Research Article |
Corresponding author: Li-Song Wang ( wanglisong@mail.kib.ac.cn ) Academic editor: Ester Gaya
© 2020 Yan-Yun Zhang, Xin-Yu Wang, Li-Juan Li, Christian Printzen, Einar Timdal, Dong-Ling Niu, An-Cheng Yin, Shi-Qiong Wang, Li-Song 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:
Zhang Y-Y, Wang X-Y, Li L-J, Printzen C, Timdal E, Niu D-L, Yin A-C, Wang S-Q, Wang L-S (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
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New collections of six Squamarina species from type localities in China were studied. The comparison of morphological characteristics and secondary metabolites with those of the type specimens and phylogenetic analyses suggest that S. callichroa and S. pachyphylla belong to Rhizoplaca, S. semisterilis belongs to Lobothallia and S. chondroderma should be retained in Lecanora temporarily. Only two species, S. kansuensis and S. oleosa, remain in Squamarina. The new combinations Lobothallia semisterilis (H. Magn.) Y. Y. Zhang, Rhizoplaca callichroa (Zahlbr.) Y. Y. Zhang and R. pachyphylla (H. Magn.) Y. Y. Zhang are proposed. Detailed descriptions to aid the identification of these species, distributions and phylogenetic trees, based on multiple collections, are presented. The generic concept of Squamarina is recircumscribed in this study.
Squamarinaceae, Petroplaca, Rhizoplaca, Lobothallia, Lecanora, type study
The genus Squamarina Poelt was first erected by
Nine species of Squamarina have so far been reported from China (
Type specimens were loaned from the Museum of Natural History Vienna (W) and the Swedish Museum of Natural History (S). The fresh material collected for this study is deposited in Kunming Institute of Botany, Chinese Academy of Sciences (
Total DNA was extracted from dry or fresh specimens using the DNeasy Plant Mini Kit (Qiagen, Germany), according to the manufacturer’s instructions. Amplifications were performed in a 25 μl volume containing 12.5 μl 2 × MasterMix (TaqDNA Polymerase [0.1 units/μl], 0.4 mM MgCl2, 0.4 nM dNTPs) (Aidlab Biotechnologies Co. Ltd.), 0.5 μl of each primer, 10 μl ddH2O and 1 μl of DNA. The PCR settings and the primers of nrITS (ITS1-5.8S-ITS2), nrLSU, RPB1, RPB2 and mtSSU follow
Sequences were assembled and edited using SeqMan 7.1 (DNAstar packages). An nrITS matrix of Lobothallia (Clauzade & Cl. Roux) Hafellner, an nrLSU matrix of Squamarina and a 5-locus (nrITS, nrLSU, RPB1, RPB2 and mtSSU) concatenated matrix of Rhizoplaca Zopf and related genera were generated using Geneious R8. Single-gene analyses were conducted, based on the Maximum Likelihood (ML) method to assess the conflict amongst individual genes and no significant incongruence was detected. Matrices were aligned with MAFFT, using the web service (http://mafft.cbrc.jp/alignment/server/index.html). Ambiguous positions were removed, using the web service of Guidance (http://guidance.tau.ac.il/ver2/). MrModeltest2.3 (
A total of 84 sequences of the nrITS, nrLSU, RPB1, RPB2 and mtSSU were newly generated for the species Squamarina chondroderma, S. semisterilis, S. callichroa, S. pachyphylla, S. gypsacea (Sm.) Poelt, S. kansuensis and S. oleosa in this study (Table
Specimens and DNA sequences for nrITS, nrLSU, RPB1, RPB2 and mtSSU used in this study, with the corresponding voucher information from GenBank indicated. Sequences, newly obtained in this study, are indicated in boldface.
Species | Locality* | Voucher specimens | Accession number* | ||||
---|---|---|---|---|---|---|---|
nrITS | nrLSU | RPB1 | RPB2 | mtSSU | |||
Aspicilia cinerea | Sweden | Nordin 6213 (UPS) | JF703115 | – | – | – | – |
A. epiglypta | Sweden | Nordin 6105 (UPS) | HQ259262 | – | – | – | – |
Cladia aggregata | Australia | HTL 19970f (F) | – | GQ500969 | – | – | – |
C. deformis | Australia | HTL 19994d (F) | – | GQ500967 | – | – | – |
Cladonia digitata | na | Ekman 3424 (BG) | – | AY756319 | – | – | – |
C. stipitata | na | AFTOL-ID 1657 (DUKE) | – | DQ973026 | – | – | – |
C. sulcata | Australia | HTL 19975i (F) | – | GQ500959 | – | – | – |
Herteliana schuyleriana | USA: North Carolina | 1885671 | – | MH887488 | – | – | – |
H. taylorii | na | Hertel 39599 (UPS) | – | AY756351 | – | – | – |
Heterodea muelleri | Australia | Elix 39643 (CANB) | – | GQ500962 | – | – | – |
Lecanora achroa | Thailand | Papong 6458 (F) | JN943714 | na | JN987926 | KT453937 | JQ782663 |
L. caesiorubella | Australia | Lumbsch 19974k (F) | JN943728 | JN939508 | JN987920 | na | na |
L. chondroderma 1 | China: Yunnan | 16-54907 ( |
MK778053 | MK778013 | MK766421 | MK766441 | na |
L. chondroderma 2 | China: Xizang | 16-52925 ( |
MK778052 | MK778012 | MK766420 | MK766440 | MN192155 |
L. chondroderma 3 | China: Xizang | 16-53527 ( |
MK778056 | MK778016 | MK766423 | MK766443 | MN192156 |
L. chondroderma 4 | China: Yunnan | 17-55591 ( |
MK778057 | MK778017 | MK766424 | MK766444 | na |
L. conizaeoides | na | K. Molnar U0505/M (DUKE) | na | na | KJ766862 | KJ766956 | KJ766418 |
L. contractula | na | AFTOL-ID 877 (DUKE) | HQ650604 | DQ986746 | DQ986817 | DQ992428 | DQ986898 |
L. dispersa | USA: Illinois | Leavitt 12-002 | KT453733 | na | KT453888 | KT453921 | na |
L. farinacea | Australia | Lumbsch 20003 (F) | JN943725 | JN939513 | JN987924 | na | JQ782672 |
L. flavopallida | Australia | Lumbsch 19972d (F) | JN943723 | JN939516 | JN987925 | KT453938 | JQ782673 |
L. formosa | China: Xinjiang | ZX 20129045-2 (SDNU) | KT453771 | KT453773 | na | KT453978 | KT453819 |
L. hybocarpa | na | Lumbsch s.n. (F) | EF105412 | EF105421 | EF105430 | na | EF105417 |
L. intricata | na | U166 (GZU) | AF070022 | DQ787345 | na | na | DQ787346 |
L. novomexicana | USA | 55026 (BRY-C) | HM577257 | na | KU935390 | KU935136 | na |
L. polytropa | na | AFTOL-ID 1798 (DUKE) | HQ650643 | DQ986792 | na | DQ992418 | DQ986807 |
L. saligna | USA | Leavitt 5702 (BRY-C) | KU934539 | na | KU935293 | KU935036 | na |
L. tropica | Kenya | Lumbsch 19573a (F) | JN943718 | JN939537 | JN987936 | na | na |
Lecidella carpathica | China: Xinjiang | ZX 20140367-2 (SDNU) | KT453741 | KT453784 | KT453905 | KT453944 | KT453831 |
L. stigmatea | China: Xinjiang | ZX 20140838 (SDNU) | KT453766 | KT453803 | KT453918 | KT453971 | KT453849 |
L. tumidula | China: Xinjiang | ZX XL0009 (SDNU) | – | KT453810 | – | – | – |
Lepraria bergensis | na | Tonsberg 28875 (BG) | – | AY756324 | – | – | – |
L. incana | na | AFTOL-ID 1792 (DUKE) | – | DQ986795 | – | – | – |
Lobothallia alphoplaca | China | 20117616 (SDNU) | JX499233 | – | – | – | – |
L. alphoplaca | China | 20117646 (SDNU) | JX476025 | – | – | – | – |
L. crassimarginata | China | 20122565 (SDNU) | JX476026 | – | – | – | – |
L. crassimarginata | China | 20122583 (SDNU) | KC007439 | – | – | – | – |
L. helanensis | China | 20122517 (SDNU) | JX476030 | – | – | – | – |
L. helanensis | China | 20122791 (SDNU) | JX476031 | – | – | – | – |
L. melanaspis | Sweden | Nordin 6622 (UPS) | HQ259272 | – | – | – | – |
L. melanaspis | Norway | Owe-Larsson 8943a (UPS) | JF825524 | – | – | – | – |
L. praeradiosa | China | 20126314 (SDNU) | JX499232 | – | – | – | – |
L. praeradiosa | China | 20126613 (SDNU) | JX499234 | – | – | – | – |
L. pruinosa | China | 20123278 (SDNU) | JX476028 | – | – | – | – |
L. pruinosa | China | 20123630 (SDNU) | JX476027 | – | – | – | – |
L. radiosa | Sweden | Nordin 5889 (UPS) | JF703124 | – | – | – | – |
L. recedens | Sweden | Nordin 6035 (UPS) | HQ406807 | – | – | – | – |
L. semisterilis | China: Qinghai | 18-59262 ( |
MK778040 | MK778009 | na | na | na |
L. semisterilis | China: Qinghai | 18-59322 ( |
MK778039 | MK778008 | MK766413 | na | na |
L. semisterilis | China: Qinghai | 18-59345 ( |
MK778042 | MK778011 | MK766415 | na | na |
L. semisterilis | China: Gansu | 18-59596 ( |
MK778041 | MK778010 | MK766414 | na | na |
Metus conglomeratus | Australia | HTL 19982b (F) | – | GQ500958 | – | – | – |
Miriquidica complanata | Poland: Karkonosze Mts | Szczepanska 935 (herb. Szczepanska) | KF562187 | KF562179 | KF601233 | na | KR995349 |
M. garovaglii | Slovakia: Karpaty Mts | Szczepanska 538 (herb. Szczepanska) | KF562188 | na | KF601234 | na | na |
Mycoblastus affinis | na | AFTOL-ID 1047 (DUKE) | na | KJ766601 | na | KJ766958 | na |
M. sanguinarius | na | AFTOL-ID 196 (DUKE) | DQ782842 | DQ912333 | na | DQ782867 | DQ912276 |
Paralecia pratorum | Italy | M-0045925 (M) | – | KP224503 | – | – | – |
Pilophorus cereolus | na | na | – | AY340559 | – | – | – |
P. strumaticus | na | na | – | AY340560 | – | – | – |
Protoparmeliopsis achariana | na | U525 | na | DQ787341 | DQ973051 | DQ973088 | DQ972976 |
P. garovaglii | USA | Leavitt 106 (BRY-C) | KU934546 | na | KU935300 | KU935043 | na |
P. muralis | na | K. Molnar U0501/AO (EGR) | na | KJ766634 | KJ766830 | KJ766943 | KJ766466 |
P. peltata | Iran | MS014622 | KT453723 | na | KT453892 | KT453927 | na |
P. zareii | Iran | SK 480 | KP059049 | na | na | na | KP059055 |
Ramboldia gowardiana | na | Bjork 9447 (UBC) | na | KJ766649 | KJ766889 | na | KJ766483 |
R. sanguinolenta | Australia: Queensland | Elix 28835 (F) | EU075548 | EU075523 | KT453920 | na | EU075534 |
Rhizoplaca callichroa 1 | China: Sichuan | 14-43348 ( |
MK778045 | na | na | na | na |
R. callichroa 2 | China: Sichuan | 14-43357 ( |
MK778046 | na | na | na | na |
R. callichroa 3 | China: Sichuan | 14-43359 ( |
MK778043 | na | na | na | na |
R. callichroa 4 | China: Yunnan | 14-43308 ( |
MK778044 | na | na | na | na |
R. chrysoleuca 1 | USA | 55000 (BRY-C) | HM577233 | KT453812 | KU935335 | KU935084 | KT453856 |
R. chrysoleuca 2 | Iran | MS014636 | KT453731 | na | KT453898 | KT453934 | na |
R. huashanensis | China | Wei 18357 (HAMS) | AY530885 | AY648104 | na | na | na |
R. marginalis 1 | USA: California | Leavitt 739 (BRY-C) | KT453732 | na | KT453901 | KT453936 | na |
R. marginalis 2 | USA | 0020826b (BRY-L) | KU934655 | na | KU935370 | KU935123 | na |
R. melanophthalma | Iran | MS014628 (H) | JX948271 | na | JX948317 | JX948355 | na |
R. pachyphylla 1 | China: Gansu | 18-59466 ( |
MK778048 | na | MK766417 | MK766436 | MN192152 |
R. pachyphylla 2 | China: Gansu | 18-59446 ( |
MK778047 | na | MK766416 | MK766435 | MN192151 |
R. pachyphylla 3 | China: Gansu | 18-59482 ( |
MK778049 | na | MK766416 | MK766437 | MN192153 |
R. pachyphylla 4 | China: Gansu | 18-59561 ( |
MK778050 | na | MK766419 | MK766438 | MN192154 |
R. polymorpha | USA | 55095 (BRY-C) | HM577326 | na | KU935411 | KU935159 | na |
R. porterii | USA | 55149 (BRY-C) | HM577380 | na | JX948341 | JX948380 | na |
R. shushanii | USA | 55065 (BRY-C) | HM577286 | na | JX948334 | JX948372 | na |
R. subdiscrepans | Russia | Vondrak 9408 (PRA) | KU934898 | na | KU935435 | KU935187 | na |
Squamarina cartilaginea | na | AFTOL-ID 1281 | – | DQ986763 | – | – | – |
S. gypsacea | Greece | O-L-196249 (O) | na | MK778021 | na | na | na |
S. gypsacea | Greece | O-L-196255 (O) | na | MK778020 | na | na | na |
S. gypsacea | Greece | O-L-59266 (O) | na | MK778019 | na | na | na |
S. gypsacea | Spain | O-L-16444 (O) | na | MK778022 | na | na | na |
S. kansuensis | China: Xizang | 16-54052 ( |
MK778059 | MK778023 | MK766425 | MK766446 | na |
S. kansuensis | China: Ningxia | 14-09-1429 (NXAC) | MK778060 | MK778024 | MK766426 | MK766447 | na |
S. kansuensis | China: Xinjiang | 20139103 (XJU) | MK778061 | MK778025 | MK766427 | MK766448 | na |
S. kansuensis | China: Qinghai | 18-59260 ( |
MK778062 | MK778026 | MK766428 | MK766449 | na |
S. kansuensis | China: Gansu | 18-59601 ( |
na | MK778031 | na | na | na |
S. lentigera | na | Haugan & Timdal 4801 (O) | – | AY756363 | – | – | – |
S. oleosa | China: Yunnan | 19-66398 ( |
MN904892 | MN904896 | na | MN923191 | MN915135 |
S. oleosa | China: Yunnan | 19-66399 ( |
MN904893 | MN904897 | MN923189 | MN923192 | MN911318 |
S. oleosa | China: Yunnan | 19-66401 ( |
MN904894 | MN904898 | MN923190 | MN923193 | MN915136 |
Stereocaulon alpinum | Austria | AT1194 (HBG) | – | JN941201 | – | – | – |
S. sasakii | Japan | AT1187 (TUR) | – | JN941206 | – | – | – |
S. tomentosum | Finland | AT1084 (TUR) | – | JN941203 | – | – | – |
Lecanora chondroderma (= Squamarina chondroderma) is sister to the genus Rhizoplaca, but differs in growing on moss and meadow and the presence of numerous rhizinose strands that are never present in its related genera. It is also distinct from the genus Squamarina by the Lecanora-type ascus and the strongly gelatinised lower cortex. This species could belong to a genus separate from Lecanora s. str. and closely related to the genera Rhizoplaca and Protoparmeliopsis, but as only one species from this group was included here, further exploration is needed in the future and we prefer to retain this species in Lecanora here. The remaining two species, Squamarina kansuensis and S. oleosa, proved to belong in Squamarina. Squamarina kansuensis is sister to S. lentigera, but differs in the larger thallus and the presence of psoromic and 2’-O-demethylpsoromic acids. Squamarina oleosa is a basal clade of the genus, which is close to the species S. cartilaginea (With.) P. James and S. gypsacea.
We revised the previously reported ascus structure for the two sections of Squamarina (
Lecanora semisterilis H. Magn., Lichens from Central Asia 1: 123–124 (1940) (Basionym). ≡ Squamarina semisterilis (H. Magn.) J.C. Wei, Enumeration of Lichens in China: 232 (1991). Type: China, Gansu Province, 2450–2600 m elev., on soil, 1931, Birger Bohlin 38L (S–Holotype!).
Thallus to 5 cm across, areolate centrally, with irregularly elongate lobes at the margin, closely to loosely attached to soil; areoles angular, plane to slightly convex, continuous to crowed, ca. 1 mm across; marginal lobes ca. 1 mm wide and 2–3 mm long; upper surface white to grey, pruinose, the pruina on the marginal lobes becoming granular; lower surface white, attached to soil directly with medullary hyphae. Upper cortex colourless with pale brown upper part, 22–55 μm high; epinecral layer colourless, 10–20 μm high; algal layer ca. 95 μm high, not continuous, the interval between different groups of algae 16–32 μm wide; medulla filled with grey granules, lower cortex lacking.
Apothecia rounded, sessile, constricted at the base, up to 2 mm in diam.; disc plane to slightly convex, blackish-brown, non-pruinose; thalline margin entire, concolorous with thallus; hymenium colourless, ca. 60 μm high; subhymenium and hypothecium colourless, I + blue; epihymenium consisting of brown granules, ca. 15 μm high; paraphyses simple, slightly thickened at the apex, ca. 3 μm in diam.; asci Aspicilia-type, 8-spored; ascospores colourless, ellipsoid, 9–13 × 5–9 μm.
Pycnidia prominent, sometimes protruding from the thallus-like apothecia, with blackish-brown ostioles, numerous, 0.1–0.4 mm across; conidia bacilliform, 5.5–6.5 × ca. 1 μm.
Upper cortex K + red, C-, P-, medulla K + red, C-, P + yellow; norstictic acid.
Growing on soil in very dry habitats at elevations of 1760–3151 m. This species was previously only known from Gansu Prov. and is reported here as new to Qinghai Prov., China.
The holotype consists of numerous fragments on soil, without apothecia but numerous pycnidia. This species was originally described as a Lecanora by
The genus Lobothallia is a small genus mainly growing on rocks, containing twelve species (
Lecanora callichroa Zahlbr., in Handel-Mazzetti, Symb. Sinic. 3: 172–173 (1930) (Basionym) ≡ Squamarina callichroa (Zahlbr.) Poelt, Mitt. Bot. Staatssamml., München 1–20: 527 (1958). Type: China, Yunnan Province, 2100 m elev., on rock, 1914, Heinrich Frh. von Handel-Mazzetti 35 (W–Isotype!)
Thallus saxicolous, to 4 cm across, squamulose to placodioid; squamules pruinose on the edges, more or less umbilicate when young; central squamules scattered to continuous, closely attached to the substrate, 1–2 mm across; marginal squamules larger than those in the centre, 2–4 mm across, with 1–2 mm free margin; upper surface yellowish-brown, smooth, plane to slightly convex; lower surface pale to pale brown, without rhizinose strands. Upper cortex filled with yellowish-brown granules dissolving in KOH, ca. 32 μm high; epinecral layer also filled with yellowish-brown granules, ca. 15 μm; algal layer continuous, 64–80 μm high; medulla thick, filled with grey to pale brown granules; lower cortex of free margin poorly developed, non-gelatinised, ca. 30 μm.
Apothecia lecanorine, laminal, dispersed, sessile, becoming slightly constricted at the base, round to irregular, 0.5–1.5 mm; disc orange, covered with pale pruina, plane to slightly convex; thalline margin entire and thick when young, becoming thin and occasionally flexuose with age; hymenium with scattered orange granules, I+ blue, ca. 80 μm high; thalline margin with evenly thick cortex, ca. 26 μm thick; epihymenium yellowish-brown, ca. 10 μm high; subhymenium and hypothecium colourless; ascus Lecanora-type, 8-spored; paraphyses slightly branched, without anastomoses; ascospores subfusiform to ellipsoid, 9.5–13.5 × 6–9 μm. Pycnidia immersed in the thallus, with pale brown ostioles; conidia filiform, straight to slightly curved, 19–26 × ca. 0.7 μm.
Rhizoplaca callichroa (A–D
Upper cortex K-, C-, P-, medulla K + yellow, C-, P-; usnic and placodiolic acids.
Growing on rock in arid environments at elevations of 984–2100 m. Previously only known from Yunnan Prov., here reported as new to Sichuan Prov., China.
The isotype grows on quartzitic rock ca. 2 cm across, containing several intact apothecia. The spore size of “Squamarina callichroa”, given in the protologue, is 15–20 × 8–9 μm (
China: Sichuan Province: Huili Co., beside Jiaopingdu bridge, near to the Jinsha river, 1550 m elev., 26°18'N, 102°22'E, on rock, 2014, Li-Song Wang et al. 14-43348, 14-43357, 14-43359; Yunnan Province: Luquan Co., beside Jiaopingdu bridge, 984 m elev., 26°18'N, 102°22'E, on rock, 2014, Li-Song Wang et al. 14-43308.
Lecanora pachyphylla H. Magn., Lichens from Central Asia 1: 120–121 (1940) (Basionym) ≡ Squamarina pachyphylla (H. Magn.) J.C. Wei, Enumeration of Lichens in China: 232 (1991).Type: China, Gansu Province, 3800–3850 m elev., on rock, 1932, Birger Bohlin (S–Holotype!).
Thallus saxicolous, areolate without lobate margin, to 4 cm across, to 5 mm thick; areoles continuous, plane to slightly convex, 1–2 mm across; upper surface yellow, densely shallow rimose; lower side with thick, grey to white hypothallus. Upper cortex uneven, filled with yellowish-brown granules dissolving in KOH, 32–48 μm thick, algal layer continuous, variable in height, 80–128 μm; medulla very thick, filled with grey to pale brown granules; lower cortex lacking.
Apothecia common, usually densely grouped, irregular in shape, up to 5 mm in diam.; disc black, pruinose at the centre, plane when young, strongly concave with age; thalline margin thin and crenate, strongly bending towards inside with age; hymenium colourless, I+ blue, ca. 50 μm high; epihymenium containing yellowish-brown granules, ca. 9.5 μm high; subhymenium and hypothecium colourless; paraphyses evenly septate, simple, 2–3 μm in diam., apex more or less swollen and bluish-green, ca. 4.5 μm in diam.; ascus Lecanora-type, 8-spored; ascospores regular in shape, ellipsoid, colourless, 5.8–8 × 3–4.5 μm.
Upper cortex K-, C-, P-, medulla K-, C-, P-; usnic acid and traces of unknown substances.
Growing on rock at elevations of 3291–3909 m. Only known from Gansu Prov., China.
The holotype grows on rock with Lecidea tessellata Flörke, Lecanora asiatica H. Magn. and Xanthoria elegans (Link) Th. Fr. and contains numerous apothecia.
This species was originally described as a Lecanora by
China: Gansu Province: Shubei Co., Mengke Glacier, 3942 m elev., 39°12'N, 95°23'E, on rock, 2018, Li-Song Wang et al. 18-59446, 18-59466, 3785 m elev., on rock, 2018, Li-Song Wang et al. 18-59482; Yumen Ci., Yuerhong Vi., 3291 m elev., 39°50'N, 96°45'E, on rock, 2018, Li-Song Wang et al. 18-59560, 18-59561.
≡ Squamarina chondroderma (Zahlbr.) J.C. Wei, Enumeration of Lichens in China: 231 (1991).Type: China, Sichuan Province, 3600–3900 m elev., 1914, Heinrich Frh. von Handel-Mazzetti 497 (W–holotype!)
Thallus to 6 cm across, squamulose or lobate, growing on moss over rock or on the meadow; squamules 0.5–2 mm across, convex, continuous to slightly overlapped; marginal lobes branched, convex, 0.5–2 mm wide, 2–4 mm long; the apex of squamules and lobes rounded, bent downwards; upper surface smooth, pale green to straw, covered by white pruina; lower surface pale to dark brown in the centre and white to pale brown at the margin; rhizinose strands blackish-brown. Upper cortex very thin, ca. 16 μm, filled with yellowish-brown granules dissolving in KOH; algal layer continuous, 48–60 μm thick, medulla filled with grey to pale brown granules, 129–161 μm high, medullary hyphae very loose, more or less hollow in centre; lower cortex well separated from medulla, evenly thick with strongly gelatinised and anticlinally arranged hyphae, ca. 80 μm thick, colourless, hyphae at lower part brown. Apothecia lecanorine, sessile, with constricted base, rounded, scattered or in small groups, up to 3 mm in diam.; disc pruinose, reddish to dark brown, slightly concave when young, slightly convex with age; thalline margin concolorous with thallus, entire to flexuose, forming a well-delimited cortex consisting of strongly gelatinised and anticlinally arranged hyphae; hymenium colourless, 58–80 μm; epihymenium filled with yellowish-brown granules, 10–15 μm; paraphyses simple, evenly septate; ascus Lecanora-type, 8-spored; ascospores colourless, ellipsoid to slightly ovoid, 7–13 × 6.5–9 μm.
Lecanora chondroderma (A, B
Upper cortex K-, C-, P + yellow, medulla K+ yellow, C-, P-; usnic acid and zeorin present in each sample, placodiolic and isousnic acids also present in most samples.
Growing on moss over rock or in meadow at 3600–4968 m elevation in the alpine zone. Worldwide distribution: China, India and Nepal. China: Sichuan Prov., reported here as new to Yunnan and Xizang provinces.
The holotype of Lecanora chondroderma consists of several fragments, containing numerous apothecia.
Lecanora chondroderma was originally described by
Lecanora geophila (Th. Fr.) Poelt is similar to L. chondroderma in morphology, chemistry and habitat, whereas the former forms a yellowish crustose, squamulose to placodioid thallus with loboid projections or phyllidia or terete lobes and epruinose, pale, flat to convex apothecia, including usnic acid, zeorin and methylplacodiolic acid (
China: Sichuan Province: 4650 m elev., 1915, Heinr. Frh. & Handel-Mazzetti 1353 (W). Yunnan Province: Shangri-La Co., Mt. Hong Shan, 4470 m elev., 28°07'N, 99°54'E, on soil, 2018, Li-Song Wang et al. 18-60317; Luquan Co., Mt. Jiaozi Snow, 4000 m elev., 26°05'N, 102°51'E, on moss over rock, 2016, Li-Song Wang et al. 16-54907; Lijiang Co., Mt. Laojunshan, 4036 m elev., 26°37'N, 99°44'E, on rock, 2017, Li-Song Wang et al. 17-55591. Xizang Province: Linzhou Co., Mt. Qiala, 4830 m elev., 30°06'N, 91°16'E, on the meadow, 2016, Li-Song Wang et al. 16-53527; Zuogong Co., on the way from Rumei to Zuogong, 4968 m elev., 29°43'N, 98°01'E, on the meadow, 2016, Li-Song Wang et al. 16-52925, 16-53079, on the meadow, 2016, Li-Song Wang et al. 16-52931.
Lecanora kansuensis H. Magn., Lichens from Central Asia 1: 116–117 (1940). Type: China, Gansu Province, 1500–1700 m elev., on soil, 1930, Birger Bohlin 202 (S–Holotype!) (Basionym)
Thallus terricolous, loosely to tightly adnate on soil, irregular to radiate in outline and with elongate marginal lobes, up to 10 cm in diam.; lobes 2–4(5) mm long, 1–2(3) mm wide, 0.2–0.4 mm thick, with white, thickened and slightly upturned edges, more or less overlapping; upper surface greenish to straw, pruinose and strongly cracked at least in the centre of the thallus; lower surface well delimited, milk-white to pale, without rhizines, margins usually containing sparse white tomentum. Upper cortex filled with yellowish-brown granules, turning colourless in KOH, 26–32 μm thick; epinecral layer grey to brown, 5–15 μm thick; algal layer continuous, well delimited, ca. 50 μm high; medulla grey, filled with calcium oxalate crystals; lower cortex lacking.
Apothecia frequent, rounded, single or in small groups, usually less than 2 mm in diam. Disc pale brown to reddish-brown, slightly concave to flat when young, usually becoming strongly convex with age. Thalline margin distinctive when young and disappearing with age. Hymenium colourless, I + blue, ca. 65 μm high; epihymenium yellowish-brown turning colourless in KOH, ca. 12.5 μm high; thalline margin with evenly thick cortex filled with grey granules; paraphyses septate, ca. 2.5 μm in diam.; hypothecium colourless, 75–87 μm high; algal layer below hypothecium continuous, 62–87 μm high; ascus Porpidia-type, 8-spored; ascospores colourless, ellipsoid to slightly fusiform, variable in size and shape even within one ascus, 7.5–15 × 5–7.5 μm.
Upper cortex K-, C-, P-, medulla K-, C-, P+ yellow; isousnic, usnic, psoromic and 2’-O-demethylpsoromic acids.
Growing on soil at 1310–4730 m of elevation. Previously only known from Gansu Prov. and reported here as new to Neimenggu, Ningxia, Sichuan, Xizang, Xinjiang and Yunnan provinces, China.
The holotype consists of several small fragments on soil, bearing a single small apothecium. This species was originally described as a Lecanora by
The genus Squamarina (= S. sect. Squamarina) includes eleven species (
China: Gansu Province: Jiayuguan, 1500 m–1700 m elev., 1930, Briger Bohlin, S-L60805 (S); Yumen Ci., Moshan National Geological Park, 1760 m elev., 39°57'N, 97°14'E, on soil, 2018, Li-Song Wang et al. 18-59601; Sunan Co., Binggou Danxia landform Park, 1970 m elev., 38°56'N, 99°50'E, on soil, 2018, Li-Song Wang et al. 18-59658; Ningxia Province: Mt. Helanshan, 38°40'N, 1310 m elev., 105°46'E, on soil, 2014,Dong-Ling Niuet al. 14-09-1429 (NXAC); Qinghai Province: Wulan Co., Gobi desert along the way from Chaka to Wulan, 3151 m elev., 36°52'N, 98°55'E, on soil, 2018, Li-Song Wang et al. 18-59260, along the way from Wulan to Delingha, 3039 m elev., 36°59'N, 98°12'E, on soil, 2018, Li-Song Wang et al. 18-59274, 18-59306; Delingha Ci., Chayegou Station, 2974 m elev., 37°23'N, 96°37'E, on soil, 2018, Li-Song Wang et al. 18-59344, 18-59343. Sichuan Province: Derong Co., 1960 m elev., 28°12'N, 99°20'E, on soil, 2009, Li-Song Wang & Wang Jue 09-31112, 09-31118; Xizang Province: Linzhou Co., 3780 m elev., 29°54'N, 91°14'E, on soil, 2016, Li-Song Wang et al. 16-54052; Xinjiang Province: A-ke-tao Co., Oytagh observation zone, 2850 m elev., 38°54'N, 75°14'E, on soil, 2013, Hurnisa Shahidin et al. 20139103; Yunnan Province: Deqin Co., 2110 m elev., 28°13'N, 99°19'E, on soil, 2012, Li-Song Wang et al. 12-34756. Neimenggu Province: Beli-miao, 41°30'N, 110°10'E, on soil, 1929, Briger Bohlin, S-F304837 (S).
Lecanora oleosa Zahlbr., in Handel-Mazzetti, Symb. Sinic. 3: 175 (1930) (Basionym) Type: China, Yunnan Province, Lijiang Co., Mt. Yulongxueshan, on rock, 1914, Heinrich Frh. von Handel-Mazetti 3576 (W–holotype!)
Thallus placodioid to subfoliose, rather closely attached to calcareous rocks, olive-green turning to yellowish-brown in the herbarium, up to 8 cm across and 5 mm high in the centre; lobes 2–4 mm long, 1.5–2.5 mm wide, ca. 1 mm thick, apices usually detached from the substrate with a white thickened edge; upper surface pruinose at least on the margins, matt to somewhat shiny, centrally cracked and faveolate-wrinkled, strongly convex, giving the thallus centre a bullate appearance, the base of the bullae carbonised, black; lower surface covered with pale brown to blackish-brown pulvinate hyphae, with sparse to numerous rhizinose strands; rhizinose strands brown to black, irregularly branched, up to 5 mm long. Upper cortex filled with yellowish-brown granules, turning colourless in KOH, 62–75 μm high, without epinecral; algal layer continuous, 65–70 μm thick; medulla filled with grey crystals of calcium oxalate and brick-red hyphae in lower part; lower cortex lacking.
Apothecia common but not abundant, laminal, scattered to slightly grouped, up to 4 mm in diam.; disc concave, plane to convex, light yellow, covered by yellowish pruina; thalline margin pruinose or not, darker than thallus, shiny, entire and distinctive when young, excluded with age. Hymenium 75–85 μm high, hyaline, I+ blue; epihymenium filled with yellowish-brown granules, not disperse into hymenium, turning colourless in KOH, 5–12.5 μm high; thalline margin without algae in the upper part, cortex filled with yellowish-brown granules, 112–125 μm thick; paraphyses septate, tips not swollen; hypothecium colourless, 100–162 μm thick, with pale brown granules forming a narrow line; algal layer below hypothecium continuous, 50–75 μm thick; ascus Porpidia-type, 8-spored. Ascospores ellipsoid to subfusiform, 15–20 × 5–7 μm. Pycnidia rare and small, ostioles yellow to yellowish-brown, conidia colourless, filiform, curved, 15–22.5 × ca. 0.7 μm.
Upper cortex K-, C-, P-, medulla K-, C-, P+ yellow; usnic, psoromic and 2’-O-demethylpsoromic acids.
Growing on rock at elevations of 2623–3440 m. Only known from Yunnan Prov., China.
The holotype grows on calcareous rock and bears only one apothecium.
This species was originally described as a Lecanora by
China: Yunnan Province: Lijiang Co., 3440 m elev., on rock, 2009, Li-Song Wang & Wang Jue 09-30034, Yulong Snow Mt., 26°56'N, 100°12'E, 2623 m elev., on calcareous rock, 2019, Li-Song Wang & Yan-yun Zhang 19-66398, 19-66399, 19-66401, 19-66402, 19-66404.
Greece: Corfu, hill above Troumpetas, 420 m elev., 39°74'N, 19°86'E, on exposed limestone outcrops, 2014, Rui, S. & Timdal, E., O-L-196249, Sokrati – Zigos road, 370 m elev., 39°72'N, 19°80'E, on rather shady limestone boulders in olive groove, 2014, Rui, S. & Timdal, E., O-L-196255; Kavalla, Thassos, along dirt road from Maries to Theologos, near Vatos, 590 m elev., 40°70'N, 24°66'E, on E-facing limestone wall in/above steep pine forest, 2000, Rui, S. & Timdal, E., O-L-59266. Spain: Alicante, between Callosa de Ensarria and Confrides, 260 m elev., 38°68'N, -0°21'E, 1985, Timdal, E., O-L-16444.
We would like to express our deep thanks to Dr. Arne Anderberg from the Swedish Museum of Natural History (S) and the curator of the Vienna Museum of Natural History (W) for the loan of specimens. This study was supported by grants from the National Natural Science Foundation of China (Nos. 31660007, 31970022, 31670028), Flora Lichenum Sinicorum (31750001), Youth Innovation Promotion Association CAS, Biological Resources Programme, Chinese Academy of Sciences (KFJ-BRP-017-23) and China Scholarship Council. This work benefitted from the sharing of expertise within the DFG priority programme SPP 1991 “Taxon-Omics” and support from DFG grant PR567/19-1 to CP.