eremothecella, and Synarthonia (Arthonia yunnanensis sp. nov., A. pararubella sp. nov., Eremothecella pruinocarpa sp. nov.) and three new Chinese records (A. rubella, E. calamicola, Synarthonia inconspicua) were reported in this present study. A phylogenetic analysis using maximum likelihood and Bayesian inference based on a two-gene dataset (mtSSU and RPB2) revealed that all these species form distinct independent clades. In addition, this study represents the first report of molecular sequences for the genus Eremothecella. Detailed descriptions, ecological and chemical characteristics, and illustrations for each species are provided.
1. Introduction
The family Arthoniaceae Rchb. (belonging to Ascomycota, Arthoniales) is one of the largest families of lichenized fungi, and was formally delineated by Reichenbach [1]. It exhibits two important morphological features: a reduction of the ascomatal borders and globose to clavate asci with a strongly thickened apical wall, and it is not monophyletic [2,3,4,5]. This family is mainly prominent in tropical regions with abundant corticolous and foliicolous species, and it constitutes a major component of the lichen flora across diverse forest types [6]. Including genera such as Arthonia Ach., Eremothecella Syd., and Synarthonia Müll. Arg., the family currently comprises 20 genera and approximately 750 species [7,8].
Arthonia (established by Acharius in 1806) remains one of the most understudied taxa even though it encompasses approximately 500 species in the world [9]. This genus is considered to be a heterogeneous group, according to Grube et al. [10]. The lichenized representatives of Arthonia are always crustose with irregular or stellate (rarely rounded) ascocarps and have hyaline to brownish, muriform, or transversely septate (often unequally) ascospores [11]. The genus (at least in its traditional sense) comprises all different life forms: lichens, non-lichenized fungi, lichen parasites, and parasymbionts [5].
The genus Eremothecella, established by Sydow [12], shows distinctive morphological characteristics, including dark brown to black ascomata with loose paraphysoids, globose asci frequently emerging through the ascomata surface as small warts, and applanate pycnidia producing elongated filiform conidia with multiple septa, which provide taxonomic separation from Arthonia [13,14]. The genus was merged into Arthonia by Santesson [15], but subsequent revisions by Sérusiaux [13] recognized the generic distinctness of Eremothecella and formally accepted four species within this classification: viz. E. calamicola Syd., E. macrosperma (Zahlbr). Sérus., E. palmulacea (Müll. Arg.) Sérus., and E. variratae (Aptroot & Sipman) Sérus. Lücking (2008) further affirmed its taxonomic autonomy based primarily on the presence of Eremothecella-type pycnidia [14]. Currently, ten species are recognized: eight foliicolous and two corticolous in this genus [16].
Synarthonia was established in 1891. This small genus was the subject of a detailed discussion and report by Van Den Broeck et al. [17]. It is characterized by solitary ascomata becoming mono- to pluri-carpocentral synascomata embedded in a slightly elevated to immersed pseudostroma, with a thin white thalline margin, and Arthonia-type asci producing transversely septate ascospores, with enlarged apical cell or muriform ascospores [18]. However, the phylogeny of Synarthonia was unclear, and the synonymy of Reichlingia Diederich & Scheid had been suggested, but Ertz et al. (2020) pointed out that Synarthonia can be placed in a well-supported monophyletic lineage [18,19].
During our ongoing studies on Arthoniaceae in China, specimens belonging to three genera—Arthonia, Eremothecella, and Synarthonia—were collected. Integrative taxonomic analyses based on combined phenotypic and molecular datasets were conducted [20]. For molecular phylogenetic analyses, the mitochondrial small subunit (mtSSU) and the RNA polymerase subunit II (RPB2) were used, following the methods described by Frisch et al. (2014) and Thiyagaraja et al. (2020) for Arthoniaceae [5,6]. Three new species and three new records of Arthoniaceae from China are reported here.
2. Materials and Methods
2.1. Morphological and Chemical Analyses
Specimens were collected from Yunnan Province, China, and were deposited in the Fungarium of the College of Life Sciences, Liaocheng University (LCUF). Freshly collected specimens were first subjected to DNA extraction or preserved at −20 °C. The extracted DNA was typically stored at −80 °C, and all specimens were ultimately deposited in the herbarium. An Olympus dissecting microscope (Olympus SZX16, Olympus Corporation, Tokyo, Japan) and an Olympus light microscope (Olympus BX53, Olympus Corporation, Tokyo, Japan) were used for morphological and anatomical studies. Measurements were taken from mature vertical sections of fruit bodies mounted in water to observe whether they contained crystals: the shape and size of the photobiont, the width of the hyphae, and the color, shape, and size of the asci, ascospores, pycnidia, and conidia. The amyloidity of ascospores was tested with Lugol’s solution. Spot tests with K (10% aqueous solution of potassium hydroxide), C (saturated solution of aqueous sodium hypochlorite), P (saturated solution of p-phenylenediamine in 95% ethylalcohol), and UV (fluorescence at 365 nm) were performed on the surface of the thallus [21]. The lichen substances were detected and identified using a UV light source and thin-layer chromatography (TLC) in solvent C [22].
2.2. DNA Extraction and PCR Sequencing
Genomic DNA was extracted from the ascomata and thallus of the specimens using a DNA secure Plant Kit (Tsingke, Beijing, China) according to the manufacturer’s instructions. The mtSSU and RPB2 regions were amplified using the primer pair mtSSU1/3R [23] and RPB2-7cF/11aR [24]. Polymerase Chain Reaction (PCR) cycling conditions were set to an initial denaturation at 95 °C for 5 min, followed by 35 cycles of denaturation at 95 °C for 45 s, annealing at 50 °C for 1 min, extension at 72 °C for 1.5 min, and a final extension at 72 °C for 10 min [25]. The PCR products were checked on 1% electrophoresis gels, stained with ethidium bromide, and sequenced by Tsingke Biotech Co., Ltd. (Beijing, China).
2.3. Phylogenetic Analyses
The new sequences were assembled via Geneious v.9.0.2 (Biomatters Ltd., Auckland, New Zealand) and potential sample contamination were confirmed by BLASTn searches on the NCBI website (http://www.ncbi.nlm.nih.gov/BLAST/, accessed on 17 April 2025). Then, high-similarity sequences of Arthoniaceae were selected for further analysis (Table 1), with Melarthonis piceae Frisch & G. Thor (belonging to Chrysotrichaceae, Arthoniales) as the outgroup [5]. The assembled sequences were aligned with the online version of MAFFT v.7 (https://mafft.cbrc.jp/alignment/server/, accessed on 17 April 2025). Ambiguously aligned regions were delimited using Gblocks v.0.91b with the least stringent selection [26]. Geneious v.9.0.2 was used to concatenate the mtSSU and RPB2 genes and produce a two-locus dataset. Maximum likelihood (ML) analysis and Bayesian inference (BI) were used to infer phylogenetic trees based on the concatenated mtSSU and RPB2 datasets. The maximum likelihood analysis was conducted with RAxML-HPC 2 on ACCESS v.8.2.12 employing a GTRGAMMA approximation with a rapid bootstrap analysis of 1000 replicates on the CIPRES Scientific gateway portal (http://www.phylo.org/portal2/, accessed on 17 April 2025) for verification [27,28]. For BI analysis, PartitionFinder 2 [29] was used to determine the best-fit model for each partition. Based on the results, we used the GTR+I+G model for mtSSU and the HKY+I+G model for RPB2. Bayesian Inference phylogenies were inferred using MrBayes v.3.2.7a under a partition model [30]. We ran 2 million generations, in which the initial 25% of sampled data was discarded as burn-in. Bayesian inference posterior probability (BIPP) equal to and above 95% was considered significant and shown at the nodes. Generated phylogenetic trees were visualized under Figtree v.1.4.4 and edited in Adobe Illustrator CC2019 software.
3. Results
3.1. Phylogenetic Results
A total of 150 specimens comprising 227 DNA sequences (139 mtSSU sequences and 88 RPB2 sequences), including fifteen mtSSU sequences and eight RPB2 sequences that were newly generated were used in this study (Table 1). For the two-locus (mtSSU and RPB2) dataset taxon set, the concatenated alignment contained 1544 characters (787 for mtSSU and 757 for RPB2). The RAxML tree was constructed with a final ML optimization likelihood value of −27,812.975243. The parameters for the GTR+I+G model of combined mtSSU and RPB2 were as follows: estimated base frequencies: A = 0.290705, C = 0.231758, G = 0.228597, T = 0.248940; substitution rate AC = 1.017805, AG = 5.448594, AT = 1.720312, CG = 1.388442, CT = 5.952238, and GT = 1.000000. The resulting topology was similar for maximum likelihood (ML) and Bayesian analysis. Therefore, only the Bayesian tree is presented, with bootstrap support ≥70% for ML analysis and posterior probabilities ≥0.95 for Bayesian analysis (Figure 1).
In the phylogenetic tree (Figure 1), the main well-supported lineages of Arthoniaceae follow the results obtained by Frisch et al. (2014) and agree well with previous hypotheses on the polyphyletic origins of parasitic Arthonia [6]. Our phylogenetic results also support that Eremothecella forms an independent clade within the family Arthoniaceae based on the sequence of Eremothecella pruinocarpa (YN242390-1, YN242386-2, YN242384, YN242387-2) and E. calamicola (YN242388, YN242387-1, YN242386-1, YN242390-2), which were newly generated in this study. These are also the first reported molecular sequences of species in the genus Eremothecella.
3.2. Taxonomy
Arthonia pararubella C.Y. Hao, S.H. Jiang, and Z.F. Jia, sp. nov. (Figure 2).
Fungal Name: FN 572511.
Diagnosis. The new species externally resembles Arthonia rubella (Fée) Nyl. but differs in the shorter ascospores and fewer septa.
Type. China. Yunnan Province: Mengla County, Xishuangbanna Tropic Botanical Garden, Chinese Academy of Sciences, Baihuayuan, 21°55′26″ N, 101°15′02″ E, alt. 560 m, on bark, 18 December 2024, C.Y. Hao (LCUF YN242036—holotype).
Description. Thallus greenish gray to whitish gray. Prothallus brownish-black line. Ascomata reddish-brown to blackening, innate, maculate to lirellate, sometimes branched or stellate, thinner at the branched ending, 0.2–1.65 × 0.05–0.2 mm. Excipulum carbonized, 21.5–30 μm thickness. Hymenium 67–77 µm tall, pale brown to brown, K/I+ blue. Asci clavate to subglobose, 47.5–66.5 × 28–44 µm ( = 55 × 35 μm), Arthonia-type, eight-spored, I+ orange-red, apical part thickened, usually with a distinct ocular chamber, K/I−. Ascospores oblong–ovoid, (2–)3-septate, upper and lower cells enlarged, ellipsoidal locules in immaturity, colorless, 25.5–29.5 × 10–13.5 μm ( = 28 × 12.5 μm), I+ orange-red.
Chemistry. Thallus K-, C-, KC-, UV-. No substances were detected by TLC.
Ecology and distribution. The new species is currently only known in China on the bark of trees in a tropical forest in Yunnan Province.
Etymology. The specific epithet pararubella refers to the similarity to Arthonia rubella.
Additional specimens examined. China. Yunnan Province: Jinghong City, Mengyang Town, Wild Elephant Valley, 22°09′58″ N, 100°51′49″ E, alt. 800 m, on bark, 16 December 2024, C.Y. Hao (LCUF YN241783).
Notes. Arthonia pararubella can be distinguished from similar species by its hyaline ascospores (25.5–29.5 × 10–13.5 μm) with two to three septa. Arthonia rubella is morphologically similar to this new species, but the former can be distinguished by longer ascospores with more septa (3–5-septate vs. (2–)3-septate; 25–43 × 10–15 μm vs. 25.5–29.5 × 10–13.5 μm) [31,32,33]. Further, both are far apart in the phylogenetic tree (Figure 1). Another similar species, A. compensatula Nyl. differs from this new species by its ascospores with more septa (4–5-septate vs. (2–)3-septate) and without amphicephalic [33,34].
Arthonia yunnanensis C.Y. Hao, S.H. Jiang, and Z.F. Jia, sp. nov. (Figure 3).
Fungal Name: FN 572510.
Diagnosis. The new species externally resembles Arthonia incarnata Kullh. ex Almq, but differs in the subglobose asci, color of the thallus, and that it lives on leaves.
Type. China. Yunnan Province: Mengla County, Xishuangbanna Tropic Botanical Garden, Chinese Academy of Sciences, lvshilin, 21°54′17″ N, 101°16′28″ E, alt. 625 m, on leaves, 18 December 2024, C.Y. Hao (LCUF YN242413—holotype).
Description. Thallus uncontinuous, yellowish green to green, thin, matte to weakly glossy. Prothallus absent. Photobiont trentepohlioid, cells globose to long elliptical. Ascomata dispersed, maculate, usually slightly irregularly roundish to short elliptical, adnate, flat to weakly convex, pale and often brownish to dark brown, 0.25–0.4 mm in diameter. Hymenium near hyaline to pale orange-brown, K/I+ blue. Asci subglobose, 27–32 × 18–23 μm ( = 29 × 20.5 μm), Arthonia-type, thin lateral walls, stipitate, eight-spored, usually with a distinct ocular chamber, K/I−. Ascospores clavate, 2(–3)-septate, usually with an enlarged distal cell, not curved, with slight constrictions at septa, colorless, 12.5–15.5 × 4.5–6.5 μm ( = 14 × 5.5 μm).
Chemistry. Thallus K-, C-, KC-, UV-. No substances were detected by TLC.
Ecology and distribution. The new species is currently only known in China on the leaves of trees in a tropical rainforest in Yunnan Province.
Etymology. The specific epithet yunnanensis refers to the type locality, Yunnan Province, Mengla County.
Additional specimens examined. China. Yunnan Province: Mengla County, Xishuangbanna Tropic Botanical Garden, Chinese Academy of Sciences, Greenstone Forest, 21°54′17″ N, 101°16′28″ E, alt. 625 m, on leaves, 18 December 2024, C.Y. Hao (LCUF YN242411).
Notes. Arthonia yunnanensis, the only species from southern China’s tropical region inhabiting leaves, is characterized by subglobose asci. It resembles Arthonia incarnata, A. accolens Stirt., and A. lividula Vain. in having 2-septate ascospores [4,15,35,36]. However, A. incarnata resembles the new species, having the same size ascospores (11.7–15.3 × 4.2–5.4 μm vs. 12.5–15.5 × 4.5–6.5 μm), but its thallus is often inapparent or visible as a whitish to pale olive-gray discoloration of bark; further, its asci are longer and narrower (40–50 × 14–20 μm vs. 27–32 × 18–23 μm) [4]. In the phylogenetic tree (Figure 1), although A. yunnanensis forms a sister clade to A. incarnata, the two clades are distinctly separated by a long genetic distance and exhibit substantial divergence. Therefore, A. yunnanensis as an independent lineage also supports its status as a species distinct from A. incarnata. Arthonia accolens can be distinguished by larger ascomata (often >0.5 mm diam.) [15,36]. Arthonia lividula differs from this new species in having shorter asci and smaller ascospores without constrictions at the septa (9–12 × 3–5 μm vs. 12.5–15.5 × 4.5–6.5 μm) [15,35]. Morphologically, A. yunnanensis was revealed as significantly different from all known species of Arthonia (see notes below), and therefore we describe this species here as a new species.
Eremothecella pruinocarpa C.Y. Hao, S.H. Jiang, and Z.F. Jia, sp. nov. (Figure 4).
Diagnosis. The new species externally resembles Eremothecella ajaysinghii Jagad. Ram & G.P. Sinha but differs in its fewer septa of ascospores.
Type. China. Yunnan Province: Jinghong City, tropical rainforests, 21°54′31″ N, 101°10′58″ E, alt. 635 m, on leaves, 17 December 2024, C.Y. Hao (LCUF YN242390-1—holotype).
Description. Thallus crustose, greenish gray, dispersed or continuous, matte. Prothallus absent. Photobiont trentepohlioid, cells in radiate plates, rectangular, ca. 11 × 4 μm. Ascomata rounded to irregular, dark brown to black, ± moderately to densely grayish pruinose, pruina sometimes increasingly dense along the margin, 0.2–1.15 mm diam. Asci obovate to globose, 45–60 × 36.5–45.5 μm ( = 54.5 × 41 μm), stipitate, eight-spored, K/I+ red, usually with a distinct ocular chamber, K/I+ blue. Ascospores colorless, clavate, curved, (6–)7(–9)-septate, enlarged distal cell, very slight constrictions at septa, 30–49 × 8–11.5 μm ( = 38 × 9.5 μm), I+ orange-red. Pycnidia numerous, ± rounded, irregular, black, 0.2–0.45 mm diam. Conidia colorless, filiform, multiseptate, 80–130 × 1.5–2 µm ( = 100 × 2 μm).
Chemistry. Thallus K-, C-, KC-, UV-. No substances were detected by TLC.
Ecology and distribution. The new species is currently only known in China on the leaves of trees in a tropical rainforest in Yunnan Province.
Etymology. The specific epithet pruinocarpa refers to the densely whitish gray ascomata.
Additional specimens examined. China. Yunnan Province: Jinghong City, tropical rainforests, 21°54′31″ N, 101°10′58″ E, alt. 635 m, on leaves, 17 December 2024 X. Li (LCUF YN242384; LCUF YN242386-2; LCUF YN242387-2).
Notes. Eremothecella pruinocarpa differs from similar species in its grayish pruinose ascomata and (6–)7(–9)-septate ascospores. Eremothecella ajaysinghii resembles the new species E. pruinocarpa in having grayish pruinose ascomata but differs in having ascospores with more septa [8–10(–11)-septate vs. (6–)7(–9)-septate] [16]. Eremothecella macrosperma is another similar species, particularly in ascospore size, but its ascomata are non-pruinose [15]. Eremothecella cyaneoides Lücking differs from this new species in having ascospores with fewer septa (3–5-septate vs. (6–)7(–9)-septate) [37]. Phylogenetically, four specimens of Eremothecella pruinocarpa form a strongly supported monophyletic clade, which also supports that it is a new species (Figure 1).
Arthonia rubella (Fée) Nyl. (Figure 5).
Fungal Name: FN 119459.
Basionym. Graphis rubella Fée, Essai Crypt. Exot. (Paris): 43 (1825) [1824].
Description. Thallus greenish gray to green, or whitish gray. Prothallus line brownish black. Photobiont trentepohlioid, cells globose. Ascomata reddish-brown and black, innate, maculate to lirellate, flexuous to irregularly branched, 0.25–1.2 mm long, disk occasionally opening. Asci clavate to subglobose, Arthonia-type, eight-spored, apical part thickened, usually with a distinct ocular chamber. Ascospores oblong–ovoid, (3–)4(–5)-septate, upper and the lower cell enlarged, colorless, 29–35 × 12–15 μm.
Chemistry. Thallus K-, C-, KC-, UV-. No substances were detected by TLC.
Ecology and distribution. Growing on exposed trees in tropical forests. Initially reported from America (as ‘Graphis rubella’) [31]. Newly reported in China.
Additional specimens examined. China. Yunnan Province: Puer City, Jingdong County, Jinping Town, Fork River Bridge, 24°37′30″ N, 100°45′22 ″ E, alt. 1320 m, on bark, 16 August 2024, C.Y. Hao (LCUF YN241343; LCUF YN241366).
Notes. The morphological and anatomical characteristics of this study align with Arthonia rubella from America as described by Fée (as ‘Graphis rubella’) [31]. The vast majority of records document this species on bark substrates in tropical South America [31,38]. It is closely related to A. compensatula in morphology, but the latter species has no amphicephalic ascospores [31,33,34]. It is also similar to A. acanthotheciicola Ertz & Common in ascospore morphology and septation, with both featuring enlarged upper and lower cells; however, the latter typically has smaller ascospores (14.5–17 × 6–6.5 µm) [39]. The newly generated A. rubella sequences from this study are clustered with the published sequences of the same species in the phylogenetic tree (Figure 1).
Eremothecella calamicola Syd. (Figure 6).
Fungal Name: FN 145347.
Description. Thallus crustose, dispersed or continuous, smooth, green, thin, matte. Prothallus absent. Photobiont trentepohlioid, cells rectangular to irregular, ca. 9 × 5 μm. Ascomata rounded to irregular in outline, 0.3–1 mm diam., dark brown to blackish brown, non-pruinose. Asci obovate to globose, 37.5–52.5 × 33.5–40.5 µm, stipitate, eight-spored, usually with a distinct ocular chamber, K/I+ pale blue. Ascospores colorless, clavate, curved, 5–6-septate, enlarged distal cell, with very slight constrictions at septa, (28.3–)30.5–38 × 8.5–10.5 μm, I+ orange-red.
Chemistry. Thallus K-, C-, KC-, UV-. No substances were detected by TLC.
Ecology and distribution. Grows exposed trees in the shaded parts of evergreen forests. Previously reported from the Philippines [12,15], New Guinea [15], Mexico, Costa Rica, Panama, Guyana, French Guiana, Ecuador, Peru, Brazil [14], and India [16]. Newly reported in China.
Additional specimens examined. China. Yunnan Province: Jinghong City, tropical rainforests, 21°54′31″ N, 101°10′58″ E, alt. 635 m, on leaves, 17 December 2024, X. Li (LCUF YN242386-1; LCUF YN242387-1; LCUF YN242388); C.Y. Hao (LCUF YN242390-2).
Notes.Eremothecella calamicola resembles Arthonia palmulacea (Müll. Arg.) R. Sant. but the latter has shorter ascospores with fewer septa (16–25 × 4–8 μm vs. (28.3–)30.5–38 × 8.5–10.5 μm; 3–5-septate vs. 5–6-septate) [15,16]. Eremothecella variratae is another similar species regarding ascospore size (28–42 × 7–10 µm vs. (28.3–)30.5–38 × 8.5–10.5 μm), but its ascomata often feature orange-yellow pruinose [15,16]. Another morphologically similar species, E. nicobarica Jagad. Ram and G. P. Sinha. can be distinguished by larger ascospores with more septa ((14–)15–17-septate vs. 5–6-septate; 60–72 × 10–13 µm vs. (28.3–)30.5–38 × 8.5–10.5 μm) [15,16]. In the phylogenetic tree, a well-supported monophyletic clade comprising the E. calamicola specimens is sister to the new species E. pruinocarpa (Figure 1).
Synarthonia inconspicua (Stirt.) Van den Broeck and Ertz. (Figure 7).
Fungal Name: FN 825152.
Basionym. Arthonia inconspicua Stirt., Proceedings of the Philosophical Society of Glasgow 11: 319. 1879 [40].
Description. Thallus whitish to greenish-gray, smooth, continuous to cracked. Photobiont trentepohlioid, cells 9 µm in diam., rounded, usually in chains. Ascomata usually solitary, 0.25–0.6 × ca. 0.2 mm, rounded to lirellate, often forming irregular clusters, slightly elevated above thallus level, scattered more or less evenly over the thallus; disk heavily white pruinose in margin and thinner pruinose in the middle, light brown when pruina removed, flat to convex. Excipulum 16.5–21.5 µm wide, loosely intricate hyphae, inspersed with orange-brown granules which are K+ completely dissolving. Epihymenium 3.5–24.5 µm tall, and adspersed with orange-brown granules which are K+ completely dissolving. Hymenium 41.5–57.5 µm tall, hyaline, not inspersed, K/I+ blue. Paraphysoids loosely intricate around the asci. Hypothecium 17–34 µm-thick, yellowish brown, composed of loosely intricate hyphae, inspersed with orange granules which are K+ completely dissolving, K/I+ blue. Asci 49–65 × 17–24 µm, clavate, obovoid to ellipsoid or globose, stipitate, occasionally with a distinct ocular chamber, K/I−. Ascospores 17.5–24 × 6.5–8 µm, hyaline, with enlarged apical cell, oblong–ovoid, (1–)3–4(–5)-septate, spore ontogeny macrocephalic, unidirectional.
Chemistry. Thallus K-, C-, KC-, UV+. No substances were detected by TLC.
Ecology and distribution. Growing on exposed trees in tropical forests. Previously reported in the tropics of Tamil Nadu [40], India [41,42,43], Cuba, D.R. Congo, India, Madagascar, Netherlands Antilles, Rwanda, Sierra Leone, Tanzania, Uganda, and USA [17]. Newly reported in China.
Additional specimens examined. China. Yunnan Province: Mengla County, Mengla Fairyland, Menglazi Naturn Reserve, 21°42′44″ N, 101°22′32″ E, alt. 700 m, on bark, 19 December 2024, C.Y. Hao (LCUF YN242231).
Notes. The similar morphological species Synarthonia albopruinosa Van den Broeck and Ertz. differs from S. inconspicua in smaller ascospores with fewer septa (12.5–17.5 × 5–6.5 µm vs. 17.5–24 × 6.5–8 µm; (1–)2–3-septate vs. (1–)3–4(–5)-septate) [17]. Additionally, S. fuscata Van Den Broeck and Ertz. can be distinguished from S. inconspicua by the absence of pruina on the ascomata and the presence of a K/I+ blue ring-like structure in the asci [17]. In terms of molecular structure, newly generated sequences in this study of S. inconspicua were clustered with the published sequences of this species in the phylogenetic tree (Figure 1).
4. Discussion
Molecular analyses are indispensable for the classification and species identification of many lichen species within a genus due to their scarce morphological distinctions [44,45]. Employing morphological and phylogenetic analyses, this study identified three new species: Arthonia yunnanensis, A. pararubella, and Eremothecella pruinocarpa, and three new records for China: A. rubella, E. calamicola, and Synarthonia inconspicua from Yunnan Province, China. This study indicates the high species diversity of the family Arthoniaceae in Yunnan Province, China.
In the family Arthoniaceae, many taxa possess only phenotypic data but lack genotypic information, which has led to confusion regarding the taxonomic status; for example, Eremothecella. It needs to be pointed out that previous studies on this genus were mainly based on morphology, but this research provides the first molecular data for Eremothecella. Our molecular results also show that Eremothecella (E. pruinocarpa and E. calamicola) forms an independent clade within Arthoniaceae. Furthermore, to improve the clarity of generic relationships within the Arthoniaceae, sustained taxonomic investigations should be conducted in the future. Thorough surveys of the regional lichen flora will undoubtedly yield additional specimens for analysis. Empirical evidence indicates that in-depth research will uncover rarer species of Arthoniaceae, along with a considerable number of hitherto undescribed taxa.
Author Contributions
Conceptualization, C.H., S.J. and Z.J.; methodology, C.H., S.J. and Z.J.; software, C.H.; validation, C.H. and S.J.; formal analysis, C.H. and Z.J.; investigation, C.H.; resources, Z.J.; data curation, C.H., S.J. and Z.J.; writing—original draft preparation, C.H.; writing—review and editing, S.J. and Z.J.; visualization, C.H.; supervision Z.J.; project administration, S.J. and Z.J.; funding acquisition, S.J. and Z.J. All authors have read and agreed to the published version of the manuscript.
Funding
This research was funded by the National Natural Science Foundation of China (Grant No. 32270007) and the Chinese Academy of Sciences (Program CAS-TAX-24-022).
Institutional Review Board Statement
Not applicable.
Informed Consent Statement
Not applicable.
Data Availability Statement
Publicly available datasets were analyzed in this study. These data can be found at http://www.ncbi.nlm.nih.gov/ (accessed on 25 April 2025).
Acknowledgments
The authors are grateful to Wei Li, Can Cui, Tao Jia, and Xue Li for collecting specimens in the field.
Conflicts of Interest
The authors declare no conflicts of interest.
References
- Reichenbach, H.T.L. Der Deutsche Botaniker; Her barienbuch, D., Ed.; Arnoldische Buchandlung: Leipzig, Germany; Dresden, Germany, 1841; Volume 1, p. 240. [Google Scholar]
- Tehler, A. A new approach to the phylogeny of Euascomycetes with a cladistic outline of Arthoniales focussing on Roccellaceae. Can. J. Bot. 1990, 68, 2458–2492. [Google Scholar] [CrossRef]
- Grube, M. Classification and Phylogeny in the Arthoniales (Lichenized ascomycetes). Bryologist 1998, 101, 377–391. [Google Scholar] [CrossRef]
- Frisch, A.; Thor, G.; Moon, K.H.; Ohmura, Y. Arthonia incarnata (Arthoniaceae), a rare and poorly known old-growth forest lichen new to Asia. Nord. J. Bot. 2017, 35, 587–594. [Google Scholar] [CrossRef]
- Thiyagaraja, V.; Lücking, R.; Ertz, D.; Wanasinghe, D.N.; Karunarathna, S.C.; Camporesi, E.; Hyde, K.D. Evolution of non-lichenized, saprotrophic species of Arthonia (Ascomycota, Arthoniales) and resurrection of Naevia, with notes on Mycoporum. Fungal Divers. 2020, 102, 205–224. [Google Scholar] [CrossRef]
- Frisch, A.; Thor, G.; Ertz, D.; Grube, M. The Arthonialean challenge: Restructuring Arthoniaceae. Taxon 2014, 63, 727–744. [Google Scholar] [CrossRef]
- Aptroot, A.; Cáceres, M.E.D.S.; Santos, L.A.D. The taxonomy of sterile Arthoniaceae from Brazil: White crusts on overhanging tropical trees can be named. Lichenologist 2024, 56, 1–13. [Google Scholar] [CrossRef]
- Lücking, R.; Hodkinson, B.P.; Leavitt, S.D. The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota—Approaching one thousand genera. Bryologist 2017, 119, 361. [Google Scholar] [CrossRef]
- Sundin, R.; Thor, G.; Frisch, A. A literature review of Arthonia s. lat. Bibl. Lichenol. 2012, 108, 257–290. [Google Scholar]
- Grube, M.; Matzer, M.; Hafellner, J. A Preliminary Account of the Lichenicolous Arthonia Species with Reddish, K+ Reactive Pigments. Lichenologist 1995, 27, 25–42. [Google Scholar] [CrossRef]
- Cannon, P.; Ertz, D.; Frisch, A.; Aptroot, A.; Chambers, S.; Coppins, B.; Sanderson, N.; Simkin, J.; Wolseley, P. Arthoniales: Arthoniaceae, including the genera Arthonia, Arthothelium, Briancoppinsia, Bryostigma, Coniocarpon, Diarthonis, Inoderma, Naevia, Pachnolepia, Reichlingia, Snippocia, Sporodophoron, Synarthonia and Tylophoron. Revis. Br. Ir. Lichens 2020, 1, 3–48. [Google Scholar]
- Sydow, H. Beitrag zur Kenntnis der Pilzflora der Philippinen-Inseln. Ann. Mycol. 1917, 15, 165–268. [Google Scholar]
- Sérusiaux, E. Reinstatement of the lichenized genus Eremothecella Sydow. Syst. Ascomycetum 1992, 11, 39–47. [Google Scholar]
- Lücking, R. Foliicolous Lichenized Fungi; Flora Neotropica Monograph: New York, NY, USA, 2008; p. 866. [Google Scholar]
- Santesson, R. Foliicolous Lichens 1. A Revision of the Taxonomy of the Obligately Foliicolous Lichenized Fungi; Symbolae Botanicae Upsalienses: Uppsala, Sweden, 1952; Volume 12, p. 590. [Google Scholar]
- Jagadeesh Ram, T.A.M.; Sinha, G.P. New species and first records of Eremothecella (Arthoniales) from the Andaman and Nicobar Islands, India. Lichenologist 2019, 51, 507–513. [Google Scholar] [CrossRef]
- Van Den Broeck, D.; Frisch, A.; Razafindrahaja, T.; Van De Vijver, B.; Ertz, D. Phylogenetic position of Synarthonia (lichenized Ascomycota, Arthoniaceae), with the description of six new species. Plant Ecol. Evol. 2018, 151, 327–351. [Google Scholar] [CrossRef]
- Joseph, S.; Sinha, G.P. Contributions to the genus Synarthonia (lichenized Ascomycota, Arthoniaceae). Lichenologist 2015, 47, 123–130. [Google Scholar] [CrossRef]
- Ertz, D.; Aptroot, A.; Sanderson, N.; Coppins, B.; Van Den Broeck, D.; Diederich, P. A new species of Synarthonia from Luxembourg, and a new combination in the genus Reichlingia (Arthoniaceae). Lichenologist 2020, 52, 261–266. [Google Scholar] [CrossRef]
- Yao, Z.; Jiang, S.; Jia, Z. Mazosia weii sp. nov. (Roccellaceae) from China, a new species supported by molecular data. Bryologist 2021, 124, 335–342. [Google Scholar] [CrossRef]
- Cui, C.; Dou, M.; Jiang, S.; Jia, Z. The Lichen Genus Letrouitia (Brigantiaeaceae, Ascomycota) in China. Diversity 2024, 16, 254. [Google Scholar] [CrossRef]
- Orange, A.; James, P.W.; White, F.J. Microchemical Methods for the Identification of Lichens; British Lichen Society: London, UK, 2001; p. 101. [Google Scholar]
- Zoller, S.; Scheidegger, C.; Sperisen, C. Pcr Primers for the Amplification of Mitochondrial Small Subunit Ribosomal DNA of Lichen-forming Ascomycetes. Lichenologist 1999, 31, 511–516. [Google Scholar] [CrossRef]
- Liu, Y.J.; Whelen, S.; Hall, B.D. Phylogenetic relationships among ascomycetes: Evidence from an RNA polymerse II subunit. Mol. Biol. Evol. 1999, 16, 1799–1808. [Google Scholar] [CrossRef] [PubMed]
- Shi, K.; Jia, Z.; Zhao, X. A New Species and Two New Records of the Lichen Genus Fissurina from China. Diversity 2023, 15, 959. [Google Scholar] [CrossRef]
- Castresana, J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol. Biol. Evol. 2000, 17, 540–552. [Google Scholar] [CrossRef] [PubMed]
- Miller, M.A.; Pfeiffer, W.; Schwartz, T. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In Proceedings of the 2010 Gateway Computing Environments Workshop (GCE), New Orleans, LA, USA, 14 November 2010. [Google Scholar]
- Stamatakis, A. RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 2014, 30, 1312–1313. [Google Scholar] [CrossRef] [PubMed]
- Lanfear, R.; Frandsen, P.B.; Wright, A.M.; Senfeld, T.; Calcott, B. PartitionFinder 2: New Methods for Selecting Partitioned Models of Evolution for Molecular and Morphological Phylogenetic Analyses. Mol. Biol. Evol. 2017, 34, 772–773. [Google Scholar] [CrossRef]
- Ronquist, F.; Teslenko, M.; Van Der Mark, P.; Ayres, D.L.; Darling, A.; Höhna, S.; Larget, B.; Liu, L.; Suchard, M.A.; Huelsenbeck, J.P. MrBayes 3.2: Efficient Bayesian Phylogenetic Inference and Model Choice Across a Large Model Space. Syst. Biol. 2012, 61, 539–542. [Google Scholar] [CrossRef] [PubMed]
- Fée, A.L.A. Essai sur les Cryptogames des Écorces Exotiques Officinales; Firmin Didot père et fils: Paris, France, 1825; p. 180. [Google Scholar]
- Nylander, W. Synopsis du genre Arthonia. Mémoires Société Impériale Sci. Nat. Cherbg. 1856, 4, 85–104. [Google Scholar]
- Willey, H. A Synopsis of the Genus Arthonia; Willey: New Bedford, MA, USA, 1890; p. 62. [Google Scholar]
- Seavey, F.; Seavey, J. Caloplaca lecanorae (Teloschistaceae), a new lichenicolous lichen and several additions to the North American lichenized mycota from Everglades National Park. Bryologist 2012, 115, 322–328. [Google Scholar] [CrossRef]
- Vainio, E.A. Lichenes insularum Philippinarum, III. Ann. Acad. Sci. Fenn. Ser. A 1921, 15, 1–368. [Google Scholar]
- Wang, W.C.; Wei, J.C. Arthonia, Byssoloma, Calenia, Chroodiscus, Coenogonium, Eremothecella, and Semigyalecta spp. new to China. Mycotaxon 2018, 133, 487–497. [Google Scholar] [CrossRef]
- Lücking, R.; Streimann, H.; Elix, J.A. Further records of foliicolous lichens and lichenicolous fungi from Australasia, with an updated checklist for continental Australia. Lichenologist 2001, 33, 195–210. [Google Scholar] [CrossRef]
- Esslinger, T.L. A Cumulative Checklist for the Lichen-Forming, Lichenicolous and Allied Fungi of the Continental United States and Canada, Version 21. Opusc. Philolichenum 2016, 15, 136–390. [Google Scholar] [CrossRef]
- Diederich, P.; Common, R.S.; Braun, U.; Heuchert, B.; Millanes, A.; Suija, A.; Ertz, D. Lichenicolous fungi from Florida growing on Graphidales. Plant Fungal Syst. 2019, 64, 249–282. [Google Scholar] [CrossRef]
- Stirton, J. New and rare lichens from India and the Himalayas. Proc. Philos. Soc. Glasg. 1879, 11, 306–322. [Google Scholar]
- Nakanishi, M. Notes on lichen species of Graphis of the Yaeyama Islands, Japan. Hikobia Suppl. 1981, 1, 211–220. [Google Scholar]
- Müller, J. Lichenes. In Primitiae Florae Costaricensis; Durand, T.H., Pittier, H., Eds.; Bulletin de la Société Royale de Botanique de Belgique: Brussels, Belgium, 1891; Volume 30, pp. 49–97. [Google Scholar]
- Dodge, C.W. Some lichens of tropical Africa. Ann. Mo. Bot. Gard. 1953, 40, 271–401. [Google Scholar] [CrossRef]
- Jiang, S.H.; Lücking, R.; Xavier-Leite, A.B.; Cáceres, M.E.S.; Aptroot, A.; Portilla, C.V.; Wei, J.C. Reallocation of foliicolous species of the genus Strigula into six genera (lichenized Ascomycota, Dothideomycetes, Strigulaceae). Fungal Divers. 2020, 102, 257–291. [Google Scholar] [CrossRef]
- Lebreton, E.; Ertz, D.; Lücking, R.; Aptroot, A.; Carriconde, F.; Ah-Peng, C.; Huang, J.-P.; Chen, K.-H.; Stenger, P.-L.; Cáceres, M.E.D.S.; et al. Global phylogeny of the family Gomphillaceae (Ascomycota, Graphidales) sheds light on the origin, diversification and endemism in foliicolous lineages. IMA Fungus 2025, 16, e144194. [Google Scholar] [CrossRef]
Figure 1.
Phylogenetic tree of the family Arthoniaceae constructed from Bayesian analyses based on a two-gene dataset (mtSSU and RPB2). Support values are on the branches (posterior probability (PP)/bootstrap value (BS)). Bayesian inference posterior probabilities above 0.95 (Left) and maximum likelihood bootstrap support above 70% (Right) are shown at nodes. The new sequences in this study are in bold.
Figure 1.
Phylogenetic tree of the family Arthoniaceae constructed from Bayesian analyses based on a two-gene dataset (mtSSU and RPB2). Support values are on the branches (posterior probability (PP)/bootstrap value (BS)). Bayesian inference posterior probabilities above 0.95 (Left) and maximum likelihood bootstrap support above 70% (Right) are shown at nodes. The new sequences in this study are in bold.
Figure 2.
Arthonia pararubella (LCUF YN242036) (A) thallus with ascomata (B) apothecium section (C,D) asci, and with eight ascospores (E) asci in 10% KOH stained with Lugol’s solution (F–I) ascospores in water. Scale bars: 0.5 mm (A); 50 µm (B); 20 µm (C–I).
Figure 2.
Arthonia pararubella (LCUF YN242036) (A) thallus with ascomata (B) apothecium section (C,D) asci, and with eight ascospores (E) asci in 10% KOH stained with Lugol’s solution (F–I) ascospores in water. Scale bars: 0.5 mm (A); 50 µm (B); 20 µm (C–I).
Figure 3.
Arthonia yunnanensis (LCUF YN242413) (A,B) thallus with ascomata (C,D) asci, and with eight ascospores (E) asci in 10% KOH stained with Lugol’s solution (F,G) ascospores in water. Scale bars: 0.5 mm (A,B); 10 µm (C–G).
Figure 3.
Arthonia yunnanensis (LCUF YN242413) (A,B) thallus with ascomata (C,D) asci, and with eight ascospores (E) asci in 10% KOH stained with Lugol’s solution (F,G) ascospores in water. Scale bars: 0.5 mm (A,B); 10 µm (C–G).
Figure 4.
Eremothecella pruinocarpa (LCUF YN242390-1) (A) thallus, with ascomata (B) thallus, pycnidia (C) asci, and eight ascospores (D) asci in 10% KOH stained with Lugol’s solution (E) ascospores in water. (F) conidia in water. Scale bars: 1 mm (A); 0.5 mm (B); 20 µm (C–F). Fungal name: FN 572512.
Figure 4.
Eremothecella pruinocarpa (LCUF YN242390-1) (A) thallus, with ascomata (B) thallus, pycnidia (C) asci, and eight ascospores (D) asci in 10% KOH stained with Lugol’s solution (E) ascospores in water. (F) conidia in water. Scale bars: 1 mm (A); 0.5 mm (B); 20 µm (C–F). Fungal name: FN 572512.
Figure 5.
Arthonia rubella (LCUF YN241343) (A) thallus with ascomata (B) apothecium section (C) asci, and with eight ascospores (D) asci in 10% KOH stained with Lugol’s solution (E) ascospores in water. Scale bars: 1 mm (A); 50 µm (B); 20 µm (C–E).
Figure 5.
Arthonia rubella (LCUF YN241343) (A) thallus with ascomata (B) apothecium section (C) asci, and with eight ascospores (D) asci in 10% KOH stained with Lugol’s solution (E) ascospores in water. Scale bars: 1 mm (A); 50 µm (B); 20 µm (C–E).
Figure 6.
Eremothecella calamicola (LCUF YN242388) (A,B) thallus with ascomata (C) asci, with eight ascospores (D) asci in 10% KOH stained with Lugol’s solution (E) ascospores in water. Scale bars: 2 mm (A); 1 mm (B); 20 µm (C–E).
Figure 6.
Eremothecella calamicola (LCUF YN242388) (A,B) thallus with ascomata (C) asci, with eight ascospores (D) asci in 10% KOH stained with Lugol’s solution (E) ascospores in water. Scale bars: 2 mm (A); 1 mm (B); 20 µm (C–E).
Figure 7.
Synarthonia inconspicua (LCUF YN242231): (A,B) thallus with ascomata (C) apothecium section (D) apothecium section (in polarized light) showing distinct patches of calcium oxalate crystals (E) excipulum, K+ completely dissolving (F,G) asci, with eight ascospores (H) asci in 10% KOH stained with Lugol’s solution (I–K) ascospores in water. Scale bars: 2 mm (A); 1 mm (B); 50 µm (C–E); 20 µm (F, G); 50 µm (H); 20 µm (I–K).
Figure 7.
Synarthonia inconspicua (LCUF YN242231): (A,B) thallus with ascomata (C) apothecium section (D) apothecium section (in polarized light) showing distinct patches of calcium oxalate crystals (E) excipulum, K+ completely dissolving (F,G) asci, with eight ascospores (H) asci in 10% KOH stained with Lugol’s solution (I–K) ascospores in water. Scale bars: 2 mm (A); 1 mm (B); 50 µm (C–E); 20 µm (F, G); 50 µm (H); 20 µm (I–K).
Table 1.
Specimens of Arthoniaceae and the outgroup were used in the phylogenetic analyses.
| Species | Specimen Voucher | GenBank Accession Number | |
|---|---|---|---|
| mtSSU | RPB2 | ||
| Arthonia atra | L10170 | KY983978 | KY983985 |
| Arthonia calcarea | Thor 11/6a | KJ850974 | KJ851105 |
| Arthonia calcarea | Ertz 7545 | EU704063 | EU704027 |
| Arthonia didyma | Ertz 7587 | EU704047 | EU704010 |
| Arthonia didyma | Frisch 11/Se41 | - | KJ851106 |
| Arthonia granithophila | Frisch 10/Se74 | KJ850981 | KJ851107 |
| Arthonia graphidicola | Frisch 10/Jp102 | KJ850980 | - |
| Arthonia ilicina | McCune 31067 | KJ850982 | - |
| Arthonia incarnata | Frisch 10/Jp94 | KY983972 | KY983980 |
| Arthonia incarnata | Frisch 12/Jp289 | KY983973 | KY983981 |
| Arthonia incarnata | Frisch 13/Jp215 | KY983974 | KY983982 |
| Arthonia maculiformis | Wedin 9393b | - | KF707658 |
| Arthonia pararubella | YN242036 | PV569153 | PV590068 |
| Arthonia pararubella | YN241783 | PV569152 | - |
| Arthonia physcidiicola | Frisch 11/Ug318 | KF707646 | KF707657 |
| Arthonia radiata | Frisch 10/Se29 | KJ850968 | KJ851108 |
| Arthonia radiata | Frisch 11/Se25 | KJ850969 | KJ851109 |
| Arthonia rubella | 49024 | MH308714 | - |
| Arthonia rubella | YN241343 | PV569139 | - |
| Arthonia rubella | YN241366 | PV569140 | - |
| Arthonia subfuscicola | Thor 11/1 | KJ850971 | KJ851110 |
| Arthonia subfuscicola | Frisch 11/Se15 | KJ850972 | KJ851111 |
| Arthonia thoriana | Sanderson 2176 | MG207687 | - |
| Arthonia thoriana | Sanderson 2174 | MG207685 | - |
| Arthonia yunnanensis | YN242413 | PV569142 | PV575682 |
| Arthonia yunnanensis | YN242411 | PV569141 | PV575683 |
| Arthonia atra | L10170 | KY983978 | KY983985 |
| Arthonia calcarea | Thor 11/6a | KJ850974 | KJ851105 |
| Arthonia calcarea | Ertz 7545 | EU704063 | EU704027 |
| Arthonia didyma | Ertz 7587 | EU704047 | EU704010 |
| Arthonia didyma | Frisch 11/Se41 | - | KJ851106 |
| Arthonia granithophila | Frisch 10/Se74 | KJ850981 | KJ851107 |
| Arthonia graphidicola | Frisch 10/Jp102 | KJ850980 | - |
| Arthonia ilicina | McCune 31067 | KJ850982 | - |
| Arthonia incarnata | Frisch 10/Jp94 | KY983972 | KY983980 |
| Arthonia incarnata | Frisch 12/Jp289 | KY983973 | KY983981 |
| Arthonia incarnata | Frisch 13/Jp215 | KY983974 | KY983982 |
| Arthonia maculiformis | Wedin 9393b | - | KF707658 |
| Arthonia pararubella | YN242036 | PV569153 | PV590068 |
| Arthonia pararubella | YN241783 | PV569152 | - |
| Arthonia physcidiicola | Frisch 11/Ug318 | KF707646 | KF707657 |
| Arthonia radiata | Frisch 10/Se29 | KJ850968 | KJ851108 |
| Arthonia radiata | Frisch 11/Se25 | KJ850969 | KJ851109 |
| Arthonia rubella | 49024 | MH308714 | - |
| Arthonia rubella | YN241343 | PV569139 | - |
| Arthonia rubella | YN241366 | PV569140 | - |
| Arthonia subfuscicola | Thor 11/1 | KJ850971 | KJ851110 |
| Arthonia subfuscicola | Frisch 11/Se15 | KJ850972 | KJ851111 |
| Arthonia thoriana | Sanderson 2176 | MG207687 | - |
| Arthonia thoriana | Sanderson 2174 | MG207685 | - |
| Arthonia yunnanensis | YN242413 | PV569142 | PV575682 |
| Arthonia yunnanensis | YN242411 | PV569141 | PV575683 |
| Arthothelium galapagoense | Ertz 11790 | - | HQ454657 |
| Arthothelium galapagoense | Ertz 11654 | - | HQ454658 |
| Arthothelium norvegicum | McCune 31061 | - | KJ851114 |
| Arthothelium orbilliferum | TRH-L-15449 | KY983977 | - |
| Arthothelium punctatum | KoLRI 044205 | MF616614 | - |
| Arthothelium punctatum | KoLRI 044206 | MF616615 | - |
| Arthothelium ruanum | KoLRI 038018 | MF616609 | MF616619 |
| Arthothelium ruanum | KoLRI 038257 | MF616610 | MF616620 |
| Arthothelium ruanum | KoLRI 038261 | MF616611 | MF616621 |
| Arthothelium ruanum | KoLRI 038275 | MF616612 | - |
| Arthothelium ruanum | KoLRI 038275 | MF616613 | - |
| Arthothelium ruanum | Zimmerman 1117 | GU327683 | - |
| Arthothelium sp. | Joensson Guyana 10 | KJ850957 | KJ851094 |
| Arthothelium sp. | Joensson Guyana 8 | KJ850958 | KJ851095 |
| Arthothelium spectabile | TNS:Frisch12Jp179a | KP870144 | KP870160 |
| Briancoppinsia cytospora | Diederich 16849 | JF830771 | - |
| Briancoppinsia cytospora | Ertz 15244 | JF830772 | - |
| Bryostigma apatetica | Svensson 2017 | KJ850992 | KJ851125 |
| Bryostigma apatetica | Svensson 1939 | KJ850993 | KJ851126 |
| Bryostigma apotheciorum | Frisch 11/Se23 | KJ850970 | KJ851148 |
| Bryostigma biatoricola | Thor 24350 | KJ850990 | KJ851149 |
| Bryostigma lapidicola | Westberg Frisch 11/Se47 | KJ850997 | KJ851119 |
| Bryostigma lobariicola | Frisch 10/Jp124 | KJ851002 | KJ851127 |
| Bryostigma lobariicola | Frisch 10/Jp737 | KJ851001 | KJ851128 |
| Bryostigma molendoi | Frisch 11/Se36 | KJ851000 | KJ851117 |
| Bryostigma molendoi | Frisch 11/Se22 | MH177777 | MH177770 |
| Bryostigma muscigenum | Thor 26206 | KJ850991 | KJ851124 |
| Bryostigma neglectula | Frisch 10/Se91 | KJ850989 | KJ851118 |
| Bryostigma parietinaria | Frisch 17/No25 | MH177778 | MH177771 |
| Bryostigma peltigerina | Westberg Frisch 11/Se46 | KJ850998 | KJ851122 |
| Bryostigma phaeophysciae | Ertz 11/2 | - | KJ851112 |
| Bryostigma sp. | Svensson 2324 | KJ851019 | KJ851123 |
| Bryostigma sp. | SK L06 | MH510022 | MH510019 |
| Bryostigma sp. | Svensson 2148 | KJ850995 | KJ851120 |
| Bryostigma sp. | SK L04 | MH510023 | MH510020 |
| Bryostigma stereocaulina | Westberg Frisch 11/Se48 | KJ850999 | - |
| Bryostigma toensbergii | Frisch N4-2-Pa4-1 | MH177775 | - |
| Bryostigma toensbergii | Frisch N4-2-Pa6-1 | MH177776 | - |
| Coniocarpon aff. cinnabarinum | Frisch 11/Ug3 | KJ850978 | KJ851102 |
| Coniocarpon cinnabarinum | Johnsen111003 | KJ850976 | KJ851103 |
| Coniocarpon cinnabarinum | Frisch11Ug296 | KP870158 | KP870170 |
| Coniocarpon fallax | LD:L10075 | KJ850979 | KJ851101 |
| Coniocarpon rubrocinctum | Nelsen 4010 | GU327684 | - |
| Crypthonia aff. vandenboomii | Frisch 11/Ug21 | KJ850960 | KJ851085 |
| Crypthonia palaeotropica | Frisch 11/Ug457 | KJ850961 | KJ851084 |
| Crypthonia palaeotropica | Frisch11Ug26B | KP870145 | KP870161 |
| Cryptophaea phaeospora | Van den Broeck 5809 | KX077541 | - |
| Cryptophaea phaeospora | Van den Broeck 5964 | KX077540 | - |
| Cryptothecia assimilis | Lumbsch 198151 | GU327688 | - |
| Cryptothecia austrocoreana | KoLRI No. 041892 | MF769374 | - |
| Cryptothecia austrocoreana | KoLRI No. 044721 | MF769375 | - |
| Cryptothecia punctosorediata | F:Nelsen 4038 | JX046450 | - |
| Cryptothecia sp. | Frisch 11/Ug39 | KJ850954 | KJ851086 |
| Cryptothecia sp. | Frisch 11/Ug18 | KJ850955 | KJ851092 |
| Cryptothecia sp. | Frisch 11/Ug194 | KJ850956 | KJ851093 |
| Cryptothecia subnidulans | v.d.Boom 40613 | KJ850952 | KJ851087 |
| Cryptothecia subnidulans | Joensson Guyana 6a | KJ850953 | KJ851088 |
| Diarthonis spadicea | Thor 26200 | - | KJ851116 |
| Diarthonis spadicea | Frisch 11/Se31 | - | KJ851115 |
| Eremothecella pruinocarpa | YN242390-1 | PV569148 | PV575678 |
| Eremothecella pruinocarpa | YN242384 | PV569149 | PV575679 |
| Eremothecella pruinocarpa | YN242386-2 | PV569150 | PV575680 |
| Eremothecella pruinocarpa | YN242387-2 | PV569151 | PV575681 |
| Eremothecella calamicola | YN242386-1 | PV569146 | - |
| Eremothecella calamicola | YN242387-1 | PV569147 | - |
| Eremothecella calamicola | YN242388 | PV569144 | - |
| Eremothecella calamicola | YN242390-2 | PV569145 | PV575677 |
| Glomerulophoron mauritiae | BR Ertz19164 | KP870153 | KP870167 |
| Herpothallon inopinatum | Rudolphi 12 | KJ850964 | KJ851099 |
| Herpothallon kigeziense | Frisch 11/Ug26 | KF707644 | KF707654 |
| Herpothallon rubrocinctum | Rudolphi 5 | KF707643 | KF707655 |
| Herpothallon rubrocinctum | Nelsen 4006 | GU327693 | - |
| Herpothallon sp. | Frisch 11/Ug401 | KF707645 | KF707653 |
| Inoderma afromontanum | Frisch 11/Ug164 | KJ850963 | KJ851090 |
| Inoderma byssaceum | Thor 25952 | KJ850962 | KJ851089 |
| Inoderma byssaceum | Lif 186 | - | KJ851091 |
| Inoderma nipponicum | TNS:Frisch12Jp227 | KP870146 | KP870162 |
| Inoderma nipponicum | TNS:Frisch13Jp1 | KP870147 | KP870163 |
| Inoderma sorediatum | Kukwa 15631 & Lubek | MG207690 | - |
| Inoderma subabietinum | Ertz16885 | KP870150 | KP870164 |
| Leprantha cinereopruinosa | Kukwa 17127 & Lubek | MG207692 | - |
| lnoderma sorediatum | Kukwa 15630 & Lubek | MG207689 | - |
| Melarthonis piceae | Thor 25995 | KJ851016 | - |
| Myriostigma candidum | Ertz 9260 | EU704052 | EU704015 |
| Myriostigma candidum | Frisch 11/Ug125 | KJ850959 | KJ851096 |
| Myriostigma miniatum | Silva T2A29 | KP843606 | - |
| Naevia aff punctiformis | Thor 24702 | KJ850975 | - |
| Naevia dispersa | UPSC 2583 | AY571383 | - |
| Naevia pinastri | MFLU 17-3497 | MN842780 | - |
| Naevia punctiformis | Thor 26158 | KJ850973 | KJ851113 |
| Pachnolepia pruinata | Tehler 9139 | - | HQ454650 |
| Pachnolepia pruinata | Frisch 11/Se34 | KJ850967 | KJ851098 |
| Reichlingia leopoldii | Ertz 13293 | JF830773 | HQ454722 |
| Reichlingia leopoldii | Ertz 13294 | JF830774 | HQ454723 |
| Reichlingia syncesioides | Frisch 11/Ug14 | KF707651 | KF707656 |
| Reichlingia zwackhii | Thor 11/3 | KF707652 | KF707662 |
| Reichlingia zwackhii | Ertz 10928 | - | HQ454655 |
| Snippocia nivea | Ertz 17437 | MG207695 | - |
| Snippocia nivea | Sanderson 2180 | MG207694 | - |
| Snippocia nivea | Sanderson 2179 | MG207693 | - |
| Sporodophoron cretaceum | Thor27720 | KP870159 | - |
| Sporodophoron gossypinum | TNS:Frisch12Jp186 | KP870154 | KP870168 |
| Sporodophoron gossypinum | TNS:Frisch12Jp197 | KP870156 | - |
| Sporodophoron primorskiense | TNS:Y.Ohmura10607 | LC086299 | - |
| Sporodophoron primorskiense | TNS:Ohmura10509 | KP870157 | KP870169 |
| Stirtonia neotropica | Cáceres & Aptroot 11112 | KP843611 | - |
| Stirtonia sp. | Frisch 11/Ug325 | KJ850965 | - |
| Synarthonia albopruinosa | VDB 6086 | MH251873 | - |
| Synarthonia aurantiacopruinosa | VDB 5764 | MH251874 | - |
| Synarthonia fuscata | VDB 6101 | MH251875 | - |
| Synarthonia inconspicua | VDB 7013B | MH251881 | - |
| Synarthonia inconspicua | ERTZ 19739A | MH251879 | - |
| Synarthonia inconspicua | YN242231 | PV569143 | - |
| Synarthonia josephiana | ERTZ 19739B | MH251876 | - |
| Synarthonia muriformis | Frisch 11/Ug41 | KJ851025 | - |
| Synarthonia muriformis | ERTZ 19344 | MH251877 | - |
| Synarthonia ochracea | VDB 6653 | MH251884 | - |
| Synarthonia pilosella | ERTZ 7808 | MH251883 | - |
| Tylophoron galapagoense | Bungartz 8749 | JF830776 | - |
| Tylophoron galapagoense | Bungartz 8750 | JF830777 | - |
| Tylophoron hibernicum | Diederich 16335 | JF830779 | - |
| Tylophoron hibernicum | Frisch 11/Ug220 | KJ850966 | KJ851097 |
| Tylophoron moderatum | Ertz 14504 | JF830780 | - |
| Tylophoron stalactiticum | Ertz 10880 | JF830781 | - |
Notes: Newly generated sequences are shown in bold.
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MDPI and ACS Style
Hao, C.; Jiang, S.; Jia, Z. Three New Species and Three New Records of Arthoniaceae (Ascomycota, Arthoniales) from China. J. Fungi 2026, 12, 42. https://doi.org/10.3390/jof12010042
AMA Style
Hao C, Jiang S, Jia Z. Three New Species and Three New Records of Arthoniaceae (Ascomycota, Arthoniales) from China. Journal of Fungi. 2026; 12(1):42. https://doi.org/10.3390/jof12010042
Chicago/Turabian StyleHao, Chengyue, Shuhua Jiang, and Zefeng Jia. 2026. "Three New Species and Three New Records of Arthoniaceae (Ascomycota, Arthoniales) from China" Journal of Fungi 12, no. 1: 42. https://doi.org/10.3390/jof12010042
APA StyleHao, C., Jiang, S., & Jia, Z. (2026). Three New Species and Three New Records of Arthoniaceae (Ascomycota, Arthoniales) from China. Journal of Fungi, 12(1), 42. https://doi.org/10.3390/jof12010042
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