Updated Taxonomy of Chinese Phylloporus (Boletaceae, Boletales) Six New Taxa and Four Redescribed Species

Mycological Progress (2021) 20:1243–1273
https://doi.org/10.1007/s11557-021-01722-8
ORIGINAL ARTICLE
Updated taxonomy of Chinese Phylloporus (Boletaceae, Boletales):

six new taxa and four redescribed species
Lu-Ling Wu 1 & Zhi-Qun Liang 2 & Ming-Sheng Su 3 & Yu-Guang Fan 1 & Ping Zhang 4 & Shuai Jiang 5 & Yan-Liu Chen 6 &
Yan-Jia Hao 7 & Nian-Kai Zeng 1
Received: 23 February 2021 / Revised: 18 June 2021 / Accepted: 21 June 2021

# German Mycological Society and Springer-Verlag GmbH Germany, part of Springer Nature 2021
Abstract
Species of Phylloporus (Boletaceae, Boletales) in China were investigated using morphology and molecular phylogenetic
analyses of DNA sequences from the nuclear 28S rDNA D1–D2 domains, rDNA internal transcribed spacer region, and
translation elongation factor 1-α gene. Ten species are recognized from China in the present study. Among them,
P. alboinfuscatus, P. grossus, P. microsquamus, P. nigrisquamus, P. nigrobrunneus, and P. tenuissimus are new to science;
and P. bogoriensis, P. castanopsidis, P. luxiensis, and P. subrubeolus are previously described. Phylloporus megacystidiatus,
originally described from southwestern China, is likely conspecific with P. yunnanensis. A key to the Chinese species of
Phylloporus is provided. A preliminary biogeographical analysis showed that the affinities of Phylloporus species between
subtropical/tropical China and Thailand were evident, both regions share seven common taxa; P. nigrisquamus and
P. microsquamus are closely related to taxa from North/Central America.
Keywords Bolete . Molecular phylogeny . Morphology . New taxa . Taxonomy
Introduction Halling 2010; Zeng et al. 2011, 2013; Neves et al. 2012;
Hosen and Li 2015, 2017; Pradeep et al. 2015; Zhao et al.
Phylloporus Quél. (Boletaceae, Boletales), typified by 2018; Chuankid et al. 2019; Montoya et al. 2019; Zhang
P. pelletieri (Lév.) Quél., is one genus in Boletaceae with et al. 2019), showing high species diversity. The edibility of
lamellate hymenophore (Corner 1970, 1974; Neves and the genus has also been noted (Dai et al. 2010). For example,
Halling 2010; Neves et al. 2012; Zeng et al. 2013). Modern collections of P. luxiensis M. Zang are sold as edibles in mar-
phylogenetic analyses confirm it is a monophyletic genus kets in Yunnan Province, southwestern China (Fig. 1).
(Neves et al. 2012; Zeng et al. 2013). The genus In China, species of Phylloporus that are new to China or
Phylloporus is widely distributed worldwide with approxi- new to science have been recently reported (Zeng et al. 2013;
mately 100 taxa uncovered (Corner 1970, 1974; Neves and Zhao et al. 2018; Chuankid et al. 2019; Zhang et al. 2019).
Section Editor: Zhu-Liang Yang
* Zhi-Qun Liang 4
College of Life Science, Hunan Normal University,
lizhqu1980@126.com Changsha 410081, China
* Nian-Kai Zeng
5
niankaiz@163.com Yinggeling Branch of Hainan Tropical Rainforest National Park,
Baisha, Hainan 572800, China
1
Key Laboratory of Tropical Translational Medicine of Ministry of
Education, Transgenic Laboratory, School of Pharmacy, Hainan 6
Key Laboratory of State Forestry Administration on Tropical
Medical University, Haikou 571199, China Forestry Research, Research Institute of Tropical Forestry, Chinese
2
College of Science, Hainan University, Haikou 570228, China Academy of Forestry, Guangzhou 510520, China
3
Key Laboratory of Modern Preparation of TCM of Ministry of 7
Education, Jiangxi University of Traditional Chinese Medicine, School of Horticulture, Anhui Agricultural University,
Nanchang 330004, China Hefei 230036, China
1244 Mycol Progress (2021) 20:1243–1273
Fig. 1 Collections of Phylloporus luxiensis sold as edibles in the market of Yunnan Province, southwestern China
However, many forests in the country still have not been ex- Molecular procedures
tensively examined for this genus. In recent years, many col-
lections of Phylloporus from China have been made and are Total genomic DNA was obtained from materials
available for molecular phylogenetic analyses and morpho- dried with silica gel with the Plant Genomic DNA
logical studies. In the present study, we described six new Kit (TIANGEN Biotech Co., China) according to the
species and four previous taxa of Phylloporus from China manufacturer’s instructions. Primer pairs used for am-
and updated the taxonomy of the genus. plifying the nuclear 28S rDNA D1–D2 domains
(28S), rDNA internal transcribed spacer (ITS) region
(ITS1-5.8S-ITS2), and translation elongation factor
1-α gene (TEF1) were LROR/LR5 (Vilgalys and
Materials and methods Hester 1990; James et al. 2006), ITS5/ITS4 (White
et al. 1990), and EF1-2F/EF1-2R (Zeng et al. 2013),
Morphological studies respectively. Polymerase chain reaction (PCR) was
performed in a total volume of 25 μL containing 13
Specimens were collected and photographed in the field and μ L of G o Ta q ® G r e e n M a s t e r M i x (T I A N G EN
deposited in the Fungal Herbarium of Hainan Medical Biotech Co., China), 2 μL of each primer (10 μM),
University (FHMU), Haikou City, Hainan Province, China, 2 μL of DNA template, and 8 μL of nuclease-free
or the Herbarium of Cryptogams, Kunming Institute of water. PCRs were performed with an initial denatur-
Botany, Chinese Academy of Sciences (KUN-HKAS). ation at 95 °C for 5 min, followed by 35 cycles of
Color codes from Kornerup and Wanscher (1981) were used. denaturation at 94 °C for 30 s, annealing at appropri-
For observations of the pileipellis, a radial-vertical section was ate temperature (50 °C for 28S and ITS, 53 °C for
made approximately halfway between the margin and the cen- TEF1) for 30 s, extension at 72 °C for 30 s and a
ter of the pileus. Sections of the squamules were taken from final extension at 72 °C for 7 min. PCR products
the middle part along the longitudinal axis of the stipe. were checked in 1% (w/v) agarose gels, and positive
Microscopic structures were rehydrated in H2O or 5% KOH reactions with a bright, single band were purified and
and drawn freehand. Basidiospores of dried specimens were directly sequenced at BGI (Guangzhou, China) using
also examined with a Quanta 250 (FEI, USA) scanning elec- a 3730xl DNA Analyzer (Applied Biosystems, USA)
tron microscope (SEM) (Zeng et al. 2013). The number of with the same primers used for PCR amplifications.
measured basidiospores was given as n/m/p, where n, m, and Forward or reverse sequences were compiled with
p represented the total number of basidiospores, basidiomata, BioEdit (Hall 1999). Edited sequences were deposited
and collections measured, respectively. Dimensions of basid- in GenBank (Table 1).
iospores were given as (a–) b–c (–d), where the range b–c
represented a minimum of 90% of the measured values (5th Dataset assembly
to 95th percentiles) and extreme values (a and d), whenever
present (a < 5th percentile, d > 95th percentile), were given in One hundred and six sequences (47 of 28S, 34 of ITS, 25 of
parentheses. Q referred to the length/width ratio of basidio- TEF1) from 56 collections of Phylloporus were generated
spores; Qm referred to the average Q of basidiospores and was (Table 1). For the concatenated dataset, 28S, ITS, and TEF1
given with a sample standard deviation. sequences were aligned with selected sequences from
Mycol Progress (2021) 20:1243–1273 1245
Table 1 Taxa, vouchers, locations, and GenBank accession numbers of the DNA sequences used in this study
Accepted names Names from previous Voucher Locality GenBank accession No. References
studies/GenBank
28S ITS TEF1
Hourangia cheoi Hourangia cheoi Zhu130 Yunnan, SW KP136950 KP137001 KP136929 Zhu et al. (2015)
(KUN-HKAS China
68306)
Hourangia cheoi Hourangia cheoi Wu203 Yunnan, SW KP136948 KP136995 KP136939 Zhu et al. (2015)
(KUN-HKAS China
59790)
Phylloporus — Y.L. Chen DX010 Guangdong, MW588630 MW588666 — Present study
alboinfuscatus (FHMU5883) southern
China
Phylloporus — N.K. Zeng4179 Hunan, central MW588626 MW588663 — Present study
alboinfuscatus (FHMU3276) China
Phylloporus Phylloporus bellus JXSB1620 Jiangxi, eastern MK765818 — — Unpublished
alboinfuscatus China
Phylloporus Phylloporus MAN022 Costa Rica JQ003678 JQ003624 — Neves et al. (2012)
alborufus alborufus
Phylloporus Phylloporus JT27954 USA JQ003704 — — Neves et al. (2012)
arenicola arenicola
Phylloporus Phylloporus KUN-HKAS Bangladesh KR094781 KR094777 KR094791 Hosen and Li (2017)
attenuatus attenuatus 76168
attenuatus attenuatus 76167
Phylloporus bellus Phylloporus aff. OR0473 China — — MH58078 Chuankid et al.
bellus (2019)
Phylloporus bellus Phylloporus bellus KUN-HKAS Yunnan, SW JQ967196 JQ967239 JQ967153 Zeng et al. (2013)
56763 China
Phylloporus bellus Phylloporus bellus KUN-HKAS Yunnan, SW JQ967197 JQ967240 JQ967154 Zeng et al. (2013)
42850 China
Phylloporus bellus Phylloporus bellus MCA559 Japan AY612817 — — Unpublished
Phylloporus bellus — Y.G Fan2750 Yunnan, SW MW588589 MW588655 — Present study
(FHMU3285) China
Phylloporus sp. Phylloporus bellus REH7733 Costa Rica Q003661 — — Neves et al. (2012)
Phylloporus sp. Phylloporus bellus REH8710 Eastern USA JQ003686 JQ003618 — Neves et al. (2012)
Phylloporus Phylloporus DED7785 Indonesia JQ003680 JQ003625 — Neves et al. (2012)
bogoriensis bogoriensis
Phylloporus — N.K. Zeng761 Hainan, MW588592 — — Present study
bogoriensis (FHMU459) southern
China
Phylloporus Phylloporus KUN-HKAS Yunnan, SW JQ967198 JQ967241 JQ967155 Zeng et al. (2013)
brunneiceps brunneiceps 56903 China
Phylloporus Phylloporus KUN-HKAS Chongqing, SW JQ967200 JQ967243 JQ967157 Zeng et al. (2013)
Phylloporus Phylloporus KUN-HKAS Chongqing, SW JQ967201 JQ967244 JQ967158 Zeng et al. (2013)
Phylloporus — Y.J. Hao1427 Anhui, eastern MW588582 — MW574465 Present study
brunneiceps (FHMU3292) China
Phylloporus Phylloporus JXSB1264 Jiangxi, eastern MK765819 — — Unpublished
brunneiceps brunneiceps China
Phylloporus Phylloporus OR050 Thailand — — KT824034 Raspé et al. (2016)
brunneiceps brunneiceps
Phylloporus Phylloporus MAN104 Thailand JQ003689 JQ003642 — Neves et al. (2012)
castanopsidis castanopsidis
castanopsidis castanopsidis
Phylloporus Phylloporus N.K. Zeng2278 Hainan, MT829130 MT822942 — Zeng and Jiang
castanopsidis castanopsidis (FHMU1517) southern (2020)
China
1246 Mycol Progress (2021) 20:1243–1273
Table 1 (continued)
studies/GenBank
28S ITS TEF1
Phylloporus Phylloporus REH7906 Panama JQ003662 JQ003638 — Neves et al. (2012)

caballeroi caballeroi
catenulatus catenulatus 76157
catenulatus catenulatus 76156
centroamericanus centroamericanus
centroamericanus centroamericanus
Phylloporus Phylloporus REH8681 Australia JQ003684 JQ003621 — Neves et al. (2012)
cyanescens cyanescens
dimorphus dimorphus
Phylloporus Phylloporus JLM1677 Eastern USA JQ003687 JQ003641 — Neves et al. (2012)
foliiporus foliiporus
Phylloporus gajari Phylloporus gajari KUN-HKAS Bangladesh KP780423 KP780419 — Hosen and Li (2015)
81585
Phylloporus gajari Phylloporus gajari KUN-HKAS Bangladesh KP780422 KP780418 — Hosen and Li 2015
76166
Phylloporus gajari Phylloporus gajari KUN-HKAS Bangladesh KP780421 KP780417 — Hosen and Li (2015)
76161
Phylloporus grossus — N.K. Zeng4159 Hunan, central MW588625 MW588662 — Present study
(FHMU3277) China
Phylloporus grossus — N.K. Zeng239 Hainan, MW588583 — — Present study
(FHMU2469) southern
China
Phylloporus grossus — N.K. Zeng4158 Hunan, central MW588624 — — Present study
(FHMU3280) China
Phylloporus grossus — N.K. Zeng3309 Hainan, MW588586 MW588642 MW574462 Present study
(FHMU2270) southern
China
(FHMU3136) southern
China
(FHMU2937) southern
China
imbricatus imbricatus 54647 China
Phylloporus Phylloporus KUN-HKAS Yunnan, SW KF112398 — KF112299 Wu et al. (2014)
Phylloporus Phylloporus MAN123 Thailand JQ003695 — — Neves et al. (2012)
infuscatus infuscatus
Phylloporus Phylloporus MB00-043 Eastern USA JQ003677 JQ003628 — Neves et al. (2012)
leucomycelinus leucomycelinus
Phylloporus Phylloporus KUN-HKAS Eastern USA JQ967206 JQ967249 JQ967163 Zeng et al. (2013)
leucomycelinus leucomycelinus 74678
Phylloporus — N.K. Zeng3439 Hainan, — — MW574467 Present study
luxiensis (FHMU3095) southern
China
Phylloporus — N.K. Zeng3442 Hainan, MW588605 — MW574469 Present study
China
Mycol Progress (2021) 20:1243–1273 1247
Table 1 (continued)
studies/GenBank
28S ITS TEF1
Phylloporus — N.K. Zeng3642 Yunnan, SW MW588609 — MW574466 Present study

luxiensis (FHMU2996) China
Phylloporus — N.K. Zeng2784 Yunnan, SW MW588608 — — Present study
Phylloporus — N.K. Zeng4196 Hunan, Central MW588628 — — Present study
Phylloporus — N.K. Zeng1364 Fujian, SE MW588607 MW588643 — Present study
Phylloporus — N.K. Zeng3444 Hainan, MW588611 MW588644 MW574468 Present study
China
Phylloporus luxiensis Phylloporus luxiensis KUN-HKAS Yunnan, SW JQ967207 JQ967250 JQ967164 Zeng et al. (2013)
57036 China
Phylloporus luxiensis Phylloporus luxiensis KUN-HKAS Yunnan, SW JQ967209 JQ967252 JQ967166 Zeng et al. (2013)
57048 China
Phylloporus luxiensis Phylloporus luxiensis KUN-HKAS Yunnan, SW KF112490 — KF112298 Wu et al. (2014)
75077 China
Phylloporus luxiensis Phylloporus sp. 2 KUN-HKAS Fujian, SE JQ967229 JQ967272 JQ967185 Zeng et al. (2013)
74680 China
Phylloporus luxiensis Phylloporus sp. 5 KUN-HKAS Fujian, SE JQ967232 JQ967275 JQ967188 Zeng et al. (2013)
74684 China
maculatus maculatus 56683 China
maculatus maculatus 59730 China
Phylloporus Phylloporus OR0285 China — — MH580800 Chuankid et al.
maculatus maculatus (2019)
Phylloporus — Y.G. Fan2752 Yunnan, SW MW588618 MW588656 — Present study
maculatus (FHMU3287) China
Phylloporus Phylloporus sp. OR0258 China — — MH580809 Chuankid et al.
microsquamus 2019
Phylloporus — N.K. Zeng2571 Hainan, MW588599 MW588648 — Present study
microsquamus (FHMU1678) southern
China
Phylloporus — N.K. Zeng189 Hainan, MW588594 — — Present study
China
China
China
China
Phylloporus — N.K. Zeng3560 Hainan MW588614 MW588654 MW574474 Present study
China
China
China
1248 Mycol Progress (2021) 20:1243–1273
Table 1 (continued)
studies/GenBank
28S ITS TEF1

China
Phylloporus — Y.G. Fan2819 Yunnan, SW MW588590 — — Present study
nigrisquamus (FHMU3271) China
Phylloporus — Y.G. Fan2834 Yunnan, SW MW588591 — — Present study
nigrisquamus (FHMU3272) China
Phylloporus Phylloporus sp. KUN-HKAS Yunnan, SW KF112396 — — Present study
nigrisquamus 63001 China
Phylloporus Phylloporus sp. MAN131 Thailand JQ003698 JQ003649 — Neves et al. (2012)
nigrisquamus
Phylloporus Phylloporus sp. 4 KUN-HKAS Yunnan, SW JQ967231 JQ967274 JQ967187 Zeng et al. (2013)
nigrobrunneus 74683 China
nigrobrunneus 74682 China
Phylloporus Phylloporus REH4582 Eastern USA JQ003679 — — Neves et al. (2012)
nigrisquamus leucomycelinus
Phylloporus Phylloporus OR0448 China — — MG212598 Vadthanarat et al.
nigrobrunneus yunnanensis (2018)
nigrobrunneus (FHMU3269) China
orientalis orientalis
orientalis orientalis
pachycystidiatus pachycystidiatus 54540 China
pachycystidiatus pachycystidiatus 54541 China
Phylloporus — N.K. Zeng3622 Yunnan, SW MW588616 MW588658 MW574459 Present study
pachycystidiatus (FHMU2834) China
Phylloporus Phylloporus KUN-HKAS Yunnan SW JQ967214 JQ967257 JQ967171 Zeng et al. (2013)
parvisporus parvisporus 54768 China
Phylloporus Phylloporus KUN-HKAS Fujian, SE JQ967213 JQ967256 JQ967170 Zeng et al. (2013)
parvisporus parvisporus 59725 China
Phylloporus pelletieri Phylloporus pelletieri K 128205 England JQ967215 JQ967258 — Zeng et al. (2013)
Phylloporus pelletieri Phylloporus pelletieri Q7199c Slovakia JQ003668 JQ003639 — Neves et al. (2012)
Phylloporus Phylloporus MAN064 Costa Rica JQ003670 — — Neves et al. (2012)
phaeoxanthus phaeoxanthus
Phylloporus Phylloporus REH7388 Costa Rica JQ003671 — — Neves et al. (2012)
phaeoxanthus var. phaeoxanthus var.
simplex simplex
pruinatus pruinatus 74687 China
Phylloporus Phylloporus KUN-HKAS Anhui, eastern MH624125 MH624124 MH624126 Zhao et al. (2018)
pruinatus pruinatus 101929 China
purpurellus purpurellus
Phylloporus pusillus Phylloporus pusillus OR0484 Yunnan, SW — — MH580802 Chuankid et al.
China (2019)
Mycol Progress (2021) 20:1243–1273 1249
Table 1 (continued)
studies/GenBank
28S ITS TEF1
Phylloporus pusillus Phylloporus pusillus OR1158 Thailand — — MH580803 Chuankid et al.

(2019)
Phylloporus pusillus Phylloporus pusillus OR1310 Thailand — — MH580804 Chuankid et al.
(2019)
Phylloporus Phylloporus Montoya_5239 Mexico MK226558 MK226550 — Montoya et al.
quercophilus quercophilus (2019)
Phylloporus Phylloporus MAN075 Eastern USA JQ003674 — — Neves et al. (2012)
rhodoxanthus rhodoxanthus
Phylloporus Phylloporus REH8714 Eastern USA JQ003675 JQ003629 — Neves et al. (2012)
Phylloporus Phylloporus MAN099 Eastern USA JQ003676 — — Neves et al. (2012)
Phylloporus Phylloporus JLM1808 Eastern USA JQ003688 JQ003654 — Neves et al. (2012)
Phylloporus rimosus Phylloporus rimosus Montoya_4834 Mexico MK226554 MK226546 MK314102 Montoya et al.
(2019)
Phylloporus rubeolus Phylloporus rubeolus KUN-HKAS Yunnan, SW JQ967216 JQ967259 JQ967172 Zeng et al. (2013)
52573 China
Phylloporus rubeolus Phylloporus rubeolus KUN-HKAS Yunnan, SW JQ967218 JQ967261 JQ967174 Zeng et al. (2013)
54543 China
Phylloporus rubeolus Phylloporus rubeolus OR0251 China — — MH580806 Chuankid et al.
(2019)
Phylloporus rubeolus Phylloporus rubeolus N.K. Zeng2459 Hainan, MT829131 MT822943 — Zeng and Jiang
(FHMU1585) southern (2020)
China
rubeolus (FHMU3301) China
rubiginosus rubiginosus
rubiginosus rubiginosus
Phylloporus Phylloporus BC063 Thailand — — MH580807 Chuankid et al.
rubiginosus rubiginosus (2019)
Phylloporus Phylloporus OR0169 China — — MH580808 Chuankid et al.
rubiginosus rubiginosus (2019)
Phylloporus Phylloporus N.K. Zeng1267 Fujian, SE MK116510 MK116513 MK116514 Zhang et al. (2019)
rubiginosus rubiginosus (FHMU824) China
Phylloporus Phylloporus N.K. Zeng589 Fujian, SE MK116509 MK116512 — Zhang et al. (2019)
rubiginosus rubiginosus (FHMU2482) China
Phylloporus Phylloporus N.K. Zeng3175 Hainan, MK116511 — — Zhang et al. (2019)
rubiginosus rubiginosus (FHMU2136) southern
China
Phylloporus Phylloporus N.K. Zeng3055 Hainan, MT829149 MT822964 MW574454 Zeng and Jiang
rubiginosus rubiginosus (FHMU2016) southern (2020); present
China study
rubrosquamosus rubrosquamosus 54542 China
Phylloporus Phylloporus KUN-HKAS Yunnan, SW KF112391 — KF112289 Wu et al. (2014)
Phylloporus Phylloporus KUN-HKAS Hainan, JQ967220 JQ967263 JQ967176 Zeng et al. (2013)
rufescens rufescens 59722 southern
China
Phylloporus Phylloporus KUN-HKAS Hainan, JQ967221 JQ967264 JQ967177 Zeng et al. (2013)
rufescens rufescens 59723 southern
China
1250 Mycol Progress (2021) 20:1243–1273
Table 1 (continued)
studies/GenBank
28S ITS TEF1
Phylloporus — N.K. Zeng1904 Hainan, — MW588645 MW574453 Present study

rufescens (FHMU1227) southern
China
Phylloporus sp. — N.K. Zeng3445 Hainan, — MW588634 MW574461 Present study
(FHMU3283) southern
China
Phylloporus sp. — N.K. Zeng4088 Guangdong, MW588621 — — Present study
(FHMU3363) southern
China
Phylloporus sp. — N.K. Zeng2724 Hainan, MW588581 MW588632 — Present study
(FHMU1718) southern
China
Phylloporus sp. — N.K. Zeng3558 Yunnan, SW — — MW574456 Present study
(FHMU3362) China
Phylloporus sp. — N.K. Zeng4228 Hainan, MW588629 MW588665 — Present study
(FHMU3282) southern
China
Phylloporus sp. Phylloporus JXSB1345 Jiangxi, eastern MK765820 — — Unpublished
brunneiceps China
Phylloporus sp. Phylloporus OR052 Thailand — — KT824035 Raspé et al. (2016)
castanopsidis
Phylloporus sp. Phylloporus BC074 Thailand — — MH580799 Chuankid et al.
castanopsidis (2019)
Phylloporus sp. Phylloporus WU17978 Austria — — MH580805 Chuankid et al.
rhodoxanthus (2019)
Phylloporus sp. Phylloporus SAR 89.457 Eastern USA U11925 — — Chapela et al.
rhodoxanthus (1994)
Phylloporus sp. Phylloporus sp. REH8729 Australia JQ003699 JQ003650 — Neves et al. (2012)
Phylloporus sp. Phylloporus sp. MAN105 Thailand JQ003690 — — Neves et al. (2012)
Phylloporus sp. Phylloporus sp. 1 KUN-HKAS Hunan, central JQ967228 JQ967271 JQ967184 Zeng et al. (2013)
74679 China
Phylloporus sp. Phylloporus sp. 3 KUN-HKAS Hainan, JQ967227 JQ967270 JQ967183 Zeng et al. (2013)
74681 southern
China
Phylloporus sp. Phylloporus sp. 8 KUN-HKAS Shandong, JQ967234 JQ967277 — Zeng et al. (2013)
74686 eastern China
Phylloporus sp. Phylloporus sp. 9 KUN-HKAS Yunnan, SW JQ967233 JQ967276 JQ967189 Zeng et al. (2013)
74685 China
Phylloporus Phylloporus sp. OR0989 Thailand — MH686277 MH580830 Chuankid et al.
subbacillisporus (2019)
Phylloporus — Y.J. Hao1553 Anhui, eastern MW588601 MW588635 MW574460 Present study
subbacillisporus (FHMU3293) China
Phylloporus Phylloporus BC022 Thailand — — MH580813 Chuankid et al.
subrubeolus subrubeolus (2019)
Phylloporus Phylloporus OR0612 Thailand — — MH580814 Chuankid et al.
subrubeolus subrubeolus (2019)
Phylloporus — N.K. Zeng4027 Hainan, MW588620 — MW574475 Present study
subrubeolus (FHMU3275) southern
China
Phylloporus — N.K. Zeng4195 Hunan, central MW588627 MW588664 MW574476 Present study
subrubeolus (FHMU3273) China
Phylloporus — N.K. Zeng4089 Guangdong, MW588622 — — Present study
China
Phylloporus — N.K. Zeng4095 Guangdong, MW588623 MW588661 — Present study
China
Mycol Progress (2021) 20:1243–1273 1251
Table 1 (continued)
studies/GenBank
28S ITS TEF1
subrubeolus 74688 China
Phylloporus Phylloporus sp.10 KUN-HKAS Hainan, JQ967237 JQ967280 JQ967192 Zeng et al. (2013)
tenuissimus 74689 southern
China
tenuissimus (FHMU2964) southern
China
yunnanensis yunnanensis 52225 China
Note. New sequences are shown in bold. SE, southeast; SW, southwest
GenBank and previous studies (Table 1). Hourangia cheoi posterior probabilities (PPs) were then calculated for a majority
(W.F. Chiu) Xue T. Zhu & Zhu L. Yang was chosen as the consensus tree of the retained Bayesian trees.
outgroup as inferred by Zhao et al. (2018). Sequences were
aligned using MUSCLE v3.6 (Edgar 2004) and manually ad-
justed where necessary. In order to test for phylogenetic con-
flict among the different genes in the concatenated dataset, Results
single-gene phylogenies based on 28S, ITS, or TEF1 were
analyzed. The results of these analyses indicated that the dif- Molecular data
ferent genes were not in conflict. Therefore, the sequences of
all three genes were concatenated using Phyutility v2.2 (Smith The concatenated dataset (28S + ITS + TEF1) included 172
and Dunn 2008) for further analyses. taxa of Boletaceae with 2021 nucleotide sites, and the align-
ment was deposited in TreeBASE (S27775). A phylogram
with branch lengths inferred with RAxML, including the sup-
Phylogenetic analyses port values, is shown (Fig. 2). Based on the concatenated
dataset, the topologies of phylogenetic trees generated from
The concatenated dataset (28S + ITS + TEF1) was analyzed with the ML and BI analyses were almost identical, while statistical
maximum likelihood (ML) and Bayesian inference (BI). ML tree support showed minor differences. The collections of
generation and bootstrap (BS) analyses were performed with Phylloporus in China were grouped into 33 independent line-
RAxML v7.2.6 (Stamatakis 2006) running 1000 replicates com- ages, and these taxa were labeled as lineages 1–33 (Fig. 2).
bined with an ML search. Bayesian analysis was performed with Related taxa that were not from China were also labeled for
MrBayes v3.1 (Huelsenbeck and Ronquist 2005) implementing the present study (Fig. 2).
the Markov chain Monte Carlo (MCMC) technique and param- In the present study, we paid particular attention to lineages
eters predetermined with MrModeltest v2.3 (Nylander 2004). 1, 7, 9, 11, 12, 16, 24, 29, 30, and 32 (Fig. 2). Lineage 1, with
For the three-gene concatenated dataset, GTR + I + G, K80 + I strong statistical support (BS = 98%, PP = 1.0), included three
+ G, and SYM + G were chosen as best-fit likelihood models for collections of P. luxiensis and four specimens (FHMU1771,
28S, ITS, and TEF1 partitions, respectively. Bayesian analysis FHMU1773, FHMU2935, and FHMU2996) from southwest-
was repeated for 30 million generations and sampled every 100 ern China, two specimens labeled as “P. sp. 2” and “P. sp. 5”
generations. Trees sampled from the first 25% of the generations and one collection (FHMU915) from southeastern China,
were abandoned as burn-in. The average standard deviation of t h r e e c o l l e c t i o n s ( F H M U 3 0 9 5 , F H M U 3 1 0 1 , an d
split frequencies was restricted below 0.01, and Bayesian FHMU3105) from southern China, one specimen
1252 Mycol Progress (2021) 20:1243–1273
Fig. 2 Phylogram inferred from a three-locus (rDNA 28S, ITS regions, and TEF1) dataset using RAxML. BS ≥ 70% and PP ≥ 0.95 are indicated above
or below the branches as RAxML BS/PP. SW, southwest; SE, southeast
(FHMU3281) from central China, and the holotype of Lineage 11, with BS of 99%, included the holotype of
P. dimorphus M.A. Neves & Halling from Thailand. P. castanopsidis M.A. Neves & Halling from Thailand and
Lineage 7, with BS of 71%, included one specimen one specimen (FHMU1517) from southern China. Lineage
(FHMU5883) from southern China, one collection 12, with high statistical support (BS = 100%, PP = 1.0), in-
(FHMU3276) from central China, and one sample identified cluded the holotype and one collection of P. subrubeolus
as P. bellus (Massee) Corner from eastern China. Lineage 9, Chuankid, K.D. Hyde & Raspé from Thailand, one collection
with high statistical support (BS = 84%, PP = 0.99), included (FHMU3273) from central China, one specimen labeled as
two specimens (FHMU3277 and FHMU3280) from central “P. sp. 7” from southwestern China, and three specimens
China and four collections (FHMU2270, FHMU2469, (FHMU3274, FHMU3275, and FHMU3278) from southern
FHMU2937, and FHMU3136) from southern China. China. Lineage 16, with high statistical support (BS = 90%,
Mycol Progress (2021) 20:1243–1273 1253
Fig. 2 continued.
PP = 1.0), included two samples (FHMU3271 and FHMU1640, FHMU1678, FHMU2467, FHMU3012,
FHMNU3272) from southwestern China, one collection iden- FHMU3018, FHMU3064, and FHMU3300) from southern
tified as P. leucomycelinus Singer from the eastern USA, one China. Lineage 32, with high statistical support (BS = 99%,
specimen labeled as “P. sp.” from Thailand, and one specimen PP = 1.0), included one sample of P. bogoriensis Höhn. from
labeled as “P. sp.” from southwestern China. Lineage 24, with Indonesia and one specimen (FHMU459) from southern
high statistical support (BS = 85%, PP = 1.0), included three China.
specimens (FHMU3268, FHMU3269, and FHMU3270), two
samples labeled as “P. sp. 4” and one specimen identified as
P. yunnanenis, all from China. Lineage 29, with high statisti- Taxonomy
cal support (BS = 100%, PP = 0.99), included one collection
labeled as “P. sp. 10” and one specimen (FHMU2964) from Phylloporus alboinfuscatus N.K. Zeng, L.L. Wu, Y.L. Chen
southern China. Lineage 30, with high statistical support (BS & P. Zhang, sp. nov.
= 100%, PP = 1.0), included one specimen labeled as “P. sp.” Figures 3a–d, 4a, and 5
from China and nine collections (FHMU1285, FHMU1416, MycoBank: MB838739
1254 Mycol Progress (2021) 20:1243–1273
Fig. 3 Basidiomata of
Phylloporus species. a–d
P. alboinfuscatus (a, d from
FHMU5883; b, c from
FHMU3276, holotype). e, f P.
bogoriensis (FHMU459). g, h
P. castanopsidis (FHMU1517).
i–k P. grossus (i, j from
FHMU3280; k from FHMU3277,
holotype). l–n P. luxiensis (l, n
from FHMU915; m from
FHMU3095). Photos: a, d Y.L.
Chen; b, c, e–n N.K. Zeng
Diagnosis: Phylloporus alboinfuscatus is characterized by lamellate, decurrent. Lamellae up to 0.7 cm broad, subdistant,
a pileus that is pale brown to brown when young and dirty commonly anastomosing, yellow (1A6), changing bluish
white when old, a stipe covered with reddish brown when injured; lamellulae common, attenuate, concolor with
squamules, thick-walled (1–2 μm) pleurocystidia and a lamellae. Stipe 2–3 × 0.5–0.7 cm, central, subcylindrical; sur-
pileipellis composed of more or less inflated hyphae. face dry, densely covered with minute, reddish brown
Etymology: Latin epithet “alboinfuscatus” refers to a pileus squamules; upper part usually ribbed by the decurrent lines
that is pale brown when young and dirty white when old. of the lamellae; context yellowish white (30A2), unchanging
Holotype: CHINA. Hunan Province: Chenzhou City, in color when injured; annulus absent. Basal mycelium white
Mangshan National Nature Reserve, elev. 1200 m, 29 (8A1). Odor not distinct.
July 2019, N.K. Zeng4179 (FHMU3276) Basidiospores [40/2/2] 9.5–13 × (4–) 4.5–5.5 μm, Q =
Basidiomata small to medium-sized. Pileus 3–7 cm in 1.73–2.40 (–3.00), Q m = 2.19 ± 0.04, subfusoid and
diam., convex to applanate; margin decurved when young, inequilateral in side view with slightly suprahilar depression,
then slightly uplifted; surface dry, tomentose, then squamu- subfusoid in ventral view, slightly thick-walled (up to 1 μm
lose, white, pale brown to brown; context approximately thick), yellowish brown in KOH, smooth under the light mi-
0.7 cm in thickness in the center of the pileus, yellowish white croscope but with bacillate ornamentation under SEM.
(30A2), unchanging in color when injured. Hymenophore Basidia 27–39 × 9–10 μm, clavate, thin-walled, 4-spored,
Mycol Progress (2021) 20:1243–1273 1255
Fig. 4 Basidiospores of
Phylloporus species from
herbarium materials under SEM.
a P. alboinfuscatus (FHMU3276,
holotype). b P. bogoriensis
(FHMU459). c P. castanopsidis
(FHMU1517). d P. grossus
(FHMU3277, holotype). e
P. luxiensis (FHMU3095). f
P. microsquamus (FHMU1678,
holotype). g P. nigrisquamus
(FHMU3271, holotype). h
P. nigrobrunneus (FHMU3268,
holotype). i P. subrubeolus
(FHMU3273). j P. tenuissimus
(FHMU2964, holotype). Note
basidiospores with bacillate
surface ornamentation. Bars = 3
μm
Fig. 5 Microscopic features of

Phylloporus alboinfuscatus
(FHMU3276, holotype). a
Basidia. b Basidiospores. c
Cheilocystidia. d Pleurocystidia.
e Pileipellis. f Stipitipellis. Scale
bars = 10 μm. Drawings by L.L.
Wu
1256 Mycol Progress (2021) 20:1243–1273
colorless to yellowish in KOH; sterigmata 4–7 μm in length. In our phylogenetic analyses, P. alboinfuscatus is closely
Hymenophoral trama phylloporoid, composed of thin- to related to P. brunneiceps N.K. Zeng, Zhu L. Yang & L.P.
slightly thick-walled (up to 0.5 μm) hyphae, 5–16 μm wide, Tang and one collection labeled as “P. sp.” (KUN-
colorless in KOH. Cheilocystidia 54–70 × 14–21 μm, abun- HKAS74681) (Fig. 2). However, P. brunneiceps has a stipe
dant, subfusiform or subclavate, slightly thick-walled (1 μm), without reddish tinge, yellowish basal mycelium, and thin- to
yellowish in KOH, no encrustations. Pleurocystidia 66–104 slightly thick-walled (up to 1 μm) pleurocystidia (Zeng et al.
× 13–24 μm, abundant, subfusiform or fusiform, thick-walled 2013). The morphological differences between
(1–2 μm), colorless to yellowish in KOH, no encrustations. P. alboinfuscatus and “P. sp.” (KUN-HKAS74681) are un-
Pileipellis a trichoderm approximately 230 μm, composed of clear because the morphological features of the latter were not
slightly interwoven, 6–15 μm in diam., thin-walled, colorless provided (Zeng et al. 2013).
to yellowish in KOH, occasionally branched hyphae; terminal
cells 20–60 × 6–15 μm, narrowly clavate or subcylindrical, Phylloporus bogoriensis Höhn., Sber. Akad. Wiss. Wien
with obtuse apex. Pileal trama made up of hyphae 5–19 μm (Math.-naturw. Kl., Abt. I) 123: 89, 1914 Figures 3e–f, 4b, and 6
in diam., thin-walled, colorless to yellowish in KOH. Basidiomata small to medium-sized. Pileus 3.5–6 cm in
Stipitipellis a hymeniform 60–70 μm thick, composed of col- diam., convex to applanate, slightly infundibuliform when
orless to yellowish in KOH, 3–16 μm wide, thin- to slightly old; surface dry, subtomentose, slightly rimose in age, pale
thick-walled (up to 1 μm) hyphae; terminal cells 20–40 × 8– brown, brown to dark brown; context 0.6–0.9 cm in thickness
15 μm, broadly or narrowly clavate, with suborbicular apex. in the center of the pileus, yellowish white, changing reddish
Stipe trama composed of longitudinally arranged, parallel quickly, then blackening when injured. Hymenophore lamel-
hyphae 4–18 μm wide, cylindrical, thin-walled, colorless to late, decurrent. Lamellae up to 0.5 cm broad, subdistant, com-
yellowish in KOH. Clamp connections absent in all tissues. monly anastomosing, yellow, changing greenish blue quickly,
Habitat: Solitary on the ground, in forests dominated by then reddish, finally blackening when injured; lamellulae
fagaceous trees common, attenuate, concolor with lamellae. Stipe 4.5–5 ×
Known distribution: Central and southern China 0.6–1 cm, central, subcylindrical; surface dry, densely cov-
Additional specimen examined: CHINA. Guangdong ered with reddish brown (9E5) squamules; context white
Province: Renhua County, Danxiashan National Nature (8A1), changing reddish (9E6) quickly, then blackening
Reserve, elev. 350 m, 5 June 2019, Y.L. Chen DX010 (12E1) when injured; annulus absent. Basal mycelium white
(FHMU5883) (8A1). Odor not distinct.
Comments: Phylloporus centroamericanus Singer & L.D. Basidiospores [40/1/2] (8–) 8.5–10.5 (–11.5) × 4–5 (–5.5)
Gómez, P. grossus N.K. Zeng, L.L. Wu & P. Zhang (see μm, Q = (1.64–) 1.80–2.50 (–2.75), Qm = 2.08 ± 0.24,
below), P. pachycystidiatus N.K. Zeng, Zhu L. Yang & L.P. subfusoid to elliptical and inequilateral in side view with
Tang, P. rubiginosus M.A. Neves & Halling and P. tunicatus slightly suprahilar depression, subfusoid in ventral view,
Corner also have thick-walled cystidia (Corner 1970; Singer slightly thick-walled (up to 0.5 μm), pale yellowish to yellow-
and Gómez 1984; Neves and Halling 2010; Neves et al. 2012; ish brown in KOH, smooth under the light microscope but
Zeng et al. 2013). However, P. centroamericanus has a small with bacillate ornamentation under SEM. Basidia 31–35 ×
basidioma, a stipe without reddish tinge, longer basidiospores 8–11 μm, clavate, thin-walled, 4-spored, yellowish in KOH;
(8–16 × 3.5–5.5 μm), longer cystidia (up to 138 μm), and a sterigmata 3–5 μm in length. Hymenophoral trama
distribution in Central America (Neves and Halling 2010); phylloporoid, composed of slightly thick-walled (up to 1
P. grossus has a hymenophore that is unchanging in color μm) hyph ae, 4–14 μm w i d e , c o l o r l e s s i n KO H .
when injured and shorter basidiospores (9–11 × 4–5.5 μm, Cheilocystidia 48–64 × 15–19 μm, subfusiform or fusiform,
see below); P. pachycystidiatus has a pileus tinged with red thin- to slightly thick-walled (up to 1 μm), colorless or golden
and longer pleurocystidia with thicker walls (up to 4 μm) yellow in KOH, no encrustations. Pleurocystidia 65–80 ×
(Zeng et al. 2013); P. rubiginosus has a brownish red to red- 10–19 μm, fusiform or subfusiform, thin- to slightly thick-
dish pileus, hymenophore and context that turn bluish oliva- walled (up to 1 μm), colorless, yellowish, or brownish yellow
ceous, then change red and finally blacken when injured, and in KOH, no encrustations. Pileipellis a trichoderm 180–210
smooth basidiospores (Neves et al. 2012; Zhang et al. 2019); μm thick, composed of hyphae more or less vertically ar-
P. tunicatus has a small basidioma, a context that turns blue ranged, thin- to slightly thick-walled (up to 1 μm), 8–15 μm
when injured and longer cystidia (Corner 1970). Moreover, wide, yellowish to golden yellow in KOH; terminal cells 30–
P. alboinfuscatus is genetically distant from 70 × 8–15 μm, narrowly clavate or subcylindrical, with obtuse
P. centroamericanus, P. grossus, P. pachycystidiatus, and apex. Pileal trama made up of hyphae 5–10 μm in diam.,
P. rubiginosus (Fig. 2). The phylogenetic relationship be- thin-walled, yellowish in KOH. Stipitipellis a hymeniform
tween P. alboinfuscatus and P. tunicatus is unknown due to approximately 135 μm thick, composed of thin- to slightly
a lack of sequences from the latter species. thick-walled (up to 1 μm) hyphae 4–10 μm wide, yellowish
Mycol Progress (2021) 20:1243–1273 1257

Phylloporus bogoriensis
(FHMU459). a Basidia. b
Basidiospores. c Cheilocystidia. d
Pleurocystidia. e Pileipellis. f
Stipitipellis. Scale bars = 10 μm.
Drawings by L.L. Wu
or yellowish brown in KOH; terminal cells 30–52 × 7–10 μm, Comments: Our molecular phylogenetic analyses indi-
narrowly clavate or subcylindrical, with obtuse apex. Stipe cated that one new Chinese collection and one specimen
trama composed of longitudinally arranged, parallel hyphae of P. bogoriensis from Indonesia, which was identified by
4–15 μm wide, cylindrical, thin- to slightly thick-walled (up to Neves et al. (2012), grouped together with strong statisti-
1 μm), yellowish in KOH. Clamp connections absent in all cal support (Fig. 2). Morphologically, our new collection
tissues. also has the unusual feature of a red to black color change
Habitat: Solitary or scattered on the ground in forests dom- of the context when injured, which matches with the de-
inated by Pinus latteri Mason scriptions of specimens of P. bogoriensis from southeast-
Known distribution: Southern China, Malaysia, Singapore ern Asia (Corner 1970). We also noticed that some mor-
(Corner 1970), and Indonesia (Höhnel 1914; Neves et al. phological features are different between the material
2012) from China and that from southeastern Asia. For example,
Specimen examined: CHINA. Hainan Province: the Chinese collection has a blue to red to black color
Wuzhishan City, campus of Qiongzhou College, elev. 350 change of the hymenophore when injured; whereas,
m, 30 July 2010, N.K. Zeng761 (FHMU459). southeastern Asian collections just have a blue color
1258 Mycol Progress (2021) 20:1243–1273
change (Corner 1970). Moreover, the cystidia of the collec- dry, densely covered with pale brown, brown to dark brown
tions from southeastern Asia are longer (60–150 × 7–20 μm). squamules; context approximately 0.5 cm in thickness in the
Although there are some differences between them, the center of the pileus, yellowish, changing bluish when injured.
Chinese collection is identified as P. bogoriensis according Hymenophore lamellate, decurrent. Lamellae up to 0.4 cm
to the present data. Phylogenetically, P. bogoriensis is closely broad, subdistant, commonly anastomosing, changing bluish
related to P. rufescens Corner (Fig. 2). However, P. rufescens when injured; lamellulae common, attenuate, concolor with
has the surface of the pileus and the stipe tinged with red, lamellae. Stipe 3–4 × 0.5–0.7 cm, central, subcylindrical; sur-
crowded lamellae, and a blue to red to black color change of face dry, densely covered with reddish brown squamules; con-
the context when injured (Corner 1970; Zeng et al. 2013). text yellowish, changing bluish when injured; annulus absent.
Basal mycelium yellowish. Odor not distinct.
Phylloporus castanopsidis M.A. Neves & Halling, Fungal Basidiospores [20/1/1] 10–14.5 × 4–5.5 μm, Q = 1.82–
Divers. 55: 113, 2012 3.38, Qm = 2.46 ± 0.33, subfusoid and inequilateral in side
Figures 3g–h, 4c, and 7 view with slightly suprahilar depression, subfusoid in ventral
Basidiomata small-sized. Pileus approximately 4.3 cm in view, slightly thick-walled (up to 0.5 μm), yellowish to
diam, convex to applanate, center slightly depressed; surface

Phylloporus castanopsidis
Drawings by L.L. Wu
Mycol Progress (2021) 20:1243–1273 1259
yellowish brown in KOH, smooth under the light microscope Etymology: Latin epithet “grossus” refers to the walls of
but with bacillate ornamentation under SEM. Basidia 26–31 the cystidia which are thick.
× 8–10.5 μm, clavate, thin-walled, 4-spored, yellowish in Holotype: CHINA. Hunan Province: Yizhang County,
KOH; sterigmata 4–5 μm in length. Hymenophoral trama Mangshan National Nature Reserve, elev. 1300 m, 29
phylloporoid, composed of slightly thick-walled (up to 1 μm) July 2019, N.K. Zeng 4159 (FHMU3277)
hyphae, 4.5–18.5 μm wide, colorless in KOH. Cheilocystidia Basidiomata small to medium-sized. Pileus 2.5–7.5 cm in
65–78 × 12–16 μm, fusiform or subfusiform, thin-walled, diam., convex to applanate, center slightly depressed, margin
colorless in KOH, no encrustations. Pleurocystidia 89–115 inrolled; surface dry, subtomentose, then squamulose, pale
× 14–17 μm, fusiform or subfusiform, thin- to slightly thick- brown (5B2), brown (6B2) to dark brown (8E5); context
walled (up to 1 μm), colorless in KOH, no encrustations. 0.2–0.8 cm in thickness in the center of the pileus, yellowish
Pileipellis a trichoderm 80–120 μm thick, composed of hy- white (1A2), unchanging in color when injured.
phae more or less vertically arranged, thin- to slightly thick- Hymenophore lamellate, decurrent. Lamellae up to 0.6cm
walled (up to 1 μm), 8–17 μm-wide, deep yellow to yellowish broad, subdistant, commonly anastomosing, yellow (1A6),
brown in KOH; terminal cells 25–58 × 9–17 μm, clavate or unchanging in color when injured; lamellulae common, atten-
subcylindrical, sometimes subfusiform, with obtuse apex. uate, concolor with lamellae. Stipe 4–5 × 0.4–0.7 cm, central,
Pileal trama made up of hyphae 6–16 μm in diam., thin- to subcylindrical; surface dry, densely covered with pale brown
slightly thick-walled (up to 1 μm), yellowish in KOH. (5B2), brown (6B2) to reddish brown (9D6) squamules; con-
Stipitipellis a hymeniform approximately 125 μm thick, com- text yellowish white (8A2), unchanging in color when injured;
posed of thin- to slightly thick-walled (up to 1 μm) emergent annulus absent. Basal mycelium white (8A1). Odor
hyphae, yellowish to dark yellow in KOH; terminal cells 15– indistinct.
50 × 6–11 μm, clavate or subfusiform. Stipe trama composed Basidiospores [80/4/4] (8–) 9–11 (–12) × 4–5.5 μm, Q
of longitudinally arranged, parallel hyphae 4–16 μm wide, = (1.73–) 1.80–2.38 (–2.67), Qm = 2.15 ± 0.26, subfusoid
cylindrical, thin- to slightly thick-walled (up to 1 μm), yellow- to elliptical and inequilateral in side view with slightly
ish in KOH. Clamp connections absent in all tissues. suprahilar depression, subfusoid in ventral view, slightly
Habitat: Solitary on the ground, in forests dominated by thin-walled (up to 0.5 μm), yellowish brown in KOH,
fagaceous trees smooth under the light microscope but with bacillate or-
Known distribution: Southern China and Thailand (Neves namentation under SEM. Basidia 25–35 × 6–10 μm, cla-
et al. 2012) vate, thin-walled, 4-spored, colorless to yellowish in
Specimen examined: CHINA. Hainan Province: KOH; sterigmata 3–5 μm in length. Hymenophoral
Yinggeling of Hainan Tropical Rainforest National Park, elev. trama phylloporoid, composed of slightly thick-walled
650 m, 31 July 2015, N.K. Zeng2278 (FHMU1517). (up to 1 μm) hyphae, 5–20.5 μm wide, colorless in
Comments: One Chinese collection and the holotype of KOH. Cheilocystidia 40–72 × 7–22 μm, abundant,
P. castanopsidis, a species originally described from subclavate, subfusiform or fusiform, thin- to thick-
Thailand (Neves et al. 2012), grouped together phylogeneti- walled (up to 1.5 μm), colorless or yellowish in KOH,
cally with strong statistical support (Fig. 2), which indicates no encrustation. Pleurocystidia 55–93 × 13–20 μm,
that the specimen from China should be recognized as abundant, subclavate, fusiform or subfusiform, thin- to
P. castanopsidis. It is a species new to China. thick-walled (up to 2 μm), colorless or yellowish in
Morphologically, several features of our collection also match KOH, no encrustation. Pileipellis a trichoderm approxi-
the protologue of P. castanopsidis; however, the materials of mately 320 μm thick, composed of hyphae more or less
P. castanopsidis from Thailand have smaller basidiospores vertically arranged, 6–19 μm wide, thin- to slightly thick-
(7.7–10.5 × 3.5–4.2 μm) and smaller cystidia (47.6–59.5 × walled (up to 1 μm), colorless to yellowish brown in
9.8–12.6 μm) (Neves et al. 2012). KOH; terminal cells 20–63 × 5–24 μm, clavate or
subcylindrical, with obtuse apex. Pileal trama made up
Phylloporus grossus N.K. Zeng, L.L. Wu & P. Zhang, sp. of hyphae 4–25 μm in diam., thin- to slightly thick-walled
nov. (up to 1 μm), colorless in KOH. Stipitipelli a hymeniform
Figures 3i–k, 4d, and 8 40–50 μm thick, composed of thin- to slightly thick-
MycoBank: MB838777 walled (up to 1 μm), golden yellowish in KOH, emergent
Diagnosis: Phylloporus grossus is characterized by a hyphae; terminal cells 20–85 × 5–10 μm, clavate,
brown pileus; a hymenophore that is unchanging in color subclavate, subfusiform or subcylindrical. Stipe trama
when injured; a stipe densely covered with pale brown, brown composed of longitudinally arranged, parallel hyphae 3–
to reddish brown squamules; white basal mycelium; thick- 14 μm wide, cylindrical, thin- to slightly thick-walled (up
walled cystidia (up to 2 μm) and a pileipellis composed of to 1 μm), colorless in KOH. Clamp connections absent
more or less inflated hyphae. in all tissues.
1260 Mycol Progress (2021) 20:1243–1273

Phylloporus grossus
Wu
Habitat: Solitary or gregarious on the ground in forests P. centroamericanus has a small basidioma, a hymenophore
dominated by fagaceous trees that changes blue when injured, longer basidiospores (8–16 ×
Known distribution: Central and southern China 3.5–5.5 μm), longer cystidia (up to 138 μm), and a distribu-
Other specimens examined: CHINA. Hainan Province: tion in North/Central America (Neves and Halling 2010);
Yinggeling of Hainan Tropical Rainforest National Park, elev. P. pachycystidiatus has longer basidiospores (11–14 × 4.5–5
650 m, 6 May 2018, N.K. Zeng3309, 3334 and 3335 μm) and longer cystidia with thicker walls (up to 4 μm) (Zeng
(FHMU2270, FHMU2937 and FHMU3136); Limushan of et al. 2013); P. rubiginosus has a hymenophore and context
Hainan Tropical Rainforest National Park, elev. 850 m, 3 that turn bluish olivaceous, then red and finally black, and
June 2009, N.K. Zeng239 (FHMU2469). Hunan Province: smooth, longer basidiospores (10–14 × 4.5–5.5 μm) (Neves
Yizhang County, Manshan National Nature Reserve, elev. et al. 2012; Zhang et al. 2019); P. tunicatus has a small
1200 m, 29 July 2019, N.K. Zeng4158 (FHMU3280). basidioma, a context that turns blue when injured, longer ba-
Comments: Phylloporus alboinfuscatus (see above), sidiospores (10.5–14 × 4.3–5 μm) and longer cystidia (Corner
P. centroamericanus, P. pachycystidiatus, P. rubiginosus 1970). The morphological differences between P. grossus and
and P. tunicatus also have thick-walled cystidia (Corner P. alboinfuscatus have been discussed above. Moreover,
1970; Singer and Gómez 1984; Neves and Halling 2010; P. grossus is genetically distant from P. alboinfuscatus,
Neves et al. 2012; Zeng et al. 2013). However, P. centroamericanus, P. pachycystidiatus, and
Mycol Progress (2021) 20:1243–1273 1261
P. rubiginosus (Fig. 2). The phylogenetic relationship be- brown; context 0.2–0.9 cm in thickness in the center of the
tween P. grossus and P. tunicatus is unknown due to a lack pileus, yellowish white, unchanging in color when injured.
of sequences from the latter species. In addition, phylogenetic Hymenophore lamellate, decurrent. Lamellae 0.3–1-cm
analyses also demonstrated that P. grossus is closely related to broad, subdistant, commonly anastomosing, yellow, unchang-
one collection, misidentified as P. brunneiceps, from eastern ing in color when injured; lamellulae common, attenuate,
China (Fig. 2); the morphological differences between them concolor with lamellae. Stipe 5–8 × 0.4–1.5 cm, central,
are unclear because the morphological features of the material subcylindrical; surface dry, covered with reddish brown
identified as P. brunneiceps are unknown. squamules; context yellowish white, unchanging in color
when injured. Basal mycelium white. Odor indistinct.
Phylloporus luxiensis M. Zang, Acta Microbiol. Sin. Basidiospores [160/8/8] 8–12 × 4–5.5 μm, Q = 1.80–
18(4): 283, 1978 2.63, Q m = 2.16 ± 0.26, subfusoid to elliptical and
Figures 3l–n, 4e, and 9 inequilateral in side view with slightly suprahilar depres-
Basidiomata small-, medium- or large-sized. Pileus 2.3– sion, subfusoid in ventral view, slightly thick-walled (up
9.5 cm in diam., convex to applanate, center slightly de- to 0.5 μm), yellowish to yellowish brown in KOH,
pressed, margin decurved; surface dry, subtomentose, some- smooth under the light microscope but with bacillate or-
times slightly rimose in age, pale brown, brown to dark namentation under SEM. Basidia 30–38 × 9.5–11.5 μm,

Phylloporus luxiensis
Drawings by L.L. Wu
1262 Mycol Progress (2021) 20:1243–1273
clavate, thin-walled, 4-spored, colorless to yellowish in relationship between P. luxiensis and P. dimorphus should
KOH; sterigmata 3–5 μm in length. Hymenophoral be further studied.
trama phylloporoid, composed of slightly thick-walled
(up to 1 μm) hyphae, 5–16 μm wide, colorless in KOH. Phylloporus microsquamus N.K. Zeng, L.L. Wu, S. Jiang
Cheilocystidia 40–93 × 9–25 μm, abundant, subfusiform & Zhi. Q. Liang, sp. nov.
or fusiform, colorless or yellowish in KOH, no encrusta- Figures 10a–b, 4f, and 11
tions. Pleurocystidia 61–93 × 13–21 μm, abundant, fusi- MycoBank: MB838778
form or subfusiform, colorless or yellowish in KOH, no Diagnosis: Phylloporus microsquamus is characterized by
encrustations. Pileipellis a trichoderm 130–135 μm thick, a pileus covered with reddish brown to brownish red, minute
composed of slightly interwoven, colorless to yellowish squamules; a yellowish white context that is unchanging in
brown in KOH, 5–9 μm wide, thin- to slightly thick- color when injured; a hymenophore that turns bluish when
walled (up to 1 μm) hyphae; terminal cells 28–48 × 6–8 injured; a stipe covered with reddish brown to brownish red
μm, clavate or subcylindrical, with obtuse apex. Pileal squamules at the middle part and a pileipellis composed of
trama composed of 3–10 μm wide, colorless in KOH, uninflated hyphae.
slightly thick-walled (up to 1 μm) hyphae. Stipitipellis a Etymology: Latin epithet “microsquamus” refers to the
hymeniform approximately 205 μm thick, composed of pileal surface that is covered with minute squamules.
thin- to slightly thick-walled (up to 1 μm) hyphae 4–16 Holotype: CHINA. Hainan Province: Yinggeling of
μm wide, deep yellow to yellowish brown in KOH; ter- Hainan Tropical Rainforest National Park, elev. 820 m, 5
minal cells 25–45 × 6–15 μm, clavate or subcylindrical, August 2015, N.K. Zeng 2571 (FHMU1678)
with obtuse apex. Stipe trama composed of longitudinal- Basidiomata small- to medium-sized. Pileus 1–6 cm in
ly arranged, parallel hyphae 6–15 μm wide, cylindrical, diam., convex, then applanate, finally center slightly de-
thin- to slightly thick-walled (up to 1 μm), colorless in pressed; margin sometimes slightly inrolled; surface pale yel-
KOH. Clamp connections absent in all tissues. lowish (30A2), covered with reddish brown (7C4) to brown-
Habitat: Solitary or occasionally caespitose on the ground, ish red, usually conical, minute squamules; context 0.2–
in forests dominated by fagaceous trees 0.4 cm in thickness in the center of the pileus, yellowish white,
Known distribution: Southwestern, central, southeastern unchanging in color when injured. Hymenophore lamellate,
and southern China and Thailand (Neves et al. 2012) decurrent. Lamellae 0.15–0.6 cm broad, subdistant, common-
Specimens examined: CHINA. Fujian Province: ly anastomosing, yellow (30A3), changing bluish quickly
Zhangping County, Xinqiao Town, Chengkou Village, elev. when injured; lamellulae common, attenuate, concolor with
350 m, 3 August 2013, N.K. Zeng1364 (FHMU915). Hainan lamellae. Stipe 2–6 × 0.2–0.7 cm, central, subcylindrical;
Province: Jianfengling of Hainan Tropical Rainforest National apex often ribbed, middle part covered with minute, reddish
Park, elev. 870 m, 28 June 2018, N.K. Zeng3439, 3442 and brown (7D7) to brownish red squamules, base villous with
3444 (FHMU3095, FHMU3101, and FHMU3105). Hunan mycelium; context yellowish white, unchanging in color
Province: Yizhang County, Mangshan National Nature when injured; annulus absent. Basal mycelium white. Odor
Reserve, elev. 800 m, 30 July 2019, N.K. Zeng4196 not distinct.
(FHMU3281). Yunnan Province: Yimen, bought from mar- Basidiospores [280/14/8] (7–) 8–11 × 4–5.5 μm, Q =
ket, 16 August 2016, N.K. Zeng2784 (FHMU1773); Wuding (1.40–) 1.56–2.50 (–2.75), Qm = 2.06 ± 0.24, subfusoid to
County, bought from market, 19 August 2016, N.K. elliptical and inequilateral in side view with slightly suprahilar
Zeng2789 (FHMU2935); Yuxi City, Eshan County, bought depression, subfusoid in ventral view, slightly thick-walled
from market, 15 July 2018, N.K. Zeng3642 (FHMU2996). (up to 1 μm), pale yellowish brown in KOH, smooth under
Comments: The concept of P. luxiensis, a species originally the light microscope but with bacillate ornamentation under
described from Yunnan Province of southwestern China, has SEM. Basidia 31–36 × 7–10.5 μm, clavate, thin-walled, 4-
been previously defined (Zeng et al. 2011). Besides south- spored; colorless to yellowish in KOH; sterigmata 3–6 μm in
western China, the present study indicates that P. luxiensis length. Hymenophoral trama phylloporoid, composed of
also occurs in central, southeastern, and southern China. We slightly thick-walled (up to 1 μm) hyphae, 4–15 μm wide,
also noted that the holotype of P. dimorphus from Thailand colorless in KOH. Cheilocystidia 60–87 × 10–17 μm,
and Chinese collections of P. luxiensis grouped together with subfusiform or fusiform, thin-walled, colorless to pale yellow
strong statistical support (Fig. 2); however, the holotype of the in KOH, no encrustations. Pleurocystidia 88–110 × 14–23
Thai species was not nested inside the lineage of P. luxiensis μm, fusiform or subfusiform, thin-walled, colorless to pale
as in a previous study (Zeng et al. 2013). Moreover, the yellow in KOH, no encrustations. Pileipellis a trichoderm
hymenophore and pileal context of P. dimorphus stain blue approximately 230 μm thick, composed of thin-walled hy-
(Neves et al. 2012); whereas, those of P. luxiensis do not phae 7.5–13 μm wide, yellowish to brilliant yellow in KOH;
change in color when injured. Thus, the taxonomic terminal cells 20–40 × 8–13 μm, narrowly subcylindrical,
Mycol Progress (2021) 20:1243–1273 1263
Fig. 10 Basidiomata of
Phylloporus species. a, b P.
microsquamus (FHMU1678,
holotype). c, d P. nigrisquamus
(FHMU3271, holotype). e–g P.
nigrobrunneus (FHMU3268,
holotype). h, i P. subrubeolus
(FHMU3273). j–l P. tenuissimus
(j, k from FHMU2964, holotype;
l from KUN-HKAS 74689).
Photos: a, b, h–l N.K. Zeng; c–g,
Y.G. Fan
with obtuse apex. Pileal trama made up of hyphae 7–15 μm August 2015, N.K. Zeng2523 (FHMU1640); Limushan of
in diam., thin-walled, colorless to yellowish in KOH. Hainan Tropical Rainforest National Park, elev. 850 m, 20
Stipitipellis a hymeniform approximately 215 μm thick, com- May 2009, N.K. Zeng189 (FHMU2467) and 26 June 2015,
posed of 5–13.5 μm wide, cylindrical hyphae, with narrowly N.K. Zeng 1990 (FHMU1285). Guangxi Zhuang
clavate terminal cells 38–70 × 8–11 μm, yellow to yellowish Autonomous Region: Nanning City, Damingshan National
brown in KOH. Stipe trama composed of longitudinally ar- Nature Reserve, elev. 1200 m, 16 July 2015, N.K.
ranged, parallel hyphae 6–13 μm wide, cylindrical, thin- to Zeng2154 (FHMU1416).
slightly thick-walled (up to 1 μm), colorless to yellowish in Comments: Morphologically, P. microsquamus is similar
KOH. Clamp connections absent in all tissues. to P. bellus, P. pruinatus Kuan Zhao & N.K. Zeng, and
Habitat: Solitary to gregarious on the ground, in forests P. rubeolus N.K. Zeng, Zhu L. Yang & L.P. Tang.
dominated by fagaceous trees However, P. bellus, a species originally described from
Known distribution: Southwestern and southern China Singapore, has a villous then subtomentose pileus and a
Specimens examined: CHINA. Yunnan Province: Jinping pileipellis composed of more or less inflated hyphae (Corner
County, Fenshuiling National Nature Reserve, elev. 2200 m, 1970; Zeng et al. 2013); P. pruinatus has a smaller basidioma
12 July 2018, N.K. Zeng3549, 3551 and 3556 (FHMU3012, (pileus 2–3 cm), a pileus with a thin pruina and a pileipellis
FHMU3064, and FHMU3018). Hainan Province: Yinggeling composed of more or less inflated hyphae (Zhao et al. 2018);
of Hainan Tropical Rainforest National Park, elev. 820 m, 4 P. rubeolus has a tomentose pileus, narrow pleurocystidia and
1264 Mycol Progress (2021) 20:1243–1273

Phylloporus microsquamus
Wu
a pileipellis composed of inflated hyphae with narrow termi- Phylloporus nigrisquamus N.K. Zeng, L.L. Wu & Y.G.
nal cells (Zeng et al. 2013). Moreover, P. microsquamus is Fan, sp. nov.
genetically distant from P. bellus, P. pruinatus, and Figures 10c–d, 4g, and 12
P. rubeolus (Fig. 2). MycoBank: MB838779
Phylogenetically, P. microsquamus is related to Diagnosis: Phylloporus nigrisquamus is characterized by a
P. purpurellus Singer and P. rimosus Bandala, Montoya pileus covered with brownish black to black squamules, a
& Garay (Fig. 2). However, P. purpurellus has a tiny stipe densely covered with reddish brown squamules, a
basidioma (pileus up to 2.6 cm), narrower basidiospores hymenophore that does not change color when injured and
(7.5–11.3 × 3.3–4 μm) and shorter and narrower cystidia long cystidia.
(48–65 × 8.5–12 μm) (Singer 1973); P. rimosus has a Etymology: Latin epithet “nigrisquamus” refers to the
hymenophore that changes blue then reddish when injured, pileal surface that is covered with black squamules.
a yellow context that stains pinkish or pinkish-brown, large Holotype: CHINA. Yunan Province: Baonshan City,
basidiospores (9.5–14 × 3.5–5 μm), and a distribution in Baihualing Ecotourism Zone, elev. 2200 m, 9 June 2018,
Mexico (Montoya et al. 2019). Y.G. Fan 2819 (FHMU3271)
Mycol Progress (2021) 20:1243–1273 1265

Phylloporus nigrisquamus
Wu
Basidiomata small-sized. Pileus 2.3–3 cm in diam., bacillate ornamentation under SEM. Basidia 30–34 × 9–10
infundibuliform; surface dry, yellow (3A7), covered with μm, clavate, thin-walled, 4-spored, colorless to yellowish in
brownish black (6F4) to black squamules; context 0.2– KOH; sterigmata 3.5–5.5 μm in length. Hymenophoral
0.5 cm in thickness in the center of the pileus, yellowish trama phylloporoid, composed of slightly thick-walled (up
(1A2), unchanging in color when injured. Hymenophore la- to 1 μm) hyphae, 5–20 μm wide, colorless in KOH.
mellate, decurrent. Lamellae up to 0.5 cm broad, subdistant, Cheilocystidia 70–100 × 11–19 μm, abundant, thin- to slight-
yellow (3A6), unchanging in color when injured; lamellulae ly thick-walled (up to 1 μm), colorless in KOH, no encrusta-
common, attenuate, concolor with lamellae. Stipe 1.2–1.7 × tions. Pleurocystidia 125–175 × 14–18 μm, abundant, fusi-
0.3–0.5 cm, central, subcylindrical; surface dry, densely cov- form or subfusiform, thin- to slightly thick-walled (up to 1
ered with reddish brown (6C8) squamules; context pale yel- μm), colorless to yellowish in KOH, no encrustations.
low (1A2), unchanging in color when injured; annulus absent. Pileipellis a trichoderm approximately 270 μm thick, com-
Basal mycelium white (8A1). Odor not distinct. posed of hyphae more or less vertically arranged, yellowish
Basidiospores [40/2/2] (8.5–) 9–11.5 (–12) × 4–5 (–5.5) brown in KOH, 7–15 μm wide, thin- to slightly thick-walled
μm, Q = (1.80–) 2.00–2.50 (–2.63), Qm = 2.25 ± 0.17, (up to 1 μm); terminal cells 40–85 × 10–16 μm, clavate or
subfusoid to elliptical and inequilateral in side view with subcylindrical, with obtuse apex. Pileal trama made up of
slightly suprahilar depression, subfusoid in ventral view, hyphae 5–18 μm in diam., thin- to slightly thick-walled (up
slightly thick-walled (up to 0.5 μm), yellowish to yellowish to 1 μm), colorless to yellowish in KOH. Stipitipellis a
brown in KOH, smooth under the light microscope but with hymeniform 60–80 μm thick, composed of thin- to slightly
1266 Mycol Progress (2021) 20:1243–1273
thick-walled (up to 1 μm) hyphae 9–14 μm wide, yellowish to central, subcylindrical; surface dry, densely covered with yel-
brownish yellow in KOH; terminal cells 26–40 × 6–13 μm, low to yellowish brown squamules (2A2); context yellowish
clavate, with suborbicular apex. Stipe trama composed of white (8A1), unchanging in color when injured; annulus ab-
longitudinally arranged, parallel hyphae 3–15 μm wide, cy- sent. Basal mycelium white (8A1). Odor not distinct.
lindrical, thin- to slightly thick-walled (up to 1 μm), colorless Basidiospores [60/3/3] (8.5–) 9–11.5 (–12) × 3–4.5 (–5)
to yellowish in KOH. Clamp connections absent in all μm, Q = (2.00–) 2.33–3.29 (–3.5), Q m = 2.73 ± 0.27,
tissues. subfusoid and inequilateral in side view with slightly
Habitat: Solitary on the ground in forests dominated by suprahilar depression, subfusoid in ventral view, slightly
fagaceous trees thick-walled (up to 0.5 μm), yellow to yellowish brown in
Known distribution: Southwestern China KOH, smooth under the light microscope but with bacillate
Other specimen examined: CHINA. Yunnan Province: ornamentation under SEM. Basidia 28–38 × 9–10 μm, cla-
Baoshan City, Baihualing Ecotourism Zone, elev. 2200 m, 9 vate, thin-walled, 4-spored, colorless to yellowish in KOH;
June 2018, Y.G. Fan2834 (FHMU3272) sterigmata 3–5 μm in length. Hymenophoral trama
Comments: Interestingly, one collection identified as phylloporoid, composed of slightly thick-walled (up to 1
P. leucomycelinus (REH4582) from the USA occurred in the μm) hyphae, 4.5–16 μm wide, colorless in KOH.
lineage of P. nigrisquamus (lineage 16) (Fig. 2), which was Cheilocystidia 45–60 × 13–15 μm, abundant, subclavate or
also noted by Neves et al. (2012). This suggests possible con- subfusiform, thin- to slightly thick-walled (up to 1 μm), col-
tamination of the isolated DNA since the material has been orless in KOH, no encrustations. Pleurocystidia 63–88 × 11–
observed and is correctly identified. Morphologically, the un- 18 μm, abundant, subfusiform or fusiform, thin- to slightly
usual color of squamules on the pileus easily distinguishes P. thick-walled (up to 1 μm), colorless in KOH, no encrustations.
nigrisquamus from other taxa of Phylloporus. Phylloporus Pileipellis a trichoderm approximately 290 μm thick, com-
nigrisquamus is phylogenetically related to one lineage (spe- posed of hyphae more or less vertically arranged, colorless
cies level) that is composed of collections labeled as to yellowish in KOH, 9–18 μm wide, thin- to slightly thick-
“P. bellus” and “P. rhodoxanthus (Schwein.) Bres.,” which walled (up to 1 μm); terminal cells 25–69 × 9–19 μm, clavate
likely represents a new species (Fig. 2). However, this poten- or subcylindrical, with obtuse apex. Pileal trama made up of
tial new species has a dark brown to yellow-orange pileus, a hyphae 6–16 μm in diam., thin- to slightly thick-walled (up to
hymenophore and context that sometimes stain blue, shorter 1 μm), colorless to yellowish in KOH. Stipitipellis a
cystidia and a distribution in North and Central America hymeniform approximately 80 μm thick, composed of thin-
(Neves and Halling 2010). to slightly thick-walled (up to 1 μm) hyphae 5–18 μm wide,
colorless to yellowish in KOH; terminal cells 15–40 × 8–18
Phylloporus nigrobrunneus N.K. Zeng, L.L. Wu & Y.G. μm, clavate, with suborbicular apex. Stipe trama composed
Fan, sp. nov. of longitudinally arranged, parallel hyphae 4–15 μm wide,
Figures 10e–g, 4h, and 13 cylindrical, thin-walled, yellowish in KOH. Clamp
MycoBank: MB838780 connections absent in all tissues.
Diagnosis: Phylloporus nigrobrunneus is characterized by Habitat: Solitary or gregarious on the ground in forests
a pileus that is blackish brown in the center and pale brown to dominated by fagaceous trees
brown at the margin, a stipe covered with yellow to yellowish Known distribution: Southwestern China
brown squamules, narrow basidiospores, and a pileipellis Other specimens examined: CHINA. Yunnan Province:
composed of slightly inflated hyphae. Baoshan City, Baihualing Ecotourism Zone, elev. 1800 m, 8
Etymology: Latin epithet “nigrobrunneus” refers to the June 2018, Y.G. Fan2749 and 2753 (FHMU3270 and
center of the pileal surface that is colored blackish brown. FHMU3269).
Holotype: CHINA. Yunnan Province: Baoshan City, Comments: Morphologically, P. nigrobrunneus is similar
Baihualing Ecotourism Zone, elev. 1800 m, 8 June 2018, to P. arenicola A.H. Sm. & Trappe and P. catenulatus Iqbal
Y.G. Fan2754 (FHMU3268) Hosen & T.H. Li. However, P. arenicola has a pileus tinged
Basidiomata small-sized. Pileus 1.5–2 cm in diam., con- with olive and is associated with Pinus in North America
vex to applanate, infundibuliform when old; surface dry, (Neves and Halling 2010). Bangladeshi P. catenulatus has
subtomentose, blackish brown in center, pale brown to brown surfaces of the pileus and the stipe tinged with red, a
(5E7) at margin; context approximately 0.2 cm in thickness in hymenophore and context that stain blue when injured, shorter
the center of the pileus, yellowish white (2A2), unchanging in basidiospores (8–9.5 × 4–5 μm), and a pileipellis composed of
color when injured. Hymenophore lamellate, decurrent. short chains of 3–4 inflated cells (Hosen and Li 2017).
Lamellae up to 0.15 cm broad, subdistant, yellow (4A6), Phylloporus subbacillisporus Raspé, K.D. Hyde &
unchanging in color when injured; lamellulae common, atten- Chuankid, a species also described from China, has narrower
uate, concolor with lamellae. Stipe 1.4–2.6 × 0.3–0.8 cm, basidiospores as well; however, P. subbacillisporus has a
Mycol Progress (2021) 20:1243–1273 1267

Phylloporus nigrobrunneus
Wu
pileus without a black tinge, a hymenophore that stains blue commonly anastomosing, pale yellow, unchanging in color
then reddish, clavate to subclavate cystidia and basidiospores when injured; lamellulae common, attenuate, concolor with
with a smooth or merely rugulose surface under SEM lamellae. Stipe 1–2 × 0.1–0.2 cm, central, subcylindrical; sur-
(Chuankid et al. 2019). Phylogenetically, P. nigrobrunneus face dry, nearly white, densely covered with pale reddish
is not closely related to P. arenicola, P. catenulatus, or brown, minute squamules; context white, unchanging in color
P. subbacillisporus (Fig. 2). when injured; annulus absent. Basal mycelium white. Odor
not distinct.
Phylloporus subrubeolus Chuankid, K.D. Hyde & Raspé, Basidiospores [60/3/3] (5.5–) 6–7.5 (–8) × (3.5–) 4–5 (–
Mycol. Prog. 18: 609, 2019 5.5) μm, Q = (1.2–) 1.3–1.75 (–1.88), Qm = 1.50 ± 0.17,
Figures 10h–i, 4i, and 14 ellipsoid in side view with slightly suprahilar depression, el-
Basidiomata very small-sized. Pileus 1–1.5 cm in diam., lipsoid in ventral view, slightly thick-walled (up to 0.5 μm),
convex to applanate, margin decurved; surface dry, pale yellowish brown in KOH, smooth under the light microscope
brown, brown, reddish brown to dark brown; context approx- but with bacillate ornamentation under SEM. Basidia 30–36
imately 0.2 cm in thickness in the center of the pileus, white, × 9–11 μm, clavate, thin-walled, 4-spored, colorless to yel-
unchanging in color when injured. Hymenophore lamellate, lowish in KOH; sterigmata 3–4 μm in length. Hymenophoral
decurrent. Lamellae up to 0.15 cm broad, subdistant, trama phylloporoid, composed of thin- to slightly thick-
1268 Mycol Progress (2021) 20:1243–1273

Phylloporus subrubeolus
Drawings by L.L. Wu
walled (up to 0.5 μm) hyphae, 4.5–13 μm wide, colorless in hyphae; terminal cells 25–51 × 9–13 μm, clavate, with
KOH. Cheilocystidia 48–64 × 9–15.5 μm, abundant, suborbicular apex. Stipe trama longitudinally arranged, par-
subclavate or subfusiform, thin- to slightly thick-walled (up allel hyphae 4–19 μm wide, cylindrical, thin- to slightly thick-
to 1 μm), yellowish in KOH, no encrustations. Pleurocystidia walled (up to 1 μm), colorless to yellowish in KOH. Clamp
61–73 × 10–13 μm, abundant, fusiform or subfusiform, thin- connections absent in all tissues.
to slightly thick-walled (up to 1 μm), yellowish in KOH, no Habitat: Solitary on the ground in forests dominated by
encrustations. Pileipellis a trichoderm approximately 170 μm fagaceous trees
thick, composed of pale yellow to yellowish brown in KOH, Known distribution: Southern and central China and
8–13 μm wide, thin- to slightly thick- walled (up to 1 μm) Thailand (Chuankid et al. 2019)
hyphae; terminal cells 20–35 × 9–11 μm, subclavate or Specimens examined: CHINA. Hainan Province:
subcylindrical, with obtuse apex. Pileal trama made up of Yinggeling of Hainan Tropical Rainforest National Park, elev.
hyphae 5–14 μm in diam., thin-walled, yellowish to deep 650 m, 24 April 2019, N.K. Zeng 4027 (FHMU3165).
yellow in KOH. Stipitipellis a hymeniform approximately Guangdong Province: Renhua County, Danxiashan National
170 μm thick, composed of 5–13 μm wide, colorless to yel- Nature Reserve, elev. 350 m, 4 June 2019, N.K. Zeng 4095
lowish in KOH, thin- to slightly thick-walled (up to 1 μm) (FHMU3164). Hunan Province: Yizhang County, Mangshan
Mycol Progress (2021) 20:1243–1273 1269
National Nature Reserve, elev. 720 m, 30 June 2019, N.K. Diagnosis: Phylloporus tenuissimus is characterized by a
Zeng 4195 (FHMU3273). small basidioma, a brown pileus without reddish tinge, a thin
Comments: Our molecular phylogenetic analyses showed context, a hymenophore and context that turn bluish when
that these new Chinese collections and the holotype of injured, and a pileipellis composed of more or less inflated
P. subrubeolus, a species first described from Thailand hyphae.
(Chuankid et al. 2019), grouped together with strong statistical Etymology: Latin epithet “tenuissimus” refers to the pileal
support (Fig. 2). This indicates that our specimens should be context that is very thin.
recognized as P. subrubeolus (Chuankid et al. 2019). Holotype: Hainan Province: Limushan of Hainan Tropical
Morphologically, several features of our collections also Rainforest National Park, elev. 750 m, 29 June 2018, N.K.
match the protologue of P. subrubeolus; however, the speci- Zeng 3514 (FHMU2964)
mens of P. subrubeolus from Thailand have larger basidio- Basidiomata small-sized. Pileus 2–3 cm in diam.,
spores (7.2–9.7 × 4.5–5.8 μm) (Chuankid et al. 2019). applanate, sometimes center depressed; margin decurved,
sometimes uplifted; surface dry, covered with pale brown to
Phylloporus tenuissimus N.K. Zeng, L.L. Wu & Zhi Q. brown (5C6) squamules; context approximately 0.2 cm in
Liang, sp. nov. thickness in the center of the pileus, white, changing bluish
Figures 10j–l, 4j, and 15 when injured. Hymenophore lamellate, decurrent. Lamellae
MycoBank: MB838781 up to 0.5 cm broad, subdistant, commonly anastomosing,

Phylloporus tenuissimus
Wu
1270 Mycol Progress (2021) 20:1243–1273
yellow (30A3), changing bluish when bruised; lamellulae and narrower basidiospores (9.5–11.5 [–12.5] × 3.7–4.2 μm)
common, attenuate, concolor with lamellae. Stipe 1.8–4 × (Corner 1974). Phylloporus subbacillisporus, a species origi-
0.3–0.6 cm, central, subcylindrical; surface dry, densely cov- nally described from Thailand, has a blue to reddish color
ered with reddish brown (6D4) squamules; context yellowish change of the pileal context and hymenophore and narrower
white, changing bluish when injured on the upper part of the basidiospores (9.2–11.9 × 3–3.7 μm) (Chuankid et al. 2019).
stipe. Basal mycelium white (8A1). Odor not distinct. Moreover, P. tenuissimus is genetically distant from P.
Basidiospores [60/3/2] (9–) 10–12 (–12.5) × 4–5 μm, Q = castanopsidis, P. cyanescens, and P. subbacillisporus (Fig.
1.80–2.67 (–3.3), Qm = 2.38 ± 0.03, subfusoid in side view 2). The phylogenetic relationship between P. tenuissimus
with slightly suprahilar depression, subfusoid in ventral view, and P. stenosporus is unknown due to a lack of sequences
slightly thick-walled (up to 0.5 μm), yellowish to yellowish from the latter species. We also noted that P. tenuissimus is
brown in KOH, smooth under the light microscope but with closely related to P. gajari Iqbal Hosen & T.H. Li (Fig. 2);
bacillate ornamentation under SEM. Hymenophoral trama however, Bangladeshi P. gajari has a larger basidioma (pileus
phylloporoid, composed of thin- to slightly thick-walled (up to up to 6.5 cm), a pileus tinged with red when young, and a
0.5 μm) hyphae, 4–17 μm wide, colorless in KOH. Basidia thicker context (0.6–1.2 cm) (Hosen and Li 2015).
29–38 × 8–10 μm, clavate, thin-walled, 4-spored, colorless in Based on previous studies (Zeng et al. 2011, 2013; Zhao
KOH; sterigmata approximately 5 μm in length. et al. 2018; Chuankid et al. 2019; Zhang et al. 2019) and our
Cheilocystidia 22–65 × 7–16 μm, subclavate, fusiform or morphological examinations, the described species within
subfusiform, thin-walled, colorless to yellowish in KOH, no Phylloporus from China can be summarized in the following
encrustations. Pleurocystidia 56–89 × 10–20 μm, subclavate, key.
fusiform or subfusiform, thin- to slightly thick-walled (up to 1
μm), colorless in KOH, no encrustations. Pileipellis a Key to the species of Phylloporus from China
trichoderm approximately 190 μm thick, composed of hyphae 1a. Basal mycelium yellowish................................................2
more or less vertically arranged, thin- to slightly thick-walled 1b. Basal mycelium whitish....................................................6
(up to 1 μm), 5–17 μm wide, colorless to pale yellowish in 2a. Pileus brown, without reddish tinge when mature.............3
KOH; terminal cells 30–56 × 5–14 μm, clavate or 2b. Pileus reddish to reddish brown.......................................5
subcylindrical, with obtuse apex. Pileal trama made up of 3a. Hymenophore unchanging in color when
hyphae 5–16 μm in diam., thin- to slightly thick-walled (up injured.....................................................................P. luxiensis
to 1 μm), colorless in KOH. Stipitipellis a hymeniform 55–70 3b. Hymenophore turning blue when injured........................4
μm thick, composed of thin- to slightly thick-walled (up to 1 4a. Context unchanging in color when injured, stipe yellow to
μm) emergent hyphae, dark brownish yellow in KOH; termi- yellowish brown................................................P. brunneiceps
nal cells 25–60 × 8–17 μm, clavate or subclavate. Stipe 4b. Context changing bluish when injured, stipe reddish
trama composed of longitudinally arranged, parallel hyphae brown..............................................................P. castanopsidis
5–15 μm wide, cylindrical, parallel, thin- to slightly thick- 5a. Pileus 4.5–11 cm, stipe 0.3–1.5 cm diam., associated with
walled (up to 1 μm), colorless in KOH. Clamp connections subalpine to alpine tree host……...............…... P. imbricatus
absent in all tissues. 5b. Pileus 4–6.5 cm, stipe 0.4–0.7 cm diam., associated with
Habitat: Solitary on the ground, in forests dominated by subtropical or tropical tree hosts..........................P. yunnanensis
fagaceous trees 6a. Cystidia thick-walled (≥ 2 μm)............P. pachycystidiatus
Known distribution: Southern China 6b. Cystidia thin- to moderately thick-walled (≤ 2 μm)……7
Other specimen examined: Hainan Province: Ledong 7a. Cystidia moderately thick-walled (up to 2 μm)…………8
County, Jianfengling of Hainan Tropical Rainforest National 7b. Cystidia thin- to slightly thick-walled (up to 1 μm)……10
Park, elev. 870 m, 6 August 2009, N.K. Zeng 478 (KUN- 8a. Pileus brownish red to reddish, hymenophore and context
HKAS74689) turning bluish olivaceous, then turning red and finally black-
Comments: Morphologically, P. tenuissimus looks like P. ening when injured........................................... P. rubiginosus
castanopsidis, P. cyanescens (Corner) M.A. Neves & Halling, 8b. Pileus white, pale brown, brown to dark brown; context
P. stenosporus Corner and P. subbacillisporus. However, P. unchanging in color when injured; hymenophore changing
castanopsidis has a yellow basal mycelium, larger basidio- bluish or unchanging in color when injured...........................9
spores (10–14.5 × 4–5.5 μm), and longer pleurocystidia 9a. Hymenophore changing bluish when injured, basidio-
(89–115 × 14–17 μm) (Neves et al. 2012). Singaporean P. spores larger (9.5–13 × 4.5–5.5 μm)………P. alboinfuscatus
cyanescens (P. bellus var. cyanescens Corner) has a pileus 9b. Hymenophore unchanging in color when injured, basid-
tinged with red, longer cystidia, larger basidiospores (10–13 iospores smaller (9–11 × 4–5.5 μm)....................... P. grossus
× 4.5–5.5 μm), and sometimes grows on rotten wood (Corner 10a. Context pale brownish fuliginous or pale
1970; Neves et al. 2012). Malaysian P. stenosporus has a stipe fuscous...............................................................P. parvisporus
without a reddish tinge, longer cystidia (60–120 × 14–20 μm) 10b. Context whitish, cream-colored or yellowish...............11
Mycol Progress (2021) 20:1243–1273 1271
11a. Context turning reddish, then blackening or changing 33 phylogenetic species are recognized from China (Fig. 2).
blue, then red, finally blackening when injured.....................12 Among them, six are described as new (lineages 7, 9, 16, 24,
11b. Context changing bluish or unchanging in color when 29, and 30): P. alboinfuscatus, P. grossus, P. microsquamus,
injured ...................................................................................13 P. nigrisquamus, P. nigrobrunneus, and P. tenuissimus.
12a. Lamellae crowded, blue-red-black color change of Eighteen species correspond with previous morphology-
context....................................................................P. rufescens based taxa (lineages 1, 4–6, 11–14, 18–21, 23, 25, 27, 28,
12b. Lamellae subdistant, red-black color change of 31, and 32): P. bellus, P. bogoriensis, P. brunneiceps,
context................................................................P. bogoriensis P. castanopsidis, P. imbricatus N.K. Zeng, Zhu L. Yang &
13a. Hymenophore and context unchanging in color when L.P. Tang, P. luxiensis, P. maculatus N.K. Zeng, Zhu L. Yang
injured ...................................................................................14 & L.P. Tang, P. pachycystidiatus, P. parvisporus Corner,
13b. Hymenophore changing bluish when injured...............18 P. pruinatus, P. pusillus Raspé, K.D. Hyde & Chuankid,
14a. Pileus tinged with black.................................................15 P. rubeolus, P. rubiginosus, P. rubrosquamosus N.K. Zeng,
14b. Pileus tinged with red....................................................16 Zhu L. Yang & L.P. Tang, P. rufescens, P. subbacillisporus,
15a. Pileus subtomentose, cystidia shorter (up to 90 P. subrubeolus, and P. yunnanensis N.K. Zeng, Zhu L. Yang
μm)................................................................P. nigrobrunneus & L.P. Tang. The remaining ten species (lineages 2, 3, 8, 10,
15b. Pileus covered with squamules, cystidia longer (up to 15, 17, 22, 26, and 33) are not described due to paucity of
175 μm)...........................................................P. nigrisquamus material. Except for one collection in lineage 17 from temper-
16a. Basidiospores longer (≥ 8 μm).....................................17 ate areas, all of these taxa are from subtropical and tropical
16b. Basidiospores shorter (≤ 8 μm)................P. subrubeolus regions of China, showing the unexpected species diversity of
17a. Pileus with a thin, white pruina; basidiospores with Qm = Phylloporus in subtropical and tropical China. Besides these
2.23........................................................................P. pruinatus recognized species, P. megacystidiatus L. Ye, Mortimer, J.C.
17b. Pileus without a pruina, basidiospores with Qm = Xu & K.D. Hyde, P. orientalis var. brevisporus Corner, and
1.99...........................................................................P. pusillus P. orientalis var. orientalis Corner were also reported in China
18a. Pileus surface with dark-colored spots.....…P. maculatus (Chen et al. 2002; Zeng et al. 2013). Phylloporus
18b. Pileus surface without spots..........................................19 megacystidiatus, originally described from Yunnan Province
19a. Hyphae uninflated in the pileipellis..............................20 of southwestern China, is likely conspecific with
19b. Hyphae more or less inflated in the pileipellis.............21 P. yu nn an en si s b ec au s e P . m e g a c y s t i d i a t us an d
20a. Basidiospores longer (11–12.5 × 4.5–5 P. yunnanensis are in the same lineage, at the species level,
μm)............................................................P. rubrosquamosus based on ITS (data not shown). Moreover, the morphological
20b. Basidiospores shorter (8–11 × 4–5.5 differences between P. megacystidiatus and P. yunnanensis
μm)................................................................P. microsquamus provided by Ye et al. (2014) seem to be infraspecific variation.
21a. Basidiospores narrower (≤ 3.9 μm), smooth under Phylloporus orientalis var. brevisporus and P. orientalis var.
SEM............................................................P. subbacillisporus orientalis, originally described from Singapore and Malaysia,
21b. Basidiospores wider (≥ 4 μm), with bacillate ornamen- respectively, were reported as occurring in China, but this
tation under SEM..................................................................22 remains to be confirmed due to the absence of recently col-
22a. Pileus somewhat reddish, terminal cells of pileipellis with lected specimens and DNA sequences from the holotype lo-
acute apex...............................................................P. rubeolus cations for these two taxa.
22b. Pileus yellowish brown to reddish brown, terminal cells It is worth noting that the holotypes of our new species
of pileipellis with obtuse apex.....................................P. bellus P. nigrobrunneus and P. subbacillisporus grouped together
based on ITS with strong statistical support (data not shown).
Moreover, Chuankid et al. (2019) also noted that the ITS
Discussion sequences of P. subbacillisporus showed 100% similarity to
one collection of P. nigrobrunneus (“P. sp. 4,” KUN-
The fungi of Phylloporus are easily recognized by mycolo- HKAS74682). However, specimens of P. nigrobrunneus
gists at the genus level. However, species in the genus are were not grouped together with the holotype of
difficult to distinguish morphologically. Thus, only one spe- P. subbacillisporus based on our concatenated dataset (28S
cies of Phylloporus was described from China before 2013 + ITS + TEF1) and instead formed an independent lineage
(Zang and Zeng 1978; Li et al. 1992; Bi et al. 1997). With (lineage 24) (Fig. 2). Morphological differences between col-
the rapid development of molecular phylogenetic analyses, lections of P. subbacillisporus and P. nigrobrunneus were
many taxa in the genus have been reevaluated or newly de- also observed (see above). Thus, the new taxon
scribed, enabling us to better understand the diversity of the P. nigrobrunneus was proposed.
genus in China (Zeng et al. 2011, 2013; Zhao et al. 2018; Interestingly, our molecular data showed that the collec-
Chuankid et al. 2019; Zhang et al. 2019). In the present study, tions named P. castanopsidis occurred in two different parts
1272 Mycol Progress (2021) 20:1243–1273
of the tree (Fig. 2). The holotype of P. castanopsidis was Science Foundation of China (Nos. 31760008 and 31560005), and the
Innovative Research Projects for Graduate Students in Hainan Province,
nested in lineage 11 (Fig. 2), indicating that this lineage cor-
China (No. Hys2019-285).
responds with true P. castanopsidis; whereas, the other line-
age that included collections of P. castanopsidis should be Data availability The sequence data generated in this study are deposited
redefined (Raspé et al. 2016; Chuankid et al. 2019). in NCBI GenBank.
Moreover, the molecular data indicated that North American
P. rhodoxanthus occurred in three different parts of the tree Declarations
(Fig. 2). The true P. rhodoxanthus was previously defined
(Neves and Halling 2010; Neves et al. 2012) and included in Conflict of interest The authors declare no competing interests.
our molecular tree (Fig. 2); whereas, the collection of
P. rhodoxanthus from Europe likely represents an
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Updated Taxonomy of Chinese Phylloporus (Boletaceae, Boletales) Six New Taxa and Four Redescribed Species

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Updated Taxonomy of Chinese Phylloporus (Boletaceae, Boletales) Six New Taxa and Four Redescribed Species

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Mycological Progress (2021) 20:1243–1273

Updated taxonomy of Chinese Phylloporus (Boletaceae, Boletales):

Received: 23 February 2021 / Revised: 18 June 2021 / Accepted: 21 June 2021

Keywords Bolete . Molecular phylogeny . Morphology . New taxa . Taxonomy

Section Editor: Zhu-Liang Yang

Phylloporus Phylloporus REH7906 Panama JQ003662 JQ003638 — Neves et al. (2012)

Phylloporus — N.K. Zeng3642 Yunnan, SW MW588609 — MW574466 Present study

Phylloporus — N.K. Zeng3551 Hainan, MW588596 MW588652 MW574470 Present study

Phylloporus pusillus Phylloporus pusillus OR1158 Thailand — — MH580803 Chuankid et al.

Phylloporus — N.K. Zeng1904 Hainan, — MW588645 MW574453 Present study

Fig. 5 Microscopic features of

Fig. 6 Microscopic features of

Fig. 7 Microscopic features of

Fig. 8 Microscopic features of

Fig. 9 Microscopic features of

Fig. 11 Microscopic features of

Fig. 12 Microscopic features of

Fig. 13 Microscopic features of

Fig. 14 Microscopic features of

Fig. 15 Microscopic features of

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