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Article

Taxonomic Revision of Solorina (Peltigeraceae, Ascomycota), Reveals a New Genus and Three New Species

by
Ting Zheng
1,2,3,
Lisong Wang
1,4,
Min Ai
1,4,
Yuxin Gan
1,4,
Rong Fan
1,
Yingjun Zhang
1,
Fiona Ruth Worthy
1,4,
Jizhen Jin
1,4,5,
Wenping Meng
2,3,
Shengbang Zhang
6 and
Xinyu Wang
1,4,*
1
State Key Laboratory of Phytochemistry and Natural Medicines, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China
2
Guizhou Botanical Garden, Guiyang 550004, China
3
Key Laboratory for Biodiversity Conservation in Karst Mountain Area of Southwestern China of the National Forestry and Grass Administration, Guiyang 550004, China
4
Yunnan Key Laboratory for Fungal Diversity and Green Development, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China
5
Key Laboratory of Plant Stress Research, College of Life Sciences, Shandong Normal University, Jinan 250014, China
6
Qinghai Shanshui Natural Resources Survey Institute, Xining 810008, China
*
Author to whom correspondence should be addressed.
J. Fungi 2025, 11(3), 169; https://doi.org/10.3390/jof11030169
Submission received: 13 January 2025 / Revised: 12 February 2025 / Accepted: 18 February 2025 / Published: 20 February 2025
(This article belongs to the Section Fungal Evolution, Biodiversity and Systematics)
Browse Figures
Versions Notes

Abstract

:
The lichen genus Solorina exhibits significant morphological and chemical variations between species. Recent molecular studies have demonstrated that Solorina is polyphyletic, underscoring the need for a comprehensive taxonomic revision. Phylogenetic analyses employing Bayesian methods and Maximum Likelihood approaches based on three molecular loci (nrITS, nrLSU, mtSSU) revealed that species of Solorina segregate into two distinct clades. The first clade includes species characterized by bright orange lower surfaces that contain secondary metabolites, notably solorinic acid. The type species, Solorina crocea, is retained in the genus Solorina. The second clade encompasses species with white or brownish lower surfaces; most species lack secondary metabolites and are now classified as a new genus, Pseudosolorina. As a result of this taxonomic revision, two species: S. crocea and S. crocoides remain in the genus Solorina. Five species with white or brownish lower surfaces were transferred to the new genus Pseudosolorina, which consists of three newly described species and five new combinations. Four species previously described as Solorina: S. embolina, S. fuegiensis, S. octospora, and S. platycarpa have morphology consistent with Pseudosolorina, but are currently retained in Solorina due to the absence of supporting DNA sequence data. A key to Solorina and Pseudosolorina is provided. The spores of S. crocea exhibit wall ornamentation featuring rounded papillae, which are distinct from those of Pseudosolorina. Molecular data and morphological characters also indicate that both Solorina and Pseudosolorina engage in symbiotic associations with photobionts cyanobacteria Nostoc and chlorophytes Coccomyxa or Asterochloris.

1. Introduction

The lichen genus Solorina Ach. belongs to Peltigeraceae, Peltigerales, Lecanoromycetidae, Lecanoromycetes, Ascomycota [1,2]. Since Linné first reported the lichen species Lichen croceus L. and Lichen saccatus L. in the 1750s [3,4], the taxonomic placement of these species has garnered significant interest among lichenologists. After several modifications, Acharius established the new genus Solorina Ach. to accommodate these two species, with S. crocea (L.) Ach. designated as the type species. The genus Solorina is characterized by its fragile foliose thallus, paraplectenchymatous upper cortex, and large, rounded apothecia that are immersed in the upper surface, without a margin, typically exhibiting a concave disc that ranges from dark red-brown to nearly black [1]. Later lichenologists, including Nylander and Gyelnik, assigned additional species to this genus [5,6,7,8,9,10,11,12]. Currently, Solorina comprises approximately ten species [13], most of which inhabit calcareous soils in high-altitude mountainous regions and exhibit a bipolar distribution. Six species and one variant have been reported from China [14].
Species of Solorina exhibit considerable variation in thallus morphology and the composition of their photobiont layers. Differences in these photobiont layers have been considered a taxonomically distinctive trait. Nylander described the genus Solorinina Nyl. as containing a cyanobacterial photobiont layer, with Solorina possessing a chlorophyte photobiont layer [15]; Hue further classified Solorina into three sections, Pleurothea Hue with two separate photobiont layers, Solorina accommodating three species with a single cyanobacterial photobiont layer, and Eusolorina Hue accommodating species featuring a single chlorophyte photobiont layer [12]. Gyelnik reclassified Solorina into two subgenera: Eusolorina Hue and Solorinina (Nyl.) Hue, and further divided them into four sections (Protosolorina Gyeln., Neosolorina Gyeln., Protosolorinina Gyeln., and Neosolorinina Gyeln.), based on variations in photobiont layers and the size of thallus [16]. Räsänen proposed the section Neosolorina (Gyeln.) Räsänen as a new genus [17]. However, these classifications are no longer widely recognized.
Currently, the species of Solorina are primarily distinguished based on the morphology of their thallus and ascospores. Ascospore numbers differ between species, with formations including 1, 2, 4, and 8 spores per ascus; the spores are thick-walled and appear roughened under light microscopy. Some studies classify species of Solorina based on the number and ornamentation of the spores [18,19,20]. Since most Solorina species lack secondary metabolites, only four species have been distinguished based on their chemical characteristics, notably S. crocea, which contains numerous secondary metabolites, including solorinic acid and norsolorinic acid [21,22]. The morphology of S. crocea and S. crocoides (Nyl.) Gyeln. differ significantly from other Solorina species by the bright orange color of their lower surface.
Recent molecular studies indicated that S. crocea diverged from all other species of Solorina [2,23]. DNA data were lacking, and records were scarce for S. crocoides. Because S. crocoides was initially reported from the Himalayas, for the purposes of this study we collected new samples from Xizang, China, from which DNA sequences were obtained. We also collected a further 36 specimens of other Solorina spp., from which we obtained DNA sequences. The phylogenetic result confirms that Solorina is polyphyletic and introduces a new genus, Pseudosolorina T. Zheng and Li S. Wang.

2. Materials and Methods

2.1. Morphological, Anatomical and Chemical Characters

Most materials for this study were collected in China, and specimens were stored in the Herbarium, Kunming Institute of Botany, Chinese Academy of Sciences (KUN). Type specimens of S. embolina, S. octospora, and S. simensis were borrowed from the University of Helsinki (H). High-resolution photographs of type or lectotype specimens of S. crocea, S. bispora, S. fuegiensis, and S. saccata were provided by the curators of Linnaean Herbarium (LINN) or obtained from the website of Global Plants (https://plants.jstor.org/, accessed on 12 September 2023).
The specimens were examined using standard microscopy techniques. Morphological descriptions were based on observations using a Nikon SMZ 745T (Minato City, Japan) dissecting microscope and a Zeiss Axio Scope A1 stereomicroscope (Oberkochen, Germany). Scanning electron microscopy ZEISS SIGMA 300 (Oberkochen, Germany) was conducted on selected specimens to visualize spore ornamentation. The identification of secondary metabolites was performed using thin-layer chromatography (TLC) [24,25] and gradient-elution high-performance liquid chromatography (HPLC) [26].

2.2. DNA Extraction, PCR Amplification and Sequencing

Genomic DNA was extracted from dry or fresh specimens following the manufacturers’ instructions using the DNA secure Plant Kit (Tiangen, Beijing, China). Three gene loci of Solorina were amplified using the following primers: ITS1F [27], ITS4 [28], LR0R [29], LR5 [30], mrSSU1 and mrSSU3R [31]. The 16S rRNA genes of cyanobacteria were amplified with primers CYA359F and CYA781R-a [32]. The nrITS genes of the chlorophytes were amplified with primers nr-SSU-1780-5′ and ITS4T [33,34]. The 25 μL PCR mixture consisted of 2 μL DNA template, 1 μL of each primer, 12.5 μL 2 × Taq PCR MasterMix (Aidlab, Hong Kong) (Taq DNA Polymerase [0.1 unit/mL]; 4 mM MgCl2; and 0.4 mM dNTPs) and 8.5 μL ddH2O. The PCR settings and primer profile followed Zhao et al. [35]. Polymerase chain reaction (PCR) products were sequenced by Sangon Biotech (Shanghai, China).

2.3. Molecular Phylogenetic Analyses

The raw sequences were assembled and edited using SeqMan v.7.0 (DNAstar packages). Sequences extracted from new materials corresponding to each gene locus were aligned with additional sequences available from GenBank (Table 1) using the Concatenate Sequence feature and MAFFT v.7.505 [36] to generate nrITS-nrLSU-mtSSU matrices. The concatenated alignments were estimated by PartitionFinder 2.1.1 [37], based on the Bayesian Information Criterion (BIC), to find the most appropriate nucleotide substitution model for each of the three loci Phylogenetic relationships were inferred using Bayesian Inference (BI) and Maximum Likelihood (ML) methods, with the genus Pannaria as the outgroup. ML analyses were performed with IQ-TREE v2.2.0 [38] using the model SYM + I + G for nrITS, GTR + G for mtSSU and nrLSU. The Bayesian method was performed with MrBayes v.3.2.7 [39]. Four Markov chains were run with 2 million generations for each dataset, and trees were sampled every 100 generations. Convergence was assessed by ensuring that the average standard deviation of split frequencies was less than 0.01. Posterior probabilities greater than 0.9 and bootstrap support values above 80% were considered to provide significant support for the phylogenetic relationships. All the trees were visualized using FigTree v. 1.4.0 [40]. The identity of cyanobacteria and chlorophyte sequences were determined through BLAST analysis on the NCBI online server (https://blast.ncbi.nlm.nih.gov/Blast.cgi, accessed on 22 November 2024).

3. Results

3.1. Phylogenetic Analysis

In this study, we generated 92 new sequences from 39 specimens and downloaded 78 additional sequences from GenBank; the final alignment for the combined regions dataset (620 bp for nuITS sequences; 552 bp for nrLSU sequences; 605 bp for mtSSU sequences) comprised 66 terminals (Table 1). Results from phylogenetic analyses presented here clearly indicate that the species of Solorina were divided into two well-supported clades: clade 1 and clade 2 (Figure 1). Clade 1 contains the type species of Solorina, S. crocea, so we retain this clade as the revised genus Solorina. The revised Solorina clade contains two species S. crocea and S. crocoides; these both have a bright orange lower surface, which is distinctly veined.
All species in clade 2, are characterized by a white or brownish lower surface. Clade 2 forms a monophyletic group with strong support for its division from clade 1. Consequently, we propose a new genus, Pseudosolorina, to classify this clade. Here, we present three new species and five new combinations for the new genus Pseudosolorina. We further subdivide the genus Pseudosolorina into three groups. The first group contains three species, P. hepatizon, P. simensis, and P. tenuior, which share thallus and apothecium morphological characters and have a single photobiont layer containing both cyanobacteria and chlorophytes. Group 2 contains two species, P. parmigera and P. saccata, with a green, chlorophyte photobiont layer and thallus over 2 cm in diam. Group 3 contains three species, P. bispora, P. bispora var. monospora, and P. spongiosa, with a green, chlorophyte photobiont layer and thallus less than 2 cm in diam. The smallest thallus diameter is for P. spongiosa, which is reduced to a circle around the apothecium.

3.2. Chemical Analysis

Solorina crocea contained several secondary metabolites, including solorinic acid, averantin, 6-O-methylaverantin, and gyrophoric acid. S. crocoides shared the same chemical constituents as S. crocea, except for the absence of gyrophoric acid. In the genus Pseudosolorina, the species P. tenuior and P. simensis contained gyrophoric acid, methyl gyrophorate, tenuiorin and 2′-O-methyltenuiorin. Other species in the genus: P. bispora, P. bispora var. monospora, P. hepatizon, P. parmigera, P. saccata and P. spongiosa lacked secondary metabolites.

3.3. Scanning Electron Microscopy for Spore Ornamentation

Analysis of spores by SEM (Figure 2) showed that Solorina crocea has a relatively smooth spore surface characterized by nipple-like ornamentations (Figure 2A). Conversely, the species of Pseudosolorina have a rough spore surface adorned with generally broad reticulating ridges and irregular lacunae ornamentations (Figure 2B–I). Although there are some differences in ornamentations among the various species of Pseudosolorina, distinguishing them solely based on this character would be challenging. No apothecia have been observed for S. crocoides, so no images are presented for the spores of this species.

3.4. Composition of the Photobionts

Molecular evidence suggests that Solorina and Pseudosolorina share photobionts, comprising cyanobacteria Nostoc and chlorophytes Coccomyxa or Asterochloris (Table 2). Morphological examinations of specimens of S. crocea reveal the presence of two distinct layers of photobionts: an upper bright green layer and a discontinuous blue lower layer. Previous reports indicated that S. crocea also contains the photobionts Coccomyxa and Nostoc [41], suggesting that they have an upper layer of Coccomyxa and a lower layer of Nostoc. In contrast, specimens from the species S. crocoides, exhibited only a single photobiont layer with both blue and bright green cells and did not possess cephalodia. Molecular data showed that these photobionts were Nostoc and Asterochloris, indicating that S. crocoides contains both cyanobionts and chlorobionts within the same photobiont layer. Within Pseudosolorina: P. hepatizon, P. simensis, and P. tenuior share characteristics of the photobiont layer found for S. crocoides: they contain Nostoc and Asterochloris within the same single photobiont layer. In contrast, P. bispora, P. bispora var. monospora, P. parmigera, P. saccata and P. spongiosa contain Nostoc and Coccomyxa, but with a single green photobiont layer only contain Coccomyxa, and Nostoc present only in cephalodia.

3.5. Revised Boundary for the Genus Solorina

The genus Solorina has traditionally been recognized by its large, rounded, impressed apothecia that are immersed in the upper surface and typically concave, without a thalline margin. These traits distinguish Solorina from its sister groups, Peltigera Willd. and Sinuicella D. F. Stone, McCune & Miądl. However, significant morphological and chemical variations exist among the species within this genus. Furthermore, molecular studies suggest that Solorina is polyphyletic [2,23], indicating that these traditional characteristics are inadequate for reliably distinguishing this genus.
Our three-loci-based analysis confirms previous findings regarding the polyphyly of Solorina [2,23], revealing the division of this genus into two distinct clades. Based on the phylogenetic analyses and morphological and chemical characters, we define these two clades as representing two genera. The clade with a bright orange lower surface, which includes the type species S. crocea, was designated as the revised Solorina. According to this re-circumscription, species within Solorina are characterized by a bright orange lower surface and the presence of distinctive secondary metabolites, including solorinic acid. These species have apothecia which are flat or convexly immersed in the upper surface of the thallus (or apothecia absent), spores with nipple-like ornamentations, photobiont layer containing both cyanobacteria Nostoc and chlorophyte Coccomyxa or Asterochloris, secondary metabolites including solorinic acid, averantin, 6-O-methylaverantin, and gyrophoric acid. The combination of these secondary metabolites produces the characteristic orange lower surface.
Within the second well-defined monophyletic clade, all species have white or brownish lower thallus surfaces. This clade was established as a new genus Pseudosolorina to accommodate the species which were excluded from the revised definition of Solorina. The species within Pseudosolorina all exhibit apothecia immersed in the upper surface and slightly to deeply concave; their spores are distinguished by having broad reticulating ridges and irregular lacunae ornamentations. The photobiont layer contains Nostoc and Asterochloris or solely Coccomyxa, with Nostoc confined to cephalodia. Most species in Pseudosolorina lack secondary metabolites. This taxonomic revision provides new classification boundaries for the genus Solorina and Pseudosolorina, based on morphological, phylogenetic, and chemical characteristics.
There are four other species previously defined as Solorina, for which genetic material was unavailable. Of these, S. octospora Arnold and S. platycarpa Hue had been reported from China, whereas S. embolina Nyl. and S. fuegiensis C.W. Dodge were reported from elsewhere. These four species have white or brownish lower surfaces; thus, morphologically, they could be assigned to the new genus Pseudosolorina. However, due to the absence of sequence data, they must currently be retained within Solorina. Further research is needed to clarify the phylogenetic position of these species.

3.6. Taxonomy

Solorina Ach., K. Vetensk-Acad. Nya Handl. 29: 228 (1808)
Type species: Solorina crocea (L.) Ach., K. Vetensk-Acad. Nya Handl. 29: 228 (1808)
Description: Thallus: large foliose, fragile, wide-spreading, dorsiventral, heteromerous, lobate; Lobes: rounded, margins raised; Upper surface: olive-green to green-gray when wet, red-brown to gray-brown when dry, smooth to scabrid, one species with soredia; Upper cortex: paraplectenchymatous; Photobiont: Coccomyxa and Nostoc; Lower cortex: distinct lower cortex lacking; Lower surface: bright orange, distinctly brownish veined, tomentose, with clusters of simple or branched rhizines; Apothecia: present or absent, large, rounded, irregularly scattered, not or only a little depressed into the thallus, flat or convex, thalline margin absent; Asci: clavate, Peltigera-type, 8-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall surface ornamented with rounded papillae.
Chemistry: Solorinic acid, averantin, 6-O-methylaverantin and gyrophoric acid (detected by TLC and HPLC).
Ecology: Distributed in high-altitude areas and mainly grows on calcareous soil.
Note: Species with apothecia immersed into the upper surface and without margins were previously included in Solorina. However, a three-loci phylogenetic analysis confirmed that this genus was polyphyletic. The species with white or brown lower surfaces were excluded from Solorina to establish a new genus named Pseudosolorina. Solorina is now characterized by its immersed apothecia, bright orange lower surface, rounded papillate ornamentation of the spores, and distinctive secondary metabolites, notably solorinic acid. Members of this genus are typically found in alpine habitats.
Solorina crocea (L.) Ach., K. Vetensk-Acad. Nya Handl. 29: 228 (1808) Figure 3
≡ Lichen croceus L., Sp. pl. 2: 1149 (1753)
= Peltigera crocea (L.) Hoffm., Descript. et Adumbr. Plant. Lich. 2(3): 60 (1794)
= Peltidea crocea (L.) Ach., Methodus, Sectio post. (Stockholmiæ): 290 (1803)
= Arthonia crocea (L.) Ach., Neues J. Bot. 1(3. Stück): 20 (1806)
= Parmelia crocea (L.) Spreng., Syst. veg., Edn 16 4(1): 280 (1827)
= Solorina crocea (L.) Ach., Magy. Bot. Lapok, 29: 30 (1930)
Type: LINN-HL1273-189 (lectotype, designated by Howe, Bull. Torrey Bot. Club 39: 201 (1912))
Description: Thallus: large foliose, wide-spreading, dorsiventral, heteromerous, lobate, up to 10 cm in diam.; Lobes: rounded, margins raised; Upper surface: olive green when wet, red-brown when dry, smooth to scabrid; Upper cortex: paraplectenchymatous, colorless or slightly yellow, measuring 70–150 µm; Medulla: 217–280 µm thick; Photobiont: comprises two photobiont layers, the upper layer contains Coccomyxa, exhibits a serrated shape and 40–110 µm thick, the lower layer, the Nostoc layer, is discrete, 20 or 70 µm thick; Lower cortex: lacking; Lower surface: bright orange, tomentose with a reticulate pattern of brown veins, with rhizoidal structures over 3 mm long, and internal cephalodia; Apothecia: large, rounded, irregularly scattered, not or rarely depressed in the upper surface, flat or convex, thalline margin absent; Asci: clavate, Peltigera-type, 8-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall surface ornamented with rounded papillae, measuring 35–45 × 10–12 µm.
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Figure 3. Solorina crocea ((A): Bruce McCune 18613 KUN, (B–G): Theodoer Esslinger 11831 KUN); (A), foliose thallus when dry; (B), smooth margin of the lobe; (C), upper surface of the apothecium; (D), bright orange lower surface distinctly brownish veined; (E), transversal sections of the thallus; (F), transversal sections of the apothecium; (G), ascus. Scale bars: 10 mm (A); 5 mm (B); 3 mm (C,D); 100 µm (E); 400 µm (F); 40 µm (G).
Figure 3. Solorina crocea ((A): Bruce McCune 18613 KUN, (B–G): Theodoer Esslinger 11831 KUN); (A), foliose thallus when dry; (B), smooth margin of the lobe; (C), upper surface of the apothecium; (D), bright orange lower surface distinctly brownish veined; (E), transversal sections of the thallus; (F), transversal sections of the apothecium; (G), ascus. Scale bars: 10 mm (A); 5 mm (B); 3 mm (C,D); 100 µm (E); 400 µm (F); 40 µm (G).
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Chemistry: Solorinic acid, averantin, 6-O-methylaverantin and gyrophoric acid (detected by TLC and HPLC).
Ecology and distribution: Distributed in high-altitude areas and mainly grows on calcareous soil. Its widespread distribution includes: Alps [42], America [43], Canada [44], Finland [45], France [46], Germany [47], Greenland [48], Iberian Peninsula and Balearic Islands [49], Iceland [50], Ireland [51], Italy [52], Japan [53], New Zealand [54], Norway [55], Poland [56], Russia [57], Sweden [58], the Czech Republic [59], Ukraine [60].
Note: S. crocea is an easily recognizable species of Solorina due to the bright orange lower surface and the two layers of photobionts. This unique thallus structure serves as an identification characteristic. This species has a wide distribution and mainly occurs in the northern hemisphere.
Specimens examined: USA, Wyoming, Park Co., Alpine tundra, Beartooth Plateau North of West Summit of Beartooth Highway, Alt. 3150 m, July 1992, 44°59′ N, 109°26′ W, 1992, B. McCune, 18613; Whatcom Co.: vicinity of the Ross Lake overlook on N base of Ruby Mt.; ca. Alt. 2200 m in a Douglas Fir-lodgepole pine forest, in soil, 1980, Leg. Theodoer and L. Esslinger, 11831.
Solorina crocoides (Nyl.) Gyeln., Magy. Bot. Lapok, 29: 30 (1930) Figure 4
≡ Solorinina crocoides Nyl. Naturaliste, 6. Année: 387 (1884)
= Solorina crocea (L.) Ach. Lichenology, 4. 1–6 (2005)
Type: Himalaya, 3657 m (no date, no collector recorded), H-1662
Description: Thallus: large foliose, wide-spreading, dorsiventral, heteromerous, lobate, 2–8 cm in diam.; Lobes: rounded, margins raised, 5–15 mm wide, usually contiguous to adjacent or slightly overlapping; Upper surface: green-gray when wet, gray-brown when dry, smooth to scabrid, with soredia growing on the margin; Soredia: black, granulose; Upper cortex: paraplectenchymatous, 25–200 µm thick, colorless; Medulla: composed of variably orientated linear hyphae; Photobiont: a single photobiont layer c. 100–200 µm thick, mixed chlorophyte and cyanobacteria cells, contains Asterochloris and Nostoc; Lower cortex: lacks distinct lower cortex; Lower surface: bright orange, distinctly brownish veined, tomentose, and with clusters of simple or branched rhizines; Apothecia: not seen.
Chemistry: Solorinic acid, averantin, 6-O-methylaverantin (detected by TLC and HPLC).
Ecology and distribution: This species is typically found in high-altitude areas, primarily inhabiting calcareous soils on rocky substrates. It often coexists with herbaceous plants and bryophytes, with the absence of tall trees in the vicinity, resulting in favorable lighting conditions. It was reported from the Himalayan region [11] and China (Sichuan and Xizang).
Note: Solorina crocoides is similar to S. crocea. A distinguishing feature between these two species is that the upper surface of S. crocea appears olive green when wet, whereas S. crocoides is gray-brown. The thallus of S. crocea possesses smoother edges, and the thallus margin of S. crocoides has black soredia. Another difference lies in their photobiont layers—S. crocea has two separate photobiont layers, whereas S. crocoides features only one photobiont layer containing both chlorophytes and cyanobacteria.
Specimens examined: China, Xizang Prov., Linzhi city, Bomi Co., 29°46′16.93″ N, 95°41.’57.39″ E, Alt. 3903 m, 9 August 2023, T. Zheng, 23-75455, 23-75456, 23-75432, 23-75431, 23-75453, 23-75450; 29°45′57.53″ N, 95°41′50.64″ E, Alt. 3798 m, 9 August 2023, T. Zheng, 23-75433, 23-75434, 23-75437; 29°46′16.93″ N, 95°41′57.39″ E, Alt. 3839 m, 9 August 2023, T. Zheng, 23-75449; 29°45.630′ N, 95°42.052′ E, Alt. 4065 m, 19 September 2014, L. S. Wang et al., 14-45996. Sichuan Prov., Luding Co. Gongga Mt., 29°20′ N, 101°30′ E, Alt. 3000 m, on rock, 1996, L. S. Wang, 96-16177.
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Figure 4. Solorina crocoides (T. Zheng 23-75450 KUN); (A), habit of S. crocoides; (B), margin of the lobe with black soredia; (C), bright orange lower surface distinctly brownish veined; (D), transversal sections of the thallus showing photobiont layer with both bright green chlorophyte cells and blue cyanobacteria cells. Scale bars: 2 mm (B); 5 mm (C); 200 µm (D).
Figure 4. Solorina crocoides (T. Zheng 23-75450 KUN); (A), habit of S. crocoides; (B), margin of the lobe with black soredia; (C), bright orange lower surface distinctly brownish veined; (D), transversal sections of the thallus showing photobiont layer with both bright green chlorophyte cells and blue cyanobacteria cells. Scale bars: 2 mm (B); 5 mm (C); 200 µm (D).
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Pseudosolorina T. Zheng and Li S. Wang, gen. nov.
Fungal Names: FN 572274
Type species: Pseudosolorina parmigera T. Zheng and Li S. Wang, sp. nov.
Etymology: Referring to its similarity to Solorina.
Diagnosis: This genus has scattered round, immersed, slightly to very deeply concaved apothecia without a thalline margin. Thallus foliose, lower surface white or pale brown, indistinctly veined, it is similar to Solorina in external morphology, but differs by the white to pale brown lower surface.
Description: Thallus: small to large foliose, dorsiventral, heteromerous, lobate; Lobes: rounded, margins flush with the substrate or slightly raised; Upper surface: apple green to green-gray when wet, white or pale-green to brown when dry, smooth to scabrid; Upper cortex: paraplectenchymatous; Photobiont: Coccomyxa and Nostoc; Lower cortex: lacking; Lower surface: white or pale brown, indistinctly veined, tomentose, and with clusters of simple or branched rhizines; Apothecia: rounded, irregularly scattered, impressed to immersed in the upper surface, red-brown to black, slightly to very deeply concave, thalline margin absent; Asci: clavate, Peltigera-type, 1-, 2- or 4-spored; Ascospores: red-brown or dark brown, 1- or 2-(3-)septate with median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented broad reticulating ridges and irregular lacunae.
Chemistry: Two species contain gyrophoric acid, methyl gyrophorate, tenuiorin, and 2′-O-methyltenuiorin, but six species lack secondary metabolites (detected by TLC and HPLC).
Ecology: On soil, markedly calcicole. The members of this genus often occur in alpine habitats.
Note: Previously, species of this genus were classified under Solorina based on the morphology of the apothecia. However, phylogenetic analyses indicate strong support for its status as a monophyletic genus. Pseudosolorina is set apart from related genera by its white or pale brownish lower surface, with apothecia irregularly scattered and immersed in the upper surface of the thallus, usually concave. In comparison to Solorina, its spore wall is rougher and ornamented with broad reticulating ridges and irregular lacunae. Most species lack secondary metabolites.
Pseudosolorina bispora (Nyl.) T. Zheng and Li S. Wang, comb. nov. Figure 5
≡ Solorina bispora Nyl., Syn. meth. lich. (Parisiis) 1(2): 331 (1860)
= Solorina saccata var. bispora (Nyl.) Arnold, Verh. Kaiserl.-Königl. Zool.-Bot. Ges. Wien, 21(3–4): 1118 (1871)
= Solorina bispora Nyl., Magy. Bot. Lapok, 29: 29 (1930)
Fungal Names: FN 572279
Type: France, Midi-Pyrénées, Hautes-Pyrénées, Nylander W., H9506085 (Holotype)
Description: Thallus: small foliose, fragile, dorsiventral, heteromerous, lobate, 1.5–2 cm in diam.; Lobes: rounded or irregular, 5–10 mm; Upper surface: bright apple-green when wet, dark green or pale grey to tinged brown when dry, usually covered with white-pruinose; Upper cortex: paraplectenchymatous, colorless, 50–100 μm thick; Medulla: 40–150 μm thick; Photobiont: Coccomyxa and Nostoc, one Coccomyxa green algal layer, 40–110 μm thick, Nostoc present in internal cephalodia which can be seen on the underside of the thallus as dark spots; Lower cortex: lacking; Lower surface: pale brown, densely tomentose, not or indistinctly veined, rhizines; Apothecia: large, rounded, irregularly scattered, sunk in depressions in the upper surface, red-brown to black, deeply concave, thalline margin absent; Asci: clavate, Peltigera-type, 2-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented broad reticulating ridges and irregular lacunae, 65–88 × 33–42 µm.
Chemistry: No lichen products detected by TLC.
Ecology and distribution: Typically found in humid soil rich in humus, often growing in association with herbaceous plants. Distributed in Alps [42], America [61], Canada [44], France [6,46], Germany [47], Greenland [62], Iberian Peninsula and Balearic Islands [49], Italy [52], New Mexico [63], Norway [64], Poland [56], Ukraine [60], and China.
Note: This species resembles Pseudosolorina saccata and was previously classified as a high-altitude variant of P. saccata. In comparison to P. saccata, its thallus is relatively small (<2 cm in diam.), often exhibiting a covering of pruina; it has two ascospores.
Specimens examined: Austria, Salzburg, Tamsweg, Grossek-Speiereck, 47°7′54″ N, 13°38′17″ E, Alt. 2162 m, 31 August 2019, Y. Y. Zhang, ZYY-46; China, Sichuan Prov., Hongyuan city, 32°13′45.83″ N, 102°35′44.75″ E, Alt. 4278 m, 9 April 2020, L. S. Wang et al., 20-66618; Yunnan Prov., Kunming city, Tangdan town, Yaojingtang, 26°15′27″ N, 102°95′79″ E, Alt. 4000 m, 15 September 2024, M. Ai, 24-76716.
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Figure 5. Pseudosolorina bispora ((A): L. S. Wang et al. 20-66618 KUN, (B): M. Ai 24-76716 KUN, (C–F): Y. Y. Zhang ZYY-46 KUN); (A), habit of P. bispora; (B), apothecium; (C), white lower surface with brownish tomentum; (D), transversal sections of the thallus; (E), transversal sections of the apothecium; (F), ascus. Scale bars: 2 mm (B,C); 100 µm (D); 300 µm (E); 40 µm (F).
Figure 5. Pseudosolorina bispora ((A): L. S. Wang et al. 20-66618 KUN, (B): M. Ai 24-76716 KUN, (C–F): Y. Y. Zhang ZYY-46 KUN); (A), habit of P. bispora; (B), apothecium; (C), white lower surface with brownish tomentum; (D), transversal sections of the thallus; (E), transversal sections of the apothecium; (F), ascus. Scale bars: 2 mm (B,C); 100 µm (D); 300 µm (E); 40 µm (F).
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Pseudosolorina bispora var. monospora (Gyeln.) T. Zheng and Li S. Wang, comb. nov. Figure 6
≡ Solorina monospora Gyeln., Magy. Bot. Lapok, 29: 29 (1930)
= Solorina bispora var. monospora (Gyeln.) Frey, Ergebn. wiss. Unters. schweiz. NatnParks, N.S. 3(27): 377 (1952).
Fungal Names: FN 572281
Description: Thallus: small foliose, fragile, dorsiventral, heteromerous, lobate, ca 1 cm in diam.; Lobes: rounded or irregular, 2–8 mm; Upper surface: bright apple-green when wet, dark green or pale grey to tinged brown when dry, usually covered with white pruina, pale grey to brown-grey, often pruinose; Upper cortex: paraplectenchymatous, colorless, 40–80 μm thick; Medulla: 40–150 μm thick; Photobiont: Coccomyxa and Nostoc, one Coccomyxa green-algal layer, c. 20–80 μm thick, Nostoc exist in internal cephalodia which can be seen on the underside of the thallus as dark spots; Lower cortex: lacking; Lower surface: pale brown, densely tomentose, not or indistinctly veined, rhizines, cephalodia internal, rarely external; Apothecia: large, rounded, irregularly scattered, sunk in depressions in the upper surface, red-brown to black, deeply concave, thalline margin absent; Asci: clavate, Peltigera-type, 1-spored; Ascospores: red-brown or dark brown, 2- (3-) septate with median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented with broad reticulating ridges and irregular lacunae, 100–160 × 35–50 µm.
Chemistry: No lichen products detected by TLC.
Ecology and distribution: This species is typically found in alpine environments, where it grows on basic soils formed over shaded limestone or calcareous schists. It is distributed in France [46], Iberian Peninsula and Balearic Islands [49], Switzerland [65], and China.
Note: This species closely resembles a variant of P. bispora (= Solorina bispora var. monospora (Gyeln.) Frey) in appearance, containing only 1 spore in the ascus. However, phylogenetic analysis suggests that it should be recognized as an independent species. Unfortunately, we did not obtain the type specimen or molecular data related to the origin of S. bispora var. monospora, so we can only describe this specimen as a similar species to S. bispora var. monospora, revised to Pseudosolorina bispora var. monospora (Gyeln.) T. Zheng and Li S. Wang. Further research is necessary to determine whether they are indeed the same species.
Specimens examined: China, Qinghai Prov., Zhiduo Co., 33°48′03.32″ N, 95°13′57.24″ E, Alt. 4436 m, 18 September 2020, X. Y. Wang et al., XY20-828; Maduo Co., 35°06′16.62″ N, 98°54′22.97″ E, Alt. 4260 m, 14 September 2020, X. Y. Wang et al., XY20-2943; Maduo Co., 34°49′40.56″ N, 99°02′03.48″ E, Alt. 4578 m, 13 September 2020, L. S. Wang et al., 20-67082.
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Figure 6. Pseudosolorina bispora var. monospora (L. S. Wang et al. 20-67082 KUN); (A), habit of P. bispora var. monospora; (B), apothecium; (C), white lower surface with brownish tomentum; (D), transversal sections of the thallus; (E), transversal sections of the apothecium; (F), ascus. Scale bars: 2 mm (B); 3 mm (C); 100 µm (D); 200 µm (E); 40 µm (F).
Figure 6. Pseudosolorina bispora var. monospora (L. S. Wang et al. 20-67082 KUN); (A), habit of P. bispora var. monospora; (B), apothecium; (C), white lower surface with brownish tomentum; (D), transversal sections of the thallus; (E), transversal sections of the apothecium; (F), ascus. Scale bars: 2 mm (B); 3 mm (C); 100 µm (D); 200 µm (E); 40 µm (F).
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Pseudosolorina hepatizon T. Zheng and Li S. Wang, sp. nov. Figure 7
Fungal Names: FN 572275
Type: China. Yunnan Prov., Shangrila city, Birong Valley, 26°39′05.72″ N, 99°44′15.57″ E, Alt. 3089 m, 15 July 2023, L. S. Wang et al., 23-75286 (Holotype KUN)
Etymology: Referring to its hepaticolor thallus.
Diagnosis: This species has a hepaticolor upper surface when dry, while most other species of Pseudosolorina display brown, greyish-brown, or pale hues. P. hepatizon has a single photobiont layer containing both cyanobacteria and chlorophytes and lacks secondary metabolites.
Description: Thallus: large foliose, dorsiventral, heteromerous, lobate, up to 9 cm in diam.; Lobes: rounded, 10–20 mm; Upper surface: green-gray when wet, hepaticolor or gray-brown when dry, smooth to scabrid; Upper cortex: paraplectenchymatous, colorless, 60–150 μm thick; Medulla: 200–300 μm thick; Photobiont: Asterochloris and Nostoc, single photobiont layer with both blue and bright green cells, 100–220 μm thick; Lower cortex: lacking; Lower surface: white, indistinctly veined, brownish tomentose, without cephalodia, with clusters of simple or branched rhizines; Apothecia: large, rounded, irregularly scattered, sunk in depressions in the upper surface, red-brown to black, slightly concave, thalline margin absent; Asci: clavate, Peltigera-type, 4-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented broad reticulating ridges and irregular lacunae, measured 42–53 × 18–25 µm.
Chemistry: No lichen products detected by TLC.
Ecology and distribution: It is distributed in high-altitude areas and primarily found on calcareous soils on stones, typically growing alongside or on bryophytes. This species is only known from Yunnan Province of China.
Note: Specimens representing this species were formerly categorized as Solorina simensis based on their morphological characteristics. In this study they have been divided from P. simensis to form the new species P. hepatizon. Despite distinct chemical characteristics, P. simensis contains secondary metabolites such as methyl gyrophorate and tenuiorin, whereas P. hepatizon lacks any secondary metabolites; it was previously identified as an acid-deficient variant of S. simensis (Krog and Swinscow 1986). DNA evidence supports its classification as a distinct species in the realm of scientific research.
Specimens examined: China, Yunnan Prov., Kunming city, Dongchuan Dist., Guniuzhai Mt., 26°09′50.25″ N, 103°13′30.86″ E, Alt. 3100 m, 10 May 2017, L. S. Wang et al., 17-55094; Shangrila city, Birong Valley, 26°39′05.72″ N, 99°44′15.57″ E, Alt. 3089 m, 15 July 2023, L. S. Wang et al., 23-75286.
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Figure 7. Pseudosolorina hepatizon (L. S. Wang et al. 23-75286 KUN); (A), habit of P. hepatizon; (B), foliose thallus when dry; (C), white lower surface with brownish tomentum and rhizines; (D), transversal sections of the thallus; (E), transversal sections of the apothecium; (F), ascus. Scale bars: 10 mm (B); 2 mm (C); 50 µm (D); 100 µm (E); 20 µm (F).
Figure 7. Pseudosolorina hepatizon (L. S. Wang et al. 23-75286 KUN); (A), habit of P. hepatizon; (B), foliose thallus when dry; (C), white lower surface with brownish tomentum and rhizines; (D), transversal sections of the thallus; (E), transversal sections of the apothecium; (F), ascus. Scale bars: 10 mm (B); 2 mm (C); 50 µm (D); 100 µm (E); 20 µm (F).
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Pseudosolorina parmigera T. Zheng and Li S. Wang, sp. nov. Figure 8
Fungal Names: FN 572277
Type: China, Yunnan Prov., Shangrila city, road side of x236, 26°38′01.25″ N, 99°43′07.92″ E, Alt. 3035 m, 15 July 2023, L. S. Wang et al., 23-75751 (Holotype KUN)
Etymology: Referring to its shield-like shaped thallus.
Diagnosis: This species has a shield-like shape of the thallus and one green photobiont layer similar to P. saccata, but P. parmigera has extensive white pruina and no external cephalodia on the upper surface, whereas P. saccata has external cephalodia and rarely white pruina on the upper surface.
Description: Thallus: large foliose, fragile, dorsiventral, heteromerous, lobate, 2–6 cm in diam.; Lobes: rounded, 5–15 mm; Upper surface: bright apple-green when wet, dark green to pale grey to tinged brown when dry, white-pruinose, smooth to scabrid; Upper cortex: paraplectenchymatous, colorless, 30–80 μm thick; Medulla: 100–250 μm thick; Photobiont: Coccomyxa and Nostoc, single Coccomyxa green-algal layer, c. 40–110 μm thick, Nostoc present in internal cephalodia which can be seen on the underside of the thallus as dark spots; Lower cortex: lacking; Lower surface: pale brown, densely tomentose, not or indistinctly veined, rhizines abundant, length: 10 mm; Apothecia: large, rounded, irregularly scattered, sunk in depressions in the upper surface, red-brown to black, medially concave, thalline margin absent; Asci: clavate, Peltigera-type, 4-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented broad reticulating ridges and irregular lacunae, dimensions: 35–50 × 17–23 µm.
Chemistry: No lichen products detected by TLC.
Ecology and distribution: Growing in calciferous soil rich in humus and terricolous mosses at approximately 3000 m in the alpine zone. This species is known from Yunnan, Sichuan, and Qinghai Provinces of China.
Note: This species shares the most similar morphological and chemical characteristics with P. saccata; the difference between these two species is that P. saccata has external cephalodia and is rarely pruinose on the upper surface, but the thallus of P. parmigera has dense pruina and lacks external cephalodia on the upper surface.
Specimens examined: China, Yunnan Prov., Lijiang city, Yulong Snow Mt. 27°07′36.82″ N, 100°13′40.70″ E, Alt. 2938 m, 8 July 2019, L. S. Wang et al., 19-62636; Shangrila city, road side of x224, 26°38′07.56″ N, 99°43′00.12″ E, Alt. 3062 m, 15 July 2023, L. S. Wang et al., 23-75739, 23-75734; Shangrila city, road side of x236, 26°38′01.25″ N, 99°43′07.92″ E, Alt. 3035 m, 15 July 2023, L. S. Wang et al., 23-75751; Dali city, Cangshan National Geopark, 25°41′42.47″ N, 100°06′30.73″ E, Alt. 3075 m, 19 May 2024, L. S. Wang and M. Ai, 24-76096. Sichuan Prov., Liangshan Yi Autonomous Prefecture Muli Co. 27°41′51.41″ N, 101°13′52.86″ E, Alt. 3177 m, 10 September 2021, X. Y. Wang et al., 21-70363, 21-70423, XY21-272, XY21-275. Qinghai Prov., Menyuan Co., 37°12′34″ N, 102°0′39″ E, Alt. 2711 m, 24 July 2022, S. B. Zhang et al., ZSB22-133; Manma Co., 32°46′50″ N, 101°11′2″ E, Alt. 3553 m, 14 July 2022, S. B. Zhang et al., ZSB22-256.
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Figure 8. Pseudosolorina parmigera (A: L. S. Wang et al. 23-75751 KUN, B,C: L. S. Wang and M. Ai 24-76096 KUN, (D–G) X. Y. Wang et al. 21-70423 KUN); (A), habit of P. parmigera; (B), upper surface with white pruina; (C), apothecium; (D), white lower surface with brownish tomentum and rhizines; (E), transversal sections of the thallus; (F), transversal sections of the apothecium; (G), ascus. Scale bars: 2 mm (B,D); 3 mm (C); 200 µm (E,F); 40 µm (G).
Figure 8. Pseudosolorina parmigera (A: L. S. Wang et al. 23-75751 KUN, B,C: L. S. Wang and M. Ai 24-76096 KUN, (D–G) X. Y. Wang et al. 21-70423 KUN); (A), habit of P. parmigera; (B), upper surface with white pruina; (C), apothecium; (D), white lower surface with brownish tomentum and rhizines; (E), transversal sections of the thallus; (F), transversal sections of the apothecium; (G), ascus. Scale bars: 2 mm (B,D); 3 mm (C); 200 µm (E,F); 40 µm (G).
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Pseudosolorina saccata (L.) T. Zheng and Li S. Wang, comb. nov. Figure 9
≡ Lichen saccatus L., Fl. Suec., Edn 2: 419 (1755)
= Lobaria saccata (L.) Hoffm., Deutschl. Fl., Zweiter Theil (Erlangen): 147 (1796)
= Peltidea saccata (L.) Ach., Methodus, Sectio post. (Stockholmiæ): 290 (1803)
= Peltigera saccata (L.) DC., in Lamarck and de Candolle, Fl. franç., Edn 3 (Paris) 2: 408 (1805)
= Arthonia saccata (L.) Ach., Neues J. Bot. 1(3. Stück): 21 (1806)
= Solorina saccata (L.) Ach., K. Vetensk-Acad. Nya Handl. 29: 228 (1808)
= Platysma saccatum (L.) Frege, Deutsch. Botan. Taschenb. 2: 165 (1812)
= Solorina saccata (L.) Ach., Magy. Bot. Lapok, 29: 29 (1930)
Fungal Names: FN 572280
Type: LINN-HL1273-197 (Lectotype, designated by Jørgensen et al., Bot. J. Linn. Soc. 115: 381. 1994)
Description: Thallus: large foliose, fragile, dorsiventral, heteromerous, lobate, 3–6 cm in diam.; Lobes: rounded, 4–20 mm; Upper surface: bright apple-green when wet, dark green to pale grey to tinged brown when dry, white-pruinose, having external cephalodia; Upper cortex: paraplectenchymatous, colorless, 50–100 μm thick; Medulla: 200–350 μm thick; Photobiont: Coccomyxa and Nostoc, single Coccomyxa green-algal layer, c. 50–110 μm thick, Nostoc present in external cephalodia on the upper surface and internal cephalodia which can be seen on the underside of the thallus as dark spots; Lower cortex: lacking; Lower surface: pale brown, densely tomentose, not or indistinctly veined, rhizines abundant and long, up to 18 mm; Apothecia: large, rounded, irregularly scattered, sunk in depressions in the upper surface, red-brown to black, medially concave, thalline margin absent; Asci: clavate, Peltigera-type, 4-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented with broad reticulating ridges and irregular lacunae, dimensions: 32–60 × 18–27 µm.
Chemistry: No lichen products detected by TLC.
Ecology and distribution: This species grows in calcareous soil rich in humus and terrestrial mosses in a cool-temperate alpine region, often inhabiting rock crevices and also commonly found along the edges of groves. Its widespread distribution includes: Alps [52], USA [66], Canada [44], England [67], France [46], Germany [47], Greece [68], Iberian Peninsula and Balearic Islands [49], India [69], Italy [52], Japan [53], Norway [64], Poland [56], Siberia [70], the Czech Republic [59], Ukraine [60] and China.
Note: Pseudosolorina saccata is widely distributed and morphologically variable. P. simensis and P. bispora were previously classified as variants under this species. P. saccata is characterized by a green upper surface, a white lower surface, a concave apothecium, and 4-spored ascus. Notably, it closely resembles P. parmigera; however, distinct attributes for P. saccata include external cephalodia and rare white pruina on the upper surface of the thallus, whereas P. parmigera typically exhibits a white pruinose covering on its upper surface and lacks external cephalodia.
Specimens examined: China, Yunnan Prov., Shangrila city, Birong Valley, 26°39′04.17″ N, 99°44′13.48″ E, Alt. 3082 m, 15 July 2023, L. S. Wang et al., 23-75282; road side of X241, 26°37′54.86″ N, 99°43′14.07″ E, Alt. 3051 m, 15 July 2023, L. S. Wang et al., 23-75756, 23-75757, 23-75755; Birong Valley, 26°39′03.98″ N, 99°44′13.12″ E, Alt. 3486 m, 15 July 2023, X. Y. Wang and Y. X. Gan, XY23-383; Birong Valley, 26°39′04.26″ N, 99°44′13.77″ E, Alt. 3047 m, 15 July 2023, L. S. Wang et al., 23-75281(a), 23-75280; Dali city, Cangshan National Geopark, 25°41′41.09″ N, 100°06′32.38″ E, Alt. 3070 m, 19 May 2024, L. S. Wang et al., 24-76098 (b). Qinghai Prov., Xunhua Co., 35°41′48″ N, 102°26′26.18″ E, Alt. 2688 m, 6 August 2022, S. B. Zhang and H. L. Kang, ZSB22-67; Banma Co., 32°47′15″ N, 101°4′45″ E, Alt. 3680 m, 14 July 2022, S. B. Zhang and H. L. Kang, ZSB22-263; Huzhu Co., 36°44′41″ N, 102°32′47″ E, Alt. 2759 m, 16 August 2022, S. B. Zhang and H. L. Kang, ZSB22-340. Gansu Prov., Luqu Co., 34°08′27.58″ N, 102°11′57.16″ E, Alt. 3588 m, 9 July 2022, L. S. Wang et al., 22-73302, 22-73303. Sichuan Prov., Jiulong Co., Wuxuhai Lake, 29°09′06.14″ N, 101°24′42.58″ E, Alt. 3691 m, 9 June 2023, L. S. Wang et al., 23-75120.
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Figure 9. Pseudosolorina saccata ((A): X. Y. Wang and YX Gan XY23-383 KUN, (B): L. S. Wang et al. 23-75281 KUN, C–G: L. S. Wang et al. 23-75757 KUN); (A), habit of P. saccata; (B), upper surface with external cephalodia; (C), white lower surface with rhizines; (D), apothecium; (E), transversal sections of the thallus; (F), transversal sections of the apothecium; (G), ascus. Scale bars: 1 mm (B); 4 mm (C); 3 mm (D); 100 µm (E); 200 µm (F); 40 µm (G).
Figure 9. Pseudosolorina saccata ((A): X. Y. Wang and YX Gan XY23-383 KUN, (B): L. S. Wang et al. 23-75281 KUN, C–G: L. S. Wang et al. 23-75757 KUN); (A), habit of P. saccata; (B), upper surface with external cephalodia; (C), white lower surface with rhizines; (D), apothecium; (E), transversal sections of the thallus; (F), transversal sections of the apothecium; (G), ascus. Scale bars: 1 mm (B); 4 mm (C); 3 mm (D); 100 µm (E); 200 µm (F); 40 µm (G).
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Pseudosolorina simensis (Hochst. ex Flot.) T. Zheng and Li S. Wang, comb. nov. Figure 10
≡ Solorina simensis Hochst. ex Flot., Linnaea 17: 17 (1843)
= Solorina saccata var. simensis (Hochst. ex Flot.) Nyl., Mém. Soc. Imp. Scity Nat. Cherbourg, 5: 101 (1858)
= Solorinina simensis (Hochst. ex Flot.) Nyl., Naturaliste, 6. Année: 387 (1884)
= Solorina simensis Hochst. ex Flot., Magy. Bot. Lapok, 29: 29 (1930)
Fungal Names: FN 572282
Type: Ethiopia, Amhara Region, Simien Mountains, W. Schimper, H-NYL 32,912 (Holotype H-NYL)
Description: Thallus: large foliose, fragile, flat, dorsiventral, heteromerous, lobate, ca 5 cm in diam.; Lobes: rounded, 5–8 mm; Upper surface: light blue when wet, gloomy to brown when dry, rarely pruinose on the margin; Upper cortex: paraplectenchymatous, colorless, 40–80 μm thick; Medulla: pale, 200–280 μm thick; Photobiont: Asterochloris and Nostoc, both present in a single photobiont layer, 70–150 μm thick in thallus and 60–100 µm in apothecium; Lower cortex: lacking; Lower surface: pale to brown, indistinctly veined, tomentose, without cephalodia, with clusters of simple or branched rhizines; Apothecia: large, rounded, irregularly scattered, sunk in depressions in the upper surface, red-brown to black, slightly concave, thalline margin absent; Asci: clavate, Peltigera-type, 4-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented with broad reticulating ridges and irregular lacunae, dimensions: 35–65 × 16–23 µm.
Chemistry: Gyrophoric acid, methyl gyrophorate, tenuiorin, and 2′-O-methyltenuiorin (detected by TLC and HPLC).
Ecology and distribution: It grows in high-altitude areas and is mainly found in calcareous soil on stone, usually growing together with or on bryophytes. It is distributed in East Africa [71], Ethiopia [5], Mexico [72], New Guinea [73], and China.
Note: This species could be distinguished by its morphological characters of the thallus and the single photobiont layer with both cyanobacterial and chlorophyte cells. It is similar to P. tenuior, but P. simensis has a thicker photobiont layer up to 150 μm in thallus and 100 µm in apothecium.
Specimens examined: Ethiopia, Amhara Region, Simien Mountains, W. Schimper, H-NYL 32,912 (holotype); H-NYL 32914. China, Sichuan Prov., Liangshan Yi Autonomous Prefecture, Muli Co., 27°43′30.82″ N, 101°14′11.52″ E, Alt. 3010 m, 11 September 2021, X. Y. Wang et al., XY21-274, 21-70422; Litang Co., 30°25′30.27″ N, 101°18′07.81″ E, Alt. 3296 m, 12 July 2022, D. Liu, LD22-696, LD22-697; Yunnan Prov., Shangrila city, road side of X246, 26°37′56.13″ N, 99°43′31.28″ E, Alt. 3110 m, 15 July 2023, L. S. Wang et al., 23-75761.
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Figure 10. Pseudosolorina simensis (A,C: X. Y. Wang et al. XY21-274 KUN, B: W. Schimper 32,914 H-NYL, (D–G): X. Y. Wang et al. 21-70422 KUN); (A), habit of P. simensis; (B), type of P. simensis; (C), foliose thallus when dry; (D), apothecium; (E), white lower surface with rhizines; (F), transversal sections of the thallus; (G), transversal sections of the apothecium; (H), ascus. Scale bars: 5 mm (B,C); 2 mm (D,E); 50 µm (F); 200 µm (G); 30 µm (H).
Figure 10. Pseudosolorina simensis (A,C: X. Y. Wang et al. XY21-274 KUN, B: W. Schimper 32,914 H-NYL, (D–G): X. Y. Wang et al. 21-70422 KUN); (A), habit of P. simensis; (B), type of P. simensis; (C), foliose thallus when dry; (D), apothecium; (E), white lower surface with rhizines; (F), transversal sections of the thallus; (G), transversal sections of the apothecium; (H), ascus. Scale bars: 5 mm (B,C); 2 mm (D,E); 50 µm (F); 200 µm (G); 30 µm (H).
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Pseudosolorina spongiosa (Sm.) T. Zheng and Li S. Wang, comb. nov. Figure 11
≡ Lichen spongiosus Sm., in Smith and Sowerby, Engl. Bot. 20: tab. 1374 (1805)
= Collema spongiosum Ach., Lich. Univ.: 661 (1810)
= Lichen furvus * spongiosum (Ach.) Lam., Encycl. Méth. Bot., Suppl. (Paris) 3(2): 416 (1813)
= Polychidium spongiosum (Ach.) Gray, Nat. Arr. Brit. Pl. (London) 1: 402 (1821)
= Lecanora limbata Sommerf., Suppl. Fl. lapp. (Oslo): 105 (1826)
= Parmelia spongiosa (Ach.) Spreng., Syst. veg., Edn 16 4(1): 277 (1827)
= Peltigera saccata var. limbata (Sommerf.) Fr., Summa veg. Scand., Sectio Prior (Stockholm): 104 (1845)
= Solorina saccata ß limbata (Sommerf.) Schaer., Enum. critic. lich. europ. (Bern): 23 (1850)
= Peltigera limbata (Sommerf.) Nyl., Not. Sällsk. Fauna et Fl. Fenn. Förh. 2: 217 (1852)
= Solorina saccata f. limbata (Sommerf.) Nyl., Mém. Soc. Imp. Sci. Nat. Cherbourg 3: 173 (1855)
= Solorina saccata var. spongiosa (Ach.) Nyl., Syn. meth. lich. (Parisiis) 1(2): 331 (1860)
= Solorina limbata (Sommerf.) Mudd, Man. Brit. Lich.: 85 (1861)
= Solorina spongiosa (Sm.) Anzi, Comm. Soc. crittog. Ital. 1(fasc. 3): 136 (1862)
= Solorinina simensis var. limbata (Sommerf.) Nyl., in Hue, Nouv. Arch. Mus. Hist. Nat., Paris, 3 sér. 2: 312 (1890)
= Solorina spongiosa (Sm.) Anzi, Magy. Bot. Lapok, 29: 29 (1930)
Fungal Names: FN 572276
Type: BM000975901 (type, collected by Anon. on 27 April 1803)
Diagnosis: This species has a reduced thallus, often with white pruina; this morphological character is shared with Solorina embelina, but P. spongiosa has four spores per ascus, whereas Solorina embelina only has one.
Description: Thallus: foliose, fragile, dorsiventral, heteromerous, poorly developed, restricted to a continuous or lacerate narrow rim surrounding the apothecium, 0.1–0.7 cm in diam.; Upper surface: bright green when wet, dark green to pale grey when dry, usually covered with white pruina; Upper cortex: paraplectenchymatous, colorless, 10–20 μm thick; Medulla: 40–80 μm thick; Photobiont: Coccomyxa and Nostoc, one Coccomyxa green-algal layer, 20–60 μm thick, Nostoc present in internal cephalodia which can be seen on the underside of the thallus as dark spots; Lower cortex: lacking; Lower surface: pale brown, densely tomentose, not or indistinctly veined, rhizines; Apothecia: rounded, irregularly scattered, sunk in depressions in the upper surface, red-brown to black, medially concave, thalline margin absent; Asci: clavate, Peltigera-type, 4-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented broad reticulating ridges and irregular lacunae, measured 30–50 × 18–23 µm.
Chemistry: No lichen products detected by TLC.
Ecology and distribution: This species typically thrives in moist calcareous soils within the alpine zone and is commonly found beneath the edges of groves. It is distributed in America [43], Austria [74], Canada [44], France [46], Germany [47], Greenland [48], Iberian Peninsula and Balearic Islands [49], Ireland [51], Poland [56], Russia [57], the Czech Republic [59], Ukraine [60].
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Figure 11. Pseudosolorina spongiosa, (A: L. S. Wang et al. 24-76099 KUN, B–E: L. S. Wang et al. 23-75119 KUN); (A), habit of P. spongiosa; (B), apothecium; (C), white lower surface with brownish tomentum; (D), transversal sections of the apothecium; (E), ascus. Scale bars: 2 mm (B); 1 mm (C); 200 µm (D); 40 µm (E).
Figure 11. Pseudosolorina spongiosa, (A: L. S. Wang et al. 24-76099 KUN, B–E: L. S. Wang et al. 23-75119 KUN); (A), habit of P. spongiosa; (B), apothecium; (C), white lower surface with brownish tomentum; (D), transversal sections of the apothecium; (E), ascus. Scale bars: 2 mm (B); 1 mm (C); 200 µm (D); 40 µm (E).
Jof 11 00169 g011
Note: This species exhibits morphological similarities with Solorina embelina, particularly in their thallus structure, where reduced thalli form a distinct small circle around the apothecium. However, the two species differ in terms of spore number: S. embelina contains only one spore per ascus, whereas P. spongiosa possesses four spores per ascus. Through a meticulous review and comparison of the original literature on Lichen spongiosus, we have identified discrepancies between the initial documentation of this species and the present specimens. The original records describe the species as having a clustered and branched thallus; apothecia that are scattered, concave, brown, externally spongy, and pale, with a thin, upright margin. In contrast, current samples of P. spongiosa feature a small thallus, lacking lobes and apothecia that are scattered, concave, brown to black, without any margin. However, due to limited access to the holotype of Lichen spongiosus, we are unable to define this species. However, our samples are phylogenetically and morphologically identical to the currently accepted species Solorina spongiosa. Consequently, we have opted to retain the classification of this species as P. spongiosa.
Specimens examined: China, Yunnan Prov., Shangrila city, Birong Valley, 26°39′04.26″ N, 99°44′13.77″ E, Alt. 3047 m, 15 July 2023, L. S. Wang et al., 23-75281(b); Dali city, Cangshan Geopark, 25°41′42.09″ N, 100°06′29.97″ E, Alt. 3092 m, 19 May 2024, L. S. Wang et al., 24-76095, 24-76098 (a), 24-76097, 24-76099. Xizang Prov., Linzhi, Nyingchi Sexiula Mountain G318 Roadside, 28 June 2021, CMY-42. Sichuan Prov., Liangshan Yi Autonomous Prefecture, Muli Co., 27°43′30.93″ N, 101°14′11.34″ E, Alt. 3070 m, 11 September 2021, X. Y. Wang et al., 21-70424; Jiulong Co., Wuxuhai Lake, 29°09′06.00″ N, 101°24′42.99″ E, Alt. 3722 m, 9 June 2023, L. S. Wang et al., 23-75118, 23-75119, 23-75121, 23-75122; Heishui Co., 32°13′57.72″ N, 102°36′21.75″ E, Alt. 3956 m, 4 September 2020, X. Y. Wang et al., XY20-399. Gansu Prov., Luqu Co., 34°08′23.63″ N, 102°11′55.61″ E, Alt. 3584 m, 9 July 2022, X. Y. Wang et al., XY22-1164. Austria, Salzburg, Lungau Eastern Alps, Niedere Tauern, Schladminger Tauern, Weißpriachtal NW of the village Mariapfarr, a short distance from Lahnbrücke, 47°13′18″ N, 13°39′24″ E, Alt. 1280 m, 3 September 2019, Y. Y. Zhang, ZYY-126.
Pseudosolorina tenuior T. Zheng and Li S. Wang, sp. nov. Figure 12
Fungal Names: FN 572278
Type: China, Yunnan Prov., Shangrila city, Birong Valley, 26°39′05.36″ N, 99°44′15.57″ E, Alt. 3406 m, 15 July 2023, L. S. Wang et al., 23-75283 (Holotype KUN)
Etymology: Referring to its thin thallus and photobiont layer.
Diagnosis: This species contains a single photobiont layer with both cyanobacterial and chlorophyte cells. It has the same four secondary metabolites as P. simensis, but P. simensis has a thicker photobiont layer: more than 70 μm in thallus and more than 60 μm in apothecia. P. tenuior has a thinner photobiont layer: less than 60 μm in thallus and less than 30 μm in apothecia.
Description: Thallus: large foliose, fragile, dorsiventral, heteromerous, lobate, 3–7 cm in diam.; Lobes: rounded, 3–18 mm; Upper surface: green-gray when wet, gray-brown when dry, rare pruinose on the margin; Upper cortex: paraplectenchymatous, colorless, 30–80 μm thick; Medulla: 200–270 μm thick; Photobiont: Asterochloris and Nostoc, in a single photobiont layer, measured 20–60 μm, photobiont layer in apothecium measuring 12–30 µm; Lower cortex: lacking; Lower surface: pale to brown, indistinctly veined, tomentose, without cephalodia, with clusters of simple or branched rhizines; Apothecia: large, rounded, irregularly scattered, sunk in depressions in the upper surface, red-brown to black, slightly concave, thalline margin absent; Asci: clavate, Peltigera-type, 4-spored; Ascospores: red-brown or dark brown, 1-septate with a median constriction, ellipsoid to fusiform, wall non-uniformly thickened, surface ornamented with broad reticulating ridges and irregular lacunae, dimensions: 40–55 × 18–23 µm.
Chemistry: Gyrophoric acid, methyl gyrophorate, tenuiorin, and 2′-O-methyltenuiorin (detected by TLC and HPLC).
Ecology and distribution: It is distributed in high-altitude areas and is mainly found in calcareous soil on stone, usually growing together with or on bryophyte. This species is known from Yunnan and Sichuan Provinces of China.
Note: This species is similar to Pseudosolorina simensis in thallus morphology, appearance of photobiont layer, and chemical characteristics, but DNA sequence data support their description as separate species. They differ in the thickness of the photobiont layer both in thallus and in apothecium: the photobiont layer of P. tenuior is 20–60 μm in thallus and 12–30 µm in apothecium, but the photobiont layer of P. simensis is 70–150 μm in thallus and 60–100 µm in apothecium.
Specimens examined: China, Sichuan Prov., Liangshan Yi Autonomous Prefecture, Muli Co., 27°43′30.80″ N, 101°14′10.55″ E, Alt. 2993 m, 11 September 2021, X. Y. Wang et al., XY21-270; Yunnan Prov., Shangrila city, Birong Valley, 26°39′05.36″ N, 99°44′15.57″ E, Alt. 3406 m, 15 July 2023, L. S. Wang et al., 23-75283; Shangrila city, Birong Valley, 26°37′50.66″ N, 99°43′20.06″ E, Alt. 3080 m, 15 July 2023, L. S. Wang et al., 23-75759.
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Figure 12. Pseudosolorina tenuior (L. S. Wang et al. 23-75283 KUN); (A), habit of P. tenuior; (B), white lower surface with brownish tomentum and rhizines; (C), apothecia; (D), transversal sections of the thallus; (E), transversal sections of the apothecium; (F), ascus. Scale bars: 5 mm (B); 2 mm (C); 100 µm (D); 200 µm (E); 40 µm (F).
Figure 12. Pseudosolorina tenuior (L. S. Wang et al. 23-75283 KUN); (A), habit of P. tenuior; (B), white lower surface with brownish tomentum and rhizines; (C), apothecia; (D), transversal sections of the thallus; (E), transversal sections of the apothecium; (F), ascus. Scale bars: 5 mm (B); 2 mm (C); 100 µm (D); 200 µm (E); 40 µm (F).
Jof 11 00169 g012

4. Conclusions

The revised circumscription of Solorina now comprises two distinct species, S. crocea and S. crocoides, which are distinguished by the unique characteristics of their thallus, photobiont layers, and chemistry. Specifically, S. crocea can be identified by its smooth upper surface with apothecia and the presence of two photobiont layers. S. crocoides is characterized by dark granules along the thallus margin and only a single photobiont layer containing both cyanobacteria and chlorophyte cells. No apothecia have been observed. These two species share similar secondary metabolites, but S. crocea contains gyrophoric acid, which is absent in S. crocoides.
Species within the new genus Pseudosolorina can be categorized into three distinct groups based on characteristics, including color and size of their thalli and formation of the photobiont layer.
The first group, P. hepatizon, P. simensis, and P. tenuior comprises species with greyish-brown upper surfaces and a single photobiont layer containing both cyanobacteria and chlorophytes. P. hepatizon differs from the other two species by its lack of secondary metabolites. P. simensis and P. tenuior both contain gyrophoric acid, methyl gyrophorate, tenuiorin, and 2′-O-methyltenuiorin. P. simensis differs from P. tenuior by the thickness of its photobiont layer (70–150 μm) relative to P. tenuior (20–60 μm). The second group features species with green upper surfaces, relatively large thalli (2–5 cm in diam.), and possession of a single green photobiont layer containing solely chlorophytes. This group consists of two species: P. parmigera and P. saccata. P. parmigera exhibits a more pruinose upper surface without external cephalodia, whereas P. saccata is rarely pruinose and exhibits external cephalodia on the upper surface. The third group comprises three species with relatively small thalli (<2 cm in diam.). Thalli of P. spongiosa form only a small circle around the apothecium. The most significant distinction between the three species is their number of ascospores; specimens of P. bispora have two spores, P. bispora var. monospora has one large spore, and P. spongiosa has four spores.
Key to species of Solorina and Pseudosolorina
1a. Lower surface bright orange, containing solorinic acid and averantin2
1b. Lower surface white or pale brown, without solorinic acid and averantin3
2a. Thallus with smooth margin, contains two separate photobiont layersS. crocea
2b. Thallus with sorediate margin, contains a single photobiont layerS. crocoides
3a. Upper surface gray-brown, with single photobiont layer containing cyanobacteria and chlorophytes4
3b. Upper surface apple green, with single photobiont layer containing chlorophytes6
4a. Lacking secondary metabolitesP. hepatizon
4b. Containing secondary metabolites5
5a. Photobiont layer 20–60 µm in thallus and 12–40 µm in apotheciumP. tenuior
5b. Photobiont layer 70–150 µm in thallus and 60–100 µm in apotheciumP. simensis
6a. Thallus well-developed, 2–6 cm in diam.7
6b. Thallus small, less than 2 cm in diam.8
7a. Upper surface pruinose, without external cephalodiaP. parmigera
7b. Upper surface rarely pruinose, with external cephalodiaP. saccata
8a. Thallus less than 0.7 cm in diam., asci 4-sporedP. spongiosa
8b. Thallus 1–2 cm in diam., asci (1-)2-spored9
9a. Asci 1-sporedP. bispora var. monospora
9b. Asci 2-sporedP. bispora

Author Contributions

Conceptualization, L.W. and Y.Z.; Methodology, M.A. and R.F.; Investigation, Y.G., J.J. and S.Z.; Resources, W.M.; Writing—original draft, T.Z.; Writing—review & editing, F.R.W. and X.W. All authors have read and agreed to the published version of the manuscript.

Funding

This study was supported by the Second Tibetan Plateau Scientific Expedition and Research Program (STEP) (2019QZKK0503), Yunnan Fundamental Research Project (202401AT070196), Yunnan Young & Elite Talents Project (YNWR-QNBJ-2020-224), Science and Technology Fund of Guizhou (ZK [2023]-236, [2020] 1Y074), Guizhou Forestry Research Project ([2023] 07), Doctoral Fund of Guizhou Academy of Sciences ([2023] 01), Youth Innovation Promotion Association CAS (2020388), and National Natural Science Foundation of China (31750001, 31970022, 32460429).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The original contributions presented in the study are included in the article, further inquiries can be directed to the corresponding author.

Acknowledgments

We express our sincere thanks to the University of Helsinki and Linnaean herbarium for providing type specimens and digital images.

Conflicts of Interest

The authors declare no conflict of interest.

References

  1. Acharius, E. Forteckning pa de i sverige vaxande arter af Lafvarnes familji. Kongliga Vetensk-Acad. Nya Handl. 1808, 29, 228. [Google Scholar]
  2. Miadlikowska, J.; Lutzoni, F. Phylogenetic classification of peltigeralean fungi (Peltigerales, Ascomycota) based on ribosomal RNA small and large subunits. Am. J. Bot. 2004, 91, 449. [Google Scholar] [CrossRef]
  3. Linné, C.V. Species Plantarum, 2nd ed.; Laurentll Salvll: Berlin, Germany, 1753; p. 1149. [Google Scholar]
  4. Linné, C.V. Flora Suecica, 2nd ed.; Sumtu & Literis Laurentii Salvii: Stockholm, Sweden, 1755; p. 419. [Google Scholar]
  5. Flotow, J.V. Lichen des reisevereins und des hochstetterschen herbarii. Linnaea 1843, 17, 17. [Google Scholar]
  6. Nylander, W. Synopsis Methodica Lichenum, 2nd ed.; Ex Typis L. Martinet: Paris, France, 1860; p. 331. [Google Scholar]
  7. Anzi, M. Manipulus lichenum rariorum vel novorum quos in Longobardia et Etruria collegit et enumeravit. Comment. Soc. Crittog. Ital. 1862, 1, 136. [Google Scholar]
  8. Dodge, C.W. Lichenological notes on the flora of the Antarctic Continent and the subantarctic islands IX–XI. Nova Hedwig. 1971, 19, 494. [Google Scholar]
  9. Fée, A.L.A. Supplement a l’essai sur les cryptogames des ecorces exotiques officinales. Essai Cryptogames Écorces Exot. Off. 1837, 2, 1–12. [Google Scholar]
  10. Arnold, F. Lichenologische ausfluge in Tirol. In Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien; Kaiserlich-Königliche Zoologisch-Botanische Gesellschaft in Wien: Vienna, Austria, 1876; Volume 26, p. 371. [Google Scholar]
  11. Nylander, W. Lichenes novi e freto behringii. Flora Bot. Ztg. 1884, 67, 219. [Google Scholar]
  12. Hue, A. Trois lichens nouveaux. Bull. Soc. Bot. Fr. 1907, 54, 419–420. [Google Scholar] [CrossRef]
  13. Kirk, P.M.; Cannon, P.F.; Minter, D.W.; Stalpers, J.A. Dictionary of the Fungi, 10th ed.; CABI Publishing: London, UK, 2008; p. 643. [Google Scholar]
  14. Wei, J.C. An Enumeration of Lichens in China, 2nd ed.; China Forestry Publishing House: Jilin, China, 2020; pp. 448–449. [Google Scholar]
  15. Nylander, W. Classification des Peltigeres. J. Nat. 1884, 6, 387–389. [Google Scholar]
  16. Gyelnik, V. Líchenologíai kozlemények. Magyar Bot. Lapok 1930, 29, 30. [Google Scholar]
  17. Räsänen, V. Das System der Flechten. Übersicht mit Bestimmungstabellen der natürlichen Flechtenfamilien, ihrer Gattungen, Untergattungen, Sektionen und Untersektionen. Acta Bot. Fenn. 1943, 33, 12–42. [Google Scholar]
  18. Thomson, N.; Thomson, J.W.W. Spore ornamentation in the lichen genus Solorina. Bryologist 1984, 87, 151–153. [Google Scholar] [CrossRef]
  19. Krog, H.; Swinscow, T.D.V. Solorina simensis and S. saccata. Lichenologist 1986, 18, 57–62. [Google Scholar] [CrossRef]
  20. Martínez, I.; Burgaz, A.R. Revision of the genus Solorina (lichenes) in Europe based on spore size variation. Ann. Bot. Fenn. 1998, 35, 137–142. [Google Scholar]
  21. Ebizuka, Y.; Sankawa, U.; Shibata, S. The constituents of Solorina crocea: Averythrin 6-monomethyl ether and methyl gyrophorate. Phytochemistry 1970, 9, 2061–2063. [Google Scholar] [CrossRef]
  22. Wang, L.S.; Harada, H.; Narui, T.; Culberson, C.F. Cyanomorph of Solorina crocea (lichenized Ascomycota, Peltigeraceae) from Sichuan, China. Lichenology 2005, 4, 1–6. [Google Scholar]
  23. Stone, D.F.; McCune, B.; Pardo-De, H.C.J.; Magain, N.; Miadlikowska, J. Sinuicella denisonii, a new genus and species in the Peltigeraceae from western North America. Lichenologist 2021, 53, 185–192. [Google Scholar] [CrossRef]
  24. Culberson, C.F. Supplement to “chemical and botanical guide to lichen products”. Bryologist 1970, 73, 177–377. [Google Scholar] [CrossRef]
  25. Orange, A.; James, P.W.; White, F.J. Microchemical Methods for the Identification of Lichens, 2nd ed.; British Lichen Society: London, UK, 2010; p. 101. [Google Scholar]
  26. Feige, G.B.; Lumbsch, T.; Huneck, S.; Elix, J. Identification of lichen substances by a standardize high-performance liquid chromatography method. J. Chromatogr. A 1993, 646, 417–427. [Google Scholar] [CrossRef]
  27. Larena, I.; Salazar, O.; Gonzalez, V.; Julian, M.C.; Rubio, V. Design of a primer for ribosomal DNA internal transcribed spacer with enhanced specificity for ascomycetes. J. Biotechnol. 1999, 75, 187–194. [Google Scholar] [CrossRef]
  28. White, T.; Bruns, T.; Lee, S.; Taylor, F.; White, T.J.; Lee, S.H.; Taylor, L.; Shawetaylor, J. PCR Protocols: A Guide to Methods and Applications; Academic Press Inc.: New York, NY, USA, 1990; pp. 315–322. [Google Scholar]
  29. Rehner, S.A.; Samuels, G.J. Taxonomy and phylogeny of Gliocladium analysed from nuclear large subunit ribosomal DNA sequences. Mycol. Res. 1994, 98, 625–634. [Google Scholar] [CrossRef]
  30. Vilgalys, R.; Hester, M. Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. J. Bacteriol. 1990, 172, 4239–4246. [Google Scholar] [CrossRef]
  31. 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]
  32. Nubel, U.; Garcia-Pichel, F.; Muyzer, G. PCR primers to amplify 16S rRNA fenes from cyanobacteria. Appl. Environ. Microbiol. 1997, 63, 3327–3332. [Google Scholar] [CrossRef] [PubMed]
  33. Piercey-Normore, M.D.; Depriest, P.T. Algal switching among lichen symbioses. Am. J. Bot. 2001, 88, 1490–1498. [Google Scholar] [CrossRef] [PubMed]
  34. Kroken, S.; Taylor, J.W. Phylogenetic species, reproductive mode, and specificity of the green alga Trebouxia forming lichens with the fungal genus Letharia. Bryologist 2000, 103, 645–660. [Google Scholar] [CrossRef]
  35. Zhao, X.; Zhang, L.L.; Zhao, Z.T.; Wang, W.C.; Leavitt, S.D.; Lumbsch, H.T. A Molecular phylogeny of the lichen genus Lecidella focusing on species from mainland China. PLoS ONE 2015, 10, 0139405. [Google Scholar] [CrossRef] [PubMed]
  36. Katoh, K.; Standley, D.M. MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Mol. Biol. Evol. 2013, 30, 772–780. [Google Scholar] [CrossRef] [PubMed]
  37. Lanfear, R.; Frandsen, P.B.; Wright, A.M.; Senfeld, T.; Calcott, B. PartitionFinder 2: New methods for selecting partitioned models of evolution formolecular and morphological phylogenetic analyses. Mol. Biol. Evol. 2017, 34, 772–773. [Google Scholar] [CrossRef]
  38. Nguyen, L.T.; Schmidt, H.A.; Haeseler, V.A.; Minh, B.Q. IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol. Biol. Evol. 2015, 32, 268–274. [Google Scholar] [CrossRef] [PubMed]
  39. Ronquist, F.; Teslenko, M.; 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]
  40. Rambaut, A. FigTree, v. 1.4.0; Institute of Evolutionary Biology, University of Edinburgh: Edinburgh, UK, 2012. [Google Scholar]
  41. Vecherskii, M.V.; Khayrullin, D.R.; Shadrin, A.M.; Lisov, A.V.; Zavarzina, A.G.; Zavarzin, A.A.; Leontievsky, A.A.; Stajich, J.E. Metagenomes of lichens Solorina crocea and Peltigera canina. Microbiol. Resour. Announc. 2022, 11, e01000-21. [Google Scholar] [CrossRef] [PubMed]
  42. Reisigl, H.; Pitschmann, H. Obere Grenzen von flora und vegetation in der Nivalstufe der Zentralen Ötztaler Alpen (Tirol). Vegetatio 1958, 8, 93–129. [Google Scholar] [CrossRef]
  43. Kiener, W. A list of algae chiefly from the alpine aone of Longs Peak, Colorado. Madroño 1946, 8, 161–173. [Google Scholar]
  44. Hale, M.E., Jr. Lichens from Baffin Island. Am. Midl. Nat. 1954, 51, 232–264. [Google Scholar] [CrossRef]
  45. Egan, R.S. Recent literature on lichens 102. Bryologist 1979, 82, 314–320. [Google Scholar]
  46. Roux, C. Liste des lichens et champignons lichénicoles de France. Bull. Société Linnéenne Provence 2012, 16, 91. [Google Scholar]
  47. Wirth, V.; Hauck, M.; Schultz, M. Die Flechten Deutschlands; Eugen Ulmer KG: Stuttgart, Germany, 2013; p. 1244. [Google Scholar]
  48. Daniëls, F.J.A. Vegetation classification in Greenland. J. Veg. Sci. 1994, 6, 781–790. [Google Scholar] [CrossRef]
  49. Llimona, X.; Hladun, N.L. Checklist of the lichens and lichenicolous fungi of the Iberian Peninsula and Balearic Islands. Bocconea 2001, 14, 5–581. [Google Scholar]
  50. Hadač, E. Plant communities of the Kaldidalur area, WSW Iceland. Part 1. Syntaxonomy. Folia Geobot. Phytotax. 1985, 20, 113–175. [Google Scholar] [CrossRef]
  51. Hadač, E. Snow-land communities of Reykjanes Peninsula, SW. Iceland (plant communities of Reykjanes Peninsula, Part 4). Folia Geobot. Phytotax. 1971, 6, 105–126. [Google Scholar] [CrossRef]
  52. Martellos, S.; Conti, M.; Nimis, P.L. Aggregation of Italian lichen data in ITALIC 7.0. J. Fungi 2023, 9, 556. [Google Scholar] [CrossRef] [PubMed]
  53. Sato, M. Range of The Japanese Lichens (VI)–(VI). Bull. Fac. Lib. Arts 1960, 11, 53–62. [Google Scholar]
  54. Galloway, D.J. Vegetaion studies on the Humboldt mountains Fiordland Part 2: The lichens. N. Z. Ecol. Soc. 1966, 13, 19–23. [Google Scholar]
  55. Vetaas, O.R. Primary succession of plant assemblages on a glacier Foreland-Bodalsbreen, Southern Norway. J. Biogeogr. 1994, 21, 297–308. [Google Scholar] [CrossRef]
  56. Fałtynowicz, W.; Czarnota, P.; Krzewicka, B.; Wilk, K.; Jabłońska, A.; Oset, M.; Ossowska, E.A.; Śliwa, L.; Kukwa, M. Lichens of Poland: A Fifth Annotated Checklist; W. Szafer Institute of Botany, Polish Academy of Sciences: Kraków, Poland, 2024; pp. 496–497. [Google Scholar]
  57. Koroleva, N.E. Phytosociological survey of the tundra vegetation of the Kola Peninsula, Russia. J. Veg. Sci. 1994, 5, 803–812. [Google Scholar] [CrossRef]
  58. Jonasson, S. Plant communities and species distribution of low alpine betula nana heaths in Northernmost Sweden. Vegetatio 1981, 44, 51–64. [Google Scholar] [CrossRef]
  59. Liška, J.; Palice, Z.; Slavíková, Š. Checklist and red list of lichens of the Czech Republic. Preslia 2008, 80, 151. [Google Scholar]
  60. Kondratyuk, S.; Popova, L.P.; Khodosovtsev, O.Y.; Lőkös, L.; Fedorenko, N.M.; Kapets, N.V. The fourth checklist of Ukrainian lichen-forming and lichenicolous fungi with analysis of current additions. Acta Bot. Hung. 2021, 63, 97–163. [Google Scholar] [CrossRef]
  61. Newberry, C.C.; St. Clair, L.L. Additions to the lichen flora of Utah. I. Bryologist 1991, 94, 154–156. [Google Scholar] [CrossRef]
  62. Hansen, E.S.; Obermayer, W. Notes on Arthrorhaphis and its lichenicolous fungi in Greenland. Bryologist 1999, 102, 104–107. [Google Scholar] [CrossRef]
  63. Egan, R.S. Additions to the lichen flora of New Mexico. Bryologist 1970, 73, 143–145. [Google Scholar] [CrossRef]
  64. Lynge, B. Some General results of recent Norwegian research work on arctic lichens. Rhodora 1934, 36, 133–171. [Google Scholar]
  65. Frey, E. Die Flechtenflora und -Vegetation des Nationalparks im Unterengadin. Ergeb. Wiss. Untersuch. Schweiz. Natl. 1952, 3, 377. [Google Scholar]
  66. Moser, T.J.; Nash, T.H., III; Thomson, J.W. Lichens of Anaktuvuk Pass, Alaska, with emphasis on the impact of Caribou Grazing. Bryologist 1979, 82, 393–408. [Google Scholar] [CrossRef]
  67. Watson, W. The bryophytes and lichens of British woods. Part II, other woodland types. J. Ecol. 1936, 24, 446–478. [Google Scholar] [CrossRef]
  68. Christensen, S.N. Fulgensia klementii and other lichens from Mt. Olimbos, Makedhonia, Greece. Willdenowia 1995, 25, 283–288. [Google Scholar]
  69. Burnham, S.H. The flora of Indian ladder and vicinity: Together with descriptive notes on the scenery. J. Torrey Bot. Soc. 1918, 18, 101–116. [Google Scholar]
  70. Nadezhda, V.M. Floristic classification and ecology of tundra vegetation of the Taymyr Peninsula, Northern Siberia. J. Veg. Sci. 1994, 5, 813–828. [Google Scholar]
  71. Killmann, D.; Fischer, E. New records for the lichen flora of Rwanda, East Africa. Willdenowia 2005, 35, 193–204. [Google Scholar] [CrossRef]
  72. Sipman, H.J.M. Lichenotheca Latinoamericana a museo botanico berolinensi edita, fasciculum tertium. Willdenowia 1997, 27, 273–280. [Google Scholar] [CrossRef]
  73. Vitikainen, O. Range extension of Solorina simensis to Papua New Guinea. Lichenologist 1989, 21, 87. [Google Scholar] [CrossRef]
  74. Sarnthein, L.G. Flora von Oesterreich-Ungarn. (Schluss.). Oesterr. Bot. Z. 1894, 44, 350–361. [Google Scholar] [CrossRef]
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Figure 1. Phylogenetic relationships within Peltigerales were assessed using Bayesian Inference (BI), analyzing combined sequence data from nrITS, nrLSU, and mtSSU. Nodes supported by Bayesian posterior probability/ML bootstrap values ≥ 0.80/80% are denoted in the results. Maximum Likelihood bootstrap values and posterior probabilities are displayed proximal to the respective nodes. The red branches represent Peltigeraceae.
Figure 1. Phylogenetic relationships within Peltigerales were assessed using Bayesian Inference (BI), analyzing combined sequence data from nrITS, nrLSU, and mtSSU. Nodes supported by Bayesian posterior probability/ML bootstrap values ≥ 0.80/80% are denoted in the results. Maximum Likelihood bootstrap values and posterior probabilities are displayed proximal to the respective nodes. The red branches represent Peltigeraceae.
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Figure 2. Scanning electron microscopy pictures of the spore of the Solorina and Pseudosolorina species. (A), S. crocea (Theodore and Esslinger 11831 KUN); (B), P. hepatizon (L. S. Wang et al. 23-75286 KUN); (C), P. tenuior (L. S. Wang et al. 23-75283 KUN); (D), P. simensis (X. Y. Wang et al. 21-70422 KUN); (E), P. parmigera (X. Y. Wang et al. 21-70423 KUN); (F), P. saccata (L. S. Wang et al. 23-75757 KUN); (G), P. bispora (Y. Y. Zhang ZYY-46 KUN); (H), P. bispora var. monospora (L. S. Wang et al. 20-67082 KUN); (I), P. spongiosa (L. S. Wang et al. 23-75119 KUN). Scale bars: 10 µm (A–F,I); 30 µm (G,H).
Figure 2. Scanning electron microscopy pictures of the spore of the Solorina and Pseudosolorina species. (A), S. crocea (Theodore and Esslinger 11831 KUN); (B), P. hepatizon (L. S. Wang et al. 23-75286 KUN); (C), P. tenuior (L. S. Wang et al. 23-75283 KUN); (D), P. simensis (X. Y. Wang et al. 21-70422 KUN); (E), P. parmigera (X. Y. Wang et al. 21-70423 KUN); (F), P. saccata (L. S. Wang et al. 23-75757 KUN); (G), P. bispora (Y. Y. Zhang ZYY-46 KUN); (H), P. bispora var. monospora (L. S. Wang et al. 20-67082 KUN); (I), P. spongiosa (L. S. Wang et al. 23-75119 KUN). Scale bars: 10 µm (A–F,I); 30 µm (G,H).
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Table 1. Taxon sampling and mycobiont data with corresponding GenBank numbers used in phylogenetic analyses of Peltigerales. Sequences newly acquired for Solorina and Pseudosolorina are indicated in bold.
Table 1. Taxon sampling and mycobiont data with corresponding GenBank numbers used in phylogenetic analyses of Peltigerales. Sequences newly acquired for Solorina and Pseudosolorina are indicated in bold.
TaxonVoucherOriginGenBank No.
nrITSnrLSUmtSSU
Collema flaccidum2000120ChinaMW453229.1EU982618.1EU982578.1
C. furfuraceumMA-16260SpainGQ396263.1EU982608.1EU982567.1
Leptogium hibernicumBjelland TB_RB_4NorwayKX013723.1KX013759.1KX013684.1
L. saturninumMA-Lichen 16024FranceDQ466043.1EU982610.1KX027281.1
Lobaria amplissima1996 Stocker-Worgotter 1717Norway: HordalandAF524923.1AY340546.1AY340500.1
L. pulmonariaSpribille 39,224 (MSC)USA: AlaskaMN483125.1AF183934.2AY340503.1
Nephroma arcticumGoffinet 1310USAAF014109.1DQ973040.1DQ972989.1
N. bellum133 LG 214CanadaHQ455058.1HQ394180.1HQ423261.1
N. expallidumCONN 9370USAHQ455059.1HQ394183.1HQ423263.1
Pannaria conopleaO-L-196369NorwayMK811685.1AY424209.1MT068515.1
P. hookeriMJK3314ArgentinaMT755913.1JX464118.1MN634245.1
P. rubiginosaLG:R1008ReunionKF704259.1JX494269.1JX494244.1
Peltigera aphthosa CanadaU73492.1AF286759.1AY340515.1
P. leucophlebiaGoward 98-CUSAAF158651.1AF286753.1AY124166.1
P. malacea CanadaU73491.1AF286756.1MH792883.1
Pseudosolorina bisporaP6109Canada: NunavutMT522618.1MT573442.1
P. saccataKL18-0005KoreaMK503160.1MK506118.1MK508904.1
P. simensis 1L1045France: Reunion IslandMT522616.1MK517842.1
P. simensis 2L915RwandaMT522617.1MK517843.1
P. spongiosa08792 (HBG)Austria: SchultzMZ708642.1 KJ766495.1
Psoroma cinnamomeumO-L-184538NorwayMK811978.1OL331762.1OM103956.1
P. hypnorumPasso 20 (BCRU 4914)ArgentinaEU885309.1AY340565.1AY340523.1
Sinuicella denisoniiOSC:Stone 10024USA: OregonNR_182350.1MT942675.1MT942641.1
Solorina croceaP6110FinlandMT522619.1MT573443.1
S. crocea 2AFTOL-ID 1619USA: Durham DQ973043.1
S. crocea 3McCune 23785USA: Chicago AF286824.1
Sticta beauvoisiiLg3303USA: NCKT281725.1DQ986769.1KT281681.1
S. weigelii1997 Stenroos 4816 (TUR)GuyanaAF524905.1EU558794.1EU558865.1
Vahliella leucophaeaEkman 3202 (BG)NorwayAF429266.1JX464125.1HQ268596.1
Pseudosolorina bispora20-66618 (KUN)China: SichuanPQ627847PQ623371PQ623429
P. bisporaZYY-46 (KUN)Austria: SalzburgPQ627812 PQ623428
P. bispora24-76716 (KUN)China: YunnanPQ627820 PQ623414
P. bispora var. monospora20-67082 (KUN)China: QinghaiPQ627846 PQ623427
P. bispora var. monosporaXY20-2943 (KUN)China: QinghaiPQ627817PQ623369PQ623425
P. bispora var. monosporaXY20-828 (KUN)China: QinghaiPQ627816PQ623370PQ623426
P. hepatizon17-55094 (KUN)China: YunnanPQ627849PQ623379PQ623444
P. hepatizon23-75286 (KUN)China: YunnanPQ627835PQ623378PQ623443
P. parmigera19-62636 (KUN)China: YunnanPQ627850 PQ623437
P. parmigera21-70423 (KUN)China: SichuanPQ627843 PQ623435
P. parmigera23-75739 (KUN)China: YunnanPQ627831PQ623374PQ623436
P. parmigera21-70363 (KUN)China: SichuanPQ627845 PQ623434
P. parmigera24-76096(KUN)China: YunnanPQ627824PQ623363PQ623418
P. parmigeraXY21-272 (KUN)China: SichuanPQ627819 PQ623413
P. saccata23-75280 (KUN)China: YunnanPQ627840
P. saccata23-75757 (KUN)China: YunnanPQ627828
P. saccata23-75281a (KUN)China: YunnanPQ627839PQ623365PQ623421
P. saccataXY23-383 (KUN)China: YunnanPQ627813PQ623372PQ623430
P. saccata23-75282 (KUN)China: YunnanPQ627837 PQ623433
P. saccata23-75755 (KUN)China: YunnanPQ627830PQ623373PQ623432
P. saccata23-75756 (KUN)China: YunnanPQ627829 PQ623431
P. saccata24-76098b (KUN)China: YunnanPQ627822 PQ623416
P. simensis21-70422 (KUN)China: SichuanPQ627844
P. simensisXY21-274 (KUN)China: SichuanPQ627815 PQ623438
P. simensis23-75761 (KUN)China: YunnanPQ627826PQ623375PQ623439
P. spongiosa21-70424 (KUN)China: SichuanPQ627842 PQ623424
P. spongiosa23-75118 (KUN)China: SichuanPQ627841PQ623368
P. spongiosa23-75281b (KUN)China: YunnanPQ627838PQ623364PQ623420
P. spongiosa24-76095 (KUN)China: YunnanPQ627825 PQ623419
P. spongiosa24-76098a (KUN)China: YunnanPQ627823 PQ623417
P. spongiosaCMY-42 (KUN)China: XizangPQ627818PQ623367PQ623423
P. spongiosaXY22-1164 (KUN)China: GansuPQ627814PQ623366PQ623422
P. spongiosa24-76099 (KUN)China: YunnanPQ627821PQ623359PQ623415
P. tenuior23-75283 (KUN)China: YunnanPQ627836 PQ623442
P. tenuiorXY21-270 (KUN)China: SichuanPQ627848PQ623376PQ623440
P. tenuior23-75759 (KUN)China: YunnanPQ627827PQ623377PQ623441
Solorina crocoides23-75431 (KUN)China: XizangPQ627834PQ623362
S. crocoides23-75450 (KUN)China: XizangPQ627832PQ623360
S. crocoides23-75449 (KUN)China: XizangPQ627833PQ623361
Table 2. The photobionts sequenced from Solorina and Pseudosolorina, with corresponding GenBank numbers. The horizontal line signifies that the information is not yet available.
Table 2. The photobionts sequenced from Solorina and Pseudosolorina, with corresponding GenBank numbers. The horizontal line signifies that the information is not yet available.
HostChlorophytaCyanobacteria
TaxonVoucherTaxonGenBank No.TaxonGenBank No.
ITS16S RNA
Pseudosolorina bispora24-76716Coccomyxa sp.PV085834Nostoc sp.PV083155
Pseudosolorina bispora var. monospora20-67082Coccomyxa sp.PV085836Nostoc sp.PV083158
Pseudosolorina hepatizon17-55094Asterochloris sp.PV085839Nostoc sp.PV083157
Pseudosolorina spongiosa24-76095Coccomyxa sp.PV085835––
Pseudosolorina parmigera24-76096Coccomyxa sp.PV085837––
Pseudosolorina saccataXY23-383Coccomyxa sp.PV085838––
Pseudosolorina simensis23-75761––Nostoc sp.PV083160
Pseudosolorina tenuior23-75283––Nostoc sp.PV083159
Solorina crocoides23-75450Asterochloris sp.PV085840Nostoc sp.PV083156
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Zheng, T.; Wang, L.; Ai, M.; Gan, Y.; Fan, R.; Zhang, Y.; Worthy, F.R.; Jin, J.; Meng, W.; Zhang, S.; et al. Taxonomic Revision of Solorina (Peltigeraceae, Ascomycota), Reveals a New Genus and Three New Species. J. Fungi 2025, 11, 169. https://doi.org/10.3390/jof11030169

AMA Style

Zheng T, Wang L, Ai M, Gan Y, Fan R, Zhang Y, Worthy FR, Jin J, Meng W, Zhang S, et al. Taxonomic Revision of Solorina (Peltigeraceae, Ascomycota), Reveals a New Genus and Three New Species. Journal of Fungi. 2025; 11(3):169. https://doi.org/10.3390/jof11030169

Chicago/Turabian Style

Zheng, Ting, Lisong Wang, Min Ai, Yuxin Gan, Rong Fan, Yingjun Zhang, Fiona Ruth Worthy, Jizhen Jin, Wenping Meng, Shengbang Zhang, and et al. 2025. "Taxonomic Revision of Solorina (Peltigeraceae, Ascomycota), Reveals a New Genus and Three New Species" Journal of Fungi 11, no. 3: 169. https://doi.org/10.3390/jof11030169

APA Style

Zheng, T., Wang, L., Ai, M., Gan, Y., Fan, R., Zhang, Y., Worthy, F. R., Jin, J., Meng, W., Zhang, S., & Wang, X. (2025). Taxonomic Revision of Solorina (Peltigeraceae, Ascomycota), Reveals a New Genus and Three New Species. Journal of Fungi, 11(3), 169. https://doi.org/10.3390/jof11030169

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