2018, Vol. 26
The genus Coenogonium Ehrenb. including more than 130 species which are mainly found in tropical and subtropical habitats, belonging to the lichenized fungi family Coenogoniaceae (Ascomycota, Lecanoromycetes, Ostropomycetidae, Ostropales), is characterized by crustose or filamentous thallus, biatorine (rarely zeorine), yellow to orange or brown apothecia with paraplectenchymatous excipulum, partially amyloid hymenium (I+ blue then quickly sordid green then red-brown), 8-spored, thin-walled unitunicate asci, 1-septate or rarely non-septate hyaline ascospores and trentepohlioid photobiont. The Coenogonium usually grow on organic substrata, including bark, bryophytes and leaves, and many species are typically or at least facultatively foliicolous[1-6]. It was established by Ehrenberg in 1820 based on type species C. linkii Ehrenb., originally only including species with filamentous thallus, has recently been emended to include Dimerella Trev. which shares all apothecial characters with Coenogonium and differs only by the crustose thallus[3]. Based on DNA sequences of the nuclear small and large subunit ribosomal RNA genes, molecular studies supported the synonymy of Coenogonium and Dimerella[7]. Recent evidence based on ontogenetic, anatomical and morphological studies also showed that the combination of the two different morphotypes into same family Coenogoniaceae is natural and accurate[7-10].
In China, nine species of Coenogonium s.l. (including Dimerella) were sporadically reported[11-27]. A new record species Coenogonium disjunctum is found in our study, and all the ten species are described in the present paper.
The specimens are deposited in the Herbarium Mycologicum Academiae Sinicae-Lichenes (HMAS-L), the Lichen Herbarium of the College of Life Sciences, Liaocheng University (LCU) and the Herbarium, Kunming Institute of Botany (KUN-L). Stereomicroscope (OLYMPUS SZX12) and a light microscope (OLYMPUS BX53) were used for the morphological and anatomical studies. Sections of thallus and ascomata were mounted in water, 10% KOH (K) and Lugol's solution (I), respectively. The lichen substances were examined by thin-layer chromatography[28-30].
1 Coenogonium dilucidum(Kremp.) Kalb & Lücking亮绒衣In Lücking & Kalb, Bot. Jb. 122 (1): 32 (2000).
≡ Lecidea dilucida Kremp., J. Mus. Godeffroy 1 (4): 103 (1873). — Dimerella dilucida (Kremp.) R. Sant., Symb. Bot. Upsal. 12 (1): 394 (1952).
Thallus foliicolous, rarely corticolous, glabrous. Apothecia small, rounded, 0.1-0.4 mm in diam., disc pale wax-colored. Ascospores broadly ellipsoid, 1-septate, 10-12.5 μm×2.5 μm, about 4-5 times as long as wide, usually uniseriate. Pycnidia not found.
Chemistry: No lichen compounds detected.
Substrate: On leaves.
Distribution and ecology: Pantropical[4]. The species is a foliicolous (on living leaves) crustose species previously reported from Hong Kong[22] (Aptroot et al. 2001, as Dimerella dilucida)[23]. Worldwide, it was also reported from Thaliand[31], Vietnam[32] and South America[6].
Specimens examined: China. Hainan: Changjiang County, Mt. Bawangling, Yajia, 820 m, 26/XI/ 2010, J. C. Wei 54 (HMAS-L 119014), 55 (HMAS-L 119012), 51 (HMAS-L 119017); Ledong County, Mt. Jianfengling, elevation unrecorded, 27/XI/2010, J. C. Wei & W. Guo 71 (HMAS-L 119032); Lingshui County, Mt. Diaoluoshan, elevation unrecorded, 01/Ⅻ/2010, J. C. Wei & W. Guo 282 (HMAS-L 119045). Guangxi: Shangsi County, Mt. Shiwandashan, 463 m, J. Li, GX15221 (LCU).
Notes: Coenogonium dilucidum is similar to C. subluteum in morphology, but differs in having more brightly orange apothecia and rare or no pycnidia. However, pycnidia are usually abundant in C. subluteum, contributing to a verrucose appearance in thallus.
2 Coenogonium disjunctum Nyl.疏羽绒衣Fig. 1: A-C
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Fig. 1 Coenogonium disjunctum (L. S. Wang & X. Y. Wang 15-49036). A Thalli; B: Section of ascomata; C: An ascus with 8 ascospores. |
Bot. Zeitschr. 20: 178 (1862).
Thallus filamentous, sometimes forming closely appressed mat, loosely or closely attached to substrate, prostrate or pannose, yellow-green. Apothecia rounded, substipitate, 0.3-0.6 mm in diam., 150-200 μm high. Disc flat and slightly concave, yellow-orange, margin thick, cream-colored, smooth. Exciple colorless, 6070 μm high. Hypothecium 30-50 μm high, palebrown. Hymenium 50-70 μm high, colorless, I+ blue. Asci 8-spored, clavate, 57.5-72.5 μm×5-7 μm. Ascospores irregularly biseriate, ellipsoid, 1-septate, 10-14 μm×2.5 μm. Pycnidia not found.
Chemistry: No lichen compounds detected.
Substrate: On bark.
Specimens examined: China. Taiwan: Taizhong, Guguan, Mt. Baxian, 1 053 m, 19/Ⅸ/2015, L. S. Wang & X. Y. Wang 15-49036 (KUN-L 52034).
Distribution and ecology: Tropical[4]. Worldwide, it was reported from the United States[33]. It is the first report from China.
Notes: Coenogonium disjunctum is characterized by filamentous, prostrate or pannose thallus, rounded and substipitate apothecia, and irregularly biseriate ascospores, 10-14 μm×2.5 μm in size. It is similar to C. interplexum, in filamentous thallus, and in some extent, they are difficult to be distinguished from the outer morphology. But C. disjunctum has much larger ascospores (10-14 μm×2.5 μm) than C. interplexum (6-9 μm×2.5-3.75 μm). It is also similar to C. implexum, but the apothecia of C. implexum are sessile.
3 Coenogonium interplexum Nyl.网绒衣Annls Sci. Nat., Bot., sér. 4 16: 92 (1862).
Thallus filamentous, prostrate and effuse, irregularly woven, with large interspaces between individual filaments, yellow-green. Apothecia rounded, 0.3-1.0 mm in diam., substipitate, 130-150 μm high; disc flat, yellow-orange; margin thin, not prominent, smooth, cream-colored. Asci 60-70 μm×4-6 μm. Ascospores uniseriate, ellipsoid, 1-septate, 6-9 μm× 2.5-3.75 μm, 2-3 times as long as wide. Pycnidia not found.
Chemistry: No lichen compounds detected.
Substrate: On bark and on rock.
Specimens examined: China. Yunnan: Simao County, Laiyang River, 1 570 m, 07/Ⅱ/2009, L. S. Wang 09-30131 (KUN-L); Puer, Lahu Autonomous County of Lancang, Macheng Village, 1 450 m, 14/ XI/2014, L. S. Wang & M. X. Yang 15-49918 (KUNL 53197). Taiwan: Taizhong, Guguan, Mt. Baxian, 1 053 m, 19/Ⅸ/2015, L. S. Wang & X. Y. Wang 1549032 (KUN-L 52030).
Distribution and ecology: Pantropical[4]. The species was previously reported from Taiwan[14-15]. Worldwide, it was also reported from North, Central and South American, Thaliand and Australia[31, 34].
Notes: Coenogonium interplexum is distinguished by the prostrate thallus formed from irregularly woven filaments which is not easily detached from the substrate (mostly bark). Compared to C. implexum, it has shorter and narrower paraphyses. Besides, the main differences between the two species are the pannose thallus with very densely arranged filaments[35] and the larger ascospores in C. implexum.
4 Coenogonium isidiatum (G. Thor & Vězda) Lücking, Aptroot & Sipman裂芽绒衣In Rivas Plata, Lücking, Aptroot, Sipman, Chaves, Umaña & Lizano, Fungal Diversity 23: 297 (2006).
≡ Dimerella isidiata G. Thor & Vězda, Folia Geobot. Phytotax. 19(1): 72 (1984).
Thallus crustose, corticolous, continuous, thin, smooth, partly shiny, yellowish green to greenish grey, 5-30 mm in diam.; prothallus absent, isidia crowded, 0.2-0.4 mm high, 0.05 mm in diam., simple or rarely branched, covered by colorless, warty papillae. Apothecia sessile, rounded in outline, 1.2-1.5 mm in diam., 380-410 μm high; disc plane, later slightly convex, orange-red; margin thick, prominent. Asci 80-95 μm×6-10 μm. Ascospores oblique-uniseriate, narrowly ellipsoid, 1-septate, 9-14 μm×3-4 μm, 3-4 times as long as wide. Pycnidia not found.
Chemistry: No lichen compounds detected.
Substrate: On bark and on rock.
Specimens examined: No specimens are listed here and the original materials have not been studied. The descriptions and notes are cited from literatures [26, 36].
Distribution and ecology: Pantropical[4]. The species was previously reported from Xizang and Sichuan[26]. Worldwide, it was known from the Neotropics and eastern Paleotropics[4], and North America[36].
Notes: Coenogonium isidiatum is characterized by large (up to 2.5 mm), flat, orange apothecia with a slightly pale margin, and dispersed, rarely branched, up to 0.3 mm long isidia. It is similar to C. isidiosum, but its ascospores (2.5-3 μm wide) are little narrower than C. isidiatum (3-4 μm wide).
5 Coenogonium leprieurii (Mont.) Nyl.勒氏绒衣Ann. Sci. Nat. Bot. sér. 4 16: 89, pl. 12, fig. 15-19 (1862).
≡ Coenogonium linkii var. leprieurii Mont., Annls Sci. Nat., Bot., sér. 3, 16: 47 (1851).
Thallus filamentous, mostly corticolous, occasionally foliicolous, on old perennial leaves, usually forming hemispherical and shelf-like structures projecting horizontally from the substrate, yellowgreen to bright green, up to 40 mm across, ecorticate. Apothecia rounded, 0.25-0.6 mm in diam., substipitate or conical, 100-150 μm high; disc flat, pale orange to yellow-orange; margin thin, not prominent, smooth, cream-colored. Ascospores irregularly biseriate, ellipsoid, non-septate, 5-8 μm×2-3 μm, 2.5-3 times as long as wide.
Chemistry: No lichen compounds detected.
Substrate: On bark and on rock.
Specimens examined: China. Yunnan: Puer, National Forest Park, 1 596 m, 18/Ⅻ/2013, X. Y. Wang & D. Liu 13-41444 (KUN-L 22326), Fujian: Wuyishan, Mt. Wuyi, Taoyuanyu, 640 m, 28/Ⅴ/2015, L. S. Wang & X. Y. Wang 15-47138 (KUN-L 50025), 15-47157 (KUN-L 50044), 15-47142 (KUN-L 50029); L. S. Wang & X. Y. Wang 15-47121 (KUN-L 50008), 15-47133 (KUN-L 50020); Wuyishan, Mt. Wuyi, Pikeng, 430 m, 25/Ⅴ/2015, L. S. Wang & X. Y. Wang 15-46799 (KUN-L 49686), 15-46759 (KUN-L 49646), 15-46746 (KUN-L 49633).
Distribution and ecology: Pantropical[4]. The species was previously reported from Fujian[11]. Worldwide, it was reported from South and Central America, Thailand[31].
Notes: Coenogonium leprieurii is characterized by its filamentous and shelf-shaped thallus which projects from the substrate, apothecia on the underside of the thallus and non-septate ascospores[5]. It is very similar to C. linkii, but distinguished in non-septate ascospores.
6 Coenogonium linkii Ehrenb.林氏绒衣In Nees von Esenbeck (ed.), Horae Phys. Berol.: 120, pl. 27 (1820).
= Coenogonium boninense Sato, J. Japan. Bot. 8: 390 (1933).
Thallus filamentous, very regularly shelf-like, rather thin and compact, with few and narrow interspaces between individual filaments, usually bright green, ecorticate. Apothecia rounded, 0.2-1.0 mm in diam., substipitate, 150-200 μm high; disc flat to convex, yellow-orange; margin thin, not prominent, smooth, cream-colored. Ascospores irregularly biseriate, ellipsoid, 1-septate, 6.5-9 μm×2-2.5 μm, 2.5-3.5 times as long as wide. Pycnidia not found.
Chemistry: No lichen compounds detected.
Substrate: On bark.
Specimens examined: China. Guangxi: Jingxi County, Tongling Cayon, 511 m, 06/Ⅵ/2011, L. S. Wang 11-32199, 11-32209 (KUN-L). Hainan: Ledong County, Mt. Jianfengling, 920 m, 11/Ⅻ/2014, R. D. Liu & J. H. Wang HN2014187 (HMAS-L 0132558).
Distribution and ecology: The species was previously reported from Taiwan[12-15]; (Yoshimura 1994, as C. boninense)[19] and Hong Kong[22] (Seaward & Aptroot 2005, as C. cf. linkii)[27]; (Pfister 1978, as C. disjunctum)[16]; (Aptroot et al. 2001, as C. linkii)[23]. Worldwide, it distributes in Neotropical lowland rain forests[4] and the United States[33].
Notes: Coenogonium linkii is characterized by filamentous and shelf-like thallus, rounded, yelloworange disc and smooth margin, substipitate apothecia and the ascospores irregularly biseriate, 1-septate, 5-7 μm×1.5-2.5 μm. It is very similar to C. leprieurii, but C. leprieurii has non-septate ascospores.
7 Coenogonium luteum (Dicks.) Kalb & Lücking金黄绒衣In Lücking & Kalb, Bot. Jb. 122 (1): 32 (2000).
≡ Lichen luteus Dicks., Fasc. Pl. Crypt. Brit. (London) 1: 11 (1785). — Dimerella lutea (Dicks.) Trevis., Rend. Reale Ist. Lomb., Milano, sér. 2: 129 (1836).
Thallus crustose, corticolous, continuous, thin, smooth, pale greenish. Apothecia sessile, rounded to slightly irregular, 0.5-2 mm in diam; disc concave, orange; margin thin, slightly prominent, smooth, creamcolored. Ascospores irregularly uniseriate, broadly ellipsoid, 1-septate, 7.5-8.75 μm×2.5 μm, 3-3.5 times as long as wide.
Chemistry: No lichen compounds detected.
Substrate: On bark.
Specimens examined: China. Hainan: Changjiang County, Mt. Bawangling, Yajia, 800 m, 26/XI/ 2010, S. N. Cao CSN067 (HMAS-L 118099). Fujian: Wuyishan, Mt. Longjing, 200 m, 22/Ⅶ/2016, Q. D. Wang FJ16004, FJ16005, FJ16006 (LCU); Mt. Wuyi, Zhongzheng Park, 400 m, 23/Ⅶ/2016, Z. F. Jia FJ16122, FJ16123, FJ16134, FJ16135, FJ16136 (LCU). Anhui: Huangshan, Mt. Huangshan, Banshan Temple, 1 440 m, X. H. Wu AH17146 (LCU). Hainan: Lingshui County, Mt. Diaoluo, 970 m, 01/Ⅻ/2010, Z. F. Jia 10-644 (HMAS-L 117951).
Distribution and ecology: The species was previously reported from Hubei[17], Zhejiang[37], Fujian[11] and Hong Kong[22], (Seaward & Aptroot 2005, as Dimerella lutea)[27]; (Pfister 1978, as Gyalecta lutea)[16]; (Aptroot et al. 2001, as D. lutea)[23]. Corticolous and muscicolous in upper montane rain forest and subparamo, widespread in temperate regions[4].
Notes: Coenogonium luteum is characterized by large apothecia with more or less concave and orange disc, excipulum with rather small, thick-walled cells, and broadly ellipsoid ascospores. The most similar species are C. eximium (with ascospores 2-2.5 μm wide) and C. fallaciosum (with apothecia plane, pale yellow to pale orange).
8 Coenogonium pineti (Ach.) Lücking & Lumbsch皮氏绒衣In Lücking, Stuart & Lumbsch, Mycologia 96 (2): 290 (2004).
≡ Lecidea pineti Ach., Lich. Univ.: 195 (1810). — Dimerella pineti (Ach.) Vězda, Lichenes Selecti Exsiccati: no. 1279 (1975).
Thallus crustose, mostly corticolous, also on mosses, continuous, sometimes barely visible, smooth, greyish green to greenish black, 5-20 mm in diam., without a distinct corticiform layer; prothallus absent. Apothecia sessile, rounded, 0.2-0.5 mm diam.; disc plane to concave, white to pinkish white, very rarely with an orange tinge; margin prominent, relatively thick, whitish to cream-colored. Asci 65-75 μm×911 μm. Ascospores (irregularly) uniseriate, ellipsoid, 1-septate, 9-14 μm×2.3-4.5 μm, 2.5-3.5 times as long as wide. Pycnidia wart-shaped, 0.1-0.2 mm in diam., whitish. Conidia oblong, non-septate, 6-8 μm×1.8-2.6 μm.
Chemistry: No lichen compounds detected.
Substrate: On bark.
Specimens examined: No specimens are listed here and the original materials have not been studied. The descriptions and notes are cited from literatures.
Distribution and ecology: The species was previously reported from Taiwan (Aptroot and Sparrius, 2003, as D. pineti)[25] and Hong Kong (Aptroot et al. 2001, as D. pineti)[23]. Cosmopolitan with a preference for temperate regions, rare in the tropics[4, 31]. Worldwide, it was reported from South Korea[34] and northeastern United States[38].
Notes: Coenogonium pineti is distinguished by small, pale apothecia with a wide margin and relative large, and broadly ellipsoid ascospores. It is similar to C. stramineum, but differs in having apothecia with a smooth margin (denticulate in C. stramineum) and asci with uniseriate ascospores (biseriately arranged in C. stramineum)[31].
9 Coenogonium subluteum (Rehm) Kalb & Lücking亚深黄绒衣In Lücking & Kalb, Bot. Jb. 122(1): 34 (2000).
≡ Biatorina sublutea Rehm, Philipp. J. Sci., C, Bot. 8 (5): 404 (1913).
= Biatorinopsis epiphylla Müll. Arg., Flora, Regensburg 64(7): 103 (1881). — Dimerella epiphylla (Müll. Arg.) Malme, Ark. Bot. 26A (13): 9 (1935) [1934]. — Microphiale epiphylla (Müll. Arg.) Zahlbr., Cat. Lich. Univers. 2: 696 (1924).
= Microphiale brachyspora (Müll. Arg.) Zahlbr., in Rechinger, Denkschr. Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 88: 22 (1911). — Biatorinopsis brachyspora Müll. Arg., Lichenes Epiphylli Novi: 16 (1890).
= Microphiale lutea f. foliicola Zahlbr., in Rechinger, Denkschr. Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 81: 247 (1907).
Thallus crustose, foliicolous, rarely muscicolous, continuous, thin, smooth, green, greenish grey or brownish grey, dull. Apothecia sessile, rounded, 0.20.5 mm in diam.; disc plane or concave, pale waxcolored to pale orange; margin comparatively thick, prominent, smooth, concolorous to disc. Ascospores uniseriate or irregularly biseriate, ellipsoid, 1-septate, 6-12 μm×3-3.5 μm, 2-3 times as long as wide.
Chemistry: No lichen compounds detected.
Substrate: On living leaves.
Specimens examined: China. Yunan: Mengla County, Wangtianshu, 700 m, 19/Ⅻ/2013, X. Y. Wang & D. Liu 13-41532 (KUN-L 22414), 13-41528 (KUN-L 22410); 691 m, 05/Ⅵ/2013, L. S. Wang & X. Y. Wang 13-38032 (KUN-L).
Distribution and ecology: Pantropical[4]. The species was previously reported from Yunnan (Zahlbrückner 1930, as Microphiale brachyspora, M. epiphylla, and M. lutea f. foliicola)[11, 18, 20, 24], Taiwan[24] and Hong Kong (as D. epiphylla)[22-23]. Worldwide, it was reported from Thaliand[31].
Notes: Coenogonium subluteum is characterized by crustose and foliicolous thallus, sessile and rounded apothecia, 0.2-0.5 mm in diam., and uniseriate or irregularly biseriate, ellipsoid, ascospores 1-septate, 6-12 μm×3-3.5 μm. It is similar to C. zonatum, but differs in having a thallus with white prothallus and very rare pycnidia[4, 31].
10 Coenogonium zonatum (Müll. Arg.) Kalb & Lücking带绒衣In Lücking & Kalb, Bot. Jb. 122(1): 34 (2000).
≡ Biatorinopsis zonata Müll. Arg., Lich. Epiphylli Novi: 16 (1890).
Thallus glabrous, foliicolous or corticolous, smooth, with white prothallus. Apothecia biatorine, lacking thalline margin, epiphyllous; Apothecia medium-sized to large, 0.3-0.8 mm diam. Disc wax-colored to pale yellow or pale orange, margin smooth; ascospores broadly ellipsoid, usually uniseriat, 1-septate, 6-12 μm×1.5-4 μm. Conidia unknown.
Chemistry: No lichen compounds detected.
Substrate: On living leaves.
Specimens examined: No specimens are listed here and the original materials have not been studied. The descriptions and notes are cited from literature [6].
Distribution and ecology: Pantropical[4]. The species was previously reported from Yunnan[24]. Worldwide, it was reported from southern South America[6].
Notes: Coenogonium zonatum is characterized by foliicolous or corticolous thallus, wax-colored to pale yellow or pale orange apothecia, smooth margin, and broadly ellipsoid, usually uniseriate ascospores, 6-12 μm×1.5-4 μm. It is similar to C. subluteum, but the latter species lacks prothallus and has abundant pycnidia[4, 31].
Key to species of the genus Coenogonium from China
1a. Isidia present; ascospores 1-septate, 9-14 μm×3-4 μm ·········································································· C. isidiatum
1b. Isidia absent································································ 2
2a. Thallus filamentous; ascospores 1-septate or non-septate······································································· 3
2b. Thallus crustose or glabrous; ascospores 1-septate······································································· 6
3a. Thallus prostrate and effuse························································· 4
3b. Thallus shelf-like·········································································5
4a. Ascospores 1-septate, 6-9 μm×2.5-3.75 μm······································································ C. interplexum
4b. Ascospores 1-septate, 10-14 μm×2.5 μm······································································ C. disjunctum
5a. Ascospores uniseriate, non-septate, 5-8 μm×2-3 μm ········································································ C. leprieurii
5b. Ascospores biseriate, 1-septate, 6.5-9 μm×2-2.5 μm······································································ C. linkii
6a. Foliicolous·································································7
6b. Corticolous·································································9
7a. Apothecia shorter, 0.1-0.4 mm in diam; ascospores 10-12.5 μm×2.5 μm········································································· C. dilucidum
7b. Apothecia longer, 0.3-2 mm in diam; ascospores 6-12 μm×1.5-4 μm········································································· 8
8a. Thallus with white prothallus; pycnidia rare; ascopores 6-12 μm×3-3.5 μm········································································· C. subluteum
8b. Thallus without prothallus; pycnidia frequent; ascopores 6-12 μm×1.5-4 μm············································································ C. zonatum
9a. Apothecia orange, 0.5-2 mm in diam.; ascopores 7.5-8.75 μm×2.5 μm············································································· C. luteum
9b. Apothecia pale yellow to pale orange-yellow, 0.2-0.5 mm in diam.; ascopores 9-14 μm×2.3-4.5 μm························· C. pineti
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