Phylogenetic analyses
Forty-three ITS and 37 nrLSU sequences were generated for this study. The concatenated ITS-LSU dataset contained 101 ITS and 107 nrLSU sequences from 107 samples representing 86 Phanerochaetaceae taxa and the outgroup, while the ITS dataset contained 71 samples representing 21 Phlebiopsis s.s. taxa, a sample of Irpex vellereus and the outgroup (Table 1). The concatenated dataset had an aligned length of 2339 characters, of which 554 were parsimony-informative. MP analysis yielded one equally parsimonious tree (TL = 3603, CI = 0.360, RI = 0.695, RC = 0.250, HI = 0.640). The ITS dataset had an aligned length of 726 characters, of which 178 were parsimony-informative. MP analysis yielded 92 equally parsimonious trees (TL = 658, CI = 0.579, RI = 0.870, RC = 0.504, HI = 0.421). jModelTest suggested GTR + I + G and HKY + G were the best-fit models of nucleotide evolution for the concatenated ITS-LSU and ITS datasets, respectively. The average standard deviation of split frequencies of BI was 0.009223 and 0.007710 at the end of the run. ML and BI analyses resulted in almost identical tree topologies compared to the MP analysis. The MP trees are shown in Fig. 1 and Fig. 2 with the parsimony bootstrap values (≥ 50%, first), Bayesian posterior probabilities (≥ 0.95, second) and likelihood bootstrap values (≥ 50%, third) labelled along the branches.
In the Phanerochaetaceae ITS-LSU tree (Fig. 1), Phlebiopsis, Phaeophlebiopsis, Hapalopilus, and Rhizochaete formed a strongly supported clade (98/1/100). Within this clade, the Phlebiopsis species clustered together with relatively strong support values (69/1/98), and species of Phaeophlebiopsis, Hapalopilus and Rhizochaete were in the sister subclades. In the Phlebiopsis ITS tree (Fig. 2), 24 lineages were resolved including 21 taxa of Phlebiopsis and ‘Irpex vellereus’. Samples of P. crassa were distributed in three distinct lineages. The six new species, P. albescens, P. brunnea, P. cylindrospora, P. magnicystidiata, P. membranacea and P. sinensis, formed distinct, strongly supported lineages.
Phlebiopsis albescens Y.N. Zhao & S.H. He, sp. nov. Figure 3
MycoBank: MB836023
Type – Sri Lanka, Avissawella, Salgala Forest, on fallen angiosperm twig, 3 March 2019, He 5805 (BJFC 030672, holotype).
Etymology – Refers to the white basidiomata.
Fruiting body – Basidiomata annual, resupinate, widely effused, closely adnate, inseparable from substrate, ceraceous to crustose, first as small patches, later confluent up to 15 cm long, 1 cm wide, up to 80 µm thick in section. Hymenophore smooth, white (6A1), orange white (6A2) to pale orange (6A3), unchanged in KOH, not cracking on drying; margin indistinct, concolorous with hymenophore. Context white.
Microscopic structures – Hyphal system monomitic; generative hyphae simple-septate. Subiculum indistinct to absent. Subhymenium well developed; hyphae hyaline, thin- to slightly thick-walled, tightly agglutinated, 2.5–4 µm in diam. Lamprocystidia abundant, conical, hyaline to pale yellow, thick-walled, heavily encrusted with crystals along entire length, embedded or slightly projecting beyond hymenium, with one or two secondary septa, with a basal simple septum, 25–40 ⋅ 8–12 µm (without encrustations). Basidia clavate to cylindrical, hyaline, thin-walled, with a basal simple septum and four sterigmata, 10–16 × 3–4.5 µm; basidioles numerous, similar to basidia but slightly smaller. Basidiospores oblong ellipsoid to short cylindrical, hyaline, thin-walled, smooth, IKI–, CB–, 3.5–5 ⋅ 2–2.2 (–2.5) µm, L = 4.4 µm, W = 2.1 µm, Q = 2.1 (n = 30/1).
Distribution – Sri Lanka
Notes – Phlebiopsis albescens is characterized by thin, white to pale orange basidiomata, an indistinct subiculum, short lamprocystidia (< 40 µm long) and basidia (< 16 µm long), and small basidiospores (< 5 µm long). Phlebiopsis punjabensis G. Kaur, Avn.P. Singh & Dhingra, from India, also has thin, white basidomata and short lamprocystidia but larger basidiospores, 5.3–8.5 × 2.5–4 µm (Kaur et al. 2015). Another species with short basidiospores, P. yunnanensis C.L. Zhao, from southern China, has thicker basidiomata, 100–500 µm thick, with a smooth to odontoid hymenophore, and ellipsoid basidiospores, 2.5–3.5 µm broad (Zhao et al. 2018). In the ITS phylogenetic tree (Fig. 2), P. albescens formed its lineage and was not closely related to any other species.
Phlebiopsis brunnea Y.N. Zhao & S.H. He, sp. nov. Figure 4
MycoBank: MB836024
Type – Sri Lanka, Western Province, Mitirigala Nissarana Vanaya Forest Monastery, on fallen angiosperm branch, 4 March 2019, He 5822 (BJFC 030689, holotype).
Etymology – Refers to the brown context of basidiomata.
Fruiting body – Basidiomata annual, resupinate, widely effused, closely adnate, inseparable from substrate, coriaceous, developing as small patches then confluent, up to 20 cm long, 5 cm wide, up to 350 µm thick in section. Hymenophore smooth, brownish grey (6C2–6D2), brownish orange (6C3) to greyish brown (6D3), unchanged in KOH, not cracking on drying; margin thinning out, indistinct, concolorous or darker than hymenophore. Context pale brown.
Microscopic structures – Hyphal system pseudodimitic; generative hyphae simple-septate. Subiculum well-developed, a non-agglutinated, loosely interwoven tissue; skeletocystidia (skeletal hyphae) brown, distinctly thick-walled, slightly encrusted, up to 120 µm long, 14 µm wide; hyphae hyaline to pale yellowish brown, thick-walled, smooth, moderately branched at right angles, frequently septate, 2–5 µm in diam. Subhymenium thin; skeletocystidia as in subiculum but shorter and more heavily encrusted; generative hyphae hyaline, thin- to thick-walled, moderately branched, frequently septate, loosely interwoven, 2–4.5 µm in diam. Lamprocystidia subulate to fusiform, hyaline, thin- to thick-walled, distal end encrusted with small crystals, projecting up to 30 µm beyond hymenium, with an obtuse or acute tip, with a basal simple septum, 35–65 × 7–10 µm. Basidia clavate to subcylindrical, hyaline, thin-walled, with a basal simple septum and four sterigmata, 20–33 × 4.5–6 µm; basidioles numerous, similar to basidia but slightly smaller. Basidiospores oblong ellipsoid to subcylindrical, hyaline, thin-walled, smooth, IKI–, CB–, 6.5–7.5 (–8) × 3–3.6 (–4) µm, L = 7.3 µm, W = 3.3 µm, Q = 2.2 (n = 30/1).
Distribution – Sri Lanka.
Notes – Phlebiopsis brunnea is characterized by a coriaceous basidiomata with a smooth hymenophore and brown context, abundant, brown skeletocystidia in the subiculum and subhymenium, lamprocystidia, and oblong ellipsoid to subcylindrical basidiospores. Hjortstamia bambusicola (Berk. & Broome) Hjortstam & Ryvarden is similar with its grayish brown hymenophore and pseudodimitic hyphal system with brown skeletocystidia but with narrower basidiospores (2.5–3 µm wide) and grows on bamboo in Australia (Hjortstam and Ryvarden 2005). Phlebiopsis brunneocystidiata (Sheng H. Wu) Miettinen has narrower lamprocystidia (5–8 µm wide) with brown walls and a host preference for Pandanaceae in Taiwan (Wu 2004). Another similar species, P. crassa differs from P. brunnea by having effused-reflexed basidiomata with a more or less purple hymenophore and larger lamprocystidia, 50–120 × 8–20 µm (Burdsall 1985; Hjortstam and Ryvarden 1990). Phlebiopsis brunnea formed weakly supported sister lineages to P. brunneocystidiata or P. crassa group B and C in the ITS-LSU and ITS trees, respectively (Figs. 1, 2).
Phlebiopsis cylindrospora Y.N. Zhao & S.H. He, sp. nov. Figure 5
MycoBank: MB836025
Type – China, Hainan Province, Lingshui County, Diaoluoshan Nature Reserve, on dead, small diameter bamboo, 2 July 2019, He 5984 (BJFC 030860, holotype).
Etymology – Refers to the cylindrical basidiospores.
Fruiting body – Basidiomata annual, resupinate, widely effused, closely adnate, inseparable from substrate, coriaceous, first as small patches, later confluent up to 20 cm long, 4 cm wide, up to 150 µm thick in section. Hymenophore smooth, orange white (6A2), orange grey (6B2) to greyish orange (6B3), turning purple in KOH, not cracking on drying; margin thinning out, indistinct, slightly fimbriate, paler than or concolorous with hymenophore. Context gray.
Microscopic structures – Hyphal system monomitic; generative hyphae simple-septate. Subiculum distinct, a somewhat agglutinated, compact tissue, arranged more or less parallel to substrate; hyphae hyaline, thick-walled, encrusted with yellow, resinous granules, infrequently branched, moderately septate, 2–4.5 µm in diam. Subhymenium indistinct; hyphae thin- to slightly thick-walled, heavily encrusted with yellow, resinous granules, frequently septate, more or less agglutinated, 2–4 µm in diam. Lamprocystidia numerous, subfusiform, hyaline, thick-walled, apically encrusted with small crystals, embedded or slightly projecting beyond hymenium, 20–30 (–40) × 5–7 µm. Basidia clavate to subcylindrical, hyaline, thin-walled, with a basal simple septum and four sterigmata, 12–16 × 4–5 µm; basidioles numerous, similar to basidia but slightly smaller. Basidiospores cylindrical, hyaline, thin-walled, smooth, IKI–, CB–, 5.5–7 × 2–2.8 (–3) µm, L = 6.2 µm, W = 2.3 µm, Q = 2.7 (n = 30/1).
Additional specimens examined – China, Hainan Province, Qiongzhong County, Limushan Nature Reserve, on fallen angiosperm twig, 8 June 2016, He 3831 (BJFC 022333); on dead, small diameter bamboo, 8 June 2016, He 3882 (BJFC 022384, CFMR); Wuzhishan County, Wuzhishan Nature Reserve, on dead, small diameter bamboo, 10 June 2016, He 3926 (BJFC 022428); 30 June 2019, He 5922 (BJFC 030797), He 5932 (BJFC 030807), He 5936 (BJFC 030811) & He 5938 (BJFC 030813); Lingshui County, Diaoluoshan Nature Reserve, on dead, small diameter bamboo, 2 July 2019, He 5981 (BJFC 030857); 5 July 2019, He 6054 (BJFC 030930), He 6061 (BJFC 030937) & He 6063 (BJFC 030939); on fallen angiosperm branch, 5 July 2019, He 6038 (BJFC 030914). Thailand, Chiang Rai, Doi Pui, on rotten bamboo, 23 July 2016, He 4080 (BJFC 023521), He 4083 (BJFC 023524) & He 4094 (BJFC 023535, CFMR).
Distribution – China and Thailand.
Notes – Phlebiopsis cylindrospora is characterized by pale-colored, smooth hymenophore that turns purple in KOH, a monomitic hyphal system with generative hyphae encrusted with yellow, resinous granules, small subfusiform lamprocystidia, cylindrical basidiospores, and habit on bamboo and woody angiosperms. It is similar to P. punjabensis that also has a pale-colored, smooth hymenophore and short lamprocystidia, but the latter species does not react with KOH and develops longer basidia (14–26 µm long), and slightly larger basidiospores (5.3–8.5 × 2.5–4 µm, Kaur et al. 2015). Phlebiopsis albescens differs from P. cylindrospora by its white hymenophore that is unchanged in KOH and distinctly smaller basidiospores (3.5–5 × 2–2.2 µm). The hymenophore in P. friesii (Lév.) Spirin & Miettinen turns purple in KOH also but is distinct from P. cylindrospora by having effused-reflexed basidiomata, a pseudodimitic hyphal system, and larger lamprocystidia, up to 80 ⋅ 20 µm (Hjortstam and Ryvarden 1990). Although the phylogenetic trees (Figs. 1, 2) show that P. cylindrospora and P. pilatii are closely related, the latter species is distinct morphologically for it lacks lamprocystidia and develops finely branched dendrohyphidia and larger basidiospores, 8–10 × 4–4.5 µm (Parmasto 1965; Larsen and Gilbertson 1977; Duhem and Michel 2009).
Phlebiopsis magnicystidiata Y.N. Zhao & S.H. He, sp. nov. Figure 6
MycoBank: MB836026
Type – China, Hunan Province, Guzhang County, Gaowangjie Nature Reserve, on dead angiosperm branch, 4 August 2018, He 5648 (BJFC 026710, holotype).
Etymology – Refers to the large lamprocystidia.
Fruiting body – Basidiomata annual, resupinate, widely effused, closely adnate, inseparable from substrate, ceraceous to coriaceous, up to 15 cm long, 5 cm wide, up to 400 µm thick in section. Hymenophore smooth to slightly odontoid with scattered tubercles, pruinose from projecting cystidia, greyish orange [6B(3–5)], brownish orange [6C(3–5)] to light brown [6D(4–6)], unchanged in KOH, sometimes sparsely and deeply cracked with age; margin thinning out, indistinct, concolorous with hymenophore. Context white.
Microscopic structures – Hyphal system monomitic; generative hyphae simple-septate. Subiculum indistinct to absent. Subhymenium thickening, well-developed; hyphae hyaline, thin- to slightly thick-walled, frequently septate, slightly agglutinated, vertically arranged, 2–4.5 µm in diam. Lamprocystidia numerous, fusiform to subulate, hyaline, thick-walled, heavily encrusted with crystals, embedded or projecting beyond hymenium up to 40 µm, with a basal simple septum, apex subacute, 40–80 × (7–) 9–13 (–15) µm (without encrustations). Basidia clavate, hyaline, thin-walled, with a basal simple septum and four sterigmata, 20–30 × 5–6 µm; basidioles numerous, similar to basidia but slightly smaller. Basidiospores broadly ellipsoid to subglobose, hyaline, thin-walled, smooth, IKI–, CB–, 4.5–6.5 × (3.5–) 3.8–4.8 µm, L = 5.6 µm, W = 4.4 µm, Q = 1.3 (n = 30/1).
Additional specimens examined – China, Yunnan Province, Mengla County, Wangtianshu Forest Park, on fallen angiosperm branch, 19 July 2014, He 20140719-18 (BJFC 019145); Taiwan Province, Taichung, Tunghai University, on dead branch of Cassia siamea, 5 August 1989, Wu 890805-1 (TNM F0022186).
Distribution – Hunan, Yunnan, and Taiwan Provinces in southern China.
Notes – Phlebiopsis magnicystidiata is characterized by large lamprocystidia and broadly ellipsoid to subglobose basidiospores. It is morphologically similar to and phylogenetically closely related to P. flavidoalba (Cooke) Hjortstam (Figs. 1, 2) that has smooth hymenophore, slightly longer ellipsoid basidiospores (6–7.5 µm long) and a distribution in North and South America (Burdsall 1985; Gilbertson and Blackwell 1985). Phlebiopsis gigantea and P. magnicystidiata have similar lamprocystidia but the former differs in its well-developed subiculum, narrowly ellipsoid basidiospores, 5–7 × 2.5–3.5 µm, and often occurs on gymnospermous wood in the North Hemisphere (Eriksson et al. 1981; Bernicchia and Gorjón 2010). Except for developing a distinct subiculum, P. darjeelingensis and P. magnicystidiata have similar sized lamprocystidia, basidia, and basidiospores (Dhingra 1987). Reports of P. flavidoalba from India (Rattan 1977) and Taiwan (Wu 1990) need to be confirmed for they may be P. magnicystidiata instead.
Phlebiopsis membranacea Y.N. Zhao & S.H. He, sp. nov. Figure 7
MycoBank: MB836027
Type – China, Hainan Province, Qiongzhong County, Limushan Nature Reserve, on dead, small diameter bamboo, 8 June 2016, He 3849 (BJFC 022351, holotype).
Etymology – Refers to the membranaceous basidiomata.
Fruiting body – Basidiomata annual, resupinate, widely effused, adnate, separable from substrate, membranaceous, up to 20 cm long, 5 cm wide, up to 250 µm thick in section. Hymenophore smooth, orange white (6A2), orange grey (6B2), greyish orange [6B(3–5)] to brownish orange [6C(3–5)], unchanged in KOH, sometimes sparsely and finely cracked with age; margin thinning out, fimbriate, concolorous with hymenophore. Context gray.
Microscopic structures – Hyphal system pseudodimitic; generative hyphae simple-septate. Subiculum well-developed, a non-agglutinated, loosely interwoven tissue; skeletocystidia abundant, fusiform to clavate, brown, thick-walled, smooth, with an acute or obtuse apex, embedded, (30–) 40–70 × 9–13 µm; hyphae hyaline, moderately to distinctly thick-walled, smooth, rigid, frequently branched at right angles, frequently septate, 3–5 µm in diam. Subhymenium thin; hyphae hyaline, thin-walled, smooth, somewhat agglutinated, interwoven, 2–4.5 µm in diam. Hymenial cystidia scattered, similar to skeletocystidia in shape and size but with paler, thinner walls, and sparse encrustations at apex. Basidia clavate, hyaline, thin-walled, with a basal simple septum and four sterigmata, 15–22 × 4–5 µm; basidioles numerous, similar to basidia but slightly smaller. Basidiospores oblong ellipsoid to subcylindrical, hyaline, thin-walled, smooth, IKI–, CB–, 4.5–6 × 2–3 µm, L = 5.4 µm, W = 2.4 µm, Q = 2.3 (n = 30/1).
Additional specimens examined – China, Hainan Province, Qiongzhong County, Limushan Nature Reserve, on dead, small diameter bamboo, 8 June 2016, He 3842 (BJFC 022344); Lingshui County, Diaoluoshan Nature Reserve, on dead, small diameter bamboo, 5 July 2019, He 6062 (BJFC 030938).
Distribution – Hainan Province, southern tropical China.
Notes – Phlebiopsis membranacea is characterized by membranaceous basidiomata with well-developed subicula, brown, smooth, thick-walled skeletocystidia, without lamprocystidia, and habit on bamboo in tropical China. Like P. membranacea, Hjortstamia novae-granatae (A.L. Welden) Hjortstam & Ryvarden, from Columbia, grows on bamboo but its brown, smooth skeletocystidia are tubular in shape and its basidiospores are larger, 5.5–7 × 3–4 µm (Hjortstam and Ryvarden 1990). Phlebiopsis laxa (Sheng H. Wu) Miettinen like P. membranacea has membranaceous basidiomata and loosely arranged subicular hyphae but differs in having lamprocystidia and larger basidiospores, 8–10 × 4–5 µm (Wu 2000). In the phylogenetic trees (Figs. 1, 2), P. membranacea is sister to P. laxa, though their relationship is not strongly supported.
Phlebiopsis sinensis Y.N. Zhao & S.H. He, sp. nov. Figure 8
MycoBank: MB836028
Type – China, Sichuan Province, Wanyuan County, Huaeshan Nature Reserve, on fallen angiosperm branch, 17 July 2013, He 4673 (BJFC 024192, holotype).
Etymology – Refers to the distribution in China.
Fruiting body – Basidiomata annual, resupinate to effused-reflexed with reflexed edges elevated and incurved with age, loosely adnate, easily detached from substrate, coriaceous, first as small patches, later confluent up to 15 cm long, 5 cm wide, up to 300 µm thick in section. Pileus projecting up to 1.5 mm; upper surface gray, slightly sulcate. Hymenophore smooth, brownish orange [6C(3–5)], greyish brown [6(D–F)3] to brown [6E(4–6)], unchanged in KOH, sometimes finely cracked with age; margin thinning out, distinct, white to gray, silky, slightly fimbriate, up to 1 mm wide. Context gray to yellowish brown.
Microscopic structures – Hyphal system pseudodimitic; generative hyphae simple-septate. Tomentum and cortex (a dark line between the tomentum and subiculum) present. Subiculum well-developed, a non-agglutinated tissue; skeletocystidia brown, thick-walled, encrusted at apex, embedded, intermediate forms between skeletocystidia and lamprocystidia observed; hyphae hyaline to pale yellow, moderately to distinctly thick-walled, smooth, rarely branched, moderately septate, easily separated, more or less parallel to substrate, 3–6 µm in diam. Subhymenium indistinct. Lamprocystidia abundant, broadly fusiform to broadly subulate, usually with a long, curved stalk and resembling skeletocystidia, hyaline to brown, thick-walled, heavily encrusted, 30–60 × 8–13 µm, projecting up to 30 µm. Basidia clavate, hyaline, thin-walled, with a basal simple septum and four sterigmata, 20–30 × 4.5–5.5 µm; basidioles numerous, similar to basidia but slightly smaller. Basidiospores oblong ellipsoid to subcylindrical, hyaline, thin-walled, smooth, IKI–, CB–, (5.5–) 6–7.5 × 2.5–3.2 (–3.5) µm, L = 6.5 µm, W = 2.8 µm, Q = 2.3 (n = 30/1).
Additional specimens examined – China, Gansu Province, Pingliang County, Kongtongshan Nature Reserve, on construction wood, 3 August 2015, He 2416 (BJFC 020870, CFMR); Hubei Province, Wufeng County, Houhe Nature Reserve, on dead angiosperm branch, 16 August 2017, He 5081 (BJFC 024599); Hunan Province, Yongshun County, Xiaoxi Nature Reserve, on dead angiosperm branch, 6 August 2018, He 5662 (BJFC 026724); Inner Mongolia, Chifeng, Aohan County, Daheishan Nature Reserve, on fallen Quercus mongolia branch, 3 September 2015, Tiezhi Liu et al. (CFSZ 10714), on fallen Pinus tabuliformis branch, 19 September 2016, Tiezhi Liu et al. (CFSZ 12436); Jiangxi Province, Ji’an County, Jinggangshan Nature Reserve, on dead Rhododendron branch, 11 August 2016, He 4295 (BJFC 023737, CFMR); Liaoning Province, Zhuanghe County, Xianrendong Forest Park, on dead Quercus branch, 5 August 2017, He 4665 (BJFC 024184); Shaanxi Province, Foping County, Foping Nature Reserve, on fallen Betula branch, 11 September 2013, He 1907 (BJFC 016374); Sichuan Province, Baoxing County, Fengtongzhai Nature Reserve, on fallen angiosperm trunk, 18 September 2012, He 20120918-3 (BJFC 014609).
Distribution – Gansu, Hubei, Hunan, Jiangxi, Liaoning, Shaanxi and Sichuan Provinces and Inner Mongolia Autonomous Region of China.
Notes – Phlebiopsis sinensis is characterized by effused to effused-reflexed, coriaceous basidiomata with well-developed subicula, brown skeletocystidia, lamprocystidia, and a temperate distribution. Submembranaceous-pellicular basidiomata, narrower cystidia (5–8 µm wide), and a tropical distribution distinguish P. brunneocystidiata from P. sinensis (Wu 2004). Both P. crassa and P. sinensis develop effused-reflexed basidiomata, but the former species has a purple-tinted hymenophore, larger lamprocystidia, 50–120 × 8–20 µm, and a tropical distribution (Hjortstam and Ryvarden 1990). Although the ITS tree (Fig. 2) shows that P. sinensis and P. friesii are sister taxa, P. friesii is distinct morphologically with a hymenophore that turns purple in KOH and has a dimitic hyphal system with hyaline to yellow skeletal hyphae (Hjortstam and Ryvarden 1990).
Phaeophlebiopsis mussooriensis (Priyanka, Dhingra & N. Kaur) Nakasone & S.H. He, comb. nov.
MycoBank: MB836029
Phlebiopsis mussooriensis Priyanka, Dhingra & N. Kaur, Mycotaxon 115: 255, 2011.
Notes – This species is characterized by a grayish yellow hymenophore, well-developed subiculum, thin-walled generative hyphae, lamprocystidia, and ellipsoid basidiospores (Priyanka et al. 2011). As mentioned in the protologue, P. mussooriensis is quite similar to P. himalayensis, now Phaeophlebiopsis himalayensis (Dhingra) Zmitr., differing primarily in basidiospore size and color change of hymenophore in KOH. Based on Priyanka et al.’s (2011) description, illustration, and comments, we propose the transfer of P. mussooriensis into Phaeophlebiopsis.
Phlebiopsis bambusicola (Berk. & Broome) Nakasone & S.H. He, comb. nov.
MycoBank: MB836030
Corticium bambusicola Berk. & Broome, Transactions of the Linnaean Society of London 2: 64, 1882.
Peniophora bambusicola (Berk. & Broome) Sacc., Sylloge Fungorum 6: 647, 1888.
Hjortstamia bambusicola (Berk. & Broome) Hjortstam & Ryvarden, Synopsis Fungorum 20: 37, 2005.
Notes – This Australian species is known only from the type and is characterized by a grayish brown hymenophore, a dimitic hyphal system, large, brown skeletocystidia, lamprocystidia, narrowly ellipsoid to allantoid basidiospores, and a habit on bamboo (Hjortstam and Ryvarden 2005). Although similar to P. crassa, P. bambusicola has narrower basidiospores, 2.5–3 µm broad and is restricted by host preference and distribution.
Phlebiopsis dregeana (Berk.) Nakasone & S.H. He, comb. nov.
MycoBank: MB836031
Corticium dregeanum Berk., London Journal of Botany 5: 3, 1846.
Hymenochaete dregeana (Berk.) Massee, Botanical Journal of the Linnean Society 27: 114, 1890.
Terana dregeana (Berk.) Kuntze, Revisio generum plantarum 2: 872, 1891.
Lopharia dregeana (Berk.) P.H.B. Talbot, Bothalia 6: 57, 1951.
Irpex dregeanus (Berk.) P.H.B. Talbot, Bothalia 6: 344, 1954.
Australohydnum dregeanum (Berk.) Hjortstam & Ryvarden, Synopsis Fungorum 4: 61, 1990.
Notes – This is a poorly understood species that has been interpreted differently by various researchers. We take a narrow concept of P. dreageana based on studies of the type specimen and specimens restricted to Africa as described and illustrated by Massee (1891), Talbot (1951), Reid (1975), and Hjortstam and Ryvarden (1990). The ellipsoid basidiospores based on these studies are approximately 6.5–8 × 4–5 µm in size. Note that the cylindrical basidiospores illustrated by Reid (1975) are questionable for Hjortstam (1989) noted that basidia and spores were not observed in the type. Hjortstam and Ryvarden (1990) took a broad interpretation of A. dreageanum when they placed Hydnum griseofuscescens Reichardt from Australia and Irpex vellereus Berk. & Broome from Sri Lanka in synonymy; see below for further discussion of these two taxa. Although A. dregeanum has since been reported from India (De 1998, as Oxyporus vellereus), South Korea (Lim 2001; Lim and Jung 2003), New Zealand (Buchanan and Ryvarden 2000), Portugal (Melo and Hjortstam 2002), Israel (Tura et al. 2011), and Italy (Saitta et al. 2014), the basidiospore size, when given, is significantly smaller than the African collections.
Sequences from authentic specimens of the species are not available at present, but ITS sequences labelled “Australohydnum dregeanum” in GenBank, from U.S.A., Korea and Sri Lanka, formed a strongly supported lineage within Phlebiopsis (Fig. 2). The identity of the taxa in this lineage needs further study.
Phlebiopsis griseofuscescens (Reichardt) Nakasone & S.H. He, comb. nov.
MycoBank: MB836032
Hydnum griseofuscescens Reichardt, Verhandlungen der Zoologisch-Botanischen Gesellschaft Wien 16: 374, 1866.
Irpex griseofuscescens (Reichardt) D.A. Reid, Kew Bulletin 17 (2): 273, 1963.
Australohydnum griseofuscescens (Reichardt) Jülich, Persoonia 10 (1): 138, 1978.
Irpex vellereus Berk. & Broome, Journal of the Linnean Society. Botany 14: 61, 1875.
Xylodon vellereus (Berk. & Broome) Kuntze, Revisio generum plantarum 3 (2): 541, 1898.
Hirschioporus vellereus (Berk. & Broome) Teng, Zhong Guo De Zhen Jun [Fungi of China]: 761, 1963.
Oxyporus vellereus (Berk. & Broome) A. Roy & A.B. De, J. Mycopathol. Res.: 41, 1998.
Phlebiopsis lacerata C.L. Zhao, Phytotaxa 440 (4): 274, 2020.
Hydnochaete philippinensis Lloyd (as “philippensis”), Mycological Writings 7 (67): 1154, 1922.
Trichaptum venustum (Berk.) G. Cunn., Bulletin of the New Zealand Department of Scientific and Industrial Research 164: 97, 1965.
Specimens examined – Sri Lanka, Western Province, Ingiriya, Dombagaskanda Forest Reserve, on fallen angiosperm branch, 27 February 2019, He 5734 (BJFC 030601). China, Sichuan Province, Miyi County, Haita Village, on fallen Quercus trunk, 13 September 2015, Cui 12629 (BJFC 028408) & Cui 12637 (BJFC 028416).
Notes – Hydnum griseofuscescens was described from Australia and is the type of Australohydnum (Jülich 1978). It is characterized by resupinate to effused-reflexed basidiomata with a hydnoid, purplish brown hymenophore, a pseudodimitic hyphal system with simple-septate, hyaline, generative hyphae, 4–9 µm broad, encrusted hymenial cystidia with hyaline walls, and small ellipsoid basidiospores, 4–6 × 2.5–3 µm (Reid 1956 as Irpex vellerus, Jülich 1978). We follow Reid (1956, 1963) who determined that H. griseofuscescens and I. vellereus, described from Sri Lanka, were synonyms after studying the types of both species. Reid (1967) also reported that T. venustum sensu Cunningham (1965) is H. griseofuscescens. Based on morphological studies and sequence analyses, we determined that P. lacerata described from southern China (Xu et al. 2020) is conspecific with P. griseofuscescens.
Gilbertson and Adaskaveg (1993) described and illustrated I. griseofuscescens from Hawaii, but this species lacks encrusted hymenial cystidia and has small basidiospores, 4–4.5 × 2–2.5 µm. Similarly, De’s (1998) description of O. vellereus from India appears to represent a different species with a monomitic hyphal system of hyaline to pale brown hyphae and cylindrical basidiospores, 5.2–7 × 2–3 µm. One of the specimens cited, VBMN 80451, is also at CBS, CBS 515.92, and its ITS sequence is available from GenBank (AF479670) as “Irpex vellereus”. This sequence was included in Lim and Jung (2003) and Fig. 2, herein, where it is on a long branch, sister to P. griseofuscescens.
Phlebiopsis novae-granatae (A.L. Welden) Nakasone & S.H. He, comb. nov.
MycoBank: MB836033
Lopharia novae-granatae A.L. Welden [as ‘nova-granata’], Mycologia 67: 540, 1975.
Porostereum novae-granatum (A.L. Welden) Hjortstam & Ryvarden [as ‘nova-granatum’], Synopsis Fungorum 4: 41, 1990.
Phanerochaete novae-granatae (A.L. Welden) Sheng H. Wu [as ‘nova-granata’], Mycotaxon 88: 375, 2003.
Hjortstamia novae-granatae (A.L. Welden) Hjortstam & Ryvarden [as ‘nova-granata’], Synopsis Fungorum 25: 19, 2008.
Notes – Reported from Colombia on bamboo, this species is characterized by a pale brown hymenophore and smooth skeletocystidia but lacking lamprocystidia (Welden 1975; Hjortstam and Ryvarden 1990). Because of its morphological similarity to P. crassa, the transfer of P. novae-granatae is proposed.
Phlebiopsis crassa species complex Fig. 9
Specimens examined – Phlebiopsis crassa group A: Vietnam, Ho Chi Minh City, the Botanical Garden Padua, on fallen angiosperm trunk, 13 October 2017, He 5205 (BJFC 024723). Sri Lanka, Central Province, Kandy, Peradeniya Botanic Garden, on fallen angiosperm branch, 2 March 2019, He 5763 (BJFC 030630). China, Guangdong Province, Renhua County, Danxiashan Nature Reserve, on fallen angiosperm trunk, 4 June 2019, He 5855 (BJFC 030730, Fig. 10a); Yunnan Province, Qiubei County, Puzhehei Nature Reserve, 17 November 2019, He 6300 (BJFC, Fig. 10c), He 6301 (BJFC, Fig. 10d), He 6303 (BJFC, Fig. 10b) & He 6304 (BJFC); Ximeng County, Mengsuolongtan Forest Park, on fallen angiosperm branch, 15 April 2005, Wu 0504 − 22 (TNM F0018719).
Phlebiopsis crassa group B: China, Guangdong Province, Renhua County, Danxiashan Nature Reserve, on fallen angiosperm branch, 4 June 2019, He 5866 (BJFC 030741, Fig. 10f); Yunnan Province, Lushui County, Gaoligongshan Nature Reserve, on fallen angiosperm trunk, 29 November 2015, He 3349 (BJFC 021744, Fig. 10g–h); Maguan County, Gulinqing Nature Reserve, on fallen angiosperm branch, 14 November 2019, He 6266 (BJFC, Fig. 10e).
Phlebiopsis crassa group C: U.S.A., Arizona, Pima County, Santa Rita Experimental Range, on Fouquieria splendens, 31 July 1976, K.K. Nakasone, KKN-86-sp (CFMR); Illinois, Coles County, Fox Ridge State Park, on hardwood, 24 September 1990, A.S. Methven, FP-1024996-sp (CFMR); Mississippi, Harrison County, Harrison Experimental Forest, on Quercus sp., 26 March 1976, H.H. Burdsall, Jr., HHB-8834-sp (CFMR).
Notes – Our phylogenetic analyses showed that samples of P. crassa group A from Vietnam, Sri Lanka and southern China formed a distinct lineage and may represent P. crassa s.s., for the type was described from Vietnam (Figs. 1, 2). Collections from southern China and Japan, group B, and the U.S.A., group C, clustered into two lineages in the ITS tree (Fig. 2,). All three lineages of P. crassa are morphologically similar, however. Unraveling this species complex is beyond the scope of this study, involving a number of presumed synonyms of P. crassa; see Lentz (1955) and Burdsall (1985).
Phlebiopsis darjeelingensis Dhingra, Nova Hedwigia 44: 222, 1987.
Phanerochaete lamprocystidiata Sheng. H. Wu, Mycotaxon 90: 426, 2004.
Phlebiopsis lamprocystidiata (Sheng H. Wu) Sheng H. Wu & Hallenb., Fungal Diversity 42: 116, 2010.
Notes – Because P. darjeelingensis, from India, and P. lamprocystidiata, from Taiwan, are nearly identical morphologically — basidiomata ceraceous when fresh then corneous when dried, well-developed subiculum of compactly packed, hyaline hyphae, and cystidia and basidiospores of similar shape and size (Dhingra 1987; Wu 2004), we consider P. lamprocystidiata to be a later synonym of P. darjeelingensis. Zmitrovich (2018) transfer Phlebiopsis lamprocystidiata to Phaeophlebiopsis based on morphology, our phylogenetic analyses show that it belongs to Phlebiopsis s.s., however.