Phytotaxa 597 (4): 287–296 ISSN 1179-3155 (print edition)

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Article PHYTOTAXA
Copyright © 2023 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.597.4.4

Three new species of Herpothallon (Lichenized Ascomycota) from Southern China

LINLIN LIU1,3, QIJIA ZUO1,4, JUNXIA XUE1,5, ZHAOJIE REN2,6 & LULU ZHANG1,7*
1
Institute of Environment and Ecology, Shandong Normal University, Jinan, 250014, P. R. China.
2
Shandong Museum, Jinan, 250014, P. R. China.
3�
[email protected]; https://orcid.org/0000-0003-3571-7119
4�
[email protected]; https://orcid.org/0000-0001-8910-1809
5�
[email protected]; https://orcid.org/0000-0002-6387-0148
6�
[email protected]; https://orcid.org/0000-0003-3206-1484
7�
[email protected]; https://orcid.org/0000-0001-8011-4451
*Corresponding author: [email protected]

Abstract

This paper describes three species of Herpothallon new to science from southern China: H. glaucescens, H. lilacinum and
H. tomentosum. The three species all possess a non-pigmented thallus, hypothallus and prothallus. Herpothallon glauces-
cens has a white, whitish grey to greyish green thallus and swollen, subglobose to ± vermiform pseudisidia, rounded at the
top. Herpothallon lilacinum has subglobose or irregularly cushion-shaped, fluffy-felty pseudisidia, white at the base, lilac
to lilac grey at their tips. Herpothallon tomentosum has globular pseudisidia, felty with many projecting hyphae, sometimes
containing a central pycnidium. Detailed descriptions for all three new species are provided with an updated key to the genus
Herpothallon in China. Additionally, a phylogenetic tree based on Bayesian and ML analyses of mtSSU data shows the posi-
tion of the new species in Herpothallon.

Keywords: Arthoniaceae, byssoid thalli, sterile lichens, taxonomy, mtSSU

Introduction

Herpothallon Tobler (1937: 446) is a widespread genus in tropical and subtropical regions, growing mostly in humid
and sheltered habitats within forests. The lichen genus is recognized by the usually loosely appressed thallus with a
byssoid hypothallus and prothallus that is often a different colour compared to the upper, algal-containing part of the
thallus; pseudisidia or pseudisidioid structures are typically present, and the different species contain a diverse array
of lichen secondary metabolites (Aptroot et al. 2009). The genus was first established by Tobler (1937), subsequently
treated as a synonym of Cryptothecia Stirton (1876 [1877]: 164), but reinstated by Aptroot et al. (2009), recognizing
29 species worldwide. So far approximately 50 species are known world-wide, eight have been reported from China
(Jagadeesh Ram et al. 2009; Jagadeesh Ram & Sinha 2009; Frisch et al. 2010; Jagadeesh Ram & Sinha 2011; Cheng et
al. 2012; Bungartz et al. 2013; Frisch et al. 2014; Jagadeesh Ram 2014; Aptroot et al. 2017; Sipman 2018; Aptroot &
Souza 2021; Chen et al. 2022; Lendemer & James C. 2022). Almost all Herpothallon species are sterile (only H. fertile
Aptroot & Lücking (2009: 40) and H. inopinatum Frisch & G. Thor (2014: 64) are known to produce ascospores),
therefore species boundaries within the genus are mainly based on morphological and chemical characteristics (Frisch
et al. 2014).
Due to its ecological and climatic preferences, the genus Herpothallon presents its highest diversity in tropical and
subtropical regions. Here we focused on lichen surveys in Southern China, which is characterized by a subtropical and
tropical, warm and humid monsoon climate. During our survey, three new species were found and are now described,
using mitochondrial small subunit (mtSSU) rDNA data, to also construct a phylogenetic tree showing the position of
these new species, supporting the delimitation of the new taxa.

Accepted by Frank Bungartz: 29 Apr. 2023; published: 17 May 2023 287

Material & methods

Investigation of lichen specimens:—The specimens studied were collected in Fujian, Guizhou, Zhejiang Provinces
and Guangxi Zhuang Autonomous Region, China, and are preserved in the Lichen Section of the Botanical Herbarium,
Shandong Normal University, Jinan, China (SDNU).
Morphology and Chemistry:—The morphological and anatomical characters were examined under a
stereomicroscope (COIC XTL7045B2) and a polarizing compound microscope (Olympus CX41). For identification
thallus and medulla were tested with the spot test reagents K (a 10% aqueous solution of potassium hydroxide), C
(a saturated solution of aqueous sodium hypochlorite), P (a saturated solution of p-phenylenediamine in 95% ethyl
alcohol), I (a 3% solution of Lugol’s iodine). Polarized light microscopy (pol) was used to locate crystals in thallus
sections. H2SO4 (a 10% solution of sulfuric acid) was then applied to tests if the characteristic needle shaped crystals
formed when calcium oxalate is present in the thallus. Lichen secondary metabolites were identified using standardized
thin-layer chromatography (TLC) with solvent systems A, B’ and C (Orange et al. 2010). Photos were taken in the
Olympus SZX16 and BX61 microscope with a DP72 camera system.
DNA Extraction:—Molecular materials were extracted directly from the clean growing portions of the thallus
(e.g.,prothallus hyphae, pseudisidia, pseudisidioid structure) of Herpothallon specimens, pigmented or carbonized
portions were removed as far as possible. Genomic DNA was extracted using the Sigma-Aldrich REDExtract-N-Amp
Plant PCR Kit (St. Louis, Missouri, U.S.A.) following the manufacturer’s instructions, except only 30 μl of extraction
buffer and 30 μl dilution buffer were used.
PCR and Sequencing:—The following primers were used for PCR amplification: mrSSU1 and mrSSU3R (Zoller
et al. 1999). The 50 μl PCR mixture consisted of 2 μl DNA, 2 μl of each primer, 25 μl 2 × Taq PCR MasterMix (Taq
DNA Polymerase [0.1 unit/μl]; 3 mM MgCl2; 100 mM KCl; 0.5 mM dNTPs; and 20 mM Tris-HCl [pH 8.3]) (Tiangen,
Beijing, China) and 19 μl dd H2O. PCR cycling conditions were 94 ℃ for 10 minutes, followed by 34 cycles of 95 ℃
for 45 seconds, 50 ℃ for 45 seconds, and 72 ℃ for 90 seconds, followed by a final extension at 72 ℃ for 10 minutes.
Sequencing was performed by the company of BioSune Biological Technology (Shanghai).
Sequence alignments:—The mtSSU sequences were assembled and edited using SeqMan v.7.0 (DNAstar packages).
Sequences were aligned using the online version of MAFFT v.7.0.26 and MEGA v.7.0. The algorithm of MAFFT choose
Auto (FFT-NS-1, FFT-NS-2, FFT-NS-i or L-INS-i; depends on data size). The species Cryptothecia austrocoreana J.J.
Woo et al. (2017: 341) was chosen as outgroup (Woo et al. 2017).
Phylogenetic analyses:—The alignments were analyzed using maximum likelihood (ML) and Bayesian
approaches. The ML analyses were performed using the program MEGA v.7.0. support values were assessed using
the‘rapid bootstrapping’ option with 1000 replicates. For the Bayesian analysis, the best substitution models were
estimated using jModelTest 2.1.7 (Darriba et al. 2012). Based on the result, we used TVM+G model for mtSSU. Four
Markov chains were run with 2 million generations for this dataset. Trees were sampled every 100 generations, with
the first 25% of trees discarded as burn-in. Stationarity of analysis was determined by examining the standard deviation
of split frequencies (< 0.01). Posterior probabilities above 0.9 and bootstrap support above 70% were considered
significant supporting values. Generated phylogenetic trees were visualized with FigTree v. 1.4.2 (Rambaut 2012).
Results & Discussion
Molecular phylogeny:—A total of 9 sequences of mtSSU were newly generated from 4 specimens, and 5
sequences downloaded from NCBI (Table 1). The aligned mtSSU region comprised 929 sites.
The phylogenetic trees obtained from maximum likelihood (ML) and Bayesian inference analysis (BI) exhibited
the same topology; we therefore present only the ML tree, with bootstrap support ≥ 70% for the ML analysis and
posterior probabilities ≥ 0.95 for the Bayesian analysis (Fig. 1).
Through evolutionary distances and support, the taxonomic positions of those three new species can be confirmed,
they indeed belong to Herpothallon. And we can clearly see that there are two main clades in Herpothallon: the three
new taxa and H. echinatum Aptroot et al. (2009: 38) belong to clade A, H. inopinatum, H. kigeziense Frisch & G. Thor
(2014: 303) and H. rubrocinctum (Ehrenb.) Aptroot, Lücking & G. Thor (2009: 61) belong to clade B. The four species
belong to clade A without chiodectonic acid, the three species belong to clade B all possess chiodectonic acid; In clade
A, H. glaucescens and H. lilacinum clusters with H. echinatum, they all possess psoromic acid as major substance, and
H. tomentosum which contains confluentic acid as major substance forms a separate lineage, indicating that secondary
metabolites are valuable in the classification of Herpothallon. Additional studies are necessary, including extended
sampling of more variable loci and material to confirm these preliminary results.

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TABLE 1. Taxon, locality, GenBank accession numbers and voucher for the specimen of Herpothallon included in the
phylogenetic analysis. Newly generated sequences are in bold. * = outgroup.
Taxon Locality GenBank accession no. Voucher
H. echinatum 1 China OQ676528 20220048 (SDNU)
H. echinatum 2 China OQ676537 20220494 (SDNU)
H. echinatum 3 China OQ676534 20220455 (SDNU)
H. glaucescens China OQ676531 20220069 (SDNU)
H. inopinatum Mexico KJ850964 Rudolphi 12 (UPS)
H. kigeziense Uganda KF707644 Frisch 11/Ug26 (UPS)
H. lilacinum 1 China OQ676532 20220090 (SDNU)
H. lilacinum 2 China OQ676529 20220240 (SDNU)
H. rubrocinctum Mexico KF707643 Rudolphi 5 (UPS)
H. rubrocinctum America GU327693 Nelsen 4006 (F)
H. tomentosum 1 China OQ676538 20220565 (SDNU)
H. tomentosum 2 China OQ676533 20220463 (SDNU)
H. tomentosum 3 China OQ676539 20222313 (SDNU)
Cryptothecia* austrocoreana South Korea MF769374 KoLRI No.041892

FIGURE 1. Phylogenetic tree constructed from Maximum Likelihood analysis of Herpothallon species, based on the mtSSU dataset.
Maximum likelihood (ML) bootstrap value ≥ 70, and Bayesian posterior probabilities (PP) value ≥ 0.95 are shown above the branches.
Bootstrap values are shown in the order of ML, PP in the tree. Newly described species are marked in bold. Scale = 0.02 substitution per
site.

New species

Herpothallon glaucescens L.L. Liu & Lu L. Zhang, sp. nov. Mycobank number: 843998 (Fig. 2)
Type:—CHINA. Zhejiang Province: Lishui City, Jingning County, Baiyun Protection Station. 1131.3 m elev., 27°72′57.76″ N, 119°64′12.09″
E, on bark of Cunninghamia R. Br., 2 December 2020, C.G. Zhao & Lu L. Zhang 20211617 (Holotype in SDNU).

Thallus corticolous, up to 3 cm across, suborbicular, sometimes flaking off, loosely to firmly appressed to the substrate,
felty to byssoid, dull, white in the center, whitish grey to greyish green along the margin, in section up to 200 µm thick,
with abundant calcium oxalate crystals throughout the thallus (insoluble in KOH, dissolving and recrystallizing as
colourless, needle-shaped crystals in 10% H2SO4), with 1–2 µm wide hyphae. Hypothallus whitish, byssoid, composed
Three new species of Herpothallon Phytotaxa 597 (4) © 2023 Magnolia Press • 289
of 1–2 µm wide hyphae. Prothallus up to 2 mm broad, whitish, indistinct, byssoid, composed of interwoven and radiating
hyphae. Pseudisidia numerous, sparse to dense, dispersed or 2–3 in coralloid aggregations, often branched, swollen,
subglobose to ±vermiform, rounded at the top, of the same colour as the thallus, compact with few projecting hyphae,
0.15–0.18 mm in diam., or 0.22–0.3 × 0.11–0.18 mm. Photobiont trentepohlioid, single or a few cells aggregated; cells
yellowish, 10–12.5 × 7–8 μm. Asci and pycnidia not seen.

FIGURE 2. Herpothallon glaucescens (holotype, SDNU 20211617). A. Thallus and prothallus. Scale = 2 mm. B. Pseudisidia. Scale =
600 µm.

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 Chemistry and spot tests: Thallus and prothallus K−, C−, P+ bright yellow, UV−, I+ blue in medulla. TLC:
psoromic acid (major), 2’-O-demethylpsoromic acid (minor).
Etymology: The epithet “glaucescens” refers to the whitish grey colour of the thallus and the pseudisidia.
Ecology and distribution: The new species was found growing on bark of Cunninghamia R. Br. at the Baiyun
Protection Station in the Zhejiang Province and the bark of other trees in the Daming Mountain National Nature
Reserve of the Guangxi Zhuang Autonomous Region.
Note: This species is characterized by a thallus with swollen, compact, subglobose to ±vermiform, often branched
pseudisidia, with few projecting hyphae, and the presence of psoromic and 2’-O-demethylpsoromic acids. Although
H. globosum G. Thor (2009: 42) also has globose pseudisidia and contains psoromic acid, it lacks calcium oxalate
crystals, has a red hypothallus and prothallus, the upper parts of its pseudisidia are dark red and often with black spots
(Aptroot et al. 2009). Herpothallon biacidum Frisch, Elix & G. Thor (2010: 286) possesses a loosely attached thallus
with abundant calcium oxalate crystals, globular to short cylindrical pseudisidia, up to 0.30 × 0.12 mm, in small
coralloid aggregations, but it has a brown to blackish brown hypothallus and produces gyrophoric and norstictic acids
(Frisch et al. 2010). Herpothallon coralloides Jagadeesh (2014: 40) is also similar to H. glaucescens because of its
firmly to loosely appressed thallus and white prothallus, cylindrical pseudisidia that are simple to irregularly branched
and coralloid, but the pseudisidia of H. coralloides are much larger, up to 1.0 × 0.1 mm. The two species also differ in
chemistry, H. coralloides containing confluentic and norstictic acids (Jagadeesh Ram 2014).
Phylogenetically, H. glaucescens is the sister taxon to H. echinatum (Fig. 1), they are similar in containing
psoromic acid as a major secondary metabolite, but H. echinatum has a much softer thallus, often seemingly farinose
due to loose soredioid fragments, and its pseudisidia are cylindrical, more elongated, up to 0.5 × 0.1 mm, and are
mostly unbranched, felty with projecting hyphae.
Additional specimen examined: CHINA. Guangxi Province: Nanning City, Wuming County, Daming Mountain
National Nature Reserve, 592.0 m elev., 23°30′12.336″ N, 108°26′08.231′′ E, on bark of a tree, 30 December 2020, X.
Zhang et al. 20211618 (SDNU).

Herpothallon lilacinum L.L. Liu & Lu L. Zhang, sp. nov. Mycobank number: 845790 (Figs 3, 4)
Type:—CHINA. Guizhou Province: Tongren City, Yang Jia Ao township, Bi Er Tang village. 874 m elev., 27°52’27.59” N, 107°58’10.71”
E, on rock, 10 June 2022, L.L. Liu, Y.X. Bi, Z.H. Jiang & D.C. Yan 20220232 (Holotype in SDNU).

Thallus corticolous or saxicolous, up to 3.5 cm across, irregular shaped, sometimes flaking off, loosely appressed to
the substrate, rather soft, felty, often seemingly farinose, dull, white to cream white, in section up to 200 µm thick, with
many calcium oxalate crystals throughout the thallus (insoluble in KOH, dissolving and recrystallizing as colourless,
needle-shaped crystals in 10% H2SO4), with 1–2 µm wide hyphae. Hypothallus whitish, byssoid, composed of 1–2
µm wide hyphae. Prothallus up to 0.8 mm broad, whitish, indistinct, byssoid to cottony, composed of interwoven and
radiating hyphae. Pseudisidia numerous, unbranched, soft, whitish subglobose or irregularly cushion-shaped, fluffy-
felty with many projecting hyphae, basally of the same colour as the thallus, upper parts often lilac to lilac grey,
0.1–0.45 mm in diam.. Photobiont trentepohlioid, in short, irregular threads; cells yellowish green, 10–15 × 5–8 μm.
Asci and pycnidia not seen.
Chemistry and spot tests: Thallus and prothallus K−, C−, P+ bright yellow, UV−, I+ blue in medulla, the lilac to lilac
grey parts K+ black blue, C−. TLC: psoromic acid (major), an unknown substance (minor), 2’-O-demethylpsoromic
acid (minor).
Etymology: The epithet “lilacinum” refers to the lilac to lilac grey pseudisidia.
Ecology and distribution: The new species was found growing on rock wall by the roadside and bark of a tree in
Guizhou Province.
Notes: This species is characterized by the subglobose or irregularly cushion-shaped, lilac to lilac grey, fluffy-
felty pseudisidia, 0.1–0.45 mm in diam., the psoromic and 2’-O-demethylpsoromic acids, and an unknown substance
chemistry. Herpothallon lilacinum is most similar to H. weii Yuliang Chen & Haiying Wang (2012: 440): both contain
psoromic acid and the similar unknown substance, but H. weii has a tightly appressed thallus, an I− medulla, a distinct
prothallus, pinkish and larger, and not whitish subglobose pseudisidia, up to 1 × 0.5 mm (Cheng et al. 2012). Crypthonia
albida (Fée) Frisch & G. Thor (2010: 290) also has fluffy-felty pseudisidia and contains psoromic acid as its major
substance, but has loosely byssoid, whitish pseudisidia, up to 1.0 × 1.0 mm (Frisch & G. Thor, 2010). Herpothallon
himalayanum Jagadeesh & Sinha (2009: 40) and H. capilliferum Pengfei Chen & Lulu Zhang (2022: 02) are also
similar to H. lilacinum in producing fluffy-felty pseudisidia, but they differ in secondary chemistry: Herpothallon
himalayanum contains gyrophoric acid as its major substance (Jagadeesh & Sinha, 2009), and H. capilliferum only
contains norstictic acid (Chen et al. 2022).

Three new species of Herpothallon Phytotaxa 597 (4) © 2023 Magnolia Press • 291
FIGURE 3. A–B. Herpothallon lilacinum growing on rock, (holotype, SDNU 20220232). A. Thallus and prothallus. Scale = 1 mm.. B.
Pseudisidia. Scale = 400 µm. C–D. The new species H. lilacinum growing on bark, (paratype, SDNU 20220090). C. Thallus. Scale = 1.5
mm. D. Pseudisidia. Scale = 600 µm.

FIGURE 4. TLC of H. lilacinum with C system (at: atranorin; U: the unknown substance; P: psoromic acid).

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 Phylogenetically, H. lilacinum clusters with H. echinatum and H. glaucescens (Fig. 1), they all possess psoromic
acid as major, but H. echinatum and H. glaucescens have subglobose or cylindrical pseudisidia, all without the unknown
substance.
Additional specimen examined: CHINA. Guizhou Province: Tongren City, Yang Jia Ao township, Bi Er Tang
village. 874 m elev., 27°52’27.59” N, 107°58’10.71” E, on rock, 10 June 2022, L.L. Liu, Y.X. Bi, Z.H. Jiang & D.C.
Yan 20220231, 20220237, 20220238, 20220239, 20220240, 20220253 (SDNU); Guizhou Province: Tongren City, Xu
Jia Ba town, Zhang Jia Gou village, along the stream. 851 m elev., 27°55’33.38” N, 108°1’59.13” E, on bark of a tree,
11 June 2022, L.L. Liu, Y.X. Bi, Z.H. Jiang & D.C. Yan 20220090 (SDNU).

Herpothallon tomentosum L.L. Liu & Lu L. Zhang, sp. nov. Mycobank number: 845791 (Fig. 5)
Type:—CHINA. Fujian Province: Longyan City, Dongxiao National Forest Park, Barbecue field. 450 m elev., 24°58’25.33” N, 117°0’56.67”
E, on bark of a tree, 12 July 2022, L.L. Liu, J.X. Xue & L. Wang 20220468 (Holotype in SDNU).

Thallus corticolous, up to 2 cm across, suborbicular to sometimes irregular, not flaking off, loosely to firmly appressed
to the substrate, soft, minutely felty, dull, blue green to greenish grey, in section up to 120 µm thick, with few calcium
oxalate crystals throughout the thallus (insoluble in KOH, dissolving and recrystallizing as colourless, needle-shaped
crystals in 10% H2SO4), with 1–2 µm wide hyphae. Hypothallus whitish, byssoid, composed of 1–2 µm wide hyphae.
Prothallus up to 0.9 mm broad, whitish, distinct, byssoid, composed of interwoven and radiating hyphae. Pseudisidia
numerous, unbranched, globular, of the same colour as the thallus, soft, felty with many projecting hyphae, 0.06–0.12
mm in diam.. Photobiont trentepohlioid, single or a few cells aggregated; cells yellowish green, 12.5–15 × 5–10
μm. Asci not seen. Pycnidia embedded in the tips of some pseudisidia, opening with a apical pore, pigmentation
occasionally extending along the pycnidial wall. Conidia simple, hyaline, short bacilliform, 3–4 × 1–1.5 μm.
Chemistry and spot tests: Thallus and prothallus K−, C−, P−, UV−, I− in medulla. TLC: confluentic acid (major),
2’-O-methylmicrophyllinic acid (minor).
Etymology: The epithet “tomentosum” refers to the pseudisidia felty with many projecting hyphae.
Ecology and distribution: The new species was found growing on bark of trees beside a stream of Dongxiao National
Forest Park and on bark of trees beside a mountain path of Tianzhu Mountain Forest Park of Fujian Province.
Notes: This species is characterized by the globular pseudisidia each containing a pycnidium, 0.06–0.16 mm in
diam., and the presence of confluentic and 2’-O-methylmicrophyllinic acids. Herpothallon tomentosum is most similar
to H. cinereum G. Thor (2009: 34) in its minutely felty thallus, the white, byssoid-felty prothallus and the presence of
confluentic and 2’-O-methylmicrophyllinic acid in its thallus. However, H. cinereum has a loosely appressed thallus,
up to 200 μm thick, and cylindrical pseudisidia up to 0.5 × 0.1 mm, without pycnidia at their tips. Herpothallon
tomentosum has the same chemistry as H. confluenticum Aptroot & Lücking (2009: 36); both have pycnidia at the
tips of their pseudisidia, but the latter has a rather firm thallus delimited by a dirty whitish prothallus, on a whitish
to brownish hypothallus, and cylindrical pseudisidia that are partly cauliflower-like at the tips, up to 0.6 × 0.2 mm
(Aptroot et al. 2009). The specimens of H. echinatum that Bungartz et al. collected from Ecuador (2013: 752) also have
a non-pigmented thallus, prothallus and hypothallus, globular pseudisidia with pycnidia at the tips, and bacilliform
conidia (3–4 × 1–1.5 μm), but they contain psoromic acid (Bungartz et al. 2013). Two other morphologically similar
species are H. biacidum and H. subglobosum Pengfei Chen & Lulu Zhang (2022: 07): both have a minutely felty
thallus and globular pseudisidia, but differ in chemistry: H. biacidum contains gyrophoric and norstictic acids (Frisch
et al. 2010), whereas H. subglobosum contains gyrophoric, lecanoric and umbilicaric acids (Chen et al. 2022).
Phylogenetically, H. tomentosum belongs into a different monophyletic clade from the other species in clade B
(Fig. 1), demonstrating that it is a distinct species. As part of our survey, we collected multiple specimens of the new
species in two areas. These specimens are similar in morphological and anatomical characters except that some of the
specimens lack pycnidia. In our phylogenetic analysis this material appears on the same branch, with a relatively close
evolutionary distance, strongly supported (BS=100, PP=1.00).
Additional specimen examined: CHINA. Fujian Province: Longyan City, Dongxiao National Forest Park,
Barbecue field. 450 m elev., 24°58’25.33” N, 117°0’56.67” E, on bark of a tree, 12 July 2022, L.L. Liu, J.X. Xue &
L. Wang 20220477 (SDNU); Fujian Province: Longyan City, Dongxiao National Forest Park, Bajiao forest to Suoluo
group. 558 m elev., 24°58’21.42” N, 117°1’1.71” E, on bark of a tree, 12 July 2022, L.L. Liu, J.X. Xue & L. Wang
20220565, 20220582, 20220587 (SDNU); Fujian Province: Xiamen City, Tianzhu Mountain Forest Park, No. 3 branch
road near the air monitoring station. 183 m elev., 24°35’46.33” N, 117°54’31.06” E, on bark of a tree, 11 July 2022,
L.L. Liu, J.X. Xue & L. Wang, 20220442, 20220443, 20220462, 20220463 (SDNU).

Three new species of Herpothallon Phytotaxa 597 (4) © 2023 Magnolia Press • 293
FIGURE 5. Herpothallon tomentosum (holotype, SDNU 20220468), A. Thallus and prothallus. Scale = 3 mm. B. Pseudisidia. Scale = 200
µm. C. Pycnidia. Scale = 40 µm. D. Conidia. Scale = 20 µm.

Key to the species of Herpothallon known from China

1. Prothallus and pseudisidia with red pigment, K+ purple in pigmented parts...........................................................H. rubrocinctum
Prothallus and pseudisidia without red pigment (rarely pinkish or lilac pigments present in pseudisidia)........................................2
2. Thallus C+ red; gyrophoric acid major, lecanoric acid and some other substances minor.................................................................3
Thallus C–; gyrophoric and lecanoric acids absent.............................................................................................................................5
3. Thallus K+ yellow; an unknown substance present (RF close to atranorin in solvent C) present; pseudisidia cylindrical (0.2 × 0.1
mm)................................................................................................................ H. viridi-isidiatum P.F. Chen & L.L. Zhang (2022: 07)
Thallus K–; the unknown minor substance absent; pseudisidia globular or cylindrical.....................................................................4
4. Pseudisidia globular (0.1 × 0.1 mm)..........................................................................................................................H. subglobosum
Pseudisidia cylindrical (1.0 × 0.1 mm)...........................................................H. philippinum (Vain.) Aptroot & Lücking (2009: 43)
5. Thallus K+ yellow or K+ yellow then red, P+ orange-red; stictic or norstictic acids present............................................................6
Thallus K–, P– or P+ yellow; stictic and norstictic acids absent........................................................................................................7
6. Thallus K+ yellow then red; norstictic acid present; pseudisidia irregularly cushion-shaped (0.4 × 0.2 mm)........... H. capilliferum
Thallus K+ yellow; stictic acid present; pseudisidia cylindrical (0.3 × 0.1 mm)..................................................................................
.......................................................................................................................... H. polyisidiatum P.F. Chen & L.L. Zhang (2022: 02)
7. Thallus P–; psoromic acid absent........................................................................................................................................................8
Thallus P+ yellow; psoromic acid present..........................................................................................................................................9
8. Confluentic acid major; globular pseudisidia (0.06–0.12 mm in diam.)..................................................................... H. tomentosum
Perlatolic acid major; minute, irregular, soredia-like granular pseudisidia (0.05 × 0.05 mm in diam.)...............................................
........................................................................................................................H. granulare (Sipman) Aptroot & Lücking (2009: 43)
9. Pseudisidia cylindrical (0.5 × 0.1 mm)........................................................................................................................... H. echinatum
Pseudisidia subglobose to irregularly cushion-shaped......................................................................................................................10

 Wei J.C. (2020) The Enumeration of Lichenized Fungi in China. In: Beijing, China Forestry Publishing House, China, 44–47.
Specimen examined: CHINA. Guizhou Province: Leishan County, Leigongshan National Forest Park, 892 m elev., on bark of a tree, 2
Nov. 2009, Q. Tian, 20102819 (SDNU). Discussion: The identification of this material as Herpothallon rubrocinctum appears problematic,
because its prothallus is an orange to red (rather than bright scarlet red), the thallus lacks secondary metabolites and the pseudisidia of the
specimen are granular. Molecular data could not successfully be obtained from the material. The report of H. rubrocinctum may thus refer
to a still undescribed species, which needs to be further investigates, when fresh specimens are collected.

294 • Phytotaxa 597 (4) © 2023 Magnolia Press LIU et al.

10. Pseudisidia without pinkish or lilac pigments; subglobose (0.22–0.3 × 0.11–0.18 mm).............................................H. glaucescens
Pseudisidia with pinkish or lilac pigments; subglobose to mostly irregularly cushion-shaped........................................................11
11. Thallus tightly appressed, medulla I–; pseudisidia irregularly cushion-shaped, pinkish, large (1 × 0.5 mm)..........................H. weii
Thallus loosely appressed, medulla I+ blue; pseudisidia whitsh subglobose or irregularly cushion-shaped, basally white, upper
parts often lilac to lilac grey, small (0.1–0.45 mm in diam.)............................................................................................H. lilacinum

Acknowledgements

We thank Prof. André Aptroot (UFMS University, Campo Grande, Brazil) for the valuable suggestions and helping to
check a new Herpothallon species: H. glaucescens. This work was supported by the Emergency Management Project
of the National Natural Science Foundation of China (32170002, 31750001).

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