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ORIGINAL ARTICLE
Species diversity and molecular systematics of Fibroporia
(Polyporales, Basidiomycota) and its related genera
Yuan-Yuan Chen
1
&Fang Wu
1
&Min Wang
1
&Bao-Kai Cui
1
Received: 7 December 2016 /Revised: 22 February 2017 /Accepted: 24 February 2017 /Published online: 11 March 2017
#German Mycological Society and Springer-Verlag Berlin Heidelberg 2017
Abstract In this study, phylogenetic analysis of Fibroporia
and its related genera was carried out based on multiple loci of
ITS, nLSU, nSSU, mtSSU, tef1 and rpb2. Combined with the
morphological characters, a new genus, Pseudofibroporia
gen. nov., is established. Phylogenetically, Pseudofibroporia
is closely related to Fibroporia, but morphologically, the for-
mer is different from the latter in its distinct pileate
basidiomata with entire margin and lacking rhizomorphs. In
addition, three new species, Fibroporia bambusae sp. nov.,
F. ceracea sp.nov.andPseudofibroporia citrinella sp. nov.,
are described from China. Fibroporia bambusae is different
from other species by its cream to pinkish buff pore surface,
cinnamon buff rhizomorphs and a dimitic hyphal system with
both clamped and simple-septate generative hyphae.
F. ceracea is distinguished by its ceraceous pore surface and
a monomitic hyphal system with clamped generative hyphae.
Pseudofibroporia citrinella is characterized by white to lemon
yellow pileal surface, lemon yellow to curry buff pores, a
dimitic hyphal system with clamped generative hyphae and
oblong to broadly ellipsoid, and slightly thick-walled basidio-
spores (4–4.5 × 2.2–2.8 μm). Illustrated descriptions of the
novel species and a key to known species of Fibroporia from
China are provided.
Keywords Brown-rot fungi .Fomitopsidaceae .
Multi-marker analysis .Phylogeny .Taxo n omy
Introduction
Fibroporia Parmasto was described by Parmasto (1968)with
Polyporus vaillantii (DC.) Fr. as the generic type, to accom-
modate species with fimbriate to rhizomorphic margin and
ellipsoid and slightly thick-walled basidiospores, and five spe-
cies were transferred into the genus: F. destructor (Fr.)
Parmasto, F. gossypium (Speg.) Parmasto, F. overholtsii
(Pilát.) Parmasto, F. radiculosa (Peck) Parmasto and
F. vaillantii (DC.) Parmasto. However, Ryvarden (1991)
regarded the rhizomorphs as an adaptive character, which is
not enough to justify a generic separation, and synonymised
Fibroporia with Antrodia P. Karst.
Phylogenetic analyses show that Antrodia is polyphyletic,
and taxa in Antrodia sensu lato have been treated as three
different genera: Antrodia sensu stricto, Fibroporia Parmasto
and Amyloporia Singer (Kim et al. 2003;Yuetal.2010;
Rajchenberg et al. 2011; Bernicchia et al. 2012;Cui2013;
Spirin et al. 2013). Recent molecular studies support the sep-
aration of Fibroporia as a distinct genus from Antrodia
(Rajchenberg et al. 2011; Bernicchia et al. 2012;Cui2013;
Ortiz-Santana et al. 2013;Spirinetal.2013;Chenetal.
2015b). The principal characteristics of Fibroporia include a
fimbriate to rhizomorphic margin, a monomitic to dimitic hy-
phal system scarcely with irregularly thick-walled generative
hyphae either in the basidiomata or in culture, uninucleated,
elliposoid basidiospores with slightly thickened walls and
tetrapolar sexuality (Parmasto 1968; Rajchenberg 2006;
Rajchenberg et al. 2011).
In this study, phylogenetic analysis of Fibroporia and its re-
lated genera was carried out based on DNA sequences of multi-
ple loci including the internal transcribed spacer (ITS) regions,
the large subunit nuclear ribosomal RNA gene (nLSU), the small
subunit nuclear ribosomal RNA gene (nSSU), the small subunit
mitochondrial rRNA gene sequences (mtSSU), the translation
Section Editor: Yu-Cheng Dai
*Bao-Kai Cui
baokaicui2013@gmail.com
1
Institute of Microbiology, Beijing Forestry University,
Beijing 100083, China
Mycol Progress (2017) 16:521–533
DOI 10.1007/s11557-017-1285-1
elongation factor 1-αgene (tef1) and the second subunit of RNA
polymerase II (rpb2). Combining morphological and molecular
evidence, a new genus is proposed, and three new species are
described.
Materials and methods
Morphological studies
The studied specimens are deposited at the herbaria of the
Institute of Microbiology, Beijing Forestry University, China
(BJFC), the Institute of Applied Ecology, Chinese Academy
of Sciences, China (IFP), Botanical Museum of University of
Helsinki, Finland (H), and Prague Museums, Czech Republic
(PRM). The microscopic routines followed Li et al. (2014)
and Zhou et al. (2016). Sections were studied at a magnifica-
tion up to × 1000 using a Nikon E80i microscope and phase
contrast illumination (Nikon, Tokyo, Japan). Drawings were
made with the aid of a drawing tube. Microscopic features,
measurements and drawings were made from slide prepara-
tions stained with Cotton Blue and Melzer’s reagent. Spores
were measured from sections cut from the tubes. In presenting
the variation in the size of the spores, 5% of measurements
were given in parentheses. In the text the following abbrevia-
tions were used: IKI = Melzer’sreagent,IKI–=indextrinoid
and inamyloid, KOH = 5% potassium hydroxide, CB =
Cotton Blue, CB–= acyanophilous, L = mean spore length
(arithmetic average of all spores), W = mean spore width
(arithmetic average of all spores), Q = variation in the L/W
ratios between the specimens studied, n = number of spores
measured from given number of specimens. Color terms
followed Petersen (1996).
Molecular study and phylogenetic analyses
A cetyl trimethylammonium bromide rapid plant genome ex-
traction kit (Demeter Biotechnologies Co., Ltd, Beijing) was
used to extract total genomic DNA from dried specimens, and
performed the polymerase chain reaction (PCR) according to
the manufacturer’s instructions with some modifications
(Chen et al. 2015a,2016). The ITS regions were amplified
with primer pairs ITS5 and ITS4 (White et al. 1990). The
nLSU regions were amplified with primer pairs LR0R and
LR7 (http://www.biology.duke.edu/fungi/mycolab/primers.
htm). The nSSU regions were amplified with primer pairs
NS1 and NS4 (White et al. 1990). The mtSSU regions were
amplified with primer pairs MS1 and MS2 (White et al. 1990).
Part of tef1 was amplified with primer pairs EF1-983F and
EF1-1567R. Rpb2 was amplified with primer pairs bRPB2-
6F and bRPB2-7R (Matheny 2005). The PCR cycling sched-
ule for ITS, mtSSU and tef1 included an initial denaturation at
95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 54 °C
for ITS and mtSSU, 54–59 °C for tef1 for 45 s, 72 °C for 1
min, and a final extension at 72 °C for 10 min. The PCR
cycling schedule for nLSU and nSSU included an initial de-
naturation at 94 °C for 1 min, followed by 35 cycles at 94 °C
for 30 s, 50 °C for nLSU and 53 °C for nSSU for 1 min, 72 °C
for 1.5 min, and a final extension at 72 °C for 10 min. The
PCR cycling schedule for rpb2 followed Kim et al. (2007)
with slight modifications: initial denaturation at 95 °C for 10
min, followed by 39 cycles at 94 °C for 1 min, 56 °C for 1 min
and 72 °C for 1 min, and a final extension at 72 °C for 10 min.
The PCR products were purified and sequenced at Beijing
Genomics Institute (China), with the same primers.
One hundred and two new sequences were generated and
deposited at GenBank (Table 1). Additional sequences were
downloaded from GenBank (Table 1). All sequences were
aligned using ClustalX (Thompson et al. 1997) and manually
adjusted in BioEdit (Hall 1999). The missing sequences are
coded as BN^. Ambiguous nucleotides were coded as BN^.
The final concatenated sequence alignment was deposited in
TreeBase (http://purl.org/phylo/treebase;submissionID
18824).
Most parsimonious phylogenies were inferred from the
combined 6-gene dataset (ITS+nLSU+nSSU+mtSSU+tef1+
rpb2), and their congruences were evaluated with the incon-
gruence length difference (ILD) test (Farris et al. 1994)imple-
mented in PAUP* 4.0b10 (Swofford 2002), under heuristic
search and 1000 homogeneity replicates. Phylogenetic analy-
sis approaches followed Zhao et al. (2014,2015). Sequences
of Trametes suaveolens (L.) Fr. and Coriolopsis polyzona
(Pers.) Ryvarden obtained from GenBank were used as
outgroups to root trees following Han et al. (2016).
Maximum parsimony analysis was applied to the combined
multiple genes datasets, and the tree construction procedure
was performed in PAUP* version 4.0b10. All characters were
equally weighted and gaps were treated as missing data. Trees
were inferred using the heuristic search option with TBR
branch swapping and 1000 random sequence additions.
Max-trees were set to 5000, branches of zero length were
collapsed and all parsimonious trees were saved. Clade ro-
bustness was assessed using a bootstrap (BT) analysis with
1000 replicates (Felsenstein 1985). Descriptive tree statistics
tree length (TL), consistency index (CI), retention index (RI),
rescaled consistency index (RC), and homoplasy index (HI)
were calculated for each Most Parsimonious Tree (MPT) gen-
erated. RAxML v.7.2.8 was used to construct a maximum
likelihood (ML) tree with a GTR+G+I model of site substitu-
tion including estimation of Gamma-distributed rate heteroge-
neity and a proportion of invariant sites (Stamatakis 2006).
The branch support was evaluated with a bootstrapping meth-
od of 1000 replicates (Hillis and Bull 1993). Phylogenetic
trees were visualized using Treeview (Page 1996).
MrModeltest 2.3 (Posada and Crandall 1998; Nylander
2004) was used to determine the best-fit evolution model for
522 Mycol Progress (2017) 16:521–533
Tab l e 1 A list of species, specimens and GenBank accession numbers of sequences used in this study
Species name Sample no. Locality GenBank accession numbers
ITS nLSU nSSU mtSSU tef1 rpb2
Amylocystis lapponica HHB-13400-Sp USA KC585237 KC585059 ————
Am. lapponica OKM-4418-Sp USA KC585238 KC585060 ————
Amyloporia carbonica Cui 12212 China KR605816 KR605755 KR605917 KR606017 KR610745 —
Am. xantha Cui 11544 China KR605817 KR605756 KR605918 KR606018 KR610746 KR610836
Antrodia heteromorpha Dai 12755 USA KP715306 KP715322 KR605908 KR606009 KP715336 KR610828
A. malicola Cui 9491 China KT968824 KT968828 KU550522* KU550502* —KT988994
A. malicola Cui 10473 China KU550482* KU550483* KU550523* KU550503* —KU550541*
A. serialis Cui 9706 China KR605811 KR605750 KR605910 KR606010 KR610741 KR610829
A. serialis Cui 10519 China KP715307 KP715323 KR605911 KR606011 KP715337 KR610830
A. serpens Dai 7465 China KR605813 KR605752 KR605913 KR606013 KR610742 KR610832
A. tanakae Cui 9743 China KR605814 KR605753 KR605914 KR606014 KR610743 KR610833
A. variiformis CBS 309.82 USA DQ491418 AY515344 —DQ491445 —DQ491391
A. variiformis FP-90100-SP USA KC585311 KC585136 ————
Coriolopsis polyzona Cui 11040 China KR605824 KR605767 KR605932 KR606029 KR610760 KR610849
Daedalea dickinsii Yuan 1090 China KR605790 KR605729 KR605878 KR605981 KR610711 KR610802
D. quercina Dai 12659 Finland KP171208 KP171230 KR605887 KR605990 KR610719 KR610810
Dacryobolus sudans FP-100190-Sp USA KC585331 KC585156 ————
Da. sudans FP-150381 Jamaica KC585333 KC585158 ————
Fibroporia albicans Cui 9464 China KC456250 KR605758 KR605920 KR606019 KR610748 KR610838
F. albicans Cui 9495 China KC456252 KU550484* KU550524* KU550505* KU550557* KU550542*
F. albicans Cui 9504 China KC456251 KU550485* KU550525* KU550504* KU550558* KU550543*
F. albicans Dai 10595 China KC456249 KR605759 KR605921 KR606020 KR610749 KR610839
F. bambusae Dai 16210 China KU550479* KU550486* KU550526* KU550506* KU550559* KU550544*
F. bambusae Dai 16211 China KU550480* KU550487* KU550527* KU550507* KU550560* KU550545*
F. bambusae Dai 16212 China KU550481* KU550488* KU550528* KU550508* KU550561* KU550546*
F. bohemica Vampola 21.8.1992 Czech Republic KF112876 KF112876 ————
F. bohemica PRM 859138 Czech Republic KT895883 KU550489* KU550529* KU550509* KU550562* KT895900
F. c e r a c e a Dai 13013 China KU550476* KU550490* KU550530* KU550510* KU550563* KU550547*
F. citrina Cui 9683 China KC456253 KU550491* KU550531* KU550511* KU550564* KU550548*
F. citrina Cui 10497 China KT895886 KT988993 KU550532* ——KU550549*
F. citrina Cui 11604 China KU550473* KU550492* KU550533* KU550512* KU550568* KU550551*
F. citrina HUBO 7237 France HM542006 —————
F. citrina HUBO 7715 Italy GU991573 —————
F. citrina HUBO 7887 Italy HM542005 —————
F. citrina LY BR 4205 Luxembourg KT895884 KU550493* —KU550513* KU550566* KT895901
F. gossypium Cui 9472 China KU550474* KU550494* KU550534* KU550514* KU550567* KU550550*
F. gossypium HUBO 7725 Italy GU991576 —————
F. gossypium HUBO 7952 Italy HM590881 —————
F. gossypium LY BR 3914 France KU550475* KU550495* —KU550515* KU550569* KU550552*
F. gossypium LY BR 3994 Argentina KT895885 KU550496* KU550535* KU550516* KU550570* KT895902
Mycol Progress (2017) 16:521–533 523
Tab l e 1 (continued)
Species name Sample no. Locality GenBank accession numbers
ITS nLSU nSSU mtSSU tef1 rpb2
F. gossypium X1403 Finland KC595905 KC595905 ————
F. norrlandica 4115 Finland KC595907 KC595907 ————
F. norrlandica 4122 Finland KC595908 KC595908 ————
F. norrlandica 4151 Finland KC595909 KC595909 ————
F. pseudorennyi X1377 France KC595927 KC595927 ————
F. pseudorennyi X1384 Russia KC595928 KC595928 ————
F. radiculosa Cui 2796 China KC456247 KU550497* KU550536* KU550517* KU550571* KU550553*
F. radiculosa Cui 11404 China KP145011 KR605760 KR605922 KR606021 KR610750 KR610840
F. radiculosa Dai 6473 China FJ644283 KU550498* KU550537* KU550518* KU550572* KT895899
F. radiculosa Dai 13041 China KP145010 KU550499* KU550538* KU550519* KU550573* KU550554*
F. radiculosa FP-105309-R USA KC585339 KC585164 ————
F. radiculosa FP-90848-T USA KC585340 KC585165 ————
F. vaillantii MR 11674 USA JF713076 —————
F. vaillantii Ostrow 2120 Italy HM590884 —————
F. vaillantii FP-90877-R Argentina KC585345 KC585170 ————
F. vaillantii HUBO 8160 Germany HM590885 —————
F. vaillantii RWD-63-263 USA KC585346 KC585172 ————
Fomitopsis betulinus Cui 10756 China KR605797 KR605736 KR605894 KR605997 KR610725 KR610815
Fo. pinicola Cui 10312 China KR605781 KR605720 KR605856 KR605960 KR610689 KR610780
Laetiporus sulphureus Dai 12154 China KF951295 KF951302 KR605924 KR606023 KR610752 KR610841
L. sulphureus Dai 12826 China KR605819 KR605762 KR605925 KR606024 KR610753 KR610842
Piptoporellus soloniensis Cui 11390 China KR605803 KR605742 KR605901 KR606003 KR610733 KR610822
Pi. soloniensis Dai 11872 China KR605804 KR605743 KR605902 KR606004 KR610731 KR610823
Postia duplicata Cui 10366 China KF699124 KJ684975 KR605927 KR606026 KR610755 KR610844
Po. lactea Cui 12206 China KR605820 KR605763 KR605926 KR606025 KR610754 KR610843
Pseudofibroporia citrinella He 20120721-15 China KU550477* KU550500* KU550539* KU550520* KU550574* KU550555*
P. citrinella Yuan 6181 China KU550478* KU550501* KU550540* KU550521* KU550575* KU550556*
Pycnoporellus fulgens CA-20 USA KC585385 KC585218 ————
Py. fulgens FP-133367-Sp USA KC585386 KC585219 ————
Sparassis crispa MBUH-DORISLABER/ss25 Germany AY218442 AY218404 ————
S. crispa X950 Finland KC595954 KC595954 ————
Trametes suaveolens Cui 11586 China KR605823 KR605766 KR605931 KR606028 KR610759 KR610848
*Newly generated sequences for this study
524 Mycol Progress (2017) 16:521–533
the combined multi-gene dataset for Bayesian inference (BI).
Bayesian inference was calculated with MrBayes 3.1.2 with a
general time reversible (GTR) model of DNA substitution and
a gamma distribution rate variation across sites (Ronquist and
Huelsenbeck 2003). Four Markov chains were run for two
runs from random starting trees for 10 million generations
(ITS+nLSU+nSSU+mtSSU+tef1+rpb2) and trees were sam-
pled every 100 generations. The first one-fourth generations
were discarded as burn-in. A majority rule consensus tree of
all remaining trees was calculated. Branches that received
bootstrap support for maximum parsimony (MP), maximum
likelihood (BS) and Bayesian posterior probabilities (BPP)
greater than or equal to 75% (MP and BS) and 0.95 (BPP)
were considered as significantly supported, respectively.
Results
Molecular phylogeny
The combined six gene (ITS+nLSU+nSSU+mtSSU+tef1+
rpb2) sequences dataset had an aligned length of 4757 char-
acters, of which 3014 characters were constant, 190 were var-
iable and parsimony-uninformative, and 1553 were parsimo-
ny-informative. MP analysis yielded 19 equally parsimonious
trees (TL = 6514, CI = 0.446, RI = 0.755, RC = 0.337, HI =
0.554). Best model for the combined ITS+nLSU+nSSU+
mtSSU+tef1+rpb2 sequences dataset estimated and applied
in the Bayesian analysis was: GTR+I+G with equal frequency
of nucleotides. ML analysis resulted in a similar topology as
MP and Bayesian analyses, and the ML topology is shown in
Fig. 1.
The phylogeny inferred from multiple genes of the com-
bined ITS+nLSU+nSSU+mtSSU+tef1+rpb2 sequences was
obtained from 72 fungal samples representing 33 taxa of
Fibroporia and its related genera in the antrodia clade.
Thirty-nine samples representing ten taxa of Fibroporia clus-
tered together and separated from species of Antrodia and
other related genera. A new genus, Pseudofibroporia
gen. nov. is established with strong support. Three
new species, Fibroporia bambusae,F. ceracea and
Pseudofibroporia citrinella are described from China;
and they are distant from other species in the phyloge-
netic tree (Fig. 1).
Taxonomy
Fibroporia Parmasto, Consp. System. Corticiac. (Tartu): 176,
1968.
MycoBank no.: MB 17585
Type species:Fibroporia vaillantii (DC.) Parmasto.
Basidiomata annual, resupinate, often widely effused, mar-
gin often fimbriate with rhizomorphs. Pore surface white to
apricot-orange, pores circular to angular, dissepiments thin,
entire to slightly lacerate. Subiculum soft and cottony, white
to apricot-orange. Hyphal system monomitic to dimitic, gen-
erative hyphae mostly with clamps and thin-walled, occasion-
ally thick-walled, rarely thick-walled with simple septa, skel-
etal hyphae dominant in subiculum and rhizomorphs, solid to
thick-walled, rarely dichotomously branched, IKI–,CB–.
Cystidia absent, cystidioles present or absent. Basidiospores
oblong to broadly ellipsoid, slightly thick-walled, hyaline,
smooth, IKI–,CB–. Grows on angiosperm or gymnosperm
wood.
Fibroporia bambusae Yu a n Y. C h en & B . K . C u i , sp . n o v.
(Figs. 2b, c and 3)
MycoBank no.: MB 816304
Basidiomata annual, resupinate, with cream to pinkish buff
pore surface, cinnamon buff rhizomorphs. Hyphal system
dimitic; generative hyphae with both clamp connections and
simple septa. Long clavate cystidioles present. Basidiospores
oblong to broadly ellipsoid, hyaline, slightly thick-walled,
IKI–,CB–.
Holotype. CHINA. Hainan Province, Wuzhishan,
Wuzhishan Nature Reserve, on rotten bamboo, 15.XI.2015,
Dai 12610 (BJFC).
Etymology:bambusae (Lat.): refers to the habit of growing
on bamboo.
Basidiomata. Annual, resupinate and widely effused, soft
to cottony, without odor or taste when fresh, corky upon dry-
ing, up to 6 cm long, 4 cm wide, 1.5 mm thick at centre,
margin thin and white, finely fimbriate. Pore surface cream
to pinkish buff when fresh, becoming white to cream upon
drying; pores round to angular, 3–4per mm; dissepiments
thin, entire to slightly lacerate. Subiculum white and cottony,
up to 0.5 mm thick. Tubes concolorous with pore surface,
corky, up to 1 mm long. Rhizomorphs present, buff to cinna-
mon buff, penetrating deep into the substrate.
Hyphal structure.Hyphalsystemdimitic;generativehy-
phae bearing clamp connections or simple septa; skeletal hy-
phae IKI–,CB–; tissues unchanged in KOH.
Subiculum. Generative hyphae with clamp connections, hy-
aline, mostly thin-walled, occasionally slightly thick-walled,
rarely branched, 2–5.5 μm in diam; simple-septate generative
hyphae occasionally present, hyaline, thick-walled with a wide
lumen, unbranched, 3–4μm in diameter; skeletal hyphae hya-
line, thick-walled with a wide or narrow lumen, rarely
branched, flexuous, interwoven, 2.5–5μm in diameter.
Tubes. Generative hyphae dominant, with clamp connec-
tions, hyaline, thin-walled, occasionally branched, 2–5μmin
diameter; skeletal hyphae hyaline, thick-walled with a dis-
tinctly wide or narrow lumen, occasionally branched, flexu-
ous, interwoven, 4–6μm in diam. Cystidia absent; cystidioles
present, long clavate, hyaline, thin-walled, 18–22 × 2.5–4μm.
Basidia clavate, bearing four sterigmata and a basal clamp
Mycol Progress (2017) 16:521–533 525
connection, 13–24 × 5–6μm; basidioles in shape similar to
basidia, but slightly smaller.
Spores. Basidiospores oblong to broadly ellipsoid, hyaline,
slightly thick-walled, smooth, usually with a big guttule, IKI–,
CB–, (3.5)3.8–5 × (2.3)2.5–3μm, L = 4.15 μm, W = 2.8 μm,
Q = 1.45–1.5 (n = 90/3).
Additional specimens (paratypes)examined. CHINA.
Hainan Province, Wuzhishan, Wuzhishan Nature Reserve,
on rotten bamboo, 15.XI.2015, Dai 12609, 12611 & 12612
(BJFC).
Notes:Fibroporia bambusae is characterized by resupinate
basidiomata with cream to pinkish buff pore surface and cin-
namon buff rhizomorphs, a dimitic hyphal system with both
clamped and simple-septate generative hyphae, long clavate
cystidioles, and oblong to broadly ellipsoid basidiospores
(3.8–5 × 2.5–3μm). Fibroporia citrina may be confused with
F. bambusae by sharing the cream pore surface and yellow
rhizomorphs, but F. citrina has smaller pores (4–5 per mm,
Bernicchia et al. 2012) and only clamped generative hyphae.
Fibroporia ceracea Yuan Y. Chen & B.K. Cui, sp. nov.
(Figs. 2d, e and 4)
MycoBank MB no.: 816305
Basidiomata annual, resupinate; pore surface cream to pale
mouse-grey, ceraceous; rhizomorphs absent. Hyphal system
monomitic; generative hyphae thin-walled, with clamp
Fig. 1 Maximum likelihood tree illustrating the phylogeny of Fibroporia
and its related genera in the antrodia clade based on combined sequences
dataset of ITS+nLSU+nSSU+mtSSU+tef1+rpb2. Branches are labeled
with maximum likelihood bootstrap higher than 50%, parsimony
bootstrap proportions higher than 50% and Bayesian posterior
probabilities more than 0.95
526 Mycol Progress (2017) 16:521–533
connections. Cystidioles present, hyaline, fusoid.
Basidiospores oblong to broadly ellipsoid, slightly thick-
walled, hyaline, IKI–,CB–.
Holotype. CHINA. Yunnan Province, Kunming, on bark of
Pinus, 22.IV.2012, Dai 13013 (BJFC).
Etymology. ceracea (Lat.): refers to the ceraceous pore
surface.
Basidiomata. Annual, resupinate, soft corky with a
ceraceous surface, without odor or taste when fresh, corky
upon drying, up to 10 cm, 7.4 cm wide, 1 cm thick at centre,
margin entire to slightly fimbriate, white and corky. Pore sur-
face cream to pale mouse-grey when young, growing to pale
ash-grey in mature, becoming cinnamon-buff to greyish
brown upon drying; pores angular, 2–4 per mm; dissepiments
thin, entire. Subiculum white, corky, up to 0.5 mm thick.
Tubes concolorous with pore surface, ceraceous, up to
0.5 mm long. Rhizomorphs absent.
Hyphal structure. Hyphal system monomitic; generative
hyphae bearing clamp connections IKI–,CB–; tissues un-
changed in KOH.
Subiculum. Generative hyphae hyaline, thin-walled, fre-
quently branched, flexuous, interwoven, 2.5–5μmindiam.
Tubes. Generative hyphae hyaline, thin-walled, frequently
branched, interwoven, 2–4μm in diam. Cystidia absent;
cystidioles fusoid, hyaline, thin-walled, 15.5–19 × 5–6.5
μm. Basidia clavate, bearing four sterigmata and a basal
clamp connection, 18–23 × 6–7μm; basidioles in shape sim-
ilar to basidia, but slightly smaller.
Spores. Basidiospores oblong to broadly ellipsoid, hyaline,
slightly thick-walled, smooth, IKI–,CB–, (4)4.2–5(5.8) ×
2.5–3μm, L = 4.6 μm, W = 2.77 μm, Q = 1.66 (n = 60/1).
Notes:Fibroporia ceracea is characterized by resupinate
basidiocarps with cream to pale mouse-grey and ceraceous
pore surface, absence of rhizomorphs, a monomitic hyphal
system with clamped generative hyphae, fusoid cystidioles,
and oblong to broadly ellipsoid basidiospores (4.2–5 × 2.5–3
μm). It can be easily distinguished from other species of
Fibroporia by the entire to slightly fimbriate margin and a
monomitic hyphal system in both subiculum and trama.
Pseudofibroporia Yuan Y. Chen & B.K. Cui, gen. nov.
Fig. 2 Basidiomata of Fibroporia and Pseudofibroporia from China. (a).
Fibroporia albicans;(b,c). Fibroporia bambusae;(d,e). Fibroporia
ceracea;(f). Fibroporia citrina;(g). Fibroporia gossypium;(h).
Fibroporia radiculosa;(i). Pseudofibroporia citrinella.—Scale bars: a,
d,h=2cm;b=1.5cm;c=0.3cm;e,f,g=1cm;i=4cm
Mycol Progress (2017) 16:521–533 527
MycoBank MB no.: 816306
Basidiomata annual, pileate; pilei flabelliform to dimidiate,
fleshy when fresh, crumbly or chalky and light in weight
after drying. Pileal surface white to lemon yellow when
fresh, becoming cream to cinnamon buff upon drying.
Pore surface lemon yellow to curry buff when fresh, be-
coming cinnamon buff to clay buff upon drying. Pores
angular, dissepiments thin and entire. Context white to
lemon yellow, fleshy when fresh, cottony or corky after
drying. Tubes concolorous with pore surface, crumbly or
chalky after dying. Hyphal system dimitic in context,
monomitic in trama, generative hyphae with clamp connec-
tions, skeletal hyphae IKI–,CB–. Cystidia absent, but bottle-
shaped cystidioles occasionally present. Basidiospores oblong
to broadly ellipsoid, hyaline, slightly thick-walled, smooth,
IKI–,CB–. Grows on angiosperm wood.
Type species:Pseudofibroporia citrinella Yuan Y. Chen &
B.K. Cui.
Etymology. Pseudofibroporia (Lat.): refers to its close rela-
tionship with Fibroporia in phylogeny.
Pseudofibroporia citrinella Yuan Y. Chen & B.K. Cui, sp.
nov. (Figs. 2i and 5)
MycoBank no.: MB 816307
Basidiomata annual, pilei flabelliform to dimidiate, white to
lemon yellow when fresh, becoming cream to cinnamon buff
upon drying; margin acute. Pore surface lemon yellow to curry
buff. Hyphal system dimitic; generative hyphae with clamp
Fig. 3 Microscopic structures of
Fibroporia bambusae (drawn
from the holotype). (a).
Basidiospores; (b). Basidia and
basidioles; (c). Cystidioles; (d).
Hyphae from trama; (e). Hyphae
from subiculum. —Scale bars: a
=5μm; b–e=10μm
528 Mycol Progress (2017) 16:521–533
connections. Cystidioles bottle-shaped, hyaline. Basidiospores
oblong to broadly ellipsoid, slightly thick-walled, IKI–,CB–.
Holotype. CHINA. Guangxi Autonomous Region,
Longzhou, Nonggang Nature Reserve, on dead angiosperm
tree, 21.VII.2012, He 20120721-15 (BJFC).
Etymology. citrinella (Lat.): refers to the lemon yellow pore
surface.
Basidiomata. Annual, pileate, broadly attached, imbricate,
fleshy when fresh, crumbly or chalky and light in weight when
dry, pilei flabelliform to dimidiate, projecting up to 8 cm,
12 cm wide and 2 cm thick at base; pileal surface white to
lemon yellow when fresh, becoming cream to cinnamon buff
upon drying, rough, uneven; margin lemon yellow when
juvenile, becoming white with age, acute; pore surface lemon
yellow to curry buff when fresh, becoming cinnamon buff to
clay buff upon drying; sterile margin cinnamon buff to reddish
brown, up to 1 mm wide; pores angular, 3–4permm;dissep-
iments thin, entire; context white to lemon yellow, becoming
cream to cinnamon buff upon drying, fleshy, cottony or corky
after drying, up to 1.5 cm thick; tubes concolorous with pore
surface, crumbly or chalky, up to 5 mm long.
Hyphal structure. Hyphal system dimitic in context,
monomitic in trama; generative hyphae with clamp connec-
tions; skeletal hyphae IKI–,CB–; tissues unchanged in KOH.
Context. Generative hyphae frequent, with clamp connec-
tions, hyaline, thin- to thick-walled with a wide lumen, rarely
Fig. 4 Microscopic structures of
Fibroporia ceracea (drawn from
the holotype). (a). Basidiospores;
(b). Basidia and basidioles; (c).
Cystidioles; (d). Hyphae from
trama; (e). Hyphae from
subiculum. —Scale bars: a = 5
μm; b–e=10μm
Mycol Progress (2017) 16:521–533 529
branched, 3–7.5 μm in diameter; skeletal hyphae hyaline,
thick-walled with a wide or narrow lumen, occasionally
branched, interwoven, 2–5μmindiameter.
Tubes. Generative hyphae hyaline, thin-walled, occasionally
branched, 2.5–6μm in diam. Cystidia absent; cystidioles bottle
shaped, hyaline, thin-walled, 19–25 × 5–6μm. Basidia clavate,
bearing four sterigmata and a basal clamp connection, 17–20 ×
5–6μm; basidioles dominant, in shape similar to basidia but
slightly smaller.
Spores. Basidiospores oblong to broadly ellipsoid, hyaline,
slightly thick-walled, smooth, usually with a small guttule,
IKI–,CB–,(3.8)4–4.5(4.8) × 2.2–2.8(3) μm, L = 4.14 μm,
W = 2.5 μm, Q = 1.65–1.68 (n = 60/2).
Additional specimen (paratype)examined. CHINA,
Gugangxi Autonomous Region, Longzhou, Nonggang
Nature Reserve, on dead angiosperm tree, 21.VII.2012,
Yuan 6181 (IFP).
Notes:Pseudofibroporia citrinella is characterized by
flabelliform to dimidiate basidiomata, white to lemon yellow
pileal surface when fresh becoming cream to cinnamon buff
upon drying, lemon yellow to curry buff pores, acute margin,
a dimitic hyphal system in context and monomitic in trama
Fig. 5 Microscopic structures of
Pseudofibroporia citrinella
(drawn from the holotype). (a).
Basidiospores; (b). Basidia and
basidioles; (c). Cystidioles; (d).
Hyphae from trama; (e). Hyphae
from context. —Scale bars: a = 5
μm; b–e=10μm
530 Mycol Progress (2017) 16:521–533
with clamped generative hyphae, bottle shaped cystidioles,
and oblong to broadly ellipsoid, slightly thick-walled basidio-
spores (4–4.5 × 2.2–2.8 μm).
Other specimens examined:
Fibroporia albicans B.K. Cui & Yuan Y. Chen. CHINA.
Jiangxi Province, Jinggangshan, Jinggangshan Nature Reserve,
on rotten stump of Pinus, 23. IX.2008, Dai 10595 (BJFC);
Xizang Autonomous Region (Tibet), Bomi County, Yupu, on
stump of Pinus, 19.IX.2010, Cui 9464, 9495 & 9504 (BJFC).
Fibroporia bohemica Bernicchia, Vampola & Prodi.
CZECH REPUBLIC. Bohemia-Moravian Uplands,
27.VIII.2008, PRM 859138 (PRM).
Fibroporia citrina (Bernicchia & Ryvarden) Bernicchia &
Ryvarden. CHINA. Heilongjiang Province, Yichun, on root of
living tree of Pinus, 26.VIII.2014, Cui 11604 (BJFC); Xizang
Autonomous Region (Tibet), Linzhi, Sejila Mountain, on fall-
en trunk of Abices, 25.IX.2010, Cui 9683 (BJFC); Yunnan
Province, Weixi County, Laojunshan Nature Reserve, on fall-
en trunk of Picea, 22.IX.2011, Cui 10480 & 10497 (BJFC).
FRANCE. Fontoy, 6.IX.2011, LY BR 4206 (BJFC).
LUXEMBOURG. 28.VIII.2011, LY BR 4205 (BJFC).
Fibroporia gossypium.ARGENTINA.23.III.2010,LYBR
3394 (BJFC). CHINA. Xizang Autonomous Region (Tibet),
Bomi County, Yupu, on stump of Pinus, 19.IX.2010, Cui
9472 (BJFC); Milin County, on fallen branch of Cupressus
gigantean, l2.VIII.2004, Dai 5611 (BJFC). FRANCE.
Pledeliac, 5.X.2010, LY BR 3914 (BJFC).
Fibroporia radiculosa. CHINA. Fujian Province, Jianou,
Wanmulin Nature Reserve, on fallen trunk of Cunnihamia,
30.VIII.2006, Cui 4244 (BJFC); Hunan Province, Changsha,
Yuelu Mountain, on rotten wood of Pinus, 5.XII.2002, Dai
3577 (IFP); 14.VII.2011, Dai 12451 & 12454 (BJFC); Jilin
Province, Antu County, Changbaishan Nature Reserve, on
fallen trunk of Pinus, 9.X.2009, Dai 11404 (BJFC); Yunnan
Province, Kunming, Heilongtan Park, on stump of Pinus,
23.IX.2012, Dai 13041 (BJFC); Zhejiang Province, Linan,
Tianmushan Nature Reserve, on rotten stump of Pinus,
13.X.2005, Cui 2790, 2796, 2797 & 2812 (BJFC);
17.X.2004, Dai 6473 (BJFC).
Fibroporia vaillantii. CHINA. Jilin Province, Antu
County, Changbaishan Nature Reserve, on rotten wood of
Quercus, 18.IX.2002, Dai 3755 & 3759 (IFP & H);
26.VIII.2005, Dai 7002 (BJFC).
Discussion
In the phylogenetic analysis based on combined ITS+nLSU+
nSSU+mtSSU+tef1+rpb2 sequences, Pseudofibroporia was
suggested as the closest relative of Fibroporia, but morpho-
logically, Pseudofibroporia is different from Fibroporia in its
distinct pileate basidiomata with entire margin and lacking of
rhizomorphs. In the antrodia clade, Piptoporellus B.K. Cui,
Tab l e 2 A comparison of taxa in Fibroporia from China
Species F. albicans F. bambusae F. ceracea F. citrina F. gossypium F. radiculosa F. vaillantii
Substrate Pinus Bamboo Pinus Gymnosperm Gymnosperm Gymnosperm Gymnosperm
Pore surface in fresh White to cream Cream to pinkish buff Cream to ash-grey Cream to pale yellow White to cream Bright orange yellow White to cream
Margin Fimbriate Fimbriate Entire to slightly fimbriate Fimbriate Fimbriate Fimbriate Fimbriate
Rhizomorphs White to cream Buff to cinnamon buff Absent Bright yellow White Bright yellow White to cream
Pores (per mm) 6–83–42–44–53–61–22–4
Hyphal system in
trama
Dimitic, skeletal
hyphae dominant
Dimitic, generative
hyphae dominant
Monomitic Dimitic, generative
hyphae dominant
Monomitic Monomitic Dimitic, skeletal
hyphae dominant
Hyphal system in
Subiculum
Dimitic Dimitic Monomitic Dimitic Dimitic Dimitic Dimitic
Generative hyphae in
Subiculum
Clamped Clamped and simple septate Clamped Clamped Clamped Clamped Clamped
Cystidioles (μm) Fusoid, 10–16 × 3–3.5 Long clavate, 18–22 × 2.5–4 Fusoid, 15.5–19 × 5–6.5 Fusoid, 11–19.5 × 5–6 Absent Absent Fusoid, 20–25 × 4–6
Basidia (μm) 10–17 × 5–713–24 × 5–618–23 × 6–717–23 × 6.5–7.5 18–23 × 6–715–30 × 4–6.5 20–28 × 6–8
Basidiospores (μm) (3.5–)4–5.2(–6) × 3–3.8(–4) (3.5)3.8–5 × (2.3)2.5–3(4–)4.2–5(–5.8) × 2.5–3(4–)4.2–5(–5.8) × 2.5–34.5–6×2.2–2.8 6–8×3–45–7×3–4
Mycol Progress (2017) 16:521–533 531
M.L. Han & Y.C. Dai, Pycnoporellus Murrill, Postia Fr.,
Daedalea Pers., Fomitopsis P. Karst., Laetiporus Murrill and
Amylocystis Bondartsev & Singer all have distinct pileus, and
maybeconfusedwithPseudofibroporia.ButPiptoporellus is
different from Pseudofibroporia by its thin-walled basidio-
spores (Han et al. 2016); Pycnoporellus differs from
Pseudofibroporia in its monomitic hyphal system with mostly
encrusted and simple-septate generative hyphae (Ryvarden and
Melo 2014); Postia is different from Pseudofibroporia in its
monomitic hyphal system and thin-walled, allantoid to cylin-
drical or ellipsoid basidiospores (Wei and Dai 2006;Shenetal.
2015); Daedalea differs from Pseudofibroporia by its perennial
basidiocarps and light ochraceous brown skeletal hyphae
(Ryvarden and Melo 2014); Fomitopsis sensu lato is different
from Pseudofibroporia by its white to tan or pinkish pore sur-
face and context and thin-walled, subglobose to cylindrical ba-
sidiospores (Ryvarden and Melo 2014); Laetiporus can be sep-
arated from Pseudofibroporia by its dimitic hyphal system with
simple-septate generative hyphae (Ryvarden and Melo 2014;
Song et al. 2014); Amylocystis separates from Pseudofibroporia
by its monomitic hyphal system and amyloid generative hyphae
(Ryvarden and Melo 2014).
In our phylogenetic analysis, 39 samples representing ten
species of Fibroporia clustered together in one clade.
Combined with the morphological characters, two new spe-
cies of Fibroporia are described. Fibroporia bambusae is
closely related to F. radiculosa, but the latter has larger pores
(1–2 per mm) and baisidiospores (6–8×3–4μm) and a
monomitic hyphal system in tubes (Gilbertson and Ryvarden
1986;Chenetal.2015b). Fibroporia citrina produces cream
to pale yellow or lemon olivaceous basidiomata with conspic-
uous bright yellow rhizomorphs, which may be confused with
F. bambusae, but differs in its broader basidiospores (4–5×3–
3.5 μm, Bernicchia et al. 2012); moreover, the two species are
distinctly different in the phylogenetic analysis. Fibroporia
vaillantii is different from F. bambusae by having white to
cream rhizomorphs, bigger basidiospores (5–7×3–4μm,
Ryvarden and Gilbertson 1993) and a dimitic hyphal system
with skeletal hyphae dominating in tubes. Fibroporia
gossypium maybeconfusedwithF. ceracea by having white
to cream basidiomata, but differs in its distinctly fimbriate
margin, a dimitic hyphal system in subiculum and the soft to
cottony pore surface (Ryvarden and Gilbertson 1993).
Fibroporia albicans also has white to cream basidiomata,
and it can be separated from F. ceracea by its smaller pores
(6–8 per mm), broader basidiospores (4–5.2 × 3–3.8 μm,
Chen et al. 2015b) and a dimitic hyphal system in both tubes
and subiculum. So far, seven species of Fibroporia have been
recorded in China: F. a l b i c a n s ,F. bambusae,F. ceracea,
F. citrina,F. gossypium,F. radiculosa and F. vaillantii.The
major characteristics of the seven species are shown in
Tab le 2, and a key to known species of Fibroporia from
China is provided.
Key to known species of Fibroporia from China
1. Hyphal system monomitic .............................. F. ceracea
1. Hyphal system dimitic ................................................... 2
2. Rhizomorphs buff to bright yellow ................................ 3
2. Rhizomorphs white to cream ......................................... 5
3. Pores 1–2 per mm ....................................... F. radiculosa
3. Pores 3–5 per mm .......................................................... 4
4. Subicular hyphae bearing clamp connections and simple
septa ............................................................ F. bambusae
4. Subicular hyphae bearing clamp connections ... F. citrina
5. Generative hyphae dominant in trama ........ F. gossypium
5. Skeletal hyphae dominant in trama ............................... 6
6. Pores 6–8 per mm; basidiospores 4–5.2 × 3–3.8 μm
....................................................................... F. albicans
6. Pores 3–6 per mm; basidiospores 5–7×3–4μm
...................................................................... F. vaillantii
Acknowledgments The authors are grateful to Prof. Yu-Cheng Dai
(BJFC, China), Dr. Shuang-Hui He (BJFC, China) and Prof. Hai-Sheng
Yuan (IFP, China) for collecting specimens. The research was financed by
the Fundamental Research Funds for the Central Universities (Project
Nos. BLYJ201610 and 2016ZCQ04) and the National Natural Science
Foundation of China (Project Nos. 31422001 and 31170018).
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