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Revision of Immersaria and a new lecanorine genus in Lecideaceae (lichenised Ascomycota, Lecanoromycetes)

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The species Immersaria cupreoatra has been included in Bellemerea. This caused us to reconsider the relationships between Bellemerea and the lecanorine species of Immersaria and to question the monophyly of Immersaria. Amongst 25 genera of the family Lecideaceae, most have lecideine apothecia, the exceptions being Bellemerea and Koerberiella, which have lecanorine apothecia. According to previous classifications, Immersaria included species with both lecanorine and lecideine apothecia. A five-loci phylogenetic tree (nrITS, nrLSU, RPB1, RPB2, and mtSSU) for Lecideaceae showed that Immersaria was split into two clades: firstly, all the lecideine apotheciate species and secondly, all the lecanorine apotheciate species. The latter clade was closely related to the remaining lecanorine apotheciate genera: Bellemerea and Koerberiella. Therefore, the genus concept of Immersaria is revised accordingly and a new lecanorine genus Lecaimmeria is proposed. Furthermore, four new species for Immersaria and seven new species and three new combinations for the new genus Lecaimmeria are proposed. Keys to Immersaria and the new genus Lecaimmeria are provided.
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Revision of Immersaria and a new
lecanorine genus in Lecideaceae
(lichenised Ascomycota, Lecanoromycetes)
Cong-Miao Xie1, Li-Song Wang2, Zun-Tian Zhao1,
Yan-Yun Zhang2, Xin-Yu Wang2, Lu-Lu Zhang3
1Key Laboratory of Plant Stress Research, College of Life Sciences, Shandong Normal University, Jinan, Shan-
dong, 250014, China 2CAS Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming
Institute of Botany, Heilongtan, Kunming, Yunnan, 650204, China 3Institute of Environment and Ecology,
Shandong Normal University, Jinan, Shandong, 250014, China
Corresponding authors: Xin-Yu Wang (wangxinyu@mail.kib.ac.cn), Lu-Lu Zhang (612038@sdnu.edu.cn)
Academic editor: Gerhard Rambold|Received 5 August 2021|Accepted 17 January 2022|Published 15 February 2022
Citation: Xie C-M, Wang L-S, Zhao Z-T, Zhang Y-Y, Wang X-Y, Zhang L-L (2022) Revision of Immersaria and a
new lecanorine genus in Lecideaceae (lichenised Ascomycota, Lecanoromycetes). MycoKeys 87: 99–132. https://doi.
org/10.3897/mycokeys.87.72614
Abstract
e species Immersaria cupreoatra has been included in Bellemerea. is caused us to reconsider the
relationships between Bellemerea and the lecanorine species of Immersaria and to question the monophyly
of Immersaria. Amongst 25 genera of the family Lecideaceae, most have lecideine apothecia, the exceptions
being Bellemerea and Koerberiella, which have lecanorine apothecia. According to previous classications,
Immersaria included species with both lecanorine and lecideine apothecia. A ve-loci phylogenetic tree
(nrITS, nrLSU, RPB1, RPB2, and mtSSU) for Lecideaceae showed that Immersaria was split into two
clades: rstly, all the lecideine apotheciate species and secondly, all the lecanorine apotheciate species.
e latter clade was closely related to the remaining lecanorine apotheciate genera: Bellemerea and
Koerberiella. erefore, the genus concept of Immersaria is revised accordingly and a new lecanorine genus
Lecaimmeria is proposed. Furthermore, four new species for Immersaria and seven new species and three
new combinations for the new genus Lecaimmeria are proposed. Keys to Immersaria and the new genus
Lecaimmeria are provided.
Keywords
China, generic classication, lecanorine apothecia, lichen, phylogeny, taxonomy
MycoKeys 87: 99–132 (2022)
doi: 10.3897/mycokeys.87.72614
https://mycokeys.pensoft.net
Copyright Cong-Miao Xie et al. 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.
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Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
100
Introduction
e lichen genus Immersaria Rambold & Pietschm. (Rambold 1989) was originally
split from the genus Lecidea Ach. in order to accommodate the species Immersaria
athroocarpa (Ach.) Rambold & Pietschm. e genus Immersaria was characterised by
its brown thallus with an epinecral layer, a pruinose margin and an amyloid medulla,
immersed apothecia with a somewhat reduced proper margin and Porpidia-type asci
with eight, simple, halonate ascospores (Rambold 1989). Subsequently, Calatayud and
Rambold (1998) enlarged the scope of the genus by including the lecanorine species,
Immersaria mehadiana Calat. & Rambold and I. cupreoatra (Nyl.) Calat. & Rambold,
based on morphological characters only. Currently, eight species of Immersaria are
known worldwide (Lücking et al. 2017), three of which have lecanorine apothecia. Four
of these species were previously reported from China (Hertel 1977; Zhang et al. 2015).
e species Immersaria cupreoatra (Nyl.) Calat. & Rambold (= Lecanora cupreoatra
Nyl.) was previously included in Bellemerea Hafellner & Cl. Roux (Clauzade and Roux
1984), then into Immersaria by Calatayud and Rambold (1998). is caused us to
reconsider the relationships between Bellemerea and the lecanorine species of Immersaria
and to question the monophyly of Immersaria. e family Lecideaceae Chevall
originally included all the crustose lecideoid genera, but now only 25 genera have been
retained. Most of these are monospecic genera or small genera with under ve species
(Fryday and Hertel 2014; McCune et al. 2017). Most genera in Lecideaceae have
lecideine apothecia. ree exceptions are Bellemerea, Immersaria and Koerberiella Stein,
which have lecanorine apothecia. Only Immersaria has both lecanorine and lecideine
apothecia, according to the previous circumscription (Calatayud and Rambold 1998;
Valadbeigi et al. 2011). Calatayud and Rambold (1998) indicated that the presence
of “two types of ascomata” represent dierent stages of ontogeny. However, there was
no molecular evidence that could clarify the species-level phylogenetic relationships
within Immersaria. In the two-loci phylogenetic tree of Buschbom and Mueller
(2004), the lecideine species Immersaria usbekica (Hertel) M. Barbero, Nav.-Ros. &
Cl. Roux was related to Lecidea tessellata Flörke. However, because only two loci of one
lecideine species were included, this tree was insucient to clarify the relationship of
the lecanorine apotheciate species in Immersaria.
In this study, a phylogenetic tree of Lecideaceae, based on ve loci, is established in
order to verify the monophyly of Immersaria. e results show that Immersaria is split
into two clades. One clade includes all the lecideine apotheciate species, which is sister
to Lecidea tessellata, L. auriculata . Fr., Cyclohymenia epilithica McCune & M.J. Cur-
tis and the Porpidia albocaerulescens group and the Porpidia speirea group. e second
clade contains all the lecanorine apotheciate species and is closely related to the rest
of the lecanorine apotheciate genera within this family: Bellemerea and Koerberiella.
erefore, the genus concept of Immersaria is revised, retaining only the species with
lecideine apothecia. e lecanorine species of Immersaria are excluded and proposed as
a new genus, Lecaimmeria C.M. Xie, Lu L. Zhang & Li S. Wang. Furthermore, four
Revision of Immersaria and a new genus in Lecideaceae 101
new species for Immersaria and seven new species and three new combinations for the
new genus Lecaimmeria are proposed, based on the four-loci phylogenetic trees. Keys
to Immersaria and the new genus are provided below.
Methods
Morphological analysis
All the materials for this study were collected in mainland China, mostly from the Qinghai-
Tibetan Plateau, during the authors’ participation in e Second Tibetan Plateau Scientic
Expedition and Research Program. ese specimens were stored in the Herbarium of the
Kunming Institute of Botany, Chinese Academy of Sciences (KUN) and the Lichen Section
of the Botanical Herbarium, Shandong Normal University (SDNU). Type specimens
were loaned from the University of Helsinki (H) and Universität Wien (WU). High-
resolution photographs of type specimens were provided by the curators of H or obtained
from the website Global Plants (https://plants.jstor.org/). Morphological descriptions were
made from under a dissecting microscope COIX. Anatomical descriptions were based
on observations made from hand-cut sections, mounted in water and using a NIKON
microscope. Usually, twenty ascospores were measured and the values of measurement means
smallest measured-largest measured, with outlying values in brackets. Photographs were
captured with a NIKON Eclipse 50i microscope, equipped with a NIKON digital camera
(DSFi2 high-denition colour camera head, NIKON, Japan). e specimens were tested
with a 10% aqueous solution of potassium hydroxide (K), a solution of aqueous sodium
hypochlorite (C) and 3% Lugol’s iodine (I) in the medulla and the surface of the thallus.
Secondary metabolites of all the specimens were examined by thin-layer chromatography
(TLC) methods, using Solvents A, B and C, as described by Orange et al. (2001).
Phylogenetic analysis
Molecular analysis was carried out on the selected specimens. Genomic DNA was
extracted from dry or fresh specimens using a DNAsecure Plant Kit (Tiangen), following
the manufacturer’s instructions. Five gene loci were amplied by using the following
primers: ITS1F (Larena et al. 1999), ITS4 (White et al. 1990), LR0R (Rehner and
Samuels 1994), LR5 (Vilgalys and Hester 1990), gRPB1a (Stiller and Hall 1997),
fRPB1c (Matheny et al. 2002), RPB2–6f, RPB2–7cr (Liu et al. 1999), mrSSU1 and
mrSSU3R (Zoller et al. 1999). e 25 µl PCR mixture consisted of 2 µl DNA, 1 µl of
each primer, 12.5 µl 2 × Taq PCR MasterMix (Aidlab) (Taq DNA Polymerase [0.1unit/
ml]; 4 mM MgCl2; and 0.4 mM dNTPs) and 8.5 µl ddH2O. Conditions for PCR of
nrITS, nrLSU and mtSSU were set for an initial denaturation at 94 °C for 10 min,
followed by 34 cycles of denaturation at 95 °C for 45 s, annealing at 50 °C for 45 s,
extension at 72 °C for 90 s and a nal extension at 72 °C for 10 min. For RPB1 and
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
102
RPB2, the parameters were set to an initial denaturation at 94 °C for 10 min, followed by
34 cycles of denaturation at 95°C for 45 s, annealing at 52 °C for 50 s, extension at 72°C
for 60 s and a nal extension at 72 °C for 5 min. e PCR products were sequenced
using Sanger technology by the company of Tsingke Biological Technology (Beijing).
e raw sequences were assembled and edited using SeqMan v.7.0 (DNAstar
packages). Sequences extracted from new materials with each gene locus were aligned with
additional sequences that were available from GenBank (Suppl.material1:TableS1),
by using MEGA v.10.0 and an online version of MAFFT v.7.0 to generate nrITS-
nrLSU-RPB1-RPB2-mtSSU or nrITS-nrLSU-RPB1-RPB2 matrices. e ve or
four gene matrices were combined by SequenceMatrix v.1.7.8. and the concatenated
alignments were estimated by PartitionFinder 2 (Lanfear et al. 2017), based on the
Bayesian Information Criterion (BIC), to nd the most appropriate nucleotide
substitution model for each of the ve loci.
Phylogenetic relationships were inferred using Bayesian Inference (BI) and Maximum
Likelihood (ML). ML analyses were performed with RAxMLHPC using the general time
reversible model of nucleotide substitution with the gamma model of rate heterogeneity
(GTRGAMMA or GTRCAT). e analyses were run with a rapid bootstrap analysis using
1000 replicates with data partitioned. e Bayesian method was performed with MrBayes
v.3.1.2 (Huelsenbeck and Ronquist 2001). Four Markov chains were run with 2 million
generations for each dataset and trees were sampled every 100 generations. It was ensured
that the average standard deviation of split frequencies was lower than 0.01. Posterior
probabilities above 0.9 and bootstrap support above 70% were considered signicant
supporting values. All the trees were visualised with FigTree v. 1.4.0 (Rambaut 2012).
Results
A total of 172 sequences of the nrITS, nrLSU, RPB1, RPB2, and mtSSU were
generated from 61 specimens representing 57 species. Although the ve-loci tree only
poorly resolved the hierarchy of genera within the family Lecideaceae and the split
between the lecanorine and lecideine genera of Lecideaceae was without robust support,
nonetheless the results revealed that the genus Immersaria was not a monophyletic
lineage. Rather, it was divided into two distant and well-supported lineages: clade 1
which contained the lecideine apotheciate species and clade 2 which contained the
lecanorine apotheciate species (Fig. 1).
Clade 1, together with Amygdalaria Norman, Cyclohymenia McCune & M.J.
Curtis, Lecidea s str. and Porpidia Körb. (Fig. 1) formed a well-supported clade (93%
MLBS and 0.99 PP), all of which have lecideine apothecia. Amygdalaria, Porpidia
and Lecidea s str. were nested together, which was consistent with the results of previous
research (Buschbom and Mueller 2004; Fryday et al. 2014). However, the relationships
between these genera still need further research. ere was a high level of support for a
monophyletic lineage of lecideine apotheciate species of Immersaria, with these being
Revision of Immersaria and a new genus in Lecideaceae 103
sister to the Lecidea tessellata, L. auriculata, Cyclohymenia, the Porpidia albocaerulescens
group and the Porpidia speirea group. e type species of Immersaria (I. athroocarpa) was
included in the lineage. us, only those Immersaria species with lecideine apothecia
belong to Immersaria s. str. is revised concept of the genus Immersaria is as follows:
a glossy surface of thallus with an epinecral layer, immersed lecideine apothecia with a
reduced margin and Porpidia-type asci with halonate ascospores.
Figure 1. Phylogenetic tree constructed from Maximum Likelihood analyses in Lecideaceae, based on the
concatenated nrITS-nrLSU-RPB1-RPB2-mtSSU dataset. Maximum Likelihood bootstrap probabilities
above 70% (left) and Bayesian Inference posterior probabilities above 0.9 (right) are given at the nodes.
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
104
ere was also a high level of support for clade 2 as a monophyletic lineage (100%
MLBS and 1.00 PP), which was clustered with other genera of Lecideales with lecanorine
apothecia: Bellemerea and Koerberiella (Fig. 1). Bellemerea could be distinguished from
Clade 2 by its amyloid ascospores and Koerberiella by its adnate apothecia. Although the
topology of clade 2 for Bellemerea and Koerberiella is not robust, there are conspicuous
dierences in their morphology and signicant dierences between the bases in their
nucleotide sequences. Since clade 2 is monophyletic with strong support, a new genus,
Lecaimmeria, is proposed to accommodate clade 2. e new genus has immersed
lecanorine apothecia with a white margin and a distinct plectenchyma developed on
top of the orange epihymenium.
Two additional phylogenetic trees were constructed, based on four loci (nrITS,
nrLSU, RPB1, and RPB2), in order to assess the phylogenetic position of species
within Immersaria and Lecaimmeria, respectively. e phylogenetic tree of Immersaria
was comprised of one highly supported clade with ve separate lineages, based on 105
sequences from 37 specimens (Fig. 2). All the species with brown, orange, irregular
or aggregate thalli formed respective monophyletic lineages. Immersaria shangrilaensis
C.M. Xie & Lu L. Zhang formed a well-supported clade and the aggregate areolae
clearly distinguished I. shangrilaensis from other species. Immersaria ferruginea C.M.
Xie & Li S. Wang also formed a well-supported clade and diered from other species
by its greyish-brown thallus. It seems that Immersaria shangrilaensis is sister to I. fer-
ruginea, but the nodes were without support. In addition, the morphology is distinct
between Immersaria shangrilaensis and I. ferruginea. e robust lineage Immersaria
aurantia C.M. Xie & Li S. Wang was distinguished by its irregular, conspicuously
orange thallus and green epihymenium. Immersaria athroocarpa was sister to I. venusta
C.M. Xie & Xin Y. Wang, but diered in its convex, polygon areolae and densely
crowded apothecia.
e phylogenetic tree of Lecaimmeria was comprised of one well-supported clade
with nine separate lineages, based on 140 sequences from 61 specimens (Fig. 3).
Immersaria” sp. and Lecaimmeria iranica (Valadb., Sipman & Rambold) C.M. Xie
comprised the basal group. “Immersaria” sp. has only been recorded from Macedonia
and Lecaimmeria iranica has been recorded from Inner Mongolia in China and from
Iran. Lecaimmeria tuberculosa C.M. Xie & Xin Y. Wang was sister to L. qinghaiensis
C.M. Xie & Li S. Wang, but conspicuously diered in its tuberculiform conidiomata.
Lecaimmeria mongolica C.M. Xie & Lu L. Zhang formed a well-supported monophyletic
lineage and its population was mainly recognised by its orange, irregular areolae and
gyrophoric acid content. Lecaimmeria botryoides C.M. Xie & Li S. Wang formed a
highly supported sister group to L. orbicularis C.M. Xie & Lu L. Zhang, L. lygaea
C.M. Xie & Lu L. Zhang and L. tibetica C.M. Xie & Xin Y. Wang, but diered in its
crowded apothecia. Lecaimmeria orbicularis formed a highly supported sister group to
L. lygaea and L. tibetica, but diered in its round apothecia and the white margin of
the apothecia. Lecaimmeria lygaea was seemingly sister to L.tibetica and diered in its
areolae having a black margin and with a well-developed prothallus between areolae.
Revision of Immersaria and a new genus in Lecideaceae 105
Discussion
Revised boundaries of Immersaria
Formerly, the boundaries for Immersaria species were: lecanorine or lecideine type of
immersed apothecia, production of conuentic acid and gyrophoric acid and ostiole
or stellate shapes of conidiomata. However, these characters were not good characters
by which to distinguish this genus. e lecanorine species Immersaria cupreoatra was
previously included in Bellemerea. Based on many specimens from China, it was also
discovered that the ostiole or stellate shapes of conidiomata appeared in dierent stages
of ontogeny. e main substances produced in the genus are conuentic acid and
gyrophoric acid; conuentic acid only occurs in lecideine species, whereas gyrophoric
acid appears in lecanorine species, with the exception of one lecideine species
Immersaria usbekica. Furthermore, these characters, the types of apothecia and the
shapes of conidiomata could not be applied as proper delimitations to classify species
within Immersaria, neither were they supported by the phylogeny.
Figure 2. Phylogenetic tree constructed from Maximum Likelihood analyses in Immersaria, based on
the concatenated nrITS-nrLSU-RPB1-RPB2 dataset. Maximum Likelihood bootstrap probabilities above
70% (left) and Bayesian Inference posterior probabilities above 0.9 (right) are given at the nodes. Solid
brown rectangle: thallus brown; solid orange rectangle: thallus yellow brown to orange brown; solid red
rectangle: thallus rusty; hollow brown rectangle: thallus pale yellow brown. Solid green circle: green
epihymenium; solid grey circle: without apothecia; hollow green circular: brown epihymenium.
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
106
e ve-loci based analysis (Fig. 1) was incompatible with previous circumscriptions
of the genus Immersaria, the members of which in this study are dened by their
lecideine immersed apothecia, brown areolae with an epinecral layer and brown/green
epihymenium without a plectenchyma. Consequently, a new genus, Lecaimmeria,
Figure 3. Phylogenetic tree constructed from Maximum Likelihood analyses in Lecaimmeria, based on
the concatenated nrITS-nrLSU-RPB1-RPB2 dataset. Maximum Likelihood bootstrap probabilities above
70% (left) and Bayesian Inference posterior probabilities above 0.9 (right) are given at the nodes. Solid
purple rectangle: areolae margin white; solid grey rectangle: areolae margin black; hollow purple rectangle:
areolae margin jade green pigmented. Solid blue circle gyrophoric acid; hollow blue circle: none.
Revision of Immersaria and a new genus in Lecideaceae 107
is established to accommodate the excluded lecanorine species. e new taxonomic
system, proposed here, revised the classication boundaries between Immersaria and
Lecaimmeria, but it may still be dicult to distinguish between them in cases when
apothecia are absent. In this case, they could be distinguished by the substances
produced or by molecular methods.
Diagnostic traits within species of Immersaria and Lecaimmeria
Species of Immersaria could be identied by their dierent thallus colours
(indicated in Fig. 2). Immersaria ferruginea has a conspicuously greyish-brown
thallus, whereas I. athroocarpa, I. aurantia, I. shangrilaensis and I. venusta have a
reddish-brown thallus. Immersaria athroocarpa (indicated in Fig. 2) is the species
that mostly has a green epihymenium, whereas the other species mostly have a
brown epihymenium. Almost all these species contain conuentic acid, which
is often accompanied by 2’-O-methylmicrophyllinic acid. Planaic acid, which
is newly reported from this genus, is only presented in specimens of Immersaria
aurantia, I. shangrilaensis and I. venusta. All the characters discussed above were
supported by the phylogeny, thus could be used as key characters to dierentiate
species in Immersaria.
Species of the new genus Lecaimmeria could be delimited by the colours of their
areolae and margins, the existence of an apothecial margin and usually by the lack of
substances. e margin of areolae (indicated in Fig. 3) was usually white, but rarely
black or jade green. Lecaimmeria lygaea could be easily distinguished by the black
margin of the areolae. e jade green margin occurs in Lecaimmeria tuberculosa,
which grows on Qilian jade. e areolae margin of Lecaimmeria qinghaiensis is white,
but is occasionally pigmented with very slightly green colour. e margin of the
apothecia is absent in Lecaimmeria tuberculosa and L. iranica, whereas the apothecia
of the other species have white margins. Most species of Lecaimmeria lack secondary
metabolites, while gyrophoric acid was detected only in Lecaimmeria botryoides,
L. iranica and L. mongolica (indicated in Fig. 3). In addition, an orange thallus
appeared only in Lecaimmeria mongolica and L. tibetica, whereas the remaining
species were brownish.
Taxonomy
Immersaria Rambold & Pietschm., Bibliotheca Lichenologica 34: 239 (1989).
Type species. Immersaria athroocarpa (Ach.) Rambold & Pietschm., in Rambold,
Biblioth. Lichenol. 34: 240 (1989).
Description. allus crustose, yellow-brown, red-brown, orange-brown or
brown, sometimes rust coloured, continuous; areolae irregular or tending to
rectangular, with a glossy surface (atrobrunnea-type) caused by a layer of dead,
colourless cells above the upper cortex, areolae sometimes aggregate with black
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
108
prothallus and forming larger areolae; margin pruinose; prothallus distinct at
the margin of thallus or absent. Upper cortex orange pigmented; epinecral layer
colourless; algal layer continuous; medulla lled with grey granules. Apothecia
lecideine, immersed, sometimes aggregate, round or irregular; disc black, at, less
concave, sometimes slightly raised, often poorly developed in section, pruinose
or not; margin reduced. Exciple almost absent, sometimes developed, brown.
Hymenium colourless; paraphyses simple, rarely branched, anastomosing or not;
epihymenium brown, green or brown green, without plectenchyma; subhymenium
colourless, sometimes pale brown; hypothecium pale brown to brown. Asci Porpidia-
type, cylindrical, eight-spored; ascospores ellipsoid, halonate, non-amyloid.
Conidiomata present or not, immersed, linear or stellate, black, margin pruinose;
conidia bacilliform.
Chemistry. allus K–, C–. Medulla I+ violet. Conuentic acid, often accompa-
nied with 2’-O-methylmicophyllinic acid, planaic acid or no substances detected by
TLC. e compound planaic acid is newly reported in this genus.
Ecology and distribution. In China, growing on bare rock, sandstone or granite,
from elevations of 3800 to 4500 m in the alpine zone of west China and elevations of
1200 to 1900 m in the steppe of north China. Worldwide distribution.
Notes. Species with lecanorine apothecia were previously included in
Immersaria (Calatayud and Rambold 1998; Valadbeigi et al. 2011), but the ve-loci
phylogenetic analysis excluded these species from Immersaria. is exclusion entails
a restricted concept of the genus. Immersaria is now dened by its orange-brown,
yellow-brown, sometimes rusty coloured thallus, the amyloid medulla, the glossy
surface of areolae with a pruinose margin, the black immersed lecideine apothecia
with a reduced proper margin, the brown epihymenium and the Porpidia-type asci
with eight halonate and non-amyloid ascospores. e members of this genus occur
in alpine habitats.
Species of Sporastatia A. Massal. might be misidentied as members of Immersaria
because of eld observations of their glossy areolae and the immersed lecideine
apothecia. However, they are characterised by multi-spored asci and their yellow-
brown thallus. Additionally, Miriquidica Hertel & Rambold resembles Immersaria by
its glossy areolae and the lecideine apothecia, but diers in its black brown thallus,
its Lecanora-type asci with non-halonate ascospores and often containing miriquidic
acid. e immersed apothecia of Immersaria may resemble Aspicilia A. Massal. and
Acarospora A. Massal., but Aspicilia has a white or grey thallus, the Aspicilia-type asci
with non-halonate ascospores; Acarospora has multi-spored asci.
Although four known species, Immersaria carbonoidea (J.W. omson) Esnault
& Cl. Roux, I. fuliginosa Fryday, I. olivacea Calat. & Rambold and I. usbekica,
currently lack molecular data, they are temporarily left in Immersaria due to
their morphology which corresponds to that of Immersaria. Our morphological
comparisons were based on high-resolution photographs of type materials and the
original descriptions.
Revision of Immersaria and a new genus in Lecideaceae 109
Immersaria athroocarpa (Ach.) Rambold & Pietschm., in Rambold, Biblioth.
Lichenol. 34: 240 (1989).
Figure 4a–e
Type. S [no locality, no date, no collector], H9508237 (H-Ach-lectotype!–
designated in Hertel 1977). High-resolution photographs seen.
Description. allus areolate, yellow-brown, orange-brown, continuous; areolae
0.2–1.0 mm across, often convex, regular polygons, tends to be squamalose at the
margin, epruinose; margin pruinose; prothallus black, not distinct, sometimes absent.
Upper cortex ca. 32.0 µm thick, yellow-brown; epinecral layer ca. 7.0 µm thick;
algal layer ca. 82.0 µm thick, cells 8.0–10.0 × 7.5 µm in diam., ellipsoid. Apothecia
frequent, densely crowded, immersed, 0.3–1.3 mm in diam.; disc black, rare pruinose,
at, epruinose; margin reduced. Exciple sometimes developed, 25.0–30.0 µm wide,
brown. Hymenium 100–115 µm thick, colourless; paraphyses 1.0–2.0 µm wide,
branched, not anastomosing; epihymenium 20.0–25.0 µm thick, brown, rarely green;
subhymenium ca. 90.0 µm thick, colourless; hypothecium pale brown to brown. Asci
Porpidia-type, cylindrical, eight-spored; ascospores 17.5–20.0 × 10.0 µm, ellipsoid,
halonate. Conidiomata immersed, stellate, black, margin pruinose; conidia 7.5–10.0 ×
1.0 µm in diam., bacilliform.
Chemistry. allus K–, C–. Medulla I+ violet. Chemotype : Conuentic acid.
Chemotype : Unknown substance.
Ecology and distribution. In China, growing on granite in arid and semi-arid
steppe habitats at elevations of 1200–1950 m. Worldwide distribution. is species
is known from Inner Mongolia and Mt. Changbai (Hertel and Zhao 1982) in China.
Notes. e lectotype grows on siliceous rock and contains several intact apothecia.
e materials from Inner Mongolia are identical with the lectotype, based on compari-
sons with high-resolution photographs and the description given by Hertel (1977). It is,
therefore, treated as Immersaria athroocarpa at present. Some Inner Mongolian materials
contain an unknown substance, but form a well-supported clade with other materials.
Immersaria athroocarpa is characterised by the convex, yellow-brown areolae and the large
sizes of ascospores. In this genus, only this species has ascospores up to 20.0 µm long.
Immersaria usbekica is similar to I. athroocarpa in its brown thallus and dense apo-
thecia, but diers in its at areolae, the brown epihymenium and the presence of con-
uentic acid and gyrophoric acid. By comparison with high-resolution photographs
and the original descriptions (Hertel 1977) of Immersaria usbekica, we discovered that
previous reports of this species from China (Zhang et al. 2015) were due to misiden-
tication. It is known from Algeria, Iran, Spain, and the USSR (Barbero et al. 1990).
Specimens examined (SDNU). C. Inner Mongolia: Chifeng City, Balin
Youqi, Hongshilazi, 1403.2 m elev., 44°13'N, 118°44'E, on rock, 2019, Ling Hu et
al. SDNU20190035; Rongshen, Wangfengou, 1217.4 m elev., 44°16'N, 118°22'E,
Ling Hu et al. SDNU20190140, SDNU20190143; Erlinba, 1915.2 m elev., 44°26'N,
118°41'E, Ling Hu et al. SDNU20190227.
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
110
Figure 4. Immersaria athroocarpa (a–e SDNU20190227): a–b thallus c apothecial anatomy d ascus
eascospores. I. aurantia (f–j KUN XY19–1290): f–g thallus h apothecial anatomy i ascus j ascospores.
Scale bars: 1 mm (a–b, f–g); 20 µm (c, h ); 10 µm (d–e, i–j).
Revision of Immersaria and a new genus in Lecideaceae 111
Immersaria aurantia C.M. Xie & Li S. Wang, sp. nov.
MycoBank No: 839738
Figure 4f–j
Etymology. e name “aurantia” refers to the orange thallus.
Type. C. Tibet: Sajia Co., Mula Village, 4752 m elev., 28°40'N, 88°45'E, on
rock, 28 Jun 2019, Xin-Yu Wang et al. XY19–1814 (KUN-holotype).
Description. allus areolate, orange, dark orange, pale orange to pale red-
brown, continuous; areolae 0.7–1.3 mm across, at, epruinose, irregular; margin thin
pruinose; prothallus not seen. Upper cortex 25.0–45.0 µm thick, orange; epinecral
layer (12.0–) 37.0–63.0 µm thick, uneven; algal layer 50.0–93.0 µm thick, cells
5.0–15.0 × 5.0–10.0 µm in diam., round to ellipsoid. Apothecia frequent, scattered,
immersed or isolated from areolae, 0.3–1.3 mm in diam.; disc black, at or concave,
sometimes pruinose; margin reduced. Exciple sometimes developed, ca. 30.0 µm wide,
brown. Hymenium 55.0–83.0 µm thick, colourless; paraphyses 2.0–3.0 µm wide, only
branched and anastomosing at apex; epihymenium ca. 20.0 µm thick, green or green-
brown; subhymenium colourless, not distinct or absent; hypothecium brown. Asci
Porpidia-type, cylindrical, eight-spored; ascospores 8.0–15.0 × 5.0–7.5 µm in diam.,
ellipsoid, halonate. Conidiomata rare, immersed, oblate, black, margin white; conidia
7.5 × 1.0 µm, bacilliform.
Chemistry. allus K–, C–. Medulla I + violet. Chemotype : Conuentic acid,
often accompanied with 2’-O-methylmicrophyllinic acid. Chemotype : Planaic acid.
Chemotype : none (rare).
Ecology and distribution. In China, growing on rock at elevations of 3900–
4300m in the alpine zone. is species is known from Qinghai, Sichuan Province and
Tibet of China.
Notes. Immersaria aurantia is characterised by its distinct orange, irregular areolae
and the mostly green epihymenium. Immersaria athroocarpa and I. venusta are similar
to I. aurantia, but I. athroocarpa diers in the convex, regularly polygonal areolae and
the more crowded apothecia; I. venusta diers in having yellow-brown, often rusty,
cracked areolae and at apothecia. Additionally, conuentic acid and planiaic acid do
not appear simultaneously in Immersaria aurantia, whereas I. venusta always contains
both compounds.
Specimens examined (KUN). C. Qinghai Province: Banma Co., 3933 m
elev., 32°40'N, 100°48'E, on rock, 2020, Li-Song Wang et al. 20–66886, 3932 m
elev., Li-Song Wang et al. 20–66897; Jiuzhi Co., Baiyu Village, 4285 m elev., 33°14'N,
100°58'E, Li-Song Wang et al. 20–67809. Sichuan Province: Rangtang Co., Mt. Haizi,
4223 m elev., 32°20'N, 101°25'E, on rock, 2020, Li-Song Wang et al. 20–66701,
4229 m elev., Li-Song Wang et al. 20–66693, 4217 m elev., Li-Song Wang et al. 20–
66680, 4221 m elev., Li-Song Wang et al. 20–66692. Tibet: Changdu City, Mangkang
Co., Luoni Village, 4145 m elev., 29°56'N, 98°33'E, on rock, 2020, Li-Song Wang
et al. 20–69091, 4138 m elev., Li-Song Wang et al. 20–69091, 20–69094; Gatuo
Town, 29°39'N, 98°35'E, 3831 m elev., Li-Song Wang et al. 20–69114, 3850 m elev.,
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
112
Li-Song Wang et al. 20–69122; Gongga Co., Jiangtang Town, 29°12'N, 90°38'E,
2019.7.23, 4560 m elev., Xin-Yu Wang et al. XY19–1287, 4556 m elev., XY19–1290;
Sajia Co., Mula Village, 28°40'N, 88°45'E, 2019.7.28, 4752 m elev., Xin-Yu Wang
et al. XY19–1814; Angren Co., Kerangla, 29°19'N, 87°01'E, 4530 m elev., Li-Song
Wang et al. 19–63635.
Immersaria ferruginea C.M. Xie & Li S. Wang, sp. nov.
MycoBank No: 839739
Figure 5a–c
Etymology. e name “ferruginea” refers to the rusty brown colour of the thallus.
Type. C. Tibet: Changdu City, Mangkang Co., Quzika Village, 4093 m elev.,
29°15'N, 98°40'E, on rock, 25 Sept 2020, Li-Song Wang et al. 20–69144 (KUN-
holotype).
Description. allus areolate, greyish-brown, continuous; areolae 0.5–1.3mm
across, at, less often convex, rectangular to polygonal, epruinose; margin pruinose;
prothallus black, not distinct. Upper cortex 50.0–68.0 µm thick, brown; epinecral
layer 17.0–40.0 µm thick; algal layer 75.0–78.0 µm thick, cells (4.0–) 7.0–13.0µm
diam., round. Apothecia frequent, densely crowded, immersed, 0.7–1.3 mm
in diam.; disc black, at, pruinose; margin pruinose, slightly raised. Exciple
sometimes developed, 25.0–28.0 µm wide, brown. Hymenium 57.0–100.0µm
thick, colourless; paraphyses 1.0–3.0 µm wide, not branched, anastomosing;
epihymenium 15.0–33.0 µm thick, brown; subhymenium 25.0–63.0 µm thick,
colourless to pale brown, rusty or dark pink; hypothecium pale brown. Asci
Porpidia-type, cylindrical; ascospores rare, 7.5–10.0 × 5.0 µm in diam., ellipsoid,
halonate. Conidiomata not seen.
Chemistry. allus K–, C–. Medulla I+ violet. Conuentic acid, often accompa-
nied with 2’-O-methylmicrophyllinic acid.
Ecology and distribution. In China, growing on quartz sandstone or granite at
elevations of 3800–4300 m in the alpine zone. is species is known from Sichuan
Province and Tibet of China.
Notes. Immersaria ferruginea is characterised by its brown, rusty thallus, its densely
crowded apothecia and its brown epihymenium. e morphology of Immersaria
ferruginea resembles I. carbonoidea, but the latter diers in its dark black-brown thallus
containing norstictic acid and black-brown hypothecium.
Specimens examined (KUN). C. Sichuan Province: Rangtang Co., Mt.
Haizi, 4227 m elev., 32°20'N, 101°25'E, on rock, 2020, Li-Song Wang et al. 20–
66697, 4221 m elev., Li-Song Wang et al. 20–67670. Tibet: Changdu City, Mangkang
Co., Quzika Village, 4093 m elev., 29°15'N, 98°40'E, Li-Song Wang et al. 20–69144,
4101, Li-Song Wang et al. 20–69146, 4122 m elev., Li-Song Wang et al. 20–69148;
Gatuo Town, 3848 m elev., 29°39'N, 98°35'E, Li-Song Wang et al. 20–69105.
Revision of Immersaria and a new genus in Lecideaceae 113
Immersaria shangrilaensis C.M. Xie & Lu L. Zhang, sp. nov.
MycoBank No: 839741
Figure 5d–f
Etymology. e name “shangrilaensis” refers to the location at which the holotype was
collected: “Shangri-La”, a county of Yunnan Province in China.
Type. C. Yunnan Province: Shangri-La County., Mt. Hong Shan, 4363 m elev.,
28°7'N, 99°54'E, on rock, 18 Aug 2018, Li-Song Wang et al. 18–60447 (KUN-holotype).
Description. allus areolate, yellow-brown, orange-brown, often appears as
greyish-brown, generally heavily pruinose, continuous, 5.7–10.0 cm across; areolae
aggregated by 4–14 small areolae (often surrounded by black prothallus), small areolae
up to 0.1 mm across, concave or at, irregular, pruinose; margin pruinose; prothallus
black, distinct. Upper cortex 32.0–50.0 µm thick, yellow-brown granules pigmented;
epinecral layer 15.0–20.0 µm thick; algal layer 47.5–65.0 µm thick, cells 7.5–8.0
× 5.0 µm in diam., ellipsoid. Apothecia frequent, crowded, immersed or isolated
from areolae, 0.3–0.8 mm in diam.; disc black, concave to at, aggregated, cracked
once mature, thin pruinose; margin reduced, slightly raised. Exciple almost absent.
Hymenium 100.0–138.0 µm thick, colourless; paraphyses ca. 2.5 µm wide, branched,
anastomosing or not; epihymenium ca. 15.0 µm thick, brown; subhymenium ca.
55.0 µm thick, colourless; hypothecium pale brown to brown. Asci Porpidia-type,
cylindrical, eight-spored; ascospores 7.0–9.0 × 3.0–4.0 µm, ellipsoid, halonate
(sometimes not distinct). Conidiomata immersed, oblate, black, margin heavily
pruinose; conidia 7.5 × 1.0 µm, bacilliform.
Chemistry. allus K–, C–. Medulla I+ violet. Conuentic acid, planaic acid and/
or 2’-O-methylmicophyllinic acid.
Ecology and distribution. In China, growing on granite at elevations of 4300–
4500 m in the alpine zone. is species is known from Yunnan Province of China.
Notes. e materials of Immersaria athroocarpa from the Shangri-La County of
Yunnan Province are morphologically identical with the specimen Hertel (1977) re-
ported from the same locality, but dier from the lectotype in its aggregate areolae, the
aggregate apothecia and the smaller size of ascospores (7.0–9.0 × 3.0–4.0 µm). Based
on the phenotypic and phylogenetic results, the material from Shangri-La is treated as
a new species, Immersaria shangrilaensis. It is characterised by its large thallus, up to
10.0 cm in diam., the aggregate areolae and apothecia and the small size of ascospores.
Specimens examined. C. Yunnan Province: Shangri-La County, 4350–4500
m elev., on rock, 1915, Handel-Mazzetti no. 6945 = WU-Lichenes0037752 (WU);
Mt. Hong Shan, 4363 m elev., 28°7'N, 99°54'E, on rock, 2018, Li-Song Wang et al.
18–60430 (KUN), Li-Song Wang et al. 18–60447 (KUN) 4503.1 m elev., Chun-
Xiao Wang et al. SDNU20181696 (SDNU), 4361.9 m elev., Chun-Xiao Wang et
al. SDNU20181675 (SDNU); Luquan Co., Mt. Jiaozixueshan, 3800 m elev., 2008,
Hai-Ying Wang SDNU20082253 (SDNU); Lijiang City, Mt. Laojun, 3981 m elev.,
26°37'N, 99°43'E, 2018, Li-Song Wang et al. 18–60555, 18–60602 (KUN).
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
114
Immersaria venusta C.M. Xie & Xin Y. Wang, sp. nov.
MycoBank No: 839742
Figure 6a–d
Etymology. e name “venusta” refers to the beautiful appearance of the thallus.
Type. C. Qinghai Province: Maqing Co., Xueshan Village, 4187 m elev.,
34°37'N, 99°42'E, on rock, 11 Sept 2020, Li-Song Wang et al. 20–67969 (KUN-
holotype).
Figure 5. Immersaria ferruginea (a–c KUN 20–69144): a–b thallus c apothecial anatomy. I. shangrilaen-
sis (d–f KUN 18–60430): d–e thallus f apothecial anatomy. Scale bars: 1 mm (a–b, d–e); 20 µm (c , f ).
Revision of Immersaria and a new genus in Lecideaceae 115
Description. allus areolate, brown, orange-brown, more or less rusty, continuous;
areolae 0.5–1.3 mm across, at or slightly convex, irregular, tending to rectangular,
often cracked, sometimes pruinose; margin pruinose; prothallus not seen. Upper cortex
ca. 38.0 µm thick, yellow brown granules pigmented; epinecral layer ca. 12.0µm thick;
algal layer ca. 128.0 µm thick, cells 5.0–10.0 × 5.0–7.5 µm in diam., round to ellipsoid.
Apothecia frequent, crowded, immersed or isolated from areolae, 0.6–1.0 mm in diam.;
disc black, at, epruinose; margin reduced, sometimes developed. Exciple sometimes
developed, ca. 35.0 µm wide, brown. Hymenium 92.0–113.0 µm thick, colourless;
paraphyses ca. 2.0 µm wide, anastomosing; epihymenium 27.5–30.0µm thick, brown;
subhymenium ca. 62.0 µm thick, colourless; hypothecium brown. Asci Porpidia-type,
cylindrical, eight-spored; ascospores 10.0–12.5 × 5.0–7.5 µm, ellipsoid, halonate.
Conidiomata immersed, linear, black, margin pruinose; conidia not seen.
Chemistry. allus K–, C–. Medulla I+ violet. Chemotype : Conuentic acid,
often accompanied with 2’-O-methylmicrophyllinic acid. Chemotype : Planaic acid.
Chemotype : none (rare).
Ecology and distribution. In China, growing on quartz sandstone or granite at
elevations of 3900–4300 m in the alpine zone. is species is known from Qinghai
Province of China.
Notes. Immersaria venusta is characterised by its yellow-brown, cracked areolae,
its at apothecia and brown epihymenium. It resembles Immersaria shangrilaensis
by its cracked areolae, but its areolae have the tendency to split into several
patches, but not aggregate like those of I. shangrilaensis. Immersaria athroocarpa is
similar to I. venusta in the brown appearance of its thallus and in forming a sister
group in the phylogenetic tree, but it diers in its yellow brown thallus, convex
areolae, densely crowded apothecia and larger ascospores (17.5–20.0 × 10.0 µm).
Immersaria venusta is also similar to I. aurantia (see notes for I. aurantia). e
brown thallus of Immersaria venusta possibly resembles that of I. olivacea, but the
latter diers in its simple or one-septate ascospores, pyriform conidia and dark
bluish-green epihymenium.
Specimens examined (KUN). C. Qinghai Province: Maqing Co., Xueshan
Village, 4187 m elev., 34°37'N, 99°42'E, on rock, 2020, Li-Song Wang et al. 20–
67969, 20–67965; Banma Co., Yaertang Village, 3930 m elev., 32°42'N, 100°42'E,
Li-Song Wang et al. 20–66940. Sichuan Province: Shiqu Co., Xinrong Village, 4043
m elev., 32°59'N, 98°19'E, on rock, 2020, Li-Song Wang et al. 20–68802; Rangtang
Co., Mt. Haizi, 4246 m elev., 32°21'N, 101°24'E, Li-Song Wang et al. 20–66721,
20–66725.
Selected additional comparative material was examined.
Bellemerea alpina (Sommerf.) Clauzade & Cl. Roux R, Lps. Petsamo, Pummangin
vuonon N-puoli, 1938, Räsänen, V., H9503269 (H); Lps. Petsamo, inter Vaitolahti
et Kervanto, 1938, Räsänen, V., H9503270 (H).
Bellemerea cinereorufescens (Ach.) Clauzade & Cl. Roux F, Ob. Simo.
Anteroinen. Rantakivellä, 1920, Räsänen, V., H9503267 (H); Le. Enontekiö,
Kirkonkylä, 1925, Kari, L.E., H9503268 (H).
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
116
High-resolution photographs seen.
Immersaria carbonoidea (J.W. omson) Esnault & Cl. Roux USA, Alaska, along the
Pitmegea River, 15 miles upstream from Cape Sabine, 1958, omson, J.W.,
M0082171 (M-isotype!), G00126754 (G-isotype!).
Immersaria olivacea Calat. & Rambold S, Espana, Castelló: Benicàssim, Parreta
Alta, 390 m elev., 1993, Calatayud, V., M0101779 (M-isotype!).
Immersaria usbekica (Hertel) M. Barbero, Nav.-Ros. & Cl. Roux A Algerie-Atlas
Tellieu, Larba, Piste de Bougara á Tablat au S-E de l’arboretum de Meindja, 1985,
Esnault, J., M0101787 (M-paratype!).
Lecaimmeria C.M. Xie, Lu L. Zhang & Li S. Wang, gen. nov.
MycoBank No: 839743
Etymology. e name “Lecaimmeria” refers to the immersed lecanorine apothecia of
all known species.
Type species. Lecaimmeria orbicularis C.M. Xie & Lu L. Zhang, sp. nov.
Description. allus crustose, red-brown, orange-brown or dark brown,
continuous or not; areolae irregular or tending to rectangular, with a glossy surface
(atrobrunnea-type) caused by a layer of dead, colourless cells above the upper cortex;
margin white or black; prothallus distinct at the margin of thallus or absent, sometimes
developed between areolae. Upper cortex orange; epinecral layer colourless; algal layer
continuous; medulla lled with grey granules. Apothecia lecanorine, immersed, round
or irregular; disc red-brown, dark red-brown or dark orange-brown, at or concave;
margin present or absent, black or white, rarely green, pruinose or not. Exciple reduced,
tissue at the lateral sides of the hymenium corresponding to the upper cortex and the
algal layer of the vegetative areolae and to hypothecial hyphal cells when apothecia
reach the margin of the areole (indicated in Figs. 7c, g, 8c, 9c, h, 10c, h). Hymenium
colourless; paraphyses simple, rarely branched, anastomosing or not; epihymeinum
orange, orange-brown, rarely brown, with a plectenchyma. Asci Porpidia-type
(indicated in Fig 9d), cylindrical, eight-spored; ascospores ellipsoid, halonate, non-
amyloid. Conidiomata present or absent, immersed, rarely convex, linear or stellate,
rarely tuberculiform; conidia bacilliform.
Chemistry. allus K–, C+/–. Medulla I+ violet. Gyrophoric acid, 4-O-demethyl-
planaic acid or no substances detected by TLC.
Ecology and distribution. In China, growing on rock, sandstone, granite or
Qilian jade (rare), from elevations of 3100 to 4800 m in the alpine zone of west China
and from 1200 to 1900 m in the steppe of north China. is genus is known from
China, Europe, Iran, Mongolia, Romania, Russia, and USA.
Notes. e ve-loci phylogenetic analysis showed that the species with lecanorine
apothecia formed a novel lineage and should be excluded from Immersaria; thus, they
are here treated as a new genus Lecaimmeria. Lecaimmeria is distinguished from related
genera by its glossy surface, orange or red-brown areolae with margins, the amyloid
Revision of Immersaria and a new genus in Lecideaceae 117
medulla, the red-brown immersed lecanorine apothecia, the orange epihymenium
with a plectenchyma and the Porpidia-type asci with eight halonate and non-amyloid
ascospores. In China, the genus is distributed in alpine areas, high altitude desert-
steppe areas or high latitude steppe. Almost all the species of Lecaimmeria grow on
granite or sandstone, with the exception of one species, L. tuberculosa, which grows on
jade. Interestingly, the margin of conidiomata and areolae of Lecaimmeria tuberculosa
appear with heavily jade-green pruinose.
e immersed apothecia and brown thallus of Lecaimmeria often resemble those
of Immersaria, but Lecaimmeria diers in its red-brown lecanorine apothecia, often
with a white margin, their orange epihymenium with a plectenchyma and the thallus
containing gyrophoric acid. is genus might be confused with Bellemerea by its
lecanorine apothecia and the Porpidia-type asci with halonate ascospores, but the latter
genus diers in its amyloid ascospores.
ree species, previously included in Immersaria, I. cupreoatra, I. iranica and I.
mehadiana, have lecanorine apothecia, but two of these, I. cupreoatra and I. mehadiana,
currently lack molecular sequences. We suggest that these three species should be
transferred to Lecaimmeria, based on the following factors. eir morphology is consistent
with Lecaimmeria according to molecular results and comparisons with type specimens,
high-resolution photographs of the type materials and the original descriptions. One
unknown “Immersaria” species from Macedonia is sister to Lecaimmeria iranica in
Figure 6. Immersaria venusta (a–d KUN 20–66725): a–b thallus c apothecial anatomy d ascus. Scale
bars: 1 mm (a–b); 20 µm (c); 10 µm (d).
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
118
the phylogenetic tree (Fig. 3), but comparison with high-resolution photograph and
previous records (Malíček and Mayrhofer 2017) show that it diers in its black margin
of areolae and absence of gyrophoric acid. is unknown species with lecanorine
apothecia is possibly a member of Lecaimmeria, but descriptions are lacking and the
specimens were not seen. us, this species is temporarily retained in Immersaria.
Lecaimmeria botryoides C.M. Xie & Li S. Wang, sp. nov.
MycoBank No: 839744
Figure 7a–d
Etymology. e name “botryoides” refers to the crowded apothecia while immature.
Type. C. Sichuan Province: Aba City, Rangtang County, Haizi Mt., 4225m
elev., 32°21'N, 101°24'E, on rock, 6 Sept 2020, Li-Song Wang et al. 20–66730
(KUN-holotype).
Description. allus areolate, red-brown, discontinuous; areolae 0.2–1.0 mm
across, at, slightly concave or convex, pruinose, polygonal, tending to be rectangular,
margins heavily pruinose. Prothallus black, distinct in the margin of thallus. Upper
cortex 20.0–25.0 µm thick, brown; epinecral layer 22.0–48.0 µm thick; algal layer
ca. 37.0 µm thick, cells 7.5–10.0 µm diam., round. Apothecia frequent, irregular,
densely crowded while immature (3–6/areolae), aggregate once mature, immersed,
0.2–1.3mm in diam.; disc red-brown, at, or concave, epruinose; margin pruinose,
slightly raised. Hymenium 67.0–100.0 (–155.0) µm thick, colourless; paraphyses ca.
2.0 µm wide, simple, only branched at the top, not anastomosing; epihymenium 25.0–
30.0 µm thick, orange; plectenchyma 2.0–8.0 µm thick; subhymenium 17.0–38.0 µm
thick, colourless; hypothecium pale brown to brown. Asci Porpidia-type, cylindrical,
eight-spored; ascospores 7.5–8.0 × 4.0–6.0 µm in diam., ellipsoid, halonate. Conidi-
omata not seen; conidia not seen.
Chemistry. allus K–, C+/–. Medulla I+ violet. Chemotype : Gyrophoric acid.
Chemotype : none.
Ecology and distribution. In China, growing on rock at elevations of 3100–4300 m
in the alpine zone. is species is known from Qinghai and Sichuan Provinces of China.
Notes. Lecaimmeria botryoides is characterised by its discontinuous thallus, densely
crowded apothecia while immature and the orange epihymenium. Lecaimmeria or-
bicularis is similar to L. botryoides in its red-brown thallus, but diers in its round, at
apothecia and continuous thallus. e red-brown thallus of Lecaimmeria botryoides
resembles L. cupreoatra, but the latter diers in the black margin of its apothecia and
its distinct black prothallus between areolae.
Specimens examined (KUN). C. Qinghai Province: Banma Co., 3958 m
elev., 32°40'N, 100°48'E, on rock, 2020, Li-Song Wang et al. 20–66900, 3932 m elev.,
Li-Song Wang et al. 20–66898, 3935 m elev., Li-Song Wang et al. 20–66891, 3178
m elev., Li-Song Wang et al. 20–66765. Sichuan Province: Rangtang Co., Mt. Haizi,
Revision of Immersaria and a new genus in Lecideaceae 119
4256m elev., 32°21'N, 101°24'E, on rock, 2020, Li-Song Wang et al. 20–66721,
4300 m elev., Li-Song Wang et al. 20–67706, 4276 m elev., Li-Song Wang et al.
20–66706, 4255m elev., Li-Song Wang et al. 20–66707, 4274 m elev., Li-Song Wang
et al. 20–66713, 4274 m elev., Li-Song Wang et al. 20–66711, 20–66705, 4225 m
elev., Li-Song Wang et al. 20–66730, 4220 m elev., 32°20'N, 101°25'E, Li-Song Wang
et al. 20–66683.
Lecaimmeria cupreoatra (Nyl.) C.M. Xie, comb. nov.
MycoBank No: 839745
Basionyms. Lecanora cupreoatra Nyl., Lichens Lapponiae orientalis: 181 (1866).
Type. R. “Medvæschiigora, ad Onegam”, 13 June 1863, . Simming,
H9508237 (H-lectotype!).
Description. Nylander (1866) and Clauzade and Roux (1985).
Notes. e lectotype grows on siliceous rock and contains several intact apothecia.
As “Immersaria cupreoatra has lecanorine apothecia and is related to I. lygeae in our
phylogeny, it is, therefore, transferred to Lecaimmeria. is species has not been cor-
rectly recorded in China (see notes for Lecaimmeria mongolica). e species is known
from Europe, Mongolia, Russia, and USA (Calatayud and Rambold 1998).
Specimens examined (H). R. Kl. Kurkijoki, Kuuppala, Himohirsi, 12 May
1934, Räsänen, V., H9503417, H9510194.
Lecaimmeria iranica (Valadb., Sipman & Rambold) C.M. Xie, comb. nov.
MycoBank No: 839746
Basionyms. Immersaria iranica Valadb., Sipman & Rambold, Lichenologist 43(3):
204 (2011).
Type. I. Mazandaran, Haraz Road, 20 km to Aamol, 36°17'N, 52°21'E, on
calcareous rock, 1475 m, 7 Apr 2006, T. Valadbeigi 9008 (TARI-holotype; B, hb.
Valadbeigi-isotype). Not seen.
Description. Valadbeigi et al. (2011).
Notes.Immersaria iranica has lecanorine apothecia, a distinct epinecral layer and
halonate ascospores (Valadbeigi et al. 2011). e materials from China are in accordance
with the materials of Iran, based on comparisons with the original descriptions and
the photographs given by Valadbeigi et al. (2011). e characters of this species are
consistent with the new genus and the phylogenetic results showed that it was clustered
with species of Lecaimmeria. erefore, it was transferred to Lecaimmeria. is species
is currently known from Iran and China.
Specimens examined (SDNU). C. Xinjiang: Urumqi, Mt. Tianshan-glacier
No.1, alt. 3800 m, on rock, 2011, Z.L. Huang SDNU20126106, SDNU20129049.
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
120
Figure 7. Lecaimmeria botryoides (a–d KUN 20–66730): a–b thallus c apothecial anatomy. L. lyagea
(e–h KUN 20–69070): e–f thallus g apothecial anatomy h scospores. Scale bars: 1 mm (a–b, e–f);
20µm (c, g ); 10 µm (d , h ).
Revision of Immersaria and a new genus in Lecideaceae 121
Lecaimmeria lygaea C.M. Xie & Lu L. Zhang, sp. nov.
MycoBank No: 839747
Figure 7e–h
Etymology. e name “lygaea” refers to the dark appearance of the thallus.
Type. C. Tibet: Changdu City, Mangkang County, Luoni Village, 4127m
elev., 29°56'N, 98°33'E, on rock, 24 Sept 2020, Li-Song Wang et al. 20–69072
(KUN-holotype).
Description. allus areolate, dark red-brown, dark brown, continuous; areolae
0.5–1.0 mm across, at, epruinose, irregular pentagonal, sometimes rectangular,
ssures between areolae often lled with black prothallus; margin black, thinly
pruinose; prothallus black, developed between areolae, also distinct in the margin.
Upper cortex ca. 20.0 µm thick, orange-brown; epinecral layer ca. 15.0 µm thick;
algal layer ca. 50.0 µm thick, cells 7.0–13.0 µm in diam., round. Apothecia frequent,
round, crowded, immersed, 0.2–0.8 mm in diam.; disc red-brown, at, or concave,
epruinose; margin black, moderately thick, pruinose, raised. Hymenium 75.0–
93.0 µm thick, colourless; paraphyses ca. 2.0 µm wide, simple, unbranched, not
anastomosing; epihymenium 25.0–38.0 µm thick, orange brown; plectenchyma ca.
7.0 µm thick; subhymenium 20.0–25.0 µm thick, colourless; hypothecium brown
Asci Porpidia-type, cylindrical, eight-spored; ascospores 12.5–20.0 × 5.0–7.5 µm in
diam., ellipsoid, halonate. Conidiomata immersed, stellate, black, margin pruinose;
conidia 5.0 × 1.0 µm, bacilliform.
Chemistry. allus K–, C–. Medulla I+ violet. Unknown fatty acid by TLC.
Ecology and distribution. In China, growing on sandstone at elevations of
4000–4200 m in the alpine zone. is species is known from the Tibet Region of
China.
Notes. Lecaimmeria lygaea is characterised by its dark brown thallus, black
margin of its areolae, black prothallus which lls the ssures between areolae, dark
orange apothecia and its orange brown epihymenium. Lecaimmeria cupreoatra and
L.mehadiana are similar to L. lygaea, but L. cupreoatra has a discontinuous thallus,
with each areola surrounded by black prothallus, dark red-brown to black-brown
apothecia without a margin. Lecaimmeria mehadiana has areolae with a white margin,
black-brown apothecia, brown epihymenium and contains 4-O-demethylplanaic
acid. e phylogenetic results show that Lecaimmeria tibetica is the sister species to
L. lygaea. ey are similar in chemistry, but dier in its orange-brown thallus and
dark orange brown apothecia.
Specimens examined (KUN). C. Tibet: Changdu City, Mangkang Co.,
Luoni Village, 4099 m elev., 29°56'N, 98°33'E, on rock, 2020, Li-Song Wang et al.
20–69054, 4131 m elev., Li-Song Wang et al. 20–69070, 4127 m elev., Li-Song Wang
et al. 20–69072, 4095 m elev., Li-Song Wang et al.20–69053.
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
122
Lecaimmeria mehadiana (Calatayud & Rambold) C.M. Xie, comb. nov.
MycoBank No: 839748
Basionyms. Immersaria mehadiana Calat & Rambold, Lichenologist 30(3): 233 (1998).
Type. R. Caras-Severin Comitat, Mehadía, Strájot Mtn., on rock, 1994,
Rambold, G.W., M0101781 (M-holotype!), M0101780, M0101782, M0101783
(M-isotype!). High-resolution photographs seen.
Description. Calatayud and Rambold (1998).
Notes. As “Immersaria mehadiana has lecanorine apothecia and resembles
L.lygaea and L. cupreoatra, having a dark brown thallus, it is, therefore, transferred to
Lecaimmeria. is species is characterised by its greyish prothallus, dark brown apo-
thecia and the brown epihymenium. It is only known from Romania (Calatayud and
Rambold 1998).
Lecaimmeria mongolica C.M. Xie & Lu L. Zhang, sp. nov.
MycoBank No: 839749
Figure 8a–d
Etymology. e name “mongolica” refers to the collection of the holotype within
Inner Mongolia, an autonomous region of China.
Type. C. Inner Mongolia: Chifeng City, Balinyouqi, Han Mountain, 1445m
elev., 44°11'N, 118°44'E, on rock, 22 Jul 2019, Zun-Tian Zhao et al. SDNU20190354
(SDNU-holotype).
Description. allus areolate, orange, continuous; areolae 0.4–0.8 mm across, epru-
inose, neatly arranged, irregular, tending to be rectangular, margin pruinose; prothallus
black, not distinct. Upper cortex ca. 20.0 µm thick, brown; epinecral layer 5.0–8.0 µm
thick; algal layer ca. 87.0 µm thick, cells 7.5–12.5 µm diam., round. Apothecia frequent,
crowded, immersed or isolated from areolae, 0.2–0.8 mm in diam.; disc red-brown,
at, slightly convex, epruinose; margin pruinose. Hymenium 62.0–83.0 µm thick, col-
ourless; paraphyses ca. 2.0 µm wide, unbranched, not anastomosing; epihymenium ca.
42.0 µm thick, orange; plectenchyma 5.0–10.0 µm thick; subhymenium 30.0–38.0 µm
thick, colourless; hypothecium brown. Asci Porpidia-type, cylindrical, eight-spored; as-
cospores 10.0–17.5 × 6.0–7.5 µm in diam., ellipsoid, halonate. Conidiomata immersed,
oblate, rare ellipsoid, black, margin pruinose; conidia 5.0 × 1.0 µm, bacilliform.
Chemistry. allus K–, C+. Medulla I+ violet. Gyrophoric acid.
Ecology and distribution. In China, growing on granite at elevations of 1400–
2000 m in steppe or mountains. is species is known from Inner Mongolia of China.
Notes. is species was once reported as “Immersaria cupreoatra from China
(Zhang et al. 2015), but after comparing our collections with the type material, this was
found to be a misclassication. Additionally, the phylogenetic results showed that these
collections formed a well-supported lineage belonging to Lecaimmeria. erefore, it is
here treated as a new species, Lecaimmeria mongolica, characterised by its orange-brown
thallus, red-brown apothecia with a distinct white margin and the thallus containing
Revision of Immersaria and a new genus in Lecideaceae 123
gyrophoric acid. Lecaimmeria cupreoatra resembles L. mongolica by containing
gyrophoric acid, but it diers in its dark black brown thallus and the black margin of its
apothecia. Lecaimmeria tibetica is similar to L. mongolica in its orange thallus, but diers
in its smaller, dark orange apothecia and that no substance can be detected by TLC.
Specimens examined (SDNU). C. Inner Mongolia: Chifeng City, Balin
Youqi, Mt. Qingyangcheng, 1445 m elev., 43°35'N, 117°30'E, on rock, 2019, Zun-
Tian Zhao et al. SDNU20190350; Han Shan, 1563 m elev., 44°11'N, 118°44'E, on
rock, Zun-Tian Zhao et al. SDNU20190354; A’ershan City, Mt. Jiguan, 1500 m
elev., on rock, 2011, Yu-Liang Cheng SDNU20124912, 1400 m elev., Dai-Feng Jiang
SDNU20124859; Ke Qi, Huanggangliang, 2000 m elev., on rock, Pan-Meng Wang
SDNU20117613, Xing-Ran Kou SDNU20117399.
Lecaimmeria orbicularis C.M. Xie & Lu L. Zhang, sp. nov.
MycoBank No: 839750
Figure 9a–e
Etymology. e name “orbicularis” refers to the round shape of the apothecia.
Type. C. Sichuan Province: Rangtang Co., Gangmuda Village, 3800 m elev.,
32°18'N, 101°3'E, on rock, 7 Sept 2020, Li-Song Wang et al. 20–66753 (KUN-holotype).
Figure 8. Lecaimmeria mongolica (a–d SDNU20190354): a–b thallus c apothecial anatomy dascospores.
Scale bars: 1 mm (a–b); 20 µm (c); 10 µm (d).
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
124
Description. allus areolate, red-brown, rarely orange-brown, continuous;
areolae 0.2–1.0 mm across, at, occasionally wrinkled, tending to rectangular, ssures
between areolae often lled with black prothallus, margin pruinose; prothallus black,
developed between areolae, also distinct in the margin. Upper cortex 42.0–58.0µm
thick, brown; epinecral layer 5.0–20.0 µm thick; algal layer 70.0–113.0µm thick,
cells 10.0–15.0 × 7.5–10.0 µm in diam., ellipsoid to round. Apothecia frequent,
scattered, immersed or isolated from areolae, 0.5–1.3 mm in diam.; disc red-brown,
at, round, epruinose; margin white, slightly raised. Hymenium 75.0–113.0 µm
thick, colourless; paraphyses ca. 2.0 µm wide, simple, unbranched, not anastomosing;
epihymenium 17.5–30.0 µm thick, orange; plectenchyma 5.0–15.0 µm thick;
subhymenium 30.0–63.0 µm thick, colourless; hypothecium brown. Asci Porpidia-
type, cylindrical, eight-spored; ascospores 12.5–15.0 × 5.0–6.0 µm, ellipsoid,
halonate. Conidiomata not seen.
Chemistry. allus K–, C–. Medulla I+ violet. None.
Ecology and distribution. In China, growing on granite or sandstone at eleva-
tions of 3700–4200 m in the alpine zone. is species is known from Qinghai and
Sichuan Provinces of China.
Notes. Lecaimmeria orbicularis is characterised by its orange brown thallus, neatly
arranged areolae and round, at apothecia. Lecaimmeria botryoides is similar to L. or-
bicularis (see notes for L. botryoides). Lecaimmeria mongolica might be confused with
L.orbicularis due to its large apothecia with a white margin, but diers in its red-brown
thallus and distribution in steppes. e red-brown thallus of Lecaimmeria cupreoatra
resembles that of L. orbicularis, but diers in the black margin of its apothecia and its
distinct black prothallus between areolae.
Specimens examined (KUN). C. Qinghai Province: Jiuzhi Co., Nianbaoy-
uze, 4200 m elev., 33°14'N, 100°58'E, on rock, 2020, Li-Song Wang et al. 20–66811,
20–66829, 20–66801, 20–66826A, 20–66821, 20–66805, 20–66833, 20–66817,
20–66841; Banma Co., Nianbaoyuze, 3930 m elev., 32°40'N, 100°48'E, Li-Song
Wang et al. 20–66909, 20–66908, 20–66896, 20–66886B, 20–66899, 20–66935,
20–66943; Zhiqingsongduo Town, 3712 m elev., 33°24'N, 101°25'E, Li-Song Wang
et al. 20–66965; Suohurima Village, 4029 m elev., 33°23'N, 100°57'E, Li-Song Wang
et al. 20–66979. Sichuan Province: Rangtang Co., Gangmuda Village, 3800 m elev.,
32°18'N, 101°3'E, on rock, 2020, Li-Song Wang et al. 20–66753, 20–66750, 3793 m
elev., Li-Song Wang et al. 20–66747, Shangrangtang Village, 3730 m elev., 32°16'N,
101°21'E, Li-Song Wang et al. 20–66743.
Lecaimmeria qinghaiensis C.M. Xie & Li S. Wang, sp. nov.
MycoBank No: 839751
Figure 9f–i
Etymology. e name “qinghaiensis” refers to the location in which the holotype was
collected, in “Qinghai”, a province of China.
Revision of Immersaria and a new genus in Lecideaceae 125
Figure 9. Lecaimmeria orbicularis (a–e KUN 20–66753): a–b thallus c apothecial anatomy d ascus (Lu-
gol’s iodine) e ascospores. L. qinghaiensis (f–i KUN 20–68696): f–g thallus h apothecial anatomy iascus.
Scale bars: 1 mm (a–b, f–g); 20 µm (c, h ); 10 µm (d , e , i ).
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
126
Type. C. Qinghai Province: Yushu City, Zaduo County, Sahuteng Town,
4634 m elev., 32°55'N, 95°46'E, on rock, 20 Sept 2020, Li-Song Wang et al. 20–
68698 (KUN-holotype).
Description. allus areolate, yellow-brown, rusty, continuous; areolae 0.5–
1.5 mm across, at, epruinose; margin pruinose, occasionally green pigmented;
prothallus black, distinct at the margin. Upper cortex 27.0–38.0 µm thick, brown;
epinecral layer 12.0–20.0 µm thick; algal layer 57.0–93.0 µm thick, cells 7.5–12.5 ×
5.0–12.5 µm in diam., ellipsoid to round. Apothecia frequent, immersed or isolated
from areolae, round or somewhat irregular while immature, ellipsoid and tending to
be rectangular or occupying the whole areolae once mature, 0.2–1.3 mm in diam.;
disc brown, dark red-brown, at, occasionally with a ssure when mature, epruinose;
margin white, slightly raised. Hymenium 52.0–63.0 µm thick, colourless; paraphyses
2.0–3.0 µm wide, unbranched, not anastomosing; epihymenium 25.0–30.0 µm thick,
dark orange-brown; plectenchyma 7.0–18.0 µm thick; subhymenium 50.0–63.0µm
thick, colourless; hypothecium brown. Asci Porpidia-type, cylindrical, eight-spored;
ascospores 8.0–15.0 × 5.0–7.5 µm in diam., ellipsoid, not distinctly halonate.
Conidiomata rare, immersed, at, slightly convex, liner, stellate, graphidoid once
mature, black, margin pruinose; conidia not seen.
Chemistry. allus K–, C–. Medulla I+ violet. None.
Ecology and distribution. In China, growing on rock at elevations of 4600–
4900m in the alpine zone. is species is known from Qinghai Province of China.
Notes. Lecaimmeria qinghaiensis is characterised by the yellow-brown, rusty thal-
lus, the red-brown apothecia often occupying the whole areolae at maturity and the
dark orange-brown epihymenium. e phylogenetic results showed that Lecaimmeria
tuberculosa is the sister species to L. qinghaiensis which is similar in the appearance
of the thallus, but diers in the brown, never rusty thallus, the red-brown apothecia
and the green, tuberculiform conidiomata. e red-brown apothecia of Lecaimmeria
qinghaiensis resembles L. iranica, but diers in the rusty thallus and the white margin
of the apothecia.
Specimens examined (KUN). C. Qinghai Province: Zaduo Co., Sahuteng
Town, 4634 m elev., 32°55'N, 95°46'E, on rock, 2020, Li-Song Wang et al. 20–68698,
4637 m elev., Li-Song Wang et al. 20–68687, 4622 m elev., Li-Song Wang et al. 20–
68696; 4790 m elev., 33°31'N, 95°8'E, Xin-Yu Wang et al. 20–3115, 4791 m elev.,
Xin-Yu Wang et al. 20–3127; Zaqing Village, 4815 m elev., Xin-Yu Wang et al. 20–849.
Lecaimmeria tibetica C.M. Xie & Xin Y. Wang, sp. nov.
MycoBank No: 839752
Figure 10a–e
Etymology. e name “tibetica” refers to the location from which the holotype was
collected: “Tibet”, an autonomous region of China.
Type. C. Tibet: Gongga Co., Jiangtang Town, 4557 m elev., 29°12'N,
90°38'E, on rock, 9 Sept 2019, Xin-Yu Wang et al. XY19–1291 (KUN-holotype).
Revision of Immersaria and a new genus in Lecideaceae 127
Description. allus areolate, orange-brown, epruinose; areolae 0.3–0.5 mm
across, irregular, upper surface uneven, margin lacking, pruinose; prothallus black,
distinct at the margin. Upper cortex 17.0–33.0 µm thick, brown; epinecral layer
10.0–20.0 µm thick; algal layer ca. 75.0 µm thick, cells 7.0–10.0 diam., round.
Apothecia rare, immersed or isolated from areolae, 0.2–0.5 mm in diam.; disc dark
orange-brown, epruinose, at, slightly convex; margin pruinose. Hymenium 105.0–
138.0 µm thick, colourless; paraphyses ca. 2.0 µm wide, unbranched, not anasto-
mosing; epihymenium ca. 25.0 µm thick, orange; plectenchyma ca. 12.0 µm thick;
subhymenium almost absent, colourless; hypothecium brown. Asci Porpidia-type,
cylindrical, eight-spored; ascospores 12.5–15.0 × 5.0–6.0 µm, ellipsoid, halonate.
Conidiomata immersed, oblate, black, margin pruinose; conidia 5.0 × 1.5–2.0 µm
in diam., bacilliform.
Chemistry. allus K–, C–. Medulla I+ violet. None.
Ecology and distribution. In China, growing on quartz sandstone at elevations of
4300–4600 m in the alpine zone. is species is known from Tibet, China.
Notes. Lecaimmeria tibetica is characterised by the orange-brown thallus, the black
pigmentation of the areolae margin and the dark orange-brown and small size of the
apothecia. Lecaimmeria tibetica is similar to L. mongolica (see notes for L. mongolica).
e red-brown apothecia of Lecaimmeria cupreoatra resembles L. tibetica, but that spe-
cies diers in its dark red-brown thallus and the presence of gyrophoric acid.
Specimens examined (KUN). C. Tibet: Gongga Co., Jiangtang Town,
4583 m elev., 29°12'N, 90°38'E, on rock, 2019, Xin-Yu Wang et al. XY19–1288,
4557 m elev., XY19–1291, 4560 m elev., XY19–1280; Dingri Co., Zhaguozhong,
4310 m elev., 28°35'N, 86°53'E, Li-Song Wang et al. 19–64071.
Lecaimmeria tuberculosa C.M. Xie & Xin Y. Wang, sp. nov.
MycoBank No: 839754
Figure 10f–i
Etymology. e name “tuberculosa” refers to the tuberculiform conidiomata.
Type. C. Gansu Province: Zhangye City, Sunan Co., Along the way from
Sunan to Qilian, 3928 m elev., 38°37'N, 99°28'E, on rock, 30 May 2018, Li-Song
Wang et al. 18–58865 (KUN-holotype).
Description. allus areolate, red-brown, continuous; areolae 0.5–1.3 mm across,
slightly convex, epruinose; margin pruinose, often jade-green pigmented; prothallus
not distinct. Upper cortex ca. 27.0 µm thick, orange; epinecral layer up to 28.0 µm
thick, uneven, sometimes absent; algal layer ca. 50.0 µm thick, cells 6.0–10.0 µm
diam., round. Apothecia frequent, scattered, immersed, 0.3–0.6 mm in diam.; disc red-
brown, concave, epruinose; margin absent. Hymenium 55.0–83.0 µm thick, colourless;
paraphyses ca. 2.0 µm wide, unbranched, not anastomosing; epihymenium 15.0–
30.0µm thick, orange; plectenchyma ca. 5.0 µm thick, discontinuous; subhymenium
ca. 38.0 µm thick, colourless; hypothecium brown. Asci Porpidia-type, cylindrical,
eight-spored; ascospores 6.0–12.5 × 3.0–5.0 µm in diam., ellipsoid, halonate.
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
128
Figure 10. Lecaimmeria tibetica (a–e KUN XY19–1288): a–b thallus c apothecial anatomy d ascus
e ascospore. L. tuberculosa (f–i KUN 18–58864): f thallus g conidiomata h apothecial anatomy
iascospores. Scale bars: 1 mm (a–b, f–g); 20 µm (c); 50 µm (h); 10 µm (d , e, i).
Revision of Immersaria and a new genus in Lecideaceae 129
Conidiomata stellate, strongly convex, rarely immersed, forming tuberculiform, black,
margin pruinose, jade-green pigmented; conidia 3.0–4.5 × 1.0 µm in diam., oblong
to bacilliform.
Chemistry. allus K–, C–. Medulla I+ violet. No substances by TLC.
Ecology and distribution. In China, growing on the Qilian jade or sandstone at
elevations of 3900–4200 m in the alpine zone. is species is known from Qinghai
Province and Gansu Province of China.
Notes. Lecaimmeria tuberculosa is characterised by its red-brown thallus, the jade-
green pruinose colour at the margin of its areolae, its red-brown, concave apothecia
without a proper margin and tuberculiform conidiomata. Lecaimmeria qinghaiensis is
similar to L. tuberculosa (see notes for L. qinghaiensis). Lecaimmeria tuberculosa usually
grows on jade and, interestingly, the margin of the conidiomata and areolae of the
species are heavily jade-green pigmented. Lecaimmeria iranica resembles L. tuberculosa
by the absence of an apothecial margin, but diers in its immersed conidiomata and
the white margin of its areolae.
Specimens examined (KUN). C. Qinghai Province: Gande Co., Qingzhen
Village, 4124 m elev., 34°11'N, 100°12'E, on rock, 2020, Li-Song Wang et al. 20–
68077, 4145 m elev., Li-Song Wang et al. 20–68055. Gansu Province: Zhangye City,
Sunan Co., along the way from Sunan to Qilian, 3928 m elev., 38°37'N, 99°28'E, on
rock, 2018, Li-Song Wang et al. 18–58856, 18–58857, 18–58865, 18–59835.
Key to species of Immersaria in China
1 allus greyish-brown; apothecia crowded ................................I. ferruginea
allus reddish-brown; apothecia rarely crowded ........................................2
2 allus orange; areolae irregular .................................................. I. aurantia
allus not orange; areolae irregular, polygonal or rectangular .................... 3
3 allus large, 6–10 cm across; areolae aggregated by several smaller areolae
and black prothallus ............................................................I. shangrilaensis
allus smaller, 2–5 cm across; areolae not aggregated ................................ 4
4 allus areolae convex, not rusty, not cracked; ascospores over 15 µm long ..
..............................................................................................I. athroocarpa
allus areolae at, often rusty, cracked; ascospores never over 15 µm long ..
..................................................................................................... I. venusta
Key to species of Lecaimmeria in China
1 Prothallus distinct and lling the ssures between areolae ...............L. lygaea
Prothallus only distinct at the margin .........................................................2
2 allus orange ............................................................................................3
allus reddish-brown ................................................................................4
3 Apothecia red-brown; containing gyrophoric acid .................... L. mongolica
Apothecia dark orange; no substance detected by TLC ................ L. tibetica
Cong-Miao Xie et al. / MycoKeys 87: 99–132 (2022)
130
4 Apothecia margin absent ............................................................................. 5
Apothecia margin present ...........................................................................6
5 Areolae margin white; epihymenium brown; pycindia immersed, linear or
stellate...........................................................................................L. iranica
Areolae margin green; epihymenium orange; pycindia convex, tuberculi-
form ......................................................................................L. tuberculosa
6 allus rusty; apothecia occupy the whole areolae .................L. qinghaiensis
allus not rusty; apothecia do not occupy the areolae ...............................7
7 Apothecia irregular, crowded while immature, aggregate when mature ..........
................................................................................................ L. botryoides
Apothecia round, rarely crowded, not aggregate ......................L. orbicularis
Acknowledgements
We are very grateful to Dr. Saara Velmala and Dr. Leena Myllys from the University
of Helsinki (H) and Dr. Water Till from the Universität Wien (WU) for the loan of
type materials, to Dr. Fiona Ruth Worthy, a post-doctoral researcher from Kunming
Institute of Botany, Centre for Mountain Ecosystem Studies, for carefully correcting
our English and to Dr. Einar Timdal (Natural History Museum, Norway) and Dr.
Alan Fryday (Michigan State University) for the very careful review and the useful
comments on the manuscript. is study was supported by the National Natural
Science Foundation of China (31750001, 31970022, 32170002), the Second Tibetan
Plateau Scientic Expedition and Research Program (STEP) (2019QZKK0503), the
Youth Innovation Promotion Association CAS (2020388) and the State Key Laboratory
of Phytochemistry and Plant Resources in West China (P2020-KF08).
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Supplementary material 1
Table S1
Authors: Cong-Miao Xie, Li-Song Wang, Zun-Tian Zhao, Yan-Yun Zhang, Xin-Yu
Wang, Lu-Lu Zhang
Data type: docx. le
Explanation note: A list of specimens and GenBank accession number of sequences
used in this study. e new sequences generated are in bold.
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/mycokeys.87.72614.suppl1
... This study is based primarily on specimens collected in December 2020 and follow up surveys carried out in 2021 during field trips in the northeastern part of Kurdistan region of Iraq, Mawat and Gapelon districts (Fig. 1). The specimens were morphologically examined by standard stereomicroscope and compound microscopic with standard identification methods (Gaya 2010, Westberg and Sohrabi 2012, Nimis and Martellos 2016, Jason 2019, Xie et al. 2022. All specimens cited here were determined, revised or confirmed by the author and verified by Dr Mohammad Sohrabi from Iran. ...
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