Available via license: CC BY 4.0
Content may be subject to copyright.
Citation: Zhong, Q.; Ai, M.; Worthy,
F.R.; Yin, A.; Jiang, Y.; Wang, L.;
Wang, X. Rediscovery of Five
Rinodina Species Originally Described
from Southwest China and One New
Species. Diversity 2023,15, 705.
https://doi.org/10.3390/d15060705
Academic Editor: Gothamie
Weerakoon
Received: 26 April 2023
Revised: 22 May 2023
Accepted: 23 May 2023
Published: 25 May 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
diversity
Article
Rediscovery of Five Rinodina Species Originally Described
from Southwest China and One New Species
Qiuyi Zhong 1,2 , Min Ai 1,2, Fiona Ruth Worthy 1,2 , Ancheng Yin 1,2 , Yi Jiang 3, Lisong Wang 1,2
and Xinyu Wang 1,2,*
1Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany,
Chinese Academy of Sciences, Kunming 650201, China
2Yunnan Key Laboratory for Fungal Diversity and Green Development, Kunming Institute of Botany,
Chinese Academy of Sciences, Kunming 650201, China
3Yunnan Institute of Microbiology, Chenggong Campus of Yunnan University, Kunming 650500, China
*Correspondence: wangxinyu@mail.kib.ac.cn
Abstract:
Rinodina is a lichenized fungal genus belonging to the Physciaceae, with c. 300 species
worldwide. Nearly a century ago, Zahlbruckner described five species of the genus Rinodina from
Southwest China. The type collections were the only records for these species. In the present study,
new records for four of these species: Rinodina cornutula,R. globulans,R. handelii, and R. setschwana,
and a recently described species, R. pluriloculata, are documented based on specimens collected
from the holotype localities. Furthermore, one new species was discovered: Rinodina hengduanensis,
characterized by areolate to subsquamulose thallus, jigsaw-like areoles, lecanorine apothecium,
and Dirinaria-type ascospores. Rinodina setschwana is transferred to the genus Buellia based on its
morphology, chemistry, and phylogeny and proposed as Buellia setschwana. We provide detailed
morphological descriptions, pictures, and molecular phylogenetic analyses.
Keywords: lichenized Ascomycetes; new species; Physciaceae; taxonomy
1. Introduction
Among lichenized fungi with a Holarctic distribution, Rinodina (Ach.) Gray is one
of the better understood genera. It has been studied in the Iberian Peninsula [
1
], Scan-
dinavia [
2
], South Europe [
3
,
4
], the Old World [
5
], North America [
6
,
7
], and Northeast
Asia [
8
]. However, molecular data have shown that Rinodina is a polyphyletic genus [
9
–
12
].
Species-level concepts within Rinodina are becoming better established, with clear outlines
of species groups emerging in conjunction with a refined understanding of crustose mi-
crolichen transitions [
8
,
12
]. However, there remains a substantial knowledge gap for East
Asia, especially for Southwest China, a region that has played an important role in the evo-
lution of higher plants [
8
,
13
–
16
]. The inclusion of specimens and molecular data from East
Asia is vital for the generation of testable hypotheses regarding lichen biogeography [8].
A total of 51 Rinodina species have been reported from China [
5
,
17
–
38
]. In the early 20th
century, five of these species were described from Southwest China by Zahlbruckner [
19
].
The chemistry of their lichen substances was not recorded. More recently, Aptroot and
Sparrius described a new species, R. pluriloculata Aptroot and Sparrius, from the same
area, including the chemistry of its lichen substances but not molecular data [
29
]. Since the
initial publication of these species, no further studies have been conducted. Inter-specific
comparisons of chemical compounds are lacking, and the phylogenetic position of these
species remains unclear.
During the Second Tibetan Plateau Scientific Expedition and Research Program (STEP),
we conducted a thorough survey of the lichen diversity of western China, with a particular
focus on the Qinghai-Tibetan Plateau. With the addition of specimens that we previously
collected in Yunnan Province, over 150 specimens that were confirmed as Rinodina species
Diversity 2023,15, 705. https://doi.org/10.3390/d15060705 https://www.mdpi.com/journal/diversity
Diversity 2023,15, 705 2 of 22
have been stored in the lichen Herbarium at the Kunming Institute of Botany (KUN).
We surveyed the relevant literature, examined the morphology of type specimens, and
compared these to our newly collected materials from the type localities. This confirmed
that we had obtained fresh collections of the species, which were originally described from
southwestern China nearly a century ago [
19
] and twenty years ago [
29
]. In addition, we
discovered one new species of Rinodina. In the present paper, we characterize these species
using morphological, chemical, and molecular evidence and provide a key to the species
included in this study.
2. Materials and Methods
2.1. Morphological and Chemical Study
All the new materials from this study have been deposited in the Herbarium of the Kun-
ming Institute of Botany (KUN). Observations of specimens were made using a Nikon SMZ
745T dissecting microscope (Nikon Corp., Tokyo, Japan). Vertical sections of apothecia and
thalli were cut with a razor blade, then mounted in GAW (
glycerol:ethanol:water = 1:1:1
)
and examined under a Nikon Eclipse 50i stereomicroscope. Measurements and descriptions
of the sections and ascospores were conducted under the microscope and stereomicro-
scope at either
×
10 or
×
40 magnification to an accuracy of 10
µ
m and 2.5
µ
m, respectively.
Photographs were all taken using a Nikon digital camera head, the DS-Fi2. Secondary
metabolites were identified by spot testing and by thin-layer chromatography (TLC), fol-
lowing the methods of Culberson [
39
] and Orange et al. [
40
], using solvent system C
(toluene:acetic acid = 85:15).
2.2. DNA Extraction, Purification, and Sequencing
Total genomic DNA was extracted from 26 specimens using DNAsecure Plant Kits
(Tiangen Biotech, Beijing, China), following the manufacturer’s protocol. Three loci were
amplified: the mitochondrial SSU gene with primer pairs mtSSU1 and mtSSU3R [
41
],
the internal transcribed spacer region (nrITS) with ITS1F [
42
] and ITS4 [
43
], and the 5
0
portion end of the 28S, including the D1–D2 domain [
44
], with LR0R [
45
] and LR5 [
46
].
PCR amplification was performed in 25
µ
L reactions containing 12.5
µ
L of 2
×
MasterMix
(0.1 units/
µ
L TaqDNA polymerase, 4 mM MgCl
2
, and 0.4 nM dNTPs; Aidlab Biotechnolo-
gies Co. Ltd., Beijing, China), 0.5
µ
L of each primer, 9.5
µ
L of ddH
2
O, 2
µ
L of each primer,
and 1
µ
L of DNA, using the PCR settings and primer profile issued by Zhao et al. [
47
]. Poly-
merase chain reaction (PCR) products were sequenced by TsingKe Biological Technology
(Kunming, China).
2.3. Phylogenetic Analysis
The DNA sequences were aligned using MAFFT v. 7.107 in GENEIOUS v. 8.0.2 with
the following parameters: algorithm = auto; scoring matrix = 200 PAM/k = 2; gap open
penalty = 1.53; offset value = 0.123 [
48
]. A 3-locus (nrITS, mtSSU, and nrLSU) concatenated
matrix was generated using GENEIOUS v. 8.0.2. This matrix contained both Rinodina
species and species belonging to related genera in the Physciaceae and Caliciaceae. In
addition to our newly generated sequences, other related sequences were downloaded
from GenBank and added to the matrix (Table 1). Subsequently, a maximum likelihood
(ML) tree and a Bayesian inference (BI) tree were constructed. Based on the lowest Bayesian
information criterion (BIC), the partitioned analysis was selected using ModelFinder in
IQ-TREE v.1.6.12 for the subsequent ML and BI analyses [
49
,
50
]: In ML analysis, GTR
+ F + I + G4 is for ITS1 and ITS2, TNe + I + G4 for 5.8S rRNA and nrLSU, and TPM3u
+ F + I + G4 for mtSSU; in BI analysis, GTR + F + I + G4 is for ITS1 and ITS2, SYM +
I + G4 for 5.8S rRNA and nrLSU, and HKY + F + I + G4 for mtSSU. The ML analyses
were performed by the IQ-TREE web server [
51
–
53
]. The BI analyses were performed by
MrBayes v. 3.2.7 [
54
], using three Markov chains running for 10 million generations for
the concatenated dataset. The trees were sampled every 100 generations, and the first 25%
of the trees were discarded as burn-ins. Posterior probabilities (PPs) were obtained from
Diversity 2023,15, 705 3 of 22
the 95% majority rule consensus tree of all saved trees. The tree files were visualized with
FigTree 1.4.2 [55].
Table 1.
Specimens and sequences used for phylogenetic analyses. Newly generated sequences are
in bold.
Species Locality Voucher
Specimens
GenBank Numbers
References
nrITS mtSSU nrLSU
Acolium karelicum unknown UPS Hermansson
16472 KX512897 na KX512879 [56]
Acroscyphus sphaerophoroides
1unknown UPS Obermayer
6077 KX512898 KX512984 na [56]
A. sphaerophoroides 2 unknown TNS Shimizu AY450562 na na [57]
Allocalicium adaequatum
UPS Spribille 14143
KX512906 KX512986 KX512859 [56]
Amandinea punctata 1 Japan, Tokyo TNS YO6843 LC669587 na na [58]
A. punctata 2 Japan, Tokyo TNS YO6855 LC669588 na na [58]
Anaptychia bryorum Bhutan, Thimphu Soehting 8378 EF582777 EF582825 na [59]
A. ciliaris Spain MAF-Lich 17758 KC559095 na na [60]
Baculifera remensa S Prieto na KX512962 KX512881 [56]
Buellia disciformis 1 Jämtland, Klaxåsen Wedin 6155 AY143392 AY143401 na [61]
B. disciformis 2 Sweden UPS A. Nordin
4429 AF540498 na na [9]
B. elegans unknown S Hansen KX512901 KX512993 na [56]
B. erubescens unknown S Wetmore 95879 KX512902 KX512969 KX512874 [56]
B. frigida unknown S Westberg KX512903 KX512992 KX512852 [56]
B. polita 1 China, Sichuan KUN-L XY20-341 OP526814 OP526788 OP526772
B. polita 2 China, Yunnan KUN-L 15-47741 OP526792 OP526774 OP526763
B. polita 3 China, Sichuan KUN-L 19-62813 OP526800 OP526780 OP526768
B. polita 4 China KUN-L 14-43546 MN615679 na na
B. polita 5 China KUN-L 15-48012 MN615678 na na
B. polita 6 China KUN-L 13-41328 MN615677 na na
B. setschwana 1 China, Yunnan KUN-L 15-48032 OP526796 na na
B. setschwana 2 China, Yunnan KUN-L 15-47744 OP526794 na na
B. setschwana 3 China, Yunnan KUN-L 15-47743 OP526793 OP526775 OP526764
B. tesserata unknown S Tehler 7323 KX512904 na KX512885 [56]
Calicium abietinum unknown UPS Tibell 25061 KX512905 KX512971 KX512872 [56]
C. corynellum unknown S Prieto KX512908 KX512985 KX512855 [56]
Coscinocladium gaditanum 1 Spain MAF 9855 AY449720 AY464073 na [62]
C. gaditanum 2 Spain MAF 9856 AY449721 AY464074 na [62]
Culbersonia nubila 1 South Africa ALV14224
MH121317
na
MH121319
[63]
C. nubila 2 South Africa ALV14225
MH121318
na
MH121320
[63]
Dimelaena oreina unknown S Lendemer 4193 KX512922 KX512976 KX512867 [56]
Diplotomma alboatrum unknown S Prieto 3034 KX512924 KX512966 KX512877 [56]
Heterodermia diademata Bolivia, Depto. Beni
B K. Bach, M.
Kesseler and
Portugal 389
AF540518 na na [9]
H. vulgaris unknown DUKE AFTOL-320
HQ650704
na DQ883798 [64,65]
Hyperphyscia adglutinata unknown BCN-Lich 17031 AF250795 GU247189 na [66]
H. crocata South Korea 120413
MN150490
na na [67]
Kashiwadia orientalis 1 Eastern Asia Hur 040044 na
KM397366
na [68]
K. orientalis 2 Eastern Asia Hur 040164 na
KM397365
na [68]
Diversity 2023,15, 705 4 of 22
Table 1. Cont.
Species Locality Voucher
Specimens
GenBank Numbers
References
nrITS mtSSU nrLSU
Mobergia calculiformis 1 Mexico Moberg 10412 AF224359 na na [69]
M. calculiformis 2 unknown M231 AF250796 na na [66]
Oxnerella safavidiorum Iran KW-L 70300
KM410153 KM410156
na [70]
Phaeophyscia endococcinodes South Korea 130163
MN150503
na na [67]
P. sciastra Norway,
Sor-Trondelag O-L-196352
MK812372
na na [71]
Phaeorrhiza nimbosa China, Xizang KUN-L 19-65695
MW133637 MW133652 MW133660
[72]
P. sareptana China, Gansu KUN-L 18-59809
MW133625 MW133640 MW133654
[72]
Physcia adscendens Norway, Vestfold O-L-198947
MK812201
na na [71]
P. tenella 1 Gästrikland S Odelvik and
Hellstrom 0827 KX512932 KX512974 KX512869 [56]
P. tenella 2 Finland Lohtander 650 AF224425 EF582800 na [59]
Physciella chloantha 1 Iberian Peninsula BCN-Lich 15525 GU247166 GU247200 na
P. chloantha 2 Iberian Peninsula BCN-Lich 17033 GU247164 GU247198 na
Physconia enteroxantha Norway, Akershus O-L-196328
MK812142
na na [71]
P. grisea Morocco, Imli Staro192 LS483208 na na [71]
Pseudothelomma ocellatum 1 unknown S Tehler 8063 KX512934 KX512957 KX512862 [56]
P. ocellatum 2 unknown UPS Hermansson
18662 KX512935 KX512952 KX512891 [56]
Pyxine himalayensis China, Yunnan KUN-L 12-36055 KY611881 KY751388 na [73]
P. subcinerea China, Taiwan KUN-L 15-49012 KY611867 KY751374 na [73]
Rinodina cana unknown Sipman 63008
MN587029
na na [74]
R. conradii China, Yunnan KUN-L 13-40531 OP526791 OP526773 na
R. cornutula 1 China, Yunnan KUN-L XY20-3571 OP526815 OP526789 na
R. cornutula 2 China, Yunnan KUN-L XY20-3572 OP526816 OP526790 na
R. gennarii Netherlands, Utrecht B H. Sipman 44435 AJ544187 na na [9]
R. globulans 1 China, Yunnan KUN-L 15-47871 OP526795 OP526776 na
R. globulans 2 China, Yunnan KUN-L XY20-171 OP526807 OP526785 OP526770
R. globulans 3 China, Yunnan KUN-L XY20-117 OP526805 na na
R. globulans 4 China, Sichuan KUN-L XY20-279 OP526810 na na
R. globulans 5 China, Sichuan KUN-L XY20-280 OP526811 na na
R. handelii 1 China, Yunnan KUN-L XY20-224 OP526808 na na
R. handelii 2 China, Yunnan KUN-L XY20-238-1 OP526809 na na
R. mniaroea 1 USA, Idaho GZU Spribille
15242 KX015687 KX015706 na [12]
R. mniaroea 2 USA, Montana GZU Spribille
20391 KX015692 KX015711 na [12]
R. mniaroea 3 Norway, Svalbard TROM_L_565871
MK812098
na na [71]
R. mniaroea 4 unknown M249 AF250811 na na [66]
R. mniaroeiza Canada GZU V. Wagner
15.07.06/1 KX015691 KX015710 na [12]
R. moziana 1Australia,
Queensland
GZU H. Mayrhofer
11742 DQ849306 na na [75]
R. moziana 2New Zealand,
Nelson
GZU M. Lambauer
0214 DQ849305 na na [75]
R. oxydata 1Australia,
Queensland
GZU H. Mayrhofer
11790 DQ849311 na na [75]
R. oxydata 2Australia,
Queensland
GZU H. Mayrhofer
11406 DQ849313 na na [75]
Diversity 2023,15, 705 5 of 22
Table 1. Cont.
Species Locality Voucher
Specimens
GenBank Numbers
References
nrITS mtSSU nrLSU
R. oxydata 3New Zealand,
Nelson
GZU M. Lambauer
0206 DQ849310 na na [75]
R. oxydata 4 Austria, Steiermark GZU H. Mayrhofer
13.930 AF540548 na na [9]
R. oxydata 5 Austria, Styria GZU H. Mayrhofer
15761 DQ849312 na na [75]
R. pluriloculata 1 China, Yunnan KUN-L XY20-162 OP526806 OP526784 na
R. pluriloculata 2 China, Yunnan KUN-L 20-66417 OP526802 OP526782 na
R. sophodes 1 Austria, Styria GZU 000272661 GU553304 GU553321 na [10]
R. sophodes 2 Austria, Steiermark GZU P. Bilovitz 968 AF540550 na na [9]
R. orientalis 1 South Korea KBA
BDNA-L-0000284
MW832807
na na [76]
R. orientalis 2 South Korea KBA
BDNA-L-0000653
MW832808
na na [76]
R. salicis 1 South Korea KBA
BDNA-L-0000558
MW832810
na na [76]
R. salicis 2 South Korea KBA
BDNA-L-0000560
MW832811
na na [76]
R. hengduanensis 1 China, Sichuan KUN-L 20-66506 OP526803 OP526783 na
R. hengduanensis 2 China, Yunnan KUN-L XY20-44 OP526804 na na
R. hengduanensis 3 China, Sichuan KUN-L XY20-290 OP526813 OP526787 na
R. hengduanensis 4 China, Sichuan KUN-L XY20-287 OP526812 OP526786 OP526771
R. zeorina 1 South Korea KBA
BDNA-L-0000642
MW832812
na na [76]
R. zeorina 2 South Korea KBA
BDNA-L-0000646
MW832813
na na [76]
Rinodinella controversa 1 unknown M281 AF250814 na na [66]
R. controversa 2 Greece, Kreta GZU Mayrhofer
and Ertl 13.74 AJ421423 na na [77]
Tetramelas pulverulentus unknown UPS Nordin 6368 KX512940 KX512983 KX512860 [56]
Texosporium sancti-jacobi unknown UPS Rosentreter
and De Bolt 6514 KX512941 KX512981 KX512863 [56]
Thelomma mammosum unknown UPS Tibell 23775 KX512942 KX512954 KX512888 [56]
T. santessonii unknown UPS Nordin 4011 KX512944 KX512951 KX512889 [56]
Tholurna dissimilis unknown UPS Wedin 6330 AY143397 AY143407 KX512893 [56]
Tornabea scutellifera 1 Spain AFTOL-ID 1061 JQ301698 na DQ973037 [11,65]
T. scutellifera 2 unknown UPS Tibell 23833 KX512946 KX512970 KX512873 [56]
Xanthoria aureola Sweden, Bohuslän E. Gaya etc. (BCN) JQ301690 JQ301526 JQ301585 [12]
Xanthoria parietina Sweden, Bohuslän E. Gaya etc. (BCN) JQ301691 JQ301530 JQ301589 [12]
3. Results and Discussion
By re-collecting these species from their holotype localities and conducting morpho-
logical and phylogenetic analyses, we confirmed the identities of Rinodina cornutula,R.
globulans,R. handelii,Buellia setschwana (
≡
R. setschwana), and R. pluriloculata. Detailed fig-
ures of morphology and spores, together with information regarding chemical compounds,
have been provided. Furthermore, the phylogenetic positions of these species have been
confirmed. An additional species has been described as new to science: R. hengduanensis,
for which morphological descriptions and molecular data have been provided.
Diversity 2023,15, 705 6 of 22
The sequence alignment comprised 105 terminals, of which 25 were newly gener-
ated sequences from this study (Table 1). Phylogenetic trees that were based on datasets
generated from either ML or BI analyses had almost identical topologies, with minimal
differences in the level of statistical support. The phylogenetic analysis showed that the
families Physciaceae and Caliciaceae both formed monophyletic clades. Rinodina cornu-
tula,R. globulans,R. handelii, and R. pluriloculata belong to the Physciaceae, while Buellia
setschwana (syn. R. setschwana) belongs to the Caliciaceae.
Our results indicate that R. pluriloculata is phylogenetically close to R. conradii. Both
species have 3–8-septate ascospores, but R. pluriloculata can be distinguished by its pale
to olive green thallus, punctiform, discrete, orbicular, convex to flattened soralia, bluish,
granular soredia, and submuriform-type ascospores. R. cornutula is phylogenetically closely
related to R. oxydata. The morphological characters of these two species are largely similar:
both are characterized by Mischoblastia-type spores and a thallus with atranorin in the
cortex. They can be separated by their thallus color and habitat. The molecular data
available for R. oxydata showed that this species is not monophyletic, so further research is
required based on samples from its type locality. R. handelii appears close to R. orientalis and
R. zeorina in the phylogenetic tree, but the three species can be separated by their ascospore
morphology and their chemistry. R. handelii lacks secondary metabolites, and it has Milvina-
to Mischoblastia-type ascospores (up to 27
µ
m), which are longer than the spores of the other
two species (both less than 20
µ
m). R. orientalis has Physcia-type spores, whereas R. zeorina
contains zeorin and has Dirinaria-type ascospores. R. globulans forms a well-supported clade
(100/100/1.00), which is also supported by its morphological, chemical, and geographic
characteristics. It differs from other species by its round, capitate, and scattered thallus
areoles covered with soralia. Interestingly, Buellia setschwana (syn. R. setschwana) forms a
monotypic lineage that is close to B. polita. These two species can be distinguished by the
characters of their apothecia: B. polita has a reddish brown hypothecium and immersed
lecideine apothecia, whereas B. setschwana has a hyaline hypothecium and cryptolecanorine
apothecia. The apothecia characters of B. setschwana are identical to those of other species
within Rinodina, which was probably the basis for its previous placement within the
genus Rinodina.
The new species Rinodina hengduanensis formed a single clade, represented by a SH-
aLRT support of 100, an ultrafast bootstrap support of 100, and a posterior probability
of 1 for the branch (Figure 1). Its species status is further supported by its distinctive
morphological, chemical, and geographic characteristics.
Diversity 2023,15, 705 7 of 22
Diversity 2023, 15, x FOR PEER REVIEW 6 of 20
Figure 1. Phylogenetic tree generated from maximum likelihood (ML) analysis based on combined
nrITS, mtSSU, and nrLSU sequences. SH-aLRT support (%) for ML greater than 80%, ultrafast boot-
strap support (%) greater than 95%, and Bayesian posterior probabilities (PPs) greater than 0.95 are
given above the nodes. Newly generated sequences are indicated in bold.
Figure 1.
Phylogenetic tree generated from maximum likelihood (ML) analysis based on combined
nrITS, mtSSU, and nrLSU sequences. SH-aLRT support (%) for ML greater than 80%, ultrafast
bootstrap support (%) greater than 95%, and Bayesian posterior probabilities (PPs) greater than 0.95
are given above the nodes. Newly generated sequences are indicated in bold.
Diversity 2023,15, 705 8 of 22
4. Taxonomy
Rinodina cornutula Zahlbr., in Handel-Mazzetti, Symb. Sinic. 3: 233 (1930), Figure 2.
Type
: China, Yunnan Prov., Manhao nahe der Grenze von Tonkin, Tonschieferfelsen
am Flußufer in der tr. St., 200 m, 2 March 1915, Handel-Mazzetti no. 5858 (WU, holotype!).
Description
:Thallus crustose, thin, areolate, continuous at the central part, scattered
near the margins, margins not distinct; areoles irregular in shape, 0.2–0.6 mm wide; upper
surface flat, brown to olive brown, mottled, dull, non-pruinose; prothallus absent or de-
limiting the thallus as a black outline; vegetative propagules absent. Apothecia common,
usually abundant and contiguous, lecanorine (becoming lecideine), broadly attached to
slightly innate, 0.2–0.6 mm in diam.; disc black, plane and roundish, non-pruinose; margin
concolorous with thallus, entire, persistent, usually becoming carbonized and concolorous
with the disc when mature; exciple hyaline to dark brown, with algal cells; hymenium
hyaline, colorless, not inspersed, 75–125
µ
m high; paraphyses simple, conglutinate, apices
expanded, light brown; epihymenium pale brown to dark brown; hypothecium hyaline,
colorless, 35–75
µ
m deep; asci 8-spored, Lecanora-type; ascospores 1-septate, Type A devel-
opment, Mischoblastia-type, pale brown, lumina triangle- or heart-shaped, walls thickened
at both ends and the septum, 18–25
×
7.5–12.5
µ
m, torus indistinct. Pycnidia were not seen.
Chemistry
: Cortex K+ yellow; KC+ yellow or orange; C
−
, P+ faint yellow; medulla I+
blue. Containing atranorin (confirmed by TLC).
Ecology and distribution
: This species is distributed across subtropical to temperate
regions, growing on siliceous rocks, rhyolite, or basalt, at elevations between 950 and
2500 m. Endemic to China, previously reported in Sichuan and Yunnan [19,36].
Notes
:Rinodina cornutula is characterized by its flat, areolate, ochre to olive thallus,
broadly attached to slightly innate apothecia, Mischoblastia-type ascospores, and containing
atranorin. This species was originally collected from Manhao town, Yunnan province,
China. The specimens that we collected from the same area were compared to type
specimens and confirmed as identical to the holotype. This species is similar to Rinodina
cana (Arnold) Arnold in morphology, but R. cana lacks atranorin (spot tests are all negative),
and its apothecia are usually entirely innate, with Milvina-type ascospores. It might be
confused with Rinodina moziana (Nyl.) Zahlbr., but the latter species can be distinguished by
its verruciform or granular thallus and its constricted ascospores when mature. R. oxydata
(A. Massal.) A. Massal differs from this species by its gray to ochraceous thallus and shorter
ascospores (<18
µ
m long). R. cornutula would key out at couplet 9 in Lee and Hur’s paper,
as it has a plane thallus, Mischoblastia-type, 18–25
×
7.5–12.5
µ
m ascospores, and is rarely
swollen at the septum [76].
Specimens examined
: China, Yunnan Prov.: Lijiang Co., Yufeng Temple, 2500 m, on
rock, August 15, 1982, Wang Lisong 82-1099; Lijiang City, Yulong Naxi Autonomous Co.,
on the way from Lijiang to Ninglang, Jinsha River side, 1846–1883 m, on rock, April 9, 2019,
Wang Lisong et al. 19-62703, 19-62705, 19-62706; Honghe prefecture, Lvchun Co., Niukong
Vil., 972 m, on rock, December 17, 2020, Wang Xinyu et al. XY20-3571, XY20-3572.
Diversity 2023,15, 705 9 of 22
Diversity 2023, 15, x FOR PEER REVIEW 8 of 20
Figure 2. Morphology and anatomy of Rinodina cornutula. (A) Thallus and apothecia. (B)
Mischoblastia-type ascospores. (C) Section of apothecia. Scale bars: 2 mm (A); 10 µm (B); 20 µm (C).
Rinodina globulans Zahlbr., in Handel-Mazzetii, Symb. Sinic. 3: 233 (1930), Figure 3.
Type: China, Sichuan Prov., Eisenschüssiger Sandstein in der tp. St. Um den Paß
Dsiliba im Daliangschan e von Ningyüen, ±3000 m, 26 April 1914, Handel-Mazzei no.
1771 (WU, holotype!).
Description: Thallus crustose, round, capitate to areolate, scaered; areoles 0.1–0.4
mm in diam.; surface pale green (greyish white to pale after storage in herbarium), with
white to greyish white pruina, covered by abundant soralia, globular, pale green;
prothallus prominent, black, persistent; Apothecia lecanorine, usually scaered, rarely
contiguous, sessile, 0.6–0.8 mm in diam.; disc brown to dark brown, concave, round, non-
pruinose; margin concolorous with thallus, thick, distinctly raised, entire, persistent;
Figure 2.
Morphology and anatomy of Rinodina cornutula. (
A
) Thallus and apothecia. (
B
)Mischoblastia-
type ascospores. (C) Section of apothecia. Scale bars: 2 mm (A); 10 µm (B); 20 µm (C).
Rinodina globulans Zahlbr., in Handel-Mazzetii, Symb. Sinic. 3: 233 (1930), Figure 3.
Type
: China, Sichuan Prov., Eisenschüssiger Sandstein in der tp. St. Um den Paß
Dsiliba im Daliangschan e von Ningyüen,
±
3000 m, 26 April 1914, Handel-Mazzetti no.
1771 (WU, holotype!).
Description
:Thallus crustose, round, capitate to areolate, scattered; areoles
0.1–0.4 mm
in diam.; surface pale green (greyish white to pale after storage in herbarium), with white
to greyish white pruina, covered by abundant soralia, globular, pale green; prothallus
prominent, black, persistent; Apothecia lecanorine, usually scattered, rarely contiguous,
sessile, 0.6–0.8 mm in diam.; disc brown to dark brown, concave, round, non-pruinose;
margin concolorous with thallus, thick, distinctly raised, entire, persistent; exciple hyaline
to pale brown, with algal cells, persistent; hymenium hyaline, colorless, not inspersed,
80–130
µ
m high, I+ blue; paraphyses simple or merely branched at the apices; epihyme-
Diversity 2023,15, 705 10 of 22
nium olive to greyish brown; hypothecium hyaline, colorless, 40–65
µ
m deep; asci 8-spored,
Lecanora-type; ascospores 1-septate, Type A development, Mischoblastia- to Pachysporaria-
type, brown, lumina triangle- or trapezoid-shaped; walls thicken at both ends and at the
septum, 15–25
×
7.5–12.5
µ
m; torus absent with oil droplets when over-mature. Pycnidia
were not seen.
Chemistry
: Cortex and medulla K+ yellow, KC
−
, C
−
, P
−
. Atranorin, zeorin, and
stictic acid are detected by TLC.
Ecology and distribution
: This species is saxicolous and often grows on sandstone at
elevations between 2000 and 3600 m. It is endemic to China, previously reported in Sichuan
province [19,36]. This is a new record for Yunnan Province, China.
Notes
:R. globulans is a unique species with numerous, scattered, globular, pale green
soralia and a prominent black prothallus, which can be reliably distinguished from other
Rinodina species in China. Apothecia are rare in this species, and the ascospores are
Mischoblastia-Pachysporaria-type. A recently published species, Rinodina punctosorediata
Aptroot and Sparrius, is highly similar to this species, with identical external and anatomic
morphology [
29
]. However, R. punctosorediata grows both on trees (Castanopsis sp.) and
rocks, whereas the holotype of R. globulans and all our collections grew on rocks. Further-
more, R. punctosorediata has smaller ascospores (<18
µ
m long) than the Milvina-type. R.
globulans would be keyed out at couplet 2 in Lee and Hur’s paper because it has scattered
thallus areoles covered with abundant, globular, pale green soralia [76].
Specimens examined
: China, Sichuan Prov.: Liangshan Yi Autonomous Prefecture,
Yanyuan Co., Weicheng Town, 2816–2831 m, on sand rock, 28 2020, Wang Xinyu et al.
XY20-280, XY20-279. Yunnan Prov.: Diqing Tibetan Autonomous Prefecture, Shangri-La
City, Jiantang Town, Pudacuo National Park, Bita, 3496 m, on rock, 25 July 2020, Wang
Xinyu et al. XY20-117; Lijiang City, Ninglang Yi Autonomous Co., Xichuan Vil., next to
Lining Highway, 2509 m, on rock, 26 July 2020, Wang Xinyu et al. XY20-171; Dali Prefecture,
Nanjian Co., Leqiu Vil., Awuhe Vil., Dayao Reservoir, 2095 m, on rock, 2 July 2015, Ye Xin
and Wang Weicheng 15-47871; Luquan Co., Jiaozi Snow Mt., 3600 m, on rock, September
18, 2003, Wang Lisong 03-22628; Dali Co., Cangshan Mt., Zhonghe Temple, 2580 m, on rock,
August 30, 2005, Wang Lisong et al. 05-25172.
Diversity 2023,15, 705 11 of 22
Diversity 2023, 15, x FOR PEER REVIEW 10 of 20
Figure 3. Morphology and anatomy of Rinodina globulans. (A) Thallus and apothecia. (B)
Pachysporaria-type ascospores. (C) Section of apothecia. Scale bars: 2 mm (A); 10 µm (B); 20 µm (C).
Chemistry: Spot tests were all negative; no metabolites were detected by TLC.
Ecology and distribution: This species is corticolous, growing on bark or branches
of Pinus, Quercus, Rhododendron, etc., in coniferous forest or mixed coniferous broad-
leaved forest, at elevations between 2700 and 3600 m. It has only previously been reported
from Yunnan province, China [19,36]. This is a new record for Sichuan province.
Notes: R. handelii is a corticolous species with a thin, rimose to continuous, rough
thallus; sessile apothecia; a dark brown disc; characterized by ascospores with torus,
always inspersed with oil droplets when mature; a size of 20–27 × 7–12.5 µm. It could be
confused with other corticolous species such as Rinodina bolanderi and R. subminuta, but R.
bolanderi can be distinguished by containing atranorin and zeorin, and Teichophila-type
ascospores, and R. subminuta can be distinguished by containing zeorin and Physcia-type
Figure 3.
Morphology and anatomy of Rinodina globulans. (
A
) Thallus and apothecia. (
B
)Pachyspo-
raria-type ascospores. (C) Section of apothecia. Scale bars: 2 mm (A); 10 µm (B); 20 µm (C).
Rinodina handelii Zahlbr., in Handel-Mazzetti, Symb. Sinic. 3: 232 (1930),
Type
: China, Yunnan Prov., Lebende Rinde von Pflaumenbäumen in Gärten der wtp.
St. in Yünnanfu, 1920 m, February 24, 1914, Handel-Mazzetti no. 275 (WU, holotype;
US, isotype).
Description
:Thallus crustose, thin, rimose to continuous, somewhat verrucose; are-
oles 0.1–0.2 mm wide; upper surface rough to verrucous, grey, brown, or pale green,
dull, non-pruinose; prothallus absent; vegetative propagules absent. Apothecia common,
scattered or contiguous, lecanorine, broadly attached, 0.2–0.6 mm in diam.; disc ochre
brown to dark brown, plane to convex, round, non-pruinose; margin concolorous with
thallus, thin, entire, disappeared when mature; exciple hyaline, with abundant algal cells;
hymenium hyaline, colorless, not inspersed, 50–100
µ
m high, I+ blue; paraphyses simple,
somewhat conglutinate, apices expanded, dark pigmented; epihymenium dark brown; hy-
Diversity 2023,15, 705 12 of 22
pothecium hyaline, colorless or pale brown, 50–100
µ
m deep; asci 8-spored, Lecanora-type;
ascospores 1-septate, Type A development, pale brown to brown, Milvina- to Mischoblastia-
type,
20–27 ×7–12.5 µm
, torus present, always inspersed with oil droplets when mature.
Pycnidia were not seen.
Chemistry: Spot tests were all negative; no metabolites were detected by TLC.
Ecology and distribution
: This species is corticolous, growing on bark or branches of
Pinus,Quercus, Rhododendron, etc., in coniferous forest or mixed coniferous broad-leaved
forest, at elevations between 2700 and 3600 m. It has only previously been reported from
Yunnan province, China [19,36]. This is a new record for Sichuan province.
Notes
:R. handelii is a corticolous species with a thin, rimose to continuous, rough
thallus; sessile apothecia; a dark brown disc; characterized by ascospores with torus,
always inspersed with oil droplets when mature; a size of 20–27
×
7–12.5
µ
m. It could be
confused with other corticolous species such as Rinodina bolanderi and R. subminuta, but
R. bolanderi can be distinguished by containing atranorin and zeorin, and Teichophila-type
ascospores, and R. subminuta can be distinguished by containing zeorin and Physcia-type
ascospores. R. handelii also resembles Rinodina subleprosula Jatta in external morphology,
which is a corticolous species previously described by Jatta from China [
17
]. However,
R. subleprosula has larger ascospores (27.5–30.5
×
12–15
µ
m). Another corticolous species,
Rinodina cinereovirens (Vain.), has similar ascospores to R. handelii, but it can be distinguished
by its medulla, which is UV + blue-white and contains sphaerophorin. R. handelii would
key out at couplet 60 in Lee and Hur’s paper because the margins of the apothecia are not
radially cracked and there are Milvina- to Mischoblastia-type ascospores [76].
Specimens examined
: China, Sichuan Prov.: Liangshan Prefecture, Yanyuan Co.,
Weicheng Town, 3210 m, on Pinus densata, 28 July 2020, Wang Lisong et al. 20-66526.
Yunnan Prov.: Diqing Prefecture, Shangri-La City, Jiantang Pudacuo National Park, Bita,
3540 m, on Quercus bark, Wang Xinyu et al. XY20-134; Gongshan Co., Yeniu Valley, 2700 m,
on bush, 30 May 2000, Wang Lisong 00-19328; Lijiang City, Ninglang Co., Hongqiao Vil., c.
3100 m, on branch, 27 July 2020, Wang Xinyu et al. XY20-224, XY20-238-1. Figure 4.
Diversity 2023,15, 705 13 of 22
Diversity 2023, 15, x FOR PEER REVIEW 11 of 20
ascospores. R. handelii also resembles Rinodina subleprosula Jaa in external morphology,
which is a corticolous species previously described by Jaa from China [17]. However, R.
subleprosula has larger ascospores (27.5–30.5 × 12–15 µm). Another corticolous species,
Rinodina cinereovirens (Vain.), has similar ascospores to R. handelii, but it can be distin-
guished by its medulla, which is UV + blue-white and contains sphaerophorin. R. handelii
would key out at couplet 60 in Lee and Hur’s paper because the margins of the apothecia
are not radially cracked and there are Milvina- to Mischoblastia-type ascospores [76].
Specimens examined: China, Sichuan Prov.: Liangshan Prefecture, Yanyuan Co.,
Weicheng Town, 3210 m, on Pinus densata, July 28, 2020, Wang Lisong et al. 20-66526. Yun-
nan Prov.: Diqing Prefecture, Shangri-La City, Jiantang Pudacuo National Park, Bita, 3540
m, on Quercus bark, Wang Xinyu et al. XY20-134; Gongshan Co., Yeniu Valley, 2700 m, on
bush, May 30, 2000, Wang Lisong 00-19328; Lijiang City, Ninglang Co., Hongqiao Vil., c.
3100 m, on branch, July 27, 2020, Wang Xinyu et al. XY20-224, XY20-238-1.
Figure 4.
Morphology and anatomy of Rinodina handelii. (
A
) Thallus and apothecia. (
B
)Mischoblastia-
type ascospores. (C) Section of apothecia. Scale bars: 1 mm (A); 10 µm (B); 20 µm (C).
Rinodina pluriloculata
Aptroot and Sparrius, Fungal Diversity 14: 42 (2003), Figure 5.
Type:
CHINA, Yunnan Prov., 5 km W of Kunming, just outside of the city, 1750 m alt.,
48RTN604714, on bark of tree base of living Eucalyptus globulus, 16 October 2002, Aptroot
no. 55505 (B, holotype; KUN isotype).
For a detailed description, see [29].
Notes
:R. pluriloculata is characterized by its continuous, verrucous, pale green to olive
thallus; punctiform, discrete, orbicular, convex to flattened soralia; bluish, granular soredia;
lecanorine, sessile apothecia; and 3–8-septate, Submuriform-type, pale brown ascospores
27–35
×
15–17.5
µ
m. Spot tests: cortex and medulla K
−
or + yellow, KC
−
, C
−
, P
−
;
containing skyrin and
±
atranorin by TLC. This species was previously reported in Yunnan,
growing on rock or bark at elevations around 2500 m. According to Aptroot and Sparrius,
the holotype of this species grows on the tree trunk base of Eucalyptus globulus, with an
Diversity 2023,15, 705 14 of 22
additional cited specimen growing on soil and shale [
29
]. The specimens that we collected
all grew on rocks. Similar to this species, Rinodina conradii Körb also has 3–8-septate
ascospores, but it can be differentiated by its brown thallus, lack of vegetative propagules,
and Type B development, Conradii-type ascospores. R. pluriloculata would key out at couplet
20 in Lee and Hur’s paper because it has Type A development ascospores (apical wall
thickening after septum formation) and contains skyrin [76].
Specimens examined
: China, Yunnan Prov.: Lijiang City, Ninglang Yi Autonomous
Co., Xichuan Vil., beside Lining Highway, 2504–2520 m, on rock, 26 July 2020, Wang Xinyu
et al. XY20-162, 20-66417.
Diversity 2023, 15, x FOR PEER REVIEW 13 of 20
Figure 5. Morphology and anatomy of Rinodina pluriloculata. (A) Thallus and apothecia. (B) Submu-
riform-type ascospores. (C) Section of apothecia. Scale bars: 2 mm (A); 10 µm (B); 20 µm (C).
Notes: Buellia setschwana is characterized by its thin, rimose thallus; black and prom-
inent prothallus; innate cryptolecanorine to lecideine; black apothecia; and containing
atranorin and norstictic acid as main compounds. It resembles Buellia polita in its external
morphology and chemical compounds but could be distinguished by the hyaline hypo-
thecium and cryptolecanorine apothecia. Both B. setschwana and the globally distributed
Buellia halonia contain the secondary metabolite arthothelin. This could lead to confusion
between the two species. However, R. halonia has exciples with typical aeruginose
pigments (HNO3+ purple), a reddish brown hypothecium, and adnate apothecia (innate
in B. setschwana). Furthermore, R. halonia is always maritime, whereas B. setschwana is con-
tinental.
Figure 5.
Morphology and anatomy of Rinodina pluriloculata. (
A
) Thallus and apothecia. (
B
)Submuri-
form-type ascospores. (C) Section of apothecia. Scale bars: 2 mm (A); 10 µm (B); 20 µm (C).
Buellia setschwana (Zahlbr.) Q. Y. Zhong and Xin Y. Wang, comb. nov., Figure 6.
=Rinodina setschwana Zahlbr., in Handel-Mazzetti, Symb. Sinic. 3: 231 (1930)
Diversity 2023,15, 705 15 of 22
Type:
China, Sichuan Prov., Sandsteinfelsen in der str. St. bei Dötschang im Pjientschang,
1450 m, 4 April 1914, Handel-Mazzetti no. 1171 (WU, holotype; US, isotype).
MycoBank No. 848787
Description
:Thallus crustose, thin, plane, closely attached to the substrate, rimose;
upper surface pale gray to pale yellowish brown, dull, non-pruinose; prothallus black,
prominent; vegetative propagules absent. Apothecia common, usually scattered or some-
times contiguous, cryptolecanorine to lecideine, innate, 0.3–0.5 (0.7) mm in diam.; disc
black, plane, round, or somewhat irregular; margin concolorous with thallus, entire, distinct
when young, sometimes becoming carbonized when mature; exciple prominent, brown;
hymenium colorless, not inspersed, 80–90
µ
m high; paraphyses simple, contiguous; apices
expanded and septate, brown pigmented; epihymenium brown; hypothecium hyaline,
colorless or pale brown, 50–60
µ
m deep; asci 8-spored, Bacidia-type; ascospores 1-septate,
Buellia-type, brown, 12.5–18
×
5–7.5
µ
m; mature spores with obtuse ends, slightly con-
stricted at septum; torus absent. Pycnidia common, dark brown to black, immersed; ostioles
brown pigmented; conidia hyaline, bacilliform, 4–6 ×1µm.
Chemistry
: Cortex K+ yellow; KC+ yellow or orange; C
−
, P
−
; medulla I+ blue.
Atranorin, arthothelin, norstictic acid, and sometimes stictic acid are detected by TLC.
Ecology and distribution
: This species grows on rocks at elevations between 1100 m
and 2000 m. It has previously been reported from Sichuan Province, China [
19
]. This is a
new record for Yunnan Province.
Notes
:Buellia setschwana is characterized by its thin, rimose thallus; black and promi-
nent prothallus; innate cryptolecanorine to lecideine; black apothecia; and containing
atranorin and norstictic acid as main compounds. It resembles Buellia polita in its external
morphology and chemical compounds but could be distinguished by the hyaline hypothe-
cium and cryptolecanorine apothecia. Both B. setschwana and the globally distributed Buellia
halonia contain the secondary metabolite arthothelin. This could lead to confusion between
the two species. However, R. halonia has exciples with typical aeruginose pigments (HNO
3
+
purple), a reddish brown hypothecium, and adnate apothecia (innate in B. setschwana).
Furthermore, R. halonia is always maritime, whereas B. setschwana is continental.
Rinodina setschwana was originally collected in Sichuan Province, Dechang County.
It was placed in the genus Rinodina based on its cryptolecanorine apothecia and hyaline
hypothecium [
19
]. After the collection of fresh specimens from the same locality and com-
parison with the type specimen, we confirmed that our collections were morphologically
identical to the holotype. However, phylogenetic analyses demonstrated that this species
should be placed within the genus Buellia rather than Rinodina. We consider that the type
specimens were previously misassigned as species of Rinodina. Based on the morphol-
ogy, chemistry, and phylogeny of specimens from the type locality, a new combination is
proposed here, Buellia setschwana.
Specimens examined
: China, Sichuan Prov.: Liangshan Prefecture, Huili Co., on the
way from Dechang to Huili, beside Beijing-Kunming Railway, 1879 m, on rock, April 11,
2019, Wang Lisong et al. 19-62862, 19-62864; Jinyang Co., Dashaba Vil., 640 m, on rock,
12 May 2017, Wang Lisong et al. 17-55142. Yunnan Prov.: Xinping Co., Mopanshan Mt.,
2540 m, on rock, 20 December 2008, Wang Lisong et al. 08-29962; Kunming Ci., Yinmin
Town, 1100 m, on rock, 16 May 2014, Wang Lisong et al. 14-43770; Nanjian Co., Zhonghua
Vil., Kongquedu Port, 2799 m, on rock, 4 July 2015, Ye Xin et al. 15-47743,15-47744.
Diversity 2023,15, 705 16 of 22
Diversity 2023, 15, x FOR PEER REVIEW 15 of 20
Figure 6. Morphology and anatomy of Buellia setschwana. (A) Thallus and apothecia of the holotype
(W-1171). (B–D) Specimen collected from the type locality (KUN 15-47744). (B) Thallus and apothe-
cia. (C) Buellia-type of ascospores. (D) Section of apothecia. Scale bars: 2 mm (A,B); 10 µm (C); 20
µm (D).
Notes: R. hengduanensis is characterized by subsquamulose, greyish-brown to oliva-
ceous thallus, jigsaw-like areoles, and Dirinaria-type ascospores. Rinodina gennarii Bagl. is
a saxicolous species with the same type of ascospores that has been reported from China
[26,27,29,78]. It differs from this new species by having a whitish thallus, crowded brown
apothecia, and smaller ascospores (length <17.5 µm). Rinodina pycnocarpa H. Magn. was
Figure 6.
Morphology and anatomy of Buellia setschwana. (
A
) Thallus and apothecia of the holotype
(W-1171). (
B
–
D
) Specimen collected from the type locality (KUN 15-47744). (
B
) Thallus and apothecia.
(
C
)Buellia-type of ascospores. (
D
) Section of apothecia. Scale bars: 2 mm (
A
,
B
); 10
µ
m (
C
); 20
µ
m (
D
).
New species
Rinodina hengduanensis Q. Y. Zhong and Xin Y. Wang, sp. nov., Figure 7.
Diversity 2023,15, 705 17 of 22
Type:
China, Sichuan Prov., Liangshan Yi Autonomous Prefecture, Yanyuan Co.,
Weicheng Town, on sandstone, 2819 m elev., 27
◦
30
0
N, 101
◦
41
0
E, 28 July 2020, Wang Xinyu,
Zhong Qiuyi, and Wang Luting no. XY20-290 (KUN, holotype).
MycoBank No. 847637
Characterized by saxicolous, subsquamulose, greyish brown to olivaceous thallus,
lecanorine apothecium, and Dirinaria-type ascospores.
Etymology:
The epithet ‘hengduanensis’ refers to its distribution, which is mainly in
the Hengduan Mountains region.
Description
:Thallus crustose and areolate in the center, areoles jigsaw-like, 0.2–0.4 mm
wide, closely attached to the substrate, continuous to scattered, margins prominent with
extending squamules, appearing subsquamulose; upper surface flat, greyish brown to olive
green, slightly glossy, non-pruinose; prothallus sometimes present (when thallus scattered),
black; vegetative propagules absent. Apothecia common, usually scattered, lecanorine,
broadly attached, 0.5–0.8 mm in diam.; disc dark brown to black, plane or convex, irregular;
margin concolorous with the thallus, crenulate, persistent; exciple hyaline, hyphae parallel,
with algal cells; hymenium colorless, hyaline, not inspersed, 100–125
µ
m high; paraphyses
simple, contiguous, apices expanded and septate, brown pigmented; epihymenium ochre
brown; hypothecium hyaline, colorless, not inspersed, 80–100
µ
m deep; asci 8-spored,
Lecanora-type; ascospores 1-septate, brown or dark, Type A development, Dirinaria-type,
20–24 ×7.5–12.5 µm
, swollen at the septum. Swelling becomes more obvious upon treat-
ment with KOH. Usually with oil inspersed when mature, torus present. Pycnidia were
not seen.
Chemistry
: Spot tests for cortex were all negative; medulla I
−
contained traces of
gyrophoric acid and skyrin, confirmed by TLC.
Ecology and distribution
: This species grows on rocks at approximately 2800 m
elevation. It is reported from Sichuan and Yunnan provinces in China.
Notes
:R. hengduanensis is characterized by subsquamulose, greyish-brown to oliva-
ceous thallus, jigsaw-like areoles, and Dirinaria-type ascospores. Rinodina gennarii Bagl.
is a saxicolous species with the same type of ascospores that has been reported from
China [
26
,
27
,
29
,
78
]. It differs from this new species by having a whitish thallus, crowded
brown apothecia, and smaller ascospores (length < 17.5
µ
m). Rinodina pycnocarpa H. Magn.
was originally reported from China. It also has a subsquamulose thallus and saxicolous
habit, but it can be differentiated by its Bicincta-type and smaller ascospores (<18
µ
m).
Three other species of Rinodina have similar subsquamulose thalli: R. dolichospora Malme
can be differentiated by its Pachysporaria-type I spores and lack of substances; R. intermedia
Bagl. can be recognized by its 3-septate to submuriform ascospores with type A develop-
ment; whereas R. willeyi Sheard and Giralt can be separated by having soredia on the areole
margin (sublabriform soralia) and its pannarin and zeorin content. R. hengduanensis would
key out at couplet 4 in Lee and Hur’s paper because it has a continental distribution and
has Dirinaria-type, larger (length > 20 µm) ascospores [76].
Specimens examined
: China, Sichuan Prov.: Liangshan Prefecture, Yanyuan Co.,
Weicheng Town, 2817–2865 m, on sandy rock, 28 July 2020, Wang Lisong et al. 20-66506,
XY20-290, XY20-287. Yunnan Prov.: Diqing Prefecture, Shangri-La City, Luoji Vil., Niru Vil.,
2774–2804 m, on rock, 21 July 2020, Wang Xinyu et al. XY20-44, XY20-41.
Diversity 2023,15, 705 18 of 22
Diversity 2023, 15, x FOR PEER REVIEW 16 of 20
originally reported from China. It also has a subsquamulose thallus and saxicolous habit,
but it can be differentiated by its Bicincta-type and smaller ascospores (<18 µm). Three
other species of Rinodina have similar subsquamulose thalli: R. dolichospora Malme can be
differentiated by its Pachysporaria-type I spores and lack of substances; R. intermedia Bagl.
can be recognized by its 3-septate to submuriform ascospores with type A development;
whereas R. willeyi Sheard and Giralt can be separated by having soredia on the areole
margin (sublabriform soralia) and its pannarin and zeorin content. R. hengduanensis would
key out at couplet 4 in Lee and Hur’s paper because it has a continental distribution and
has Dirinaria-type, larger (length >20 µm) ascospores [76].
Specimens examined: China, Sichuan Prov.: Liangshan Prefecture, Yanyuan Co.,
Weicheng Town, 2817–2865 m, on sandy rock, July 28, 2020, Wang Lisong et al. 20-66506,
XY20-290, XY20-287. Yunnan Prov.: Diqing Prefecture, Shangri-La City, Luoji Vil., Niru
Vil., 2774–2804 m, on rock, July 21, 2020, Wang Xinyu et al. XY20-44, XY20-41.
Figure 7.
Morphology and anatomy of Rinodina hengduanensis. (
A
) Thallus and apothecia. (
B
)Diri-
naria-type ascospores. (C) Section of apothecia. Scale bars: 2 mm (A); 10 µm (B); 20 µm (C).
5. Conclusions
In the present study, using morphological, chemical, and phylogenetic methods, new
records for the five endemic species Rinodina cornutula,R. globulans,R. handelii,R. setschwana,
and R. pluriloculata from Southwest China were documented based on specimens collected
from the holotype localities. Rinodina setschwana was transferred to the genus Buellia and
proposed as Buellia setschwana. One new species was discovered: Rinodina hengduanensis.
Photographs, chemical compounds, and sequences support the circumscription of these
species. However, such details are lacking for many of the previously proposed species
of Rinodina. The absence of genetic material means that no molecular phylogeny can yet
be constructed that includes all species currently assigned to the genus. For those species
lacking genetic material, further new collections are required from the type localities.
Diversity 2023,15, 705 19 of 22
Currently available keys based on morphology and chemistry are either outdated,
limited in their geographic scope, or do not include the recently described Rinodina spp. A
full revision of Rinodina is required in order to produce a key that should incorporate all
described species across its full global distribution.
Author Contributions:
Conceptualization, X.W. and L.W.; methodology, X.W.; software, Q.Z. and
M.A.; validation, F.R.W., A.Y. and Q.Z.; formal analysis, Q.Z. and M.A.; investigation, Q.Z., M.A. and
X.W.; resources, M.A. and A.Y.; data curation, Q.Z. and M.A.; writing—original draft preparation,
Q.Z.; writing—review and editing, F.R.W. and X.W.; visualization, Q.Z. and A.Y.; supervision, X.W.
and L.W.; project administration, X.W. and L.W.; funding acquisition, X.W., Y.J. and L.W. All authors
have read and agreed to the published version of the manuscript.
Funding:
This research was funded by Flora Lichenum Sinicorum, grant number 31750001; the Sec-
ond Tibetan Plateau Scietific Expedition and Research Program (STEP), grant number 2019QZKK0503;
Youth Innovation Promotion Association CAS, grant number 2020388, Yunnan Young and Elite Tal-
ents Project, National Natural Science Foundation of China, grant number 31970022,32060001; and
the Government Project of Yunnan Province (YNWR-QNBJ-2018-085).
Institutional Review Board Statement: Not applicable.
Data Availability Statement:
Publicly available datasets were analyzed in this study. This data can
be found from here: https://www.ncbi.nlm.nih.gov/ (accessed on 24 September 2022).
Acknowledgments:
We would like to express our deep thanks to the herbaria WU and US for
providing type specimens or digital images. We are grateful to Zhang Yanyun, Xie Congmiao, Wang
Luting, Ye Xin, and Wang Weicheng for their help with field research.
Conflicts of Interest: The authors declare no conflict of interest.
References
1.
Giralt, M. The lichen genera Rinodina and Rinodinella (lichenized Ascomycetes, Physciaceae) in the Iberian Peninsula. Bibl. Lichenol.
2001,79, 1–160.
2.
Mayrhofer, H.; Moberg, R. Rinodina. In Nordic Lichen Flora; Ahti, T., Jørgensen, P.M., Kristinsson, H., Eds.; Nordic Lichen Society:
Uddevalla, Sweden, 2002; Volume 2, pp. 41–69.
3.
Giralt, M.; Matzer, M. The corticolous species of the genus Rinodina with biatorine or lecideine apothecia in southern Europe and
Macaronesia. Lichenologist
1994
,26, 319–332. Available online: https://www.researchgate.net/publication/231922212 (accessed
on 11 April 2023). [CrossRef]
4.
Giralt, M.; Mayrhofer, H.; Sheard, J.W. The corticolous and lignicolous sorediate, blastidiate and isidiate species of the genus
Rinodina in southern Europe. Lichenologist
1995
,27, 3–24. Available online: https://www.researchgate.net/publication/231957857
(accessed on 11 April 2023). [CrossRef]
5.
Mayrhofer, H. Die saxicolen Arten der Flechtengattungen Rinodina und Rinodinella in der Alten Welt. J. Hattori Bot. Lab.
1984
,55,
327–493.
6.
Sheard, J.W. The Lichen Genus Rinodina (Ach.) Gray (Lecanoromycetidae, Physciaceae) in North America, North of Mexico; NRC Research
Press: Ottawa, EO, Canada, 2010; pp. 1–246.
7.
Sheard, J.W. A synopsis and new key to the species of Rinodina (Ach.) Gray (Physciaceae, lichenized Ascomycetes) presently
recognized in North America. Herzogia 2018,31, 395–423.
8.
Sheard, J.W.; Ezhkin, A.K.; Galanina, I.A.; Himelbrant, D.; Kuznetsova, E.; Shimizu, A.; Stepanchikova, I.; Thor, G.; Tønsberg, T.;
Yakovchenko, L.S.; et al. The lichen genus Rinodina (Physciaceae, Caliciales) in north-eastern Asia. Lichenologist
2017
,49, 617–672.
Available online: https://www.researchgate.net/publication/321054036 (accessed on 12 April 2023). [CrossRef]
9.
Helms, G.; Friedl, T.; Rambold, G. Phylogenetic relationships of the Physciaceae inferred from rDNA sequence data and selected
phenotypic characters. Mycologia
2003
,95, 1078–1099. Available online: https://www.researchgate.net/publication/49674865
(accessed on 13 April 2023). [CrossRef]
10.
Nadyeina, O.; Grube, M.; Mayrhofer, H. A contribution to the taxonomy of the genus Rinodina (Physciaceae, lichenized
Ascomycotina) using combined ITS and mtSSU rDNA data. Lichenologist
2010
,42, 521–531. Available online: http://europepmc.
org/article/PMC/3223597 (accessed on 13 April 2023). [CrossRef]
11.
Gaya, E.; Högnabba, F.; Holguin, A.; Molnar, K.; Fernández-Brime, S.; Stenroos, S.; Arup, U.; Søchting, U.; den Boom, P.V.;
Lücking, R.; et al. Implementing a cumulative supermatrix approach for a comprehensive phylogenetic study of the Teloschistales
(Pezizomycotina, Ascomycota). Mol. Phylogenet. Evol. 2012,63, 374–387. [CrossRef]
12.
Resl, P.; Mayrhofer, H.; Clayden, S.R.; Spribille, T.; Thor, G.; Tønsberg, T.; Sheard, J.W. Morphological, chemical and species
delimitation analyses provide new taxonomic insights into two groups of Rinodina.Lichenologist
2016
,48, 469–488. Available
online: https://www.researchgate.net/publication/308793339 (accessed on 12 April 2023). [CrossRef]
Diversity 2023,15, 705 20 of 22
13.
Wen, J. Evolution of eastern Asian and eastern North American disjunct distributions of flowering plants. Annu. Rev. Eco. Evol.
Syst. 1999,30, 421–455. [CrossRef]
14.
Xiang, Q.-Y.; Soltis, D.E.; Soltis, P.S. The eastern Asian and eastern and western North American disjunction: Congruent
phylogenetic patterns in seven diverse genera. Mol. Phylogenet. Evol. 1998,10, 178–190. [CrossRef]
15.
Qian, H. Floristic relationships between eastern Asia and North America: Test of Gray’s hypothesis. Am. Nat.
2002
,160, 317–332.
Available online: https://www.researchgate.net/publication/23174933 (accessed on 12 April 2023). [CrossRef]
16.
Ding, W.; Ree, R.; Spicer, R.; Xing, Y. Ancient orogenic and monsoon-driven assembly of the world’s richest temperate alpine
flora. Science 2020,369, 578–581. [CrossRef] [PubMed]
17.
Jatta, A. Licheni Cinesi raccolti allo Shen-si negli anni 1894-1898 dal. rev. Padre Missionario, G. Giraldi. Nuovo G. Bot. Ital.
1902
,9,
477. Available online: https://bibdigital.rjb.csic.es/viewer/12810/?offset=#page=490&viewer=picture&o=bookmark&n=0&q=
(accessed on 11 April 2023).
18. Paulson, R. Lichens of Mount Everest. Bot. J. Linn. Soc. Lond. 1925,63, 189–193.
19.
Zahlbruckner, A. Lichenes (Übersicht über sämtliche bisher aus China bekannten Flechten). In Symbolae Sinicae: Botanische
Ergebnisse der Expedition der Akademie der Wissenschaften in Wien Nach Südwest-China, 1914–1918; Handel-Mazzetti, H.F., Brotherus,
V.F., Eds.; Springer: Wien, Austria, 1930; Volume 3, pp. 226–234. Available online: https://www.biodiversitylibrary.org/item/15
292#page/1/mode/1up (accessed on 12 April 2023).
20.
Magnusson, A.H. Lichens from Central Asia, I. In Reports Scientific Expedition North-West Provinces of China (the Sino-Swedish
Expedition) 13, XI, Botany, 1; Hedin, S., Ed.; Aktiebolaget Thule: Stockholm, Sweden, 1940; pp. 145–156.
21.
Magnusson, A.H. Lichens from Central Asia II. In Reports Scientific Expedition North-West Provinces of China (the Sino-Swedish
Expedition) 22, XI, Botany, 2; Hedin, S., Ed.; Aktiebolaget Thule: Stockholm, Sweden, 1944; pp. 55–57.
22. Moreau, F.; Moreau, M.F. Lichens de Chine. Rev. Bryol. Et Lichenol. 1951,20, 183–199.
23. Wang Yang, J.-R.; Lai, M.-J. A checklist of the lichens of Taiwan. Taiwania 1973,18, 83–104.
24.
Wu, J.-N.; Xiang, T.A. Primary study on lichens from Yuntai Mountain, Lianyungang, Jiangsu Province. J. Nanjing Norm. Univ.-Nat.
Sci. 1981,3, 1–11. (In Chinese)
25.
Xu, B.S. Cryptogamic Flora of the Yangtze Delta and Adjacent Regions; Shanghai Scientific and Technical Publishers: Shanghai, China,
1989; pp. 254–255. (In Chinese)
26.
Abbas, A.; Jiang, Y.C.; Wu, J.N. Lichens from the Tianshan Mountains, Xinjiang, China. J. Nanjing Norm. Univ.-Nat. Sci.
1993
,16,
74–82. (In Chinese)
27.
Abbas, A.; Wu, J.N. Lichens of Xinjiang; Sci-Tec & Hygiene Publishing House of Xinjiang: Urumqi, China, 1998; pp. 1–178.
(In Chinese)
28.
Aptroot, A.; Seaward, M.R.D. Annotated checklist of Hongkong Lichens. Trop. Bryol.
1999
,17, 57–101. Available online:
https://www.researchgate.net/publication/265989726 (accessed on 10 April 2023).
29.
Aptroot, A.; Sparrius, L.B. New microlichens from Taiwan. Fungal Divers.
2003
,14, 1–50. Available online: https://www.
researchgate.net/publication/216829813 (accessed on 10 April 2023).
30.
Obermayer, W.; Blaha, J.; Mayrhofer, H. Buellia centralis and chemotypes of Dimelaena oreina in Tibet and other Central-Asian
regions. Symb. Bot. Ups.
2004
,34, 327–342. Available online: https://www.researchgate.net/publication/242552311 (accessed on
12 April 2023).
31.
Aptroot, A.; Saipunkaew, W.; Sipman, H.J.M.; Sparrius, L.B.; Wolseley, P.A. New lichens from Thailand, mainly microlichens from
Chiang Mai. Fungal Divers.
2007
,24, 75–134. Available online: https://www.researchgate.net/publication/228515656 (accessed
on 10 April 2023).
32. Li, Y.; Chen, C.; Zhao, Z. A primary study on lichens from Mount Yi. J. Fungal Res. 2008,6, 70–73. (In Chinese)
33. Liu, M.; Wei, J.-C. Lichen diversity in Shapotou region of Tengger Desert, China. Mycosystema 2013,32, 42–50. (In Chinese)
34.
Joshi, S.; Jayalal, U.; Oh, S.-O.; Li, X.-R.; Jia, R.-L.; Hur, J.-S. New records of lichens from Shapotou area in Ningxia of Northwest
China. Mycosystema
2014
,33, 167–173. Available online: https://www.researchgate.net/publication/263655180 (accessed on 11
April 2023).
35.
Kondratyuk, S.Y.; Lökös, L.; Halda, J.P.; Moniri, M.H.; Farkas, E.; Park, J.S.; Lee, B.G.; Oh, S.-O.; Hur, J.-S. New and noteworthy
lichen-forming and lichenicolous fungi: 5. Acta Bot. Hung. 2016,58, 319–396. [CrossRef]
36. Ren, Q.; Zheng, X. Notes on the genus Rinodina in mainland China. J. Liaocheng Univ. (Nat. Sci.) 2020,33, 85–97. (In Chinese)
37.
Tursun, A.; Shahidin, H.; Tumur, A. Taxonomic study on Rinodina in Xinjiang, China. Acta Bot. Boreali-Occident. Sin.
2020
,40,
1978–1988. (In Chinese)
38. Wei, J.-C. The Enumeration of Lichenized Fungi in China; China Forestry Publishing House: Beijing, China, 2020; pp. 450–454.
39.
Culberson, C.F. Improved conditions and new data for identification of lichen products by standardized thin-layer chromato-
graphic method. J. Chromatogr. A 1972,72, 113–125. [CrossRef] [PubMed]
40.
Orange, A.; James, P.W.; White, F.J. Microchemical Methods for the Identification of Lichens; British Lichen Society: London, UK, 2001;
pp. 1–101.
41.
Zoller, S.; Scheidegger, C.; Sperisen, C. PCR primers for the amplification of mitochondrial small subunit ribosomal DNA of lichen-
forming ascomycetes. Lichenologist
1999
,31, 511–516. Available online: https://www.researchgate.net/publication/229071472
(accessed on 12 April 2023). [CrossRef]
Diversity 2023,15, 705 21 of 22
42.
Gardes, M.M.; Bruns, T.D. ITS primers with enhanced specificity for basidiomycetes-application for the identification of mycor-
rhizae and rust. Mol. Ecol. 1993,2, 113–118. [CrossRef] [PubMed]
43.
White, T.J.; Bruns, T.; Lee, S.; Taylor, J. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In
PCR Protocols: A Guide to Methods and Applications; Innis, M.A., Gelfand, D.H., Sninsky, J.J., White, T.J., Eds.; Academic Press: San
Diego, CA, USA, 1990; pp. 315–322. Available online: https://www.researchgate.net/publication/223058289 (accessed on 12
April 2023).
44.
Hassouna, N.; Mithot, B.; Bachellerie, J.P. The complete nucleotide sequence of mouse 28S rRNA gene. Implications for the
process of size increase of the large subunit rRNA in higher eukaryotes. Nucleic Acids Res.
1984
,12, 3563–3583. [CrossRef]
[PubMed]
45.
Rehner, S.A.; Samuels, G.J. Taxonomy and phylogeny of Gliocladium analysed from nuclear large subunit ribosomal DNA
sequences. Mycol. Res. 1994,98, 625–634. [CrossRef]
46.
Vilgalys, R.; Hester, M. Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several
Cryptococcus species. J. Bacteriol.
1990
,172, 4238–4246. Available online: http://europepmc.org/article/PMC/213247 (accessed
on 12 April 2023). [CrossRef]
47.
Zhao, X.; Zhang, L.L.; Zhao, Z.T.; Wang, W.C.; Leavitt, S.D.; Lumbsch, H.T. Molecular phylogeny of the lichen genus Lecidella
focusing on species from mainland China. PLoS ONE
2015
,10, e0139405. Available online: https://www.researchgate.net/
publication/283181184 (accessed on 12 April 2023). [CrossRef]
48.
Katoh, K.; Standley, D.M. MAFFT multiple sequence alignment software version 7: Improvements in performance and usability.
Mol. Biol. Evol.
2013
,30, 772–780. Available online: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3603318/pdf/mst010.pdf
(accessed on 11 April 2023). [CrossRef]
49.
Chernomor, O.A.; von Haeseler, A.; Minh, B.Q. Terrace aware data structure for phylogenomic inference from supermatrices. Syst.
Biol.
2016
,65, 997–1008. Available online: http://europepmc.org/backend/ptpmcrender.fcgi?accid=PMC5066062&blobtype=pdf
(accessed on 10 April 2023). [CrossRef]
50.
Kalyaanamoorthy, S.; Minh, B.Q.; Wong, T.K.F.; von Haeseler, A.; Jermiin, L.S. ModelFinder: Fast model selection for accurate
phylogenetic estimates. Nat. Methods
2017
,14, 587–589. Available online: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC545
3245/pdf/emss-72237.pdf (accessed on 11 April 2023). [CrossRef]
51.
Nguyen, L.-T.; Schmidt, H.A.; von Haeseler, A.; Minh, B.Q. IQ-TREE: A fast and effective stochastic algorithm for estimating
maximum likelihood phylogenies. Mol. Biol. Evol.
2015
,32, 268–274. Available online: http://europepmc.org/backend/
ptpmcrender.fcgi?accid=PMC4271533&blobtype=pdf (accessed on 11 April 2023). [CrossRef] [PubMed]
52.
Trifinopoulos, J.; Nguyen, L.-T.; von Haeseler, A.; Minh, B.Q. W-IQ-TREE: A fast online phylogenetic tool for maximum likelihood
analysis. Nucleic Acids Res.
2016
,44, W232–W235. Available online: http://europepmc.org/article/PMC/4987875 (accessed on
12 April 2023).
53.
Zhou, X.; Shen, X.-X.; Hittinger, C.T.; Rokas, A. Evaluating fast maximum likelihood-based phylogenetic programs using empirical
phylogenomic data sets. Mol. Biol. Evol.
2017
,35, 486–530. Available online: https://academic.oup.com/mbe/article/35/2/486
/4644721 (accessed on 12 April 2023). [CrossRef] [PubMed]
54.
Ronquist, F.; Teslenko, M.; van der Mark, P.; Ayres, D.; Darling, A.; Höhna, S.; Larget, B.; Liu, L.; Suchard, M.A.; Huelsenbeck,
J.P. MrBayes 3.2: Efficient bayesian phylogenetic inference and model choice across a large model space. Syst. Biol.
2012
,
61, 539–542. Available online: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3329765/pdf/sys029.pdf (accessed on 12
April 2023). [CrossRef] [PubMed]
55.
Rambaut, A. FigTree v. 1.4.2. Institute of Evolutionary Biology, University of Edinburgh. Available online: http://tree.bio.ed.ac.
uk/software/figtree/ (accessed on 12 April 2023).
56.
Prieto, M.; Wedin, M. Phylogeny, taxonomy and diversification events in the Caliciaceae. Fungal Divers.
2017
,82, 221–238.
Available online: https://link.springer.com/article/10.1007/s13225-016-0372-y (accessed on 16 April 2023). [CrossRef]
57.
Tibell, L. Tholurna dissimilis and generic delimitations in Caliciaceae inferred from nuclear ITS and LSU rDNA phylogenies
(Lecanorales, lichenized ascomycetes). Mycol. Res. 2003,107, 1403–1418. [CrossRef]
58.
Ohmura, Y.; Sugimoto, M.; Yakovchenko, L.; Davydov, E.A. Additional species and ITS rDNA data for the lichen mycota of the
Imperial Palace grounds, Tokyo, Japan. Bull. Natl. Mus. Nat. Sci.
2022
,48, 1–16. Available online: https://www.researchgate.net/
publication/358952425 (accessed on 16 April 2023).
59.
Lohtander, K.; Ahti, T.; Stenroos, S.; Urbanavichus, G. Is Anaptychia monophyletic? A phylogenetic study based on nuclear and
mitochondrial genes. Ann. Bot. Fenn.
2008
,45, 55–60. Available online: https://www.researchgate.net/publication/232273420
(accessed on 16 April 2023). [CrossRef]
60.
Molina, M.C.; Divakar, P.K.; González, N. Success in the isolation and axenic culture of Anaptychia ciliaris (Physciaceae,
Lecanoromycetes) mycobiont. Mycoscience
2015
,56, 351–358. Available online: https://www.researchgate.net/publication/2728
91236 (accessed on 16 April 2023). [CrossRef]
61.
Wedin, M.; Baloch, E.; Grube, M. Parsimony analyses of mtSSU and nITS rDNA sequences reveal the natural relationships of the
lichen families Physciaceae and Caliciaceae. Taxon
2002
,51, 655–660. Available online: https://nhm2.uio.no/botanisk/lav/RLL/
PDF20/R24173.pdf (accessed on 16 April 2023). [CrossRef]
62.
Crespo, A.; Blanco, O.; Llimona, X.; Ferencová, Z.L.; Hawksworth, D.L. Coscinocladium, an overlooked endemic and monotypic
Mediterranean lichen genus of Physciaceae, reinstated by molecular phylogenetic analysis. Taxon 2004,53, 405–414. [CrossRef]
Diversity 2023,15, 705 22 of 22
63.
Aptroot, A.; Maphangwa, K.; Zedda, L.; Tekere, M.; Alvarado, P.; Sipman, H. The phylogenetic position of Culbersonia is in the
Caliciaceae (lichenized ascomycetes). Lichenologist 2019,51, 187–191. [CrossRef]
64. Schmull, M.; Miadlikowska, J.; Pelzer, M.; Stocker-Worgotter, E.; Hofstetter, V.; Fraker, E.; Hodkinson, B.P.; Reeb, V.; Kukwa, M.;
Lumbsch, H.T.; et al. Phylogenetic affiliations of members of the heterogeneous lichen-forming fungi of the genus Lecidea sensu
zahlbruckner (lecanoromycetes, ascomycota). Mycologia
2011
,103, 983–1003. Available online: https://www.researchgate.net/
publication/51192614 (accessed on 16 April 2023). [CrossRef] [PubMed]
65.
Miadlikowska, J.; Kauff, F.; Hofstetter, V.; Fraker, E.; Grube, M.; Hafellner, J.; Reeb, V.; Hodkinson, B.P.; Kukwa, M.; Lücking, R.;
et al. New insights into classification and evolution of the Lecanoromycetes (Pezizomycotina, Ascomycota) from phylogenetic
analyses of three ribosomal RNA- and two protein-coding genes. Mycologia
2006
,98, 1088–1103. Available online: https:
//www.researchgate.net/publication/6344409 (accessed on 16 April 2023). [CrossRef] [PubMed]
66.
Grube, M.; Arup, U. Molecular and morphological evolution in the Physciaceae (Lecanorales, lichenized Ascomycotina), with
special emphasis on the genus Rinodina.Lichenologist 2001,33, 63–72. [CrossRef]
67.
Liu, D.; Hur, J.-S. Revision of the lichen genus Phaeophyscia and allied atranorin absent taxa (Physciaceae) in South Korea. Microor-
ganisms 2019,7, 242. Available online: https://www.mdpi.com/2076-2607/7/8/242 (accessed on 16 April 2023). [CrossRef]
68.
Kondratyuk, S.Y.; L˝okös, L.; Kim, J.A.; Jeong, M.-H.; Kondratiuk, A.; Oh, S.-O.; Hur, J.-S. Kashiwadia gen. nov. (Physciaceae,
lichen-forming Ascomycota), proved by phylogenetic analysis of the Eastern Asian Physciaceae. Acta Bot. Hung.
2014
,56, 369–378.
Available online: https://www.researchgate.net/publication/277924093 (accessed on 16 April 2023). [CrossRef]
69.
Lohtander, K.; Källersjö, M.; Moberg, R.; Tehler, A. The family Physciaceae in Fennoscandia: Phylogeny inferred from ITS
sequences. Mycologia
2000
,92, 728–735. Available online: https://www.researchgate.net/publication/232273357 (accessed on 16
April 2023). [CrossRef]
70.
Kondratyuk, S.Y.; L˝okös, L.; Kim, J.A.; Jeong, M.-H.; Zarei-Darki, B.; Hur, J.-S. Oxnerella safavidiorum gen. et spec. nov.
(Lecanoromycetidae, Ascomycota) from Iran (Asia) proved by phylogenetic analysis. Acta Bot. Hung.
2014
,56, 379–398. Available
online: https://go.gale.com/ps/i.do?p=AONE&u=googlescholar&id=GALE\T1\textbar{}A458953398&v=2.1&it=r&asid=e8
83ca1e (accessed on 16 April 2023). [CrossRef]
71.
Marthinsen, G.; Rui, S.; Timdal, E. OLICH: A reference library of DNA barcodes for Nordic lichens. Biodivers. Data J.
2019
,7,
e36252. Available online: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6711938/ (accessed on 16 April 2023). [CrossRef]
72.
Zhong, Q.; Zhang, Y.; Wang, X.; Timdal, E.; Gong, H.; Wang, Z.; Wang, L. Phaeorrhiza (Physciaceae), a new lichen genus record to
China. Phytotaxa 2021,510, 228–238. [CrossRef]
73.
Yang, M.-X.; Wang, X.-Y.; Liu, D.; Zhang, Y.-Y.; Li, L.-J.; Yin, A.-C.; Scheidegger, C.; Wang, L.-S. New species and records of Pyxine
(Caliciaceae) in China. MycoKeys
2019
,45, 93–109. Available online: https://mycokeys.pensoft.net/article/29374/ (accessed on
16 April 2023). [CrossRef] [PubMed]
74.
Sipman, H.; Aptroot, A. Ikaeria serusiauxii, a new Caloplaca-like lichen from Macaronesia and mainland Portugal, with a lichen
checklist for Porto Santo. Plant Fungal Syst.
2020
,65, 120–130. Available online: https://www.semanticscholar.org/paper/02f5c2
8f8a50f47d97994584d5283511e0c4c6d8 (accessed on 16 April 2023). [CrossRef]
75.
Kaschik, M. Taxonomic Studies on Saxicolous Species of the Genus Rinodina (Lichenized Ascomycetes, Physciaceae) in the Southern
Hemisphere with Emphasis in Australia and New Zealand; Bibliotheca Lichenologica: Stuttgart, Germany, 2006; pp. 1–162.
76.
Lee, B.G.; Hur, J.-S. Two new Rinodina lichens from South Korea, with an updated key to the species of Rinodina in the far
eastern Asia. Mycokeys
2022
,87, 159–182. Available online: https://europepmc.org/article/PMC/PMC8891229 (accessed on 16
April 2023). [CrossRef] [PubMed]
77.
Bhattacharya, D.; Friedl, T.; Helms, G. Vertical evolution and intragenic spread of lichen-fungal group I introns. J. Mol. Evol.
2002
,
55, 74–84. [CrossRef]
78. Abbas, A.; Wu, J.N. The lichens from Tielimaiti Pass Kuqa County, Xinjiang, China. Arid Zone Res. 1994,11, 19–23. (In Chinese)
Disclaimer/Publisher’s Note:
The statements, opinions and data contained in all publications are solely those of the individual
author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to
people or property resulting from any ideas, methods, instructions or products referred to in the content.
