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Three new species of Aleurodiscus s.l. (Russulales, Basidiomycota) from southern China
expand article infoYan Tian, Masoomeh Ghobad-Nejhad§, Shuang-Hui He, Yu-Cheng Dai
‡ Beijing Forestry University, Beijing, China
§ Iranian Research Organization for Science and Technology, Tehran, Iran
Open Access

Abstract

Three new species of Aleurodiscus s.l. with corticioid basidiomata are described and illustrated from southern China based on morphological evidence and phylogenetic analyses of ITS and nrLSU sequence data. Aleurodiscus bambusinus was collected from Jiangxi Province on bamboo and is distinct by having a compact texture, simple-septate generative hyphae, abundant acanthophyses, basidia with acanthophysoid appendages and smooth basidiospores. Aleurodiscus isabellinus was collected from Yunnan Province on both angiosperm wood and bamboo and is distinct by having soft basidiomata with yellow to yellowish-brown hymenophore, yellow acanthophyses, simple-septate generative hyphae and smooth basidiospores. Aleurodiscus subroseus was collected from Guangxi Autonomous Region and Guizhou Province on angiosperm wood and is distinct by having pinkish basidiomata when fresh, clamped generative hyphae, clavate acanthophyses and echinulate basidiospores. In the phylogenetic tree, A. bambusinus and A. isabellinus were nested within the A. cerussatus group, whilst A. subroseus was clustered with A. wakefieldiae. An identification key to 26 species of Aleurodiscus s.l. in China is provided.

Keywords

acanthophyses, corticioid fungi, Stereaceae, taxonomy, wood-inhabiting fungi

Introduction

Aleurodiscus s.l. is a large group of wood-inhabiting fungi with a broad morphological circumscription. It is characterised by having cupulate, effused or effused-reflexed basidiomata, a monomitic or dimitic hyphal system with simple-septate or clamped generative hyphae, smooth or ornamented, amyloid basidiospores and sterile organs such as acanthophyses, gloeocystidia and dendrohyphidia (Núñez and Ryvarden 1997). Although Aleurodiscus s.l. had been divided into several small genera based on different combinations of morphological characters, phylogenetic analyses did not fully support these separations (Wu et al. 2001; Dai and He 2016). Accordingly, the inter- and intra-generic phylogeny of Aleurodiscus s.l. in Stereaceae is still unclear and no reliable morphological characters can be used to recognise the small segregated genera. Thus, the broad sense concept of the genus has often been adopted by mycologists when describing new species (Núñez and Ryvarden 1997; Gorjón et al. 2013; Dai et al. 2017a, b).

A recent survey on Aleurodiscus s.l. from China (Dai and He 2016, 2017, Dai et al. 2017a, b) revealed that its species diversity is high and many species, especially those with corticioid basidiomata on both herbaceous and ligneous plants, are still undescribed. In the present study, three new species are described and illustrated from southern China, amongst which two species have abundant acanthophyses and smooth basidiospores and one species bears echinulate basidiospores. Morphological differences between new species and their relatives are discussed. Their phylogenetic positions were inferred from a combined dataset of ITS and nrLSU sequence data.

Materials and methods

Morphological studies

Voucher specimens are deposited in the herbaria of Beijing Forestry University, Beijing, China (BJFC), Centre for Forest Mycology Research, U.S. Forest Service, Madison, USA (CFMR) and Southwest Forestry University, Kunming, China (SWFC). Freehand sections were made from basidiomata and mounted in 2% (w/v) potassium hydroxide (KOH), 1% phloxine (w/v) or Melzer’s reagent. Microscopic examinations were carried out with a Nikon Eclipse 80i microscope at magnifications up to 1000×. Drawings were made with the aid of a drawing tube. The following abbreviations are used: L = mean spore length, W = mean spore width, Q = L/W ratio, n (a/b) = number of spores (a) measured from number of specimens (b). Colour names and codes follow Kornerup and Wanscher (1978).

DNA extraction and sequencing

A CTAB plant genome rapid extraction kit-DN14 (Aidlab Biotechnologies Co. Ltd, Beijing) was employed for DNA extraction and PCR amplification from dried specimens. The ITS and nrLSU gene regions were amplified with primer pairs ITS5/ITS4 (White et al. 1990) and LR0R/LR7 (http://www.biology.duke.edu/fungi/mycolab/primers.htm), respectively. The PCR procedures followed Dai and He (2016). DNA sequencing was performed at Beijing Genomics Institute and the sequences were deposited in GenBank.

Phylogenetic analyses

The molecular phylogeny was inferred from a combined dataset of ITS and nrLSU sequences of representative members of Stereaceae sensu Larsson (2007) (Table 1). The ingroup taxa sampling and outgroup selection followed Dai et al. (2017b). The sequences were aligned using MAFFT v.6 (Katoh and Toh 2008, http://mafft.cbrc.jp/alignment/server/). Alignments were optimised manually in BioEdit 7.0.5.3 (Hall 1999) and deposited at TreeBase (http://treebase.org/treebase-web/home.html, submission ID: 22474). Maximum Parsimony (MP), Bayesian Inference (BI) and Maximum Likelihood (ML) analyses were performed by using PAUP* 4.0b10 (Swofford 2002), MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003) and RAxML 7.2.6 (Stamatakis 2006), respectively. The best models of evolution for BI were estimated by using MrModeltest 2.2 (Nylander 2004). The methods and parameter settings for the three kinds of phylogenetic analyses followed Liu et al. (2018).

Species and sequences used in the phylogenetic analyses. Newly generated sequences are set in bold.

Taxa Voucher Locality ITS nrLSU
Acanthobasidium bambusicola He 2357 China KU559343 KU574833
A. norvegicum T 623 France AY039328
A. phragmitis CBS 233.86 France AY039305
A. weirii HHB 12678 USA AY039322
Acanthofungus rimosus Wu 9601-1 Taiwan AY039333
Aleurodiscus abietis T 330 Canada AY039324
A. amorphus Ghobad-Nejhad 2464 China KU559342 KU574832
A. aurantius T 621 France AY039317
A. bambusinus He 4261 China KY706207 KY706219
A. bambusinus He 4263 China KY706208 KY706218
A. bisporus T 627 Guadeloupe AY039318
A. botryosus He 2712 China KX306877 KY450788
A. canadensis Wu 1207-90 China KY706203 KY706225
A. cerussatus He 2208 China KX306874 KY450785
A. dextrinoideocerussatus He 2820 China KY706206 MH109044
A. dextrinoideophyses He 4105 China MH109050 KY450784
A. effusus He 2261 China KU559344 KU574834
A. gigasporus Wu 0108-15 China KY706205 KY706213
A. grantii He 2895 China KU559347 KU574837
A. isabellinus He 5283 China MH109052 MH109046
A. isabellinus He 5294 China MH109053 MH109047
A. lapponicus FP 100753 USA AY039320
A. lividocoeruleus MB 1825 USA AY039314
A. mesaverdensis FP 120155 USA KU559359 KU574817
A. mirabilis Dai 13281 China KU559350 KU574839
A. oakesii He 2243 USA KU559352 KU574840
A. penicillatus HHB 13223 USA KU574816
A. sp. Ghobad-Nejhad 2360 China MH109051 MH109045
A. subroseus He 4807 China MH109054 MH109048
A. subroseus He 4814 China MH109055 MH109049
A. tenuissimus He 3575 China KX306880 KX842529
A. thailandicus He 4099 Thailand KY450781 KY450782
A. tropicus He 3830 China KX553875 KX578720
A. tropicus He 3834 China KX553876 KY706221
A. verrucosporus He 4491 China KY450786 KY450790
A. wakefieldiae He 2580 China KU559353 KU874841
Boidinia macrospora Wu 9202-2 China: Taiwan AF506377 AF506377
Conferticium heimii CBS 321.66 Central African Republic AF506381 AF506381
C. ravum NH 13291 Estonia AF506382 AF506382
Gloeocystidiellum aspellum LIN 625 China: Taiwan AF506432 AF506432
Gloeocystidiopsis cryptacanthus KHL 10334 Puerto Rico AF506442 AF506442
G. flammea AH 000219 La Réunion AF506438 AF506438
Gloeodontia discolor KHL 10099 Puerto Rico AF506445 AF506445
G. pyramidata LR 15502 Columbia AF506446 AF506446
Megalocystidium chelidonium LodgeSJ 110.1 USA AF506441 AF506441
M. leucoxanthum HK 82 Denmark AF506420 AF506420
M. wakullum Oslo 930107 Tanzania AF506443 AF506443
Neoaleurodiscus fujii He 2921 China KU559357 KU574845
Stereum complicatum He 2234 USA KU559368 KU574828
S. ostrea He 2067 USA KU559366 KU574826
S. sanguinolentum He 2111 USA KU559367 KU574827
Xylobous frustulatus He 2231 USA KU881905 KU574825
X. subpileatus FP 106735 USA AY039309

Phylogeny results

The ITS-nrLSU sequences dataset contained 42 ITS and 53 nrLSU sequences from 53 samples representing 47 ingroup taxa and the outgroup (Table 1). Seven ITS and seven nrLSU sequences were generated for this study. The dataset had an aligned length of 2045 characters, of which 384 were parsimony informative. Maximum Parsimony (MP) analysis yielded 85 equally parsimonious trees. The best model estimated and applied in the Bayesian analysis was GTR+I+G. The average standard deviation of split frequencies of BI was 0.007863. ML and BI analyses resulted in almost the same tree topologies as that of MP analysis. Only the MP tree is shown in Fig. 1 with maximum likelihood and maximum parsimony bootstrap values ≥50% and BPP ≥0.95 labelled along the branches. In the tree, A. bambusinus and A. isabellinus were nested within the A. cerussatus (Bres.) Höhn. & Litsch. group (MP = 92%, BI = 1.00, ML = 87%). Aleurodiscus subroseus was clustered with A. wakefieldiae, but their relationship has no support in BI and ML analyses.

Figure 1. 

Maximum parsimony phylogeny of the combined ITS and nrLSU sequences data of Stereaceae. Branches are labelled with maximum parsimony and maximum likelihood bootstrap values ≥50% and Bayesian posterior probabilities ≥0.95 (MP/BI/ML).

Taxonomy

Aleurodiscus bambusinus S.H. He & Y.C. Dai, sp. nov.

MycoBank No: 824755
Figs 2a–b, 3

Diagnosis

The species is distinct by having corticioid basidiomata, a compact texture, simple-septate generative hyphae, abundant acanthophyses, basidia with an acanthophysoid appendage and smooth basidiospores 7–10 × 4–6 μm and growing on bamboo.

Holotype

CHINA. Jiangxi Province, Yifeng County, Guanshan Nature Reserve, alt. ca. 800 m, on fallen culms and branches of bamboo, 10 Aug 2016, He 4261 (holotype, BJFC 023703).

Etymology

Bambusinus” refers to the substrate of bamboo.

Basidiomata

Annual, resupinate, effused, closely adnate, inseparable from substrate, coriaceous, at first as small patches, later confluent up to 30 cm long and 2.5 cm wide, 180–300 μm thick. Hymenophore smooth, white (4A1) to yellowish-white (4A2) when young, becoming greyish-yellow [4B (3–4)] to brownish-orange [6C (5–8)] with age, uncracked or cracked with age; margin abrupt, indistinct, concolorous with hymenophore.

Microscopic structures

Hyphal system monomitic; generative hyphae simple-septate, colourless, thin- to thick-walled, scattered near the substrate, 2–4 μm in diam. Subiculum thin to indistinct. Subhymenium thick, with compact texture, composed of acanthophyses and gloeocystidia. Acanthophyses abundant, hyphoid or distinctly swollen in the middle part, colourless, thin-walled, with abundant spines in apex, 30–40 × 3–12 μm. Gloeocystidia abundant, flexuous or slightly moniliform with one to several constrictions, slightly thick-walled, negative in sulphobenzaldehyde, 30–55 × 8–13 μm. Basidia subclavate to subcylindrical, colourless, slightly thick-walled, usually with a lateral acanthophysoid appendage, with four sterigmata and a basal simple septum, 25–35 × 7–9 μm. Basidiospores ellipsoid to broadly ellipsoid, bearing a distinct apiculus, colourless, thin-walled, smooth, amyloid, 7–10 × 4–6 μm, L = 8.7 μm, W = 4.9 μm, Q = 1.6–1.9 (n = 90/3).

Additional specimens examined

CHINA. Jiangxi Province, Yifeng County, Guanshan Nature Reserve, alt. ca. 800 m, on fallen culms and branches of bamboo, 10 Aug 2016, He 4250 (BJFC 023692) and He 4263 (BJFC 023705).

Remarks

Aleurodiscus bambusinus is morphologically similar and phylogenetically close to A. dextrinoideophyses S.H. He and A. tropicus L.D. Dai & S.H. He that also grow on bamboo in East Asia (Dai et al. 2017a, b). Aleurodiscus dextrinoideophyses differs from A. bambusinus by having apparently dextrinoid acanthophyses and smaller basidiospores (5–7 × 3–4 μm, Dai et al. 2017b). Aleurodiscus tropicus differs from A. bambusinus by having a looser texture and slightly larger basidiospores (9–12 × 5–7.5 μm, Dai et al. 2017a). The ITS similarity between A. bambusinus (He 4261) and A. dextrinoideophyses (He 4105) is 95.6% of 434 base pairs and, between A. bambusinus (He 4261) and A. tropicus (He 3830), is 97.3% of 582 base pairs. Aleurodiscus aberrans G. Cunn. and A. rimulosus Núñez & Ryvarden are also similar to A. bambusinus, but they differ from this new species by having smooth basidia and growing on angiosperm wood outside of Asia (Núñez and Ryvarden 1997).

Figure 2. 

Basidiomata. a–b Aleurodiscus bambusinus (a He 4250 b holotype, He 4261) c A. isabellinus (holotype, KKN-2017-19) d–e A. subroseus (d He 5571 e He 4895). Scale bars: 1 cm.

Figure 3. 

Microscopic structures of Aleurodiscus bambusinus (drawn from the holotype). a Basidiospores; b Basidia c Gloeocystidia d–e Acanthophyses f Generative hyphae.

Aleurodiscus isabellinus S.H. He & Y.C. Dai, sp. nov.

MycoBank No: 824758
Figs 2c, 4

Diagnosis

The species is distinct by having soft, yellow to yellowish-brown and corticioid basidiomata, a loose texture, abundant yellow acanthophyses, simple-septate generative hyphae and smooth basidiospores 6–8.5 × 3–4 μm.

Holotype

CHINA. Yunnan Province, Dali County, Cangshan Nature Reserve, alt. ca. 2600 m, on fallen decorticated angiosperm branches, 27 Oct 2017, KKN-2017-19 (holotype in CFMR, isotype in BJFC).

Etymology

Isabellinus” refers to the yellowish-brown basidiomata.

Basidiomata

Annual, resupinate, effused, adnate, inseparable from substrate, soft, membranaceous to coriaceous, at first as small patches, later confluent up to 15 cm long and 1 cm wide, 150–300 μm thick. Hymenophore smooth, light orange [5A(4–5)] , greyish-orange[5B(5–6)], orange [5B(7–8)] to brownish-yellow [5C(7–8)], uncracked or cracked with age; margin thinning out, fimbriate, white (5A1) when juvenile, becoming abrupt, indistinct, concolorous with hymenophore when mature.

Microscopic structures

Hyphal system monomitic, generative hyphae simple-septate, colourless, thin- to slightly thick-walled, straight, loosely interwoven, frequently branched and septate, 2–4 μm in diam. Acanthophyses abundant, colourless to yellow, thick-walled, hyphoid or arising laterally or apically from a clavate or cylindrical base 30–50 × 5–7 μm, with abundant spines in upper part, some hyphoid ones near substrate with long spines (branches) resembling binding hyphae. Gloeocystidia abundant, embedded, colourless, slightly thick-walled, subcylindrical or slightly moniliform, negative in sulphobenzaldehyde, 35–110 × 5–8 μm. Basidia clavate, colourless, thin-walled, with four sterigmata and a basal simple septum, 40–55 × 6–7 μm. Basidiospores ellipsoid to oblong ellipsoid, bearing a distinct apiculus, colourless, thin-walled, smooth, amyloid, (5.5–) 6–8.5 × (2.8–) 3–4 μm, L = 7 μm, W =3.7 μm, Q = 1.9 (n = 24/1).

Additional specimens examined

CHINA. Yunnan Province, Dali County, Cangshan Nature Reserve, alt. ca. 2600 m, on small dead bamboo, 27 Oct 2017, He 5283 (BJFC 024801) and He 5287 (BJFC 024805); on fallen angiosperm branch, 27 Oct 2017, He 5294 (BJFC 024812); Jingdong County, Ailaoshan Nature Reserve, alt. 2450 m, on fallen angiosperm branch, 4 Oct 2017, C.L. Zhao 3843 (SWFC).

Remarks

All the studied specimens of A. isabellinus lack a true hymenium and only the holotype has a few basidia and basidiospores. Aleurodiscus isabellinus was nested within the A. cerussatus group (Fig. 1). In this group, Aleurodiscus thailandicus S.H. He is similar to A. isabellinus by sharing the yellow basidiomata and acanthophyses, but differs by having two types of gloeocystida and acanthophyses without a clavate or cylindrical base (Dai et al. 2017a). The ITS similarity between A. isabellinus (He 5283) and A. thailandicus (He 4099) is 93.6% of 578 base pairs. Aleurodiscus thailandicus was described from Thailand based on a fertile specimen on bamboo, but later several sterile specimens on bamboo from south-western China were identified as this species according to the sequence data. Morphologically, the soft and yellow to yellowish-brown basidiomata of A. isabellinus resemble the genus Vararia P. Karst. which belongs to Peniophoraceae according to phylogenetic analyses.

Figure 4. 

Microscopic structures of Aleurodiscus isabellinus (drawn from the isotype). a Basidiospores b A basidium and a basidiole c Gloeocystidia d–f Acanthophyses g Generative hyphae.

Aleurodiscus subroseus S.H. He & Y.C. Dai, sp. nov.

MycoBank No: 824757
Figs 2d–e, 5

Diagnosis

The species is distinct by having pinkish and corticioid basidiomata when fresh, clamped generative hyphae, moniliform gloeocystidia, presence of acanthophyses (acanthocystidia) and echinulate basidiospores 16–20 × 11–14 μm.

Holotype

CHINA. Guangxi Autonomous Region, Xing’an County, Mao’ershan Nature Reserve, alt. ca. 1600 m, on dead but still attached branch of living angiosperm tree, 13 Jul 2017, He 4807 (holotype, BJFC 024326).

Etymology

Subroseus” (Lat.) refers to the pinkish basidiomata when fresh.

Basidiomata

Annual, resupinate, effused, closely adnate, inseparable from substrate, coriaceous, at first as small irregular patches, later confluent up to 35 cm long and 3 cm wide, up to 300 μm thick. Hymenophore smooth, pinkish-white (12A2), pink (12A3), pale orange (6A3) to light orange (6A4) when fresh, becoming pale orange (6A3), light orange [6A(4–5)], greyish-orange [6B(3–6)] to brownish-orange [6C(5–6)] when dry, uncracked; margin abrupt, white and distinct when fresh, becoming concolorous or darker than hymenophore and indistinct when dry, slightly elevated when mature.

Microscopic structures

Hyphal system monomitic, generative hyphae with clamp connections. Subiculum thin to indistinct. Subhymenium thickening with age, with embedded gloeocystidia, acanthophyses and crystals. Hyphae in this layer colourless, thin-walled, frequently branched and septate, agglutinated, 2–4 μm in diam. Gloeocystidia abundant, moniliform, with one to several constrictions, smooth, slightly thick-walled, negative in sulphobenzaldehyde, 45–70 × 6–12 μm. Acanthophyses (acanthocystidia) abundant, variable in shape and size, subclavate to subcylindrical, with few to many spines at apex, colourless, slightly thick-walled, 30–60 × 6–20 μm. Hyphidia scattered, thin-walled, colourless, rarely branched. Basidia clavate, slightly sinuous, colourless, thin-walled, smooth, with four sterigmata and a basal clamp connection, 52–80 × 13–17 μm. Basidiospores ellipsoid to broadly ellipsoid, bearing a distinct apiculus, colourless, slightly thick-walled, echinulate, strongly amyloid, 16–20 × 11–14 μm, L = 18.4 μm, W = 12.6 μm, Q = 1.5 (n = 90/3) (spines excluded).

Additional specimens examined

CHINA. Guangxi Autonomous Region, Xing’an County, Mao’ershan Nature Reserve, alt. ca. 1600 m, on dead but still attached branch of living angiosperm tree, 13 Jul 2017, He 4814 (BJFC 024333); Jinxiu County, Dayaoshan Nature Reserve, Yinshan Forest Park, alt. ca. 1500 m, on fallen angiosperm branch, 16 Jul 2017, He 4895 (BJFC 024414). Guizhou Province, Jiangkou County, Fanjingshan Nature Reserve, alt. 1500–2000 m, on dead but still attached branch of living angiosperm tree, 11 Jul 2018, He 5558 (BJFC); 12 Jul 2018, He 5571, He 5577, He 5581, He 5585, He 5589 and He 5593 (BJFC).

Remarks

Aleurodiscus subroseus is morphologically similar and phylogenetically close to A. wakefieldiae Boidin & Beller (Fig. 1), but the latter differs by having longer basidia (80–180 μm) and larger basidiospores (20–28 × 14–20 μm, Núñez and Ryvarden 1997). Aleurodiscus penicillatus Burt is similar to A. subroseus, but differs by growing on gymnosperm wood and having wider basidiospores (13–17 μm, Núñez and Ryvarden 1997). Aleurodiscus mirabilis (Berk. & M.A. Curtis) Höhn. also has pinkish fresh basidiomata and is widely distributed in southern China. However, it can be easily distinguished from A. subroseus by having basally warted basidia and larger basidiospores (24–28 × 14–17 μm, Núñez and Ryvarden 1997). In the phylogenetic tree (Fig. 1), A. penicillatus and A. mirabilis are distantly related to A. subroseus. Aleurodiscus corticola Gorjón et al. from Argentina on bark of living Nothofagus dombeyi also has moniliform gloeocystidia and similar basidiospores with A. subroseus, but differs by having pulvinate and tuberculate basidiomata and absence of acanthophyses (Gorjón et al. 2013).

Figure 5. 

Microscopic structures of Aleurodiscus subroseus (drawn from the holotype). a Basidiospores; b A basidium and a basidiole c Acanthophyses d Gloeocystidia e Hyphidia f Generative hyphae.

Key to 26 species of Aleurodiscus s.l. in China

Acanthobasidium Oberw., Aleurocystidiellum P.A. Lemke and Neoaleurodiscus Sheng H. Wu are used for some species. Basidiospores data are from Núñez & Ryvarden (1997) or otherwise measured by the authors.

1 Basidiospores smooth 2
Basidiospores ornamented 11
2 Acanthophyses absent 3
Acanthophyses present 4
3 Basidiospores thick-walled, 23–27 × 16–21 μm; on Rhododendron Neoaleurodiscus fujii
Basidiospores thin-walled, 18–23 × 14–19 μm; on Quercus A. ljubarskii
4 Basidia with two sterigmata; basidiospores >12 µm long A. canadensis
Basidia with four sterigmata; basidiospores <12 µm long 5
5 Generative hyphae simple-septate 6
Generative hyphae clamped 10
6 Acanthophyses apparently dextrinoid A. dextrinoideophyses
Acanthophyses indextrinoid 7
7 Basidia smooth; acanthophyses yellow 8
Basidia with an acanthophysoid appendage; acanthophyses colourless 9
8 Gloeocystidia of two types; acanthophyses hyphoid A. thailandicus
Gloeocystidia of one type; acanthophyses hyphoid, subclavate to subcylindrical A. isabellinus
9 Texture loose; basidiospores 9–12 × 5–7.5 μm A. tropicus
Texture compact; basidiospores 7–10 × 4–6 μm A. bambusinus
10 Acanthophyses apparently dextrinoid A. dextrinoideocerussatus
Acanthophyses indextrinoid A. cerussatus
11 Acanthophyses absent 12
Acanthophyses present 19
12 Generative hyphae simple-septate 13
Generative hyphae clamped 16
13 Basidiomata discoid; basidiospores >20 µm long A. amorphus
Basidiomata corticioid; basidiospores <20 µm long 14
14 Basidiospores <8 µm long A. tenuissimus
Basidiospores >8 µm long 15
15 Basidiospores 12–17 × 10–15 µm; on angiosperm wood A. ryvardenii
Basidiospores 8–11.5 × 6–8.5 µm; on bamboo A. verrucosporus
16 Basidiospores >20 µm long A. grantii
Basidiospores <20 µm long 17
17 On Quercus Aleurocystidiellum disciforme
On gymnosperm 18
18 Encrusted skeletocystidia present; on Abies Aleurocystidiellum subcruentatum
Moniliform gloeocystidia present; on Pinus Aleurocystidiellum tsugae
19 Acanthophyses amyloid A. botryosus
Acanthophyses non-amyloid 20
20 Basidiospores globose; on bamboo Acanthobasidium bambusicola
Basidiospores ellipsoid; on wood 21
21 On gymnosperm 22
On angiosperm 23
22 Basidiospores 16–21 × 12–17 μm A. effusus
Basidiospores 26–38 × 20–28 μm A. gigasporus
23 Basidiomata white when fresh; acanthophyses rare A. microcarpus
Basidiomata pinkish when fresh; acanthophyses abundant 24
24 Basidiospores 16–20 × 11–14 μm A. subroseus
Basidiospores >20 µm long, >14 µm wide 25
25 Acanthophyses hyphoid, covered with spines at whole upper part; basidia and gloeocystidia covered with spines at basal part; basidiospores usually D-shaped A. mirabilis
Acanthophyses hyphoid to clavate, covered with spines only at apex; basidia and gloeocystidia smooth; basidiospores ellipsoid A. wakefieldiae

Acknowledgements

The authors thank Dr. Karen Nakasone (Center for Forest Mycology Research, Northern Research Station, U.S. Forest Service, Madison, USA) and Dr. Chang-Lin Zhao (Southwest Forestry University, Kunming, China) for providing their specimens. This study was supported by the National Natural Science Foundation of China (Nos. 31470144 & 31670013).

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