A new species of Albatrellus sensu stricto (Albatrellaceae, Russuales) from China

Abstract

Based on morphological and molecular data, a new species of Albatrellus sensu stricto, A. alpinus sp. nov., is described from Yunnan Province of southwestern China. Albatrellus alpinus is characterized by a buff to pale yellow pileal surface, simple septate hyphae, yellow and shapeless oily substance present in all hyphae but seldom in tramal hyphae, and small amyloid basidiospores. Phylogenetic analyses based on the nuclear ribosomal internal transcribed spacer (nrITS) and the large subunit of the nuclear ribosomal RNA gene (nLSU) indicated that the new species is nested within Albatrellus sensu stricto. A key to the Chinese species of Albatrellus sensu stricto is provided.

Full text

Phytotaxa 510 (1): 043–052
https://www.mapress.com/j/pt/
Copyright © 2021 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Genevieve Gates: 9 Jun. 2021; published: 1 Jul. 2021
https://doi.org/10.11646/phytotaxa.510.1.4
43
A new species of Albatrellus sensu stricto (Albatrellaceae, Russuales) from China
HONG-MIN ZHOU1,3, YING-DA WU1,2,4 & YU-CHENG DAI1,5*
1Institute of Microbiology, School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China
2China Fire and Rescue Institute, Beijing 102202, China
3
�
hongminzhou@foxmail.com; https://orcid.org/0000-0002-0724-5815
4
�
wydbjfu@163.com; https://orcid.org/0000-0003-1295-4015
5
�
yuchengd@yahoo.com; https://orcid.org/0000-0002-6523-0320
*Corresponding author
Abstract
Based on morphological and molecular data, a new species of Albatrellus sensu stricto, A. alpinus sp. nov., is described from
Yunnan Province of southwestern China. Albatrellus alpinus is characterized by a buff to pale yellow pileal surface, simple
septate hyphae, yellow and shapeless oily substance present in all hyphae but seldom in tramal hyphae, and small amyloid
basidiospores. Phylogenetic analyses based on the nuclear ribosomal internal transcribed spacer (nrITS) and the large subu-
nit of the nuclear ribosomal RNA gene (nLSU) indicated that the new species is nested within Albatrellus sensu stricto. A
key to the Chinese species of Albatrellus sensu stricto is provided.
Key words: Albatrellus sensu stricto, russuloid fungi, phylogeny, taxonomy
Introduction
Albatrellus Gray (1821: 645) was established based on A. albidus Gray (1821: 645), characterized by annual, terrestrial,
stipitate, fleshy basidiocarps, circular, flabellate, nephroid or irregular pilei, off-white, yellow, orange, brown, blue
to black pore surface, a monomitic hyphal system, and oval, broad oval, or teardrop-shaped, smooth basidiospores
(Pouzar 1966, 1975). Most species have a distribution in the temperate regions of North America, Europe and Asia
(Audet 2010; Vadthanarat et al. 2017). Species in China have been mostly found from Sichuan, Yunnan and Tibet in
southwest China (Cui et al. 2008; Zheng & Liu 2008).
Albatrellus was initially considered as a member of the Polyporales (Kirk et al. 2008). However, some phylogenetic
studies using molecular markers showed that the genus was polyphyletic, with most of them within the Russulales’
clade (Gardes & Bruns 1996; Hibbett et al. 1997; Hibbett & Thorn 2001; Larsson & Larsson 2003; Greslebin et al.
2004), while the saprotrophic species, e.g. Xanthoporus syringae (Parmasto) Audet (2010: 452), were nested within
the polyporoid clade referring to the homobasidiomycetes phylogenetic relationships (Hibbett & Binder 2002; Hibbett
et al. 2005; Miller et al. 2006; Zheng & Liu 2006).
Audet (2010) suggested using Scutiger Paulet (1808: 49) sensu lato instead of Albatrellus sensu lato, including
the narrow definition, Albatrellus sensu stricto. Despite Scutiger prior to Albatrellus, we still used the name Albatrellus
sensu lato, considering it was widely used and the significant differences between the two genera (Pouzar 1966b).
Albatrellus sensu lato presented ten genera: Albatrellopsis Teixeira (1993: 21), Albatrellus Gray, Polyporoletus Snell
(1936: 467), Scutiger, Laeticutis Audet (2010: 441), Neoalbatrellus Audet (2010: 442), Polyporopsis Audet (2010:
447), Polypus Audet (2010: 443), Xanthoporus Audet (2010: 451), Xeroceps Audet (2010: 452), including 36 taxa at
least. The last six genera were proposed by Audet. Among the ten genera, Albatrellus sensu stricto is characterized by
annual, centrally or laterally stipitate, solitary basidiocarps, fleshy, glabrous to squamulose pilei, circular to angular
pores (3–5 per mm), simple septate hyphae, and small, amyloid or inamyloid basidiospores (3–5.7 × 2.5–4.2 μm). So
far, ten species have been included in Albatrellus sensu stricto: A. arizonicus Gilb. (1991: 99), A. avellaneus Pouzar
(1972: 196), A. cantharellus (Lloyd) Pouzar (1972: 196), A. citrinus Ryman (2003: 1245), A. ovinus (Schaeff.) Kotl.
& Pouzar (1957: 154), A. piceiphilus B.K. Cui & Y.C. Dai (2008: 44), A. roseus J. Khan, Sher & Khalid (2018: 14),

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44 • Phytotaxa 510 (1) © 2021 Magnolia Press
A. subrubescens (Murrill) Pouzar (1972: 196), A. tianschanicus (Bondartsev) Pouzar (1966: 358) and A. zhuangii Y.C.
Dai & Juan Li (2008: 268) (Li et al. 2008; Audet 2010; Khan et al. 2018).
Of all the taxa in Albatrellus sensu lato, 14 taxa have been found in China viz., Albatrellopsis flettioides, Albatrellus
avellaneus, Albatrellus cantharellus, Albatrellus roseus, Albatrellus tianschanicus, Neoalbatrellus yasudae, Polypus
dispansus, Xanthoporus syringae, Albatrellus fumosus, Albatrellus piceiphilus, Albatrellus tibetanus, Albatrellus
zhuangii, Neoalbatrellus odorus, Xeroceps yunnanensis (Pouzar 1966a, 1966b, 1972; Zheng & Liu 2006, 2008; Cui
et al. 2008; Audet 2010; Audet & Luther 2015; Chen et al. 2017; Khan et al. 2018), the last six species have been
described from China (Cui et al. 2008; Li et al. 2008; Zheng & Liu 2008; Chen et al. 2017).
During the study on Albatrellus sensu lato from China, three unknown specimens were collected from southwestern
China. To confirm their affinity, phylogenetic analyses based on the nrITS and nLSU rDNA sequences were carried
out. Both morphological characteristics and molecular evidence demonstrated these three specimens represented a new
species of Albatrellus sensu stricto, which we describe in the present paper.
Materials and methods
Morphological studies
Specimens are deposited in the herbarium of the Institute of Microbiology, Beijing Forestry University (BJFC).
Special color terms followed Petersen (1996). Sections were examined at 1000× magnification using a Nikon Eclipse
80i and phase contrast illumination. Spores were measured from sections cut from the tubes, 5% of measurements
were excluded from each end of the range and are given in parentheses. In the text, the following abbreviations were
used: IKI = Melzer’s reagent, IKI– = neither amyloid nor dextrinoid, IKI+ = amyloid, CB = Cotton Blue, CB+ =
cyanophilous, CB– = acyanophilous, L = mean spore length (arithmetic average of all spores), W = mean spore width
(arithmetic average of all spores), Q = variation in the ratios of L/W between specimens studied, n = number of spores
measured from given number of specimens.
Molecular studies
Total genomic DNA was extracted from dried specimens using a CTAB rapid plant genome extraction kit (Aidlab
Biotechnologies Co., Ltd., Beijing, China) according to the manufacturer’s instructions with some modifications (Shen
et al. 2019).
The nuclear ribosomal internal transcribed spacer (nrITS) was amplified with primer pairs ITS5 and ITS4 (White
et al. 1990); the large subunit of nuclear ribosomal RNA gene (nLSU) was amplified with LR0R and LR7 (http://www.
biology.duke.edu/fungi/mycolab/primers.htm).
The PCR procedure for nrITS was as follows: initial denaturation at 95 °C for 3 min, followed by 35 cycles at
94 °C for 40 s, 54 °C for 45 s and 72 °C for 1 min, and a final extension of 72 °C for 10 min. The PCR procedure for
nLSU was: initial denaturation at 94 °C for 1 min, followed by 34 cycles of denaturation at 94 °C for 30 s, annealing
at 50 °C for 1 min and extension at 72 °C for 1.5 min, and a final extension at 72 °C for 10 min. DNA sequencing was
performed at the Beijing Genomics Institute. The sequences were deposited at GenBank (Table 1).
Phylogenetic analyses
Twenty-four new sequences were generated (Table 1). Additional sequences were downloaded from GenBank to
explore the phylogenetic relationships of Albatrellus sensu lato. Heterobasidion annosum (Fr.) Bref. (1889: 154) and
H. australe Y.C. Dai & Korhonen (2009: 354) were selected as outgroups for phylogenetic analyses following Albee-
Scott (2007). Sequences were aligned with ClustalX (Thompson et al. 1997), and manually adjusted in BioEdit (Hall
1999), followed by subsequent deposition at TreeBase (submission ID 27692; www.treebase. org).
The phylogenetic analyses were performed according to the protocols of Cui et al. (2019) and Sun et al. (2020).
Maximum Parsimony (MP), Maximum Likelihood (ML) and Bayesian Inference (BI) methods were employed to
perform phylogenetic analyses. MP analysis was performed using PAUP* 4.0b10 (Swofford 2002). All characters
were equally weighted, and gaps were treated as missing data. Clade robustness was assessed using a Bootstrap
(BT) analysis with 1000 replicates (Felsenstein 1985). Descriptive tree statistics viz. T Length (TL), Consistency
Index (CI), Retention Index (RI), Rescaled Consistency index (RC), and Homoplasy Index (HI), were calculated for
each Maximum Parsimonious Tree (MPT) generated; RAxML v.7.2.6 (Stamatakis 2006) was used for ML analysis,
Bootstrap (BS) replicates were performed under the auto FC option (Pattengale et al. 2010). BI was calculated with

A NEW SPECIES OF ALBATRELLUS SENSU STRICTO Phytotaxa 510 (1) © 2021 Magnolia Press • 45
MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003). For the topology, the default uniform setting was selected that puts
equal probability on all distinct, fully resolved topologies and the three phylogenetic methods resulted in similar
topologies for each dataset. Therefore, only the topology from the MP tree was presented along with statistical values
from the MP/ML/BI algorithms.
TABLE 1. A list of species, specimens and GenBank accession numbers of sequences used in this study.
Species Sample no. Locality GenBank accession no.
nrITS nLSU
Albatrellopsis confluens PV 101-93 Czech Republic AF506393 ——
Albatrellopsis flettii Dai 12873 USA MW807370*MW811451*
Albatrellopsis flettii DAVFP 27659 Unknown JF899544 JF899544
Albatrellopsis flettii UBC F-23821 Canada MF955016 MF955016
Albatrellopsis flettioides MM 72 Pakistan MT040747 ——
Albatrellopsis flettioides MM 76 Pakistan MT040748 ——
Albatrellus alpinus Cui 17023 China MW534154*MW534169*
Albatrellus alpinus Dai 20813 China MW534155*MW534170*
Albatrellus alpinus Dai 20905 China MW534156*MW534171*
Albatrellus avellaneus Dai 20193 China MW534157*MW534172*
Albatrellus avellaneus Dai 20805 China MW269671 MW269683
Albatrellus avellaneus Dai 20806 China MW269672 MW269684
Albatrellus avellaneus p817i USA EU669393 EU669436
Albatrellus avellaneus p819i USA EU669252 EU669302
Albatrellus citrinus Muskos 850928 Sweden AY198190 ——
Albatrellus citrinus Ryman 8002 Sweden AY198191 ——
Albatrellus fumosus Dai 21256 China MW807371*MW811452*
Albatrellus fumosus Dai 21257 China MW807372*MW811453*
Albatrellus ovinus Danell 3/8 2000 Sweden AY198199 ——
Albatrellus ovinus Fransson 1 Sweden AY198200 ——
Albatrellus ovinus Dai 15171 Finland MW534158*MW534173*
Albatrellus ovinus PS 11795 Finland MW269673 MW269685
Albatrellus piceiphilus Cui 2220 China DQ789397 ——
Albatrellus piceiphilus Cui 2221 China DQ789396 ——
Albatrellus roseus SWAT 000121 Pakistan MF110285 MF110295
Albatrellus roseus SWAT 0001222 Pakistan MF110297 MF110297
Albatrellus subrubescens Muskos 97-180 Sweden AY198207 ——
Albatrellus subrubescens Muskos 98-120 Sweden AY198206 ——
Albatrellus subrubescens PV 154-95 Czech Republic AF506395 AF506395
Neoalbatrellus caeruleoporus K.A. Harrison 8825 Canada AY963565 ——
Neoalbatrellus odorus Cui 14226 China KY322503 KY322501
Neoalbatrellus odorus Cui 14227 China KY322504 KY322502
Neoalbatrellus subcaeruleoporus DAVFP 28304 Canada KC862265 ——
Neoalbatrellus subcaeruleoporus OSC 66097 USA KC985126 ——
Polypus dispansus NY 7979 Costa Rica FJ439516 ——
Polypus dispansus OSC 61288 USA EU852810 ——
Scutiger ellisii JLF 1838 USA JX415333 JX415333
Scutiger ellisii States J. WYEF 22
Aug 1998
USA AY621803 ——
Scutiger pes-caprae QFB 7993 Unknown FJ439514 AF506394
Scutiger pes-caprae PV 153-95 Czech Republic AF506394 AF506394
Xanthoporus peckianus QFB 7987 Unknown FJ439513 ——
Xanthoporus syringae Dai 21848 China MW807373*MW811454*
Xanthoporus syringae Dai 21849 China MW807374*MW811455*
Xanthoporus syringae Dai 46749 China AY789078 AY684166
Xeroceps skamania OSC 70203 USA EU669326 ——
Xeroceps skamania WTU 9750 USA EU697276 EU669303
Xeroceps yunnanensis Dai 18452 China MW807375*MW811456*
Xeroceps yunnanensis Dai 20808 China MW807376*MW811457*
Heterobasidion annosum 06071/1 Italy KJ651458 KJ651516
Heterobasidion australe Cui 12602 China MT146485 MT446034
*Newly generated sequences for this study

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46 • Phytotaxa 510 (1) © 2021 Magnolia Press
Branches that received bootstrap support for Maximum Parsimony (MP), Maximum Likelihood (ML-BS) and
Bayesian Posterior Probabilities (BPP) greater than or equal to 75 % (MP and BS) and 0.95 (BPP) were considered as
significantly supported.
Results
Phylogeny
The dataset (nrITS + nLSU) included sequences from seven genera in Albatrellus sensu lato and 48 fungal specimens
representing 21 species. BI resulted in a similar consensus tree to that of the MP and ML analyses, with one million
generations and an average standard deviation of split frequencies = 0.005061.
The dataset had an aligned length of 2153 characters, of which 1505 were constant, 76 were variable but parsimony-
uninformative, and 572 were parsimony-informative. MP analysis yielded four equally most parsimonious trees (TL =
1508, CI = 0.622, RI = 0.847, RC = 0.527, HI = 0.378).
Our phylogeny based on nrITS + nLSU dataset was similar to previous studies (Cui et al. 2008; Audet 2010;
Audet & Luther 2015). All the sequences clustered in separate lineages viz., Albatrellopsis, Albatrellus sensu stricto,
Neoalbatrellus, Polypus, Scutiger, Xanthoporus and Xeroceps. The new species Albatrellus alpinus clustered in the
clade Albatrellus sensu stricto, forming a new lineage with a robust support (100/100/1.00) (Fig. 1).
FIGURE 1. Phylogeny of Albatrellus sensu lato inferred from nrITS + nLSU sequences. Topology is from MP tree and statistical values
(MP/ML/BI) are indicated for each node that simultaneously received BS from ML and MP ≥ 75%, and BPP from BI ≥ 0.95. New species
are in bold.

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Taxonomy
Albatrellus alpinus H.M. Zhou & Y.C. Dai, sp. nov. (Figs. 2–3)
MycoBank: 840110
Type:—CHINA. Yunnan Province, Lijiang, Yulong County, Laojunshan, Ninety-nine Longtan Scenic Area, on ground in forest of Pinus
and Quercus, alt. 3800 m. 15 August 2018, B.K. Cui, Cui 17023 (Holotype, BJFC 030322).
Etymology:—Alpinus (Lat.): refers to the species being found in the alpine zone of the mountains.
Description:—Basidiocarps annual, terrestrial, centrally or laterally stipitate. Pilei circular to flabelliform or
irregular-shaped, up to 2.5 cm diam, and 2 mm thick. Pileal surface buff to pale yellow when fresh, rough to faintly
squamulose, clay buff to grayish brown upon drying, azonate. Pore surface hazel to sepia when dry; pores irregular,
2–3 per mm; dissepiments thin, entire to lacerate. Context white when fresh, becoming reddish brown to black upon
drying, rigid, up to 1.6 mm thick. Tubes white when fresh, buff to curry yellow or cigar brown when dry, brittle, up
to 0.4 mm long. Stipe clay buff to fawn when fresh, becoming dingy with age and gradually becoming reddish brown
upon drying, up to 2.5 cm long, 5 mm diam.
FIGURE 2. A basidiocarp of Albatrellus alpinus (Holotype, Cui 17023). Scale bar = 1.0 cm. Photo by: B.K. Cui.

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FIGURE 3. Microscopic structures of Albatrellus alpinus (Holotype). a Basidiospores b Basidia and basidioles c Stipitipellis hyphae d
Pileipellis hyphae e Hyphae from context f Hyphae from trama. Drawings by: H.M. Zhou.

A NEW SPECIES OF ALBATRELLUS SENSU STRICTO Phytotaxa 510 (1) © 2021 Magnolia Press • 49
Hyphal system monomitic; generative hyphae thin- to thick-walled, simple septate. Pileipellis composed of
generative hyphae, hyaline, thin- to thick-walled, simple septate, IKI+, CB–, 3–10 μm diam; shapeless oily substance
exuding from the hyphae, becoming gray-black in KOH and yellow-brown to red in Melzer’s reagent. Context composed
of generative hyphae, hyaline, thin- to thick-walled, simple septate, occasionally branched, interwoven, IKI–, CB–,
3–20 μm diam; gloeoplerous hyphae thin-walled, 3–10 μm diam, shapeless oily substance exuding from the hyphae
but rarely. Stipitipellis composed of generative hyphae thin- to thick-walled, IKI+, CB–, 2.5–10 μm diam, shapeless
oily substance exuding from the hyphae. Trama of tubes composed of generative hyphae hyaline, thin-walled, simple
septate, parallel along the tubes, IKI–, CB–, 2.5–3.5 μm diam; cystidia and cystidioles absent; basidia clavate with two
or four sterigmata and a simple septum at the base, 19–24.5 × 5–6 μm.
Basidiospores ovoid to ellipsoid, hyaline, thin- to slightly thick-walled, smooth, mostly with a guttule, IKI+, CB–,
(3.5–)3.9–5 × (2.9–)3–4 μm, L = 4.21 μm, W = 3.31 μm, Q = 1.23–1.34 (n = 90/3).
Additional specimens examined (paratypes):—CHINA. Yunnan Province, Dali, Heqing County, on ground in
forest of Pinus, 31 August 2019, Y.C. Dai, Dai 20813 (BJFC 032480); Jianchuan County, Laojunshan Nature Reserve,
on ground in mixed forest, alt. 3000 m. 22 July 2019, Y.C. Dai, Dai 20905 (BJFC 032564).
Discussion
Phylogenetically Albatrellus alpinus formed a robustly supported lineage (100/100/1), clustered within Albatrellus
sensu stricto, and is closely related to A. avellaneus, A. citrinus, A. ovinus, A. piceiphilus, A. roseus and A. subrubescens
(Fig. 1). However, A. avellaneus differs from A. alpinus by inamyloid and larger basidiospores (5–6 × 3.5–4.5 μm vs.
3.9–5 × 3–4 μm, Pouzar 1972). Albatrellus citrinus is distinguished from A. alpinus by a smooth pileal surface when
fresh and orange pores when dry (Ryman et al. 2003). Albatrellus ovinus is separated from A. alpinus by inamyloid
basidiospores (Audet 2010). Albatrellus piceiphilus is different from A. alpinus by a smooth and zonate pileal surface
when fresh (Cui et al. 2008). Albatrellus roseus differs from A. alpinus by larger basidiospores (5–6 × 4–4.5 μm vs.
3.9–5 × 3–4 μm, Khan et al. 2018). Albatrellus subrubescens can be differentiated from A. alpinus by the white to
cream and smooth pileal surface when fresh and smaller basidiospores, 3.8–4.6 × 2.8–3.3 μm, L = 4.2 μm, W = 3 μm,
Q = 1.3–1.4 (Ginns 1997; Niemelä 2005).
Morphologically, Albatrellus alpinus resembles A. tibetanus H.D. Zheng & P.G. Liu (2008: 166) by having septate
hyphae and similar sized basidiospores, but A. tibetanus differs from A. alpinus by inamyloid basidiospores (Zheng
& Liu 2008). Albatrellus alpinus is similar to A. tianschanicus by squamulose pileal surface and similar pore size, but
the latter differs from the former by its black stipe and weakly amyloid basidiospores (Pouzar 1966b; Zheng & Liu
2008).
Two other Asian species Albatrellus zhuangii and A. cantharellus are easily recognized by their bigger spores
(4.9–5.7 × 3.8–4.4 μm and 5.8–6.8 ×4.8–5.5 μm, Li et al. 2008; Pouzar 1975). Neoalbatrellus yasudae is easily
distinguished from the new species by non-amyloid spores (Pouzar 1975).
Key to species of Albatrellus sensu stricto in China
1. Basidiocarps with a rudimentary stipe .................................................................................................................Albatrellus fumosus
1. Basidiocarps with a stipe ....................................................................................................................................................................2
2. Basidiospores inamyloid ..................................................................................................................................Albatrellus avellaneus
2. Basidiospores amyloid ........................................................................................................................................................................3
3. Basidiospores > 5 µm long .................................................................................................................................................................4
3. Basidiospores < 5 µm long .................................................................................................................................................................6
4. Basidiospores > 4.5 µm wide ........................................................................................................................ Albatrellus cantharellus
4. Basidiospores < 4.5 µm wide .............................................................................................................................................................5
5. Pileal surface smooth and viscid when fresh ....................................................................................................... Albatrellus zhuangii
5. Pileal surface with squamulose when fresh .............................................................................................................Albatrellus roseus
6. Gloeoplerous hyphae absent ...............................................................................................................................................................7
6. Gloeoplerous hyphae present .............................................................................................................................................................8
7. Pileal surface densely colored with blackish to deep brown squamulose when fresh ....................................... Albatrellus tibetanus
7. Pileal surface smooth when fresh .................................................................................................................... Albatrellus piceiphilus
8. Pileal surface light brown to tan when fresh, basidiocarps with black stipe when fresh ............................ Albatrellus tianschanicus
8. Pileal surface buff to pale yellow when fresh, basidiocarps with white stipe when fresh .................................... Albatrellus alpinus

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Acknowledgments
This study was supported by the Second Tibetan Plateau Scientific Expedition and Research Program (STEP Grant
No. 2019QZKK0503). We thank the two anonymous reviewers for their corrections and suggestions to improve our
work.
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