An overview of the genus Lobothallia (lichenized Ascomycota, Megasporaceae) in China

205 An overview of the genus Lobothallia (lichenized Ascomycota, Megasporaceae) in China Lun Wang1*, Yi-Shan Feng1*, Li-Song Wang2,3, Xin-Yu Wang2,3 , Yan-Yun Zhang1 1 Key Laboratory of Biodiversity Conservation and Ecological Security in the Yangtze River Basin of Anhui Province, College of Life Sciences, Anhui Normal University, 241000 Wuhu, China 2 State Key Laboratory of Phytochemistry and Natural Medicines, Kunming Institute of Botany, Chinese Academy of Sciences, 650201 Kunming, China 3 Yunnan Key Laboratory for Fungal Diversity and Green Development, Kunming Institute of Botany, Chinese Academy of Sciences, 650201 Kunming, China Corresponding authors: Xin-Yu Wang ([email protected]); Yan-Yun Zhang ([email protected]) Copyright: © Lun Wang et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Lobothallia is a species-rich genus of crustose lichens that is mainly distributed in the Northern Hemisphere. In recent years, significant advancements have been made in the taxonomy and phylogeny of this genus, leading to the description of numerous new species. However, Lobothallia in China has never been systematically revised. In this study, approximately 500 specimens of Lobothallia collected from different provinces of China were examined. Based on morphological and multi-gene phylogenetic analyses, an overview of this genus in China was provided. Eight species, L. benzilanensis, L. complanata, L. polypycnidiata, L. pseudopruinosa, L. pulchra, L. rubra, L. stipitata and L. wangii, were described as new to science. The species, Lobothallia brachyloba, was reported in China for the first time. In addition, we generated DNA sequences of the species Lobothallia hedinii from topotype specimens and clarified its phylogenetic position. Detailed descriptions, illustrations of morphological characters of the above-mentioned species, and comparisons with closely related taxa are provided. A dichotomous key to 22 species of Lobothallia from China is presented. Key words: Lichen-forming fungi, new species, Pertusariales, phylogeny, species delimitation Introduction According to the estimation of Hyde et al. (2024), fungi comprise 19 phyla, 83 classes, 1,220 families, 10,685 genera and approximately 140,000 species. Lichenization represents a major fungal lifestyle, with ca. 19,000 known lichenized species (lichens) worldwide (Lücking et al. 2017), including about 3,000 reported from China (Wei 2020). Over the past decades, while China has witnessed a significant increase in lichenological research output, studies focusing on the genus Lobothallia (Clauzade & Cl. Roux) Hafellner remain scarce (Kou et al. 2013; Paukov et al. 2019; Zhang et al. 2020, 2024a). The taxonomic concept of Lobothallia was initially established as a subgenus within Aspicilia A. Massal. (Clauzade and Roux 1984), with subsequent elevation to generic rank in Aspiciliaceae (ined.) by Hafellner (1991). Academic editor: Thorsten Lumbsch Received: 30 September 2025 Accepted: 6 November 2025 Published: 20 November 2025 Citation: Wang L, Feng Y-S, Wang L-S, Wang X-Y, Zhang Y-Y (2025) An overview of the genus Lobothallia (lichenized Ascomycota, Megasporaceae) in China. MycoKeys 125: 205–244. https://doi. org/10.3897/mycokeys.125.173554 MycoKeys 125: 205–244 (2025) DOI: 10.3897/mycokeys.125.173554 * These authors contributed equally to this work. 206 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Schmitt et al. (2006) reconstructed the phylogenetic framework of Pertusariales using nuclear large subunit (nuLSU) and mitochondrial small subunit (mtSSU) ribosomal DNA sequences, proposing the transfer of Aspicilia and Lobothallia to Megasporaceae Lumbsch and regarding Aspiciliaceae as a synonym of Megasporaceae. Nordin et al. (2010) reconstructed the phylogeny of Megasporaceae by integrating nrLSU and mtSSU data and confirmed the monophyly of both Megasporaceae and Lobothallia. Subsequent researchers have continually revised Megasporaceae, introducing several new genera (Sohrabi et al. 2013b; Haji Moniri et al. 2017; Zakeri et al. 2017; Paukov et al. 2024; Wheeler et al. 2024). To date, eleven genera within this family are accepted (Wheeler et al. 2024), with Lobothallia exhibiting the closest phylogenetic affinity to Teuvoa Sohrabi & S.D. Leav. Lobothallia is widely distributed in arid and mountainous regions of the Northern Hemisphere, though species richness varies considerably across different parts of Eurasia (Paukov et al. 2019). A few widespread taxa, such as Lobothallia alphoplaca (Wahlenb.) Hafellner, have been recorded in the Southern Hemisphere, for example in New Zealand (Galloway and Ledingham 2012). As the third most species-rich genus within Megasporaceae, Lobothallia currently encompasses 31 accepted species: L. alphoplaca, L. brachyloba Paukov & I.V. Frolov, L. cernohorskyana (Clauzade & Vězda) A. Nordin et al., L. chadefaudiana (Cl. Roux) A. Nordin et al., L. cheresina (Müll. Arg.) A. Nordin et al., L. controversa Cl. Roux & A. Nordin, L. crassimarginata Kou & Q. Ren, L. crenulata Lun Wang & Y.Y. Zhang, L. densipruinosa A. Ashraf et al., L. determinata (H. Magn.) T.B. Wheeler, L. elobulata Zulfiqar et al., L. epiadelpha Paukov & A. Nordin, L. gangwondoana S.Y. Kondr. et al., L. hedinii (H. Magn.) Paukov et al., L. hydrocharis (Poelt & Nimis) Sohrabi & Nimis, L. iqbalii Zulfiqar et al., L. kuminovae (Sedeln.) Sedeln., L. lacteola (Oxner) Şenkard. et al., L. lobulata Lun Wang & Y.Y. Zhang, L. melanaspis (Ach.) Hafellner, L. pakistanica Razzaq et al., L. peltastictoides (Hasse) T.B. Wheeler, L. platycarpa (J. Steiner) Paukov, L. praeradiosa (Nyl.) Hafellner, L. pruinosa Kou & Q. Ren, L. pulvinata R. Zulfiqar et al., L. radiosa (Hoffm.) Hafellner, L. recedens (Taylor) A. Nordin et al., L. semisterilis (H. Magn.) Y.Y. Zhang, L. subdiffracta (H. Magn.) Paukov and L. zogtii Paukov & Davydov (Hafellner 1991; Nordin et al. 2010; Roux 2012; Kou et al. 2013; Nimis 2016; Roux et al. 2016; Sedelnikova and Sedelnikov 2018; Paukov et al. 2019; Kondratyuk et al. 2020; Sipman and Aptroot 2020; Zhang et al. 2020, 2024a; Ashraf et al. 2022; Zulfiqar et al. 2022, 2023; Nascimbene et al. 2023; Wheeler et al. 2024). The taxonomic nomenclature of several species within the genus has undergone revision. Lobothallia farinosa (Flörke) A. Nordin, Savić & Tibell, originally proposed by Nordin et al. (2010) as a new combination based on the invalid basionym Aspicilia farinosa sensu Nyl., was subsequently reclassified by Roux et al. (2016) as the novel species Lobothallia controversa. Lobothallia parasitica (B. de Lesd.) Mayrhofer et al. (Mayrhofer et al. 2005) is considered a synonym of L. radiosa, since its basionym, Aspicilia parasitica B. de Lesd., was treated as Lobothallia radiosa chemotype parasitica by Paukov et al. (2019) based on isotype specimen study. Kou et al. (2013) described Lobothallia helanensis Kou & Q. Ren from Nei Mongol, China; however, subsequent phylogenetic and morphological analyses by Paukov et al. (2019) demonstrated that Lecanora subdiffracta H. Magn. and Lobothallia helanensis are conspecific. Since the epithet “subdiffracta” has nomenclatural priority, Lobothallia helanensis was recombined as L. subdiffracta (H. Magn.) Paukov. 207 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China In China, Lobothallia predominantly occurs in the arid and mountainous environments of northern and northwestern regions, as documented in both historical and contemporary studies (Magnusson 1940, 1944; Mamut et al. 2010, 2012; Kou et al. 2013; Paukov et al. 2019; Zhang et al. 2020, 2024a). Prior to this study, 13 species were known from China, with 10 species (i.e., L. crassimarginata, L. cheresina, L. crenulata, L. determinata, L. hedinii, L. lobulata, L. pruinosa, L. semisterilis, L. subdiffracta, and L. zogtii) having their type localities within the country. During our investigation of lichen diversity in China, approximately 500 Lobothallia specimens were examined and over 450 sequences (ITS, nrLSU, mtSSU) were generated. Using an integrative approach combining morphology, anatomy, chemistry, and multi-locus phylogenetic reconstruction, we conducted comprehensive molecular phylogenetic analyses and species delimitation within the genus. This investigation resulted in the discovery of eight new species and one new record for China, while also clarifying the phylogenetic position of Lobothallia hedinii for the first time. Detailed morphological descriptions with microscopic photographs are provided for these ten species, accompanied by interspecific comparative discussions. A dichotomous key to all Chinese Lobothallia species is presented. Materials and methods Sample collection, morphology and chemical examination Through fieldwork, loans, and herbarium visits, approximately 500 Lobothallia specimens were examined in this study. These specimens were collected from nine provinces in China, namely Gansu, Hebei, Nei Mongol, Ningxia, Qinghai, Sichuan, Xizang, Xinjiang and Yunnan, covering the main distribution areas of this genus in the country. Voucher specimens are deposited in the following herbaria: Botany Herbarium, College of Life Sciences, Anhui Normal University (AHUB), Lichen Herbarium, Kunming Institute of Botany, Chinese Academy of Sciences (KUN-L) and Herbarium, College of Life Sciences, Shandong Normal University (SDNU). High-resolution images of type specimens for some species were provided by the Swedish Museum of Natural History (S) and the Komarov Botanical Institute of RAS (LE) or obtained from the website of Global Plants (https://plants.jstor.org/). Specimens were examined using standard microscopy techniques. External morphological characters were observed on air-dried material under a Nikon SMZ 745T (Minato City, Japan) stereomicroscope. Anatomical features were studied using a Zeiss Axio Scope A1 light microscope (Oberkochen, Germany). Cross-sections of apothecia and thalli were cut by hand with a razor blade and observed after mounting in water, 10% aqueous solution of potassium hydroxide (K), 10% water solution of nitric acid HNO3 (N), Lugol’s iodine solutions (I), or lactophenol cotton blue (LCB). Lichen substances were preliminarily examined by spot tests using the following reagents: K, C (sodium hypochlorite solution), KC (C after pretreatment with K), and P (1,4-p-phenylenediamine). Further identification was conducted by thin layer chromatography (TLC) using solvent system C (Orange et al. 2001; Elix 2014). In Lobothallia, the subhymenium and hypothecium are typically poorly delimited, even in sections stained with LCB. To avoid confusion, we therefore adopt the term “subhymenial layers” for the area between the lower limit of the hymenium and the base of the apothecium (Ivanovich et al. 2025). The separate 208 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China plectenchyma atop the hymenium is called epithecium, which is different from the pigmented layer known as epihymenium (Bungartz 2002). Ascospore measurements are presented as: (lowest recorded–) (x – - SD) – x – – (x – + SD) (–highest recorded), where x – is the arithmetic mean and SD is the standard deviation, followed by the number of measurements (n). The length-to-width ration of ascospores is indicated as “l/w”. DNA extraction, PCR amplification and sequencing Genomic DNA was extracted from dried specimens using the DNA secure Plant Kit (Tiangen, Beijing, China). PCR amplification was performed with the primers ITS1F and ITS 4a (Gardes and Bruns 1993; Larena et al. 1999) for the ITS region; LR0R and LR5 (Vilgalys and Hester 1990; Rehner and Samuels 1994) for the LSU/28S rDNA region; and mrSSU1 and mrSSU3R (Zoller et al. 1999) for the mitochondrial small subunit (mtSSU) rRNA region. Polymerase chain reaction (PCR) was carried out in a 25 μL volume, containing 12.5 μL 2 × Taq MasterMix (Aidlab, Hong Kong), 1 μL of each primer, 9.5 μL ddH2O and 1 μL DNA. The thermal cycling conditions followed Zhang et al. (2024b). PCR products were sequenced by Sangon Biotech (Shanghai, China) using the amplification primers. Sequence alignment and phylogenetic analyses The raw sequences were initially verified using the BLAST tool on the NCBI online service (https://blast.ncbi.nlm.nih.gov/Blast.cgi) to confirm the lichen affinity. Forward and reverse sequences were assembled in Geneious Prime 2024.1 (https://www.geneious.com) and manually edited. We used newly generated nrITS, nrLSU and mtSSU sequences and additional sequences obtained from GenBank (Table 1) to construct a phylogenetic tree. The newly generated sequences have been deposited in GenBank. Matrixes of nrITS, nrLSU, mtSSU were aligned separately using MAFFT v. 7 (https://mafft.cbrc.jp/alignment/server/) with the E-INS-I strategy (Katoh and Standley 2013; Katoh et al. 2019) under default parameters. The alignments were checked in MEGA v. 7 (Kumar et al. 2016) and minor misalignments were manually adjusted. Subsequently, Maximum Likelihood (ML) single-locus trees were constructed in IQTree (Nguyen et al. 2015; Trifinopoulos et al. 2016) and compared to assess potential, strongly supported conflicts among gene trees. When all statistically supported clades (bootstrap support values≥70%) were consistent across individual gene trees, the datasets were concatenated. The three loci were combined via the Concatenate Sequence function in PhyloSuite v1.2.3 (Xiang et al. 2023). ModelFinder (Kalyaanamoorthy et al. 2017) was used to estimate the best schemes and nucleotide substitution models for maximum likelihood (ML) and Bayesian inference (BI) analyses. The best schemes and selected models are shown in Table 2. Bayesian tree inference was carried out by MrBayes 3.2 (Ronquist et al. 2012). Two independent runs of four Markov Chain Monte Carlo (MCMC) were performed each for 5 M generations, sampling every 1000 generation. To assess convergence, the average standard deviation of split frequencies (ASDSF) among runs was calculated every 200 generations, with the first 25% of sampled trees discarded as burn-in. The analyses were terminated automatically when 209 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Table 1. Voucher information and GenBank accession numbers of sequences used in this study. Newly generated sequences are in bold. ‘na’ indicates there is no sequence available. Taxa Locality Voucher nrITS nrLSU mtSSU Aspicilia angelica Montana, USA Wheeler 3535 PP965029 PP965095 PP971393 Aspicilia boykinii Montana, USA Wheeler 7263 PP975030 PP965098 PP971376 Aspicilia cinerea Finnmark, Norway Wheeler 6277 MW447391 PP965054 MW424811 Aspicilia epiglypta Västergötland, Sweden Nordin 6303 (UPS) EU057907 HM060756 HM060718 Aspicilia fumosa Montana, USA Wheeler 6802 PP965000 PP965068 PP971354 Aspicilia indissimilis Sweden Nordin 5943 (UPS) EU057909 HM060746 HM060708 Aspicilia olivaceobrunnea Arizona, USA Owe-Larsson 8771 (ASU), type PP965013 PP965084 PP971361 Aspicilia subradians Alaska, USA Wheeler 5070 PP965009 PP965080 PP971357 Aspilidea myrinii Jämtland, Sweden Nordin 7447 (WIS-0079196) MW899516 MW907112 MW907106 Aspilidea myrinii Sweden Nordin 6205 (UPS) na HM060754 HM060716 Aspilidea subadunans Alaska, USA Wheeler 4628 MW899518 MW907114 MW907108 Aspilidea subadunans Alaska, USA Wheeler 3942 MW899517 MW907113 MW907109 Aspiciliella cupreoglauca Greece Sipman & Raus 61847 (B) KY618843 KY576954 KY576930 Aspiciliella intermutans Armenia Zakeri 40503 (GLM) KY596011 KY576947 KY576923 Aspiciliella intermutans Czech Republic Palice 11394 (PRA) MH248855 MH255580 MH248888 Aspiciliella portosantana Portugal Sipman 63019 (B), type KY618852 KY576962 KY576939 Antidea brucei California, USA Owe-Larsson 9161 (ASU), isotype PP965011 PP965082 PP971359 Antidea brucei California, USA Knudsen 15069 (ASU) PP975035 PP978599 PP974318 Atrostelia magnifica Republic of Tyva, Russia Davydov 21959 (LE), isotype PP832009 PP832002 PP842134 Atrostelia magnifica Republic of Tyva, Russia Davydov 21961 (hb. Davydov), paratype PP832010 PP832003 PP842135 Circinaria arida Montana, USA Wheeler 5819 PP964998 PP978597 PP971346 C. caesiocinerea Uppland, Sweden Tibell 22612 (UPS) EU057897 HM060731 HM060693 C. esculenta Astrakhan, Russia Owe-Larsson 9796 (UPS) JQ797511 JQ797493 JQ797485 C. gibbosa Uppland, Sweden Nordin 5878 (UPS) EU057908 HM060740 HM060702 C. hoffmannii Montana, USA Wheeler 3634 PP964982 PP965052 PP971331 C. leprosescens Sweden Nordin 5906 (UPS) EU057911 HM060749 HM060711 Lobothallia alphoplaca Ukraine SK A20 (KW) KT456207 na KT456211 L. alphoplaca Norway O–L–200411 MK812484 na na L. benzilanensis China, Yunnan 18-60350 (KUN-L) PV956002 PV955950 na L. brachyloba Russia, Republic of Altai Frolov 357 (UFU), holotype MK347506 na MK348228 L. brachyloba China, Xinjiang 22-72815 (KUN-L) PV956003 na na L. cernohorskyana Iran, South Khorasan Tari 2311 (B) na JQ797496 JQ797481 L. cheresina USA, Utah Wheeler 7216 (hb. Wheeler) PP965022 na PP971366 L. complanata China, Sichuan 20-67554 (KUN-L), holotype PV956004 PV955951 PV956066 L. complanata China, Sichuan 20-66561 (KUN-L) PV956005 na PV956067 L. controversa France, Rhône-Alpes Roux 25286 (UPS), holotype na HM060761 HM060723 L. crassimarginata China, Nei Mongol Wang 20122565 (SDNU), holotype JX476026 na na L. crassimarginata China, Nei Mongol Tong 20122583 (SDNU) KC007439 na na L. crenulata China, Xizang ZYY22-301 (KUN-L), holotype PP663142 PV955952 PP663165 L. crenulata China, Xizang ZYY22-331 (AHUB) PP663141 PV955953 PP663164 L. densipruinosa Pakistan LAH 36790, holotype MZ871507 na na L. densipruinosa Pakistan LAH 36951 MZ871515 na na L. determinata USA, Montana Wheeler 3641 (hb. Wheeler) PP965020 na PP971365 L. determinata USA, Montana Wheeler 6017 (hb. Wheeler) PP965021 na PP971398 L. elobulata Pakistan LAH37153, holotype ON384441 na na L. elobulata Pakistan LAH37154 ON428667 na na L. epiadelpha Russia, Orenburg Oblast Paukov 1881 (UFU L-3189), holotype MK347505 na MK348232 L. hedinii China, Gansu 18-59518 (KUN-L) PV956006 na na L. hedinii China, Qinghai 20-68298 (KUN-L) PV956008 na PV956069 L. hedinii China, Xizang 22-71155 (KUN-L) PV956016 PV955960 PV956077 L. hedinii China, Qinghai 20-68147 (KUN-L) PV956019 PV955962 PV956080 L. hydrocharis Italy, Sardinia JN72085b (BOLO) OQ073922 na na L. hydrocharis Italy, Sardinia SMNS-STU-F-0002807 (STU) OQ073923 na na 210 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Taxa Locality Voucher nrITS nrLSU mtSSU L. iqbalii Pakistan LAH37149, holotype ON384444 na na L. iqbalii Pakistan LAH37150 ON384445 na na L. lobulata China, Sichuan ZYY22-819 (KUN-L), holotype PP663143 PV955963 PP663166 L. lobulata China, Sichuan ZYY22-829 (AHUB) PP663147 PV955964 PP663170 L. melanaspis Sweden, Jämtland Nordin 6622 (UPS) HQ259272 HM060726 HM060688 L. melanaspis Norway Owe-Larsson 8943a (UPS) JF825524 na na L. pakistanica Pakistan LAH37137, holotype ON392718 na na L. pakistanica Pakistan LAH37139 ON392720 na na L. peltastictoides USA, California Knudsen 14420 (UCR) PP965019 PP965087 PP971367 L. polypycnidiata China, Qinghai XY20-3034 (KUN-L), holotype PV956020 PV955965 PV956081 L. polypycnidiata China, Qinghai XY20-808 (KUN-L) PV956021 PV955966 PV956082 L. polypycnidiata China, Qinghai 20-68413 (KUN-L) PV956022 na PV956083 L. polypycnidiata China, Qinghai 20-67303 (KUN-L) PV956023 na PV956084 L. polypycnidiata China, Qinghai 20-67134 (KUN-L) PV956024 PV955967 PV956085 L. polypycnidiata China, Qinghai 20-68672 (KUN-L) PV956025 na PV956086 L. polypycnidiata China, Qinghai 20-67160 (KUN-L) PV956026 PV955968 PV956087 L. polypycnidiata China, Xizang 20-68878 (KUN-L) PV956027 PV955969 na L. praeradiosa China, Xinjiang ZYY22-570 (AHUB) PP663151 PV955970 PP663173 L. praeradiosa China, Xinjiang ZYY22-596 (AHUB) PP663150 PV955971 PP663172 L. praeradiosa Russia, Orenburg oblast Paukov (UFU L-1264) MK347501 na MK348229 L. pruinosa China, Qinghai 22-73170 (KUN-L) PV956030 PV955972 PV956090 L. pruinosa China, Nei Mongol Wang 20123278 (SDNU), holotype JX476028 na na L. pseudopruinosa China, Xizang 19-64075 (KUN-L), holotype PV956031 PV955973 na L. pseudopruinosa China, Qinghai 20-68331 (KUN-L) PV956032 PV955974 na L. pseudopruinosa China, Qinghai XY22-1089 (KUN-L) PV956033 PV955975 na L. pseudopruinosa China, Qinghai 22-72438 (KUN-L) PV956034 PV955976 PV956091 L. pseudopruinosa China, Xizang ZYY22-324 (KUN-L) PV956035 PV955977 na L. pseudopruinosa China, Qinghai 20-67831 (KUN-L) PV956036 PV955978 PV956092 L. pulchra China, Xizang 19-66027 (KUN-L), holotype PV956037 na PV956093 L. pulchra China, Qinghai XY20-3169 (KUN-L) PV956038 na PV956094 L. pulvinata Pakistan CH-93, paratype OP268463 na na L. pulvinata Pakistan PN-04 OP268468 na OQ772296 L. radiosa Sweden Nordin 5889 (UPS) JF703124 na na L. radiosa Czech Republic, Southern Moravia Kocourkova 10226/1 ON447612 ON391443 ON367885 L. radiosa Switzerland Lumbsch 9 Aug. 2004 (F) na DQ780306 DQ780274 L. radiosa China, Xinjiang 22-72256 (KUN-L) PX061735 na na L. recedens Portugal Sipman 62857 MN586980 na na L. recedens Sweden Nordin 6035 (UPS) HQ406807 na na L. rubra China, Xinjiang 22-72301 (KUN-L) PV956039 PV955979 PV956095 L. rubra China, Xinjiang XY22-897 (KUN-L), holotype PV956040 na PV956096 L. rubra China, Xinjiang 22-72886 (KUN-L) PV956041 na PV956097 L. semisterilis China, Gansu ZYY22-704 (AHUB) PP663161 PV955980 PP663180 L. semisterilis China, Gansu ZYY22-715 (AHUB) PP663159 PV955981 PP663178 L. subdiffracta Russia, Republic of Altai Frolov 178-1 (UFU) MK347503 na MK348233 L. subdiffracta China, Xinjiang ZYY22-628 (AHUB) PP663162 na PP663181 ‘L. helanensis’ China, Nei Mongol D.B. Tong 20122518 (SDNU) JX476030 na na ‘L. helanensis’ China, Nei Mongol D.B. Tong 20122791 (SDNU) JX476031 na na L. subdiffracta var. rimosa China, Xinjiang ZYY22-647 (KUN-L) PP663156 PV955982 PP663175 L. subdiffracta var. rimosa China, Xinjiang Wang et al. 22-72975 (KUN-L) PP663155 na PP663174 L. stipitata China, Xinjiang 22-72871 (KUN-L), holotype PV956044 na PV956100 L. stipitata China, Xinjiang ZYY22-616 (KUN-L) PV956045 PV955983 PV956101 L. stipitata China, Xinjiang ZYY22-617 (KUN-L) PV956046 PV955984 PV956102 L. stipitata China, Xinjiang XY22-887 (KUN-L) PV956047 PV955985 PV956103 L. wangii China, Xizang 22-71215 (KUN-L), holotype PV956048 na na L. wangii China, Xizang ZYY22-334 (KUN-L) PV956049 PV955986 PV956104 L. wangii China, Xizang 19-65657 (KUN-L) PV956050 na PV956105 211 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Taxa Locality Voucher nrITS nrLSU mtSSU Megaspora rimisorediata Alberta, Canada Spribille WP642 PP964997 PP978600 na M. verrucosa Antarctica Zuo NJ2018074 (hb. unknown) OL331496 OL331779 na M. verrucosa East Azerbaijan, Iran Sipman 55434 (B) na JQ797498 JQ797483 Ochrolechia subpallescens USA Lumbsch 19900a (MIN) na GU980985 GU980978 Ochrolechia upsaliensis USA Lumbsch 19916e (MIN) na GU980986 GU980979 Ochrolechia yasudae na AFTOL 882 na DQ986776 DQ986902 Oxneriaria haeyrenii Sweden Nordin 5997 (UPS) na HM060755 HM060717 Oxneriaria mashiginensis Sweden Nordin 5790 (UPS) EU057912 HM060732 HM060694 Oxneriaria permutata Sweden Nordin 6027 (UPS) EU057918 HM060747 HM060709 Oxneriaria supertegens Sweden Nordin 6023 (UPS) EU057938 HM060751 HM060713 Sagedia mastrucata Finnmark, Norway Wheeler 6467 PP975034 PP965066 PP971347 S. simoensis Troms, Norway Owe-Larsson 9000 (UPS) EU057926 HM060739 HM060701 S. simoensis Finnmark, Norway Wheeler 6288 PP975033 PP965067 PP971348 S. zonata Troms, Norway Owe-Larsson 8942 (UPS) EU057946 HM060738 HM060700 Teuvoa junipericola Utah, USA Wheeler 7390 PP965018 na PP971364 T. junipericola Utah, USA Leavitt 742 (BRY) JX306744 JX306758 na T. uxoris Castilla, Spain Rico & Pizarro 3622 (H) JX306743 JX306757 na T. uxoris Castilla, Spain Rico & Pizarro 765 (BRY) JX306745 JX306759 na the ASDSF values fell below 0.01, indicating that stationarity had been reached. ML tree analyses were performed using IQ-TREE (Nguyen et al. 2015; Trifinopoulos et al. 2016) under default settings, with branch support evaluated through 1000 standard non-parametric bootstrap replicates. Trees were visualized and graphically improved using Interactive Tree of Life webpage (iTOL) (Letunic and Bork 2021) and Adobe illustrator 2020 SP. Results Phylogenetic analysis A phylogenetic tree of Lobothallia was reconstructed based on a concatenated matrix of nrITS, nrLSU, and mtSSU sequences. Ochrolechia subpallescens Verseghy, O. upsaliensis (L.) A. Massal., and O. yasudae Vain. from the family Ochrolechiaceae R.C. Harris ex Lumbsch & I. Schmitt were selected as outgroups (Wheeler et al. 2024). The dataset comprised 127 samples, 83 of which belonged to the genus Lobothallia, including 20 type specimens and 45 newly sequenced samples (Table 1). The concatenated alignment contained 118 nrITS (36 newly generated), 78 nrLSU (30 new), and 91 mtSSU (26 new) sequences, with a total length of 2342 bp partitioned into five regions (Table 2). Phylogenetic reconstruction was performed under Maximum Likelihood (ML) and Bayesian Inference (BI) methods. Both methods produced congruent tree topologies; the BI tree is presented in the main text (Fig. 1). Table 2. The best schemes and nucleotide substitution models selected by ModelFinder for the concatenated 3-gene dataset. Partition Regions Positions Model for IQ-TREE Model for Bayes Partition 1 ITS1 1–238 TIM2e+I+G4 GTR+F+I+G4 Partition 2 5.8S 239–395 TNe+I+G4 K2P+I Partition 3 ITS2 396–575 TIM2e+I+G4 GTR+F+I+G4 Partition 4 nrLSU 576–1669 GTR+F+I+G4 GTR+F+I+G4 Partition 5 mtSSU 1670–2342 GTR+F+I+G4 GTR+F+I+G4 212 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Ochrolechia subpallescens Lumbsch 19900a USA Ochrolechia yasudae AFTOL 882 Ochrolechia upsaliensis Lumbsch 19916e USA Aspilidea myrinii Nordin 6205 Sweden Aspilidea myrinii Nordin 7447 Sweden Aspilidea subadunans Wheeler 3942 USA Aspilidea subadunans Wheeler 4628 USA Antidea brucei Owe-Larsson 9161 USA Antidea brucei Knudsen 15069 USA Teuvoa junipericola Leavitt 742 USA Teuvoa junipericola Wheeler 7390 USA Teuvoa uxoris Rico & Pizarro 765 Spain Teuvoa uxoris Rico & Pizarro 3622 Spain Lobothallia radiosa 22-72256(KKUN-L) China, Xinjiang Lobothallia radiosa Kocourkova 10226/1 Czech Republic Lobothallia radiosa Lumbsch 9 Aug. 2004 Switzerland Lobothallia radiosa Nordin 5889 Sweden Lobothallia hydrocharis JN72085b Italy Lobothallia hydrocharis SMNS-STU-F-0002807 Italy Lobothallia lobulata ZYY22-819 (KUN-L) China, Sichuan Holotype Lobothallia lobulata ZYY22-829 (AHUB) China, Sichuan Lobothallia recedens Sipman 62857 Portugal Lobothallia recedens Nordin 6035 Sweden Lobothallia cernohorskyana Tari 2311 Iran Lobothallia peltastictoides Knudsen 14420 USA Lobothallia crenulata ZYY22-301 (KUN-L) China, Xizang Holotype Lobothallia crenulata ZYY22-331 (AHUB) China, Xizang Lobothallia determinata Wheeler 3641 USA Lobothallia determinata Wheeler 6017 USA Lobothallia controversa Roux 25286 France Holotype Lobothallia cheresina Wheeler 7216 USA Lobothallia wangii 22-71215 (KUN-L) China, Xizang Holotype Lobothallia wangii ZYY22-334 (KUN-L) China, Xizang Lobothallia wangii 19-65657 (KUN-L) China, Xizang Lobothallia pseudopruinosa 19-64075 (KUN-L) China, Xizang Holotype Lobothallia pseudopruinosa ZYY22-324 (KUN-L) China, Xizang Lobothallia pseudopruinosa 20-67831 (KUN-L) China, Qinghai Lobothallia pseudopruinosa 22-72438 (KUN-L) China, Qinghai Lobothallia pseudopruinosa XY22-1089 (KUN-L) China, Qinghai Lobothallia pseudopruinosa 20-68331 (KUN-L) China, Qinghai Lobothallia densipruinosa LAH 36790, Pakistan Holotype Lobothallia densipruinosa LAH 36951, Pakistan Lobothallia pruinosa 22-73170 (KUN-L) China, Qinghai Lobothallia pruinosa Wang 20123278 China, Inner mongolia Holotype Lobothallia complanata 20-66561 (KUN-L) China, Sichuan Lobothallia complanata 20-67554 (KUN-L) China, Sichuan Holotype Lobothallia rubra 22-72301 (KUN-L) China, Xinjiang Lobothallia rubra 22-72886 (KUN-L) China, Xinjiang Lobothallia rubra XY22-897 (KUN-L) China, Xinjiang Holotype Lobothallia praeradiosa ZYY22-596 (AHUB) China, Xinjiang Lobothallia praeradiosa Paukov (UFU L-1264) Russia Lobothallia praeradiosa ZYY22-570 (AHUB) China, Xinjiang Lobothallia semisterilis ZYY22-704 (AHUB) China, Gansu Lobothallia semisterilis ZYY22-715 (AHUB) China, Gansu Lobothallia stipitata XY22-887 (KUN-L) China, Xinjiang Lobothallia stipitata ZYY22-616 (KUN-L) China, Xinjiang Lobothallia stipitata ZYY22-617 (KUN-L) China, Xinjiang Lobothallia stipitata 22-72871 (KUN-L) China, Xinjiang Holotype Lobothallia alphoplaca SK A20 Ukraine Lobothallia alphoplaca O–L–200411 Norway Lobothallia melanaspis Nordin 6622 Sweden Lobothallia melanaspis Owe-Larsson 8943a Norway Lobothallia iqbalii LAH37149 Pakistan Holotype Lobothallia iqbalii LAH37150 Pakistan Lobothallia pakistanica LAH37137 Pakistan Holotype Lobothallia pakistanica LAH37139 Pakistan Lobothallia pulvinata CH-93 Pakistan Paratype Lobothallia pulvinata PN-04 Pakistan Lobothallia polypycnidiata XY20-808 (KUN-L) China, Qinghai Lobothallia polypycnidiata 20-68413 (KUN-L) China, Qinghai Lobothallia polypycnidiata XY20-3034 (KUN-L) China, Qinghai Holotype Lobothallia polypycnidiata 20-67134 (KUN-L) China, Qinghai Lobothallia polypycnidiata 20-68672 (KUN-L) China, Qinghai Lobothallia polypycnidiata 20-67303 (KUN-L) China, Qinghai Lobothallia polypycnidiata 20-67160 (KUN-L) China, Qinghai Lobothallia polypycnidiata 20-68878 (KUN-L) China, Xizang Lobothallia hedinii 18-59518 (KUN-L) China, Gansu Lobothallia hedinii 20-68298 (KUN-L) China, Qinghai Lobothallia hedinii 22-71155 (KUN-L) China, Xizang Lobothallia hedinii 20-68147 (KUN-L) China, Qinghai Lobothallia benzilanensis 18-60350 (KUN-L) China, Yunnan Holotype Lobothallia crassimarginata Wang 20122565 China, Inner Mongolia Holotype Lobothallia crassimarginata Tong 20122583 China, Inner Mongolia Lobothallia epiadelpha Paukov 1881 Russia Holotype Lobothallia pulchra 19-66027 (KUN-L) China, Xizang Holotype Lobothallia pulchra XY20-3169 (KUN-L) China, Qinghai ‘Lobothalliahelanensis ’D.B. Tong 20122518 China, Inner Mongolia ‘Lobothallia helanensis ’D.B. Tong 20122791 China, Inner Mongolia Lobothallia subdiffracta var. rimosa ZYY22-647 China, Xinjiang Lobothallia subdiffracta var. rimosa Wang et al. 22-72975 China, Xinjiang Lobothallia subdiffracta Frolov 178-1 Russia Lobothallia subdiffracta ZYY22-628 (AHUB) China, Xinjiang Lobothallia brachyloba 22-72815 (KUN-L) China, Xinjiang Lobothallia brachyloba Frolov 357 Russia Holotype Lobothallia elobulata LAH37153 Pakistan Holotype Lobothallia elobulata LAH37154 Pakistan Atrostelia magnifica Davydov 21959 Russia Atrostelia magnifica Davydov 21961 Russia Aspiciliella cupreoglauca Sipman & Raus 61847 Greece Aspiciliella portosantana Sipman 63019 Portugal Aspiciliella intermutans Zakeri 40503 Armenia Aspiciliella intermutans Palice 11394 Czech Republic Megaspora verrucosa Zuo NJ2018074 Antarctica Megaspora verrucosa Sipman 55434 Iran Megaspora rimisorediata Spribille WP642 Canada Circinaria gibbosa Nordin 5878 Sweden Circinaria caesiocinerea Tibell 22612 Sweden Circinaria leprosescens Nordin 5906 Sweden Circinaria arida Wheeler 5819 USA Circinaria hoffmannii Wheeler 3634 USA Circinaria esculenta Owe-Larsson 9796 Russia Sagedia mastrucata Wheeler 6467 Norway Sagedia zonata Owe-Larsson 8942 Norway Sagedia simoensis Owe-Larsson 9000 Norway Sagedia simoensis Wheeler 6288 Norway Oxneriaria haeyrenii Nordin 5997 Sweden Oxneriaria mashiginensis Nordin 5790 Sweden Oxneriaria supertegens Nordin 6023 Sweden Oxneriaria permutata Nordin 6027 Sweden Aspicilia cinerea Wheeler 6277 Norway Aspicilia indissimilis Nordin 5943 Sweden Aspicilia epiglypta Nordin 6303 Sweden Aspicilia subradians Wheeler 5070 USA Aspicilia boykinii Wheeler 7263 USA Aspicilia angelica Wheeler 3535 USA Aspicilia olivaceobrunnea Owe-Larsson 8771 USA Aspicilia fumosa Wheeler 6802 USA 32/0.44 46/0.83 96/1.00 24/0.61 100/1.00 100/1.00 98/1.00 69/0.75 100/1.00 100/1.00 100/1.00 65/0.99 96/1.00 86/1.00 78/0.74 100/1.00 100/1.00 98/1.00 97/0.90 40/0.58 26/0.45 48/0.94 100/1.00 100/1.00 52/0.80 100/1.00 76/1.00 98/1.00 94/1.00 53/0.90 61/0.95 86/0.98 86/1.00 100/1.00 97/1.00 47/0.57 47/0.74 89/1.00 84/0.63 60/1.00 44/0.52 100/1.00 100/1.00 85/1.00 33/0.42 100/1.00 55/0.58 100/1.00 87/1.00 100/1.00 100/1.00 59/0.90 100/1.00 100/1.00 75/0.78 100/1.00 54/0.59 100/1.00 100/0.99 98/1.00 46/0.55 86/1.00 39/0.98 100/1.00 49/0.98 100/1.00 100/1.00 100/1.00 100/1.00 80/1.00 100/1.00 100/1.00 100/1.00 100/1.00 100/1.00 82/1.00 27/0.22 33/0.90 45/1.00 92/1.00 100/1.00 30/0.49 98/1.00 100/1.00 86/1.00 92/1.00 99/1.00 100/1.00 45/0.93 34/0.51 53/0.34 81/1.00 99/1.00 99/1.00 66/0.95 100/1.00 100/1.00 80/0.98 98/1.00 100/1.00 64/0.40 86/1.00 99/0.88 23/0.39 99/1.00 56/0.98 100/1.00 100/1.00 100/1.00 64/0.94 87/0.94 100/1.00 100/1.00 84/1.00 70/0.99 38/0.51 100/1.00 100/1.00 66/0.81 59/1.00 Tree scale: 0.05 Lobothallia Megasporaceae Atrostelia Aspiciliella Megaspora Circinaria Sagedia Oxneriaria Aspicilia Teuvoa Antidea Aspilidea Figure 1. Bayesian phylogenetic tree of Lobothallia inferred from the concatenated alignment of nrITS-nrLSU-mtSSU sequences. Maximum likelihood bootstrap values (MLBP) and Bayesian posterior probabilities (BPP) are indicated at the nodes. Thickened branches indicate MLBP ≥ 70% and BPP ≥ 0.95. Newly described species are highlighted in red, newly recorded species in China in blue, and newly generated sequences in bold. 213 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Statistical support values are labeled on the branch when the ML bootstrap proportion (MLBP) ≥ 70% and Bayesian posterior probabilities (BPP) ≥ 0.95. The phylogenetic analysis resolved 33 species of Lobothallia into three major clades (Clade I–III). Clade I (MLBP/BPP ≥96/1.00) represents the core group of Lobothallia, exhibiting phenotypic diversity spanning the genus. The basal lineage of this clade is Lobothallia brachyloba, characterized by its short marginal lobes, 1–7 apothecia per areole, and cryptolecanorine apothecia (Paukov et al. 2019). Most species within Clade I formed highly supported monophyletic groups, with the exception of L. epiadelpha and L. benzilanensis, which remained as singletons. Some backbone nodes, however, received only weak support, potentially indicating the insufficient informative sites of the sequences. Within Clade I, eight novel lineages were identified: seven described as new species, one confirmed as the species L. hedinii that was originally described from Gansu Province, China, for which no sequence was available before this research. Lobothallia pulchra and L. crassimarginata clustered together with weak support; both species contain stictic acid but differ in thallus color and apothecial type. Another new species, Lobothallia stipitata, also contains stictic acid but is distinguished by its pale brown to brown thallus, and its distinct pruinose apothecial disc with a brown thalline margin. Lobothallia polypycnidiata and L. hedinii formed sister lineages with strong support (87/1.00). Although several short-branched, highly supported subclades were observed within L. polypycnidiata, we treated them as the same species due to their consistent morphology and overlapping geographical distributions, following the species delimitation framework of Carstens et al. (2013). Lobothallia rubra and L. complanata clustered together with weak support but are morphologically distinct, differing in thallus color and thickness (white and thick vs. orangish gray and thin) and the presence versus absence of gyrophoric acid. Lobothallia pseudopruinosa, L. pruinosa, and L. densipruinosa gathered in a subclade with high support (100/1.00). Clade II consisted solely of Lobothallia elobulata. This clade is distinguished from Clades I and III by the presence of prothallus, a characteristic commonly found in the genus Aspicilia within Megasporaceae. Clade III (76/1.00) was resolved as a monophyletic lineage with Lobothallia wangii at the base. Within this clade, Lobothallia hydrocharis, L. lobulata, L. radiosa, and L. recedens formed a strongly supported subclade (100/1.00), in which all species consistently displayed gray to dark gray thalli. The remaining species, Lobothallia cernohorskyana, L. cheresina, L. crenulata, L. determinata, and L. peltastictoides, formed a distinct, well-supported subclade (70/0.99) characterized by white to off-white thalli. Taxonomy New species Lobothallia benzilanensis Lun Wang & Y. Y. Zhang, sp. nov. Fungal Names: FN 572942 Fig. 2 Etymology. The specific epithet refers to Benzilan Town, the type locality of this species. 220 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Xin-Yu Wang et al. XY19-2271 (KUN-L 73043) • Chamdo Ci., Jiangda Co., Tongpu Vil., 31°38'30.46"N, 98°26'11.62"E, alt. 3948 m, on limestone, 23 September 2020, Li-Song Wang et al. 20-68878 (KUN-L 77062), 20-68840 (KUN-L 77024). Lobothallia pseudopruinosa Lun Wang & Y. Y. Zhang, sp. nov. Fungal Names: FN 572945 Fig. 5 Etymology. The epithet indicates that this species is similar to Lobothallia pruinosa in morphology. Diagnosis. This species is characterized by its tightly adnate and thin thallus, areolate center and shortly lobate margin; plane, white or brownish gray upper surface with fine granular pruina; cryptolecanorine apothecia; large ascospores (10.0-)11.8–13.3–14.8(-17.0) × (8.0-)8.5–9.3–10.0(-11.0) µm; and its long conidia (5–)6–8(–9) × 1 µm. Holotype. China • Xizang Autonomous Region: Shigatse Ci., Dingri Co., Zhaguo Vil., 28°35'23.26"N, 86°53'55.21"E, alt. 4320 m, on rock, 27 July 2019, Li-Song Wang et al. 19-64075 (KUN-L 68571). Description. Thallus tightly adnate to the substrate, up to 3 cm across, centrally areolate, 1–2 mm thick, marginally lobate, 0.3–0.8 mm thick. Areoles (0.3–)0.6–1(–1.5) mm wide, angular to rounded, flat, not constricted at base; interspaces between areoles 0.05–0.1 mm wide. Lobes radiate, short, plane, apices equal to slightly wider than base, 1–2 mm long, base 0.5–1 mm wide, apex 0.5–1.5 mm wide. Upper surface white or pale brownish gray, pruinose, brownish where pruina thin. Upper cortex paraplectenchymatous, 30–50 µm thick, containing dark brown granules (soluble in K); epinecral layer with dense grayish-black granules (insoluble in K), 10–40(–50) µm thick. Algal layer 70– 100(–150) µm thick, discontinuous; photobiont chlorococcoid, (6–)8–20 µm diameter. Medulla 0.1–0.5 mm thick, opaque, filled with grayish-black granules (insoluble or partially soluble in K). Lower cortex absent. Apothecia cryptolecanorine, common, grouped in the center, 1–3 per areole, angular, (0.1–)0.3–0.8(–1.0) mm in diameter; disc concave, brownish to black, matt, pruinose; apothecial margin indistinct, concolorous with the thallus. Exciple narrow, widening to 10–30(–50) μm in the uppermost part. Epithecium, hymenium and subhymenial layers combined (100–)125–150(–175) µm high; epithecium 5–15 µm thick, gelatinous; epihymenium 15–25 µm high, with dark brown to brownish granules (soluble in K), N–; hymenium (70–)90–100(–110) µm high, hyaline, I+ blue; subhymenial layers 50–75 µm high, hyaline, I+ blue; algal layer relatively continuous below hypothecium, 50–70 μm high; paraphyses submoniliform, simple, septate, with 1–3 uppermost cells shorter and wider than the basal cells, 4–6 μm wide (basal cells ca. 2 μm wide); asci clavate, Aspicilia-type, hyaline, 8-spored, 60–75 × 15–25 µm; ascospores simple, hyaline, broadly ellipsoid, rarely spherical, (10.0-)11.8–13.3–14.8(-17.0) × (8.0- )8.5–9.3–10.0(-11.0) µm (n = 86), l/w ratio (1.0-)1.2–1.4–1.6(-2.1), wall ca.1 µm thick. Pycnidia rare, punctiform, slightly convex, 0.05–0.1 mm diameter, ostiole brown to dark brown, conidia bacilliform, hyaline, (5–)6–8(–9) × 1 µm. Chemistry. Cortex K+ pale yellow or orange, P–. Medulla K+ yellow to orange-red, P+ yellow, C–, KC–. Containing norstictic and connorstictic acids. 221 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Habitat and distribution. Saxicolous. Currently known only in Qinghai, Xizang Province, China. Notes. Specimens of this new species were collected from Qinghai and Xizang Provinces, China, at elevations ranging from 3000 to 4500 m. The overall morphology of these specimens is uniform, with the exception of voucher 20-67831 (KUN-L 76010), which exhibits a thinner upper cortex composed of loosely arranged hyphae. Phylogenetically, Lobothallia pruinosa and L. densipruinosa are the closest relatives of the new species. L. pruinosa can be distinguished by its lecanorine apothecia at maturity and broader lobes. L. densipruinosa differs in its whitish to light greenish gray and entirely pruinose thallus, apothecial disc dark olivaceous when wet and blackish when dry, and the absence of connorstictic acid (Kou et al. 2013; Ashraf et al. 2022). Lobothallia pakistanica resembles L. pseudopruinosa in its white, lobate thallus but differs in possessing larger central areoles, broader marginal lobes, and lacking of secondary metabolites (Zulfiqar et al. 2022). Within Lobothallia, L. brachyloba, L. platycarpa L. pulvinata and this new species share an areolate thallus with lobate margins, a whitish to whitish-gray upper surface, and the presence of norstictic acid. Nevertheless, each can be distinguished Figure 5. Lobothallia pseudopruinosa (KUN-L 68571, holotype). A. Thallus; B. Areoles and apothecia; C. Lobes; D. Section of thallus; E. Section of thallus (LCB); F. Vertical section of apothecium; G. Vertical section of apothecium (Lugol’s solution); H. Vertical section of apothecium (LCB); I. Ascus and ascospores; J. Paraphyses (LCB); K. Conidia. Scale bars:1 mm (A); 0.2 mm (B, C); 50 µm (D, F); 20 µm (E, G, H); 5 µm (I, J, K). 222 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China by specific characters: L. brachyloba has sparse thalline and 1–7 apothecia per areole (Paukov et al. 2019); L. platycarpa possesses larger areoles, fewer and non-aggregated apothecia, and limited distribution in North Africa (Algeria) (Zulfiqar et al. 2022); And L. pulvinata exhibits a thinner thallus and upper cortex (10–15 µm), and lecanorine apothecia (Zulfiqar et al. 2023). Additionally, Lobothallia cheresina, L. controversa, and L. lacteola also have a tightly adnate, whitish to whitish–gray thallus, but these species differ from L. pseudopruinosa in their absent or indistinct marginal lobes and their different secondary metabolites. Additional specimens examined. Lobothallia pseudopruinosa. China • Xizang: Shigatse Ci., Dingri Co., Zhaguo Vil., G219, 28°35'09.99"N, 87°03'42.35"E, alt. 4306 m, on surface of weathered shale in open area, 16 June 2022, LiSong Wang et al. 22-71222 (KUN-L 85846), Yan-Yun Zhang ZYY22-324 (KUN-L 81905), ZYY22-326 (KUN-L 81907) • Angren Co., 29°19'01.32"N, 87°01'59.60"E, alt. 4524 m, on rock, 19 July 2019, Li-Song Wang et al. 1963621 (KUN-L 68115) • Qinghai Prov.: Yushu Tibetan Autonomous Prefecture, Qumalai Co., Qumahe Vil., 34°54'50.67"N, 94°46'20.74"E, alt. 4396 m, on limestone, 17 September 2020, Li-Song Wang et al. 20-68331 (KUN-L 76512) • Zeku Co., Maixiu Town, 35°15'53.57"N, 101°52'30.23"E, alt. 3143–3163 m, on rock, 8 July 2022, Xin-Yu Wang and Min Ai XY22-1089 (KUN-L 84907), XY22-1088 (KUN-L 84906), An-Cheng Yin and Han-Xiang Chen 22-72438 (KUN-L 87064) • Jiuzhi Co., NianBaoYuZe National Geopark, 33°13'57.57"N, 100°55'58.56"E, alt. 4063 m, on sandy soil, 8 September 2020, Li-Song Wang et al. 20-67831 (KUN-L 76010). Lobothallia pruinosa. China • Nei Mongol: Bayanhot Town, Helan Mountain, alt. 1600 m, on rock, 17 August 2011, H.Y.Wang 20123575, 20123154, 20123282, 20123626 (SDNU) • Urat Rear Banner Co., alt. 1600 m, on rock, 19 August 2011, H.Y.Wang 20122917 (SDNU), D.B.Tong 20123276, 20123888 (SDNU) • Qinghai Prov.: Hualong Hui Ehtnic Autonomous Co., Yashiga Town, 36°03'52.78"N, 101°57'03.44"E, alt. 2002 m, on rock, 8 July 2022, Li-Song Wang et al. 22-73170 (KUN-L 87648). Lobothallia pulchra Lun Wang & Y. Y. Zhang, sp. nov. Fungal Names: FN 572946 Fig. 6 Etymology. The epithet refers to the beautiful morphology of the thallus of this species. Diagnosis. Thallus areolate with radiate lobes, upper surface white and pruinose; apothecia lecanorine, adnate and rounded, 1–2 apothecia per areole, apothecial margin distinct and persistent, disc plane to slightly convex, not to rarely with faint pruina, containing norstictic and stictic acids. Holotype. China • Xizang Autonomous Region: Shigatse Ci., Dingri Co., Zhaguo Vil., 28°35'24.49"N, 86°53'58.22"E, elev. 4293 m, on rock, 27 July 2019, Li-Song Wang et al. 19-66027 (KUN-L 70435). Description. Thallus tightly adnate to the substrate, 0.3–2 mm thick, centrally areolate, marginally lobate. Areoles 0.5–1.2 mm wide, angular to rounded, not constricted at base, interspaces between areoles 0.05–0.1 mm wide. Lobes radiate, short, 1–2.5 mm long, base width (0.3–)0.6–0.8(–1) mm, apex 223 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China width 0.6–2 mm. Upper surface moderately convex, white, partially pale brown on marginal lobes, pruinose. Upper cortex paraplectenchymatous, even, 30– 40 μm thick, containing brown granules (soluble in K); epinecral layer gelatinous, 10–20 μm thick. Algal layer 125–150 μm high, discontinuous; photobiont chlorococcoid, cells 6–18 µm diameter. Medulla 0.3–0.6 mm thick, opaque, filled with gray-black granules (insoluble in K). Lower cortex absent. Apothecia lecanorine, common, 1–2 per areole, simple to grouped in the center, orbicular to slightly angular by pressure from adjacent apothecia, (0.3– )0.6–2 mm diameter, initially immersed, later sessile, not to slightly constricted at base; disc plane, matt, brownish-black to black, epruinose or faintly pruinose, shallowly fissured when overmature; apothecial margin entire, concolorous with thallus, well-developed and persistent, (0.05–)0.1–0.2(–0.25) mm. Exciple narrow, widening to 70–80 μm in the uppermost part. Epithecium, hymenium and subhymenial layers combined 125–160 µm high; epithecium 5–15 µm high, epihymenium 12.5–20 μm high, filled with dark brown to brown granules (mostly soluble in K), N–; hymenium 90–100 μm high, hyaline, I+ dark blue; subhymenial layers ca. 50 μm high, hyaline, I+ dark blue; discontinuous algal layer beneath hypothecium; paraphyses simple, septate, submoniliform at the tips, 1–4 uppermost cells shorter and broader than basal cells, 3–5 μm wide (basal cells 2–3 μm Figure 6. Lobothallia pulchra (KUN-L 70435, holotype). A. Habitat; B. Thallus; C. Lobes; D. Apothecia; E. Vertical section of apothecium; F, G. Section of thallus (G in LCB); H. Vertical section of apothecium (LCB); I. Ascus; J. Paraphyses (LCB); K. Ascospores; L. Conidia (LCB). Scale bars: 1mm (B); 0.5 mm (C); 0.2 mm (D); 100 µm (E, F, G); 50 µm (H); 5 µm (I, J, K, L). 224 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China wide); asci clavate, Aspicilia-type, hyaline, 8-spored, 70–80 × 20–30 µm; ascospores simple, hyaline, ellipsoid to broadly ellipsoid, (10.0-)10.8–12.3–13.7(- 16.0) × (8.0-)7.8–8.5–9.1(-10.0) µm (n = 30), l/w ratio (1.1-)1.3–1.5–1.7(-1.9), wall ca.1 µm thick. Pycnidia few, punctiform, flat to slightly convex, 0.15–0.2 mm diameter; ostiole brownish black; conidia bacilliform, hyaline, 6–8 × 1 µm. Chemistry. Cortex K+ pale yellow or K–, P–; medulla K+ yellow to orange-red, P+ yellow or orange-red, C–, KC– or KC+ pale yellow; norstictic and stictic acids detected by TLC, some specimens additionally contain cryptostictic, connorstictic and constictic acids. Habitat and distribution. Saxicolous. Currently known from Xizang and Qinghai Provinces, China. Notes. Lobothallia pseudopruinosa and L. pruinosa resemble L. pulchra in having an areolate thallus with radiate lobes and a white pruinose surface, but differ in their indistinct apothecial margin, distinctly pruinose discs, and the absence of stictic acid (Kou et al. 2013). Lobothallia wangii differs from L. pulchra in its predominantly cryptolecanorine apothecia, N+ epihymenium, and the absence of both marginal lobes and stictic acid. Lobothallia pakistanica and L. pulvinata also have a white thallus with central areoles and marginal lobes, but differ from L. pulchra in their thinner thalli, apothecia that are not constricted at base, and in the lack of stictic acid (Zulfiqar et al. 2022, 2023). Additional specimens examined. China • Xizang Autonomous Region: Tingri Co., Zhaguo Vi., 28°35'24.49"N, 86°53'58.22"E, alt. 4256–4299 m, on rock, 27 July 2019, Li-Song Wang et al. 19-65686 (KUN-L 70194), 19-65652 (KUN-L 70159) • Dagze Co., Bangdui Vi, 29°44'06.51"N, 91°24'55.29"E, alt. 3700 m, on rock, 16 July 2019, Li-Song Wang et al. 19-64597 (KUN-L 69093) • Qinghai: Yushu Tibetan Autonomous Prefecture, Zaduo Co., 32°52'43.64"N, 95°20'29.61"E, alt. 4074 m, on rock, 20 September 2020, Li-Song Wang et al. 20-67281 (KUN-L 75458) • Ado Vi., 32°52'43.35"N, 95°20'30.19"E, alt. 4080 m, on rock, 20 September 2020, Xin-Yu Wang et al. XY20-3169 (KUN-L 79525). Lobothallia rubra Lun Wang & Y. Y. Zhang, sp. nov. Fungal Names: FN 572947 Fig. 7 Etymology. The specific epithet refers to the red reaction of medulla in C spot-test. Diagnosis. Thallus areolate with a lobate margin, upper surface white with pale brown tinge, pruinose; areoles raised when bearing apothecia or pycnidia; apothecia lecanorine, disc pruinose; pycnidia protruding; medulla C+ rose–red, contain gyrophoric acid. Holotype. China • Xinjiang Uygur Autonomous Region: Hejing Co., Baluntai Vil., 42°51'56.45"N, 86°27'09.16"E, alt. 2026 m, on rock, 02 July 2022, Xin-Yu Wang and Min Ai XY22-897 (holotype: KUN-L 84715). Description. Thallus closely adnate to substrate, up to 5 cm across, areolate, 1–4 mm thick, marginally lobate, 0.4–1 mm thick. Areoles (0.5–)0.8–2(– 2.4) mm wide, contiguous, angular to rounded, non-constricted at base, interspaces between areoles 0.1–0.2 mm wide. Lobes slightly broader in the apex, 2–3 mm long, 0.7–1.4 mm wide at base, 0.6–2(–2.5) mm wide at apex. Upper surface convex, cracked, pruinose, white with pale brown tinge where pruina is 225 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China thin. Upper cortex paraplectenchymatous, 20–40 μm thick, containing brown granules (soluble in K); epinecral layer composed of coarse plate-like crystals, gelatinous material, and gray-black granules (insoluble in K), 10–30 μm thick. Algal layer 75–150 μm high, discontinuous, interrupted by hyphae, 10–60 μm wide; photobiont chlorococcoid, cells 8–20 µm diameter. Medulla 0.5–1 mm high, filled with gray–black granules (generally insoluble in K). Lower cortex absent. Apothecia lecanorine, common, usually 1 per areole, (0.3–)0.6–1.5(– 2.0) mm diameter, rounded, sessile, slightly constricted at base when mature; disc slightly concave to plane, black, pruinose; apothecial margin entire, persistent, 0.1–0.25 mm wide, pruinose, concolorous with thallus. Exciple narrow, Figure 7. Lobothallia rubra (KUN-L 84715, holotype); A. Thallus; B. Apothecia; C. Pycnidia; D. Lobes; E. Section of thallus (LCB); F. Section of thallus (in K); G. Vertical section of apothecium; H. Vertical section of apothecium (LCB); I, J. Vertical section of pycnidium; K. Ascus and ascospores; L. Paraphyses (left in LCB, right in K); M. Conidia (LCB). Scale bars: 2mm (A); 0.5 mm (D); 0.2 mm (B, C); 50 µm (G, I, J); 20 µm (E, F, H); 10 µm (K, M); 5 µm (L). 226 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China up to 40 μm wide. Epithecium, hymenium and subhymenial layers combined (150–)175–225(–250) µm high; epithecium 5–20 µm high; epihymenium 20–50 μm high, filled with dark brown granules (soluble in K), N–; hymenium 100–150 μm high, hyaline, I– or I+ weakly bluish in lower part; subhymenial layers 50–100 μm high, hypothecium 40–50 μm, hyaline, I+ blue; algal layer below hypothecium discontinuous, 50–60 μm high; paraphyses simple, septate, submoniliform at the tips, with 2–3 uppermost cells shorter and broader than basal cells, 3–6 μm wide (basal cells 2–3 μm wide); asci clavate, Aspicilia-type, hyaline, 8-spored, 70–85 × 20–30 µm; ascospores simple, hyaline, broadly ellipsoid, (10.0-)11.9–13.1–14.2(-15.0) × (8.0-)8.7–9.4–10.0(-11.0) µm (n = 48), l/w ratio (1.0-)1.3–1.4–1.5(-1.8), wall ca.1 µm thick. Pycnidia common, protruding, 0.1–0.25 mm in diameter, sometimes elongated, 0.5(–0.7) × 0.2 mm (length × width); ostiole brownish to dark brown; conidia bacilliform, hyaline, 5–6 × 1 µm. Chemistry. Cortex: K+ pale yellow, P–; medulla: K+ pale yellow to orange or K–, P–, C+ rose–red, KC+ red to pale yellow; containing gyrophoric, norstictic, stictic, connorstictic and constictic acids. Habitat and distribution. Saxicolous. Currently known only from Xinjiang, China. Notes. This species is readily recognized by its C+ rose–red medullary reaction, resulting from the production of gyrophoric acid. Lobothallia semisterilis resembles this species in its white to gray thallus and protruding pycnidia, but differs in being terricolous and lacking gyrophoric and stictic acids (Zhang et al. 2020). Lobothallia pruinosa shares the characteristic white pruina on both thallus and discs, but differs in its thinner thallus, punctiform pycnidia, and the presence of norstictic and connorstictic acids as the major secondary metabolite (Kou et al. 2013). Lobothallia pulchra is similar to L. rubra in possessing white thallus and lecanorine apothecia, but differs in having epruinose discs and lacking gyrophoric acid. Additional specimens examined. China • Xinjiang Uygur Autonomous Region: Hejing Co., Baluntai Town, 42°51'56.45"N, 86°27'09.16"E, alt. 2026–2046 m, on rock, 2 July 2022, An-Cheng Yin and Han-Xiang Chen 22-72301 (KUN-L 86927) • National Highway 216, 42°51'56.39"N, 86°27'09.31"E, alt. 2100 m, on limestone, 2 July 2022, Li-Song Wang et al. 22-72886 (KUN-L 87373). Lobothallia stipitata Lun Wang & Y. Y. Zhang, sp. nov. Fungal Names: FN 572948 Fig. 8 Etymology. The epithet refers to the areoles with a constricted and stipitate base. Diagnosis. Thallus brown, thick, centrally areolate, areoles with a stipitate base, marginally lobate, margin of lobes free from the substrate with granular pruina on the upper surface; apothecia lecanorine, constricted at the base, disc with conspicuous white pruina, apothecial margin persistent, epruinose, brown to orange-brown; hymenium pale brown. Holotype. China • Xinjiang Uygur Autonomous Region: Hejing Co., along road G218, 42°54'22.90"N, 86°17'36.95"E, alt. 2228 m, on rock, 01 July 2022, Li-Song Wang et al. 22-72871 (KUN-L 87358). Description. Thallus relatively loosely attached to the substrate, up to 4–6 cm across, centrally areolate, 3–6(–8) mm thick, marginally lobate, (0.5–)1–2 mm 227 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China thick, free from the substrate. Areoles angular (0.6–1.2 mm wide) to elongated (1.5–2 × 0.6–1 mm wide), base constricted into a stipe, up to 4 mm long, interspaces between areoles 0.1–0.3 mm wide. Lobes long, simple to 2–3 branched, apices slightly wider than base, 3–5 mm long, 0.6–1.5 mm wide at base, 0.6–2.5 mm wide at apex. Upper surface flat to slightly convex, matt, brown, thinly pruinose, granular pruina present on the margin of lobes. Upper cortex paraplectenchymatous, even, 40–50 μm thick, inspersed with brownish and gray granules (soluble in K); epinecral layer gelatinous, 10–15 μm thick. Algal layer 75–150 μm high, discontinuous; photobiont chlorococcoid, cells 10–25 µm diameter. Medulla 0.6–3 mm high, filled with gray-black granules (generally insoluble in K). Lower cortex absent. Figure 8. Lobothallia stipitata (KUN-L 87358, holotype). A. Thallus; B. Profile of thallus; C. Stipe; D. Lobes; E. Apothecia; F. Pycnidium; G, H. Section of thallus (H in K reagent); I. Vertical section of apothecium (LCB); J. Vertical section of apothecium; K. Upper cortex and epinecral layer (LCB); L. Ascus; M. Paraphyses (LCB); N. Ascospores; O. Conidia. Scale bars: 2 mm (A); 1 mm (B, C); 0.5 mm (D); 0.2 mm (E, F); 100 µm (G, H); 50 µm (I, J); 20 µm (K); 10 µm (L, M, N, O). 228 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Apothecia lecanorine, common, 1–2 per areole, simple to grouped, orbicular to angular by pressure, (0.3–)0.5–1.5(–1.8) mm wide, constricted at base in maturity; disc concave to plane, brown to black, conspicuously pruinose; apothecial margin entire, persistent, epruinose, brown to orange-brown, 0.1– 0.25(–0.3) mm wide. Exciple narrow, widening to 60–80 μm in the uppermost part. Epithecium, hymenium and subhymenial layers combined 125–175 µm high; epithecium 5–15 µm high, with hyaline plate-like crystals; epihymenium 12.5–37.5 μm high, filled with dark brown granules (soluble in K), N–; hymenium 100–120 μm high, pale brown (soluble in K), I+ dark blue; subhymenial layers 50–75 μm high, hypothecium 40–50 μm, pale brown (soluble in K), I+ dark blue; algal layer below hypothecium sparse or absent; paraphyses simple, septate, submoniliform at the tips, with 2–4 uppermost cells shorter and broader than basal cells, 3–5 μm wide (basal cells ca. 2 μm wide); asci clavate, Aspicilia-type, hyaline, 8-spored, 70–85 × 18–25 µm; ascospores simple, hyaline, broadly ellipsoid to ellipsoid, (9.0-)9.8–11.4–12.9(-15.0) × (7.0-)6.8–7.6–8.4(- 11.0) µm (n = 30), l/w ratio (1.0-)1.3–1.5–1.7(-2.0), wall ca.1 µm thick. Pycnidia punctiform, plane to slightly convex, 0.1–0.25 mm diameter; ostiole brown to dark brown; conidia bacilliform, hyaline, 4–6 × 1 µm. Chemistry. Cortex K+ pale yellow, P–; medulla K+ yellow, P+ orange, C–, KC–; containing norstictic, stictic, cryptostictic and connorstictic acids. Habitat and distribution. Saxicolous. Currently only known from Xinjiang Province, China. Notes. Lobothallia praeradiosa resembles L. stipitata in having an areolate thallus with radiating lobes, a constricted apothecial base, orange-brown thallus and apothecial margins, and containing norstictic acid. However, the former differs in its epruinose disc, hyaline hymenium, and lack of stictic acid (Kou et al. 2013; Paukov et al. 2019; Ryan 2004). Lobothallia semisterilis shares narrowly elongated lobes with granular pruina on the margins, but differs in its terricolous habitat, white to gray thallus, apothecia-like pycnidia, and absence of stictic acid (Zhang et al. 2020). Additional specimens examined. China • Xinjiang Uygur Autonomous Region: Hejing Co., National Highway 218, 42°54'22.90"N, 86°17'36.95"E, alt. 2228 m, on limestone, 1 July 2022, Yan-Yun Zhang ZYY22-616 (AHUB 00470), ZYY22-617 (KUN-L 82191) • Baluntai Town, 42°54'36.56"N, 86°17'02.47"E, alt. 2171 m, on rock, 1 July 2022, Xin-Yu Wang and Min Ai XY22-887 (KUN-L 84705). Lobothallia wangii Lun Wang & Y. Y. Zhang, sp. nov. Fungal Names: FN 572949 Fig. 9 Etymology. The epithet refers to the Chinese lichenologist Li-Song Wang, who collected the type specimen. Diagnosis. Thallus areolate without marginal lobes, upper surface white, flat; apothecia common, cryptolecanorine to lecanorine, disc black, pruinose, slightly concave to plane, often shallowly fissured at maturity; epihymenium N+ weakly greenish to green, containing norstictic and connorstictic acids. Holotype. China • Xizang Autonomous Region: Shigatse Ci., Dingri Co., along road G219, 28°35'10.18"N, 87°03'43.62"E, alt. 4311 m, on rock, 06 June 2022, Li-Song Wang et al. 22-71215 (KUN-L 85839). 229 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Description. Thallus tightly adnate to the substrate, 1–4 mm thick, areolate without marginal lobes, spreading in the field up to 25 cm across, the collected specimen is fragmented, up to 4 cm across. Central areoles commonly rounded, (0.2–)0.5–1(–1.6) mm diameter, not constricted at base, interspaces between areoles 0.1–0.2 mm wide; marginal areoles rounded to slightly elongated, 0.8–1.2 × 0.5–1.2 mm. Upper surface flat, white and pruinose. Upper cortex paraplectenchymatous, 30–50 μm thick, inspersed with pale gray granules (soluble in K), uppermost part filled with dark brown granules (partly soluble in K), 20–30 μm thick; epinecral layer gray dark, 20– 50 μm thick. Algal layer 75–100 μm thick, mostly continuous, occasionally Figure 9. Lobothallia wangii (KUN-L 85839, holotype). A. Habit; B, C. Apothecia; D. Section of thallus (below in K reagent); E. Upper cortex and epinecral layer (LCB); F. Section of apothecium; G. Section of apothecium (in N reagent); H. Section of pycnidium; I. Ascus and ascospores; J. Paraphyses (LCB); K. Conidia. Scale bars: 1 mm (A); 0.2 mm (B, C); 100 µm (D, G); 20 µm (E, F, H); 10 µm (I); 5 µm (J, K). 236 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China gray to orange–brown, apothecial margin gray to orangish–brown ....... ...........................................................................................L. praeradiosa 16 Central thallus thin, less than 2 mm; lobes plane ......................................17 – Central thallus thick, more than 2 mm; lobes slightly convex ..................20 17 Apothecia not exceeding 3 per areole .......................................................18 – Apothecia commonly more than 3 per areole ........................ L. brachyloba 18 Disc black or dark brown with white pruina ...............................................19 – Disc brown with orange-gray pruina ....................................... L. complanata 19 Apothecia cryptolecanorine, 0.3–0.8 mm, apothecial margin at the same level with disc ...................................................................L. pseudopruinosa – Apothecia cryptolecanorine to lecanorine, 0.7–1.2 mm, apothecial margin somewhat higher than the disc ................................................... L. pruinosa 20 Pycnidia sparse, punctiform or slightly protruding at maturity ................21 – Pycnidia dense per areole, spot-like to elongate and curved ....................... ..............................................................................................L. polypycnidiata 21 Marginal lobes parallel, disc pruinose.............................................L. hedinii – Marginal lobes non-parallel, disc epruinose ................................. L. radiosa Discussion Species diversity Thirty-nine species of Lobothallia are accepted worldwide in this study. Among them, twenty-two are present in China, including eight new species (Figs 2–9), one newly recorded species (Fig. 10), and thirteen known species, ten of which are studied here (Figs 11, 12) (Magnusson 1940, 1944; Mamut et al. 2010, 2012; Kou et al. 2013; Paukov et al. 2019; Wheeler et al. 2024; Zhang et al. 2020, 2024a). This research significantly increases the species diversity of Lobothallia and shows that China is a distribution center of this genus, especially in the north, northwest and southwest. The Qinghai-Tibet Plateau region emerges as a significant diversity hotspot for the genus, exhibiting a substantially higher species richness compared to other areas. Unfortunately, no specimens from northeastern China were examined, but comprehensive future sampling may further enrich the species diversity of Lobothallia in the country. Morphological and microscopical characters In Megasporaceae, Lobothallia is characterized by relatively small ascospores that rarely exceed 18 μm in length, and short conidia, 3–10 μm long (Nordin et al. 2010; Kou et al. 2013; Paukov et al. 2019; Zulfiqar et al. 2022). Additional morpho-anatomical characters include: a paraplectenchymatous upper cortex, a discontinuous algal layer; an Nor rarely N+ slightly greenish epihymenium (vs. N+ emerald green in other genera of Megasporaceae) (Paukov et al. 2019); simple to rarely anastomosed paraphyses with swollen, submoniliform to moniliform tips; and a subhypothecial algal layer (Ryan 2004; Kou et al. 2013; Paukov et al. 2019), which may be poorly visible in some cases (Roux et al. 2016). Although Antidea brucei (Owe-Larss. & A. Nordin) T. B. Wheeler, Aspilidea myrinii (Fr.) Hafellner and A. subadunans (Vain.) T. B. Wheeler, J. W. McCarthy & Fryday possess ascospores and conidia of similar size to the Lobothallia species, they 237 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China are phylogenetically distant (Hafellner and Türk 2001; Owe-Larsson et al. 2007; Wheeler et al. 2024). Similarly, species of Teuvoa share anatomical similarities with Lobothallia, such as paraphyses structure, ascospore and conidia size. However, Teuvoa differs in having non-lobate thalli, lacking secondary metabolites, and its corticolous or terricolous habitat (Rico et al. 2007; Sohrabi et al. 2013b). The traditional key morphological feature for defining the genus Lobothallia— the presence of marginal lobes—exhibits transitional variation across the genus (Hafellner 1991; Ryan 2004). Although it is therefore not a reliable character for generic delimitation, it remains valuable in species-level identification (Paukov et al. 2019). Currently, approximately one-third of the species in Lobothallia lack or exhibit indistinct lobes, viz. L. benzilanensis, L. cernohorskyana, L. chadefaudiana, L. cheresina, L. controversa, L. determinata, L. elobulata, L. gangwondoana, L. lacteola, L. peltastictoides, L. recedens, L. subdiffracta and L. wangii (Magnusson 1940, 1944; Roux et al. 2016; Paukov et al. 2019; Kondratyuk et al. 2020; Zulfiqar et al. 2022; Wheeler et al. 2024). The length of lobes is a diagnostic character at species level (Paukov et al. 2019; Zulfiqar et al. 2023), ranging from short (< 2 mm) to relatively long (up to 5 mm). Most species of Lobothallia have an areolate thallus (at least in the center), with the following exceptions: L. epiadelpha, which is centrally squamulose; L. zogtii, which is centrally squamulose-areolate Figure 12. Other Lobothallia species from China examined in this study. A. L. alphoplaca (22-72816, KUN-L); B. L. crassimarginata (20122418B, SDNU); C. L. crenulata (ZYY22-301, KUN-L, holotype); D. L. lobulata (ZYY22-819, KUN-L, holotype); E. L. praeradiosa (22-71752, KUN-L); F. L. pruinosa (22-73170, KUN-L); G. L. semisterilis (ZYY22-719, KUN-L); H. L. subdiffracta (ZYY22-628, KUN-L); I. L. radiosa (22-72256, KUN-L). Scales bars: 1 mm (B–I). 238 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China (Paukov et al. 2019); and L. benzilanensis and L. gangwondoana, which contain more or less distant simple areoles (Kondratyuk et al. 2020). Species delimitation in Lobothallia is based on the integration of multiple morphological and chemical characters. Key thallus features for identification include thallus thickness (varying from < 2 mm to 8 mm), areoles size, and the presence of pruina. Apothecial type (e.g., cryptolecanorine, cryptolecanorine to lecanorine, lecanorine), disc color and pruina, and thalline margin width and development are also frequently employed for species delimitation. Additional phenotypic characters further support species delimitation, including habitat conditions and substrate type; thallus attachment, growth form and diameter, the width and depth of inter-areolar cracks; the presence or absence of lobules and of basal constriction in areoles; smooth or rimose upper surface; and the cortex and medulla thickness. Reproductive traits include the number of apothecia and pycnidia per areole; the presence or absence of a constricted apothecial base; disc shape (concave, plane, convex) and texture (shiny/matt, with or without fissures); epihymenium reactions (N− or N+ light green), hymenium and hypothecium colour (hyaline or pale brown), ascospore and conidial sizes (Magnusson 1940, 1944; Ryan 2004; Kou et al. 2013; Nimis 2016; Roux et al. 2016; Paukov et al. 2019; Kondratyuk et al. 2020; Zhang et al. 2020, 2024a; Ashraf et al. 2022; Zulfiqar et al. 2022, 2023). Secondary metabolites Species of Lobothallia can be categorized into two groups based on the presence or absence of secondary metabolites. Among species producing secondary metabolites, norstictic acid is the predominant and most frequently encountered compound (Ryan 2004; Roux 2012; Roux et al. 2016; Kou et al. 2013; Paukov et al. 2019; Kondratyuk et al. 2020; Zhang et al. 2020; Ashraf et al. 2022; Zulfiqar et al. 2022, 2023). Other metabolites include stictic acid (Kou et al. 2013; Moon and Kashiwadani 2009; Roux et al. 2016; Paukov et al. 2019), constictic acid (Ryan 2004; Kou et al. 2013), connorstictic acid (Paukov et al. 2019), salazinic acid (Ryan 2004), and terpenes (Galloway and Ledingham 2012; Roux et al. 2016). All newly described species in this study contain norstictic acid. Additional secondary metabolites detected include stictic acid in Lobothallia pulchra, L. rubra, and L. stipitata; cryptostictic acid (recorded for the first time in Lobothallia) in L. benzilanensis, L. pulchra, and L. stipitata; gyrophoric acid exclusively in L. rubra; and connorstictic acid, which frequently occurred as an accessory acid in all new species. Furthermore, thin-layer chromatography (TLC) analysis of 11 known species (10 previously reported and one newly recorded in China) indicated that all species, except L. crenulata, L. lobulata and L. subdiffracta, contain norstictic acid as the major secondary metabolite (Kou et al. 2013; Paukov et al. 2019; Zhang et al. 2024a). Stictic acid was detected in L. crassimarginata; connorstictic acid was identified in the species of L. alphoplaca, L. crassimarginata, L. hadinii, L. praeradiosa, L. pruinosa, and L. radiosa. Phylogeny Although the phylogenetic topology of Megasporaceae from our three-locus dataset is largely congruent with the five-locus results of Wheeler et al. (2024), 239 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China and both place Lobothallia as sister to Teuvoa, the statistical support for this relationship is considerably lower in our analysis. Consequently, the relationship between Lobothallia and Teuvoa remains uncertain and warrants further investigation. Our results confirm that Lobothallia comprises three primary clades (Clades I–III), consistent with previous studies (Kou et al. 2013; Paukov et al. 2019; Zulfiqar et al. 2022, 2023; Zhang et al. 2024a). However, the evolutionary relationships among these three clades remain unresolved due to weak supports at the backbone nodes (Fig. 1). The above results, to some extent, support the assumption of Wheeler et al. (2024)—if Teuvoa is accepted at the genus level, then a case could be made for erecting additional genera within Lobothallia. Whether different clades or subclades of Lobothallia warrant recognition as separate genera in the future will depend on comprehensive phylogenetic and morphological studies within the family Megasporaceae. Furthermore, this study identified six Lobothallia species for which molecular data are unavailable: L. chadefaudiana, L. gangwondoana, L. kuminovae, L. lacteola, L. platycarpa, and L. zogtii. Their phylogenetic positions remain unresolved. Lobothallia chadefaudiana is morphologically most similar to L. cernohorskyana, differing primarily in the absence of secondary metabolites (Clauzade and Vězda 1970; Roux 1977); its phylogenetic position is tentatively inferred to belong to Clade III. Lobothallia gangwondoana, characterized by a smooth, slightly brownish-gray or cacao gray thallus with indistinct marginal lobes and distant simple areoles (Kondratyuk et al. 2020), is tentatively placed near L. crassimarginata or L. benzilanensis within Clade I. Lobothallia kuminovae, reported from Siberian Russia, is morphologically more or less identical to L. praeradiosa based on its original description (Sedelnikova 1982). Further specimen examination is required to determine whether these two taxa are conspecific. Lobothallia lacteola, with its circular outline, plicate margins, and lack of long and straight radial cracks (Paukov et al. 2019), is tentatively inferred to belong to Clade III. Lobothallia platycarpa morphologically resembles the subclade comprising the three Pakistan species, L. pulvinata, L. pakistanica and L. iqbalii. Lobothallia zogtii, originally described from Xinjiang Province, China, has a brown thallus, short marginal lobes, lecanorine apothecia, and stictic acid (Paukov et al. 2019); its phylogenetic position is tentatively inferred to be within Clade I. Acknowledgements The authors thank Dr. Lulu Zhang from Shandong Normal University for the loan of specimens and permitting DNA extraction; Dr. Alexander G. Paukov, curator of Ural Federal University, for providing the high-quality morphological photos of Lobothallia zogtii. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. 240 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Use of AI No use of AI was reported. Funding This study was funded by the National Natural Science Foundation of China (32400005, 32570250), Yunnan Young & Elite Talents Project (YNWR-QNBJ-2020-224), Yunnan Fundamental Research Projects (202401AT070196), and the Second Tibetan Plateau Scientific Expedition and Research Program (STEP) (2019QZKK0503). Author contributions Conceptualization: LSW, XYW, and YYZ; Data curation: LW and YYZ; Formal analysis: LW and YSF; Funding acquisition: LSW, XYW, and YYZ; Investigation: LW and YSF; Methodology: LW and YSF; Project administration: LSW, XYW, and YYZ; Resources: LW and LSW; Software: LW and YSF; Supervision: XYW and YYZ; Validation: LW and YYZ; Visualization: LW; Writing – original draft: LW and YYZ; Review, and editing: XYW and YYZ. Author ORCIDs Xin-Yu Wang https://orcid.org/0000-0003-2166-6111 Yan-Yun Zhang https://orcid.org/0000-0002-0902-5066 Data availability All of the data that support the findings of this study are available in the main text. References Ashraf A, Habib K, Khalid AN (2022) A new pruinose lichen species in genus Lobothallia (Megasporaceae, lichen forming Ascomycota) from Pakistan. Acta Botanica Brasilica 36:e2021abb0225. https://doi.org/10.1590/0102-33062021abb0225 Bungartz F (2002) Morphology and Anatomy of the Fertile Structures. In: Nash TH III, Ryan BD, Gries C, Bungartz F (Eds) Lichen Flora of the Greater Sonoran Desert Region 1. Lichens Unlimited, Arizona State University, Arizona, 24–34. https://doi. org/10.1639/0007-2745(2002)105[0725:LFOTGS]2.0.CO;2 Carstens BC, Pelletier TA, Reid NM, Satler JD (2013) How to fail at species delimitation. Molecular Ecology 22(17): 4369–4383. https://doi.org/10.1111/mec.12413 Clauzade G, Roux C (1984) Les genres Aspicilia Massal. et Bellemerea Hafellner et Roux. Bulletin de la Société Botanique de Centre-Ouest, Nouvelle Série 15:127–141. https:// www.researchgate.net/publication/330508116 Clauzade G, Vězda A (1970) Lecanora cernohorskyana Clauzade et Vězda sp. n. Preslia 42: 215–219. https://www.preslia.cz/article/10804 Elix JA (2014) Catalogue of Standardized Chromatographic Data and Biosynthetic Relationships for Lichen Substances. Australian National University, Canberra. https:// www.cpbr.gov.au/abrs/lichenlist/TLC%20Cat%206MC.pdf Galloway DJ, Ledingham J (2012) Additional lichen records from New Zealand 48. Australasian Lichenology 70: 14–17. https://www.anbg.gov.au/abrs/lichenlist/AL_70.pdf Gardes M, Bruns TD (1993) ITS primers with enhanced specificity for Basidiomycetes-application to the identification of mycorrhizae and rusts. Molecular Ecology 2(2): 113–118. https://doi.org/10.1111/j.1365-294X.1993.tb00005.x Hafellner J (1991) Die Gattung Aspicilia, ihre Ableitungen nebst Bemerkungen über cryptolecanorine Ascocarporganisation bei anderen Genera der Lecanorales 241 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China (Ascomycetes lichenisati). Acta Botanica Malacitana 16: 133–140. https://doi. org/10.24310/abm.v16i.9153 Hafellner J, Türk R (2001) Die lichenisierten Pilze Österreichs – eine Checkliste der bisher nachgewiesenen Arten mit Verbreitungsangaben. Stapfia 76: 1–167. https://www. researchgate.net/publication/309282880 Haji Moniri M, Gromakova AB, Lőkös L, Kondratyuk SY (2017) New members of the Megasporaceae (Pertusariales,lichen-forming Ascomycota): Megaspora iranica spec. nova and Oxneriaria gen. nova. Acta Botanica Hungarica 59(3–4): 343–370. https:// doi.org/10.1556/034.59.2017.3-4.5 Hyde KD, Noorabadi MT, Thiyagaraja V, He MQ, Johnston PR (2024) The 2024 Outline of Fungi and fungus-like taxa. Mycosphere 15(1): 5146–6239. https://doi.org/10.5943/ mycosphere/15/1/25 Ivanovich C, Weber L, Palice Z, Hollinger J, Otte V, Sohrabi M, Sheehy S, Printzen C (2025) A taxonomic revision of the lichen genus Lecanoropsis (Lecanoraceae). Phytotaxa 695(1): 1–56. https://doi.org/10.11646/phytotaxa.695.1.1 Kalyaanamoorthy S, Minh BQ, Wong TK, Haeseler Av, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. https://doi.org/10.1038/nmeth.4285 Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010 Katoh K, Rozewicki J, Yamada KD (2019) MAFFT online service: Multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics 20(4): 1160–1166. https://doi.org/10.1093/bib/bbx108 Kondratyuk SY, Upreti DK, Mishra GK, Nayaka KK, Orlov OO, Kondratiuk AS, Lőkös L, Farkas E, Woo J, Hur J (2020) New and noteworthy lichen-forming and lichenicolous fungi 10. Acta Botanica Hungarica 62(1–2): 81–84. https://doi. org/10.1556/034.62.2020.1-2.6 Kou XR, Li SX, Ren Q (2013) Three new species and one new record of Lobothallia from China. Mycotaxon 123: 241–249. https://doi.org/10.5248/123.241 Kumar S, Stecher G, Tamura K (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33: 1870. https:// doi.org/10.1093/molbev/msw054 Larena I, Salazar O, González V, Julián MC, Rubio V (1999) Design of a primer for ribosomal DNA internal transcribed spacer with enhanced specificity for Ascomycetes. Journal of Biotechnology 75(2–3): 187–194. https://doi.org/10.1016/S01681656(99)00154-6 Letunic I, Bork P (2021) Interactive Tree of Life (iTOL) v5: An online tool for phylogenetic tree display and annotation. Nucleic Acids Research 49(W1): W293–W296. https:// doi.org/10.1093/nar/gkab301 Lücking R, Hodkinson BP, Leavitt SD (2017) The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota–Approaching one thousand genera. The Bryologist 119(4): 361–416. https://doi.org/10.1639/0007-2745-119.4.361 Magnusson AH (1940) Lichens from central Asia I. In: reports from the scientific expedition to the NorthWestern Provinces of China under the leadership of Dr. Sven Hedin (the Sino-Swedish expedition). Publication 13, XI, Botany. Aktiebolaget Thule, Stockholm, 1–168. Magnusson AH (1944) Lichens from central Asia II. Reports from the scientific expedition to the NorthWestern Provinces of China under the leadership of Dr. Sven Hedin 242 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China (the Sino-Swedish expedition). Publication 22, XI, Botany. Aktiebolaget Thule, Stockholm, 1–68. Mamut R, Abdulla A, Abbas A (2010) A newly recorded lichen genus (Lobothallia) and species from China. Journal of Wuhan Botanical Research 28(3): 362–364. https:// doi.org/10.3724/SP.J.1142.2010.30362 Mamut R, Suo FY, Abbas A (2012) Lobothallia radiosa: A new record of lichen genus Lobothallia from China. Zhiwu Kexue Xuebao 30(3): 305–308. https://doi.org/10.3724/ SP.J.1142.2012.30305 Mayrhofer H, Denchev CM, Stoykov DY, Nikolova SO (2005) Catalogue of the lichenized and lichenicolous fungi in Bulgaria. Mycologia Balcanica 2(1): 18. https://doi. org/10.5281/zenodo.2546852 Moon KH, Kashiwadani H (2009) Lobothallia alphoplaca (Wahlenb.) Hafellner (Megasporaceae) found in Korea. Shokubutsu Kenkyu Zasshi 84: 303–305. https://cir.nii.ac.jp/ crid/1523669553271969152 Müller AJ (1880) Enumeration lichenum Aegyptiacum. Revue mycologique, Toulouse 2(2): 75. Nascimbene J, Nimis PL, Klüßendorf J, Thüs H (2023) Freshwater lichens, including new species in the genera Verrucaria, Placopyrenium and Circinaria, associated with Lobothallia hydrocharis (Poelt & Nimis) Sohrabi & Nimis from Watercourses of Sardinia. Journal of Fungi 9(3): 380. https://doi.org/10.3390/jof9030380 Nguyen L-T, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32: 268–274. https://doi.org/10.1093/molbev/msu300 Nimis PL (2016) The lichens of Italy, A second annotated catalogue. Trieste: EUT-Edizioni Università di Trieste, Italy, 288–290. http://hdl.handle.net/10077/21022 Nordin A, Savić S, Tibell L (2010) Phylogeny and taxonomy of Aspicilia and Megasporaceae. Mycologia 102(6): 1339–1349. https://doi.org/10.3852/09-266 Orange A, James PW, White FJ (2001) Microchemical methods for the identification of lichens. British Lichen Society, London. Owe-Larsson B, Nordin A, Tibell L (2007) Aspicilia. In: Nash TH III, Gries C, Bungartz F (Eds) Lichen Flora of the Greater Sonoran Desert Region 3. Lichens Unlimited, Arizona State University, Arizona, 61–108. Paukov AG, Davydov EA, Nordin A, Roux C, Şenkardeler A, Sohrabi M, Vondrák J, Frolov IV, Teptina AY, Shiryaeva AS (2019) Three new species, new combinations and a key to known species of Lobothallia (Megasporaceae). Lichenologist 51(4): 301–322. https://doi.org/10.1017/S0024282919000264 Paukov AG, Davydov EA, Yakovchenko L, Ren Q, Sohrabi M (2024) Atrostelia—A new genus in Megasporaceae (Pertusariales, lichenized Ascomycota). Mycological Progress 23(1): 1–9. https://doi.org/10.21203/rs.3.rs-3687993/v1 Rehner SA, Samuels GJ (1994) Taxonomy and phylogeny of Gliocladium analysed from nuclear large subunit ribosomal DNA sequences. Mycological Research 98(6): 625– 634. https://doi.org/10.1016/S0953-7562(09)80409-7 Rico VJ, Arago’n G, Esnault J (2007) Aspicilia uxoris, an epiphytic species from Algeria, Morocco and Spain. Lichenologist 39: 109–119. https://doi.org/10.1017/ S0024282907006524 Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model selection across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029 243 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Roux C (1977) Aspicilia chadefaudiana Cl. Roux, sp. nov. et remarques sur le genre Aspicilia. Revue Bryologique et Lichénologique 43(2):159–172(57–70). https://www. researchgate.net/publication/330117271 Roux C (2012) Liste des lichens et champignons lichénicoles de France. Bulletin de la Société linnéenne de Provence Numéro spécial 16: 1–220. Roux C, Nordin A, Bertrand M (2016) Lobothallia controversa Cl. Roux & A. Nordin sp. nov., correspondant au Lecanora farinosa sensu Nyl. non (Flörke) Nyl. Herzogia 29(2): 586–595. https://doi.org/10.13158/heia.29.2.2016.586 Ryan BD (2004) Lobothallia. In: Nash III TH, Ryan BD, Diederich P, Gries C, Bungartz F (Eds) Lichen Flora of the Greater Sonoran Desert Region 2. Lichens Unlimited, Arizona State University, Arizona, 352–357. Schmitt I, Yamamoto Y, Lumbsch HT (2006) Phylogeny of Pertusariales (Ascomycotina): Resurrection of Ochrolechiaceae and new circumscription of Megasporaceae. The Journal of the Hattori Botanical Laboratory 100: 753–764. https://doi.org/10.18968/ jhbl.100.0_753 Sedelnikova NV (1982) Species nova lichenis generis Placodium (Ach.) Müll. Arg. Novosti Sistematiki Nizshikh Rasteniy 19: 165–166. Sedelnikova NV, Sedelnikov VP (2018) An appreciation of the participation of lichen species at high-mountain phytocoehosis of Republic Tuva. Растительный мир Азиатской России, Nº 3(31): 9–35. Sipman HJ, Aptroot A (2020) Ikaeria serusiauxii, a new Caloplaca-like lichen from Macaronesia and mainland Portugal, with a lichen checklist for Porto Santo. Plant and Fungal Systematics 65(1): 126. https://doi.org/10.35535/pfsyst-2020-0006 Sohrabi M, Stenroos S, Myllys L, Søchting U, Ahti T, Hyvönen J (2013a) Phylogeny and taxonomy of the ‘manna lichens’. Mycological Progress 12: 231–269. https://doi. org/10.1007/s11557-012-0830-1 Sohrabi M, Leavitt SD, Rico VJ, Halici MG, Shrestha G, Stenroos S (2013b) Teuvoa, a new lichen genus in Megasporaceae (Ascomycota: Pertusariales), including Teuvoa junipericola sp. nov. Lichenologist 45: 347–360. https://doi.org/10.1017/ S0024282913000108 Trifinopoulos J, Nguyen LT, von Haeseler A, Minh BQ (2016) W-IQ-TREE: A fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44(W1): W232–W235. https://doi.org/10.1093/nar/gkw256 Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172(8): 4238–4246. https://doi.org/10.1128/jb.172.8.4238-4246.1990 Wei JC (2020) The Enumeration of Lichenized Fungi in China. China Forestry Publishing House, Beijing. Wheeler T, McCarthy J, Owe-Larsson B, Fryday A (2024) Taxonomic innovations in Megasporaceae (lichenized Ascomycota, Pertusariales): Antidea, a new genus for Aspicilia brucei; two new species of Aspicilia, and new combinations in Aspilidea and Lobothallia. Lichenologist 56(5): 273–286. https://doi.org/10.1017/ S0024282924000239 Xiang CY, Gao F, Jakovlić I, Lei HP, Hu Y, Zhang H, Zou H, Wang GT, Zhang D (2023) Using PhyloSuite for molecular phylogeny and tree‐based analyses. iMeta 2(1): e87. https:// doi.org/10.1002/imt2.87 Zakeri Z, Divakar PK, Otte V (2017) Taxonomy and phylogeny of Aspiciliella, a resurrected genus of Megasporaceae, including the new species A. portosantana. Herzogia 30(1): 166–176. https://doi.org/10.13158/heia.30.1.2017.166 244 MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554 Lun Wang et al.: An overview of the genus Lobothallia in China Zhang YY, Wang XY, Li LJ, Printzen C, Timdal E, Niu DL,Yin AC, Wang SQ, Wang LS (2020) Squamarina (lichenised fungi) species described from China belong to at least three unrelated genera. MycoKeys 66: 135–157. https://doi.org/10.3897/mycokeys.66.39057 Zhang YY, Wang L, Yu XM, Cheng S, Liu JL, Wang XY (2024a) Three new taxa of the lichen genus Lobothallia (Megasporaceae, Ascomycota) from China. MycoKeys 108: 351–369. https://doi.org/10.3897/mycokeys.108.126994 Zhang YY, Yin YJ, Wang L, Printzen C, Wang LS, Wang XY (2024b) Two new species of Rhizoplaca (Lecanoraceae) from Southwest China. MycoKeys 101: 233–248. https:// doi.org/10.3897/mycokeys.101.115678 Zoller S, Scheidegger C, Sperisen C (1999) PCR primers for the amplification of mitochondrial small subunit ribosomal DNA of lichen-forming ascomycetes. Lichenologist 31(5): 511–516. https://doi.org/10.1006/lich.1999.0220 Zulfiqar R, Razzaq F, Afshan NUS, Fayyaz I, Habib K, Khalid AN, Paukov AG (2022) Three new species of Lobothallia (Megasporaceae, Pertusariales, Ascomycota) from Pakistan and a new combination in the genus. Mycological Progress 21: 80. https://doi. org/10.1007/s11557-022-01830-z Zulfiqar R, Razzaq F, Iqbal MS, Khalid AN (2023) A new species of Lobothallia (Megasporaceae, Pertusariales, Ascomycota) from Pakistan. Plant Systematics and Evolution 309: 35. https://doi.org/10.1007/s00606-023-01874-y