Research Article |
Corresponding author: Xin-yu Wang ( wangxinyu@mail.kib.ac.cn ) Academic editor: Gerhard Rambold
© 2023 An-cheng Yin, Qiu-yi Zhong, Christoph Scheidegger, Ji-zhen Jin, Fiona R. Worthy, Li-song Wang, Xin-yu Wang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Yin A-c, Zhong Q-y, Scheidegger C, Jin J-z, Worthy FR, Wang L-s, Wang X-y (2023) The phylogeny and taxonomy of Glypholecia (Acarosporaceae, lichenized Ascomycota), including a new species from northwestern China. MycoKeys 98: 153-165. https://doi.org/10.3897/mycokeys.98.104314
|
Glypholecia qinghaiensis An C. Yin, Q. Y. Zhong & Li S. Wang is described as new to science. It is characterized by its squamulose thallus, compound apothecia, ellipsoid ascospores, and the presence of rhizines on the lower surface of the thallus. A phylogenetic tree of Glypholecia species was constructed based on nrITS and mtSSU sequences. Two species G. qinghaiensis and G. scabra are confirmed in China.
cosmopolitan, lichenized fungi, morphological diversity, phylogenetic analyses, Tibetan Plateau
Glypholecia Nyl. is a genus of lichenized fungi belonging to Acarosporaceae, Acarosporales, Acarosporomycetidae, Lecanoromycetes, Ascomycota (
The first known specimen of the genus now designated Glypholecia, was collected in the summer of 1810, by Balbis G. B., who was working at the Botanical Gardens of Turin, Italy. Balbis sent this lichen specimen to Germany, where it was received by Funck H. C., who labelled the specimen as Balbis. This name was never published. Later, Persoon H. C. examined this same specimen in France, and described it as a new species Urceolaria scabra Pers. (
In 1814, Acharius E. published the lichen species Lecanora rhagadiosa Ach (
In 1853, Nylander W. established the monotypic genus Glypholecia, based on the type species Glypholecia candidissima Nyl. (
Accordingly, most lichenologists accept only one species G. scabra (Pers.) Müll. Arg. as belonging to the genus Glypholecia. The distribution of G. scabra is largely disjunct, with intercontinental populations. It occurs on siliceous rocks in desert, alpine regions, including Europe, Africa, Asia, and North America (
Previously, a second species, G. tibetanica H. Magn., was described as endemic to China. It has only been recorded from Xizang province, China (Zahlbruckner 1933). However,
During the Second Tibetan Plateau Scientific Expedition and Research Program (STEP), we collected numerous lichen specimens from across northwestern China. The expedition included many field surveys across the type locality of G. tibetanica, during which we discovered some specimens that conformed to Zahlbruckner’s (1932) description. Our new specimens differ from A. nodulosa var. reagens in the characteristics of their upper cortex, hymenium chemical reaction and their number of ascospores. The molecular sequences obtained from these new specimens demonstrate that they belong to the genus Glypholecia, rather than to Acarospora (
We examined materials of Glypholecia from the lichen herbaria of the Kunming Institute of Botany (KUN-L), the College of Life Science and Technology, Xinjiang University (
We extracted total genomic DNA from 20 specimens (19 from China and one from Switzerland) using the DNAsecure Plant Kit (Tiangen Biotech, Beijing) following the manufacturer’s protocol. We amplified the internal transcribed spacer regions (nrITS) with the primer pairs ITS1F (
We aligned DNA sequences using the program MAFFT v. 7.107 in GENEIOUS v. 8.0.2, setting the following parameters: algorithm = auto; scoring matrix = 200 PAM / k=2; gap open penalty = 1.53; offset value = 0.123 (
In the present study we generated twenty new nrITS and eighteen new mtSSU sequences. We constructed ML and BI topologies based on these nrITS and mtSSU sequences, and nine additional sequences downloaded from GenBank (Table
Specimens and sequences used for phylogenetic analyses. Newly generated sequences are in bold.
Taxon | Locality | Voucher specimens | GenBank number (nrITS) | GenBank number (mtSSU) | References |
---|---|---|---|---|---|
G. qinghaiensis | Ningxia, China | KUN-L 10-0241 | MZ330798 | OP749902 | – |
Gansu, China | KUN-L 18-58434 | MZ330797 | OP749903 | – | |
Gansu, China | KUN-L 18-59534 | MZ330793 | OP749907 | – | |
Qinghai, China | KUN-L 20-68255 | MZ330789 | OP749910 | – | |
Xinjiang, China | KUN-L 22-71630 | OP749916 | OP749899 | – | |
G. scabra | Neimenggu, China |
|
MZ330786 | – | – |
Gansu, China | KUN-L 18-58747 | MZ330796 | OP749904 | – | |
Qinghai, China | KUN-L 18-59190 | MZ330792 | OP749906 | – | |
Gansu, China | KUN-L 18-58881 | MZ330795 | OP749905 | – | |
Xizang, China | KUN-L 19-65418 | MZ330791 | OP749908 | – | |
Xizang, China | KUN-L 19-66159 | MZ330790 | OP749909 | – | |
Xinjiang, China | KUN-L XY22-856 | OP749911 | OP749895 | – | |
Xinjiang, China | KUN-L XY22-856-2 | OP749912 | OP749896 | – | |
Xizang, China | KUN-L XY22-584 | OP749913 | OP749894 | – | |
Xinjiang, China | KUN-L 22-72868 | OP749914 | OP749897 | – | |
Xinjiang, China | KUN-L 22-71693 | OP749915 | OP749898 | – | |
Xizang, China | KUN-L 22-71500 | OP749917 | OP749900 | ||
Xizang, China | KUN-L 22-71500-2 | OP749918 | OP749901 | – | |
Xizang, China | KUN-L 22-71435 | OP749919 | OP749893 | – | |
Canton of Valais, Switzerland | Scheideg-10522 | MZ330788 | – | – | |
Unknown | AFTOL 1008 | HQ650722 | – |
|
|
Oppland, Norway | S. Westberg 08-232 | LN810811 | LN810936 |
|
|
Acarospora sp. | Gansu, China | Huang Manrong GS157 | FJ919810 | – |
|
A. placodiiformis | Spain, Madrid | Westberg 10-211 | LN810795 | LN810920 |
|
A. schleicheri | Bouches-du-Rhône, France | DUKE Reeb VR5-VII-98/30 | DQ525529 | – |
|
Sichuan, China | UPS L-070426 | LN810800 | LN810925 |
|
|
Arizona, USA | UPS L-162697 | LN810801 | LN810926 |
|
|
Pleopsidium flavum | Steiermark, Austria | UPS L-105590 | AY853385 | AY853336 |
|
P. chlorophanum | Jämtland, Sweden | UPS L-179248 | LN810813 | LN810938 |
|
Our phylogenetic results indicated that Glypholecia is monophyletic in China. The genus Glypholecia is characterized by its squamulose to crustose thallus, compound apothecia forming multiple structures, multi-spored asci (usually exceeding 30 spores per ascus), spherical small ascospores and C+ red reaction of the cortex due to the presence of gyrophoric acid.
Species of Glypholecia were separated into two main clades, as inferred from the phylogenetic tree with strong support. Based on the combination of morphological characters and phylogenetic analysis, we propose a new species in Glypholecia. We divide the specimens of the genus Glypholecia collected from China into two clades, which correspond to differences in the lower surface morphology of the thallus: the specimens in Clade 1 have ellipsoid ascospores (2.5–3 × 4–6.5 μm), and rhizines on the lower surface of the thallus, whereas Clade 2 contains specimens with spherical ascospores (3–4 µm), and a central holdfast, called an umbilicus. Within Clade 2, our samples of G. scabra collected from China were clustered with the European samples, but with some genetic divergence. All Glypholecia specimens which we collected from the type locality of “G. tibetanica” were monophyletic with G. scabra, with a high support value. Within Clade 1, those specimens assigned as G. qinghaiensis lack genetic variability within the gene regions included in this study. This might be due to either the sparse population or the shared geography and environment. All of our G. qinghaiensis specimens were collected from arid to semi-arid areas of northwestern China, usually growing on sandy rock or sandy soil.
Our phylogenetic analyses showed that a specimen from Gansu province in China, which was identified as G. scabra by
Resembles G. scabra, has abundant and compound apothecia, but differs in having ellipsoid ascospores, rhizines, and a different molecular fingerprint (based on nrITS and mtSSU data).
China, Qinghai Prov., Haixi Mongolian and Tibetan Autonomous Prefecture, Dulan Co., 3066 m, on sandy soil, 2020-09-15, Wang Lisong et al. 20-68255 (holotype – KUN).
Thallus squamulose to squamulose-subfoliose, peltate-subumbilicate, up to 6 mm in diam., 6.5–7 mm thick, with margins sometimes rolling under; surface: upper surface white, pale brown to brown, cracked and wrinkled, usually warty, partly pruinose or occasionally densely pruinose at margins; lower surface white, gray to pale brown, rough, wrinkled, rhizines, umbilicate, 3–6 mm long, attached with a broad; upper cortex: paraplectenchymatous, pale brown, containing calcium oxalate crystals, 30–70 μm thick; medulla: pale, containing calcium oxalate crystals, 125–250 μm thick; hyphae loose, arachnoid, 2.5–3 μm. Apothecia very common, lecanorine, punctiform when young, but later becoming compound reddish brown to dark brown, becoming lower than the thallus surface when mature; disc with cracks and sometimes tuberculous; paraphyses septate, 2–2.5 μm in diam.; margins concolorous with the thallus; asci: clavate, c. (30–)50-spored; ascospores: ellipsoid, 4–6.5 × 2.5–3 μm, hyaline, thick wall. Pycnidia: rare, flask-shaped. Conidia bacilliform, c. 2–3 × 1 µm.
Hymenium: I+ blue; cortex and medulla: K–, C+ red, KC+ red, P–; secondary metabolites: gyrophoric acid.
Usually on sandy rocks or rarely on soil over rocks; so far only known from Gansu, Ningxia, Qinghai and Xizang provinces in China; growing in arid to semi-arid areas; distributed from 1600 to 4700 m altitude.
The epithet “qinghaiensis” refers to the holotype locality of the species.
Glypholecia qinghaiensis can be distinguished from G. scabra by having ellipsoid ascospores and rhizines. Phylogenetic analysis in this study supports the separate classification of these two species. This species has only been reported from northwestern China, including Gansu, Ningxia, Qinghai and Xizang provinces.
China (stored in KUN). Gansu Prov.: Jiuquan City, Yumen City, Yuerhong Vil., 3044 m, on rock, 2018-05-27, Wang Lisong et al. 18-59534; Subei Mongolian Autonomous Co., Suyan Line, 2376m, on soil over rock, 2018-05-23, Wang Lisong et al. 18-58434; Ningxia Prov.: Zhongwei Co., Suwumuyang Site, 1611 m, on rock, 2010-09-19, Niu Dong-Ling et al. 10-0241. Qinghai Prov.: Dulan Co., 3066 m, on sandy rock, 2020-09-15, Wang Lisong et al. 20-68255; Xinjiang Prov.: Wuqia Co., Fossil hill, 2559 m, on sandy rock, 2022-06-26, Wang Lisong et al. 22-71630.
≡ Urceolaria scabra Pers., Ann. Wetter. Gesellsch. Ges. Naturk. 2: 10, 1810.
Monte Cenisio [in Alps between France and Italy], s. d., Balbis s. n. (not seen).
Glypholecia scabra is characterized by its squamulose thallus, abundant and compound apothecia, and umbilicate lower surface, as shown in Fig.
Morphological diversity within the genus Glypholecia A–F Glypholecia scabra. Differing morphology putatively caused by different habitat conditions, e.g., high or low temperature, aridity, different altitudes (1300–5100 m) G–I G. tibetanica. Collected from type locality, thallus with numerous pycnidia. J-F Umbilicus at the lower surface, formed by fasciculate white rhizines A, J Wang Lisong KUN 18-58925 B Wang Lisong KUN 18-58814-b C Wang Lisong KUN 18-58820 D Wang Lisong KUN 19-65418 E, K Wang Lisong KUN 18-59346 F Wang Lisong KUN 18-58747 G Wang Xinyu KUN XY22-854 H Wang Lisong KUN 22-71500 I, L Wang Xinyu KUN XY22-856. Scale bars: 1 mm.
China. Gansu Prov. (stored in KUN): Zhangye City, Sunan Yugur Autonomous Co., on the way from Sunan to Qilian, hinterland of Qilian Mt., 3958 m, on rock, 2018-05-30, Wang Lisong et al. 18-58881; near Binggoudanxia Geopark, 1984 m, on rock, 2018-05-29, Wang Lisong et al. 18-58747. Qinghai Prov. (stored in KUN): Hainan Tibetan Autonomous Region, Gonghe Co., Heimahe Vil., 3429 m, on rock, 2018-05-19, Wang Lisong et al. 18-59190; Xining City, Huangyuan Co., on the way from Xining to Qinghai Lake, 2476 m, on rock, 2018-05-18, Wang Lisong et al. 18-59094. Neimenggu Prov. (stored in
Switzerland (stored by C. Scheidegger). Canton of Valais: Evolène, Mount Le Tsaté., 2492 m, on calcareous rocks, s. d., C. Scheidegger Scheideg-10522.
Aksai-Chin-Plateau, [in Xizang Prov., China], (in Botanischer Garten Zürich, holotype, lost).
These specimens were collected by Walter Bosshard in 1927 in Ritu County, Xizang Province, and then reported by H. Magnusson as a new species G. tibetanica, characterized by its squamulose thallus, C+ red cortex, abundant black pycnidia and cylindroid conidia, 3.5–4.5 × 1 μm, but the cited type specimen was sterile (Zahlbruckner 1933). Its characteristic of abundant pycnidia is not seen in G. qinghaiensis.
We found some Glypholecia specimens with numerous pycnidia, as shown in Fig.
The authors are very grateful to Hurnisa Xahidin (Xinjiang University) for providing sequence data for this study. This research was supported by the Second Tibetan Plateau Scientific Expedition and Research Program (STEP) (2019QZKK0503), grants from the Flora Lichenum Sinicorum (31750001), Youth Innovation Promotion Association CAS (2020388), Yunnan Young and Elite Talents Project and National Natural Science Foundation of China (Nos. 31970022, 31670028, 32060001).
The authors have declared that no competing interests exist.
No ethical statement was reported.
This research was funding from the Flora Lichenum Sinicorum (31750001), Youth Innovation Promotion Association CAS (2020388), Yunnan Young and Elite Talents Project and National Natural Science Foundation of China (Nos. 31970022, 31670028, 32060001), Second Tibetan Plateau Scientific Expedition and Research Program (STEP) (2019QZKK0503).
All authors contributed to this work.
An-cheng Yin https://orcid.org/0009-0000-7837-7797
Qiu-yi Zhong https://orcid.org/0000-0001-6256-6083
Christoph Scheidegger https://orcid.org/0000-0003-3713-5331
Ji-zhen Jin https://orcid.org/0009-0005-4351-8162
Fiona R. Worthy https://orcid.org/0000-0003-0042-3110
Li-song Wang https://orcid.org/0000-0003-3721-5956
Xin-yu Wang https://orcid.org/0000-0003-2166-6111
All of the data that support the findings of this study are available in the main text or Supplementary Information.