Introduction

Phylloporia Murrill was introduced as a monotypic genus, with P. parasitica Murrill growing on a living leaf as generic type (Murrill 1904). Since Wagner and Ryvarden (2002) redefined the concept of Phylloporia with descriptions of species, a key and a preliminary phylogenetic analysis, many species have been newly introduced for Phylloporia with the aid of molecular phylogeny (Valenzuela et al. 2011; Zhou and Dai 2012; Decock et al. 2013, 2015; Gafforov et al. 2014; Yombiyeni et al. 2015; Zhou 2015b, c). A total of 38 species were accepted in Phylloporia before the current study, 15 of which were originally described from China (Cui et al. 2010; Zhou and Dai 2012; Zhou 2013, 2015b, c; Liu et al. 2015).

Phylloporia is a monophyletic genus within the Hymenochaetaceae as recovered by nuclear large subunit ribosomal DNA (nLSU) sequences (Decock et al. 2015; Zhou 2015b). According to the summarization of current accepted species, the diagnostic characters for Phylloporia within the Hymenochaetaceae are a lack of setae and the presence of abundant thick-walled, colored and tiny basidiospores. However, its other morphological characters are highly diverse. For instance, the habit could be annual or perennial, the basidiocarps stipitate or sessile, the context homogeneous or duplex, the duplex context separated by a black zone or not, and the hyphal system could be monomitic or dimitic. Most species of Phylloporia appear to colonize on specific living host genera as potential forest pathogens, while a few species grow on dead wood.

Although the diversity and phylogeny of Hymenochaetaceae have been extensively explored in China, resulting in newly introducing several genera and many species (Dai 2010; Zhou and Xue 2012; Zhou and Qin 2013; Wu et al. 2015; Zhou 2015a; Zhou et al. 2016a, b), there are still many unidentified Chinese hymenochaetoid specimens at the herbaria of the Institute of Applied Ecology, Chinese Academy of Sciences (IFP) and the Institute of Microbiology, Beijing Forestry University (BJFC). When reexamining this kind of specimen, several specimens appeared to fit well with the concept of Phylloporia. After morphological examination and phylogenetic analysis, these specimens were identified to represent Phylloporia pulla (Mont. & Berk.) Decock & Yombiyeni, and two undescribed species that are newly introduced in the current study. In addition, an identification key to worldwide species of Phylloporia is provided.

Materials and methods

Morphological examination

The microscopic procedure followed Dai (2010). Specimen sections, stained in cotton blue (CB), Melzer’s reagent (IKI) and 5 % potassium hydroxide (KOH), were examined using a Nikon Eclipse 80i microscope at magnification up to 1000× under phase contrast illumination. All measurements were taken from sections under CB. When presenting basidiospore size variations, 5 % of the measurements were excluded from each end of the range and are given in parentheses. Special color terms follow Petersen (1996). Microscopic structures were drawn with the aid of a drawing tube. In the text, L stands for mean basidiospore length (arithmetic average of all basidiospores), W for mean basidiospore width (arithmetic average of all basidiospores), Q for the variation of L/W ratio between the specimens studied, and n for number of basidiospores measured from a given number of specimens.

Molecular sequencing

The nLSU region was directly amplified from herbarium specimens using the Phire® Plant Direct PCR Kit (Finnzymes Oy, Finland) according to the manufacturer’s instructions, with primers LR0R and LR7 (Vilgalys and Hester 1990). The PCR procedure was as follows: initial denaturation at 98 °C for 5 min, followed by 39 cycles at 98 °C for 5 s, 48 °C for 5 s and 72 °C for 5 s, and a final extension at 72 °C for 10 min. The PCR products were sequenced using primers LR0R, LR7, LR3 and LR3R (Vilgalys and Hester 1990) at the Beijing Genomics Institute, China. All newly generated sequences were deposited in GenBank (https://www.ncbi.nlm.nih.gov/genbank/; Fig. 1).

Fig. 1
figure 1

Phylogenetic positions of Phylloporia minutipora, P. pulla and P. radiata inferred from the nLSU data set. The topology is from the maximum likelihood algorithm with bootstrap values and Bayesian posterior probabilities, respectively, from maximum likelihood and Bayesian inference algorithms, if simultaneously above 50 % and 0.8, at the nodes. The newly sequenced specimens are in boldface

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Phylogenetic analysis

To explore the phylogenetic positions of the newly sequenced specimens, their nLSU sequences were incorporated into the data sets of previous phylogenetic studies (Decock et al. 2015; Zhou 2015b). Inonotus hispidus (Bull.) P. Karst. was selected as the outgroup (Zhou 2014). The related information to sequences included in the current nLSU data set is summarized in Table 1. The current nLSU data set was aligned using MAFFT 7.110 (Katoh and Standley 2013) with the Q-INS-i option (Katoh and Toh 2008) and the resulting alignment was deposited in TreeBASE (http://www.treebase.org; accession numbers S19032). The best-fit evolutionary model of the alignment was estimated using jModelTest 2.1.4 (Guindon and Gascuel 2003; Darriba et al. 2012), and was set for subsequent phylogenetic analysis. Phylogenetic analysis was performed under maximum likelihood (ML) and Bayesian inference (BI) algorithms. ML algorithm was inferred using raxmlGUI 1.2 (Stamatakis 2006; Silvestro and Michalak 2012). The bootstrap (BS) values were tested under the auto FC option (Pattengale et al. 2010). BI algorithm was conducted using MrBayes 3.2 (Ronquist and Huelsenbeck 2003). Two independent runs with a Metropolis-coupled Markov chain Monte Carlo method were employed. Each run with four chains for 10 million generations starts from random trees. Trees were sampled every 1000th generation. Of the sampling trees, the first 25 % were discarded as burn in, whereas all remaining trees were used to construct a 50 % majority consensus tree and for calculating Bayesian posterior probabilities (BPPs).

Table 1 Information about GenBank accession numbers for nLSU sequences, origin, hosts, and voucher numbers of the specimens used in the phylogenetic analysis
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Results

Molecular phylogeny

The nLSU sequences were newly sequenced from nine specimens (Fig. 1). The current nLSU data set of 119 sequences resulted in an alignment with 951 characters. Its best-fit evolutionary model was estimated as GTR + I + G. The BS search for ML algorithm stropped after 250 replicates. All chains converged in BI algorithm after 10 million generations, where the effective sample sizes of all parameters were more than 1000 and the potential scale reduction factors were close to 1.000. ML and BI algorithms constructed congruent topologies in main lineages, and thus only the topology from the ML algorithm is presented along with statistical values from both algorithms at the nodes.

The current phylogeny inferred from nLSU data set (Fig. 1) shows that the newly sequenced specimens fell into the strongly supported Phylloporia clade (96 %/1) as three new terminal lineages. The lineage comprising specimens Dai 9627 and Cui 5251 (100 %/1) is determined as P. pulla, while the other two, being composed of specimens Dai 9257, LWZ 20150531-13, LWZ 20150531-14 and LWZ 20150531-15 (100 %/1), and LWZ 20141122-5, LWZ 20141122-6 and LWZ 20141122-19 (99 %/1), are described as two new species.

Taxonomy

Phylloporia minutipora L.W. Zhou, sp. nov. (Figs. 2 and 3).

Fig. 2
figure 2

Basidiocarps of Phylloporia minutipora (Dai 9257). Scale bar: 2 cm

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Fig. 3
figure 3

Microscopic structures of Phylloporia minutipora (drawn from the holotype). a. Basidiospores. b. Basidia and basidioles. c. Hyphae from trama. d. Hyphae from lower context. Scale bars: a = 5 μm, b–d = 10 μm

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MycoBank no.: MB 816182

Holotype: China, Hainan, Jianfengling National Nature Reserve, on the base of a living angiosperm tree, 17 November 2007, Dai 9257 (IFP).

Etymology: minutipora (Lat.): referring to the extremely small pores.

Basidiocarps annual, sessile, imbricate, without odor or taste, woody. Pilei dimidiate to flabelliform, fused together, applanate, single pileus projecting up to 10 cm, 7 cm wide and 0.5 cm thick at base. Pileal surface yellowish brown to dark brown, velutinate, distinctly concentrically sulcate with narrow to wide zones; margin honey-yellow, obtuse. Pore surface honey-yellow, more or less shining; sterile margin distinct, curry-yellow, up to 2 mm wide; pores angular, 12–15 per mm; dissepiments thick, entire. Context up to 3 mm thick, duplex, with a black zone, lower context cinnamon-buff, woody, up to 1.5 mm thick, upper tomentum dark brown, soft, up to 1.5 mm thick. Tubes honey-yellow, woody, up to 2 mm long.

Hyphal system dimitic; generative hyphae simple septate; tissue darkening but otherwise unchanged in KOH. Context: in the lower context, generative hyphae yellowish, slightly thick-walled, rarely branched, frequently septate, 2.5–4 μm in diam; skeletal hyphae yellow, thick-walled with a wide lumen, unbranched, aseptate, interwoven, 3–5 μm in diam; in the upper tomentum, generative hyphae yellow, slightly thick-walled, unbranched, frequently septate, 2.5–4 μm in diam; skeletal hyphae brown, thick-walled with a wide lumen, unbranched, aseptate, loosely interwoven, 3.5–5.5 μm in diam; in the black zone, hyphae dark brown, distinctly thick-walled with a narrow lumen, strongly agglutinate, interwoven. Tubes: generative hyphae hyaline to yellowish, thin- to slightly thick-walled, occasionally branched, frequently septate, 2–3 μm in diam; skeletal hyphae dominant, yellow, thick-walled with a wide lumen, unbranched, aseptate, interwoven, 3–5 μm in diam. Cystidia and cystidioles absent. Basidia barrel-shaped, hyaline, thin-walled, with four sterigmata and a simple septum at the base, 5–7 × 3–4 μm; basidioles clavate, slightly smaller than basidia. Basidiospores broadly ellipsoid, pale yellowish, slightly thick-walled, smooth, IKI–, CB–, 2.5–3 × (1.5–)2–2.5 μm, L = 2.74 μm, W = 2.14 μm, Q = 1.26–1.29 (n = 120/4).

Additional specimens studied (paratypes): China, Hainan, Wuzhishan National Nature Reserve, on living angiosperm trunk, 31 May 2015, LWZ 20150531-13 (IFP), LWZ 20150531-14 (IFP), LWZ 20150531-15 (IFP).

Phylloporia radiata L.W. Zhou, sp. nov. (Figs. 4 and 5).

Fig. 4
figure 4

Basidiocarps of Phylloporia radiata (LWZ 20141122-5). a. Pileal surface. b. Pore surface. Scale bars: a–b = 1 cm

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Fig. 5
figure 5

Microscopic structures of Phylloporia radiata (drawn from the holotype). a. Basidiospores. b. Basidia and basidioles. c. Hyphae from trama. d. Hyphae from lower context. Scale bars: a = 5 μm, b–d = 10 μm

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MycoBank no.: MB 816183

Holotype: China, Guizhou, Fanjingshan National Nature Reserve, on living liana, 22 November 2014, LWZ 20141122-6 (IFP).

Etymology: radiata (Lat.): referring to the radially striate pileal surface.

Basidiocarps annual, sessile, attached by a small vertex, imbricate, rarely solitary, without odor or taste, corky. Pilei dimidiate, flabelliform or spathulate, sometimes fused together, applanate, single pileus projecting up to 2.5 cm, 3 cm wide and 0.5 cm thick at base. Pileal surface honey-yellow, velutinate, faintly concentrically sulcate with wide zones, radially striate; margin honey-yellow, sharp. Pore surface reddish brown, more or less shining; sterile margin distinct, curry-yellow to cinnamon-buff, up to 1 mm wide; pores angular, 8–10 per mm; dissepiments thin, entire. Context up to 4 mm thick, duplex, with a black zone, lower context honey-yellow, corky, up to 2 mm thick, upper tomentum concolorous with the lower context, soft, up to 2 mm thick. Tubes cinnamon-buff, corky, up to 1 mm long.

Hyphal system monomitic; generative hyphae simple septate; tissue darkening but otherwise unchanged in KOH. Context: hyphae in the lower context yellow, thick-walled with a wide lumen, unbranched, frequently septate, regularly arranged, 3–5 μm in diam; hyphae in the upper tomentum yellowish, thick-walled with a wide lumen, unbranched, frequently septate, loosely interwoven, 2.5–4 μm in diam; hyphae in the black zone dark brown, distinctly thick-walled with a narrow lumen, strongly agglutinate, interwoven. Tubes: hyphae yellow, thick-walled with a wide lumen, rarely branched, frequently septate, subparallel along the tubes, 2–4 μm in diam. Cystidia and cystidioles absent. Basidia clavate, hyaline, thin-walled, with four sterigmata and a simple septum at the base, 10–15 × 4–7 μm; basidioles in shape similar to basidia, but slightly smaller than basidia. Basidiospores broadly ellipsoid, pale yellowish, slightly thick-walled, smooth, IKI–, CB–, 2.5–3.5 × 2–2.5(−3) μm, L = 3.02 μm, W = 2.42 μm, Q = 1.24–1.26 (n = 90/3).

Additional specimens studied (paratypes): China, Guizhou, Fanjingshan National Nature Reserve, on living liana, 22 November 2014, LWZ 20141122-5 (IFP), LWZ 20141122-19 (IFP).

Other specimens studied: Phylloporia pulla. China, Hainan, Wuzhishan National Nature Reserve, on living angiosperm trunk, 22 May 2008, Dai 9627 (IFP); Jianfengling National Nature Reserve, on living angiosperm trunk, 19 November 2007, Cui 5251 (BJFC).

Discussion

Phylloporia minutipora and P. radiata lack setae and bear thick-walled, colored and tiny basidiospores, which correspond to the morphological concept of Phylloporia. Moreover, the phylogeny inferred from nLSU sequences (Fig. 1) also confirmed these two species to be members of Phylloporia.

Phylloporia minutipora is characterized in the genus by a combination of annual, sessile and imbricate basidiocarps, distinctly concentrically sulcate pileal surface with obtuse margin, angular pores, duplex context separated by a black zone, a dimitic hyphal system, and broadly ellipsoid basidiospores. Its most astonishing characters are extremely small pores of 12–15 per mm and basidiospores of 2.5–3 × 2–2.5 μm even in Phylloporia that is a genus known for tiny basidiospores.

The imbricate pilei and a dimitic hyphal system of Phylloporia minutipora bring P. fulva Yombiyeni & Decock, P. pectinata (Klotzsch) Ryvarden and P. pulla to mind. However, comparing with P. minutipora, besides larger pores and basidiospores, P. fulva and P. pulla also differ in their pendant pilei being attached to the substrata by a small vertex (Yombiyeni et al. 2015), and P. pectinata has a perennial habit (Wagner and Ryvarden 2002).

Phylloporia radiata is characterized in the genus by a combination of annual, sessile and imbricate basidiocarps, faintly sulcate and radially striate pileal surface, sharp pileal margin, angular pores of 8–10 per mm, duplex context separated by a black zone, a monomitic hyphal system, and broadly ellipsoid basidiospores of 2.5–3.5 × 2–2.5 μm.

Phylloporia clausenae L.W. Zhou resembles P. radiata by its annual and sessile basidiocarps, duplex context, a monomitic hyphal system, and broadly ellipsoid basidiospores of 3–3.5 × 2–3 μm (Zhou 2015b). However, P. clausenae differs mainly in its distinctly sulcate pileal surface, obtuse pileal margin, basal context separated by two black zones, and wider hyphae in the tomentum (4–6 μm in diam; Zhou 2015b). Moreover, P. clausenae grows on living angiosperm trunk (Zhou 2015b), whereas P. radiata is only known on living liana. Phylloporia ulloai R. Valenz. et al. is another species of Phylloporia that was found exclusively on living liana, and it shares with P. radiata annual and sessile basidiocarps, wsharp pileal margin, duplex context separated by a black zone, a monomitic hyphal system and broadly ellipsoid basidiospores (Valenzuela et al. 2011). However, P. ulloai, originating from Mexico, is distinct from P. radiata by its much larger basidiocarps (> 4 cm long, > 8 cm wide, and > 1.5 cm thick), larger pores (6–8 per mm), wider hyphae in context (> 5 μm in diam) and slightly larger basidiospores (3.2–3.6 × 2.5–3.2 μm; Valenzuela et al. 2011). The current nLSU-based phylogeny does not recover any reliable relationship at the specific level as in previous studies (Yombiyeni et al. 2015; Zhou 2015b). Therefore, it is impossible to tell whether P. radiata and P. ulloai have a common ancestor restricted to growth on living liana, or if they evolved the habit of living liana separately.

Polyporus pullus Mont. & Berk. was described from Java, Indonesia in 1844 (Montagne and Berkeley 1844). Recently, the holotype of this species has been morphologically reexamined, and its taxonomic position was set in Phylloporia (Yombiyeni et al. 2015). It is very difficult, if possible, to obtain any molecular sequence from the holotype that was collected more than 150 years ago. Therefore, according to morphological comparison, the specimens Cui 5251 and Dai 9627 from Hainan, tropical China were tentatively identified as P. pulla. Both Hainan and Java locate in tropical Asia, and more importantly, the two Chinese specimens share identical morphology in main taxonomic characters with the holotype. The only difference is that the concentric sulcus in pileal surface of the Chinese specimens is distinct and that of the holotype is faint, which might be caused by the more aged Chinese specimens. The nLSU sequences from the two Chinese specimens could represent P. pulla in future phylogenetic analyses of Phylloporia.

An identification key to 30 species of Phylloporia was recently provided by Zhou (2015b), which is essential for identifying specimens of this genus. However, during the time that paper was under review, eight more species, viz. Phylloporia afrospathulata Yombiyeni & Decock, P. dependens Y.C. Dai, P. flabelliforma Decock & Yombiyeni, P. fulva, P. gabonensis Decock & Yombiyeni, P. inonotoides Yombiyeni & Decock, P. pulla and P. yuchengii Yu.Sh. Gafforov et al., were added to Phylloporia (Gafforov et al. 2014; Decock et al. 2015; Liu et al. 2015; Yombiyeni et al. 2015). With the addition of P. minutipora and P. radiata, newly described in the current study, a total of 40 species are accepted in Phylloporia. An updated key to Phylloporia is provided below.

  • 1. Basidiocarps resupinate-----------P. parasitica

  • 1. Basidiocarps sessile or stipitate------------2

  • 2. Basidiocarps stipitate and terrestrial (on buried wood or roots)----------------------3

  • 2. Basidiocarps sessile and on aerial wood---------7

  • 3. Pores > 10 per mm-----------------4

  • 3. Pores < 10 per mm-----------------5

  • 4. Basidiospores < 3.3 μm long, < 2.3 μm wide-------P. terrestris L.W. Zhou

  • 4. Basidiospores > 3.3 μm long, > 2.3 μm wide------P. afrospathulata

  • 5. Basidiospores mostly < 3 μm long----P. minutispora Ipulet & Ryvarden

  • 5. Basidiospores > 3 μm long--------------6

  • 6. Basidiospores > 4 μm long---------P. verae-crucis (Berk. ex Sacc.) Ryvarden

  • 6. Basidiospores 3–4 μm long------P. spathulata (Hook.) Ryvarden

  • 7. Hyphal system dimitic---------------8

  • 7. Hyphal system monomitic-------------12

  • 8. Basidiocarps perennial-----------P. pectinata

  • 8. Basidiocarps annual----------------9

  • 9. Basidiocarps solitary, pores < 9 per mm----------P. nouraguensis Decock & G. Castillo

  • 9. Basidiocarps in cluster, pores > 9 per mm--------10

  • 10. Pileal surface lighter (grayish orange to pale cinnamon), pores < 11 per mm--------------P. fulva

  • 10. Pileal surface darker (yellowish brown to dark brown), pores 11–15 per mm-----------------11

  • 11. Pileus attached by a small vertex and pendant, pores < 12 per mm; basidiospores mostly > 2.5 μm wide----P. pulla

  • 11. Pileus widely attached to the substratum, pores > 12 per mm; basidiospores < 2.5 μm wide ------P. minutipora

  • 12. Pores 2–4 per mm----------------13

  • 12. Pores 4–12 per mm----------------16

  • 13. Basidiospores broadly ellipsoid to subglobose-----P. fruticum (Berk. & M.A. Curtis) Ryvarden

  • 13. Basidiospores oblong-ellipsoid, subcylindrical to cylindrical----------------------14

  • 14. Context duplex-------P. rzedowskii R. Valenz. & Decock

  • 14. Context homogeneous--------------15

  • 15. Context < 1 mm thick; on living branch---P. oblongospora Y.C. Dai & H.S. Yuan

  • 15. Context 2–4 mm thick; on living trunk----P. inonotoides

  • 16. Basidiocarps annual to perennial, dense and hard consistency---------------------17

  • 16. Basidiocarps annual, soft corky at least at tomentum layer---------------------23

  • 17. Pores 10–12 per mm; on living Tilia-------P. tiliae L.W. Zhou

  • 17. Pores 6–9 per mm; on other angiosperms-------18

  • 18. Pileal surface zonate and sulcate-----------19

  • 18. Pileal surface azonate-----------P. yuchengii

  • 19. Pores 6–7 per mm----------------20

  • 19. Pores 7–9 per mm----------------21

  • 20. Basidiospores ellipsoid; mostly on Ribes------P. ribis (Schumach.) Ryvarden

  • 20. Basidiospores subglobose; mostly on Ephedra, Cotoneaster or Jasminum----------P. ephedrae (Woron.) Parmasto

  • 21. Basidiospores > 2.7 μm wide--------P. dependens

  • 21. Basidiospores < 2.7 μm wide------------22

  • 22. Basidiospores ellipsoid to oblong-ellipsoid with a guttule; on Abelia---------P. gutta L.W. Zhou & Y.C. Dai

  • 22. Basidiospores broadly ellipsoid without a guttule; on living Crataegus-------P. crataegi L.W. Zhou & Y.C. Dai

  • 23. Basidiospores broadly ellipsoid to subglobose-----24

  • 23. Basidiospores ellipsoid, oblong-ellipsoid to cylindrical--32

  • 24. Pores 5–6 per mm----------------25

  • 24. Pores 6–11 per mm----------------27

  • 25. Context duplex-----P. ampelina (Bondartsev & Singer) Bondartseva

  • 25. Context homogeneous--------------26

  • 26. Pileus < 1.5 mm thick, margin regular----P. flabelliforma

  • 26. Pileus > 1.5 mm thick, margin irregular----P. gabonensis

  • 27. Basidiocarps > 8 cm wide, > 15 mm thick; contextual hyphae > 5 μm in diam------------P. ulloai

  • 27. Basidiocarps < 8 cm wide, < 15 mm thick; contextual hyphae < 5 μm in diam--------------28

  • 28. Contextual hyphae regularly arranged---------29

  • 28. Contextual hyphae interwoven-----------30

  • 29. Pileus distinctly sulcate, not radially striate, margin obtuse, basal context separated by two black zones; hyphae in tomemtum > 4 μm in diam; on living angiosperm trunk-----------------P. clausenae

  • 29. Pileus faintly sulcate, radially striate, margin sharp, context duplex thoroughly; hyphae in tomemtum < 4 μm in diam; on living liana------------P. radiata

  • 30. Contextual hyphae slightly thick-walled with a wide lumen, frequently septate, large rhomboid crystals absent---------------------31

  • 30. Contextual hyphae thick-walled with a narrow lumen, occasionally septate, large rhomboid crystals present in trama and context------------P. chrysites (Berk.) Ryvarden

  • 31. Pores 10–12 per mm; basidiospores < 3 μm long; on living Fontanesia-----P. fontanesiae L.W. Zhou & Y.C. Dai

  • 31. Pores 7–9 per mm; basidiospores > 3 μm long; on other angiosperms-----P. oreophila L.W. Zhou & Y.C. Dai

  • 32. Basidiospores mostly > 3 μm wide----------33

  • 32. Basidiospores mostly < 3 μm wide----------34

  • 33. Pores 4–6 per mm----P. hainaniana Y.C. Dai & B.K. Cui

  • 33. Pores 8–10 per mm-----P. capucina (Mont.) Ryvarden

  • 34. Basidiocarp solitary---------------35

  • 34. Basidiocarp imbricate---------------38

  • 35. Context homogeneous-----P. homocarnica L.W. Zhou

  • 35. Context duplex-----------------36

  • 36. Context not separated by a black zone; on living Flacourtia----------P. flacourtiae L.W. Zhou

  • 36. Context separated by a black zone; on other angiosperms--------------------37

  • 37. Pores circular; basidiospores mostly > 2.2 μm wide---P. weberiana (Bres. & Henn. ex Sacc.) Ryvarden

  • 37. Pores angular; basidiospores mostly < 2.2 μm wide-----------P. cylindrispora L.W. Zhou

  • 38. Basidiospores mostly < 2.5 μm wide---------39

  • 38. Basidiospores mostly > 2.5 μm wide----P. bibulosa (Lloyd) Ryvarden

  • 39. Context duplex, not separated by a black zone; basidiospores > 3.5 μm long, contextual hyphae interwoven; on living Nandina----P. nandinae L.W. Zhou & Y.C. Dai

  • 39. Context duplex, separated by a black zone; basidiospores < 3.5 μm long, contextual hyphae regularly arranged; on living Osmanthus------P. osmanthi L.W. Zhou