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A Contribution to the Phylogeny and Taxonomy of Phlebiopsis (Polyporales, Basidiomycota)

October 2020

October 2020

DOI:10.21203/rs.3.rs-86969/v1

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CC BY 4.0

Authors:

Ya-Nan Zhao

Beijing Forestry University

Che-Chih Chen

National Museum of Natural Science

Shi-Liang Liu

Chinese Academy of Sciences

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An in-depth study of the phylogeny and taxonomy of the corticioid genus Phlebiopsis (Phanerochaetaceae) was conducted. Phylogenetic analyses of the ITS1-5.8S-ITS2 and nrLSU sequences demonstrated that Phlebiopsis is a strongly supported clade that is distinct from its sister clade that includes Phaeophlebiopsis , Hapalopilus and Rhizochaete . Two genera, Australohydnum and Hjortstamia , are reduced to synonyms under Phlebiopsis because generic type species A. griseofuscescens and H. friesii , respectively, are embedded in the Phlebiopsis clade. Twenty-four lineages are resolved in the ITS phylogenetic tree of Phlebiopsis , including six new taxa from Sri Lanka and China — P. albescens , P. brunnea , P. cylindrospora , P. magnicystidiata , P. membranacea and P. sinensis . Five new combinations are proposed — Phaeophlebiopsis mussooriensis , Phlebiopsis bambusicola , P. dregeana , P. griseofuscescens and P. novae-granatae . Phlebiopsis crassa is a morphological species complex with three distinct lineages. Phlebiopsis lamprocystidiata is determined to be a later synonym of P. darjeelingensis . The new taxa are described, illustrated, and compared and contrasted to morphologically similar species. An emended description of Phlebiopsis is provided along with an identification key to 27 accepted species.

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A Contribution to the Phylogeny and Taxonomy of

Phlebiopsis (Polyporales, Basidiomycota)

Ya-Nan Zhao

Beijing Forestry University

Karen K. Nakasone

Center for Forest Mycology Research

Che-Chi Chen

National Museum of Nature and Science: Kokuritsu Kagaku Hakubutsukan

Shi-Liang Liu

Institute of Microbiology Chinese Academy of Sciences

Yi-Feng Cao

Beijing Forestry University

Shuang-Hui He (  shuanghuihe@yahoo.com )

Beijing Forestry University https://orcid.org/0000-0003-4702-3034

Research

Keywords: corticioid fungi, Phanerochaetaceae, phlebioid fungi, white rot, wood-decaying fungi, six new

species, ve new combinations

DOI: https://doi.org/10.21203/rs.3.rs-86969/v1

License:   This work is licensed under a Creative Commons Attribution 4.0 International License. 

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Page 2/33

Abstract

An in-depth study of the phylogeny and taxonomy of the corticioid genus

Phlebiopsis

(Phanerochaetaceae) was conducted. Phylogenetic analyses of the ITS1-5.8S-ITS2 and nrLSU sequences

demonstrated that

Phlebiopsis

is a strongly supported clade that is distinct from its sister clade that

includes

Phaeophlebiopsis

Hapalopilus

and

Rhizochaete

. Two genera,

Australohydnum

and

Hjortstamia

are reduced to synonyms under

Phlebiopsis

because generic type species

A. griseofuscescens

and

friesii

, respectively, are embedded in the

Phlebiopsis

clade. Twenty-four lineages are resolved in the ITS

phylogenetic tree of

Phlebiopsis

, including six new taxa from Sri Lanka and China —

P. albescens

brunnea

P. cylindrospora

P. magnicystidiata

P. membranacea

and

P. sinensis

. Five new combinations are

proposed —

Phaeophlebiopsis mussooriensis

Phlebiopsis bambusicola

P. dregeana

P. griseofuscescens

and

P. novae-granatae

Phlebiopsis crassa

is a morphological species complex with three distinct

lineages.

Phlebiopsis lamprocystidiata

is determined to be a later synonym of

P. darjeelingensis

.The new

taxa are described, illustrated, and compared and contrasted to morphologically similar species. An

emended description of

Phlebiopsis

is provided along with an identication key to 27 accepted species.

Introduction

In 1978, the genus

Phlebiopsis

Jülich was erected for

Thelephora gigantea

Fr. that has effused,

ceraceous basidiomata with a smooth to odontioid hymenophore, a monomitic hyphal system with

hyaline, partially agglutinated, simple-septate hyphae, lamprocystidia with thick, hyaline walls, and

basidiospores with hyaline, thin, smooth walls that do not react in Melzer’s reagent or cotton blue (Jülich

1978; Bernicchia and Gorjón 2010). Over the next 40 years, 12 species with similar morphology were

described in or transferred to the genus by Hjortstam and Ryvarden (1980), Jülich and Stalpers (1980),

Dhingra (1987), Hjortstam (1987), Gilbertson & Adaskaveg (1993), Douanla-Meli and Langer (2009), Wu

et al. (2010), Priyanka et al. (2011), Kaur et al. (2015) and Zhao et al. (2018). Morphologically,

Phlebiopsis

is similar to

Scopuloides

(Massee) Höhn. & Litsch. and some species of

Phanerochaete

Karst. It was traditionally placed in the

Phanerochaete sensu lato

group (Rattan 1977; Burdsall 1985).

Although Burdsall (1985) considered

Phlebiopsis

and

Scopuloides

to be synonyms of

Phanerochaete

most researchers at the time recognized the genera as distinct (Eriksson et al. 1978, 1981, 1984; Jülich

and Stalpers 1980; Wu 1990).

The generic circumscription of

Phlebiopsis

was expanded when molecular studies showed that

Phanerochaete crassa

(Lév.) Burds. and

Phlebiopsis gigantea

(Fr.) Jülich were closely related (de Koker et

al. 2003; Greslebin et al. 2004; Wu et al. 2010; Floudas and Hibbett 2015). With the inclusion of

P. crassa

Phlebiopsis

now also includes species with effused-reexed, coriaceous basidiomata, a dimitic hyphal

system, and lamprocystidia or skeletocystidia with light brown walls. In addition, Floudas and Hibbett

(2015) created

Phaeophlebiopsis

Floudas & Hibbett to accommodate

Phlebiopsis peniophoroides

Gilb. &

Adask. and similar species with ceraceous, beige-brown basidiomata and subicula, lamprocystidia with

brown walls, and small basidiospores. The limits of the

Phlebiopsis

clade were extended by Miettinen et

al. (2016) who transferred six species into

Phlebiopsis

. The results of their phylogenetic study showed

Page 3/33

that the type species of

Castanoporus

Ryvarden,

Merulius castaneus

Lloyd, was nested in a clade with

gigantea

and, therefore, a synonym of

Phlebiopsis

. Similarly,

Dentocorticium pilatii

(Parmasto) Duehm &

Michel,

Lopharia papyrina

(Mont.) Boidin,

Phanerochaete brunneocystidiata

Sheng H. Wu, and

P. laxa

Sheng H. Wu clustered in the

Phlebiopsis

clade. Based on the morphological similarity of

Thelephora

friesii

Lév., the type of

Hjortstamia

Boidin & Gilles, to

L. papyrina

and

P. crassa

, they also transferred

friesii

Phlebiopsis

, thereby reducing

Hjortstamia

to a synonym of

Phlebiopsis

. Earlier, Boidin and Gilles

(2003) erected the genus

Hjortstamia

for

T. friesii

and related taxa with effused-reexed to pileate

basidiomata, a dimitic hyphal system of simple-septate hyphae and thick-walled skeletal hyphae,

lamprocystidia with thick walls. Ryvarden (1991) proposed

Castanoporus

for

M. castaneus

, a resupinate

species with yellowish brown to purple pores, a monomitic hyphal system with simple-septate hyphae,

and lamprocystidia with thick, hyaline walls.

Phlebiopsis pilatii

(Parmasto) Spirin & Miettinen is unique in

the genus for it has a dimitic hyphal system of simple-septate generative and microbinding (squeletto-

ligatives) hyphae and nely branched hyphidia but lacks lamprocystidia or skeletal cystidia (Larsen and

Gilbertson 1977; Duhem and Michel 2009).

With

Hjortstamia

and

Castanoporus

as synonyms,

Phlebiopsis

became a morphologically heterogeneous

genus with effused, effused-reexed or pileate basidiomata with a membranous, ceraceous, corneous or

coriaceous texture, hymenophore smooth to tuberculate, odontoid, or poroid, hyphal system monomitic or

dimitic with a loose to compact subiculum, and typically with lamprocystidia or skeletocystidia with

hyaline to brown walls. In phylogenetic analyses of Phanerochaetaceae,

Phlebiopsis

species are in a

clade sister to

Rhizochaete

Gresl., Nakasone & Rajchenb.,

Hapalopilus

P. Karst. and

Phaeophlebiopsis

, but

distant from

Phanerochaete sensu stricto

and

Scopuloides

(Floudas and Hibbett 2015; Miettinen et al.

2016).

Another genus of interest is

Australohydnum

Jülich for it is similar to

Phlebiopsis

by its warted, irpicoid to

hydnoid hymenophore, a dimitic hyphal system with hyaline, encrusted skeletocystidia, and thin-walled,

smooth basidiospores (Jülich 1978). The morphological similarities between

Australohydnum

and

Phanerochaete s.l.

were observed by Hjortstam and Ryvarden (1990). In a limited study of

Irpex s.s.

sequences of

Australohydnum dregeanum

(Berk.) Hjortstam & Ryvarden and

Irpex vellereus

Berk. &

Broome (a possible synonym of

A. dregeanum

) clustered together in a clade sister to two

Phanerochaete

species (Lim and Jung 2003). However, the phylogenetic relationship of

Australohydnum

within the

Phanerochaetaceae remained unknown (Miettinen et al. 2016).

Among the 24 names of

Phlebiopsis

recovered in Index of Fungorum (http://www.indexfungorum.org/,

accessed 21 June 2020), four species were transferred to

Phaeophlebiospis

. Of the remaining 20 species,

11 were described originally from Asia (e.g., Dhingra 1987; Wu 2000, 2004; Priyanka et al. 2011; Kaur et

al. 2015; Zhao et al. 2018; Xu et al. 2020). More than 150 specimens of

Phlebiopsis

were collected by the

corresponding author from China and Southeast Asia in recent years. Based on these specimens and

sequences obtained from GenBank, a phylogenetic analysis and taxonomic study of

Phlebiopsis

and

related taxa in the Phanerochaetaceae were undertaken. This study is a contribution to the understanding

of the diversity and phylogenetic relationships of crust fungi in China.

Page 4/33

Materials And Methods

Morphological studies

Voucher specimens are deposited at the herbaria of Beijing Forestry University, Beijing, China (BJFC),

Centre for Forest Mycology Research, U.S. Forest Service, Madison, Wisconsin, U.S.A. (CFMR) and

National Museum of Natural Science, Taichung, Taiwan, China (TNM). Freehand sections were made

from dried basidiomata and mounted in 2% (w/v) potassium hydroxide (KOH), 1% (w/v) phloxine,

Melzer’s reagent (IKI) or cotton blue (CB). Microscopic examinations were carried out with a Nikon Eclipse

80i microscope (Nikon Corporation, Japan) at magnications up to 1000 ×. Drawings were made with the

aid of a drawing tube. The following abbreviations are used: IKI– = neither amyloid nor dextrinoid, CB– =

acyanophilous, L = mean spore length, W = mean spore width, Q = L/W ratio, n (a/b) = number of spores

(a) measured from number of specimens (b). Color codes and names follow Kornerup and Wanscher

(1978).

DNA extraction and sequencing

A CTAB plant genomic DNA extraction Kit DN14 (Aidlab Biotechnologies Co., Ltd, Beijing, China) was

used to extract total genomic DNA from dried specimens then amplied by the polymerase chain reaction

(PCR), according to the manufacturer's instructions. The ITS1-5.8S-ITS2 region was amplied with the

primer pair ITS5/ITS4 (White et al. 1990) using the following protocol: initial denaturation at 95°C for

4min, followed by 34 cycles at 94°C for 40s, 58°C for 45s and 72°C for 1min, and nal extension at

72°C for 10min. The nrLSU D1-D2 region was amplied with the primer pair LR0R/LR7

(http://www.biology.duke.edu/fungi/mycolab/primers.htm) employing the following procedure: initial

denaturation at 94°C for 1min, followed by 34 cycles at 94°C for 30s, 50°C for 1min and 72°C for

1.5min, and nal extension at 72°C for 10min. DNA sequencing was performed at Beijing Genomics

Institute, and the sequences were deposited in GenBank (Table1). BioEdit v.7.0.5.3 (Hall 1999) and

Geneious Basic v.11.1.15 (Kearse et al. 2012) were used to review the chromatograms and for contig

assembly.

Page 5/33

Table 1

Species and sequences used in the phylogenetic analyses. New species are set in bold with type

specimens indicated with an asterisk (*).

Taxa Voucher Locality ITS nrLSU Reference

Bjerkandera adusta

HHB-12826-

Sp USA KP134983 KP135198 Floudas &

Hibbett 2015

B. centroamericana

L-13104-sp Costa

Rica KY948791 KY948855 Justo et al.

2017

Crepatura ellipsospora

CLZhao 1265 China MK343692 MK343696 Ma & Zhao

2019

Donkia pulcherrima

GC 1707-11 China LC378994 LC379152 Chen et al.

2018a

Geliporus exilisporus

Dai 2172 China KU598211 KU598216 Yuan et al.

2017

Hapalopilus eupatorii

Dammrich

10744 Germany KX752620 KX752620 Miettinen et al.

2016

H. percoctus

Miettinen

2008 Botswana KX752597 KX752597 Miettinen et al.

2016

H. nidulans

JV0206/2 Sweden KX752623 KX752623 Miettinen et al.

2016

Hyphodermella

corrugata

MA-Fungi

5527 Morocco FN600372 JN939597 Telleria et al.

2010

H. poroides

Dai 10848 China KX008368 KX011853 Zhao et al.

2017

H. rosae

FP-150552 USA KP134978 KP135223 Floudas &

Hibbett 2015

Irpex vellereus

CBS 515.92 India AF479670 — Lim & Jung

2003

Odontoebula

orientalis

GC 1703-76 China LC379004 LC379156 Chen et al.

2018a

Oxychaete

cervinogilvus

Schigel-5216 Australia KX752596 KX752596 Miettinen et al.

2016

Phaeophlebiopsis

caribbeana

HHB-6990 USA KP135415 KP135243 Floudas &

Hibbett 2015

P. himalayensis

He 3854 China MT386378 MT447410 present study

P. peniophoroides

FP-150577 USA KP135417 KP135273 Floudas &

Hibbett 2015

P. ravenelii

CBS 411.50 France MH856691 MH868208 Vu et al. 2019

Page 6/33

Taxa Voucher Locality ITS nrLSU Reference

P. ravenelii

FCUG 2216 France — GQ470674 Wu et al. 2010

Phanerina mellea

Miettinen

11393 Indonesia KX752602 KX752602 Miettinen et al.

2016

Phanerochaete

arizonica

RLG-10248-

Sp USA KP135170 KP135239 Floudas &

Hibbett 2015

P. australis

HHB-7105-Sp USA KP135081 KP135240 Floudas &

Hibbett 2015

P. bambusicola

Wu 0707-2 China MF399404 MF399395 Wu et al.

2018a

P. brunnea

He 1873 China KX212220 KX212224 Liu & He 2016

P. burtii

HHB-4618-Sp USA KP135117 KP135241 Floudas &

Hibbett 2015

P. canobrunnea

CHWC 1506-

66 China LC412095 LC412104 Wu et al.

2018b

P. carnosa

HHB-9195 USA KP135129 KP135242 Floudas &

Hibbett 2015

P. chrysosporium

HHB-6251-Sp USA KP135094 KP135246 Floudas &

Hibbett 2015

P. citrinosanguinea

FP-105385-

Sp USA KP135100 KP135234 Floudas &

Hibbett 2015

P. concrescens

Spirin 7322 Russia KP994380 KP994382 Volobuev et al.

2015

P. cumulodentata

LE 298935 Russia KP994359 KP994386 Volobuev et al.

2015

P. cystidiata

Wu 1708 −

326 China LC412097 LC412100 Wu et al.

2018b

P. ericina

HHB-2288 USA KP135167 KP135247 Floudas &

Hibbett 2015

P. incarnata

WEI 16–075 China MF399406 MF399397 Wu et al.

2018a

P. inata

Dai 10376 China JX623929 JX644062 Jia et al. 2014

P. laevis

HHB-15519 USA KP135149 KP135249 Floudas &

Hibbett 2015

P. livescens

FD-106 USA KP135070 KP135253 Floudas &

Hibbett 2015

P. magnoliae

HHB-9829-Sp USA KP135089 KP135237 Floudas &

Hibbett 2015

Page 7/33

Taxa Voucher Locality ITS nrLSU Reference

P. porostereoides

He 1902 China KX212217 KX212221 Liu & He 2016

P. pseudomagnoliae

PP-25 South

Africa KP135091 KP135250 Floudas &

Hibbett 2015

P. pseudosanguinea

FD-244 USA KP135098 KP135251 Floudas &

Hibbett 2015

P. rhodella

FD-18 USA KP135187 KP135258 Floudas &

Hibbett 2015

P. robusta

Wu 1109-69 China MF399409 MF399400 Wu et al.

2018a

P. sanguinea

HHB-7524 USA KP135101 KP135244 Floudas &

Hibbett 2015

P. sanguineocarnosa

FD-359 USA KP135122 KP135245 Floudas &

Hibbett 2015

P. sordida

FD-241 USA KP135136 KP135252 Floudas &

Hibbett 2015

P. stereoides

He 2309 China KX212219 KX212223 Liu & He 2016

P. subceracea

FP-105974-R USA KP135162 KP135255 Floudas &

Hibbett 2015

P. taiwaniana

Wu 0112 − 13 China MF399412 MF399403 Wu et al.

2018a

P. velutina

Kotiranta

25567 Russia KP994354 KP994387 Volobuev et al.

2015

Phlebia rma

Edman K268 Sweden EU118654 EU118654 Larsson 2007

P. lilascens

FCUG 2005 — AF141622 AF141622 —

Phlebiopsis albobadia He 5805 * Sri Lanka MT452526 — present study

P. amethystea

URM 93248 Brazil MK993644 MK993638 Xavier de Lima

et al. 2020

P. amethystea

URM 84741 Brazil MK993645 MK993639 Xavier de Lima

et al. 2020

P. brunnea He 5822 * Sri Lanka MT452527 MT447451 present study

P. brunneocystidiata

Chen 666 China MT561707 GQ470640 Wu et al. 2010,

present study

P. brunneocystidiata

Chen 1143 China — GQ470639 Wu et al. 2010

P. castanea

Spirin-5295 Russia KX752610 KX752610 Miettinen et al.

2016

Page 8/33

Taxa Voucher Locality ITS nrLSU Reference

P. castanea

GC 1612-6 China KY688208 — Chen et al.

2018b

P. castanea

CLZhao 3501 China MK269230 — —

P. castanea

He 2489 China — MT447406 present study

P. crassa

group A He 5205 Vietnam MT452523 MT447448 present study

P. crassa

group A He 5763 Sri Lanka MT452524 MT447449 present study

P. crassa

group A He 5855 China MT452525 MT447450 present study

P. crassa

group A He 6304 China MT561714 MT598029 present study

P. crassa

group A Wu 0504 − 22 China MT561715 GQ470634 Wu et al. 2010,

present study

P. crassa

group B He 3349 China MT561712 MT447407 present study

P. crassa

group B He 5866 China MT386376 MT447408 present study

P. crassa

group B He 6266 China MT561713 MT598035 present study

P. crassa

group B CLZhao 724 China MG231790 — —

P. crassa

group B MAFF

420737 Japan AB809163 AB809163 —

P. crassa

group C KKN-86-Sp USA KP135394 KP135215 Floudas &

Hibbett 2015

P. crassa

group C FP-102496-

sp USA AY219341 — de Koker et al.

2003

P. crassa

group C HHB 8834 USA KP135393 — Floudas &

Hibbett 2015

P. crassa

group C ME 516 USA KP135395 — Floudas &

Hibbett 2015

P. cf. dregeana

SFC 980804-

4Korea AF479669 — Lim & Jung

2003

P. cf. dregeana

UOC-DAMIA-

D46 Sri Lanka KP734203 — —

P. cf. dregeana

FLAS-F-

60030 USA KY654737 — —

P. avidoalba

FD-263 USA KP135402 KP135271 Floudas &

Hibbett 2015

P. avidoalba

Miettinen

17896 USA KX752607 KX752607 Miettinen et al.

2016

Page 9/33

Taxa Voucher Locality ITS nrLSU Reference

P. avidoalba

CFMR4167 USA KX065957 — —

P. avidoalba

HHB-4617 USA KP135401 —Floudas &

Hibbett 2015

P. friesii

He 5722 Sri Lanka MT452528 MT447413 present study

P. friesii

He 5817 Sri Lanka MT452529 MT447414 present study

P. friesii

He 5820 Sri Lanka MT452530 MT447415 present study

P. gigantea

He 5290 China MT386381 MT447416 present study

P. gigantea

Miettinen

15354 Finland KX752605 —Miettinen et al.

2016

P. gigantea

CBS 935.70 Germany MH860011 MH871798 Vu et al. 2019

P. gigantea

FP-70857-Sp USA KP135390 KP135272 Floudas &

Hibbett 2015

P. griseofuscescens

He 5734 Sri Lanka MT561708 MT598032 present study

P. griseofuscescens

Cui 12629 China MT561718 — present study

P. griseofuscescens

CLZhao 3692 China MT180946 MT180950 Xu et al. 2020

P. griseofuscescens

CLZhao 3705 China MT180947 MT180951 Xu et al. 2020

P. laxa

Wu 9311-17 China MT561710 GQ470649 Wu et al. 2010,

present study

P. magnicystidiata He 5648 * China MT386377 MT447409 present study

P. magnicystidiata He

20140719_18 China MT561719 —present study

P. magnicystidiata Wu 890805-1 China MT561711 GQ470667 Wu et al. 2010,

present study

P. membranacea He 3842 China MT386400 MT447440 present study

P. membranacea He 3849 * China MT386401 MT447441 present study

P. membranacea He 6062 China MT386402 MT447442 present study

P. pilatii

He 5114 China MT386385 MT447421 present study

P. pilatii

He 5165 China MT386386 MT447422 present study

P. pilatii

Dai 17041 China KY971603 KY971604 Wu et al. 2017

P. pilatii

Spirin 5048 Russia KX752590 KX752590 Miettinen et al.

2016

Page 10/33

Taxa Voucher Locality ITS nrLSU Reference

P. sinensis He 4295 China MT386395 MT447433 present study

P. sinensis He 4665 China MT386396 MT447434 present study

P. sinensis He 4673 * China MT386397 MT447435 present study

P. sinensis He 5662 China MT386398 MT447436 present study

sp. FP-102937 USA KP135391 KP135270 Floudas &

Hibbett 2015

sp. ECS1971 USA KP135392 — Floudas &

Hibbett 2015

sp. He 3827 China — MT447437 present study

P. yunnanensis

He 2623 China MT386387 MT447423 present study

P. yunnanensis

He 3249 China MT386375 MT447425 present study

P. yunnanensis

CLZhao 3958 China MH744140 MH744142 Zhao et al.

2018

P. yunnanensis

CLZhao 3990 China MH744141 MH744143 Zhao et al.

2018

Pirex concentricus

OSC-41587 USA KP134984 KP135275 Floudas &

Hibbett 2015

Porostereum fulvum

LY: 18496 France MG649453 MG649455 —

P. spadiceum

CBS 474.48 France MH856438 MH867984 Vu et al. 2019

Rhizochaete

americana

FP-102188 USA KP135409 KP135277 Floudas &

Hibbett 2015

R. belizensis

FP-150712 Belize KP135408 KP135280 Floudas &

Hibbett 2015

R. brunnea

MR 229 Argentina AY219389 AY219389 Greslebin et al.

2004

R. violascens

KHL 11169 Norway EU118612 EU118612 Larsson 2007

R. lamentosa

HHB-3169-Sp USA KP135410 KP135278 Floudas &

Hibbett 2015

R. ava

PR 1141 Puerto

Rico KY273030 KY273033 Nakasone et

al. 2017

R. fouqueriae

KKN-121-sp USA KY948786 KY948858 Justo et al.

2017

R. radicata

FD-123 USA KP135407 KP135279 Floudas &

Hibbett 2015

Page 11/33

Taxa Voucher Locality ITS nrLSU Reference

R. sulphurina

HHB-5604 USA KY273031 GU187610 Binder et al.

2010

R. sulphurosa

URM 87190 Brazil KT003522 KT003519 Chikowski et

al. 2015

Riopa metamorphosa

Spirin 2395 Russia KX752601 KX752601 Miettinen et al.

2016

R. pudens

Cui 3238 China JX623931 JX644060 Jia et al. 2014

Terana caerulea

FP-104073 USA KP134980 KP135276 Floudas &

Hibbett 2015

Outgroup     

Ceraceomyces serpens

HHB-15692-

Sp USA KP135031 KP135200 Floudas &

Hibbett 2015

Phebia acerina

FD-301 USA KP135378 KP135260 Floudas &

Hibbett 2015

Phylogenetic analyses

Two separate datasets, the concatenated ITS-nrLSU sequences of species in the Phanerochaetaceae and

ITS only sequences of

Phlebiopsis

, were analyzed.

Ceraceomyces serpens

(Tode) Ginns and

Phlebia

acerina

Peck were selected as the outgroup for the ITS-LSU dataset, whilst

Rhizochaete radicata

(Henn.)

Gresl., Nakasone & Rajchenb. was used in the ITS dataset (Floudas and Hibbett 2015). For the

concatenated dataset, the sequences of ITS and nrLSU were aligned separately using MAFFT v.7 (Katoh

et al. 2017, http://mafft.cbrc.jp/alignment/server/) with the G-INS-I iterative renement algorithm, and

optimized manually in BioEdit v.7.0.5.3. The separate alignments were then concatenated using Mesquite

v.3.5.1 (Maddison and Maddison 2018). The datasets were deposited in TreeBase

(http://treebase.org/treebase-web/home.html, submission ID: 26529 for Phanerochaetaceae ITS-LSU,

26530 for

Phlebiopsis

ITS).

Maximum parsimony (MP), Maximum likelihood (ML) analyses and Bayesian inference (BI) were carried

out by using PAUP* v.4.0b10 (Swofford 2002), RAxML v.8.2.10 (Stamatakis 2014) and MrBayes 3.2.6

(Ronquist et al. 2012), respectively. In MP analysis, trees were generated using 100 replicates of random

stepwise addition of sequence and tree-bisection reconnection (TBR) branch-swapping algorithm with all

characters given equal weight. Branch supports for all parsimony analyses were estimated by performing

1000 bootstrap replicates with a heuristic search of 10 random-addition replicates for each bootstrap

replicate. In ML analysis, statistical support values were obtained using rapid bootstrapping with 1000

replicates, with default settings used for other parameters. For BI, the best-t substitution model was

estimated with jModeltest v.2.17 (Darriba et al. 2012). Four Markov chains were run for 5,000,000 and

3,000,000 generations for the Phanerochaetaceae ITS-LSU and

Phlebiopsis

ITS datasets, respectively,

until the split deviation frequency value was lower than 0.01. Trees were sampled every 100th generation.

Page 12/33

The rst quarter of the trees, which represented the burn-in phase of the analyses, were discarded, and the

remaining trees were used to calculate posterior probabilities (BPP) in the majority rule consensus tree.

Results

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-t 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%, rst), 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

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, rst as small patches, later conuent 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),

Page 13/33

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 conuent, 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

Page 14/33

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-reexed

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

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, rst as small patches, later conuent 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 mbriate, 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

Page 15/33

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-reexed 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 nely 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).

Page 16/33

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

avidoalba

(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. avidoalba

from

India (Rattan 1977) and Taiwan (Wu 1990) need to be conrmed 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).

Page 17/33

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 nely cracked with age; margin thinning out, mbriate,

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.

Page 18/33

Fruiting body – Basidiomata annual, resupinate to effused-reexed with reexed edges elevated and

incurved with age, loosely adnate, easily detached from substrate, coriaceous, rst as small patches, later

conuent 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 nely cracked with age; margin thinning out,

distinct, white to gray, silky, slightly mbriate, 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-reexed, 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-reexed 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

Page 19/33

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

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

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.

Page 20/33

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 signicantly 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 Scientic and

Industrial Research 164: 97, 1965.

Page 21/33

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-reexed 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

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

griseofuscescens

. Based on morphological studies and sequence analyses, we determined that

lacerata

described from southern China (Xu et al. 2020) is conspecic 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

Page 22/33

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

Phaeophlebiopsis

based on morphology, our

phylogenetic analyses show that it belongs to

Phlebiopsis

s.s., however.

Discussion

Page 23/33

The generic limits of

Phlebiopsis

has expanded over the last 40years since its introduction in 1978 to

include signicant morphological range in basidiomata habit and texture and hymenophore conguration

with the aid of molecular phylogenetic methods (e.g., Floudas and Hibbett 2015; Miettinen et al. 2016;

Zhao et al. 2018; Xavier de Lima et al. 2020; Xu et al. 2020). In this study, we emphasized sampling of

Phlebiopsis

taxa, and our overall results conrm those of Floudas and Hibbett (2015), Miettinen et al.

(2016), and Chen et al. (2018a). In Figs.1 and 2,

Phlebiopsis

, including the types of

Australohydnum

griseofuscescens

and

Hjortstamia

P. friesii

, formed a well-supported clade in the Phanerochaetaceae and

is closely related to

Phaeophlebiopsis

Hapalopilus

and

Rhizochaete

. The genera

Phlebiopsis

and

Australohydnum

were published simultaneously (Jülich 1978) but the former is favored to avoid

unnecessary name changes. Twenty-four lineages were resolved in the ITS tree of

Phlebiopsis

, among

which 18 are accepted species, including the

P. crassa

species complex and six new species described

herein. Further study is required to identify the taxa named

. cf.

dregeana

Irpex vellerus

Phlebiopsis

sp.

FP-102937 and

Phlebiopsis

sp. ECS-1971.

Among the 24 names of

Phlebiopsis

in Index Fungorum (accessed on 21 June 2020), we accept 17 taxa

Phlebiopsis s.s.

, including 11 that are supported by molecular data. Five taxa,

P. himalayensis

Dhingra,

P. mussooriensis

P. peniophoroides

Gilb. & Adask.,

P. ravenelii

(Cooke) Hjortstam, and

P. roumeguerei

(Bres.) Jülich & Stalpers were transferred to

Phaeophlebiopsis

based on morphology and sequence data.

Phlebiopsis lacerata

and

P. lamprocystidiata

are synonyms of

P. griseofuscescens

and

P. darjeelingensis

respectively, as discussed above. Thus, 27 species of

Phlebiopsis

worldwide are accepted, including the

six new species and four new combinations reported herein. An emended description of

Phlebiopsis

and

an identication key to all species in the genus worldwide are presented below.

Phlebiopsis (Jülich) Nakasone & S.H. He, emended

Castanoporus

Ryvarden, Synopsis Fungorum 5: 121, 1991.

Hjortstamia

Boidin & Gilles, Bulletin de la Société Mycologique de France 118 (2): 99, 2003.

Australohydnum

Jülich, Persoonia 10 (1): 138, 1978.

Description: Basidiomata annual, resupinate, effused, effused-reexed or pileate, ceraceous,

membranaceous to coriaceous. Pilei, when present, tomentose, gray to brown. Hymenophore smooth,

tuberculate, odontioid, hydnoid to poroid, white, gray, grayish brown, purplish brown or brown, turning

purple in KOH in two species. Hyphal system monomitic or dimitic; generative hyphae simple-septate,

hyaline or rarely pale brown, in dimitic species with skeletal or, in one species, micro-binding hyphae.

Subiculum absent to well-developed, hyaline, brown, agglutinated or not, compact to loosely interwoven.

Skeletocystidia absent or present, hyaline or brown, distinctly thick-walled, smooth or encrusted.

Hymenial cystidia or lamprocystidia typically present, hyaline or light brown, thick-walled, usually

encrusted. Dendrohyphidia present in one species, hyaline, thin-walled, smooth, branched. Basidia clavate

or subcylindrical, with 4 sterigmata and a basal simple septum. Basidiospores cylindrical, ellipsoid,

broadly ellipsoid or subglobose, hyaline, thin-walled, smooth, negative in Melzer’s reagent, acyanophilous.

Page 24/33

Type species:

Phlebiopsis gigantea

(Fr.) Jülich

Notes – The terminology relating to the cystidia observed in

Phlebiopsis

species is varied in the literature

and thus confusing. There are up to three kinds of cystidia, but intermediate forms can develop to blur

their distinctiveness. Lamprocystidia are found in most species of

Phlebiopsis

in the hymenium, often

projecting, and may become embedded as the basidiomata thickens. They are typically conical or

subfusiform with thick walls that are lightly to heavily encrusted in the upper half or apex. Skeletocystidia

are found in dimitic or pseudodimitic species in which thick-walled hyphae in the subiculum curve toward

the hymenium but remain embedded in the subiculum or subhymenium. The terminal ends may or may

not be differentiated and usually lack encrustations. Hymenial cystidia are those structures that are

similar to skeletocystidia but terminate in the hymenium and may be encrusted. In other cases, they are

formed in the subhymenium and are smaller than lamprocystidia and not conical or heavily encrusted.

Key to 27

Phlebiopsis

species

1. Hymenophore poroid, irpicoid or hydnoid........... 2

1.Hymenophore smooth, tuberculate or odontioid........... 4

2.Basidiomata resupinate; hymenophore poroid to irpicoid; on gymnosperms........

P. castanea

2.Basidiomata effused-reexed; hymenophore tuberculate to hydnoid; on angiosperms........ 3

3.Basidiospores 6.5–8 × 4–5 µm; African species.................

P. dregeana

3.Basidiospores 4.5–6 × 2.5–3 µm; Asian and Australasian species...........

P. griseofuscescens

4.Dendrohyphidia present...................

P. pilatii

4.Dendrohyphidia absent................. 5

5. Hyphal system pseudodimitic or dimitic.................... 6

5. Hyphal system monomitic.................. 13

6. Hymenophore turning purple in KOH...............

P. friesii

6. Hymenophore unchanged in KOH...................... 7

7. Basidiomata with well-developed pilei; skeletocystidia absent...........

P. papyrina

7. Basidiomata resupinate to effused-reexed; skeletocystidia present............ 8

8. Hymenophore without purple tints.................... 9

8. Hymenophore more or less purple..................... 12

Page 25/33

9. Lamprocystidia none; basidiospores ≤ 6 µm long.....................

P. membranacea

9. Lamprocystidia present; basidiospores ≥ 6 µm long...................... 10

10. On angiosperms and gymnosperms; from temperate China...................

P. sinensis

10. On angiosperms or bamboo; from tropical-subtropical Asia or Australia.............. 11

11. Basidiospores 6–7 × 2.5–3 µm; on bamboo; from Australia....................

P. bambusicola

11. Basidiospores 6.5–7.5 × 3–3.6 µm; on angiospermous wood; from Sri Lanka.........

P. brunnea

12. Lamprocystidia brown to dark brown; South American species................

P. amethystea

12. Lamprocystidia cystidia hyaline to pale brown; North American or Asian species...

P. crassa s.l.

13. Lamprocystidia none; skeletocystidia or hymenial cystidia present................. 14

13. Lamprocystidia present; skeletocystidia absent................ 15

14. Basidiospores 5.5–7 × 3–4 µm; on bamboo; from Colombia...............

P. novae-granatae

14. Basidiospores 3.7–5.5 × 2.5–3.3 µm; on hardwood; from New Zealand............

P. abulata

15. Basidiospores > 8 µm long, > 4 µm broad..................

P. laxa

15. Basidiospores < 8 µm long, < 4 µm broad................. 16

16. Lamprocystidia small, generally < 40 µm long............... 17

16. Lamprocystidia large, generally > 40 µm long................ 20

17. Hymenophore purple in KOH............

P. cylindrospora

17. Hymenophore unchanged in KOH............. 18

18. Basidiospores broadly ellipsoid, 3.5–4.5 × 2.5–3.5 µm, Q = 1.3...........

P. yunnanensis

18. Basidiospores narrowly ellipsoid to cylindrical................ 19

19. Basidiospores 3.5–5 × 2–2.2 µm................

P. albescens

19. Basidiospores 5.3–8.5 × 2.5–4 μm..............

P. punjabensis

20 .Lamprocystidia brown; on Pandanaceae; from Taiwan............

P. brunneocystidiata

20. Lamprocystidia hyaline; on other plants; from various locations....................... 21

Page 26/33

21. Subiculum indistinct to absent........................ 22

21 .Subiculum distinct to well-developed................... 24

22. Basidia mostly with 2 sterigmata................

P. bicornis

22. Basidia with 4 sterigmata.................... 23

23.Basidiospores 5.5–7.5 × 3.5–4.5 µm; from North and South America..........

P. avidoalba

23. Basidiospores 4.5–6.5 × 3.8–4.8 µm; from Asia................

P. magnicystidiata

24. Basidiospores narrowly ellipsoid to ellipsoid, ≤ 3 µm broad................. 25

24. Basidiospores broadly ellipsoid, ≥ 4 µm broad.................... 26

25. Hymenophore smooth, pale orange to rosy; lamprocystidia 40–50 × 6–7 µm; basidiospores < 2.5 µm

wide; from Argentina...............

P. erubescens

25. Hymenophore smooth to tuberculate, pale white to gray; lamprocystidia 60–90 × 10–20 µm;

basidiospores ≥ 2.5 µm wide; from Northern Hemisphere...........

P. gigantea

26. Lamprocystidia < 10 µm wide; from South America.............

P. galochroa

26. Lamprocystidia > 10 µm wide; from Asia.............

P. darjeelingensis

Conclusions

An in-depth study of the phylogeny and taxonomy of the genus

Phlebiopsis

was conducted based on

both of morphological and molecular evidence. The results show that species of

Phlebiopsis

formed a

monophyletic clade in the Phanerochaetaceae but are morphologically heterogeneous with

Australohydnum

Castanoporus

and

Hjortstamia

as its synonyms. The species diversity of the genus is

high in subtropical and tropical areas with several new species described from East Asia. The present

study also show that the DNA sequence data are very useful in exploring cryptic taxa and diversity of

corticioid fungi, a large and widely distributed group poorly studied while compared with mushrooms and

polypores.

List Of Abbreviations

ITS: internal transcribed spacer; nrLSU: nuclear ribosomal large subunit; BJFC: herbarium of Beijing

Forestry University, Beijing, China; CFMR: Centre for Forest Mycology Research, U.S. Forest Service,

Madison, Wisconsin, U.S.A.; TNM: National Museum of Natural Science, Taichung, Taiwan, China; KOH:

2% (w/v) potassium hydroxide; IKI: Melzer’s reagent; CB: cotton blue; IKI–: neither amyloid nor dextrinoid;

CB–: acyanophilous; L: mean spore length; W: mean spore width; Q: L/W ratio, n (a/b): number of spores

Page 27/33

(a) measured from number of specimens (b); CTAB: cetyltrimethylammonium bromide; DNA:

deoxyribonucleic acid; PCR: polymerase chain reaction; MP: maximum parsimony; ML maximum

likelihood; BI: Bayesian inference; TBR: tree-bisection reconnection; BPP: Bayesian posterior probability.

Declarations

Ethics approval and consent to participate

Not applicable.

Adherence to national and international regulations

Not applicable.

Consent for publication

Not applicable.

Availability of data and materials

Details of the availability of the data and materials used in this study can be found within the text. DNA

sequences were submitted to NCBI Genbank database (Table 1). Alignments were deposited at TreeBase

(submission ID: 26529 for Phanerochaetaceae ITS-LSU, 26530 for

Phlebiopsis

ITS). Voucher specimens

are deposited in the herbaria listed in the materials and methods section.

Competing interests

The authors declare that they have no competing interests

Funding

Financial support was provided by the National Natural Science Foundation of China (Nos. 31870011 &

31750001).

Authors' contributions

Shuang-Hui He designed the research, collected most of the specimens, and wrote the text. Ya-Nan Zhao

performed the phylogenetic analyses and did most of the measurement, descriptions and illustrations.

Karen K. Nakasone loaned and examined type specimens of some related species, and revised language

of the text. Che-Chi Chen provided with some specimens and sequences. Shi-Liang Liu and Yi-Feng Cao

helped in eld trips and species illustrations.

Acknowledgment

Page 28/33

The authors would like to express their deep appreciation to Profs. Sheng-Hua Wu (National Museum of

Natural Science, Taiwan, China), Tie-Zhi Liu (Chifeng University, Inner Mongolia, China), and Bao-Kai Cui

(Beijing Forestry University, Beijing, China) for allowing us to study their specimens. Dr. Hai-Xia Ma

(Institute of Tropical Bioscience and Biotechnology, Chinese Academy of Tropical Agricultural Sciences,

Hainan, China) provided us with specimens.

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Phlebiopsis lacerata sp. nov. (Polyporales, Basidiomycota) from southern China

Article

Full-text available

Apr 2020

A new white-rot species, Phlebiopsis lacerata sp. nov., is proposed based on morphological and molecular characters. It is characterized by resupinate to effuse-reflexed basidiomata, a monomitic hyphal system with simple septate generative hyphae, conical cystidia that are thick-walled, heavily encrusted with large crystal in the apical part, and ellipsoid basidiospores (4–5.5 × 3–3.5 µm). Phylogenetic analyses of the ITS and LSU nrRNA gene regions showed that P. lacerata belongs to the Phanerochaetaceae and is nested in the phlebioid clade. Further investigation based on ITS+nLSU sequences with more representative taxa in Phlebiopsis demonstrated that P. lacerata forms a monophyletic lineage with a strong support (100% BS, 100% BP, 1.00 BPP) and then groups with P. crassa.

Three new species of Phanerochaete (Polyporales, Basidiomycota)

Article

Full-text available

Oct 2018

Phanerochaete canobrunnea, P. cystidiata and P. fusca are presented as new species, supported by morphological studies and two sets of phylogenetic analyses. The 5.8S+nuc 28S+rpb1 dataset shows the generic placement of the three species within the phlebioid clade of Polyporales. The ITS+nuc 28S dataset displays relationships for the new taxa within Phanerochaete s.s. Phanerochaete canobrunnea grew on angiosperm branches in subtropical Taiwan and is characterised by greyish brown hymenial surface, brown genera-tive hyphae and skeletal hyphae and absence of cystidia. Phanerochaete cystidiata grew on angiosperm branches above 1000 m in montane Taiwan and SW Yunnan Province of China and is characterised by cream to yellowish hymenial surface and more or less encrusted leptocystidia. Phanerochaete fusca grew on angiosperm branches at 1700 m in Hubei Province of China and is characterised by dark brown hymenial surface, leptocystidia, brown subicular hyphae and colourless to brownish basidiospores.

Large-scale generation and analysis of filamentous fungal DNA barcodes boosts coverage for kingdom Fungi and reveals thresholds for fungal species and higher taxon delimitation

Article

Full-text available

May 2018

Species identification lies at the heart of biodiversity studies that has in recent years favoured DNA-based approaches. Microbial Biological Resource Centres are a rich source for diverse and high-quality reference materials in microbiology, and yet the strains preserved in these biobanks have been exploited only on a limited scale to generate DNA barcodes. As part of a project funded in the Netherlands to barcode specimens of major national biobanks, sequences of two nuclear ribosomal genetic markers, the Internal Transcribed Spaces and 5.8S gene (ITS) and the D1/D2 domain of the 26S Large Subunit (LSU), were generated as DNA barcode data for ca. 100 000 fungal strains originally assigned to ca. 17 000 species in the CBS fungal biobank maintained at the Westerdijk Fungal Biodiversity Institute, Utrecht. Using more than 24 000 DNA barcode sequences of 12 000 ex-type and manually validated filamentous fungal strains of 7 300 accepted species, the optimal identity thresholds to discriminate filamentous fungal species were predicted as 99.6 % for ITS and 99.8 % for LSU. We showed that 17 % and 18 % of the species could not be discriminated by the ITS and LSU genetic markers, respectively. Among them, ∼8 % were indistinguishable using both genetic markers. ITS has been shown to outperform LSU in filamentous fungal species discrimination with a probability of correct identification of 82 % vs. 77.6 %, and a clustering quality value of 84 % vs. 77.7 %. At higher taxonomic classifications, LSU has been shown to have a better discriminatory power than ITS. With a clustering quality value of 80 %, LSU outperformed ITS in identifying filamentous fungi at the ordinal level. At the generic level, the clustering quality values produced by both genetic markers were low, indicating the necessity for taxonomic revisions at genus level and, likely, for applying more conserved genetic markers or even whole genomes. The taxonomic thresholds predicted for filamentous fungal identification at the genus, family, order and class levels were 94.3 %, 88.5 %, 81.2 % and 80.9 % based on ITS barcodes, and 98.2 %, 96.2 %, 94.7 % and 92.7 % based on LSU barcodes. The DNA barcodes used in this study have been deposited to GenBank and will also be publicly available at the Westerdijk Institute's website as reference sequences for fungal identification, marking an unprecedented data release event in global fungal barcoding efforts to date.

Four species of polyporoid fungi newly recorded from Taiwan

Article

Full-text available

Apr 2018

Four wood-rotting polypores are reported from Taiwan for the first time: Ceriporiopsis pseudogilvescens, Megasporia major, Phlebiopsis castanea, and Trametes maxima. ITS (internal transcribed spacer) sequences were obtained from each specimen to confirm the determinations.

Additions to neotropical stereoid fungi (Polyporales, Basidiomycota): one new species of Lopharia and one new combination in Phlebiopsis

Article

Jan 2020

Stereoid fungi are an artificial group with mostly effused-reflexed to stipitate basidiomata, smooth hymenophore, and hyaline spores. From recent surveys in the Brazilian Atlantic Forest, Caatinga, and Cerrado, six specimens of this group had their identity tested with the nrITS and nrLSU sequences. Two of them were nested within the Lopharia s.s. clade and represent a new species L. erubescens, characterized by the dextrinoid reaction of the cystidia, and small basidia and spores. The other four were initially identified as Hjortstamia amethystea, but nested in the Phlebiopsis clade. Thus, we proposed the new combination, Phlebiopsis amethystea. We also provide keys to neotropical Lopharia s.l. and Phlebiopsis s.l. and allied species.

Crepatura ellipsospora gen. et sp. nov. in Phanerochaetaceae (Polyporales, Basidiomycota) bearing a tuberculate hymenial surface

Article

Jun 2019

A new wood-inhabiting fungal genus, Crepatura, typified by C. ellipsospora sp. nov., is proposed based on a combination of morphological features and molecular evidence. The genus is characterized by an annual growth habit, resupinate basidiocarps with smooth to irregularly tuberculate hymenial surface, a monomitic hyphal system with thick-walled generative hyphae bearing both clamp connections and simple septa and ellipsoid, hyaline, thick-walled, smooth, negative in Melzer’s, acyanophilous basidiospores measuring 6.5–7.5 × 4–5 μm. Sequences of ITS and LSU nrRNA gene regions of the studied samples were generated, and phylogenetic analyses were performed with maximum likelihood, maximum parsimony, and Bayesian inference methods. The phylogenetic analyses based on molecular data of ITS+nLSU sequences showed that Crepatura belonged to the Phanerochaetaceae family and nested into the Donkia clade. The result demonstrated that the genus Crepatura formed a monophyletic lineage with a strong support and phylogenetically was closely related to Pirex concentricus.

Phlebiopsis yunnanensis sp. Nov. (Polyporales, Basidiomycota) evidenced by morphological characters and phylogenetic analysis

Article

Jan 2018
NOVA HEDWIGIA

A new wood-inhabiting fungal species, Phlebiopsis yunnanensis sp. nov., is proposed based on morphological and molecular characters. The species is characterized by resupinate, membranaceous to subceraceous basidiocarps, a monomitic hyphal system with simple separated generative hyphae, cystidia conical, thick-walled, heavily encrusted with large crystals in the apical part and ellipsoid basidiospores measuring 3.5–4.5 × 2.5–3.5 μm. The internal transcribed spacer (ITS) and the large subunit (LSU) regions of nuclear ribosomal RNA gene sequences of the studied samples were generated, and phylogenetic analyses were performed with maximum likelihood, maximum parsimony and bayesian inference methods. The phylogenetic analyses based on molecular data of ITS+nLSU sequences showed that P. yunnanensis belonged to the Phanerochaetaceae and nested into the phlebioid clade. Further investigation was obtained for more representative taxa in the Phlebiopsis based on ITS+nLSU sequences, in which the result demonstrated that the species P. yunnanensis formed a monophyletic lineage with a strong support (100% BS, 100% BP, 1.00 BPP) and then grouped with P. gigantea and P. lamprocystidiata. © 2018 J. Cramer in Gebrüder Borntraeger Verlagsbuchhandlung, Stuttgart, Germany.

Lopharia

Article

May 1975

A. L. Welden

Basidiocarps of members of the Cinerascens groups of Lopharia have a well-developed cuticle and have simple clamps on their generative hyphae. Lopharia cinerascens has a cosmopolitan distribution and L. sharpiana is known only from Mexico. Basidiocarps of members of the Papyrina group lack a cuticle and do not ordinarily produce clamps on their generative hyphae (double or multiple clamps are rarely produced at some septa). Lopharia crassa, L. mexicana, L. nova-granata, L. papyracea, L. papyrina, and L. rimosissima are in the Papyrina group. All are distributed within the New World except L. papyracea, which appears to to be confined to the Old World tropics. Basidiocarps of members of the Spadicea group lack a cuticle but regularly produce clamps on their generative hyphae. All are distributed within the Old World. Lopharia cystidiosa, L. fulva, L. phellodendri, L. spadicea, and L. umbrino-alutacea are in the Spadicea group. Lopharia appears to be closely related to at least some species of Phanerochaete, to Chaetoporus, Flavodon, Irpex, and Steccherinum.

The Resupinate Aphyllophorales of the North Western Himalayas

Article

Nov 1978

Two new species of Phanerochaete (Basidiomycota) and redescription of P. robusta

Article

Dec 2017

In the current study, the new species Phanerochaete bambucicola and P. incarnata, from subtropical Taiwan are presented, while P. robusta is redescribed. We amplified the 5.8 s and LSU ribosomal genes along with RPB1. We generated an ITS dataset and a concatenated 5.8 s + nLSU + RPB1 dataset, which we analyzed using Maximum Parsimony and Bayesian methods. Our results suggest that the two new species belong in Phanerochaete s.s. P. bambucicola was collected on dead bamboo culm, characterized by the loose subiculum, the presence of leptocystidia, and the short lateral branches occasionally seen on the subicular hyphae. P. incarnata is characterized by the pink hymenial surface, and the presence of leptocystidia, which are usually covered with yellowish resinous material. The presence of enclosed, irregularly swollen, leptocystidia with oil drops is reported for P. robusta.

A Contribution to the Phylogeny and Taxonomy of Phlebiopsis (Polyporales, Basidiomycota)

Abstract

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