Immersaria Rambold & Pietschm., Bibliotheca Lichenologica 34: 239 (1989)
Type: Immersaria athroocarpa (Ach.) Rambold & Pietschm., in Rambold, Biblthca Lichenol. 34: 240 (1989).
Description
Thallus crustose, yellow brown, red brown, orange brown or brown, sometimes with rusty colored, continuous, areolae irregular or tend to rectangular, with wax glossy, margin pruinose, areolae sometimes aggregate with black prothallus and forming bigger areolae, prothallus distinct at the margin of thallus or absent. Upper cortex with orange pigmented, epinecral layer colorless, algal layer continuous, medulla filled with gray granules, lower cortex absent. Apothecia lecideine-type, immersed, sometimes aggregate, round or irregular; disc black, flat, less concave, margin reduced, sometimes slightly raised, often poorly developed in section, pruinose or not. Exciple almost absent, sometimes developed, brown. Hymenium colorless, paraphyses simple, rare branch, anastomosing or not; epihymenium brown, green or brown green, without epinecral layer; subhymenium colorless, sometimes pale brown; hypothecium pale brown to brown. Asci Porpidia-type, cylindrical, eight-spored; ascospores ellipsoid, halonate, non-amyloid. Pycnidia present or not, immersed, liner or stellate, black, margin pruinose; conidia bacilliform.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. Confluentic acid, often accompanied with 2’-O-methylmicophyllinic acid, planaic acid or no substances detected by TLC. The compound planaic acid is newly found in this genus.
Ecology and distribution
Growing on bare rock, sandstone or granite, from elevations of 3800 to 4500 m in the alpine zone of west China and elevations of 1200 to 1900 m in the steppe of north China. Worldwide distribution.
Remarks
Species with lecanorine apothecia were previously included in Immersaria (Calatayud and Rambold 1998, Valadbeigi et al. 2011), but the five-loci phylogenetic analysis excluded these species from Immersaria. This exclusion entails a restricted concept of the genus. Immersaria is defined by its orange brown, yellow brown, sometimes rusty colored thallus, the amyloid medulla, the wax glossy areolae with a pruinose margin, the black immersed lecideine apothecia with a reduced margin, the brown epihymenium and the Porpidia-type asci with eight halonate and non-amyloid ascospores. The members of this genus occur in alpine or steppe habitats.
Species of Sporastatia A. Massal. have frequently been misidentified as members of Immersaria because of field observations of their glossy areolae and the immersed lecideine apothecia. However, they are characterized by multispora and their radiated, yellow brown thallus. Also, Miriquidica Hertel & Rambold often resembles Immersaria by its glossy areolae and the lecideine apothecia, but differs in its black brown thallus, its Lecanora-type asci with non-halonate ascospores and often containing miriquidic acid. The immersed apothecia of Immersaria may resemble Aspicilia A. Massal. and Acarospora A. Massal., but Aspicilia has a white or gray thallus, the Aspicilia-type asci with non-halonate ascospores; Acarospora has multispora and a convex thallus.
Although four known species, Immersaria carbonoidea (J.W. Thomson) Esnault & Cl. Roux, I. fuliginosa Fryday, I. olivacea Calat. & Rambold and I. usbekica, currently lack molecular data, we temporarily conserved them within the Immersaria due to their morphology, which corresponds to that of Immersaria. Morphological comparisons were based on high-resolution photographs of type materials and the original descriptions.
Immersaria athroocarpa (Ach.) Rambold & Pietschm., in Rambold, Biblthca Lichenol. 34: 240 (1989). — Fig. 4a–e
Type
Sweden [no locality, no date, no collector], H9508237 (H-Ach – lectotype!). High-resolution photographs seen.
Description
Thallus areolate, yellow brown, orange brown, continuous, areolae 0.25–1.0 mm in diam., often convex, regular polygon, tends to be squamalose in the margin, epruinose; margin pruinose; prothallus black, not distinct, sometimes absent. Upper cortex 32.5 µm thick, yellow brown granules pigmented; epinecral layer 7.5 µm thick; algal layer 82.5 µm thick, cells 8.0–10.0 × 7.5 µm in diam., round; lower cortex absent. Apothecia frequent, densely crowded, immersed, 0.3–1.3 mm in diam.; disc black, rare pruinose, flat, epruinose; margin reduced. Exciple sometimes developed, 25.0 µm wide, brown. Hymenium 107.5–112.5 µm thick, colorless; paraphyses 1.0–2.0 µm wide, branched, not anastomosing; epihymenium 25.0 µm thick, brown, rare green; subhymenium 90.0 µm thick, colorless; hypothecium pale brown to brown, sometimes absent. Asci Porpidia-type, cylindrical, eight-spored; ascospores 17.5–20.0 × 10.0 µm in diam., ellipsoid, halo. Pycnidia immersed, stellate, black, margin pruinose; conidia 7.5–10.0 × 1.0 µm in diam., bacilliform.
Chemistry: Thallus K–, C–. Medulla I+, K–, C–. Ⅰ: Confluentic acid. Ⅱ: Unknown substance.
Ecology and distribution
Growing on granite in arid and semi-arid steppe habitats at elevations of 1217–1915 m. Worldwide distribution. This species is known from Inner Mongolia and Mt. Changbai (Hertel and Zhao 1982) of China.
Remarks
The lectotype grows on siliceous rock and contains several intact apothecia. The materials from Inner Mongolia are identical with the lectotype based on comparisons with high-resolution photographs and the description given by Hertel (1977). It is therefore treated as I. athroocarpa at present. Some Inner Mongolian materials contain an unknown substance but form a well–supported clade with other materials. I. athroocarpa is characterised by the convex, yellow brown areolae and the big sizes of ascospores. In this genus, only this species has ascospores up to 20.0 µm long.
Immersaria usberkica is similar to I. athroocarpa in its brown thallus and dense apothecia, but differs in its flat areolae, the brown epihymenium and the presence of confluentic acid and gyrophoric acid. By comparing with high-resolution photographs and the original descriptions (Hertel 1977) of I. usberkica, we discovered that previous reports of this species from China (Zhang et al. 2015) were due to misidentification. It is known from Algeria, Iran, Spain and the USSR (Barbero et al. 1990).
Specimens examined (SDNU). CHINA, Inner Mongolia, Chifeng City, Balin Youqi, Hongshilazi, 1403.2 m elev., 44°13′N, 118°44′E, on rock, 2019, Ling Hu et al. SDNU20190035; Rongshen, Wangfengou, 1217.4 m elev., 44°16′N, 118°22′E, Ling Hu et al. SDNU20190140, SDNU20190143; Erlinba, 1915.2 m elev., 44°26′N, 118°41′E, Ling Hu et al. SDNU20190227.
Immersaria aurantia C.M. Xie & Li S. Wang, sp. nov. — Fungal Names FN839738; Fig. 4f–j
Etymology
The epithet “aurantia” refers to the orange thallus.
Type
CHINA, Tibet, Sajia Co., Mula village, 4752 m elev., 28°40′N, 88°45′E, on rock, 28 Jun. 2019, Xin Yu Wang et al. XY19-1814 (KUN–holotype).
Description
Thallus areolate, orange, dark orange, pale orange to pale red brown, continuous, areolae 0.75–1.25 mm in diam., flat, epruinose, irregular, margin thin pruinose; prothallus not seen. Upper cortex 25.0–45.0 µm thick, orange granules pigmented; epinecral layer 12.5–37.5–62.5 µm thick, uneven; algal layer 50.0–92.5 µm thick, cells 5.0–15.0 × 5.0–10.0 µm in diam., round; lower cortex absent. Apothecia frequent, scattered, immersed or isolated from areolae, 0.3–1.3 mm in diam.; disc black, flat or concave, sometimes pruinose; margin reduced. Exciple sometimes developed, 30.0 µm wide, brown. Hymenium 55.0–82.5 µm thick, colorless; paraphyses 2.0–2.5 µm wide, only branched and anastomosing at apex; epihymenium 20.0 thick, green or green brown; subhymenium colorless, not distinct or absent; hypothecium brown. Asci Porpidia-type, cylindrical, eight-spored; ascospores 8.0–15.0 × 5.0–7.5 µm in diam., ellipsoid, halonate. Pycnidia rare, immersed, oblate, black, margin white; conidia 7.5 × 1.0 µm in diam., bacilliform.
Chemistry: Thallus K–, C–. Medulla I+, K–, C–. Ⅰ: Confluentic acid, often accompanied with 2’-O-methylmicrophylinic acid. Ⅱ: Planaic acid. Ⅲ: none (rare).
Ecology and distribution
Growing on rock at elevations of 3931–4229 m in the alpine zone. This species is known from Qinghai, Sichuan Province and Tibet of China.
Remarks: Immersaria aurantia is characterised by its distinct orange, irregular areolae and the mostly green epihymenium. I. athroocarpa and I. venusta are similar to I. aurantia, but I. athroocarpa differs in the convex, regular polygon areolae and the more crowded apothecia; I. venusta differs in having the yellow brown, often rusty, cracked areolae and the flat apothecia. Besides, confluentic acid and planiaic acid do not appear simultaneously in I. aurantia, whereas I. venusta always contains both of these two compounds together. The orange thallus of I. aurantia may resemble I. fuliginosa, but the latter differs in the black margin of areolae and the distinct thalliconidia.
Specimens examined (KUN). CHINA, Qinghai Province, Banma Co., 3933 m elev., 32°40′N, 100°48′E, on rock, 2020, Li-Song Wang et al. 20-66886, 3932 m elev., Li-Song Wang et al. 20-66897; Jiuzhi Co., Baiyu village, 4285 m elev., 33°14′N, 100°58′E, Li-Song Wang et al. 20-67809. Sichuan Province: Rangtang Co., Mt. Haizi, 4223 m elev., 32°20′N, 101°25′E, on rock, 2020, Li-Song Wang et al. 20-66701, 4229 m elev., Li-Song Wang et al. 20-66693, 4217 m elev., Li-Song Wang et al. 20-66680, 4221 m elev., Li-Song Wang et al. 20-66692. Tibet, Changdu City, Mangkang Co., Luoni village, 4145 m elev., 29°56′N, 98°33′E, on rock, 2020, Li-Song Wang et al. 20-69091, 4138 m elev., Li-Song Wang et al. 20-69091, 20-69094; Gatuo town, 29°39′N, 98°35′E, 3831 m elev., Li-Song Wang et al. 20-69114, 3850 m elev., Li-Song Wang et al. 20-69122; Gongga Co., Jiangtang town, 29°12′N, 90°38′E, 2019.7.23, 4560 m elev., Xin-Yu Wang et al. XY19-1287, 4556 m elev., XY19-1290; Sajia Co., Mula village, 28°40′N, 88°45′E, 2019.7.28, 4752 m elev., Xin-Yu Wang et al. XY19-1814; Angren Co., Kerangla, 29°19′N, 87°01′E, 4530 m elev., Li-Song Wang et al. 19-63635.
Immersaria ferruginea C.M. Xie & Li S. Wang, sp. nov. — Fungal Names FN839739; Fig. 5a–c
Etymology
The epithet “ferruginea” refers to the rusty brown color of the thallus.
Type
CHINA, Tibet, Changdu City, Mangkang Co., Quzika village, 4093 m elev., 29°15′N, 98°40′E, on rock, 25 Sept. 2020, Li-Song Wang et al. 20-69144 (KUN–holotype).
Description
Thallus areolate, grayish brown, rusty colored, continuous, areolae 0.5–1.25 mm in diam., flat, less convex, rectangle to polygon, epruinose; margin pruinose; prothallus black, not distinct. Upper cortex 50.0–67.5 µm thick, brown granules pigmented; epinecral layer 17.5–40.0 µm thick; algal layer 75.0–77.5 µm thick, cells 4.0–7.5–13.0 × 4.0–7.5–13.0 µm in diam., round; lower cortex absent. Apothecia frequent, densely crowded, immersed, 0.75–1.25 mm in diam.; disc black, flat, pruinose; margin pruinose, slightly raised. Exciple sometimes developed, 25.0–27.5 µm wide, brown. Hymenium 57.5–100.0 µm thick, colorless; paraphyses 1.0–2.5 µm µm wide, not branched, slightly anastomosing; epihymenium 15.0–32.5 µm thick, brown; subhymenium 25.0–62.5 µm thick, colorless to pale brown, rusty or dark pink; hypothecium pale brown. Asci Porpidia-type, cylindrical; ascospores not seen, or rarely seen. Pycnidia not seen.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. Confluentic acid, often accompanied with 2’-O-methylmicrophylinic acid.
Ecology and distribution
Growing on quartz sandstone or granite at elevations of 3848–4227 m in the alpine zone. This species is known from Sichuan Province and Tibet of China.
Remarks: Immersaria ferruginea is characterized by its brown and rusty thallus, its densely crowded apothecia and its brown epihymenium. I. ferruginea and I. plumbeoolivacea were previously regarded as a single species, but it was later found that I. ferruginea grows on red sandstone whereas I. plumbeoolivacea grows on granite, and the thallus of I. ferruginea is rusty. The morphology of I. ferruginea resembles I. carbonoidea, but the latter differs in its dark black brown thallus, black brown hypothecium and content of norstictic acid.
Specimens examined (KUN). CHINA, Sichuan Province, Rangtang Co., Mt. Haizi, 4227 m elev., 32°20′N, 101°25′E, on rock, 2020, Li-Song Wang et al. 20-66697, 4221 m elev., Li-Song Wang et al. 20-67670. Tibet, Changdu City, Mangkang Co., Quzika village, 4093 m elev., 29°15′N, 98°40′E, Li-Song Wang et al. 20-69144, 4101, Li-Song Wang et al. 20-69146, 4122 m elev., Li-Song Wang et al. 20-69148; Gatuo town, 3848 m elev., 29°39′N, 98°35′E, Li-Song Wang et al. 20-69105.
Immersaria plumbeoolivacea C.M. Xie & Xin Y. Wang, sp. nov. — Fungal Names FN839740; Fig. 5d–f
Etymology
The epithet “plumbeoolivacea” refers to the lead brown thallus.
Type
CHINA, Qinghai Province, Banma Co., 3948 m elev., 32°40′N, 100°48′E, on rock, 9 Sept. 2020, Li-Song Wang et al. 20-66914 (KUN–holotype).
Description
Thallus areolate, grayish brown, continuous, areolae 0.5–1.3 mm in diam., undulate, flat, convex or concave, irregular, epruinose; margin pruinose; prothallus black, not distinct. Upper cortex 40.0 µm thick, brown granules pigmented; epinecral layer 30.0 µm thick; algal layer 62.5 µm thick, cells 10.0–15.0 × 5.0–7.5 µm in diam., round; lower cortex absent. Apothecia frequent, crowded, immersed, irregular, 0.75–1.75 mm in diam.; disc black, sometimes pruinose, flat; margin thin, pruinose, slightly raised. Exciple sometimes developed, 30.0–37.5 µm wide, brown. Hymenium 67.5–100.0 µm thick, colorless; paraphyses 2.0–2.5 µm wide, not branched, slightly anastomosing; epihymenium 37.5 µm thick, green or brown green; subhymenium 50.0–67.5 µm thick, colorless to pale brown, rusty or dark pink; hypothecium pale brown. Asci Porpidia-type, cylindrical; ascospores not seen. Pycnidia rare, immersed, oblate, black, margin pruinose; conidia 7.5 × 1.0 µm in diam., bacilliform.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. Confluentic acid, often accompanied with 2’-O-methylmicrophylinic acid.
Ecology and distribution
Growing on granite at elevations of 3932–3971 m in the alpine zone. This species is known from Qinghai Province of China.
Remarks: Immersaria plumbeoolivacea is characterized by its gray brown thallus, the areolae tend to be squamulose, with pruinose apothecia and green epihymenium. The phylogenetic results showed that I. ferruginea was a sister species to I. plumbeoolivacea, which is similar in its brown thallus and crowded apothecia, but differs in its thallus with rusty color and a brown epihymenium. The morphology of I. plumbeoolivacea resembles I. carbonoidea, but the latter has a black brown, thallus and contains traces of norstictic acid.
Specimens examined (KUN). CHINA, Qinghai Province, Banma Co., 3932 m elev., 32°40′N, 100°48′E, on rock, 2020, Li-Song Wang et al. 20-66889, 3948 m elev., Li-Song Wang et al. 20-66914, 3945 m elev., Li-Song Wang et al. 20-66915; Makehe, 3971 m elev., 32°42′N, 100°42′E, Li-Song Wang et al. 20-67896.
Immersaria shangrilaensis C.M. Xie & Lu L. Zhang, sp. nov. — Fungal Names FN839741; Fig. 6a–d
Etymology: The epithet “shangrilaensis” refers to the location at which the holotype was collected: “Shangri-La", a county of Yunnan Province in China.
Type
CHINA, Yunnan Province, Shangri-La County., Mt. Hong Shan, 4363 m elev., 28°7′N, 99°54′E, on rock, 18 Aug. 2018, Li-Song Wang et al. 18-60447 (KUN–holotype).
Description
Thallus areolate, yellow brown, orange brown, often appears as grayish brown. It is generally heavily pruinose, continuous, 5.75–10.0 cm across, areolae aggregated by 4–14 small areolae (often surrounded by black prothallus), small areolae up to 0.13 mm in diam., concave, or flat, irregular, pruinose; margin pruinose; prothallus black, distinct. Upper cortex 32.5–50.0 µm thick, yellow brown granules pigmented; epinecral layer 15.0–20.0 µm thick; algal layer 47.5–65.0 µm thick, cells 7.5–8.0 × 5.0 µm in diam., round; lower cortex absent. Apothecia frequent, crowded, immersed or isolated from areolae, 0.38–0.75 mm in diam.; disc black, concave to flat, aggregated, crack once mature, thin pruinose; margin reduced, slightly raised. Exciple almost absent. Hymenium 100.0–137.5 µm thick, colorless; paraphyses 2.5 µm wide, branched, anastomosing or not; epihymenium 15.0 µm thick, brown; subhymenium 55.0 µm thick, colorless; hypothecium colorless to pale brown, sometimes absent. Asci Porpidia-type, cylindrical, eight-spored; ascospores 7.0–9.0 × 3.0–4.0 µm, ellipsoid, halonate (sometimes not distinct). Pycnidia immersed, oblate, black, margin heavily pruinose; conidia 7.5 × 1.0 µm, bacilliform.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. Confluentic acid, planaic acid and/or 2’-O-methylmicophyllinic acid.
Ecology and distribution
Growing on granite at elevations of 4361–4503 m in the alpine zone. This species is known from Yunnan Province of China.
Remarks
The materials of Immersaria athroocarpa from Shangri-La county of Yunnan Province are identical with the paratype Hertel (1977) reported from the same locality, but differ from the Lectotype in its aggregate areolae, the aggregate apothecia and the smaller size of ascospores (7.0–9.0 × 3.0–4.0 µm). Based on the phenotypic and phylogenetic results, the materials from Shangri-La were treated as a new species, I. shangrilaensis. It is characterized by its large thallus, up to 10.0 cm in diam., the aggregate areolae and apothecia and the small size of ascospores.
Specimens examined
CHINA, Yunnan Province, Shangri-La Country, 4350–4500 m elev., on rock, 1915, Handel-Mazzetti no. 6945 = WU-Lichenes0037752 (WU); Mt. Hong Shan, 4363 m elev., 28°7′N, 99°54′E, on rock, 2018, Li-Song Wang et al. 18-60430 (KUN), Li-Song Wang et al. 18-60447 (KUN) 4503.1 m elev., Chun-Xiao Wang et al. SDNU20181696 (SDNU), 4361.9 m elev., Chun-Xiao Wang et al. SDNU20181675 (SDNU); Luquan Co., Mt. Jiaozixueshan, 3800 m elev., 2008, Hai-Ying Wang SDNU20082253 (SDNU); Lijiang City, Mt. Laojun, 26°37′N, 99°43′E, 2018, 3981m, Li-Song Wang et al. 18-60555, 18-60602 (KUN).
Immersaria venusta C.M. Xie & Xin Y. Wang, sp. nov. — Fungal Names FN839742; Fig. 6e–i
Etymology
The epithet “venusta” refers to the beautiful appearance of the thallus.
Type
CHINA, Qinghai Province, Maqing Co., Xueshan village, 4187 m elev., 34°37′N, 99°42′E, on rock, 11 Sept. 2020, Li-Song Wang et al. 20-67969 (KUN–holotype).
Description
Thallus areolate, brown, orange brown, more or less rusty, continuous, areolae 0.5–1.25 mm in diam., flat or slightly convex, irregular, tend to rectangular, often cracked, sometimes pruinose; margin pruinose; prothallus not seen. Upper cortex 37.5 µm thick, yellow brown granules pigmented; epinecral layer 12.5 µm thick; algal layer 127.5 µm thick, cells 5.0–10.0 × 5.0–7.5 µm in diam., round; lower cortex absent. Apothecia frequent, crowded, immersed or isolated from areolae, 0.6–1.0 mm in diam.; disc black, flat, epruinose; margin reduced, sometimes developed. Exciple sometimes developed, 35.0 µm wide, brown. Hymenium 92.5–112.5 µm thick, colorless; paraphyses 2.0 µm wide, anastomosing; epihymenium 27.5–30.0 µm thick, brown; subhymenium 62.5 µm thick, colorless; hypothecium brown. Asci Porpidia-type, cylindrical, eight-spored; ascospores 10.0–12.5 × 5.0–7.5 µm, ellipsoid, halonate. Pycnidia immersed, linear, black, margin pruinose; conidia not seen.
Chemistry: Thallus K–, C–. Medulla I+, K–, C–. Ⅰ: Confluentic acid, often accompanied with 2’-O-methylmicrophylinic acid. Ⅱ: Planaic acid. Ⅲ: none (rare).
Ecology and distribution
Growing on quartz sandstone or granite at elevations of 3930–4256 m in the alpine zone. This species is known from Qinghai Province of China.
Remarks: Immersaria venusta is characterised by its yellow brown, cracked areolae, its flat apothecia and brown epihymenium. It resembles I. shangrilaensis by its cracked areolae, but its areolae have the tendency to split into several patches but not aggregate like those of I. shangrilaensis. I. athroocarpa is similar to I. venusta in the brown appearance of its thallus and in forming a sister group in the phylogenetic tree, but it differs in its yellow brown thallus, convex areolae, densely crowded apothecia and bigger ascospores (17.5–20.0 × 10.0 µm). This species is also similar to I. aurantia (see notes for I. aurantia). The brown thallus of I. venusta possibly resembles I. olivacea, but the latter differs in its simple or one-septate ascospores, pyriform conidia and dark bluish green epihymenium. Three specimens of I. venusta have brown thallus, which resembles I. plumbeoolivacea, but they have neatly arranged, thick areolae.
Specimens examined (KUN). CHINA, Qinghai Province, Maqing Co., Xueshan village, 4187 m elev., 34°37′N, 99°42′E, on rock, 2020, Li-Song Wang et al. 20-67969, 20-67965; Banma Co., Yaertang village, 3930 m elev., 32°42′N, 100°42′E, Li-Song Wang et al. 20-66940. Sichuan Province, Shiqu Co., Xinrong village, 4043 m elev., 32°59′N, 98°19′E, on rock, 2020, Li-Song Wang et al. 20-68802; Rangtang Co., Mt. Haizi, 4246 m elev., 32°21′N, 101°24′E, Li-Song Wang et al. 20-66721, 20-66725.
Selected additional comparative material was examined.
Bellemerea alpina (Sommerf.) Clauzade & Cl. Roux RUSSIA, Lps. Petsamo, Pummangin vuonon N-puoli, 1938, Räsänen, V., H9503269 (H–type!); Lps. Petsamo, inter Vaitolahti et Kervanto, 1938, Räsänen, V., H9503270 (H–type!).
Bellemerea cinereorufescens (Ach.) Clauzade & Cl. Roux FINLAND, Ob. Simo. Anteroinen. Rantakivellä, 1920, Räsänen, V., H9503267 (H–type!); Le. Enontekiö, Kirkonkylä, 1925, Kari, L.E., H9503268 (H–type!).
Immersaria cupreoatra (Nyl.) Calatayud & Rambold RUSSIA, Kl. Kurkijoki, Kuuppala, Himohirsi, 1934, Räsänen, V., H9503417, H9510194 (H–type!).
High-resolution photographs seen.
Immersaria carbonoidea (J.W. Thomson) Esnault & Cl. Roux USA, Alaska, along the Pitmegea River, 15 miles upstream from Cape Sabine, 1958, Thomson, J.W., M0082171 (M–Isotype!), G00126754 (G–isotype!).
Immersaria olivacea Calat. & Rambold SPAIN, Espana, Castelló: Benicàssim, Parreta Alta, 390 m elev., 1993, Calatayud, V., M0101779 (M–isotype!).
Immersaria usbekica (Hertel) M. Barbero, Nav.-Ros. & Cl. Roux ALGERIA, Algerie-Atlas Tellieu, Larba, Piste de Bougara á Tablat au S-E de l'arboretum de Meindja, 1985, Esnault, J., M0101787 (M–paratype!).
Lecaimmeria C.M. Xie, Lu L. Zhang & Li S. Wang, gen. nov. — Fungal Names FN839743;
Etymology
The epithet “Lecaimmeria” refers to the immersed lecanorine apothecia of known species.
Type: Lecaimmeria orbicularis C.M. Xie & Lu L. Zhang, sp. nov.
Description
Thallus crustose, red brown, orange brown or dark brown, continuous or not, areolae irregular or tend to rectangular, with wax glossy, margin white or black, prothallus distinct at the margin of thallus or absent, sometimes developed between areolae. Upper cortex with orange granules pigmented, epinecral layer colorless, algal layer continuous, medulla filled with gray granules, lower cortex absent. Apothecia lecanorine-type, immersed, round or irregular; disc red brown, dark red brown or dark orange brown, flat or concave, margin present or absent, black or white, rare green, pruinose or not. Exciple reduced, tissue at the lateral sides of the hymenium corresponding to the upper cortex and the algal layer of the vegetative areolae, and to hypothecial hyphal cells when apothecia reach the margin of the areole. Hymenium colorless, paraphyses simple, rare branch, anastomosing or not, epihymeinum orange, orange brown, rare brown, with epinecral layer. Asci Porpidia-type, cylindrical, eight-spored; ascospores ellipsoid, halonate, non-amyloid. Pycnidia present or absent, immersed, rare convex, liner or stellate, rare tuberculiform, conidia bacilliform.
Chemistry
Thallus K–, C–. Medulla I+, K–, C+/–. Gyrophoric acid, 4-O-demethylplanaic acid or no substances detected by TLC.
Ecology and distribution
Growing on rock, sandstone, granite or Qilian jade (rare), from elevations of 3100 to 4800 m in the alpine zone of west China and from 1200 to 1900 m in the steppe of north China. This genus is known from China, Europe, Iran, Macedonia, Mongolia, Romania and Russia.
Remarks
The five-loci phylogenetic analysis showed that these lecanorine apothecia species should be excluded from Immersaria and formed a novel lineage, thus they are here treated as a new genus Lecaimmeria. Lecaimmeria is distinguished from related genera by its glossy wax, orange or red brown areolae with margins, the amyloid medulla, the red brown immersed lecanorine apothecia with margins, the orange epihymenium with an epinecral layer and the Porpidia-type asci with eight halonate and non-amyloid ascospores. The genus is distributed in alpine areas, high altitude dessert-steppe areas or high latitude steppe. Almost all the species of Lecaimmeria grow on granite or sandstone, with the exception of one species, L. tuberculosa, which grows on jade. Interestingly, the margin of pycnidia and areolae of L. tuberculosa appear with heavily jade green pruinose.
The immersed apothecia and brown thallus of Lecaimmeria often resemble those of Immersaria, but Lecaimmeria differs in its red brown lecanorine apothecia, often with a white margin, their orange epihymenium with an epinecral layer and content of gyrophoric acid. This genus is often confused with Bellemerea in its lecanorine apothecia and the Porpidia-type asci with halonate ascospores, but the latter genus differs in its white thallus and amyloid ascospores.
Three species previously in Immersaria, I. cupreoatra, I. iranica and I. mehadiana, have lecanorine apothecia, but two of these, I. cupreoatra and I. mehadiana, currently lack molecular sequences. It is suggested that these three species should be transferred to Lecaimmeria, based on the following factors. Their morphology is consistent with Lecaimmeria according to molecular results and comparisons with type specimens, high-resolution photographs of the type materials and the original descriptions. One unknown “Immersaria” species from Macedonia is sister to L. iranica in the phylogenetic tree (Fig. 3), but differs in its black margin of areolae, absence of gyrophoric acid and comparison with high-resolution photograph and previous records (Malíček and Mayrhofer 2017). This unknown species with lecanorine apothecia is possibly a member of Lecaimmeria, but descriptions are lacking and the specimens were not seen. Thus, this species was temporarily retained in Immersaria.
Lecaimmeria botryoides C.M. Xie & Li S. Wang, sp. nov. — Fungal Names FN839744; Fig. 7a–e
Etymology
The epithet “botryoides” refers to the crowded apothecia while immature.
Type
CHINA, Sichuan Province, Aba City, Rangtang County, Haizi Mt., 4225 m elev., 32°21′N, 101°24′E, on rock, 6 Sept. 2020, Li-Song Wang et al. 20-66730 (KUN–holotype).
Description
Thallus areolate, red brown, discontinuous, areolae 0.25–1.0 mm in diam., flat, slightly concave or convex, pruinose, polygon, tend to be rectangular, margins with heavy pruinose. Prothallus black, distinct in the margin of thallus. Upper cortex 20.0–25.0 µm thick, brown; epinecral layer 22.5–47.5 µm thick; algal layer 37.5 µm thick, cells 7.5–10.0 × 7.5–10.0 µm in diam., round; lower cortex absent. Apothecia frequent, irregular, densely crowded while immature (3–6/areolae), aggregate once mature, immersed, 0.25–1.25 mm in diam.; disc red brown, flat, or concave, epruinose; margin pruinose, slightly raised. Hymenium 67.5–100.0 (–155.0) µm thick, colorless; paraphyses 2.5 µm wide, simple, only branched at the top, not anastomosing; epihymenium 25.0–30.0 µm thick, orange; epinecral layer 2.5–7.5 µm thick; subhymenium 17.5–37.5 µm thick, colorless; hypothecium colorless. Asci Porpidia-type, cylindrical, eight-spored; ascospores 7.5–8.0 × 4.0–6.0 µm in diam., ellipsoid, halonate. Pycnidia not seen; conidia not seen.
Chemistry: Thallus K–, C–. Medulla I+, K–, C+/–. Ⅰ: Gyrophoric acid. Ⅱ: none.
Ecology and distribution
Growing on rock at elevations of 3178–4300 m in the alpine zone. This species is known from Qinghai and Sichuan Province of China.
Remarks: Lecaimmeria botryoides is characterized by its discontinuous thallus, densely crowded apothecia while immature, and the orange epihymenium. L. orbicularis is similar to L. botryoides in its red brown thallus, but differs in its round, flat apothecia and continuous thallus. The red brown thallus of L. botryoides resembles L. cupreoatra, but the latter differs in its black margin of apothecia and its distinct black prothallus between areolae.
Specimens examined (KUN). CHINA, Qinghai Province, Banma Co., 3958 m elev., 32°40′N, 100°48′E, on rock, 2020, Li-Song Wang et al. 20-66900, 3932 m elev., Li-Song Wang et al. 20-66898, 3935 m elev., Li-Song Wang et al. 20-66891, 3178 m elev., Li-Song Wang et al. 20-66765. Sichuan Province, Rangtang Co., Mt. Haizi, 4256 m elev., 32°21′N, 101°24′E, on rock, 2020, Li-Song Wang et al. 20-66721, 4300 m elev., Li-Song Wang et al. 20-67706, 4276 m elev., Li-Song Wang et al. 20-66706, 4255 m elev., Li-Song Wang et al. 20-66707, 4274 m elev., Li-Song Wang et al. 20-66713, 4274 m elev., Li-Song Wang et al. 20-66711, 20-66705, 4225 m elev., Li-Song Wang et al. 20-66730, 4220 m elev., 32°20′N, 101°25′E, Li-Song Wang et al. 20-66683.
Lecaimmeria cupreoatra (Nyl.) C.M. Xie, comb. nov. — Fungal Names FN839745;
Type
RUSSIA, Ad Onegam, Medväschiigora, 1863, Simming, Th., H-NYL 25061 = H9508237 (H–lectotype!).
Description
Nylander (1866).
Remarks
The lectotype grows on siliceous rock and contains several intact apothecia. “Immersaria” cupreoatra has lecanorine apothecia and is related to I. lygeae in its phylogeny, therefore it was transferred to Lecaimmeria. This species has not been recorded in China (see notes for L. mongolica). The species is known from Europe, Macedonia, Mongolia and Russia (Calatayud and Rambold 1998).
Lecaimmeria iranica (Valadb., Sipman & Rambold) C.M. Xie, comb. nov. — Fungal Names FN839746;
Type
IRAN, Mazandaran, Haraz road, 20 km to Aamol, 36°17′N, 52°21′E, on calcareous rock, 1475 m, 7 Apr. 2006, T. Valadbeigi 9008 (TARI–holotype; B, hb. Valadbeigi–isotype). Not seen.
Description
Valadbeigi et al. (2011).
Remarks
“Immersaria” iranica has lecanorine apothecia, a distinct epinecral layer and halonate ascospores (Valadbeigi et al. 2011). The materials from China are in accordance with the materials of Iran based on comparisons with the original descriptions and the photographs given by Valadbeigi et al. (2011). The characters of this species are consistent with the new genus and the phylogenetic results showed that it was clustered with species of Lecaimmeria. Therefore, it was transferred to Lecaimmeria. This species is currently known from Iran and China.
Specimens examined (SDNU). CHINA, Xinjiang, Urumqi, Mt. Tianshan-glacier No.1, alt. 3800 m, on rock, 2011, Z.L. Huang SDNU20126106, SDNU20129049.
Lecaimmeria lygaea C.M. Xie & Lu L. Zhang, sp. nov. — Fungal Names FN839747; Fig. 7f–j
Etymology
The epithet “lygaea” refers to the dark appearance of the thallus.
Type
CHINA, Tibet, Changdu City, Mangkang County, Luoni Village, 4127 m elev., 29°56′N, 98°33′E, on rock, 24 Sept. 2020, Li-Song Wang et al. 20-69072 (KUN–holotype).
Description
Thallus areolate, dark red brown, dark brown, continuous, areolae 0.5–1.0 mm, flat, epruinose, irregular pentagonal, sometimes rectangular, fissures between areolae often filled with black prothallus; margin black, thin pruinose; prothallus black, developed between areolae, also distinct in the margin. Upper cortex 20.0 µm thick, orange brown granules pigmented; epinecral layer 15.0 µm thick; algal layer 50.0 µm thick, cells 7.5–12.5 × 7.5–12.5 µm in diam., round; lower cortex absent. Apothecia frequent, round, crowded, immersed, 0.25–0.75 mm in diam.; disc red brown, flat, or concave, epruinose; margin black, moderate thick, pruinose, raised. Hymenium 75.0–92.5 µm thick, colorless; paraphyses 2.0 µm wide, simple, unbranched, not anastomosing; epihymenium 25.0–37.5 µm thick, orange brown; epinecral layer 7.5 µm thick; subhymenium 20.0–25.0 µm thick, colorless; hypothecium absent. Asci Porpidia-type, cylindrical, eight-spored; ascospores 12.5–20.0 × 5.0–7.5 µm in diam., ellipsoid, halonate. Pycnidia immersed, stellate, black, margin pruinose; conidia 5.0 × 1.0 µm in diam., bacilliform.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. Unknown fatty acid.
Ecology and distribution
Growing on sandstone at elevations of 4095–4131 m in the alpine zone. This species is known from Tibet of China.
Remarks: Lecaimmeria lygaea is characterized by its dark brown thallus, black margin of its areolae, black prothallus which filled the fissures between areolae, dark orange apothecia and its orange brown epihymenium. L. cupreoatra and L. mehadiana are similar to L. lygaea, but L. cupreoatra has a discontinuous thallus, with each areola surrounded by black prothallus, dark red brown to black brown apothecia without margin. L. mehadiana has a white margin of areolae, black brown apothecia, brown epihymenium and contains 4-O-demethylplanaic acid. The phylogenetic results show that L. tibetica was a sister species to L. lygaea. They are similar in chemistry, but differ in its orange brown thallus and dark orange brown apothecia.
Specimens examined (KUN). CHINA, Tibet, Changdu City, Mangkang Co., Luoni village, 4099 m elev., 29°56′N, 98°33′E, on rock, 2020, Li-Song Wang et al. 20-69054, 4131 m elev., Li-Song Wang et al. 20-69070, 4127 m elev., Li-Song Wang et al. 20-69072, 4095 m elev., Li-Song Wang et al.20-69053.
Lecaimmeria mehadiana (Calatayud & Rambold) C.M. Xie, comb. nov. — Fungal Names FN839748;
Type
ROMANIA, Caras-Severin Comitat, Mehadía, Strájot Mtn., on rock, 1994, Rambold, G.W., M0101781 (M–holotype!), M0101780, M0101782, M0101783 (M–isotype!). High-resolution photographs seen.
Description
Calatayud and Rambold (1998).
Remarks
“Immersaria” mehadiana has lecanorine apothecia and is related to L. lygaea and L. cupreoatra in the appearance of dark brown thallus, therefore it has been transferred to Lecaimmeria. This species is characterized by its greyish prothallus, dark brown apothecia and the brown epihymenium. It is only known from Romania (Calatayud and Rambold 1998).
Lecaimmeria mongolica C.M. Xie & Lu L. Zhang, sp. nov. — Fungal Names FN839749; Fig. 8a-e
Etymology
The epithet “mongolica” refers to the collection of the holotype within Inner Mongolia, an autonomous region of China.
Type: CHINA, Inner Mongolia: Chifeng City, Balinyouqi, Han Mountain, 1445m elev., 44°11′N, 118°44′E, on rock, 22 Jul. 2019, Z.T. Zhao et al. SDNU20190354 (SDNU–holotype).
Description
Thallus areolate, orange, continuous, areolae 0.4–0.8 mm in diam., epruinose, neatly arranged, irregular, tend to be rectangular, margin pruinose; prothallus black, not distinct. Upper cortex 20.0 µm thick, brown granules pigmented; epinecral layer 5.0–7.5 µm thick; algal layer 87.5 µm thick, cells 7.5–12.5 × 7.5–12.5 µm in diam., round; lower cortex absent. Apothecia frequent, crowded, immersed or isolated from areolae, 0.25–0.75 mm in diam.; disc red brown, flat, slightly convex, epruinose; margin pruinose. Hymenium 62.5–82.5 µm thick, colorless; paraphyses 2.5 µm wide, unbranched, not anastomosing; epihymenium 42.5 µm thick, orange; epinecral layer 5.0–10.0 µm thick; subhymenium 30.0–37.5 µm thick, colorless; hypothecium absent. Asci Porpidia-type, cylindrical, eight-spored; ascospores 10.0–17.5 × 6.0–7.5 µm in diam., ellipsoid, halonate. Pycnidia immersed, oblate, rare ellipsoid, black, margin pruinose; conidia 5.0 × 1.0 µm in diam., bacilliform.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. Gyrophoric acid.
Ecology and distribution
Growing on granite at elevations of 1400–2000 m in steppe or mountains. This species is known from Inner Mongolia of China.
Remarks
This species was once reported as “Immersaria” cupreoatra from China (Zhang et al. 2015) but after comparing with the type materials this was found to be a misclassification. Also, the phylogenetic results showed that these materials formed a well-supported lineage belonging to Lecaimmeria. Therefore, here it has been treated as a new species, Lecaimmeria mongolica. It is characterized by its orange brown thallus, red brown apothecia with a distinct white margin and content of gyrophoric acid. L. cupreoatra resembles L. mongolica by containing gyrophoric acid, but it differs in its dark black brown thallus and the black margin of its apothecia. L. tibetica is similar to L. mongolica in its orange thallus, but differs in its smaller, dark orange apothecia and that no substance can be detected by TLC.
Specimens examined (SDNU). CHINA, Inner Mongolia: Chifeng City, Balin Youqi, Mt. Qingyangcheng, 1445 m elev., 43°35′N, 117°30′E, on rock, 2019, Zun-Tian Zhao et al. SDNU20190350; Han Shan, 1563 m elev., 44°11′N, 118°44′E, on rock, Zun-Tian Zhao et al. SDNU20190354; A’ershan City, Mt. Jiguan, 1500 m elev., on rock, 2011, Yu-Liang Cheng SDNU20124912, 1400 m elev., Dai-Feng Jiang SDNU20124859; Ke Qi, Huanggangliang, 2000 m elev., on rock, Pan-Meng Wang SDNU20117613, Xing-Ran Kou SDNU20117399.
Lecaimmeria orbicularis C.M. Xie & Lu L. Zhang, sp. nov. — Fungal Names FN839750; Fig. 9a–e
Etymology
The epithet “orbicularis” refers to the round shape of the apothecia.
Type
CHINA, Sichuan Province, Rangtang Co., Gangmuda village, 3800 m elev., 32°18′N, 101°3′E, on rock, 7 Sept. 2020, Li-Song Wang et al. 20-66753 (KUN–holotype).
Description
Thallus areolate, red brown, rare orange brown, continuous, areolae 0.25–1.0 mm, flat, occasionally wrinkled, tend to rectangular, fissures between areolae often filled with black prothallus, margin pruinose; prothallus black, developed between areolae, also distinct in the margin. Upper cortex 42.5–57.5 µm thick, brown granules pigmented; epinecral layer 5.0–20.0 µm thick; algal layer 70.0–112.5 µm thick, cells 10.0–15.0 × 7.5–10.0 µm in diam., round; lower cortex absent. Apothecia frequent, less crowded, immersed or isolated from areolae, 0.5–1.25 mm in diam.; disc red brown, flat, round, epruinose; margin white, slightly raised. Hymenium 75.0–112.5 µm thick, colorless; paraphyses 2.5 µm wide, simple, unbranched, not anastomosing; epihymenium 17.5–30.0 µm thick, orange; epinecral layer 5.0–15.0 µm thick; subhymenium 30.0–62.5 µm thick, colorless; hypothecium absent. Asci Porpidia-type, cylindrical, eight-spored; ascospores 12.5–15.0 × 5.0–6.0 µm in diam., ellipsoid, halonate. Pycnidia not seen.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. None.
Ecology and distribution
Growing on granite or sandstone at elevations of 3730–4200 m in the alpine zone. This species is known from Qinghai and Sichuan Provinces of China.
Remarks: Lecaimmeria orbicularis is characterized by its orange brown thallus, neatly arranged areolae, and round, flat apothecia. L. botryoides is similar to L. orbicularis (see notes for L. botryoides). L. mongolica might be confused with L. orbicularis due to its big apothecia with a white margin, but differs in its red brown thallus and distribution in steppes. The red brown thallus of L. cupreoatra resembles that of L. orbicularis, but differs in the black margin of its apothecia and its distinct black prothallus between areolae.
Specimens examined (KUN). CHINA, Qinghai Province, Jiuzhi Co., Nianbaoyuze, 4200 m elev., 33°14′N, 100°58′E, on rock, 2020, Li-Song Wang et al. 20-66811, 20-66829, 20-66801, 20-66826A, 20-66821, 20-66805, 20-66833, 20-66817, 20-66841; Banma Co., Nianbaoyuze, 3930 m elev., 32°40′N, 100°48′E, Li-Song Wang et al. 20-66909, 20-66908, 20-66896, 20-66886B, 20-66899, 20-66935, 20-66943; Zhiqingsongduo town, 3712 m elev., 33°24′N, 101°25′E, Li-Song Wang et al. 20-66965; Suohurima village, 4029 m elev., 33°23′N, 100°57′E, Li-Song Wang et al. 20-66979. Sichuan Province, Rangtang Co., Gangmuda village, 3800 m elev., 32°18′N, 101°3′E, on rock, 2020, Li-Song Wang et al. 20-66753, 20-66750, 3793 m elev., Li-Song Wang et al. 20-66747, Shangrangtang village, 3730 m elev., 32°16′N, 101°21′E, Li-Song Wang et al. 20-66743.
Lecaimmeria qinghaiensis C.M. Xie & Li S. Wang, sp. nov. — Fungal Names FN839751; Fig. 9f–i
Etymology
The epithet “qinghaiensis” refers to the location in which the holotype was collected, in “Qinghai”, a province of China.
Type
CHINA, Qinghai Province, Yushu City, Zaduo County, Sahuteng town, 4634 m elev., 32°55′N, 95°46′E, on rock, 20 Sept. 2020, Li-Song Wang et al. 20-68698 (KUN–holotype).
Description
Thallus areolate, yellow brown, rusty, continuous, areolae 0.5–1.5 mm in diam., flat, epruinose; margin pruinose, occasionally green pigmented; prothallus black, distinct at the margin. Upper cortex 27.5–37.5 µm thick, brown granules pigmented; epinecral layer 12.5–20.0 µm thick; algal layer 57.5–92.5 µm thick, cells 7.5–12.5 × 5.0–12.5 µm in diam., round; lower cortex absent. Apothecia frequent, immersed or isolated from areolae, round or somewhat irregular while immature, ellipsoid and tend to be rectangular for occupying the whole areolae once mature, 0.25–1.25 mm in diam.; disc brown, dark red brown, flat, occasionally with fissure when mature, epruinose; margin white, slightly raised. Hymenium 52.5–62.5 µm thick, colorless; paraphyses 2.0–2.5 µm wide, unbranched, not anastomosing; epihymenium 25.0–30.0 µm thick, dark orange brown; epinecral layer 7.5–17.5 µm thick; subhymenium 50.0–62.5 µm thick, colorless; hypothecium absent. Asci Porpidia-type, cylindrical, eight-spored; ascospores 8.0–15.0 × 5.0–7.5 µm in diam., ellipsoid, halonate not distinct. Pycnidia rare, immersed, flat, slightly convex, liner, stellate, graphidoid once mature, black, margin pruinose; conidia not seen.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. None.
Ecology and distribution
Growing on rock at elevations of 4622–4815 m in the alpine zone. This species is known from Qinghai Province of China.
Remarks: Lecaimmeria qinghaiensis is characterized by the yellow brown, rusty thallus, the red brown apothecia often occupying the whole areolae at maturity, and the dark orange brown epihymenium. The phylogenetic results showed that L. tuberculosa was sister species to L. qinghaiensis which is similar in the appearance of thallus, but differs in the brown, never rusty thallus, the red brown apothecia, and the green, tuberculiform pycnidia. The red brown thallus of L. qinghaiensis resembles L. iranica, but differs in the rusty thallus and the white margin of apothecia.
Specimens examined (KUN). CHINA, Qinghai Province, Zaduo Co., Sahuteng town, 4634 m elev., 32°55′N, 95°46′E, on rock, 2020, Li-Song Wang et al. 20-68698, 4637 m elev., Li-Song Wang et al. 20-68687, 4622 m elev., Li-Song Wang et al. 20-68696; 4790 m elev., 33°31′N, 95°8′E, Xin-Yu Wang et al. 20-3115, 4791 m elev., Xin-Yu Wang et al. 20-3127; Zaqing village, 4815 m elev., Xin-Yu Wang et al. 20–849.
Lecaimmeria tibetica C.M. Xie & Xin Y. Wang, sp. nov. — Fungal Names FN839752; Fig. 10a-e
Etymology: The epithet “tibetica” refers to the location from which the holotype was collected: “Tibet”, an
autonomous region of China.
Type
CHINA, Tibet, Gongga Co., Jiangtang town, 4557 m elev., 29°12′N, 90°38′E, on rock, 9 Sept. 2019, Xin-Yu Wang et al. XY19-1291 (KUN–holotype).
Description
Thallus areolate, orange brown, epruinose, areolae 0.3–0.5 mm in diam., irregular, upper surface uneven, margin lacking, pruinose; prothallus black, distinct at the margin. Upper cortex 17.5–32.5 µm thick, brown granules pigmented; epinecral layer 10.0–20.0 µm thick; algal layer 75.0 µm thick, cells 7.5–10.0 × 7.5–10.0 µm in diam., round; lower cortex absent. Apothecia rare, immersed or isolated from areolae, 0.25–0.5 mm in diam.; disc dark orange brown, epruinose, flat, slightly convex; margin pruinose. Hymenium 105.0–137.5 µm thick, colorless; paraphyses 2.5 µm wide, unbranched, not anastomosing; epihymenium 25.0 µm thick, orange; epinecral layer 12.5 µm thick; subhymenium almost absent, colorless; hypothecium absent. Asci Porpidia-type, cylindrical, eight-spored; ascospores 12.5–15.0 × 5.0–6.0 µm in diam., ellipsoid, halonate. Pycnidia immersed, oblate, black, margin pruinose; conidia 5.0 × 1.5–2.0 µm in diam., bacilliform.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. None.
Ecology and distribution
Growing on quartz sandstone at elevations of 4310–4583 m in the alpine zone. This species is known from Tibet of China.
Remarks: Lecaimmeria tibetica is characterized by the orange brown thallus, the black pigmented of areolae margin, and the dark orange brown and small size of apothecia. L. tibetica is similar to L. mongolica (see notes for L. mongolica). The red brown apothecia of L. cupreoatra resembles L. tibetica, but differs in its dark red brown thallus and the presence of gyrophoric acid.
Specimens examined (KUN). CHINA, Tibet, Gongga Co., Jiangtang town, 4583 m elev., 29°12′N, 90°38′E, on rock, 2019, Xin-Yu Wang et al. XY19-1288, 4557 m elev., XY19-1291, 4560 m elev., XY19-1280; Dingri Co., Zhaguozhong, 4310 m elev., 28°35′N, 86°53′E, Li-Song Wang et al. 19-64071.
Lecaimmeria tuberculosa C.M. Xie & Xin Y. Wang, sp. nov. — Fungal Names FN839754; Fig. 10f–i
Etymology
The epithet “tuberculosa” refers to the tuberculiform of the pycnidia.
Type
CHINA, Gansu Province, Zhangye City, Sunan Co., Along the way from Sunan to Qilian, 3928 m elev., 38°37′N, 99°28′E, on rock, 30 May 2018, Li-Song Wang et al. 18-58865 (KUN–holotype).
Description
Thallus areolate, red brown, continuous, areolae 0.5–1.25 mm in diam., slightly convex, epruinose; margin pruinose, often with jade green pigmented; prothallus not distinct. Upper cortex 27.5 µm thick, orange granules pigmented; epinecral layer up to 27.5 µm thick, uneven, sometimes absent; algal layer 50.0 µm thick, cells 6.0–10.0 × 6.0–10.0 µm in diam., round; lower cortex absent. Apothecia frequent, scattered, immersed, 0.3–0.6 mm in diam.; disc red brown, concave, epruinose; margin absent. Hymenium 55.0–82.5 µm thick, colorless; paraphyses 2.5 µm wide, unbranched, not anastomosing; epihymenium 15.0–30.0 µm thick, orange; epinecral layer 5.0 µm thick, discontinuous; subhymenium 37.5 µm thick, colorless; hypothecium colorless. Asci Porpidia-type, cylindrical, eight-spored; ascospores 6.0–12.5 × 3.0–5.0 µm in diam., ellipsoid, halonate. Pycnidia stellate, strongly convex, rarely immersed, forming tuberculiform, black, margin pruinose, with jade green pigmented; conidia 3.0–4.5 × 1.0 µm in diam., oblong to bacilliform.
Chemistry
Thallus K–, C–. Medulla I+, K–, C–. None.
Ecology and distribution
Growing on the Qilian jade or sandstone at elevations of 3928–4124 m in the alpine zone. This species is known from Qinghai Province and Gansu Province of China.
Remarks: Lecaimmeria tuberculosa is characterized by its red brown thallus, the jade green pruinose at the margin of its areolae, its red brown, concave apothecia without a proper margin and tuberculiform pycnidia. L. qinghaiensis is similar to L. tuberculosa (see notes for L. qinghaiensis). L. tuberculosa usually grows on jade, and interestingly the margin of pycnidia and areolae of the species are heavily jade green pigmented. L. iranica resembles L. tuberculosa by the absence of apothecia margin, but differs in its immersed pycnidia and white margin of its areolae.
Specimens examined (KUN). CHINA, Qinghai Province, Gande Co., Qingzhen village, 4124 m elev., 34°11′N, 100°12′E, on rock, 2020, Li-Song Wang et al. 20-68077, 4145 m elev., Li-Song Wang et al. 20-68055. Gansu Province, Zhangye City, Sunan Co., along the way from Sunan to Qilian, 3928 m elev., 38°37′N, 99°28′E, on rock, 2018, Li-Song Wang et al. 18-58856, 18-58857, 18-58865, 18-59835.
Key to Lecaimmeria and allied genera
1 Apothecia lecanorine-type. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 2
Apothecia lecideine-type. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .5
2 Apothecia adnate; Isidia spherical. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .Koerberiella
Apothecia immersed; Isidia absent. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . 3
3 Thallus surface wax glossy, orange or red brown. .. .. .. .. .. .. .. .. .. .. .. .. .. Lecaimmeria
Thallus surface not glossy, white. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 4
4 Asci Aspicilia-type, ascospores non-halonate. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . Aspicilia
Asci Porpidia-type, ascospores halonate, amyloid. .. .. .. .. .. .. .. .. .. .. .. .. .. .Bellemerea
5 Epinecral layer absent. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . 6
Epinecral layer present. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 8
6 Asci Lecanora-type; often containing miriquidic acid. .. .. .. .. .. .. .. .. .. .. .. . .Miriquidica
Asci not Lecanora-type; not containing miriquidic acid. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .7
7 Asci Lecidea-type, ascospores non-halonate. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .Lecidea
Asci Porpidia-type, ascospores halonate. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. Porpidia
8 Thallus surface dark brown; Asci Lecanora-type, ascospore non-halonate. .. .. .. . .Miriquidica
Thallus surface orange or red brown; Asci Porpidia-type, ascospores halonate. .. . .Immersaria
Key to species of Immersaria in this study
1 Thallus grayish brown; Apothecia crowded. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .2
Thallus reddish brown; Apothecia rarely crowded. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .3
2 Thallus rusty; Apothecia over 1.3 mm in diam.; Epihymenium brown; growing on red sandstone. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . I. ferruginea
Thallus never rusty; Apothecia never over 1.3 mm in diam.; Epihymenium green; growing on granite. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. I. plumbeoolivacea
3 Thallus orange, areolae irregular. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . I. aurantia
Thallus not orange, areolae irregular, polygon or rectangle. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 4
4 Thallus up to 10 cm across, areolae aggregated by several smaller areolae and black prothallus. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . .I. shangrilaensis
Thallus not up to 10 cm across, areolae not aggregated. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .5
5 Thallus areolae convex, not rusty, not cracked; Ascospores over 15 µm long. .. . .I. athroocarpa
Thallus areolae flat, often rusty, cracked; Ascospores never over 15 µm long.. .. .. .. I. venusta
Key to species of Lecaimmeria in this study
1 Prothallus distinct and filled the fissures between areolae. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . .2
Prothallus only distinct at the margin. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . 4
2 Areolae margin white; Apothecia black brown; containing 4-O-demethylplanaic acid. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . L. mehadiana
Areolae margin black; Apothecia dark red brown; containing gyrophoric acid or no substance detected by TLC. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . 3
3 Thallus discontinuous, prothallus continuous, distinct and encircle every areolae; Apothecia dark red brown, margin absent. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. L. cupreoatra
Thallus continuous, prothallus discontinuous, not encircle every areolae; Apothecia dark orange, margins present. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . L. lygaea
4 Thallus orange. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . 5
Thallus reddish brown. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .6
5 Apothecia red brown; containing gyrophoric acid. .. .. .. .. .. .. .. .. .. .. .. .. . .L. mongolica
Apothecia dark orange; no substance detected by TLC. .. .. .. .. .. .. .. .. .. .. .. .. L. tibetica
6 Apothecia margin absent. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. . 7
Apothecia margin present. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 8
7 Areolae margin white; Epihymenium brown; Pycindia immersed, liner or stellate.. .. .L. iranica
Areolae margin green; Epihymenium orange; Pycindia convex, tuberculiform. . .L. tuberculosa
8 Thallus rusty; Apothecia occupy the whole areolae. .. .. .. .. .. .. .. .. .. .. .. .L. qinghaiensis
Thallus not rusty; Apothecia do not occupy the areolae. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .9
9 Apothecia irregular, crowded while immature, aggregate when mature. .. .. .. .. . L. botryoides
Apothecia round, rarely crowded, not aggregate. .. .. .. .. .. .. .. .. .. .. .. .. .. L. orbicularis