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Citation: Su, L.; Yu, T.; Xue, R.;
Zhang, W.; Xu, C.; Xia, X.; Li, J.; Lei,
H.; Dong, Y.; Zhang, G.; et al. New
Contributions on Species Diversity of
Genus Hydnum and Lentaria s.l. in
China. J. Fungi 2024,10, 824.
hps://doi.org/10.3390/jof10120824
Academic Editor: Seung‑Yoon Oh
Received: 23 October 2024
Revised: 18 November 2024
Accepted: 25 November 2024
Published: 27 November 2024
Copyright: © 2024 by the authors.
Licensee MDPI, Basel, Swierland.
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4.0/).
Article
New Contributions on Species Diversity of Genus Hydnum and
Lentaria s.l. in China
Linjie Su 1, 2 , Taijie Yu 1, 2 , Rou Xue 1, 2, Wenhao Zhang 1,2, Chang Xu 1,2, Xing Xia 1,2, Jia Li 1,2, Hanchi Lei 1, 2,
Yang Dong 3, Guoli Zhang 1,2,* and Liping Tang 1,2,*
1School of Pharmaceutical Sciences and Yunnan Key Laboratory of Pharmacology for Natural Products,
Kunming Medical University, Kunming 650500, China; suull121@163.com (L.S.);
18385037921@163.com (T.Y.); 18787089431@163.com (R.X.); zhangwenhao1@kmmu.edu.cn (W.Z.);
xuchang727298@163.com (C.X.); xiaxing3790@outlook.com (X.X.); 18892800409@163.com (J.L.);
15559729789@163.com (H.L.)
2Yunnan College of Modern Biomedical Industry, Kunming Medical University, Kunming 650500, China
3Forestry Bureau of Shitai County, Shitai 245100, China; 15256009347@163.com
*Correspondence: zhangguoli2001@163.com (G.Z.); tangliping@kmmu.edu.cn (L.T.)
Abstract: Southwest China is extremely rich in fungal resources, and a large number of new taxa
have been discovered in recent years. In the present study, we examined 26 specimens of the gen‑
era Hydnum and Lentaria sensu lato, most of which were obtained in Yunnan Province. Through
ITS‑nrLSU‑tef1 phylogenetic analysis, combined with morphological studies and geographic analy‑
ses, ve new species were described, viz. H. cremeum (奶油齿菌), H. avoquamosum (黄鳞齿菌), H.
roseoalbum (粉白齿菌), H. roseotangerinum (粉橙齿菌), and L. subalpina (亚高山木瑚菌). Furthermore,
we also supplied new information on some known species, including host plants and new distri‑
bution records. We re‑examined the holotype sequences of two known taxa, H. abellatum and H.
pallidomarginatum, treating H. abellatum as a synonym of H. pallidomarginatum. Additionally, a key
to Lentaria s.l. in China was provided.
Keywords: taxonomy; species delimitation; coralloid fungi; edible mushroom; spinous fungi
1. Introduction
Yunnan Province, China, is located in the southwestern extension of the Himalayas.
It has tropical rainforests, subtropical evergreen broad‑leaved forests, and subalpine conif‑
erous forests, which provide favorable habitats and symbiotic hosts for the growth of
fungi [1–3]. As suggested by Hawksworth, the number of fungi (including lichens) oc‑
curring in a given area is roughly six times as high as the native plant populations, and
thus, a total of approximately 104,000 fungal species would be expected in Yunnan [4].
Up to date, about 3000 species of higher fungi have been reported in Yunnan [5–16]. This
means that a vast number of macrofungal resources in this region remain to be explored.
Hydnum L. belongs to Hydnaceae, Cantharellales, which was established by Linnaeus
in 1753 and typed as H. repandum L. [17,18]. Species in this genus are morphologically
characterized by white, ochraceous to brownish‑orange basidiomata with a spinous hy‑
menophore, often developing stipe, smooth, hyaline, globose, subglobose to ellipsoid ba‑
sidiospores, and clavate to subfusiform basidia with one to eight sterigmata [17–26]. Hyd‑
num is distributed worldwide and usually ectomycorrhizal fungi, which can establish sym‑
biosis with wide ranges of host plants, such as species in Dipterocarpaceae, Fagaceae, Mal‑
vaceae, Pinaceae, and Salicaceae [18,19,21–23]. Previously, Hydnum included taxa with a
spinose hymenophore, but a majority of species were later transferred to the genera Heri‑
cium Pers., Phellodon P. Karst., and Sarcodon Quél. Ex P. Karst. [17,18,20,22,27]. Accord‑
ing to molecular phylogenetic results, the genus Hydnum was divided into ve subgenera,
viz. the subgen. Alba Niskanen & Liimat., subgen. Brevispina T. Cao & H. S. Yuan, sub‑
gen. Hydnum L., subgen. Pallida Niskanen & Liimat., and subgen. Rufescentia Niskanen
J. Fungi 2024,10, 824. https://doi.org/10.3390/jof10120824 https://www.mdpi.com/journal/jof
J. Fungi 2024,10, 824 2 of 25
& Liimat. [18,20]. The species of Hydnum were mainly found in temperate regions, only
a few taxa were reported from subtropical to tropical forests [18–20,22–25,28]. Recently,
new species from the genus Hydnum were continuously reported in China. Until 2024,
22 species have been reported in China, and seven of them were distributed in Yunnan
Province [18,19,21,26,29]. However, the diversity of Hydnum in China is still seriously un‑
derestimated [4].
The genus Lentaria Corner, belonging to Lentariaceae, Gomphales, was proposed in
1950, and it was widespread from temperate to tropical zones in the world [30–34]. Species
in the genus Lentaria s.l. are characterized by coralloid basidiomata emerging from a subicu‑
lum, are supercial or immersed on the substrate, often branched, with high length/width ra‑
tio basidiospores, a monomitic hyphal system, and being saprotrophic on the dead wood of
various plants. This genus plays important roles in forest ecosystems [31,33,34]. However,
there is lile information available on it [30,31,34]. So far, only 23 records can be found in
the Index Fungorum (https://www.indexfungorum.org/, accessed on 18 November 2024).
Among them, ve species have been reported in China, viz. L. bambusina P. Zhang & Zuo
H. Chen, L. byssiseda Corner, L. patouillardii (Bres.) Corner, L. surculus (Berk.) Corner, and
L. uncispora P. Zhang & Zuo H. Chen, with only L. surculus recorded in Yunnan [31]. Addi‑
tionally, according to previous studies, this polyphyletic genus needs further study [31,33].
In the past years, some specimens from two genera, Hydnum and Lentaria s.l., have
been collected, mainly in southwestern China. Based on both morphological and molec‑
ular data, this study aims to (i) identify and describe new taxa; (ii) supply new informa‑
tion for ve known species from the genera Hydnum and Lentaria s.l.; and (iii) discuss one
doubtful species and clarify the species diversity of Hydnum and Lentaria s.l. in China.
2. Materials and Methods
2.1. Specimen Collecting
Twenty‑six specimens were collected during the rainy season from Anhui, Hainan,
Jilin, and Yunnan Provinces in China from 2015 to 2024. Fresh fruiting bodies were dried
with a dryer at 50–60 ◦C and silica gel after detailed eld records were made on the same
day. Those collections were deposited in the Mycological Herbarium of Kunming Medical
University (MHKMU). Detailed information is shown in Table 1and Figure 1.
Table 1. Information of specimens examined in this study.
No. Specimen Voucher Locality Habitat Altitude Longitude
and Latitude Date
1 MHKMU C Xu 071 China: Anhui mixed forests 955 m N 30◦00′20″,
E 117◦27′98″ 21 July 2023
2 MHKMU HY Huang 151 China: Yunnan broad‑leaved
forests 2269 m N 24◦38′57″,
E 104◦09′57″ 23 August 2018
3 MHKMU HY Huang 873 China: Yunnan mixed forests 2731 m N 27◦21′16″,
E 100◦09′37″ 24 August 2020
4 MHKMU J Zhao 225 China: Yunnan mixed forests 1710 m N 25◦19′58″,
E 98◦35′30″ 1 August 2015
5 MHKMU LJ Su 430 China: Hainan broad‑leaved
forests 245 m N 19◦17′52″,
E 109◦41′20″ 15 August 2024
6 MHKMU LJ Su 431 China: Hainan broad‑leaved
forests 245 m N 19◦17′52″,
E 109◦41′20″ 15 August 2024
7 MHKMU LP Tang 2116 China: Yunnan mixed forests 1710 m N 25◦19′58″,
E 98◦35′30″ 1 August 2015
8 MHKMU LP Tang 2886 China: Jilin mixed forests 820 m N 42◦22′19″,
E 128◦00′15″ 20 August 2019
9 MHKMU LP Tang 3319 China: Yunnan mixed forests 2890 m N 26◦46′11″,
E 100◦02′51″ 21 August 2020
10 MHKMU LP Tang 3453 China: Yunnan mixed forests 3170 m N 27◦23′14″,
E 100◦06′31″ 26 August 2020
J. Fungi 2024,10, 824 3 of 25
Table 1. Cont.
No. Specimen Voucher Locality Habitat Altitude Longitude
and Latitude Date
11 MHKMU LP Tang 3454 China: Yunnan mixed forests 3170 m N 27◦23′14″,
E 100◦06′31″ 26 August 2020
12 MHKMU LP Tang 3458 China: Yunnan mixed forests 3170 m N 27◦23′14″,
E 100◦06′31″ 26 August 2020
13 MHKMU LP Tang 3458‑1 China: Yunnan mixed forests 3170 m N 27◦23′14″,
E 100◦06′31″ 26 August 2020
14 MHKMU M Mu 740 China: Yunnan mixed forests 3180 m N 26◦58′26″,
E 100◦10′47″ 22 August 2020
15 MHKMU M Mu 791 China: Yunnan mixed forests 3240 m N 27◦23′14″,
E 100◦06′19″ 26 August 2020
16 MHKMU SD Yang 378 China: Yunnan mixed forests 2130 m N 25◦36′51″,
E 102◦09′23″ 20 August 2016
17 MHKMU SD Yang 557 China: Yunnan broad‑leaved
forests 2205 m N 24◦39′27″,
E 104◦10′17″ 24 August 2018
18 MHKMU TJ Yu 197 China: Yunnan mixed forests 3116 m N 27◦42′34″,
E 100◦31′30″ 2 October 2023
19 MHKMU TJ Yu 206 China: Yunnan mixed forests 3468 m N 27◦43′21″,
E 100◦31′54″ 2 October 2023
20 MHKMU WH Zhang 599 China: Yunnan broad‑leaved
forests 2500 m N 26◦23′41″,
E 99◦50′10″ 5 October 2020
21 MHKMU WH Zhang 606 China: Yunnan broad‑leaved
forests 2954 m N 26◦17′52″,
E 99◦46′09″ 6 October 2020
22 MHKMU WH Zhang
606‑1 China: Yunnan broad‑leaved
forests 2954 m N 26◦17′52″,
E 99◦46′09″ 6 October 2020
23 MHKMU X Na 149 China: Yunnan mixed forests 3194 m N 26◦41′52″,
E 100◦02′09″ 21 August 2020
24 MHKMU X Xia 129 China: Yunnan mixed forests 3468 m N 27◦43′21″,
E 100◦31′54″ 2 October 2023
25 MHKMU YJ Pu 361 China: Yunnan mixed forests 3180 m N 26◦58′26″,
E 100◦10′47″ 23 August 2020
26 MHKMU YR Li 004 China: Yunnan mixed forests 2090 m N 25◦36′51″,
E 102◦09′23″ 20 August 2016
J. Fungi 2024, 10, x FOR PEER REVIEW 3 of 26
18 MHKMU TJ Yu 197 China: Yunnan mixed forests 3116 m N 27°42′34″, E 100°31′30″ 2 October 2023
19 MHKMU TJ Yu 206 China: Yunnan mixed forests 3468 m N 27°43′21″, E 100°31′54″ 2 October 2023
20 MHKMU WH Zhang 599 China: Yunnan broad-leaved forests 2500 m N 26°23′41″, E 99°50′10″ 5 October 2020
21 MHKMU WH Zhang 606 China: Yunnan broad-leaved forests 2954 m N 26°17′52″, E 99°46′09″ 6 October 2020
22 MHKMU WH Zhang 606-1 China: Yunnan broad-leaved forests 2954 m N 26°17′52″, E 99°46′09″ 6 October 2020
23 MHKMU X Na 149 China: Yunnan mixed forests 3194 m N 26°41′52″, E 100°02′09″ 21 August 2020
24 MHKMU X Xia 129 China: Yunnan mixed forests 3468 m N 27°43′21″, E 100°31′54″ 2 October 2023
25 MHKMU YJ Pu 361 China: Yunnan mixed forests 3180 m N 26°58′26″, E 100°10′47″ 23 August 2020
26 MHKMU YR Li 004 China: Yunnan mixed forests 2090 m N 25°36′51″, E 102°09′23″ 20 August 2016
Figure 1. Collecting locations of specimens in China (Numbers correspond to the specimen voucher
available in Table 1).
2.2. Morphological Studies
The macroscopic morphology was described based on field records and photographs
of basidiomata and color codes referred to by Kornerup and Wanscher [35]. The size of
basidiomata is referred to by Bas C [36]. Micro-morphological structures were observed
under a Leica DM2500 light microscope (Leica Microsystems, Welar, Germany) from
dried materials, which were mounted in 5% potassium hydroxide (KOH) and stained with
1% Congo Red solution (w/v) when necessary. Melzer’s reagent was used to examine the
amyloidity of basidiospores. In this paper, [n/m/p] refers to ‘n’ basidiospores measured
from ‘m’ basidiomata of ‘p’ collections; ‘(a) b–c (d)’ refers to the length and width of ba-
sidiospores, ‘a’ and ‘d’ refer to the minimum and maximum values of measured values,
and ‘b–c’ refers to the distribution interval of 90% of measured values; ‘Q’ means the
length/width ratio of a basidiospore; and ‘Qm’ refers to mean ± standard deviations of Q
values of all basidiospores.
2.3. DNA Extraction, PCR Amplification and Sequencing
Total genomic DNA was extracted from ca. 20 mg of dried basidiomata tissue using
the modified CTAB method [37]. Three DNA loci, nuclear ribosomal DNA internal tran-
scribed space (ITS) regions, large subunit nuclear ribosomal RNA (nrLSU), and translation
elongation factor 1 (tef1) were amplified by polymerase chain reaction (PCR) using primer
pairs ITS5/ITS4, LR0R/LR5, 983F/1567R, and HEF1F/HEF1R, respectively [19,38–40].
The PCR amplification reaction system consisted of 12.5 µL 2 × Taq PCR Master mix
(Biomed, Beijing, China), 1 µL of each primer (5 µM), 1 µL template DNA, and then was
refilled to 25 µL with sterilized, double-steamed water. The PCR conditions followed the
description of Tang et al. [41], which were as follows: pre-denaturation at 94 °C for 5 min;
Figure 1. Collecting locations of specimens in China (Numbers correspond to the specimen voucher
available in Table 1).
2.2. Morphological Studies
The macroscopic morphology was described based on eld records and photographs
of basidiomata and color codes referred to by Kornerup and Wanscher [35]. The size of
basidiomata is referred to by Bas C [36]. Micro‑morphological structures were observed
under a Leica DM2500 light microscope (Leica Microsystems, Welar, Germany) from
dried materials, which were mounted in 5% potassium hydroxide (KOH) and stained with
J. Fungi 2024,10, 824 4 of 25
1% Congo Red solution (w/v) when necessary. Melzer’s reagent was used to examine the
amyloidity of basidiospores. In this paper, [n/m/p] refers to ‘n’ basidiospores measured
from ‘m’ basidiomata of ‘p’ collections; ‘(a) b–c (d)’ refers to the length and width of ba‑
sidiospores, ‘a’ and ‘d’ refer to the minimum and maximum values of measured values,
and ‘b–c’ refers to the distribution interval of 90% of measured values; ‘Q’ means the
length/width ratio of a basidiospore; and ‘Qm’ refers to mean ±standard deviations of
Q values of all basidiospores.
2.3. DNA Extraction, PCR Amplication and Sequencing
Total genomic DNA was extracted from ca. 20 mg of dried basidiomata tissue using
the modied CTAB method [37]. Three DNA loci, nuclear ribosomal DNA internal tran‑
scribed space (ITS) regions, large subunit nuclear ribosomal RNA (nrLSU), and translation
elongation factor 1 (tef1) were amplied by polymerase chain reaction (PCR) using primer
pairs ITS5/ITS4, LR0R/LR5, 983F/1567R, and HEF1F/HEF1R, respectively [19,38–40].
The PCR amplication reaction system consisted of 12.5 µL 2 ×Taq PCR Master mix
(Biomed, Beijing, China), 1 µL of each primer (5 µM), 1 µL template DNA, and then was
relled to 25 µL with sterilized, double‑steamed water. The PCR conditions followed the
description of Tang et al. [41], which were as follows: pre‑denaturation at 94 ◦C for 5 min;
35 cycles of denaturation at 94 ◦C for 40 s, annealing at 56 ◦C for 40 s, and extension at
72 ◦C for 1 min; nally, it was extended at 72 ◦C for 10 min. PCR products were examined
by electrophoresis on 1% agarose gels. The amplied PCR products were sequenced in
single direction using an ABI 3730 DNA Analyzer (Sangon, Shanghai, China) with the
same primers.
2.4. Phylogenetic Analyses
Newly generated sequences in this study were combined with those downloaded
from GenBank/UNITE for phylogenetic construction according to the previous stud‑
ies [18–21,26,31,33]. Information about sequences is presented in Table 2. The sequence
matrix of ITS, nrLSU, and tef1 was separately aligned with MAFFT v7.490 and manually
optimized on BioEdit v7.0.9 where necessary [42,43]. All positions with less than 50% site
coverage were eliminated, i.e., fewer than 50% alignment gaps, missing data, and ambigu‑
ous bases (partial deletion option).
ITS‑nrLSU‑tef1 and ITS‑nrLSU datasets were analyzed using Maximum Likelihood
(ML) and Bayesian Inference (BI) methods, respectively. ML analyses were implemented
with RAxML‑HPC v8.2.10, and 1000 rapid bootstrap replicates were performed; GTRGAMMA
was set by default as the selected model [44]. For BI analyses, the most appropriate sub‑
stitution models were selected by MrModeltest v2.3 under the Akaike information cri‑
terion (AIC) [45]. BI analyses were conducted using MrBayes v3.2.7 [46]. For Hydnum,
General Time Reversible + Proportion of Invariable Sites + Gamma (GTR + I + G) for ITS
and nrLSU, GTR + G for tef1; four Markov chains were run for 5,000,000 generations until
split deviation frequency value was <0.01, sampling every 100th generation. For Lentaria
s.l., GTR + I + G for ITS, Symmetrical Model + Proportion of Invariable Sites + Gamma
(SYM + I + G) for nrLSU; four Markov chains were run for 3,000,000 generations until split
deviation frequency value was <0.01, sampling every 1000th generation. Then, the rst
25% of sampled trees from generations were discarded as burn‑in, and Bayesian posterior
probabilities (PP) were then calculated for a majority consensus tree of retained Bayesian
trees. All trees were visualized in FigTree v1.4.2.
Table 2. Specimens and sequences used in this study.
Taxa Voucher Locality GenBank/UNITE Accession Nos. References
ITS nrLSU tef1
Clavariadelphus
khinganensis MHKMU HY Huang 368 NE China: Jilin MT447468 — — [12]
Hydnum alboaurantiacum TENN 073053 (T) USA MH379955 — — [23]
J. Fungi 2024,10, 824 5 of 25
Table 2. Cont.
Taxa Voucher Locality GenBank/UNITE Accession Nos. References
ITS nrLSU tef1
H. alboaurantiacum TENN 073051 USA MH379937 — — [23]
H. alboluteum TUMH 63988 (T) Japan LC621802 — LC622439 [22]
H. alboluteum TUMH 63989 Japan LC621803 — — [22]
H. albomagnum AFTOL‑ID 471 USA DQ218305 AY700199 DQ234568 [47]
H. albomagnum TENN 073062 (T) USA MH379943 — — [23]
H. albopallidum TUMH 63997 (T) Japan LC621807 LC717904 LC622442 [22]
H. albopallidum TUMH 63998 Japan LC621808 — LC622443 [22]
H. berkeleyanum CAL 1656 (T) India NR_158533 NG_070500 — [24]
H. berkeleyanum HKAS77834 SW China: Yunnan KU612525 KU612667 — [19]
H. berkeleyanum IFP 019484 China MW980552 MW979538 — [18]
H. berkeleyanum MHKMU YJ Pu 361 SW China: Yunnan PQ287656 PQ287738 PQ295833 This study
H. berkeleyanum MHKMU M Mu 740 SW China: Yunnan PQ287657 PQ287739 PQ295834 This study
H. boreorepandum H 6003711 (T) Finland KX388657 — — [20]
H. boreorepandum TUMH 64005 Japan LC621814 LC717880 LC622449 [22,48]
H. boreorepandum (H.
repandum)HKAS54416 NE China: Jilin KU612583 — — [19]
H. brevispinum IFP 019464 (T) C China: Hunan MW980578 MW979559 — [18]
H. brevispinum IFP 019465 C China: Hunan MW980579 MW979560 — [18]
H. cremeum MHKMU TJ Yu 197 (T) SW China: Yunnan PQ287674 PQ287755 PQ295848 This study
H. cremeum MHKMU WH Zhang
599 SW China: Yunnan PQ287673 PQ287754 PQ295847 This study
H. cremeoalbum GDGM93011 C China: Hubei OR947110 OR947129 — [26]
H. cremeoalbum TUMH 60740 (T) Japan AB906678 — — [25]
H. cremeoalbum HKAS92345 SW China:
Chongqing KU612619 KU612676 KU612764 [19]
H. cremeoalbum FHMU1631 S China: Hainan OQ656784 OQ656792 — [21]
H. cremeoalbum (H.
albomagnum)IFP 019480 C China: Hunan MW980550 MW979536 — [18]
H. erectum FHMU7689 (T) E China: Zhejiang OR722666 OR722669 — [21]
H. abellatum IFP 019459 (T) NE China: Liaoning MW980575 MW979556 — [18]
H. avidocanum IFP 019460 (T) SW China: Yunnan MW980559 MW979545 MW999440 [18]
H. avidocanum IFP 019461 SW China: Yunnan MW980560 MW979546 MW999441 [18]
H. avosquamosum MHKMU LP Tang 3454
(T) SW China: Yunnan PQ287672 PQ287753 PQ295846 This study
H. ibericum BIO Fungi: 12330 (T) Spain HE611086 — — [28]
H. ibericum MA‑fungi 3457 Spain AJ547879 — — [49]
H. jussii H 6003709 (T) Finland KX388665 — — [20]
H. jussii IFP 019485 NW China: Xinjiang MW980553 MW979539 MW999436 [18]
H. khanspurense KH‑09 (LAN88021) (T) Pakistan OQ130694 — — [50]
H. khanspurense KH‑50 (LAN29722) Pakistan OQ130695 — — [50]
H. longibasidium IFP 019462 (T) C China: Hunan MW980556 MW979541 MW999438 [18]
H. longibasidium IFP 019463 C China: Hunan MW980555 MW979542 MW999439 [18]
H. longipes GDGM82458 (T) SW China: Yunnan OR947121 — — [26]
H. melitosarx GDGM84518 SW China: Sichuan OR947117 OR947136 — [26]
H. melitosarx H7043937 (T) USA KX388683 — — [20]
H. melitosarx (H.
rufescens)HKAS92338 NE China: Jilin KU612538 KU612659 KU612784 [19]
H. microcarpum GDGM87902 (T) S China:
Guangdong OR947116 OR947134 — [26]
H. microcarpum GDGM87902‑1 S China:
Guangdong OR947115 — — [26]
H. minum TUMH 60737 (T) Japan AB906675 — — [25]
H. minum IFP 019482 C China: Hunan MW980557 MW979543 — [18]
H. minum MHKMU YR Li 004 SW China: Yunnan PQ287659 PQ287741 — This study
H. minum MHKMU SD Yang 378 SW China: Yunnan PQ287658 PQ287740 — This study
H. minum FHMU2408 S China: Hainan OQ656785 OQ656793 — [21]
H. orientalbidum GDGM93480 SW China:
Chongqing OR947108 OR947127 — [26]
H. orientalbidum GDGM91301 E China: Zhejiang OR947111 OR947130 — [26]
H. orientalbidum TUMH 62998 (T) Japan LC377875 LC717908 LC622478 [48,51]
H. orientalbidum FHMU6327 S China: Hainan OQ656787 OQ656794 — [21]
H. pallidocroceum IFP 019466 (T) NW China: Xinjiang MW980568 MW979554 MW999449 [18]
H. pallidocroceum IFP 019467 NW China: Xinjiang MW980569 MW979555 MW999450 [18]
H. pallidomarginatum IFP 019468 (T) SW China: Yunnan MW980566 MW979552 MW999447 [18]
H. pallidomarginatum MHKMU HY Huang
873 SW China: Yunnan PQ287660 PQ287742 PQ295835 This study
H. pallidomarginatum MHKMU LP Tang 3319 SW China: Yunnan PQ287661 PQ287743 PQ295836 This study
H. pallidomarginatum MHKMU LP Tang 3453 SW China: Yunnan PQ287662 PQ287744 PQ295837 This study
H. pallidomarginatum MHKMU SD Yang 557 SW China: Yunnan PQ287663 PQ287745 PQ295838 This study
H. pallidomarginatum MHKMU M Mu 791 SW China: Yunnan PQ287664 PQ287746 PQ295839 This study
H. pallidomarginatum (H.
vesterholtii)HKAS92344 NE China:
Heilongjiang KU612556 KU612649 KU612788 [19]
H. pallidomarginatum (H.
vesterholtii)HKAS56213 SW China: Yunnan KU612554 — — [19]
H. pinicola GDGM93020 C China: Hubei OR947109 OR947128 — [26]
H. pinicola GDGM83047 SW China: Yunnan OR947119 OR947137 — [26]
H. pinicola TUMH 64004 (T) Japan LC621813 — LC622448 [22]
J. Fungi 2024,10, 824 6 of 25
Table 2. Cont.
Taxa Voucher Locality GenBank/UNITE Accession Nos. References
ITS nrLSU tef1
H. repandum 031209A Slovenia KU612574 KU612655 KU612770 [19]
H. repandum H 6003710 (T) Finland NR_164553 — — [20]
H. roseoalbum MHKMU WH Zhang
606 (T) SW China: Yunnan PQ287669 PQ287751 PQ295844 This study
H. roseoalbum MHKMU WH Zhang
606‑1 SW China: Yunnan PQ287670 PQ287752 PQ295845 This study
H. roseotangerinum MHKMU LP Tang 3458
(T) SW China: Yunnan PQ287675 PQ287756 PQ295849 This study
H. roseotangerinum MHKMU LP Tang
3458‑1 SW China: Yunnan PQ287676 PQ287757 PQ295850 This study
H. sinorepandum GDGM82445 (T) SW China: Yunnan OR947122 OR947139 — [26]
H. sinorepandum GDGM82382 SW China: Yunnan OR947124 OR947141 — [26]
Hydnum sp. HMJAU60215 China OM341393 — — GenBank
Hydnum sp. HMJAU60216 China OM341394 — — GenBank
Hydnum sp. 1 (H.
vesterholtii)HKAS92342 SW China: Yunnan KU612564 KU612646 KU612786 [19]
Hydnum sp. 1 (H.
vesterholtii)HKAS77884 C China: Hubei KU612565 KU612645 KU612787 [19]
Hydnum sp. 2 (H.
vesterholtii)HKAS92341 NW China: Shaanxi KU612562 KU612647 KU612790 [19]
Hydnum sp. 3 MHKMU LP Tang 2886 NE China: Jilin PQ287671 — — This study
Hydnum sp. 3 (H.
vesterholtii)HKAS92343 SW China: Sichuan KU612563 KU612648 — [19]
H. sphaericum IFP 019470 (T) C China: Hunan MW980563 MW979549 MW999444 [18]
H. sphaericum MHKMU X Na 149 SW China: Yunnan PQ287665 PQ287747 PQ295840 This study
H. subalpinum TUMH 64013 Japan LC717913 LC717888 LC717874 [48]
H. subalpinum TUMH 64016 (T) Japan LC621871 LC717891 LC622497 [22,48]
H. subtilior TENN 073050 USA MH379918 — — [23]
H. subtilior TENN 073034 (T) USA NR_164029 — — [23]
H. tangerinum IFP 019473 (T) C China: Hunan MW980580 MW979561 — [18]
H. tangerinum IFP 019474 C China: Hunan MW980581 MW979562 — [18]
H. tenuistipitum IFP 019476 (T) C China: Hunan MW980576 MW979557 — [18]
H. tenuistipitum FHMU7644 C China: Hunan OQ913759 OQ913756 — [21]
H. tenuistipitum MHKMU HY Huang
151 SW China: Yunnan PQ287666 PQ287748 PQ295841 This study
H. tenuistipitum MHKMU LP Tang 2116 SW China: Yunnan PQ287667 PQ287749 PQ295842 This study
H. tenuistipitum MHKMU J Zhao 225 SW China: Yunnan PQ287668 PQ287750 PQ295843 This study
H. tomaense TUMH 64086 (T) Japan LC621885 LC717907 LC622509 [22,48]
H. tomaense TUMH 64085 Japan LC621884 — LC622508 [22]
H. treui TU110403 (T) Papua New Guinea UDB01304 * — — [20,52]
H. treui FHMU7690 S China: Hainan OR722667 OR722670 — [21]
H. umbilicatum CORT: 012241 (T) USA MH379890 — — [23]
H. umbilicatum TUMH 64093 Japan LC621893 — — [22]
H. umbilicatum (H.
ellipsosporum)HMJAU5985 NE China: Jilin KU612602 — — [19]
H. ventricosum IFP 019478 (T) NE China: Liaoning MW980561 MW979547 MW999442 [18]
H. ventricosum IFP 019479 NE China: Liaoning MW980562 MW979548 — [18]
H. vesterholtii BIO Fungi: 12904 (T) France HE611087 — — [28]
H. zongolicense MEXU 26248 (T) Mexico KC152121 — — [20]
Lentaria albovinacea FO 46869 Unknown DQ071734 — — [53]
L. bambusina MHHNU 6794 (T) C China: Hunan KU870448 — — [31]
L. bambusina MHHNU 7302 C China: Hunan KU324496 — — [31]
L. bambusina MHKMU C Xu 071 E China: Anhui PQ287679 PQ287760 — This study
L. byssiseda TENN61159‑C2 USA: Tennessee FJ596785 — — [54]
L. byssiseda TENN61159‑C3 USA: Tennessee FJ596786 — — [54]
L. gossypina FCME 27625 Mexico: Campeche MK253199 MK253219 — [33]
L. gossypina FCME 27624 Mexico: Campeche MK253198 MK253218 — [33]
L. micheneri RRD6 USA: Tennessee MF773634 — — GenBank
L. micheneri iNaturalis 178123461 USA: Tennessee PP526130 — — GenBank
L. patouillardii MHHNU 7829 NW China: Xinjiang KU324498 — — [31]
L. patouillardii HMJAU 26892 NW China: Inner
Mongolia KU870449 — — [31]
L. patouillardii MA‑Fungi 48032 Spain AJ292290 — — GenBank
L. subalpina MHKMU TJ Yu 206 (T) SW China: Yunnan PQ287677 PQ287758 — This study
L. subalpina MHKMU X Xia 129 SW China: Yunnan PQ287678 PQ287759 — This study
L. surculus MHHNU 8721 SW China: Yunnan KU870450 — — [31]
L. surculus FHMU 880 S China:
Guangdong KU870451 — — [31]
L. surculus MHKMU LJ Su 430 S China: Hainan PQ287680 PQ287761 — This study
L. surculus MHKMU LJ Su 431 S China: Hainan PQ287681 PQ287762 — This study
L. uncispora MHHNU 7707 (T) SW China: Sichuan KU324497 — — [31]
L. variabilis FCME 21524 Mexico: Campeche MK253189 MK253215 — [33]
L. variabilis FCME 19864 Mexico: Campeche MK253185 MK253206 — [33]
Sistotrema muscicola KHL 11721 Finland AJ606040 AJ606040 — [55]
S. muscicola taxon: 154757 Finland AJ606041 AJ606041 — [55]
Newly generated sequences are highlighted in bold. ‘T’ refers to type specimens; * refers to sequence retrieved
from UNITE; wrong names in GenBank are put in brackets, e.g., H. pallidomarginatum (H. vesterholtii); C = Central,
E = Eastern, NE = Northeastern, NW = Northwestern, S = Southern, SW = Southwestern.
J. Fungi 2024,10, 824 7 of 25
3. Results
3.1. Molecular Phylogeny
3.1.1. Genus Hydnum
In the ITS‑nrLSU‑tef1 dataset, 105 ITS, 72 nrLSU, and 46 tef1 sequences were included,
including 59 newly generated and 164 retrieved ones from GenBank/UNITE, respectively.
Sistotrema muscicola (Pers.) S. Lundell was selected as an outgroup based on recent stud‑
ies [18,21]. The combined dataset includes 104 taxa with 2357 nucleotide sites, and the
alignment is available at TreeBase (ID: 31750). The topology of the phylogenetic trees ob‑
tained from the combined and the single‑locus matrices were essentially the same, and we
only show the results of the combined matrix here; the results of the single‑locus are de‑
tailed in the Supplementary Materials. The ITS‑nrLSU‑tef1 phylogenetic tree is shown in
Figure 2.
J. Fungi 2024, 10, x FOR PEER REVIEW 8 of 26
Figure 2. Phylogenetic tree of Hydnum species based on ITS-nrLSU-tef1 sequences. Bootstrap values
≥ 70% and Bayesian posterior probabilities values ≥ 0.95 are shown on branches (newly generated
sequences in bold and new species in red).
3.1.2. Genus Lentaria s.l.
In the ITS-nrLSU dataset, 34 sequences were included, including ten newly generated
and 24 retrieved ones from GenBank. Clavariadelphus khinganensis J. Zhao, L.P. Tang & P.
Zhang was used as the outgroup based on recent studies [12,31]. The combined dataset
includes 23 taxa with 1602 nucleotide sites, and the alignment is available at TreeBase (ID:
31752). The topology of the phylogenetic trees obtained from the combined and the single-
locus matrix were essentially the same, and we only show the results of the combined
matrix here; the results of the single-locus are detailed in the Supplementary Materials.
The phylogenetic tree is shown in Figure 3.
Our topologies obtained from the ITS-nrLSU dataset showed that the genus Lentaria
s.l. was a polyphyletic group consisting of two clades. This result was similar to the phylo-
grams in previous studies [31,33]. Lentaria subalpina sp. nov was closely related to L. bys-
siseda Corner and L. micheneri (Berk. & M.A. Curtis) Corner, while this new species had no
identified sister species in this study.
Figure 2. Phylogenetic tree of Hydnum species based on ITS‑nrLSU‑tef1 sequences. Bootstrap
values ≥70% and Bayesian posterior probabilities values ≥0.95 are shown on branches (newly gen‑
erated sequences in bold and new species in red).
The topologies obtained from the ITS‑nrLSU‑tef1 datasets in our study were similar
to the phylograms from previous studies [18,21,22,26]. Newly generated sequences were
classied into ten clades with high support values (BS/BPP ≥90/0.95), including four new
J. Fungi 2024,10, 824 8 of 25
species. Among these taxa, H. cremeum sp. nov from China formed a distinct lineage
(BS/BPP = 100/1) in the subgen. Pallida, and it was a sister species of H. albopallidum R.
Sugaw. & N. Endo from Japan (BS/BPP = 60/0.96); H. avosquamosum sp. nov formed a
distinct lineage in the subgen. Pallida, but there was no resolution among the closely re‑
lated phylogenetic species; H. roseoalbum sp. nov was grouped in the subgen. Alba. s. lato
(BS/BPP = 100/1); Hydnum roseotangerinum sp. nov from China in the subgen. Rufescentia
was closely related to two Asian species, H. berkeleyanum K. Das, Hembrom, A. Baghela &
Vizzini and H. ventricosum T. Cao & H. S. Yuan.
3.1.2. Genus Lentaria s.l.
In the ITS‑nrLSU dataset, 34 sequences were included, including ten newly generated
and 24 retrieved ones from GenBank. Clavariadelphus khinganensis J. Zhao, L.P. Tang & P.
Zhang was used as the outgroup based on recent studies [12,31]. The combined dataset
includes 23 taxa with 1602 nucleotide sites, and the alignment is available at TreeBase (ID:
31752). The topology of the phylogenetic trees obtained from the combined and the single‑
locus matrix were essentially the same, and we only show the results of the combined
matrix here; the results of the single‑locus are detailed in the Supplementary Materials.
The phylogenetic tree is shown in Figure 3.
J. Fungi 2024, 10, x FOR PEER REVIEW 9 of 26
Figure 3. Phylogenetic tree of Lentaria s.l. species based on ITS-nrLSU sequences. Bootstrap values
≥ 70% and Bayesian posterior probabilities values ≥ 0.95 are shown on branches (newly generated
sequences in bold and new species in red).
3.2. Taxonomy
Hydnum berkeleyanum K. Das, Hembrom, A. Baghela & Vizzini. Cryptogam. Mycol.
39(2): 212, 2018 (Figure 4a–f)
Chinese name—贝氏齿菌 [29]
MycoBank—MB 824501
Description—Basidiomata small-sized. Pileus 4–5 cm in diam., plano-convex to plane,
irregular round, sometimes with irregular lobes or umbilicate in the center; surface dry,
irregularly bumpy, non-staining, azonate, pale orange when young, becoming yellowish
brown to orange-brown when mature, paler towards margin; margin entire, incurved.
Context 0.5–0.7 cm thick in pileal center, fleshy, and warm cream to yellowish white. Stipe
6 cm long, 0.9–1.6 cm in diam., central to eccentric, cylindrical, white to pale orange-
brown, solid interior, the base often inflated. Spines 0.4–0.5 cm long (with some shorter
spines on stipe), adnexed to decurrent, not flaened, with an acute (rarely subacute) apex,
crowded, creamy, gradually more yellowish when drying. Odor and Taste not recorded.
Basidiospores [40/2/2] 8.1–9.0 (–9.5) × 8.1–9.0 (–9.5) µm, Q = 1.00–1.06 (–1.11), Qm = 1.02
± 0.03, globose, sometimes subglobose, thin-walled, inamyloid. Basidia 31–64 (–82) × 9.5–
14 µm, subcylindric to clavate, 2- to 4-spored, clamped; sterigmata 2.5–8.5 µm long. Basid-
ioles numerous, subcylindrical or subclavate, smaller than basidia. Cystidia absent. Spine
tip sterile, composed of interwoven hyphae (3.5–11 µm wide), septate, clamped, apex sub-
clavate to clavate, becoming parallel towards the apex. Hymenophoral trama composed of
hyphae (2.5–6 µm wide), clamped, in subparallel to interwoven paern. Pileipellis com-
posed of densely interwoven, suberect hyphae (5–14 µm wide), cylindrical, thin-walled,
clamped, the terminal cell rounded at the apex, with pale yellow intracellular pigment.
Clamp connections: present.
Habitat—Solitary or concrescent on the ground of temperate to subalpine mixed for-
ests dominated by Ericaceae, Fagaceae, and Pinaceae.
Specimens examined—CHINA. Yunnan Province, Shuhe Ancient Town (束河古镇),
N 26°58′26″, E 100°10′47″, elev. 3180 m, 22 August 2020, Mu Man 740 (MHKMU M Mu
740); about 3 km from Alpine Botanical Garden (高山植物园), N 26°58′26″, E 100°10′47″,
elev. 3180 m, 22 August 2020, on the ground of subalpine mixed forests dominated by
Ericaceae, Fagaceae, and Pinaceae, Pu Yunju 361 (MHKMU YJ Pu 361).
Known distribution—Asia: Yunnan Province, China, elev. above 3000 m
[18,19,24,26]; India (type location), elev. 1800–2200 m [24]; and Pakistan, elev. 2200 m–2500
m [50].
Figure 3. Phylogenetic tree of Lentaria s.l. species based on ITS‑nrLSU sequences. Bootstrap
values ≥70% and Bayesian posterior probabilities values ≥0.95 are shown on branches (newly gen‑
erated sequences in bold and new species in red).
Our topologies obtained from the ITS‑nrLSU dataset showed that the genus Lentaria
s.l. was a polyphyletic group consisting of two clades. This result was similar to the phy‑
lograms in previous studies [31,33]. Lentaria subalpina sp. nov was closely related to L.
byssiseda Corner and L. micheneri (Berk. & M.A. Curtis) Corner, while this new species had
no identied sister species in this study.
3.2. Taxonomy
Hydnum berkeleyanum K. Das, Hembrom, A. Baghela & Vizzini. Cryptogam. Mycol.
39(2): 212, 2018 (Figure 4a–f)
Chinese name—贝氏齿菌 [29]
MycoBank—MB 824501
Description—Basidiomata small‑sized. Pileus 4–5 cm in diam., plano‑convex to plane,
irregular round, sometimes with irregular lobes or umbilicate in the center; surface dry,
irregularly bumpy, non‑staining, azonate, pale orange when young, becoming yellowish
brown to orange‑brown when mature, paler towards margin; margin entire, incurved. Con‑
text 0.5–0.7 cm thick in pileal center, eshy, and warm cream to yellowish white. Stipe 6
J. Fungi 2024,10, 824 9 of 25
cm long, 0.9–1.6 cm in diam., central to eccentric, cylindrical, white to pale orange‑brown,
solid interior, the base often inated. Spines 0.4–0.5 cm long (with some shorter spines on
stipe), adnexed to decurrent, not aened, with an acute (rarely subacute) apex, crowded,
creamy, gradually more yellowish when drying. Odor and Taste not recorded.
J. Fungi 2024, 10, x FOR PEER REVIEW 10 of 26
Notes—Hydnum berkeleyanum is characterized by its pale orange to orange-brown
colored pileus, 2- to 4-spored basidia, and globose basidiospores. It was described from
India, with reports also found in Pakistan and China [18,24,26,50]. We examined collec-
tions from Yunnan and several DNA sequences from China labeled as H. berkeleyanum in
the database. No major differences were found between our specimen and the type, except
for the size of basidiomata. Additionally, Qin et al. [21] mistakenly recorded the collecting
site of HKAS77834 as Hunan Province, China. After reviewing the literature, we deter-
mined that this specimen was taken from Yunnan Province rather than Hunan [19]. Thus,
the known distribution of H. berkeleyanum in China is limited to the southwest rather than
the central regions.
Figure 4. Macroscopic and microscopic features of Hydnum berkeleyanum. (a–c) Basidiomata, (a) from
MHKMU YJ Pu 361; (b,c) from MHKMU M Mu 740; (d) Basidiospores; (e) Basidia and Basidioles;
(f) Pileipellis. Bars (d) = 10 µm; (e) = 20 µm; (f) = 50 µm.
Hydnum cremeum L.P. Tang, L.J. Su & T.J. Yu sp. nov. (Figure 5a–f)
Chinese name—奶油齿菌
MycoBank—MB 855827
Etymology—Latin ‘cremeum’ = cream, refers to the color of basidiomata.
Diagnosis—Distinct from other species within Hydnum mainly by its small basidio-
mata with a warm cream to yellowish white pileus, globose to subglobose basidiospores,
and occurrence in subalpine mixed forests.
Holotype—CHINA. Yunnan Province, Ninglang County (宁蒗县), Gewa Village (格
瓦村), N 27°42′34″, E 100°31′′30″, elev. 3116 m, 2 October 2023, on the ground of subalpine
mixed forests dominated by Pinus yunnanensis Franch., P. densata Mast., Quercus L., and
Rhododendron decorum Franch., Yu Taijie 197 (MHKMU TJ Yu 197). GenBank accession
numbers: ITS = PQ287674, nrLSU = PQ287755, tef1 = PQ295848.
Description—Basidiomata very small- to small-sized. Pileus 2.5–3.7 cm in diam., plano-
convex to plane, irregular round, central sometimes depress, shallow infundibuliform
when old; surface dry, irregularly bumpy, non-staining, azonate, warm cream (3A1) to
yellowish white (3–4A2), with light yellow (5A5) to light grayish-orange (5B4–5) patches;
margin entire, incurved. Context 0.2–0.3 cm thick in pileal center, fleshy, warm cream,
slightly becoming yellowish white to pale orange on exposure. Stipe 3–6 cm long, 0.5–1
cm in diam., central, eccentric to lateral, cylindrical, concolorous with the pileus, solid
interior. Spines 0.4–0.5 cm long (with some shorter spines on stipe), adnexed to slightly
decurrent, not flaened, with an acute (rarely subacute) apex, crowded, cream, gradually
more yellow when drying. Odor and Tas te not recorded.
Figure 4. Macroscopic and microscopic features of Hydnum berkeleyanum. (a–c) Basidiomata, (a) from
MHKMU YJ Pu 361; (b,c) from MHKMU M Mu 740; (d) Basidiospores; (e) Basidia and Basidioles;
(f) Pileipellis. Bars (d) = 10 µm; (e) = 20 µm; (f) = 50 µm.
Basidiospores[40/2/2] 8.1–9.0 (–9.5) ×8.1–9.0 (–9.5) µm, Q = 1.00–1.06 (–1.11), Qm = 1.02 ±0.03,
globose, sometimes subglobose, thin‑walled, inamyloid. Basidia 31–64 (–82) ×9.5–14 µm,
subcylindric to clavate, 2‑ to 4‑spored, clamped; sterigmata 2.5–8.5 µm long. Basidioles nu‑
merous, subcylindrical or subclavate, smaller than basidia. Cystidia absent. Spine tip sterile,
composed of interwoven hyphae (3.5–11 µm wide), septate, clamped, apex subclavate to
clavate, becoming parallel towards the apex. Hymenophoral trama composed of hyphae
(2.5–6 µm wide), clamped, in subparallel to interwoven paern. Pileipellis composed of
densely interwoven, suberect hyphae (5–14 µm wide), cylindrical, thin‑walled, clamped,
the terminal cell rounded at the apex, with pale yellow intracellular pigment. Clamp con‑
nections: present.
Habitat—Solitary or concrescent on the ground of temperate to subalpine mixed forests
dominated by Ericaceae, Fagaceae, and Pinaceae.
Specimens examined—CHINA. Yunnan Province, Shuhe Ancient Town (束河古镇),
N 26◦58′26″, E 100◦10′47″, elev. 3180 m, 22 August 2020, Mu Man 740 (MHKMU M Mu
740); about 3 km from Alpine Botanical Garden (高山植物园), N 26◦58′26″, E 100◦10′47″,
elev. 3180 m, 22 August 2020, on the ground of subalpine mixed forests dominated by
Ericaceae, Fagaceae, and Pinaceae, Pu Yunju 361 (MHKMU YJ Pu 361).
Known distribution—Asia: Yunnan Province, China, elev. above 3000 m [18,19,24,26];
India (type location), elev. 1800–2200 m [24]; and Pakistan, elev. 2200 m–2500 m [50].
Notes—Hydnum berkeleyanum is characterized by its pale orange to orange‑brown col‑
ored pileus, 2‑ to 4‑spored basidia, and globose basidiospores. It was described from In‑
dia, with reports also found in Pakistan and China [18,24,26,50]. We examined collections
from Yunnan and several DNA sequences from China labeled as H. berkeleyanum in the
database. No major dierences were found between our specimen and the type, except for
the size of basidiomata. Additionally, Qin et al. [21] mistakenly recorded the collecting site
of HKAS77834 as Hunan Province, China. After reviewing the literature, we determined
that this specimen was taken from Yunnan Province rather than Hunan [19]. Thus, the
known distribution of H. berkeleyanum in China is limited to the southwest rather than the
central regions.
Hydnum cremeum L.P. Tang, L.J. Su & T.J. Yu sp. nov. (Figure 5a–f)
J. Fungi 2024,10, 824 10 of 25
J. Fungi 2024, 10, x FOR PEER REVIEW 11 of 26
Basidiospores [80/3/2] 8.1–9.5 × 7.1–9.0 (–9.5) µm, Q = 1.00–1.11 (–1.13), Qm = 1.05 ±
0.04, globose to subglobose, thin-walled, inamyloid. Basidia 40–57 × 9.5–13 µm, subcylin-
dric to clavate, 2- to 4-spored, clamped; sterigmata 5–9.5 µm long. Basidioles numerous,
subcylindrical or subclavate, smaller than basidia. Cystidia absent. Spine tip sterile, com-
posed of interwoven hyphae (4–11.5 µm wide), septate, clamped, apex subclavate to cla-
vate, becoming parallel towards the apex. Hymenophoral trama composed of hyphae (2.5–
6 µm wide), clamped, in subparallel to interwoven paern. Pileipellis composed of densely
interwoven, subparallel to suberect hyphae (3.5–9.5 µm wide), cylindrical, thin-walled,
clamped, the terminal cell rounded at the apex, with pale yellow intracellular pigment.
Clamp connections present.
Habitat—Solitary or concrescent on the ground of subtropical to subalpine mixed
forests dominated by Ericaceae, Fagaceae, and Pinaceae.
Additional specimens examined—CHINA. Yunnan Province, Jianchuan County (剑
川县), Shibaoshan Scenic Spot (石宝山景区), N 26°23′41″, E 99°50′10″, elev. 2500 m, 5 Oc-
tober 2020, on the ground of subtropical broad-leaved forests dominated by Fagaceae and
Ericaceae, Zhang Wenhao 599 (MHKMU WH Zhang 599).
Known distribution—China: Yunnan Province, elev. above 2500 m.
Notes—Hydnum cremeum is easily distinguished by its small-sized, cream basidio-
mata, with light yellow patches, sometimes a shallow infundibuliform pileus, 2–4-spored
basidia, and globose to subglobose basidiospores. In China, H. cremeum is often misiden-
tified with H. orientalbidum R. Sugaw. & N. Endo, H. pallidomarginatum T. C ao & H.S. Y uan,
and H. pinicola R. Sugaw. & N. Endo. However, H. orientalbidum has slightly larger basid-
iomata (pileus up to 5.5 cm in diam.), 3–5 (7)-spored basidia, and smaller basidiospores
(4.5–6 × 4–5 µm) [22]; H. pallidomarginatum has a zonate, orange white to pale orange pi-
leus, concolorous spines, 2–4-spored basidia, and longer basidiospores (8.2–9.8 × 6.5–7.8
µm) [18]; H. pinicola has pale orange spines, 4–8-spored basidia, and smaller basidiospores
(4.5–5.5 × 4–5 µm) [22,26]. Furthermore, H. sinorepandum Ming Zhang & C.Q. Wang and
H. subalpinum R. Sugaw. & N. Endo are also distributed in the subalpine regions of Asia
and have a creamy to yellowish white pileus. In contrast, H. sinorepandum has large basid-
iomata (pileus up to 12 cm in diam) with a light orange tinge, 4–8-spored basidia, and
broadly ellipsoid basidiospores [26]; H. subalpinum has robust basidiomata, 3–5-spored
basidia, relatively narrower basidiospores (Q = 1.03–1.23), and occurrence in coniferous
forests in Japan [22,26,48]. Additionally, H. cremeum is sister to H. albopallidum, but the
laer has an umbilicate pileus, (2) 3–4-spored basidia, narrower basidiospores (8–9.5 × 7–
8 µm), and is only found in Japan to date [22].
Figure 5. Macroscopic and microscopic features of Hydnum cremeum. (a–c) Basidiomata, (a,c) from
MHKMU TJ Yu 197, Holotype; (b) from MHKMU WH Zhang 599; (d) Basidiospores; (e) Basidia and
Basidioles; (f) Pileipellis. Bars (d) = 10 µm; (e) = 20 µm; (f) = 50 µm.
Chinese name—奶油齿菌
MycoBank—MB 855827
Etymology—Latin ‘cremeum’ = cream, refers to the color of basidiomata.
Diagnosis—Distinct from other species within Hydnum mainly by its small basid‑
iomata with a warm cream to yellowish white pileus, globose to subglobose basidiospores,
and occurrence in subalpine mixed forests.
Holotype—CHINA. YunnanProvince, Ninglang County (宁蒗县), Gewa Village (格瓦村),
N 27◦42′34″, E 100◦31′ ′30″, elev. 3116 m, 2 October 2023, on the ground of subalpine mixed
forests dominated by Pinus yunnanensis Franch., P. densata Mast., Quercus L., and Rhododen‑
dron decorum Franch., Yu Taijie 197 (MHKMU TJ Yu 197). GenBank accession numbers: ITS
= PQ287674, nrLSU = PQ287755, tef1 = PQ295848.
Description—Basidiomata very small‑ to small‑sized. Pileus 2.5–3.7 cm in diam., plano‑
convex to plane, irregular round, central sometimes depress, shallow infundibuliform
when old; surface dry, irregularly bumpy, non‑staining, azonate, warm cream (3A1) to
yellowish white (3–4A2), with light yellow (5A5) to light grayish‑orange (5B4–5) patches;
margin entire, incurved. Context 0.2–0.3 cm thick in pileal center, eshy, warm cream,
slightly becoming yellowish white to pale orange on exposure. Stipe 3–6 cm long, 0.5–1 cm
in diam., central, eccentric to lateral, cylindrical, concolorous with the pileus, solid interior.
Spines 0.4–0.5 cm long (with some shorter spines on stipe), adnexed to slightly decurrent,
not aened, with an acute (rarely subacute) apex, crowded, cream, gradually more yel‑
low when drying. Odor and Taste not recorded.
Basidiospores [80/3/2] 8.1–9.5 ×7.1–9.0 (–9.5) µm, Q = 1.00–1.11 (–1.13), Qm = 1.05 ±0.04,
globose to subglobose, thin‑walled, inamyloid. Basidia 40–57 ×9.5–13 µm, subcylindric to
clavate, 2‑ to 4‑spored, clamped; sterigmata 5–9.5 µm long. Basidioles numerous, subcylin‑
drical or subclavate, smaller than basidia. Cystidia absent. Spine tip sterile, composed of
interwoven hyphae (4–11.5 µm wide), septate, clamped, apex subclavate to clavate, becom‑
ing parallel towards the apex. Hymenophoral trama composed of hyphae (2.5–6 µm wide),
clamped, in subparallel to interwoven paern. Pileipellis composed of densely interwoven,
subparallel to suberect hyphae (3.5–9.5 µm wide), cylindrical, thin‑walled, clamped, the
terminal cell rounded at the apex, with pale yellow intracellular pigment. Clamp connec‑
tions present.
Habitat—Solitary or concrescent on the ground of subtropical to subalpine mixed
forests dominated by Ericaceae, Fagaceae, and Pinaceae.
J. Fungi 2024,10, 824 11 of 25
Additional specimens examined—CHINA. Yunnan Province, Jianchuan County (剑川县),
Shibaoshan Scenic Spot (石宝山景区), N 26◦23′41″, E 99◦50′10″, elev. 2500 m, 5 October
2020, on the ground of subtropical broad‑leaved forests dominated by Fagaceae and Eri‑
caceae, Zhang Wenhao 599 (MHKMU WH Zhang 599).
Known distribution—China: Yunnan Province, elev. above 2500 m.
Notes—Hydnum cremeum is easily distinguished by its small‑sized, cream basidiomata,
with light yellow patches, sometimes a shallow infundibuliform pileus, 2–4‑spored basidia,
and globose to subglobose basidiospores. In China, H. cremeum is often misidentied with H.
orientalbidum R. Sugaw. & N. Endo, H. pallidomarginatum T. Cao & H.S. Yuan, and H. pinicola
R. Sugaw. & N. Endo. However, H. orientalbidum has slightly larger basidiomata (pileus up
to 5.5 cm in diam.), 3–5 (7)‑spored basidia, and smaller basidiospores (4.5–6 ×4–5 µm) [22];
H. pallidomarginatum has a zonate, orange white to pale orange pileus, concolorous spines, 2–
4‑spored basidia, and longer basidiospores (8.2–9.8 ×6.5–7.8 µm) [18]; H. pinicola has pale
orange spines, 4–8‑spored basidia, and smaller basidiospores (4.5–5.5 ×4–5 µm) [22,26].
Furthermore, H. sinorepandum Ming Zhang & C.Q. Wang and H. subalpinum R. Sugaw. &
N. Endo are also distributed in the subalpine regions of Asia and have a creamy to yel‑
lowish white pileus. In contrast, H. sinorepandum has large basidiomata (pileus up to
12 cm in diam) with a light orange tinge, 4–8‑spored basidia, and broadly ellipsoid ba‑
sidiospores [26]; H. subalpinum has robust basidiomata, 3–5‑spored basidia, relatively nar‑
rower basidiospores (Q = 1.03–1.23), and occurrence in coniferous forests in Japan [22,26,48].
Additionally, H. cremeum is sister to H. albopallidum, but the laer has an umbilicate pileus,
(2) 3–4‑spored basidia, narrower basidiospores (8–9.5 ×7–8 µm), and is only found in
Japan to date [22].
Hydnum avosquamosum L.P. Tang & L.J. Su sp. nov. (Figure 6a–f)
J. Fungi 2024, 10, x FOR PEER REVIEW 13 of 26
related to species in the subgen. Pallida phylogenetically. But species in the subgen. Pallida
always have paler basidiomata and mostly ovoid to broadly ellipsoid basidiospores (Q >
1.25) [18,20,23].
Figure 6. Macroscopic and microscopic features of Hydnum flavosquamosum from MHKMU LP Tang
3454 (Holotype). (a–c) Basidiomata; (d) Basidiospores; (e) Basidia and Basidioles; (f) Pileipellis. Bars
(d) = 10 µm; (e) = 20 µm; (f) = 50 µm.
Hydnum pallidomarginatum T. Cao & H.S. Yuan, Stud. Mycol. 99: 100121, 2021 (Fig-
ure 7a–f)
Synonym—Hydnum flabellatum T. Cao & H.S. Yuan, Stud. Mycol. 99: 100121, 2021,
syn. nov.
Chinese name—淡缘齿菌
MycoBank—MB 839419
Description—Morphological characteristics are described in detail by Cao et al. [18].
Specimens examined—CHINA. Yunnan Province, Shizong County (师宗县), N
24°39′27′′, E 104°10′17′′, elev. 2205 m, 24 August 2018, on the ground of subtropical broad-
leaved forests dominated by Ericaceae, Fagaceae, and Pinaceae, Yang Shuda 557
(MHKMU SD Yang 557); Yulong County (玉龙县), N 26°46′11′′, E 100°02′51′′, elev. 2890 m,
21 August 2020, on the ground of subalpine mixed forests dominated by Pinus yunnanen-
sis, Quercus semecarpifolia Sm., and Rhododendron spp., Tang Liping 3319 (MHKMU LP Tang
3319); Mianshaba (棉沙坝), N 27°21′16″, E 100°09′ 37″, elev. 2731 m, 24 August 2020, on the
ground of subalpine mixed forests dominated by P. yunnanensis, Quercus spp., and Rhodo-
dendron spp., Huang Hongyan 873 (MHKMU HY Huang 873); Yi Family Village (彝家村),
N 27°23′14″, E 100°06′ 31″, elev. 3170 m, 26 August 2020, on the ground of subalpine mixed
forests dominated by Picea likiangensis, Pinus armandi, P. densata, P. yunnanensis, Q. guyavi-
folia, and R. decorum, Tang Liping 3453 (MHKMU LP Tang 3453); in the same location, N
27°23′14″, E 100°06′19″, elev. 3240 m, 26 August 2020, on the ground of subalpine mixed
forests dominated by P. armandi, Quercus spp., and Rhododendron spp., Mu Man 791
(MHKMU M Mu 791)
Known distribution—China: Heilongjiang (this study), elev. unknown, Liaoning
[18], elev. unknown, and Yunnan Province [18], elev. 2200–3300 m (this study).
Notes—In 2021, H. flabellatum and H. pallidomarginatum were described from China
[18]. The laer has a pale orange pileus with a paler margin, sometimes infundibuliform,
decurrent spines, broadly ellipsoid basidiospores, and a distribution from temperate to
subtropical [18]. In our phylogenetic analyses, H. flabellatum is grouped with H. pallido-
marginatum in the subgen. Pallida, but there were no significant genetic differences. We
Figure 6. Macroscopic and microscopic features of Hydnum avosquamosum from MHKMU LP Tang
3454 (Holotype). (a–c) Basidiomata; (d) Basidiospores; (e) Basidia and Basidioles; (f) Pileipellis. Bars
(d) = 10 µm; (e) = 20 µm; (f) = 50 µm.
Chinese name—黄鳞齿菌
MycoBank—MB 855834
Etymology—Latin ‘avo’ = yellow, ‘squamosum’ = scaly, refers to the pileus covering
with yellowish brown scales.
Diagnosis—Distinct from other species within Hydnum mainly by its light yellow to
light brownish‑orange pileus with slightly dark scales, context becoming pale brownish on
exposure, adnexed, spines, subglobose to broadly ellipsoid (Q = 1.05–1.23) basidiospores,
and erectly arranged hyphae in pileipellis.
Holotype—CHINA. Yunnan Province, Yi Family Village (彝家村), N 27◦23′14″, E 100◦06′
31″, elev. 3170 m, 26 August 2020, on the ground of subalpine mixed forests dominated by
J. Fungi 2024,10, 824 12 of 25
Picea likiangensis (Franch.) E. Pri., Pinus armandi Franch., P. densata,Quercus guyavifolia
H. Lév., and Rhododendron decorum,Tang Liping 3454 (MHKMU LP Tang 3454). GenBank
accession numbers: ITS = PQ287672, nrLSU = PQ287753, tef1 = PQ295846.
Description—Basidiomata medium‑sized. Pileus 5 cm in diam., plano‑convex to plane,
easily cracked; surface dry, azonate, light yellow (4A4–5) to light brownish‑orange (5B4–5),
covered with light orange (5A5), brownish yellow (5C7), grayish orange (5B6) to light
brown (5D8) scales; margin split, incurved. Context 0.5 cm thick in pileal center, eshy,
cream to yellowish white, slightly becoming pale brownish on exposure, eventually turns
light reddish‑brown (6C6). Stipe 2 cm long, 0.9 cm in diam., eccentric to lateral, cylindri‑
cal, cream to light yellow, solid interior. Spines 0.5 cm long, adnexed, non‑decurrent, not
aened, with an acute apex, crowded, light yellow to light brownish‑orange, gradually
more yellow when drying. Odor and Taste not recorded.
Basidiospores [40/1/1] 8.6–9.5 (–10.0) ×7.6–8.6 (–9.0) µm, Q = (1.00–) 1.05–1.23 (–1.25),
Qm = 1.12 ±0.06, subglobose to broadly ellipsoid, rarely globose, thin‑walled, inamyloid.
Basidia 37–58 ×9.5–13 µm, subcylindric to clavate, 2‑ to 5‑spored, clamped; sterigmata
4–7.5 µm long. Basidioles numerous, subcylindrical or subclavate, smaller than basidia.
Cystidia absent. Spine tip sterile, composed of interwoven hyphae (5–10 µm wide), septate,
clamped, apex subclavate to clavate, becoming parallel towards the apex. Hymenophoral
trama composed of hyphae (2–6 µm wide), clamped, in subparallel to interwoven paern.
Pileipellis composed of densely interwoven, erect hyphae (3–7 µm wide), cylindrical to sub‑
fusiform, thin‑walled, clamped, the terminal cell rounded at the apex, with pale yellow
intracellular pigment. Clamp connections present.
Habitat—Solitary on the ground of subalpine mixed forests dominated by Ericaceae,
Fagaceae, and Pinaceae.
Known distribution—China: Yunnan Province, elev. about 3100 m.
Notes—Hydnum avosquamosum has a yellow to light brownish‑colored pileus covered
with slightly dark scales, non‑decurrent spines, 2–5‑spored basidia, and subglobose to broadly
ellipsoid basidiospores. It is similar to H. berkeleyanum,H. rufescens Pers., and H. subrufescens
Niskanen & Liimat, but they can be distinguished by their darker pileus. Additionally, H.
berkeleyanum has larger and more robust basidiomata (pileus up to 8 cm in diam.), a brown
pileal center, longer spines (9 mm), 2–4‑spored basidia, and distributing in temperate to sub‑
tropical regions [24,29]; H. rufescens has a deep reddish‑orange to brownish‑orange pileus,
without scales, 3–5‑spored basidia, smaller basidiospores (7.0–8.5 ×6.0–7.2 µm), and grows
in European mixed forests [20]; H. subrufescens has a brownish ochraceous and non‑squamous
pileus, 3–4‑spored basidia, and smaller basidiospores (7.4–8.8 ×6.4–7.8 µm), with a distribu‑
tion limited to eastern North America [20]. Hydnum erectum N.K. Zeng, H.Z. Qin, W.F. Lin &
L.G. Hu also has erectly arranged hyphae in the pileipellis, but it has a creamy pileus, shorter
spines (1–2 mm long), 2–4‑spored basidia, and small basidiospores (6.5–8 ×5.5–7.5 µm) [21].
In addition, H. avosquamosum is closely related to species in the subgen. Pallida phylogenet‑
ically. But species in the subgen. Pallida always have paler basidiomata and mostly ovoid to
broadly ellipsoid basidiospores (Q > 1.25) [18,20,23].
Hydnum pallidomarginatum T. Cao & H.S. Yuan, Stud. Mycol. 99: 100121, 2021
(Figure 7a–f)
Synonym—Hydnum abellatum T. Cao & H.S. Yuan, Stud. Mycol. 99: 100121, 2021,
syn. nov.
Chinese name—淡缘齿菌
MycoBank—MB 839419
Description—Morphological characteristics are described in detail by Cao et al. [18].
Specimens examined—CHINA. Yunnan Province, Shizong County (师宗县), N 24◦39′27′′ ,
E 104◦10′17′′, elev. 2205 m, 24 August 2018, on the ground of subtropical broad‑leaved forests
dominated by Ericaceae, Fagaceae, and Pinaceae, Yang Shuda 557 (MHKMU SD Yang 557);
Yulong County (玉龙县), N 26◦46′11′′ , E 100◦02′51′′ , elev. 2890 m, 21 August 2020, on the
ground of subalpine mixed forests dominated by Pinus yunnanensis,Quercus semecarpifolia
Sm., and Rhododendron spp., Tang Liping 3319 (MHKMU LP Tang 3319); Mianshaba (棉沙坝),
J. Fungi 2024,10, 824 13 of 25
N 27◦21′16″, E 100◦09′37″, elev. 2731 m, 24 August 2020, on the ground of subalpine mixed
forests dominated by P. yunnanensis,Quercus spp., and Rhododendron spp., Huang Hongyan
873 (MHKMU HY Huang 873); Yi Family Village (彝家村), N 27◦23′14″, E 100◦06′31″, elev.
3170 m, 26 August 2020, on the ground of subalpine mixed forests dominated by Picea likian‑
gensis,Pinus armandi,P. densata,P. yunnanensis,Q. guyavifolia, and R. decorum,Tang Liping
3453 (MHKMU LP Tang 3453); in the same location, N 27◦23′14″, E 100◦06′19″, elev. 3240 m,
26 August 2020, on the ground of subalpine mixed forests dominated by P. armandi,Quercus
spp., and Rhododendron spp., Mu Man 791 (MHKMU M Mu 791)
J. Fungi 2024, 10, x FOR PEER REVIEW 14 of 26
rechecked the holotype’s ITS sequence of H. flabellatum, and it differed from H. pallidomar-
ginatum by fourteen nucleotide sites. However, nine of them appeared at non-variable
sites in the genus Hydnum. It is clear that the nucleotide site differences of this taxon arose
from the poor-quality sequence. Additionally, both taxa had overlapping distributions in
temperate forests of northeastern China. Moreover, the author also mentioned that the
two taxa had a quite high similarity in their macro- and micro-morphology with minor
differences in H. flabellatum having a scabrous pileus, longer stipes, 2–5-spored basidia,
and the dimensions of the hyphae [18]. Therefore, we proposed that H. flabellatum and H.
pallidomarginatum were con-species, and H. flabellatum should be a synonym of H. pallido-
marginatum.
Figure 7. Macroscopic and microscopic features of Hydnum pallidomarginatum. (a–c) Basidiomata, (a)
from MHKMU HY Huang 873, (b) from MHKMU M Mu 791, (c) from MHKMU LP Tang 3453; (d)
Basidiospores; (e) Basidia and Basidioles; (f) Pileipellis. Bars (d) = 10 µm; (e) = 20 µm; (f) = 50 µm.
Hydnum roseoalbum L.P. Tang, L.J. Su & W.H. Zhang sp. nov. (Figure 8a–f)
Chinese name—粉白齿菌
MycoBank—MB 855835
Etymology—Latin ‘roseo’ = pink, ‘album’ = white, refers to the color of its spines.
Diagnosis—Distinct from other species within Hydnum by its creamy to whitish pi-
leus, pale pink spines, five-spored basidia, globose basidiospores, erectly arranged hy-
phae in pileipellis, and occurrence in subalpine broad-leaved forests.
Holotype—CHINA. Yunnan Province, Jianchuan County (剑川县), N 26°17′52″, E
99°46′9″, elev. 2954 m, 6 October 2020, on the ground of subalpine broad-leaved forests
dominated by Ericaceae and Fagaceae, Zhang Wenhao 606 (MHKMU WH Zhang 606). Gen-
Bank accession numbers: ITS = PQ287669, nrLSU = PQ287751, tef1 = PQ295844.
Description—Basidiomata medium-sized. Pileus 6.5 cm in diam., plano-convex; sur-
face dry, sometimes with spines, azonate, cream to whitish; margin entire, incurved. Con-
text 0.5 cm thick in pileal center, fleshy, white. Stipe 7.5 cm long, 1.4 cm in diam., eccentric,
cylindrical, white to warm cream, solid interior. Spines 0.5–0.7 cm long (with some shorter
spines on stipe), adnexed to decurrent, not flaened, with an acute (rarely suba